THE

American Naturalist

NATURAL HISTORY

BOSTON, U. S. A.
GINN & COMPANY, PUBLISHERS

Mo.Bot.Garaen
1908

INDEX

INDEX

Miller's Families and Genera

of Bats 671

MiUikan and Gale's Physics . 41
Molisch's Purpurbakterien . 541
Montgomery's Racial Descent 719
Norton's Elements of Geology 409
Punnett's Mendelism . . .329
Rich's Feathered Game . . 726
Ridgway's Birds of North and

Middle America .... 672
Scharff's European Animals •.
Schvvarz's Longleaf Pine . . 539
Skeat and Blagden's Pagan
Races of the Malay Penin-

de Vries' Arten und Varietaten 189
Yerkes' Dancing Mouse . . 786
Breeding habits of Amhlystoma

ofSalamandra . . . .467

of ...... ^ .. 342

Cambarus, Attached young of . 253
Campbell, D. W. Studies on the

Ophioglossaceae 139

Recent studies on gymno-

sperms 801

Capitalization of specific names 525

Cataloguing museum specimens 77

ply ^.391

Chaetognathn, Distribution of . 241

Chalicotheres 733

Chrysler. M. A. See Jeffrey, E.

C. and — 355

Chubs' nests 323, 468

Cilia, Structure of 545

Color vision. The problem of . 365
Courtis, S. A. Response of toads

to sound 677

Crayfish, Attached young of . 253
Cushman, J . A. Ostracoda from

Dane, J. M. The problem of

Davenport, C. B. Modernized
Darwinism 44

Xenia in wheat .... 47

Davis. W. T. Hyla andersonii
and Rana virgatipes in New
Jersey 49

Dearborn, G. V. N. Hough and
Sedgwick's Physiology . . .194

Dillingham, F. T. The staff-
former food supply of starving
Indians 391

Distribution of related species

207, 241. 653

Diving beetles 477

Dumford, C. D. The flying-
fish problem 65

Earthworms as planters of trees 71 1

Eigenmann, C. H. Fowler's
" Heterognathous Fishes " with
a note on the Stethaprioninae . 767

Eliot, C. W. Agassiz Centen-
nial 413

Embryos, Specific characters in . 589

Entomology, Notes on

532, 597, 727, 798

Ernst, H. C. Molisch's Pur-
purbakteria 541

Exhibits at the 7th Int. Zool.
Congress 657

Fi^rasfer afflnis, Uahit^ oi . . 1
Flying-fish problem . . . 65, 347
Fowler, H. W. Records of
Pennsylvania fishes ... 5

Gastropoda, Orthogenetic vari-

Geographic distribution of close-
ly related species 207, 241, 653
Gill, T. Stone-gathering fishes 468
Gonionemus, Behavior of . . 683
Goodale, G. L. liurkett and

Poe's Cotton 405

Grabau. A. W. ( )vUu>iii-m'tu-

variuticn in ( la^tn.iHMla . . 607
Graphic inctluKl ..f ccnclating

INDEX

INDEX

Ritter, W. E. The significant j Telexomus, New species of . .

results of a decade's study of ! Termite nests, Fungi of . .

Rivellia boscii. Mating habit of . 465 | kulente Piuphorbien . . .

Rotifers, South American . . 97 i Toads, Response to soiuid . .

Ruggles, A. G. See Matheson, i Trelease, W. Bergen and Davis's

R. and • 567 Botany

Ruthven, A. G. Note on the : liotanical notes 53, 344,

garter snal

iology c

TOL. XLI, NO. 481

JANUARY, 1907

THE

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

I. Hote on the Habits of Fierasfer .... PE0FES80R EDWIN LINTON 1

n. Eecordi of Pennsylvania Fishes ...... HENEY W. FOWLEB 6

in. Specific Name of Nectnrus maculosus .... PSOFESSOK F. C. WAITE 23

IV- Volvox for laboratory nse BEETRAM G. SMITH 31

V. Ostracoda from Southeastern Massachusetts .... J. A. CUSHMAN 35
VI. Notes and Literature: Physics, A First Course in Physics; Biology, Jennings'
Behavior of the Lower Organisms, Modernized Darwinism; Momentum
in Variation, Xenia in Wheat; Zoology. Folsom's Entomology, Observa-
tions on Hyla and Rana virgatipes; Zoological Laboratory Notes,
Notes; Boiany. Bergen and Davis's Principles of Botany, Rydberg's

Flora of Colorado, Notes 41

Vn. PublicationB received 67

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The American Naturalist

ASSOCIATE EDITORS
J. A. ALLEN, Ph.D., American Museum of Natural History, New York
E. A. ANDREWS, Ph.D., Johns Hopkins University, Baltimore
WILLIAM S. BAYLEY, Ph.D., Colby University, Waterville
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell UniversUy. Ithaca.
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA, M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University
CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM, Ph.D., Lake Forest University
ARNOLD E. ORTMANN, Ph.D., €ame^ie Museum, Pittsburg
D. p. PENHALLOW^, D.Sc, F.R.S.C., McGUl University, Montreal
H. M. RICHARDS, S.D., Columbia University, New York
W. E. RITTER, Ph.D., University of California, Berkeley
ERWIN F. smith, S.D., U. S. DepaHment of Agriculture, Washington
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington
W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis
HENRY B. WARD, Ph.D., University of Nebraska, Lincoln
WILLIAM M. WHEELER, Ph.D., American Museum of Natural History,
New York

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THE

AMERICAN NATURALIST

Vol. XLI January, 1907 No. 481

xrm: ox the HAurrs of Fn:u.\sFi:n afiisis

2 THE AMERICAN NATURALIST [Vol. XLI

in diameter, was inverted over the animal, the edgeg resting on the
dorsal side and about 30 milhmeters from each end. Instead of
making any effort to escape the animal retreated beneath the finger-
bowl and at the expiration of 30 minutes had made no effort to
escape.

At this point in the experiment, my patience proved inferior to
that of the holothurian" and I lifted the finger-bowl. When this
was done a specimen of the interesting genus Fierasfer, which I
have identified as F. affinis, was found. Evidently it had been
driven to leave its host on account of the deficient supply of oxygen.
The fish, which was nearly transparent, measured 124 millimeters
in length, and was very slender, tapering almost uniformly from
the head to the tip of the long and whip-like tail.

As soon as the finger-bowl was removed the fisli beijan to swim

actively about in the dish. It kept its nose close to the surface of
the water, and at times even thrust its head above the surface in
its eagerness to get oxygen.

After a few minutes it ceased to swim at the surface but appeared
to be still uneasy. No test was made to prove what seemed to be
indicated by its actions, viz., that its sense of sight is defective.
Judging from its subsequent behavior it was even then trying to
find its customary quarters, but, to those of us who were watching,
its movements began to be somewhat aimless.

No. 481]

HABITS OF FIERASFER

3

At last, and, so far as I was able to judge, by accident, its nose
came in contact with the holothurian near the anterior end. Imme-
diately the fish appeared to become excited and he<j;im to feel its
way back toward the posterior end of rlic liDloiliiiilaii. In its
procuress it bumped its nose rapidly a<.'-aiiist ihc side of the passive
holothurian, and, as if following a scent, prociu^lcd without any
pause or regression toward the cloacal aperture. The instant
that the nose of the fish touched the edge of the aperture, which
was rather tightly closetl, the slender tail was brought around with
a very ra[)id whip-like movement, which terminated in a thrust
w^iereby about 5 millimeter^ of the tij) were darted into the narrow
slit between the apposed lip.> of the cloaca. This movement was

effected while the nose of the fish was stiU touching the cloacal

Up to this time the fish had exhibited more or less excitement
but as soon ns the tip of its tail had l)etMi inserted it straightened

host* In thi.r process the fish aj.peared to hv making xrnie use <.f
the spines of the dorsal ami ventral lins. The motion was a slow,
gliding one and was not (h'pendent on the relaxing of the cloacal
sphincter of the holotlnirian.

The lips of the cloacal aperture remained closed during the
ingress except ^o far as they were forced apart by the body of the
fish.

4

THE AMERICAN NATURALIST

[Vol. XLI

The time occupied by the fish in entering the holothurian was
not taken, but in my notes which I made immediately after my
observations, I find that I estimated the time to be probably not
more than half a minute.

In order to give some of the workers in the laboratory, who had
not seen all of the events described above, an opportunity to view
this interesting instance of animal behavior, the experiment was
repeated.

When the finger-bowl had been removed from the holothurian
a second time the fish was found to be again in the water.

Then were repeated in minutest detail the actions which I have
described above. The fish swam actively at the surface of the
water — part of the time with its mouth above the surface. Then it
began to move in a more or less aimless fashion. Finally it touched
the holothurian with its nose, this time about the middle of the
length. Then followed in exact order the reactions which liad
been observed before, viz., the rapid bumping of its nose against
the side of the holothurian, the undeviating progress towards the
posterior end, the whip-like motion and insertion of the tail while
the nose was still in contact with tiie margin of the cloacal aperture,
the leisurely straightening of the body, and the gradual retreat into
the resisting, at least not assisting, holothurian.

It should be added that tiie above account is made up from
notes which I wrote rlown immediately after the observations were
made. Except in one of two details they are confirmed by Dr.
Ulric Dahlgren who has kindly placed his notes at my disposal.

The above accomu is the story of the way one Fierasfer gained
entrance to its host. Whether every individual Fierasfer would
behave exactly tiie same, under similar circumstances, perhaps

I am indebted to Professor Cornelia M. (lapj) for reference to
an excellent article on the habits, anatomy, etc., of Fierasfer by
Dr. Carlo Emery, i Fauna uml Flora ,1rs c'nljrs mn Xraprl, vol. 1,
ISSO). J)r. Kmerv note> that Fi(M-asfer enters it> host tail first.

A somewhat similar habit i> indi<-at..l for the counnon eel l.v
xvhat the veteran and accurate obs<M-ver, \ inal X. Mdwards, tells
me, r/^., that eels go into holes tail first.

RECORDS OF PENNSYLVANIA FISHES

HENRY W. FOWLER

Two complete accounts of the fishes of Pennsylvania have
appeared, one by Cope in 1881, and the other by Dr. Bean in 1892.
The first of these is of a rather general nature, though based on the
author's studies while collecting about the State, and the other is
to a large extent little more than an elaborated compilation of the
first. In view of the desideratum of (lefim't(> rccor.l^ for the dis-
tribution of the different species in the various stn^ains lakes, etc.,
I have collected at a number of localities, and tlius am able to
offer nearly complete notes for some districts. This, and an
examination of the fine collection of Pennsylvania fishes in the
Academy of Natural Sciences of Philadelphia will complete my
records. It may be understood that no species is here included
unless known to me from the examination of a Pcniisx Ivaiiia
specimen. Further exploration will undoubtedly cxkmkI tlic dis-
tribution of many species, besides resulting in the a( (iui^ition of
some others not now known from within our limits.

The collections in the Academy embrace chiffly those made by
E. D. Cope and J. H. Slack, together with others made by T. 1).
Keim, B. W. (iriftiths, P. Lorrilliere, W. S. Sut. h, S. 1'. ( Lindsay,
D. McCadden, II. T. Wolff, Alfre.l Sattertliuait, J. S. Witmer,
W. Stone, and myself. In this conneetiou the writer wishes to
thank Mr. Wm. L. Meehan of the State Fish Commission for
permission to collect fishes in Pennsylvania with nets.

Petromyzon marinus Limueus. Lami'ukv. - Delaware River,
in the Brandywine tributaries in the l)elawan> l.aMu at Stock
Grange, Chester (\k (W. Stone); at 1 lolme^l.ur*;. Philadelphia
Co., Cornwells, Bucks Co., Dingman^ F.-rry, Pik.- Co. (H. T.
Wolff); apparently less fre.|U<-iit in the D.^laware than formerly.

Ichthyomyzon concolor < Kirtlatid >. Siia kix Lami-hkv. — .\lle-
gheny River at Port Allegany, McKean Co.

5

6

THE AMERICAN NATURALIST

[Vol. XLI

Lampetra wilderi Jordan and Evermann. Brook Lamprey.
— Kiskiminitas River (E. D. Cope); Allegheny River at Port
Allegany, McKean Co.; Susquehanna tributaries at Emporium,
■Cameron Co.

ACIPENSERID^

Acipenser sturio Linnaeus. Sturgeon. — Delaware River at
Holmesburg, Tacony, Bridesburg, and Torresdale, in Philadelphia
Co., and Cornwells, Bristol, and Tullytown in Bucks Co. I have
found it fairly abundant at times and seen some large examples.

POLYODONTID/E

Polyodon spathula (Walbaum). Paddle Fish. — Reported as
occurring occasionally in the Allegheny River below Corydon,
Warren Co., which it has been known to ascend as far as Sala-
manca and Olean in Cattaraugus Co., N. Y.

PSALLISOSTOMATID.E

Psallisostomus osseus (Linnaeus). Gar Pike.— Small exam-
ples occur in the lower Delaware River as far as Morrisville,
Bucks Co., and occasionally a large one is noted.

Amia calva Linnfeus. Bow Fix.— An example from Lake
Erie, obtained by Dr. Watson, may have been taken within our

GLOSSODONTID.E

Glossodon harengoides Rafinesque. Moon Eye. — Beaver River
(E. D. Cope); Youghiogheny River (E. D. Cope).

Clupeid.e

Pomolobus pseudoharengus (Wilson). Ale wife.— Delaware
River at Tinicum, Delaware Co., Holmesburg, Tacony, and
Torresdale, Philadelphia Co., and Cornwells, Croydon, Bristol,
Tullytown, and Morrisville, Bucks Co. Abundant in the spring.

No. 481]

PENNSYLVANIA FISHES

7

Alosa sapidissima (Wilson). Shad.— Delaware River at Tini-
cum, Delaware Co., League Island, Frankford, Tacony, Holmes-
burg, and Torresdale, Philadelphia Co., Cornwells, Eddington,
Croydon, Bristol, Tullytown, and Morrisville, Bucks Co., and
Dingmans Ferry, Pike Co. (H. T. Wolff). Some seasons - more
abundant than others. In the Susquehanna they are taken about
Peach Bottom and McCall's Ferry, Lancaster Co.

DOROSOMATID^

Dorosoma cepedianum (Le Sueur). Mjjd Shad. — Delaware
River at Torresdale, Philadelphia Co. (Dr. H. D. Senior), and
Cornwells, Bristol, and Tullytown, Bucks Co. Apparently less
abundant than formerly, though even in midwinter a few have
been hauled out of the mud.

Engraulidid.e

Anchovia mitchilli (Valenciennes). Anchovy.— ]\Iany exam-
ples were obtained many years ago below Philadelphia, some
most likely within our limits, by Dr. Colin Arrott. They occur
in the lower Delaware and are more a feature of the marine fauna.

Salmoxid.e

Salvelinus fontinalis (MitchiU). Brook Trout.— Brandywine
tributaries in Chester Co., and those of the Schuylkill near Port
Kennedy, Montgomery Co. (D. McCadden), all in the Delaware
basin; the Susquehanna basin in the Loyalsock near Lopez,
Sullivan Co., and near Galeton, Potter Co.; the Allegheny basin
above Port Allegheny in McKean Co., and Seven Bridges in
Potter Co.; the Genesee basin about Gold in Potter Co.

Argextinid.e

Osmerus mordax (^Nlitchill). Smelt. — I have seen a number
of examples taken from the Schuylkill and Delaware near Phila-
delphia.

THE AMERICAN NATURALIST

[Vol. XLl

ANGUILLID.E

Anguilla chrisypa Rafinesque. Eel. — Delaware River at Tini-
cum and Darby Creek, Delaware Co., League Island, Frankford,
Tacony, Holmesburg, Torresdale, with the Pennypack Creek basin
at Bustleton, in Philadelphia Co., Corn wells, Croydon, Bristol,
Tullytown, and ^Vlorrisville, with the Neshaminy Creek basin at
Hulmeville and Newtown, Bucks Co., and Dingmans Ferry, Pike
Co. (H. T. Wolff); Susquehanna basin at Galeton, Potter Co. I
did not meet with eels over the Allegheny divide in Potter and
McKean Counties, and the impression is prevalent that they do
not occur in the upper tributaries of either the Allegheny or the

Cyprinid.e

Campostoma anomalum (Rafinesque). Stone Roller. — Alle-
gheny River at Port Allegany, McKean Co., in June, 1906,
where I found entirely tuberculated males; Kiskiminitas River
(E. D. Cope).

Chrosomus erythrogaster eos (Cope). Red-bellied Dace. —
Cotypes of C. eos Cope examined.

Hybognathus nuchalis Agassiz. Silvery Minxow.— Kiskimin-
itas River (E. D. Cope).

Hybognathus nuchalis regius (( Jiraiil i. ivvsn.iix Sii x khv Min-
now.— Delaware River at Tacony, lloliiu'shtuu-. aii<l Torresdale,
Philadelphia Co., and Cornwells and Bri.st<,l, al>o the Xesluuniny
Creek basin at Hulmeville and Neshaminy Falls, in Bucks Co.
Many exanipl(>> were examined.

Pimephales notatus (Rafinesque). Bu xi -noskd Mixxow. —
Kiskiminitas River (E. D. Cope) and the Alleirheny River at Port
Allegany, McKean Co., June, lOOfi.

Semotilus bullaris (Rafinesque). Vw.i. Kisn. - 1 )e!aware River
basin in the Brandywine Creek basin at Kt nnett S(juare, Menden-
hall, Wilhstown Barrens, and opposite Chadds Ford, Chester
Co.; Darby Creek basin, Ddawan" (O.; Tacony, the Penny-
pack Creek basin at Holmesbiu-g and Bustleton, the Poquessing
Creek basin opposite Cornwells and Torresdale, Philadelphia

No. 481]

rj-.WSVLVAMA FISHES

9

Co.; the Neshaminy Creek basin at Croydon, I Iii1iih'\ illc. Xrslia-
miny Falls, and Newtown, Mill Creek a'oout Hii>iol. aixl

Tullytown Creek basin about Tullytowii. Uuck. Co.: Ddawaiv
Water Gap, Monroe Co. (E. D. Copej; DinfjinaiH Vvrr\\ l'ik<'
Co. (H. T. Wolff). In the Susquehanna basin I liavr it from tlu-
Northeast Creek at Nottingham, Chester Co., and tin- Conc-
stoga Creek, Lancaster Co. (E. D. Cope).

Semotilus atromaculatus (Mitchill). (^m li. - 1 )(l;i\van> Kivcr
basin in the Brandywine Creek basin at KcinuMi S(]iiarc, Mcndcii-
hall, and opposite Chadds Ford, Chester Co.; ( "ol)l)'> i 'vvck ( K.
D. Cope) and Darby Creek, Delaware Co.; Tacony Creek, the
Pennypack Creek at Holmesburg and Bustleton, the Poquessing
Creek at Torresdale and opposite Cornwells, Philadelphia Co.;
the Neshaminy Creek at Croydon, Hulmeville, Neshaminy Falls,
and Newtown, ^lill Creek at Bristol, Tullytown Cvvvk at 'rull.v-
town, and Morrisville, Bucks Co.; Dingmans Ferrv. Pike Cc
(H. T. Wolff). The Susquehanna basin in the Octoran. near
Nottingham, Chester Co., and Muncv, Lycominu C... K(av<r
River (E. D. Cope); Warren Co. (Dr. J. H. Slack : Ki-^kiminitas
River (E. D. Cope); the Allegheny \{Wvv at \Uyuunu\^, Putter
Co. I have also found it in the headwaters of the ( iciu M f aioiiud
Gold, Potter Co.

Leuciscus vandoisulus Valenciennes. Posv 1 ) a* i . ( )( torato
Creek, in the Su.squehanna ba.sin, near Xottiiiuhain. Clic-itT Co.

Leuciscus elongatus (Kirtland). Red-sidkd D a( k. - Allc-heny
River at Port Allegany, McKean Co., in June. V.m.

Brama crysoleucas (Mitchill). Roach.— 1 )ehi\van' l>a>iii in the
Brandywine Creek l)asin at Kennett S.,uare, Mendcnliall. aixl
opposite Chadds Ford. Chester (\).\ Tltiicum and Darby Creek.
Delaware Co.; Tacony (/reek, tlie Pennypack at Ilohru-ln.rg. and
Bustleton, and the Piuiiiessing at Torre>da!e and opposite Corn-
wells, Phihidelphia Co.: Xe^liaininy Creek at Croydon, llttlme-
ville, Neshaiuinv Falls, and Newtown. Mill Creek at Bristol,
Tullytown Creek at Tullytown, and M..rriMille. Bucks Co.;
Pennypack at Hatboro, Montgomery < o. ; Daltn ille. Lackawanna
Co.; Dingmans Ferry, Pike Co. U. T. Woltl'i, and in the SuMjue-
hanna basin from the Loyalsock ( feek near Lojhv, >u!livaii ( o.

Notropis bifrenatus (Cope). Bridled Minnow.— Delaware

10

THE AMERICAN NATURALIST [Vol. XLI

River in the Brandywine at Chadds Ford, Delaware Co.; the
Pennypack at Holmesburg and Bustleton in Philadelphia Co.,
and Hatboro in Montgomery Co.; the Poquessing at Cornwells,
the Neshaminy at Croydon, Hulmeville, Neshaminy Falls, and
Newtown, and Mill Creek at Bristol, Bucks Co. Abundant and
rather local.

Notropis cayuga Meek. Cayuga Minnow. — I have already
recorded the only example seen, which was taken near Port Alle-
gany in 1904.

Notropis procne (Cope). Swallow Minnow. — Cotypes of
Hybognathus procne Cope examined. I have examples also from
the Delaware basin taken in the Schuylkill (E. D. Cope) and the
Pennypack Creek near Holmesburg, Philadelphia. A small
minnow was taken on one occasion, which may be this species,
in the headwaters of Northeast Creek, near Nottingham, Chester
Co., in the Susquehanna basin.

Notropis hudsonius amarus (Girard). Eastern Gudgeon.—
Susquehanna basin at Paradise, Lancaster Co. (J. S. Witmer).
Delaware basin in the Brandy wine Creek at Chadds Ford, Dela-
ware Co. ; open river and Pennypack Creek at Holmesburg, and
the Poquessing at Torresdale, Philadelphia Co.; Neshaminy
Creek at Croydon, Hulmeville, and Neshaminy Falls, and Mill
Creek at Bristol, Bucks Co. More abundant in the larger streams.

Notropis whipplii (Girard). Silver Fin.— Youghiogheny River
(E. D. Cope) and Kiskiminitas River (E. D. Cope).

Notropis whipplii analostanus (Girard). Eastern Silver Fin.
— Delaware River basin in the Brandywine Creek at Chadds
Ford, and Darby Creek, Delaware Co.; Wissahickon Creek near
Barren Hill, and Pennypack Creek near Hatboro, Montgomery
Co.; Tacony Creek, Pennypack at Holmesburg and Bustleton,
and Poquessing Creek at Torresdale and opposite Cornwells,
Philadelphia Co.; the Neshaminy Creek at Croydon, Hulmeville,
Neshaminy Falls, Frog Hollow, and Newtown, Mill Creek at
Bristol, Tullytown Creek at Tullytown and Morrisville, Bucks
Co.; Dingmans Ferry, Pike Co. (H. T. Wolff). It is also abun-
dant in the Susquehanna basin where I received it from Paradise,
Lancaster Co. (J. S. Witmer), and the Northeast Creek at Notting-
ham, Chester Co.

Ko. 481]

PENNSYLVANIA FISHES

Notropis comutus (Mitchill). Red Fin.— Delaware llWw l.asin
in the Brandywine Creek at Kennett Square, M. n.lciiliali. aii.l
opposite Chadds Ford, Chester Co.; Darby Creek and Ridley
Creek near WiUistown Barrens, Delaware Co.; AVissahickon Creek
near Barren Hill, and the Pennypack at Hatboro, Montgomery
Co.; Tacony Creek, Pennypack at Holmesburg and Bustleton,
Poquessing Creek at Torresdale and opposite Cornwells, Phila-
delphia Co.; Neshaminy Creek at Croydon, Huliueville. Ne>lia-
miny Falls, Newtown, Frog Hollow, and Chalfont, Mill ( re< k ar
Bristol, Tullytown Creek at TuUytown, and Morrisx ille. Bucks
Co. In the Susquehanna basin in Northeast Creek at Notting-
ham, Chester Co.; Conestoga Creek (E. D. Cope) and Paradise,
Lancaster Co. (J. S. Witmer), and Meshoppen, Elk Co. (E. D.
Cope). In the Allegheny from near Croydon, Warren Co.. and
it also occurs farther up. Kiskiminitas River (E. I). Cope ).

Notropis chalybaeus (Cope). Iron-colored Mi.\.\()\\ . — Ne>!ia-
miny Creek near Newton, and Mill Creek, Bristol, l)otli in the
Delaware basin, Bucks Co.

Notropis atherinoides Rafinesque. Emei{ali) Minnow.— Beaver
River (E. D. Cope).

Notropis rubrifrons (Cope). Rosy-pac ko Minnow. — Cotypes
of Alhurnus rvbrijrons Cope examined.

Notropis photogenis (Cope). Wuite-kvku Minnow . — Cotypes
of Squalius 'photogenis Cope examined.

Notropis photogenis amcenus (Abbott). Attrac ii\ k Minnow.
— Hulmeville, in the Neshaminy Creek, Bucks Co.

Ericymba buccata Cope. Silver-mouthed Minnow.— Cotypes
of the species examined.

Rhinichthys cataract® (Valenciennes). Long-nosed Dace. —
Delaware River basin in the Brandywine tributaries near Kennett
Square and Mendenhall, Chester Co. In the Susquehanna at
Paradise, Lanca.ster Co. (J. S. Witiiierl. Beaver River (E. D.

Rhinichthys atronasus (Mitchill i. Bi .m k-nosed Dace.— Del-
aware River basin in the Brandywine Creek basin at Kennett
Square, Mendenhall, opposite Chadds Ford, Chester Co.; Darby,
Ridley, and Cobb's Creeks, Delaware Co.; Schuylkill River,
Taoony Creek, Pennypack Creek at Holmesburg, and Bustleton,

12

THE AMERICAN NATURALIST [Vol. XLI

and Poquessing Creek at Torresdale, and opposite Cornwells,
Philadelphia Co. ; Wisahickon Creek near Barren Hill and Penny-
pack at Hatboro, Montgomery Co.; Neshaminy Creek at Croydon,
Hulmeville, Neshaminy Falls, Frog Hollow, Chalfont, and New-
town, Mill Creek at Bristol, Tullytown Creek at Tullytown,
Morrisville, Bucks Co.; Dingmans Ferry, Pike Co. (H. T. Wolff);
Daleville, Lackawanna Co. In the Susquehanna basin it occurs
in the Octoraro and Northeast Creeks near Nottingham, Chester
Co.; Paradise in Lancaster Co. (J. S. Witmer); the Loyalsock
Creek near Lopez, Sullivan Co., and Pine Creek at Galeton and
above, Potter Co. In the Allegheny River it is abundant at
Coudersport and Raymonds, Potter Co. Kiskiminitas River (E.
D. Cope). Youghiogheny River (E. D. Cope). Genesee basin
near Gold, Potter Co. Potomac drainage of Fulton Co., (W.

Hybopsis kentucMensis (Rafinesque). Horxed Chub. — Sus-
quehanna basin in Elk Creek, Chester Co. (E. D. Cope); Conestoga
Creek, Lancaster Co. (E. D. Cope). Beaver River (E. D. Cope);
Youghiogheny River (E. D. Cope); Kiskiminitas River (E. D.
Cope); Warren Co. (Dr. J. H. Slack); Allegheny River at Port
Allegany, McKean Co.

Exoglossum maxillingua (Le Sueur). CrT-urs Mi wow . - Alle-
gheny River at Port Allegany, McKean Co.

Carpiodes cypiinus (Le Sueur). Eastern Carp Sucker. —
Conestoga Creek, in the Susquehanna basin, Lancaster Co. (E.
D. Cope).

Cycleptus elongatus (Le Sueur). Black Horse.— Kiskimini-
tas River (E. D. Cope).

Catostomus commersonnii (Lacepede). Common Sk keh - Del-
aware River in the Brandywine tributaries at Keruiett S(,uare,
Mendcnhall and opposite Clia.lds Ford, Chester Co.; Tinicum,
Ridlfv, (ol.l.'s and l)arl)v ( ir.-ks, 1 Maware Co. ; Pennypack
Creek af llatboro, Montgoin<'ry Co.; lacony Creek, Pennypack
Creek at Holmesburg and Bustleton, Poquessing Creek at Torres-
dale and opposite Cornwells, Philadelphia Co. ; Neshaminy Creek

No. 481]

PENNSYLVA NIA FISHES

13

at Croydon, Hulmeville, dialtnnt. Fn.<r Hollow, NcslKiminy Falls,
and Newtown, Mill Creek at Uristol, 'rullvtoun i'vwl at 'rully-
town, and Morrisville, Hiuks (\,.; Drlauarr Water Cap, Monio,-
Co. (E. D. Cope); Dinpnans Ferry, Fik.- C... ill- T- Wolll).
In the Susquehanna basin in tlu- Northeast ami Ocioraro ( rtvk^
near Nottingham, Chester Co.; Conesto<^a Creek K. D. Cope)
and Paradise, Faneaster Co. -.1. S. WitnuM -. In th.> .Mlej^^heny
River it occurs al Fort Alle-any in McKean Co.

Catostomus nigricans Fe Siu-ur. k St ( ki.k. In the

Susquehanna basin in the Cone.stoj^a Creek ul. Stanll'er) and at
Paradise, Lancaster Co. (J. S. Witiner); Octoraro Creek near
Nottingham, Chester Co. Kiskiminitas River (E. L). Cope). I
have not positively identified this from the Delaware basin.

Erimyzon sucetta oblongus (Mitchill). Mt i.i.i-.T. — Delaware
River basin in the Brandyvvine tributaries at Keimeit S.piare and
Mendenhall, Chester Co.; Brandvwine at ChM> l ord. Kidley
and Darby Creeks, Delaware Co.; Tacony Creek, ri\er an<l W uuy
pack Creek at Ilolmesburg and Bustleton, Po(jue>,sin^M "reek at
Torresdale, and opposite Cornwells, Philadelphia ( o.; .Xe.sh-
aminy Oeek at Croydon, Huhneville, Neshainiiiy Fall^, an.l New-
town, Mill Creek at Bristol, Tnllytown Creek at Tiilhtown, and
Morrisville, liucks Co. In ilir SnM,nelianna ha^i.i from Center
Co. (Dr. IF Allport). (Jenesee liiver at (...Id. Poller Co.

Moxostoma anisurum (Rafiiie>(pie). \\ ni i i.-No>i.i> .^i < kkw.
Heaver River ' F. D. Cope) and Youghiogheny Fixer F. 1). Cope).

Moxostoma aureolum ( Fe Stieur). (.oi.dkn Kkd House.
Beaver River (F. 1). Cope) and Voughiogheny River ; F. D. Cope).

Moxostoma macrolepidotum (Fe Sueur). Rkd Horse.— Cone-
stoga Creek, Lancaster Co. (E. D. Cope).

Moxostoma breviceps (Cope). Loxg-tailed Red House. -
Type of Pti/cJiosfomus hrrviccps (\)pe examined.

Placopharynx duquesnii ( Fe Sueur). Bi(;-.ta\\i.i) St ckkh. —
Beaver River (E. D. Cope).

Ictalurus punctatus Ra

Blue Cat.— Beav

(E. D. Cope).

IS (linnieus). White Cat. — Delaware Rive

14

THE AMERICAN NATURALIST

[Vol. XLI

basin in tributaries in Chester Co. (V. Bernard); Holmesburg
and Torresdale, Philadelphia Co.; Bristol, Bucks Co.; Susque-
hanna River (E. D. Cope).

Ameiurus nebulosus (Le Sueur). Yellow Cat. — Delaware
River basin in the Brandjwine tributaries at Kennett Square and
Mendenhall, Chester Co.; Tinicum, Brandywine at Chadds
Ford, Darby, Ridley, and Cobb's Creeks, Delaware Co.; Jenkin-
town, Montgomery Co. (H. Crawley); Falls of Schuylkill (Dr.
Uhler), Tacony Creek, river and Pennypack at Holmesburg and
Bustleton, Poquessing Creek at Torresdale, and opposite Corn-
wells, Philadelphia Co.; Neshaminy Creek at Croydon, Hulme-
ville, Neshaminy Falls, Chalfont, and Newtown, Mill Creek at
Bristol, Tullytown Creek at Tullytown, and Morrisville, Bucks
Co.; Dingmans Ferry, Pike Co. (H. T. Wolff); Susquehanna
River (E. D. Cope); Northeast Creek near Nottingham, Chester
Co.; Conestoga Creek, Lancaster Co. (E. D. Cope); Allegheny
River at Coudersport and Perry ville. Potter Co.

Gronias nigrilabris Cope. Blind Cat. — Cotypes of the species
examined.

Leptops olivaris (Rafinesque). Mud Cat. — Youghiogheny
River (E. D. Cope).

Schilbeodes gyrinus (Mitchill). Tadpole Stone Cat. — Dela-
ware River at Holmesburg, Philadelphia Co.; Mill Creek at
Bristol, and Tullytown, Bucks Co.; Delaware Water Gap, Monroe
Co. (E. D. Cope); Dingmans Ferry, Pike Co. (H. T. Wolff). In
the Susquehanna from the Loyalsock Creek near I^pez, Sullivan
Co. In the Genesee below Gold, Potter Co.

Schilbeodes insignia (Richardson). Margined Stone Cat. —
Delaware River in the Schuylkill and at Holmesburg, Philadel-
phia Co. ; Susquehanna basin at CarHsle, Cumberland Co. (S. F.
Baird); Conestoga Creek (J. Stauffer) and Paradise, Lancaster
Co. (J. S. Witmer).

ESOCID.E

Esox americanus (Gmelin). Banded Pickerel. — Delaware
River basin in the Brandywine tril>utaries at Kennett Square and
Mendenhall, Chester Co.; Tinicum and Darby Creek, Delaware
Co.; Tacony Creek, Pennypack Creek at Holmesburg, and Po-
quessing Creek at Torresdale and opposite Cornwells, Philadel-

No. 481]

PENNSYLVANIA FISHES

15

phia Co.; Neshaminy Creek at Neshaminy Falls and Xtnvtowii.
Mill Creek at Bristol, Tullytown, and Morrisville, \hu-k. ( o. In
the Octoraro Creek of the Susquehanna basin near Xottiii^liaiii,
Chester Co.

Esox vermiculatus Le Sueur. Wisi Pk kkkki.. - Allegheny
River in Potter County.

Esox reticulatus Le Sueur. Cirviv PiKi:.^Rock Hill Pond
and Dingmans Ferry, in the Delaware Basin, Pike Co. (H. T.
Wolff).

Esox lucius Linngeus. Pike. — Although I did not secure any
examples of this species in the Allegheny River while at Port
Allegany, in ]\IcKean Co., in 1904, it was reported as occurring
farther down stream. One from Lake Erie may have been from
within our Hmits (Dr. Watson).

Esox masquinongy ohiensis (Kirtland). Ohio River Musk-
ALLUNGE. — Reported to occur in the Allegheny as far as Cor}don,
Warren Co., and in New York to Olean. Warren Co. (Dr. J. H.
Slack).

Umbrid.e

Umbra limi pygmsea (De Kay). Mfd ^NIixxow.— Delaware
River at Philadelphia and Ilohn'ol.ur-, Phila.l.'lphia Co.; Bristol
and Tullytown, Bucks Co.; Sclmylkill Kivcr Dr. llarlanK

Fundulus heteroclitus macrolepidotus ^ W albauiii . Mi mmichog.
— Delaware River in tide-water, at Tiiiicuni, Delaware Co.;
League Island, Tacony, Holmesburg, and Torresdale, Philadel-
phia Co.; Cornwells, Croydon, Bristol, Tullytown, and Morrisville,
Bucks Co.

Fundulus diaphanus (Le Sueur). liAHHEi) Kii.i.ii- ish. — Dela-
ware River in tide-water and above, Hrandywiiic ba^in in ( 'lic^tt r
Co. at Kennett Square; Brandywine at ('iia.liis FonI, Darbv and
Ridley Creeks, Delaware Co.; Tacony Creek, Peiiiivpack ( "r.-ck
at Holmesburg and Bustleton, Poquessing Creek at Torro.ialr
and opposite Cornwells, Philadelphia Co.; Neshatniiiy ("nek at
Croydon, Hulmeville, Neshaminy Falls, and Newtown, ^Nlill

16

THE AMERICAN NATURALIST [Vol. XLI

Creek at Bristol, Tullytown Creek at Tullytown, and Morrisville,
Bucks Co. ; Montgomery Co. ; the Susquehanna basin at Paradise
(J. S. Witmer) and Lancaster Co. (E. D. Cope); Warren County
(Dr. J. H. Slack).

Mastaccembelid.^:

Tylosurus marinus (Walbaum). Green Gar. — Delaware River
(Dr. Uhler), at Bristol, Bucks Co.; Susquehanna River (E. D.
Cope).

Atherinid.k

Labidesthes sicculus (Cope). Brook Silverside. — Youghi-
ogheny River (E. D. Cope).

Gasterosterid^

Eucalia inconstans (Kirtland). Brook Stickleback. — Erie,
Erie Co.

Apeltes quadracus (Mitchill). Four-spined Stickleback. —
Delaware River, in tide-water, at Tinicum, Delaware Co.; Tacony,
Holmesburg, and Torresdale, Philadelphia Co.; Cornwells, Croy-
don, Bristol, Tullytown, and Morrisville, Bucks Co.

Aphredoderid.k

Aphredoderus sayanus ((iilliams). Pirate Perch. — Delaware
River at Tinicum, Delaware Co.; League Island, Tacony, Holmes-
burg, and Torresdale, Philadelphia Co. ; Mill Creek and the river
at Bristol, Bucks Co.

Centrarchid^

Pomoxis annularis Rafinesque. Crappie. — Kiskiminitas River
(E. D. Cope). I have an example from the Delaware at Browns-
ville, Bucks Co. (J. G. Dillin).

Ambloplites rupestris (Rafinesque). Rock Bass. — Beaver River
(E. D. Cope), Warren Co. (Dr. J. H. Slack), and Kiskiminitas
River (E. D. Cope).

No. 481] PENNSYLVANIA FISHES

17

Enneacanthus gloriosus (Holbrook). Blue-spotted Sunfish.
— Delaware River at League Island (Professor Wm. M. (labb),
and Holmesburg, Philadelphia Co.; river and Mill Creek at Bristol,
Bucks Co. In the Susquehanna from the Conestoga Creek, Lan-
caster Co.

Enneacanthus obesus (Girard). Sphagnum Sunfish.— Found
only in the ditches of the lower part of Philadelphia along the
Delaware.

Mesogonistius chsetodon (Baird). Banded Sunfish. — Dela-
ware River at Holmesburg, Philadelphia Co., and Bristol, Bucks
Co.

Lepomis auritus (Linnaeus). Red-breasted Sunfish.— Dela-
ware River basin in the Brandywine tributaries (Dr. H. Allen),
Londongrove (E. D. Cope); Chadds Ford on the Brandywine,
Darby, Ridley, and Cobb's Creeks, Delaware Co.; Montgomery
Co. (W. Cassin), and Pennypack at Hatboro; Tacony Creek,
Pennypack at Holmesburg and Bustleton, Po(iucssiiii: at Torres-
dale and opposite Cornwells, Philadelphia Co.; Xolmniiny ( r<'ek
at Croydon, Hulmeville, Neshaminy Falls, and Newtown. Mill
Creek at Bristol, TuUytown and Morrisville, Bucks Co.; Ding-
mans Ferry, Pike Co. (H. T. Wolff). In the Susquehanna basin
from the Octoraro Creek (E. D. Cope) and Conestoga Creek,
Lancaster Co. (E. D. Cope), and Paradise, Lancaster Co. (J. S.
Witmer).

Lepomis megalotis (Rafinesque). Long-eared Sunfish. —
Kiskiminitas River (E. D. Cope).

Lepomis macrochirus Rafinesque. Large-finxkd Sunfish.
— Cotypes of Lepomotis ncphelus Cope examined.

Lepomis paUadus (Mitchill). Blue-gill Slmi^h.- Warren
Co. (Dr. J. H. Slack); Ki.^kiminitas River ( K. D. ( ope i.

Eupomotis gibbosus ( Liiitiiens ). ("hmmox Sim ish.- Dela-
ware River basin in tribiitario of I^rau-lyu in.- near Kennett
Square, Chester Co. (E. 1). Cope and II. Allen): Hraiidyuiiie at
Chadds Ford, Ridley, Darl)y, and (\)bl)"s Creek, and Tinicum,
Delaware Co.; Jenkintown and Hatboro, Montgomery Co.;
Tacony Creek, Pennypack Creek at Holmesburg and Bustleton,
Poquessing at Torresdale and opposite Cornwells, Philadelphia
<'().; Xeslianiiny Creek at Croydon, Hulmeville, Neshaminy Falls,

18

THE AMERICAN NATURALIST [Vol. XLI

and Newtown, Bristol, and Mill Creek, Tullytown and Tullytown
Creek, and Morrisville, Bucks Co.; Dingmans Ferry, Pike Co.
(H. T. Wolff). In the Susquehanna from the Conestoga Creek,
Lancaster Co. (E. D. Cope); the Loyalsock near Lopez, Sulhvan
Co.; Oetoraro Creek (E. D. Cope). Erie, Erie Co. (C. Rutter).

Micropteras dolomieu Lacepede. Small-mouthed Bass. —
Youghiogheny River (E. D. Cope) and the Allegheny at Corydon,
Warren Co., and met with as far as Olean, N. Y.

Micropteras salmoides (Lacepfede). Large-mouthed Bass. —
Warren Co.

Percid.*:

Stizostedion vitreum salmoneum (Rafinesque). Blue Pike. —
Warren Co. (Dr. J. H. Slack), Beaver River (E. D. Cope), and
Youghiogheny River (E. D. Cope).

Stizostedion canadense griseum (De Kay). Sauger. — Warren
Co. (Dr. J. H. Slack), Beaver River (E. D. Cope), and Youghi-
ogheny River (E. D. Cope).

Perca flavescens (Mitchill). Yellow Perch. — Delaware River
in Darby Creek, Delaware Co.; river and Pennypack Creek at
Holmesburg, and Poquessing Creek at Torresdale, Philadelphia
Co.; Neshaminy Creek at Croydon, Hulmeville, Neshaminy
Falls and Newtown, Mill Creek at Bristol, Tullytown Creek at
Tullytown and Morrisville, Bucks Co. In the Susquehanna basin
from the Conestoga Creek in Lancaster Co. (E. D. Cope).

Percina caprodes (Rafinesque). Log Perch. — Type of Perca
nehulosa Haldeman examined, Youghiogheny River (E. D. Cope)
and Kiskiminitas River.

Hadropteras macrocephalus (Cope). Long-headed Darter.

— Cotype of Etheostoma macrocephalum Cope examined.
Hadropteras peltatus (Cope). Shielded Darter. — Type of

Etheostoma peltatum Stauffer, in Cope, examined.

Diplesion blennioides (Rafinesque). Green-sided Darter. —
Beaver River (E. D. Cope).

Boleosoma nigram (Rafinesque). Johnny Darter. — Cotypes
of B. olmstedi brevipinnis Cope examined.

Boleosoma nigrum olmstedi (Storer), Tessellated Darter.

— Delaware River basin in the Brandywine tributaries at Kennett

No. 481]

PENNSYLVANIA FISHES

19

Square, Mendenhall and opposite Chadds Ford, Chester Co.;
Brandywine at Chadds Ford, Ridley, Darby, and Cobb's Creek,
and Tiniciim, Delaware /7!%s.
Co. ; Pennypack at Hat- // / 'X

boro, and Jenkintown,

Montgomery Co.; League *
Island, Tacony Creek, 4
Frankford Creek, Penny-
pack Creek at Holmes-
burg and Bustleton, and
Poquessing Creek at ^
Torresdale, and opposite
Cornwells, Philadelphia I
Co.; Neshaminy Creek
at Croydon, Newportville,
Hulmeville, Neshaminy
Falls, Frog Hollow, and

Newtown, Mill Creek at - ^

Bristol, TuUytown Creek ' ; vj^

at TuUytown, and Morris- ^ '

ville, Bucks Co.; Diiiu-
mansFerry, Pike Co. (II.
T. Wolff). In the Sus-
quehanna basin I have it
from the Octoraro (Veek
at Nottingham in Chester
Co., the Loyalsock near
Ix>pez, SuUivan Co., and
the Pequea at Paradise,
Lancaster Co. (J. S. Wit-
mer), besides the type of
Percina minima Halde-
man. The accompany-
ing figure represents a tstoVer).
remarkable variation of

fin-rays seen in an example I secured at Holmesburg, Phila-
delphia, ^September 11th, 1898. Although I have not seen the
type of Boleosoma oesopm Cope my Loyalsock examples go far to
establish it as a pure synonym of this fish.

20

THE AMERICAN NATURALIST [Vol. XLI

Etheostoma caeruleum Storer. Blue Darter. — Kiskiminitas
River (E. D. Cope).

Etheostoma flabellare Rafinesque. Fan-tailed Darter. — Kis-
kiminitas River (E. D. Cope); Youghiogheny River (E. D.
Cope); Pittsburgh (Jacob Green); Allegheny River at Port Alle-
gany in McKean Co. during July and August of 1904 and June
of 1906, where it is abundant.

Boleichthys fusiformis erochrous (Cope). Sphagnum Darter.
— Delaware basin in Mill Creek near Bristol, Bucks Co.

Roccus lineatus (Bloch). Striped Bass. — Delaware River
basin at Tinicum, Delaware Co.; League Island, Tacony, Holmes-
burg, and Torresdale, Philadelphia Co. ; Cornwells, Bristol, Tully-
town, and Morrisville, Bucks Co. In the Susquehanna basin I
have it from the Conestoga in Lancaster Co. (E. D. Cope).

Roccus chrysops (Rafinesque). White Bass. — Reported from
just below the headwaters of the Genesee in Potter Co. near Gold.

Morone americana (Gmelin). White Perch. — Delaware River
at League Island, Tacony, Holmesburg, and Torresdale, Phila-
delphia Co., Cornwells, Bristol, Croydon, Tullytown, and Morris-
ville, Bucks Co.

Aplodinotus grunniens Rafinesque. Fresh-water Drum. —
One from Lake Erie (Dr. Watson) may have been taken in our

Cottid-E

Uranidea graciUs viscosa (Haldeman). Miller's Thumb.—
The Delaware basin in the Brandywine tributaries near West
Chester, Chester Co.; in the Schuylkill basin near Port Kennedy,
Montgomer}' Co. (D. McCadden), and Douglassville, Berks Co.
(S. N. Rhoads); in the Neshaminy basin near New Britain, Bucks
Co. (Dr. C. C. Abbott). In the Susquehanna basin in Spruce
Creek, Huntingdon Co. In the Genesee basin at Gold, Potter
Co., also other examples from the same County (E. Harris).

No. 481]

PENNSYLVANIA FISHES

21

SOLEID.E

Achirus fasciatus Lac^pede. Sole. — Schuylkill River in the
Delaware basin (Dr. Harlan) and Bristol, Bucks Co. (Dr. J. De B.
Abbott).

Gadid.e

Lota maculosa (Le Sueur). Ling. — Erie, Erie Co.; Susque-
hanna basin at Muncy, Lycoming Co. (E. D. Cope).

SPECIFIC NAME OF NECTURUS MACULOSUS

Y. C. WAITE

In view of the fact that this animal is now extensively used in
research and teaching, and since the majority of teachers and
writers follow Cope ('89) and erroneously use the name Necturus
mcwulatus, it seems worth while to call attention to the correct ter-
minology.

There has been considerable confusion in the nomenclature of
this form since it was first described. ^Nlinor variations have in
several cases received specific names, and in the earlier literature
it was frequently confused with Cryptobranchm allegheniensis of
which it was for a time considered the larva.

The first scientific description of this animal was by Schneider
in 1799 from a single specimen in the museum at Brunswick. This
specimen came from Lake Champlain. Schneider did not consider
it a new genus but put it under the European genus Salamandra
without appending any specific name.^

Lac^pede ('07) described a museum specimen, saying that it had
never been before described, evidently not knowing of Schneider s
description. He recognized that it differed from Salamandra and
therefore referred it to the genus Proteus, naming it Proteus tetra-
dactyle with the provision that if it were found to be a larva, it
should be called Salamandra tetrad actyle.

Barton ('07, pp. 196-7), describes "a large species of Salaman-
dra" which he proposes to call 5. horrida, or maxima, or gigantea.
It is evident from a reading of his paper that he has confused
Necturus and Cryptobranchus, and the general inaccuracy of his
description makes his contribution of little value.

Rafinesque ('18, p. 40), gave a brief pivliminary .lex-Hption of
this salamander under the name Sirena mnculo.m. He, however,
recognized, as the following quotation shows, that it might repre-
sent a new genus (p. 40) : " In Zoology my discoveries are par-

'The original paper is not available. I quote from Holbrook ('42).

23

24

THE AMERICAN NATURALIST [Vol. XLI

ticularly important consisting of about 25 new undescribed
quadrupeds, 30 new birds and about 32 new reptiles" * * * *
"Among so many undescribed things it must follow that several
may constitute new genera. . . .1 propose to select 8 N. G. and 10
N, Sp. in order to convey an idea of the whole.

(p. 41) I. N. Sp. Sirena maculosa, (A Reptile). Body oliva-
ceous brown, covered with large unequal blackish spots * * *
"This spotted siren bears the generic name Water Puppet along
with S. lutea and S. fusca."

A year later, after he had been able to study his collections,
Rafinesque ('19) erected the new genus Necturus to include this
form. I quote part of his description, (p. 418): "Ille Classe.
Reptiles * * * * 7. Necturus (Batracien) Different des genres
Salamandra, Triturus (Triton, Laur.), Larvarius (Proteus, Aurt.),
par queue comprimde 4 doigts separes a tons les 4 pieds, branchies

exterieures persistent communement jusqu' a la vieillesse

Esp^ces: N. maculatus, N. lutescens, N. fuscus, N. marginatus,
N. axolotes?, N. anguillaris, N. operculatus, etc."

A year later Rafinesque ('20) again describes this form, (p. 4) :

"III Class. Erpetia — the Reptiles 17. Necturus macu-

losus, olive brown covered with large unequal black spots ....
My genus Necturus (70 N. G. An.) is distinguished from Triturus
by having teeth, four toes to all the feet and external gills present

to a late period .... 18. Necturus luteus 19. Necturus phos-

phoreus" * * *.

Mitchill ('21, p. 183) says in regard to this form: "From such
survey as I have been capable of making I am inclined to consider
him a Proteus; but of a species different to that known to European
naturalists." Later, in a very extensive description with good
plate, (Mitchill, '24) he describes a specimen from Lake Erie and
recognizes that it differs from the genus Proteus, but is " averse tO'
unnecessary multipUcation of genera."

In 1823 Say (James, '23, vol. 1, p. 5, footnote) describes speci-
mens from the Allegheny River, with permanent branchiae. He says
that it is caught at Pittsl)urg but is not so abundant as S. alleghe-
niensis. He gives the new name Triton lateralis.

Harlan ('24, p. 233, pi. 16), evidently unaware of the papers of
Rafinesque, erected the new genus Menobranchus. "The Am-

No. 481]

NAME OF NECTURUS

25

phiuma, the Siren, the Proteus and the Salamaudnt will l.c acknowl-
edged by all to constitute separate genera. Tlic Idlinills and
allegheniensis not belonging to any of these will rc(|uirc appropriate
generic names.... As the most prominent feature (iistin::ni>ln"ng
the T. lateralis from the Salamandra is its persistent hranchiu',
we have preferred a name significant of this fact. MeiioKraix lius.
Generic characters. Persistent branchijB, two row^ of teeth in the
upper and one row in the lower jaw; four foott^l, tour io( ^ to each
foot, clawless." He describes two species, .1/. lafira/is and M.
tetradactylus. Since these were found to be but variations of the
same species and since the term tetradactylus describecl a generic
character, the first species, only, held.

Harlan erected the genus Abranchus to include the allcf/hrnirn-
sis, but in a note a few months later (same journal and volume, p.
270) he changed this to Menopoma having learned that the name
Abranchus was preempted for a genus of nudibraiu h iiiollux >.

Barnes ('26) calls it Proteus lateralis and says f]). 2s7) that "the
first specific name given was by Mr. Say who called it latrnili.s in
allusion to the black lateral line. The discoveries already made

show that the character from which he derived this is

variable. " He dissents from Harlan and does not think that a
new genus should be established. He evidently knew nothing
of Rafinesque's description.

In a later note (Barnes, '27, p. 68), he says: "Dr. :Mitchill has
laterly called it Proteus maculatus, which as it is a good descriptive
name. . . .1 am disposed to adopt." Mitchill had evidently gotten
the maculatus from Rafinesque's second paper ('19) but had not
seen the first ('18) or third ('20) papers.

Fitzinger ('26, p. 43) gave a new name to the genus. " Genus.
Phaenerobranchus. Aus T.acepede's Proteus tetradactylus, Say's
Triton lateraHs, aus Nord Amerika, schiif ieh die (laftunir Phae-
nerobranchus (Menobraiichus, Harlan; Necturu-, IlafiiiesipH')."
Under the list of reptiles in the Zoological Mu>eum at \ ienna
he includes (p. 66) "Phaenerobranchus cepedii i Troteus tetra-
<la< tylus. La Cepede)."

Harlan ('27) in his synopsis inclu<les (p. ;>2:{) " Mcnohranchus
lateralis" and this name w^as followed by many writers.

Wagler ('30, p. 210) returns to the generic name of Xecturus,

26

THE AMERICAN NATURALIST [\^ol. XLI

and quotes as synonyms, Proteus tetradactylus Lacepede, Triton
lateralis Say, and Menobranchus lateralis Harlan.

Tschudi ('38, p. 97) adopts Menobranchus lateralis.

Holbrook ('42, vol. 5) describes two species of Menobranchus:
(p. Ill) M. maculat2is (Barnes) in which he makes no reference to
Rafinesque, and (p. 115) M. lateralis (Say). He states that the
two species may be only geographical varieties.

DeKay ('42, p. 87, pi. 18, fig. 45) uses Menobranchus lateralis
and does not refer to Rafinesque.

Baird ('50), in his "Revision of the Tailed-Batrachia " gives cor-
rect references to Rafinesque's three papers and while adopting
his generic name, adheres to Say's specific name (which was given
five years later) giving the name of the form as Necturus lateralis
1823, which was the date of Say's description, although Say did
not use the term Necturus.

Gray ('50) gives the reference to Rafinesque's three papers and
is the first to adopt the correct name Necturus maculosus. How-
ever, in the second edition of this work (Boulenger, '82, p. 84)
reference is made only to Rafinesque's second ('19) paper and the
term Necturus maculatus is taken.

Dumeril and Bibron ('54, p. 183) use Harlan's term Meno-
branchus lateralis.

Finally, Cope ('89, p. 23) adopts the name Necturus maculatus
in spite of the fact that his references to synonyms shows that he
had consulted all of Rafinesque's papers.

The following are part of the Laws of priority as published in
the International Rules of Nomenclature (:05): —

(p. 35) ''Art 25. The valid name of a genus or species can be
only that under which it was first designated on the condition : (a)
That this name was published and accompanied by an indication
or a definition or a description and (b) That the author applied
the principles of binary nomenclature."

(p. 36) " Art. 28. A genus formed by the union of two or more
genera or subgenera takes the oldest valid generic or subgeneric
name of its components. The same rule obtains when two or
more species or subspecies are united to form a single species or
subspecies.

(p. 37) "Art. 32. A generic or a specific name once pubhshed

No. 481]

NAME OF NECTURUS

27

cannot be rejected even bv its author because of inappropriate-

Although Schneider gave the first description of this form he
did not follow the binary system nor did he recognize it as a sepa-

There can be no question that Rafinesque ('19) was the first
to erect and name a new genus to receive this form.

The specific name tetradactylus given by Lac^pede ('07) cannot
hold because it describes a generic character ("4 doigts s^par^s h.
tous les 4 pieds") in the new genus Necturus of Rafinesque.

The description by Barton ('07), is certainly so inaccurate,
including his hesitation between three specific names, that his
paper can have little weight.

The first scientific description with the use of a binary nomen-
clature is that of Rafinesque ('18). Here the specific name is
maculosa. According to Art. 32 of the rules on priority this term
could not have been changed by Rafinesque if he had wished to do
so. The term maculatus used in his 1819 paper is either an unin-
tentional slip on his part or a typographical error. At any rate the
use of the term maculosus in his '20 paper, which was published
under his immediate direction with opportunity to correct proof,
shows that he preferred the original adjective form maculosus to
the participial form maculatus. Such is certainly the better gram-
matical usage.

The confusion has arisen from the fact that the first ('18) paper
was printed in a rather obscure literary periodical where scientific
men were unlikely to see it. Likewise the third paper ('20) was
published in an obscure private publication, a serial which did not
continue and so was easily lost sight of. The second paper ('19) in
which the error occurred was in a prominent scientific journal and
thus came to be generally known.

In the past ten years although many papers have been written
on Necturus, two only have, as far as I know, used the correct
nomenclature. These are Eycleshymer (:06) and Waite ('97).

I believe that it is clear from the foreoging that the correct name
is Necturus maculosus and I hope that this may come into general

28

THE AMERICAN NATURALIST

[Vol. XLI

LITERATURE.

'06. International Rules of Nomenclature, adopted by the Inter-
national Congress of Zoology, Paris, 1905.
Baird, S. F.

'50. Revision of the North American Tailed-Batrachia with Descrip-
tions of a New Genus and Species. Journ. Acad. Nat. Sci.^
Philadelphia, ser. 2, vol. I, pp. 281-294.
Barnes, D. H.

'26. An Arrangement of the Genera of the Batracian Animals with a
Description of the more Remarkable Species including a mono-
graph of the Doubtful Reptiles. Amer. Journ. Sci. and Arts,
ser. 1, vol. 11, pp. 268-297.
Barnes, D. H.

'27. Note on the Doubtful Reptils. Amer. Journ. Sci. and Arts, ser. 1,
vol. 13, pp. 66-70.

'07. [Scientific Notes]. Phila. Med. and Physical Journ., suppl. 2,
sect. 2.

BOULENGER, G. A.

'82. Catalogue of the Batrachia Gradientia, s. Caudata and Batrachia
Apoda in the Collection of the British Museum. London, 1882.
(Second edition of Gray, '50).
Cope, E. D.

'89. The Batrachia of North America. Bull. 34, U. S. Nat. Mus.,
Washington.
DeKay, J. E.

'42. Zoology of New York. Part III. Reptiles and Amphibia.
Dum^:ril, a. M. C. et Bibron, G.

'54. Erpetologie generale ou histoire naturelle complete des Reptiles.
Tome 9, Paris.
Evcleshymer, a. C.

'06. The Habits of Nrcturus maculosus. Amer. Nat., vol. 40, pp.

Fitzin(;kk, L. I.

'26. Neuc Classification dcr Reptilion. Wien.
Gray, J. E.

'50. Catalogue of the Specimens of Amphibia in the Collection of the
British Museum. Part II. Batrachia Gradientia etc. London.

No. 481]

NAME OF NECTURUS

29

Harlan, R.

'24, Observations on the Genus Salamaiidra with tlic Anatomy <.f
the Salamandra gigantea (Barton) or S. (lUajlu himsis (.Micliaux)

pt. 2, pp. 222-234.

HoLnuooK. J. i;.

in 1819 and 1820 under the coniniaiHl of Major S. H. Long. 2
vols., Philadelphia, and 3 vols., London.

LAcf;pfcDE, M. DE.

'07. Sur une espece de quadniprdc ovipaiv non .■ncoic drcriic Ann.
Mus. Hist. Nat. Pans. vol. 10. _>;!() l';!.",, ,,]. 17.

MiTCHILL, S. L.

'21. The Proteu.s of the North Atnrricni Lalo-. .1///,/-. .Inum. S.i.
and Arts, svr. 1. vol. 1. p,.. IM Is:;.

MiTCHILL, S. L.

'24. Observations on Srvcral Itrpiilrs of \,,rtl. Ainrrica vvln.-li MTin
to belong to the fannlv I'n.iMi.^. Annr. J.nn-n. Sr,. an,} Arts,

R.\FINESQUE, C. S.

'18. Further Accounts of I )isrov(.ri(- in Xatnral History in thr WV^t-

R.\FINESQrK. C.

I'interieur drs J-tats I'nis d' Ainrri.iUi' durant I'annee 1818.
Journ. Phiisuiur. Chnuu. K Hist. Nat., vol. 88, pp. 417-429,
June 1819."
Rafinesque, C. S.

30

THE AMERICAN NATURALIST

[Vol. XLI

Wagler, Joh.

'30. Naturliches System der Amphibien mit vorgehenden Classifica-
tion der Saugethiere und Vogel. Munchen, Stuttgart und
Tubingen.
Waite, F. C.

'97. Variations in the Brachial and Lumbo-sacral Plex(uses) of
Necturus maculosus Rafinesque. Bull. Mus. Comp. ZooL, vol. 31,
pp. 69-92.

VOLVOX FOR LABORATORY USE

BERTRAM G. SMITH

In providing a supply of Volvox for class use late in the fall,
difficulty has been experienced in two respects: in keeping the
material alive in the laboratory as long as desired, and in getting
specimens containing sperm and ova. Inquiry reveals the fact
that others have had the same trouble. I have recently been able
to overcome both these difficulties, and at the suggestion of Dr.
H. H. Newman have recorded the method in some brief notes.

Species. — So far as known, the only species of Volvox that has
been found in the vicinity of Ann Arbor during the late autunm is
Volvox aureus Ehrenb. It is not very abundant, and 1 have never
found it in the sexual stage at the time it was collected. It (occurs
in small glacial pools containing Riccia and duckweed.

During the early spring Volvox glohatar Linri., and no other
species, occurs in great abundance in the same pools that later con-
tain Volvox aureus. I have occasionally found it in the sexual
stage when collected.

By the latter part of June Volvox glohator has become quite
scarce, and V. aureus has begun to appear. During the early
part of July the two species exist in the same habitats, but neither
is very abundant.

Since Volvox is so widely used for laboratory work by beginning
classes, the marked specific differences are matters of importance.
Volvox glohator is the form described in text-books bur the de-
scription is far from being applicable to V. aun iis. Since the latter
may at times be the only species available, it may be profitable to
call attention to the differences between the two species, for it is to
be suspected that Volvox aureus is sometimes used without its
species being recognized.

Volvox glohator is considerably larger than F. aiinnis: its somatic
cells are more numerous and compactly arraiiired. The somatic
cells of glohator are angular and connected l»y \erv stout proto-
plasmic strands; the somatic cells of aureus are round when seen

31

32

THE AMERICAN NATURALIST

[Vol. XLI

from the surface, and connected by very slender protoplasmic
strands, difficult for students to make out under the microscope.
Volvox glohator is monoecious: in the sexual stage both sperm
bundles and eggs may be found in the same colony at the same
time. The number of sperm bundles in a single colony is small.
Volvox aureus is either dioecious or monoecious proterogynous :
sperm and eggs are never found together in a single colony at the
same time, but the colony may contain one or the other exclusively.
However, I have found daughter colonies, some of which contained
bundles of sperms, others ova, within the same parent colony.
The number of sperm bundles in a single colony is very large.
SphcBrosira volvox Ehrenb. is an old name for the male colony of
Volvox aureus.

Kofoid ('99) gives the following key for the determination of
the two species:

Cells about 10,000 (minimum 1,500, maximum 22,000), angular with
stout connecting protoplasmic processes into which the chromato-
phore may enter. Diameter of colony about 700 fx (minimum 400,
maximum 1,200) ; diameter of ceil body 3-5 /i. V. glohator L.

Cells 500-1000 (minimum 200, maximum 4,400) ; rounded, with slender
connecting protoplasmic processes into which the chromatophore
does not enter. Diameter of colony 170-180 /i; diameter of cell
body 5-80 [i. Volvox aureus Ehrenb.

Klein ('99) gives illustrations of the general appearance of the
two species, including reproductive stages. Meyer ('96) gives
details of cell structure, with illustrations.

Volvox glohator is probably a better form for laboratory work
than V. aureus, and can be obtained in greater abundance if
secured early in the spring. Hence if Volvox is to be studied in the
fall by large classes it is w ell to preserve this species in 4 % formalin
in the spring; for many purposes the preserved material, if not
kept too long, is as good as the living. The study of preserved
specimens of V. glohator can then be supplemented bv' the living
V. aureus.

Keeping Volvox alive in the Lahoratory. — Terry ( :06) in experi-
menting on the galvanotropism of Volvox met with the usual
difficulty in keeping it alive in the laborator}% and concluded that
the organisms died of insufficient nourishment caused by improper

Xo. 481]

VOLOX FOR LABORATORY USE

food supply and poor light, but found it impossil)
either so that they would live for more than four d;
experience has convinced me that in our laborato
principal causes of the death of the organism has I
mineral substances in the tap water in which the s|
kept.

In collecting Volvox for our laboratorv it Iia> Ix t ii
bring in considerable quantitio of vcwctjihlc matcri;
Riccia, etc.) from the ponds in wliicli X'olvo.x occiirr
a little water, and place this material in shallow glas
with tap water. As fast as the organisms gathered
side of the dish they were picked off and removed to ;
tap water, it being deemed unsafe to leave them ii
dish exposed to the attacks of crustaceans. Su-
deleterious substances in the tap water, as well as
proper food supply, caused the death of the oriraiiisii
attempt to reproduce the natural couilitioii-. Wat-
Volvox was brought in in consiflcrahic (|uantinc>, to
small amount of the vegetai)l<' niatcriai. and place
glass dishes without the addition of tap uatci'. Tli

loss of water by eva|)orati()n and to k<'e|> out Itactrr'
except that when exj)osed to direct >unliulit it wa-- t'o -

to prevent a rise of temperature Ih-voikI tlic optiinuni.
was not changed at any time during- the cour>e of th -
in the majority of the atpiaria thus prepared. \'o|vo\
several weeks; in the fall of !!•().">. I'olro.r (inn tis \va>
several aquaria for from foiu- t(» ei<:ht week^; le>-
attained with Volro.v (jhihafor in the >{)ri

It was noted that I'niro.r (/Inhator chi-ters about de
larvie, perhaps on account of the presence of car
A moderate amoinit of decayiui;' plant or
water seems to be one of the essential c<»n
In its natural environment. N'olvox i> ot
stagnant water.

34

THE AMERICAN NATURALIST

[Vol. XLI

for Volvox. It is more difficult to keep the material alive in the
laboratory during warm weather because, in exposing the dishes to
sunlight, the water is likely to become warmer than that of the
ponds in which Volvox lives. This difficulty might be overcome
by placing the dishes where they will be partly immersed in the
running water of a shallow aquarium, and at the same time receive
an abundance of sunHght.

In case organisms that feed upon Volvox are too abundant, the
latter may be freed from its enemies by removing it with a pipette
when clustered at the Hghted side of the dish, to a dish of pond
water strained through bolting cloth to remove crustaceans, etc.

Obtaining Volvox in the Sexual Stage. — Both in the fall and
in the spring, material in the sexual stage was obtained in abun-
dance merely by keeping the organisms alive in the laboratory.
Several aquaria should be set up, and in some of them, specimens
in the sexual stage will usually be found in the course of one or two
weeks. It was noted by Dr. H. H. Newman that they often remain
hidden in the ooze at the bottom of the dish. When they reach the
sexual stage they seem to become less motile and consequently
drop to the bottom. This is especially true of sperm colonies in
V. aureus.

University of Michigan
Zoological Laboratory

LITERATURE

'99. Morphologisehe und biologische Studien iiber die Gattung Vol-
vox. Jahrb. f. wiss. BoL, vol. 20.
KOFOID, C. A.

'99. Plankton Studies, etc. Bull. III. State Lab. Nat. Hist., vol. 5.
Meyer, Arthur.

'96. Die Plasmaverbindungen und die Membranen von Volvox globa-
tor, aureus, und tertius, mit Ruckslcht auf die thierischen Zelleii.
Hot. Zrit. vol. 54, pp. 187-217.

:06. ( ialvaiiotropism in Volvox. Am. Journ. Physiol, vol. 15, No. 3.

OSTRACODA FROM S( )I TIIKAs rKKX MASSACHU-
SKT'IV

JOSEPH A. CUSIIMW

That the Ostracoda of New England have l.eeii u'lvaily ne^docted
may at once be seen by a reference to Miss Hailil>nir> list of tlie
New England Crustacea. At the time of its [nihlication there
was a single species reported from New Miin;laii(l and tliat from
but one locality. With a view to supplying this lack of records
in a slight measure, some collecting has been done in our ponds,
mainly about Boston. Several persons have kindly Mipplied
material which has now been placed in the collections of the Boston
Society of Natural History. Seven species are reported here^
making the number of species now known from the fresh water
of New England, nine instead of one. All of the records so far,
however, are from Massachusetts.

It has been a matter of interest to find the local distribution of
the species. Where a species is found at all it is usually abundant.
Of the three species of Cypris reported, all were found in ponds
in the vicinity of Boston and but a short distance apart, yet no one
collection contained more than a single species.

A number of other species have been collected but in immature
condition or in insufficient numbers for complete diagnosis.

The measurements given are average ones for the material
examined. As a rule, if adults alone are taken there is a noticeable
constancy in measurements but in cases where the collection con-
tains the young also, the range in measurements is considerably
greater.

It is to be hoped that more collecting will he done in the near

siderable addition to tlu" ]>re^ent list should he made. The seven
species representing live genera are given below.

36

THE AMERICAN NATURALIST

[Vol. XLI

Family Cyprididw

Subfamily Cypridinje

Genus Spirocypris Sharpe, 1903

1. Spirocypris passaica Sharj)e

Length 1.54 mm., height 0.7G mm., hreadth ().7S mm.

Spirocypris passa^m Sharpe, Proc. [ '. S. Xat. Mus., ynl 2(\, 1903, p. 9S2,
pi. 66, figs. 1-3.

This species and genus were described as new from material
in the U. S. National Museum, collected at Passaic, New Jersey.
There is no other record for it, as far as I know, up to the present.
It is especially interesting, therefore, to be able to record this spe-
cies from Massachusetts. Several specimens were obtained from
Wellesley, Mass., April 20, 1905, collected })y Mr. Irving 1.. Shaw.

The Massachusetts specimens were very sliglitly smaller than
the types but otherwise the specimens agreed veiy well. The
peculiar arrangement of the testes in concentric circles is very
apparent and striking. The original description gives the furca
as 23 times as long as wide. In the specimen measured from
Massachusetts the length was 0.486 mm. and the breadth 0.021
mm. A closer ratio of 23: 1 could hardly be obtained in such a
structure. Further collecting may show this sj)e('ies to be widely
distributed in New England.

Genus Cypkis O. F. Miiller, 1792
2. C3rpris virens ( Jnrine)

No. 481] MASSACHUSETTS OSTRACODES

37

3. Cypris fuscata ( -I urine )

Monoculus fuscatus Jurine, L c, 1S20, p. 171, I'.f. t^^. 1. 2.
Cypris fuscata Zaddach, L c, 1844, p. :r->.
Length 1.36 mm., height 0.81 mm., breadili O.:.') mm.
Near Fresh Pond, Cambridge, Mass. April 30, 100.). J. A. C,
coll.

In a very shallow pond-hole this species was very abundant on
the date given. As in the case of the preceding species, this is
very widely distributed and should be found throughout New
England.

4. Cypris reticulata Zaddach

Cypris reticulata Zaddach, /. c, 1844, p. 34.

length 1.20 mm., height 0.70 mm., breadth 0.60 mm.

Brookhne, Mass. April 10, 1905. Irving L. Shaw, coll.

Although this species is a very characteristic one and is widely
distributed in Europe, its occurrence in this country has hitherto
rested upon the single record of Dr. Sharpe. He found it in great
numbers at Normal, III., in a small grassy pool. It seems to
prefer such a habitat and therefore may be looked for in New
England in such places. The species was abundant in the col-
lection from Brookline.

Subfamily Cypridopsinjc

Cienus Cypridopsis Brady, 1868

0. Cypridopsis vidua (O. F. Miiller)

Cypris vuiua O. F. Miiller. En(^^

1792. p. 5"), tal). 4. fisrs. 7 0.
Cypridopsis vidua Brady. " A M*
coda," Trans. Li,i,i. S<>v. I.nml,
figs. 27-30, 4r>.

Length 0.64-0.75 mm., height 0.38-0.42 mm., breadth 0.42-
0.47 mm.

Small pond, West Cambridge, Mass. April 30, 1905. J. A. C,
coll.

38

THE AMERICAN NATURALIST [Vol. XLI

In tap water from Fresh Pond, Cambridge, Mass. Aug. 12,
1905. A. S. Pearse, coll.

Cohasset, Mass. Oct. 22, 1906. Owen Bryant, coll.

This species should be found everywhere in all kinds of fresh
water. It is probably the most abundant and one of the most
widely distributed of our fresh-water ostracods. It may be over-
looked on account of its small size.

Subfamily Cyclocypridinae
Genus Cypria Zenker, 1854
6. Cypria exsculpta (Fischer)

Cypris exsculpta Fischer, "Beitrag zur Kenntniss der Ostracoden."

Abhandl. math. phys. Klasse k. bayr. Akad. d. Wiss., vol. 7, 1855,

p. 18, pi. 19, figs. 36-38.
€ypria exsculpta Brady and Norman, "Monograph of the Marine and

Freshwater Ostracoda, Sec. I," Trans. Roy. Dublin Soc, ser. 2, vol. 4,

1889, p. 68, pi. 11, figs. 1-4.

Length 0.68 mm., height 0.44 mm., breadth 0.28 mm.

Woods Hole, Mass. Abundant in fresh-water pond, June 25,
1905, A. S. Pearse, coll. July 15, 1906, J. A. C, coll. Auburn-
dale, Mass., Oct. 28, 1906, C. W. Johnson, coll.

This species is almost as widely distributed and abundant as
the preceding. It should be found throughout New England.

Subfamily Candoninse

Genus Candoxa Baird, 1850

7. Candona Candida ((). F. :\Iiiller)

Cypris c(t>t(lii1n ( ). F. Miillcr, /. r., 17'.t2. p. ^Vl. tai). 0, figs. 7-9.
Candona cnndi^ln Lilljchorj:-, I), Cnistnnis , .,■ nnlinihus Tribus, 1853,

Length O.SC) !.:;<) mm., liri-lit 0.45 O.C.i; mm.

Arhngton, Ma>s.. May 7, l')()5, A. S. IVarsc, coll.; AuhunHlale,
Mass., Oct. 2.S. KKH;, (\ \V. Johnson, coU.

This species has not been reported from America as far as I
am aware. The .specimen.s seem to a,nret> well with European

No. 481] MASSACHUSETTS OSTRACODES 39

figures and descriptions and seem to be that species without doubt.
There is a very considerable range in the measurements given as
many young specimens and both sexes wen- in the lot measured.
An average measurement would be close to the inaxiuuiin given
here, if adults alone were taken.

This species, hke the others of its genus, has a crawlmg ha]>it
and may be in this way overlooked in collecting and in the exami-
nation of fresh material. It was not abundant in either of the
two collections in which it was found.

NOTES AND LITERATURE.

PHYSICS.

A First Course in Physics.^— These two books outline a thor-
oughly substantial course in elementary physics. They are obviously
intended to be used together, but each is complete in itself and either
(preferably the laboratory manual, as the authors themselves say
in their preface) could be used alone as the basis of a shorter course.

The essential feature of these books is their emphasis on the necessity
of showing a student "the hows and whys of the physical world in
which he lives" as well as the "how much" to which the reaction
from "the superficial, descriptive physics of thirty years ago" has led
us. For this reason, a great number of devices which are in common
use are explained with the help, in many cases, of admirable diagrams
of actual machines; as examples w^e may mention platform scales
for wagons, gas meters, two kinds of hydraulic elevators, the fire
engine, the railroad locomotive, hydraulic and steam turbines and gas
engines, artificial-ice and liquid-air machines, an excellent discussion
of the modem methods of heating and ventilating houses, a full descrip-
tion not only of the instruments used in telegraphy and telephony,
including the carbon transmitter, but also of the circuits themselves,
including even the new Bell central-battery system of telephony, auto-
matic signals and all, three pages of musical instruments, the Zeiss
binocular and, of course, wireless telegraphy. In the present instance,
the introduction of these illustrative digressions is governed by so just a
sense of proportion, and they are handled so well and are hacked by
so much thoroughly good physics of a more <juantitativt> sort, that the
result is much to be commended. It should always \)v iviiunibered,
however,— this is to be taken not as a criticism l)ut as a warning —
that this sort of thing may very easily become, in the hands of authors
and especially of teachers less scholarly than Pr()fes>or .Millikan
and Dr. Gale, an unfortunate return to the old-fashioned ^uprrficial.
descriptive "natural philosophy" which thev themselves so dcHnitely
deplore.

'Millikan, Robert Andrews and Gale. Henry Gordon. .4 First Course in
Physics. Boston, Ginn & Co., 1906. 8vo, viii + 488 pp.

Millikan, R. A. and Gale H. G., A Laboratory Course in Physics, for Sec-
ondary Schools. Boston, Ginn & Co., 1906. 8vo, x + 134 pp.

41

42

THE AMERICAN NATURALIST [Vol. XLI

Another interesting feature of these books is the free use which is
made in quahtative explanations of such conceptions as the kinetic
theory of gases, the ionic theory of electrolytic conduction, and the
wave front in geometrical optics. Whether or not it pays, for instance,
to displace the old ray-optics, which must, of course, be properly
interpreted, by the more valuable but also more difficult notion of
the wave front, is a question of pedagogy which each teacher must
decide for himself. Fortunately the treatment of the most danger-
ously spectacular part of our modern physics is confined to the last
twelve pages of the text-book, where there is an account, admirable
as regards both interest and conservatism, of vacuum tube phenomena
and of radio-activity, including some of the evidence for the existence
of electrons, together with brief statements of the corpuscular theory
of matter and of the disintegration theory of radio-activity.

Many other features, while not unique, are nevertheless worthy of
much praise. For instance, the experiments, both for the laboratory
and for the lecture room, are ingeniously simple and yet, so far as one
oan judge without trying them, entirely effective.

The typography is good, and the illustrations are most excellent,
both in technique and in conception; and the sixteen full-page half-
tones of eminent physicists, each with a short paragraph describing
the man's life and work, are a notable addition not only to the attrac-
tiveness but to the real value of the books.

H. N. D.

BIOLOGY.

Jennings' Behavior of the Lower Organisms.^ — It is now nearly
a decade since Professor Jennings published his first brochure on
the reactions to stimuli in unicellular organisms. The intervening
period has been one of continuous activity on his part in the study
of animal behavior, especially aiiioiii,^ the lower organisms. His
investigations have not been strictly ((.iifiiKMl lo the Protozoa for
among the score or more of titles of inipoitaiit contributions from his

' H. S. Jennings. Behavior of the Lower Organiams. Columbia University.
Biological Series, New York, The Macmillan Co., 1906, 8vo. xiv + 366 pp.,
illus. $3.00.

No. 481] NOTES AND LITERATURE

43

pen are studies of the reactions of Metridiuin and of rotifers. Nor
have his investigations been limited to the animal world alone for
groups on the border lines such as the flagellates and liiiet<Tia have
also been included. All students of these groups and especially
investigators of animal behavior and workers in the field of eoni|)ara-
tive psychology will Gjid cause for congratulation in the fact that
Professor Jennings has taken this opportunity to restirvey the whole
field of his experimental work and to summarize and restate his con-
clusions in this most important field of research. While many studio
in this field have been made primarily from the standpoint of the
psychologist, or have been of a desultory character, or are but partial
in scope, the work summarized in this book has been dominated by
the broadest scientific spirit, has been conducted with the greatest
care and thoroughness, has included in its scope all possible avenues
of approach to the analysis of animal behavior, as exemplified in the
simplest organisms, and has been carried through to a stage of com-
pletion where fundamental generalizations are possil.h-. Thv work
of others in this field whether in a,<:recinciit or not. with the author's
conclusions, is treated with liillne» and tainic-. 'i1ie hook thus
becomes an exemplification of the value of intensive r(\search, an
indispensable authority for any who wish to become familiar with the
latest results in the field of animal psychology. As illustrative of the
thoroughness with which the analysis has been carried out we find
that in Paramecium the structure and the normal movements are
described and correlated, and the reactions to chemical and mechanical
stimuli of various sorts determined, the ab-sence of reaction to liiiht
but the sensitiveness to the ultra-violet rays noted, as ar<' also the
reactions to heat and cold, to induction shocks and a constant current
of electricity, to water currents, gravity, and centrifugal force. The
relation of these actions of orientation to other reactions is carefully
analyzed. The behavior of Paramecium in daily life in the aquarium,

44

THE AMERICAN NATURALIST [Vol. XLI

haphazard, in character. The phenomena are thus similar to those
shown in the "learning" of higher organisms, save that the modifica-
tions depend upon less complex relations and last a shorter time.

Each organism is found to exhibit a set of actions made up, in the
case of the lower organisms, of a few factors combined in various ways
in a coordinated system which Professor Jennings designates as "the
action system." For the term "motor reaction" employed in his
earlier papers the phrase "avoiding reaction" is now used to designate
the stereotyped method of reaction of Infusoria to most stimuli. The
author rejects the local action theory of tropisms as a "more or less
artificial construction, made by combining certain elements of behavior
and omitting others that are of most essential significance." In its
place he proposes the method of "trial and error" as an explanation
of behavior. The stimulus interferes with definite internal processes
occurring in the organism and this interference causes a change in
behavior and varied movements which subject th(^ organism indis-
criminately to many different conditions. It merely acts in all sorts
of ways possible to it. When oih- of these new conditions thus met
relieves the organism from the existin*;; int(>rferenee with its life proc-
esses, the trials cease.

As a second cornerstone in the formulation of behavior we find the
law of "resolution of physiological states" thus stated: "The resolu-
tion of one physiological state into another becomes easier and more
rapid after it has taken place a number of times." It appears that
even in Stentor and Vorticella repetition of an action brings the second
step in a sequence in behavior more quickly upon the first. Here lie
the foundations of the phenomena which are usually designated as
habit formations, memory and learning, and the question may well
be asked w hether they are not coextensive with life and based funda-
mentally on the physical and chemical structure of colloids.

C. A. K.

Modernized Darwinism.^ — Professor Guenther has written a very
readable book on Darwinism and allied biological problems which
the tyro will find quite intelligible. The translation seems good
and the publishers have done their part well. The treatment of
the subject is rather novel, moSt of the chapters being divided tax-

' C. Guenther. Darwinism and the Problems of Life. Translated from the
third edition by Joseph McCabe. London: A. Brown & Co., 1906, Dutton
& Co., New York, American agents. 8vo, 439 pp.

No. 481]

AT)77' N . t M) 1. 1 TKRA T I HE

45

inomically under the headino-s

which to hang certain l)i()l<)<iic;
are considered: protective colo
of animals; under l)inls, sc:
reptiles and amphibians, the d
life; under fishes, the origin (
organs, and the biogenetic la
inheritance of acquired charac

arrangement does not lend itself to ,
ment of the subject but the result is (
Not only is the book interestingly
safe one. No evolutionary henv^io
simon-pure, orthodox natural selcctic
as befits a scientific book, n fereiicc
theory and that of orthogeiuvsis. 'V

we know tl.;,
orthogenesis

dation of it
scientific wo,

of the We
chapter h\
essary lini

46

THE AMERICAN NATURALIST [Vol. XLI

Momentum in Variation. — It is a little late to criticize an article
that appeared in November, 1905, nevertheless I should like to say a
word or two in regard to the paper by Mr. F. B. Loomis entitled
"Momentum in Variation." The conclusion is reached that a varia-
tion started along any line tends to carry that line of development to
its ultimate, being driven by momentum. If the feature is detrimen-
tal, the group dies out. If, however, it is merely a minor feature, it
makes a handicap.

No one doubts that in the course of evolution, specialization goes,
so far as to carry a given species or group out of existence, but that
this is of such widespread occurrence as Mr. Loomis implies, is open
to doubt. Certainly the statements which he adduces to support
this theory, are in many cases erroneous and in other instances open
to quite other deductions than are placed upon them.

The few comments here given, are not at all in the line of captious-
criticisms, but are merely intended as a protest against any such
short-cut to a solution of important problems as that taken in the
paper in question.

If we begin with the Saber-toothed Tigers, which are cited as-
examples of extinction due to overdevelopment, we may go back
about twenty years to the time when Professor Cope reached a similar
conclusion, saying in the course of some discussion that Smilodon
undoubtedly became extinct because it could not obtain food, where-
upon someone present arose and said: "Mr. Cope, what did the
Smilodon feed on"? In connection with this Dr. Matthew has
recently brought forward some facts tending to show that the long-
tusks of Smilodon were of service in cutting through the long hair and
thick hide of some of the contemporary ground sloths. Certainly if
the tusks of Smilodon caused its extinction, why does not the Walrus-
die out for a similar cause ? The Mammoth with its extreme develop-
ment of tusks is also cited to illustrate the principle of momentum in
variati6n, leading to extermination, but the great Gangetic elephant
which shows the most enormous development of tusks, became extinct
long ago, while other members of the race whose tusks were far more
recurved lived on. Nor did those mastodons in which the tusks were
greatly curved, come to an untimely end one whit sooner than their
contemporaries with fairly straight tusks. The African Elephant,,
which is much the most primitive in structure of existing species, and
more nearly resembles E. ganesa in tusk development, is the species-
that has thrived best. Moreover, the African elephant is the one in
which tusks are present in both sexes while a large proportion of the

No. 481] NOTES AND LITERATURE

females of Asiatic Elephants are tusklcss, so that here \\c li;i\<- ;i ( use
in which tusk development has gone beyond sex diliVrctitiiuioii.

If Babirussa seems to be handicapped bv its teeth, though tliere is
another side to the case, how about Mesoplodon, in one species of
which the teeth lock over the beak so that the animal can open its
mouth for a short distance only and yet shows no signs of i);is.sing out

The elongation of the snout of Teleosaurus is cited as aii..ili.T .11.-
advantageous character but the Gangetic (Javial in wliirh ihr mioih
is nearly as long, finds this of great service in calcliiiig as un-
doubtedly Teleosaurus did, aii.l Dr. Al.rl -iNvs rl,,noaiinn ..f snout
as characteristic of fresh-water (ctaccaiis.

Stegosaurus did not come to an end on account of it^ licaw aruior
but from some other cause, for llie active predatory <linosaurs. sndi
as Allosaurus, that were unincumbered by any defensive ainior, died
out just as did their heavier-plated contemporaries. The male
Narwhal which has a single long tusk lives in the sann- ^ a and jn-i
as long and happily as his tuskless spouse, and many similar in-iances
might be cited. The problem of the extinction of aiiimaf- far too
complicated to be decided in haste and few of ihc ( xamplo ( iied by
Mr. Loomis seem to be conclusive.

Xenia in Wheat.' — As everyone has noticed, when white sweet
corn is pollinated whh red corn the outer part of the grains, although
not truly part of the embryo, is red. This is a case of so called xenia.
Xenia has been observed in other ca.ses also, notably in beans and
in wheat. Tschermak has recently studied xenia in wheat in more
detail. He experimented with two races — the Hanna wlu-at and tiie
Petkus wheat. Both kinds of wheat when in bud yiehi both given
and yellow seeds;^ but yellow Hanna wheat gives 80% of yellou grains

The green Hanna and the vellow I
only about half of their own kind
that when green and yellow llanna
resulting show the color of the mot

Likewise when the green and i!ic

licher Kreuzung, II." Zcitschr. /. d. Inndw. Vrrsun
pp., Feb., 1906.

48

THE AMERICAN NATURALIST

[Vol. XLI

the seeds have only the maternal color. On the other hand, when the
opposite colors are derived from different races, and, especially, when
the father is either yellow Hanna or green Petkus, the paternal char-
acter shows strongly on the seeds. Consequently, xenia is better
manifested in wheats that are not very closely related than in those
that are.

C. B. D.

ZOOLOGY

Folsom's Entomology.^ — Dr. Folsom's new work occupies a
unique place among entomological text-books. As stated in the
preface, "the book was written in an effort to meet the growing
demand for a biological treatment of entomology." To this end the
systematic side of the subject has been confined to a mere outline of
the orders, following essentially the system of Brauer. The external
anatomy, too, has been very briefly touched upon as that has been
emphasized by the current texts.

On the other hand, there is an admirably clear-cut discussion of
the elements of internal anatomy and of physiology. The value of
this chapter lies not only in the careful organization of the material
presented but in the omission of a vast amount of detail. The author
has followed a common error in stating that the alary muscles are
unstriated. He speaks of the follicular cells of the ovary as derived
from the primitive germ cells, — a view which is not held by recent
investigators of this subject. In view of the decisive work of Petrunke-
witsch and otlicr of WcMsinann's students one is surprised to see, p.
145, the stntrnirni thai ' males may, of course, result from fertilized
eggs, as ill ilic honrv-lHc. according to Dickel."

The cluiincr on <1( \ clopincnt likewise shows the virtue of vigorous
pruning. There is a wry brief but (excellent outline of the embryo-
logical development, wliilc tli(^ <ricat( r portion of the chapter is devoted
to the postembryonic (Icvclopiiiciit.

■ Folsom, J. W. Entomology, vnth Special Reference to its Biological and
Economic Aspects. Philadelphia, Blakiston's Son & Co., 1906. 8vo, vii +
485 pp., with 1 col. pi. and 300 illustrations.

No. 481] NOTES AND LITERATURE

49

The remainder of the text is largely devoted to biological phases of
the subject. Much material which is not accessible in any other text
is here brought together and is treated from a broad l>i<)l()^'i( Ml view-
point. The subjects of color and coloration; the origin of aduptJitinns
and of species, distribution; the relation of insects U> plants and lo
other animals; their interrelations and their behavior, are treated in
a concise but most readable and interesting manner.

Though the method of treatment is professedly economic as well as
biologic, the practical aspect of the subject receives but scant attention.
The relations of insects to plants, and to other animals, are discussed
from the view-point of the biologist. Six pages are devoted to an
excellent summary of the important subject of the transmission of
disease by insects. The sixteen pages on insects in relation to man
are largely given over to a statement of the importance of the subject
and to an historical sketch of the progress of economic entomology m
America.

The illustrations are excellent and, in many cases, new and prepared
by the author. Such as have been copied are very carefully credited.
An extensive and carefully arranged bibliography will be vefy helpful
to the student.

Dr. Folsom is to be congratulated on the clear, concise, and interest-
ing presentation of his material. The book is one which is bound to
prove stimulating, and which every worker in the field of entomology
and every teacher of zoology will want in his o\vn librar>^ Whether
it will meet the present day demands for an entomological text-book
is a question. ^

Additional Observations on Hyla andersonii and Rana virgatipes
inNew Jersey.— An elfort %Na.^ made thi> past sunitn.T to add to the
observations on Jli/la amlrr.nnn and Rnna i,rqni>}»s published m

two pre\

i observed in

(■S..f /////'/

-vere attracted to a fe

50

THE AMERICAN NATURALIST [Voi.. XLI

miKklv bottom ot ii pool. The under parts are lighter and show a
gold( n t n ^^ hu h sonietunes extends uj) the sides The sindll hind
liu. .Un^^ « ..ih ..i. tIu ( nds ot the toe. the disk that are so fonspic li-
ons in the niai iiiv 1 1 vh.. The tail is spotted, and there is usually a dark

nlari)lot( lu s as on tlu tails of the tadpoles of Ihjla uisicoloi W hen
the tail IS nearly absorbed, and they leave the water, they are about
25 mm. long and of a dull olive green. Thev grow lighter, that is,
brighter gre«Mi in hue witli the disappearance oi the tail, until the
little frogs, winch ni length ol l)odv are 15 mm., reseml)le the mature
individuals. 1 h<- white ih;ii margins the green of the back and ex-
treimtus is not m) < onspK nons ds in the adults, and tlu saffron of the
under parts is wantmg m those that 1 have examined. The narrow
band of purplish brown that commences at the nose and extends
through the eyes and so down the sides is conspicuous in the little
frogs before the last remnant ot the tail has disappeared.

The adult Ihjla andinonti is amu ingU activt at night and jumjjs
about the lower limbs of the trees and on to the bushes with much
agility. They seem rarely to climb over five or six feet from the ground.
They sit upright and look pert, and if interrupted in the midst of their
song they leave their l)nbl)les blown up until such time as the intruder
goes away or stands still. In tlie day time they are usually quiet and
forthe most [>ait hide in t he da nip moss and leaves lying on the ground.

On [\\v warm elondy evening of August 10th, Hyla andersonii was

ingdale, .\. J. ^^Iiis locality is 15 miles northeast of Lakehurst,
which has been I he most northern locality for the frog heretofore
recorded.

Harm rinjatipr, niav be .alle<l the Carpenter Frog, for its note

uttered rhurk-np. rhurh-up, and tin- frog usually hammers from three
to four limes. Vur a time I was not sure of the singer, .but some
caprive iiidiv idnals under the influence of good living have uttered
this eall-non- in my room while I sat by. These frogs domineer over

would hold his head down in the meekest manncM-, and h(> becaiiM^ so
cowed that if I touched him at any time with inv finger, he assnined
the humble position. Miss Dickerson in The Fnxj Booh say> that

52

THE AMERICAN NATURALIST [Vol. XLI

("Ueber die Kolbenzellen in der Epidermis der Fische." Jen.
Zeitschr. /. Naturwiss., vol. 40, pp. 589-646, pis. 22-26) goblet cells
occur in the epidermis of cyclostomes and most physostomous teleosts.
All goblet cells are modified epithelial cells from the deepest or germinal
layer of the epidermis. They are undoubtedly specialized unicellular
glands which may have in addition some supporting function.

The Selachian Eye. From a study of the eyes of some eighteen
species of sharks and rays Franz ("Zur Anatomic, Histologic, und
functionellen Gestaltung des Selachierauges.' Jen. Zeitschr. f. Natur-
wiss., vol. 40, pp. 697-840, pi. 29), has shown that while there are
many specific differences, the eyes of this group as a whole are clearly
distinguishable from those of other vertebrates. What is especially
peculiar in them is the tapetum lucidum, an epithelial musculature in
the iris instead of the usual mesodermal one, a specialized zonula
zinnii, and the absence of a falciform process characteristic of other
fishes. The adaptations shown by the eyes of different species are
discussed at some length.

G. H. P.

BOTANY

Bergen and Davis's Principles of Botany.^ — One of the most
successful American elementary botanical text-books has been Bergen's
Foundations of Botany. With its author. Dr. Davis has been asso-
-ciated in the preparation of the present book, which is certain to find
favor with the users of its predecessor and to ^\'m many new friends
since in addition to what was best in the earlier text there is now
given a consecutive series of studies of representative spore plants
so treated as to outline the evolutionary history of the plant world.
Both authors are experienced teachers, and also familiar with research
problems at first hand, and they have brought to their task unusual
care in grouping and handling the subject matter an<l in well illus-

The book is said to furnish material for a full velar's work. It
contains, indeed, enough to occupy considerably more than this time,

1 Bergen, J. Y., and Davis, B. M. Principles of Botany. Boston. Ginn
& Co., 1906. 12mo, ix + 555 pp., 14 pL, 402 figs.

No. 481] NOTES AND LITERATURE

58

if all of its topics were thoroughly worked over; but tli(> purpose of
its authors has been to present somewhat more than is likrly to he
used, so that individual teachers may find it comprehensive enougii
to base on it courses adapted to their several needs. It is safe to say
that it will be a much consulted book even in laboratories where other
manuals are used to outline the courses given.

Rydberg's Flora of Colorado.'

1906) deals with Hioloiry

"Development ol" :\I(.rph(
Development of Biolo^ry "
ficial and Natural Selcn iio
"The Fundamental Proli
Wiesner, "The Devclopiu
of the Other Sciences '; 1
lems"; Arthur, "The flis
"Vegetable Patholog^ an
of Ecologv in Miulvvn

54

THE AMERICAN NATURALIST [Vol. XLI

The second volume of Postelsia, the Yearbook of the Minnesota
Seaside Station, issued from the Pioneer Press of St. Paul, contains
the following papers: — Rosendahl, "Observations on Plant Dis-
tribution in llenfrew District of Vancouver Island"; Butters, "The
Conifers of Vancouver Island"; Evan;,. " ne})atic<e of Vancouver";
Hone, "Sonic Wr.t.-ni Hchcilinc.c"; (;rijr<r>. -Rrn/rnria parrnh.
a New Kcli, fnmi Vniunnv.-r Islaiul": Ilcnkcl. "A Study of Tide-
pools on the W.-st (■..;isi ..f V;nic<.uv(M- Islnnd": and H.-ill. "Somc
Geological Fcalun-s ot \\\v MiinKvsom Seaside Slalion."

aerial roots, is^descril)e.l an.l liu-n,vd for Ileniii'elia by Sehoutc'iu vol.
20, part 2, of the Annah v <hi Jnnlin Hntnnninr <l, llinh ir.nn/.

An iilnstrared paper on llie ni.'dnllnry ravs ,,f ConifeiN. bv Tassi.
forms part of the ree.^ntlv i.sii,.,| vol S. fa^e 1 I, of the Hnllrltinn
del Lahnralnrn, ul Ortu linUnnn. of tin- I .n^e,.it^ ot >iena

An illnstnu.'d paper on ll.e leaf strnetinv of certain New Zealand
plant.. In Mj.. Ilrrriott. i. pnbli.hed in xol. :5s of the Tmnsa,finn^

other papers of boianieal interest

cellulose '.k<.|,t<.n "'reinaininn- after tin' .j.'liiinin. atioii ot oth.^r^. i>

^h.^som■i lintnnlrni (:nrl'!r'''^ ''''' ' ' '

re^peeti^(■l^ of the rec'eniU i..„cd fnxt and .ixtli Nohnne' of the li,^ unl

Part 11 of Koorders \ aleton-> " VdditanH'nia ad ( oonil ionein
Flone Arborea' Javaniea'" ha^ iveenilv be.^i, i.>ne.i at Batavia as no.

Ana<<onniol iheboianN ot ( -hri.t nia-^ Idand. bx l{idle^ , i. < onta ined
in iIm. rcM-Mllv i..„.d no lo ,,f ,he .hnnnni fin Slr,nt^ Hnnuh n/

A number of gcneri
signature of his Leafii t.

56

THE AMERICAN NATURALIST [Vol. XLI

The dune-fixing planting of Cape Cod is illustrated by Birge in
The American Inventor for September.

An illustrated forestal account of Sequoia is given by Sterling in
School Science and Mathematics for October.

A series of "Botaniker Portrats/' each accompanied by a short
but comprehensive biographic sketch, is being issued in quarto fascicles
by Dorfler of Vienna. Judging from the two fascicles thus far issued,
the quality of execution and accuracy of text are unimpeachable.

Kellogg contributes a paper on the scientific aspects of Luther
Burbank's work to The Popvlar Science Monthly for October.

PUBLICATIONS RECEIVED

Bergen, J. Y., and Davis, B. M. Principles of Botany. Boston and New
York, Ginn and Co., 1906. 12mo, ix + 555 pp., illus. $1.50.— Guenther,
C. Darwinism and the Problems of Life. A study of Familiar Animal Life.
London, A. Owen and Co., 1906. 8vo, 426 pp.— Holder, C. F. Half Hours
with Fishes, Reptiles, and Birds. New York, American Book Co., 1906.
12mo, 255 pp., 244 figs. 60 cts.— Hough, T., and Sedgwick, W. T. The
Human Mechanism, its Physiology and Hygiene and the Sanitation of its Sur-
roundings. Boston, Ginn and Co., 1906. 12mo, ix + 564 pp., illus.—
Howe, R. H., Jr., and M. A. Common and Conspicuous Lichens of New
England. A Fieldhook for Beginners. Part IV. Boston, W. B. Clarke and
Co., 1906. 16mo, pp. 57-71, illus. 50 cts.— Ingersoll, Ernest. The
Life of Animals. The Mammals. New York, the Macmillan Co., 1906.
8vo, xi + 555 pp., illus.— Jennings, H. S. Behavior of the Loiver Organisms.
New York, The Macmillan Co., 1906. 8vo. xiv + 366 pp.. illus. ?.3.(X).—
LiNviLLE, H. R., AND Kelly, H. A. .4 Text-book in C nn-nl Z..ul,yp,. }\u<Um.
Ginn and Co., 1906. 12mo, x + 462 pp., 233 text-fii.-. >1 Mii i ik \ \.

R. A., AND Gale, H. G. A Laboratory Course in J'h;/>ir.. /,,,• >, ,
Boston, Ginn and Co., 1906. 12mo, viii + 131 i>p ■ iHn^ ,M<.m.,mm,:,.v.
T. H. The Analysis of Racial Descent i>, Animnf... .Xcw ^ mk, Henry H.,lt
and Co., 1906. 8vo, xi + 311 pp.— Skkm. \V\i. W.. am. Hr.\>;i>i.\. ( o.
Pagan Races of the Malay Peninsula. Xt-w York. Thr MMcnullan Co., I'.HiG.
8vo, 2 vols., xl + 724, x + 775 pp., illu.s. sKiOi). W imch r. ( T. /■,././.
Laboratory, and Library Manual in Physical (r, n,im pi,,/. P,o-io!i and New
York, Ginn and Co., 1906. 8vo, xii + 178 pp. - 101 pj). »( rulo<l .<heet.s.
$1.00.

Adams, G. E., and Wheeler, H. J. Continucus Corn Culfure. R. L
Agrie. College, bull. 113. pp. 99-114.— Barhkk. H. C. Htiiiipt.ra from
Southwestern Texas. Sci. Bull. Mus. Brookh/n Inst. Arts nn,l Sri., vol. 1,

Proc. U. S. Nat. Mus., vol. 31. pp. 109-160. i)ls. 3 .V - HA iifj i;. I. \. The
Species of Botryocrinus. Ottawa Nat., vol. 20. pp. '.I3 loi. {'.kan. T. H.
A Catalogue of the Fi.shes of Bermu<la. with Xotes on a Collection made in
1905 for the Field Maseum. Field Cnl,imh,nn Mus.. v.n A. <.>r.. ^•o]. 7. no. J,
pp. 21-89.— Blatchley. W. S. The IVrrolcuni Indintry of Soutlu.a^t.-rn

les Gennadas ou Pem'Mdes bathyp«>lagiqu(vs. linlL .\l n,-. ,i. ]!..■„,...

no. 80, 13 pp.— BouviER, E. L. Observations >ur h- I'. n/ni.- .In i.rr

Haliporus Sp. Bate. Bull. Mus. Ocianogr- ^< -1^ " ' ^' " I P

BowNocKER, J. A. Salt Deposits and the Suit Itulii>try in Ohio. i,.,,/.
Surv. Ohio, ser. 4, bull. 8, xv + 42 pp., 6 figs.— Cakv, .M. On the Diurnal
Lepidoptera of the Athabaska and Mackenzie Region, Briti.sh America.

58

THE AMERICAN NATURALIST [Vol. XLI

Proc. U. S. Nat. Mus., vol. 31, pp. 425-457.— Cobb, J. N. The Commercial
Fisheries of Alaska in 1905. U. S. Bureau Fisheries, doc. 603, 46 pp.— Cobb,
N. A. Methods of Using the Microscope, Camera-Lucida and Solar Projector
for Purposes of Examination and the Production of Illustrations. Ist Ann.
Rept. Div. Path. Phys., Eccp. Sla. Hawaiian Sugar Planters' Assn., 29 pp.—
CoMERE, J. Observations sur la periodicite du developpement de la flore
algologique dans la region toulousanie. Bull. Soc. Bat. France, ser. 4, vol. 6,
pp. 390-407.— Bellinger, O. P. Locomotion of Amoeb« and Allied Forms.
Journ. Exp. Zool, vol. 3, pp. 337-358, pis. 1-2.— Evans, A. W. Notes on
Japanese Hepaticae. Proc. Washington Acad. Sci., vol. 8, pp. 141-166, pis.
6-8.— EvERMANN, B. W., AND Seale, A. Fishes Collected in the Philippine
Islands by Maj. Edgar A. Mearns. Surgeon, U. S. Army. Proc. U. S. Nat.
Mus., vol. 31, pp. 505-512.— Fairman, C. E. New or Rare Pyrenomycetese
from Western New York. Proc. Rochester Acad. Sci., vol. 4, pp. 215-224,
pis. 20-22.— Farringtox, O. C. Zoisite from Lower California. Field
Columbian Mus., geol. ser., vol. 3, no. 4, pp. 55-57, pi. 28.— Fawcett, H. S.
Variation in Ray Flowers of Anthemis cotula and Other Composites. Proc.
Iowa Acad. Sci., 1905, pp. 55-59, pis. 12-20.— Fisher, W. K. New Star-
fishes from the Pacific Coast of North America. Proc. Washington Acad. Sci.,
vol. 8, pp. 111-139.— Felt, E. P. The Gipsy and Brown Tail Moths. N. Y.
State Mus., bull. 103, 42 pp., 10 pis.— Felt, E. P. Twenty-first Report of
the State Entomologist on Injurious and Other In.sects of the State of New
York. N. Y. State Mus., bull. 104, 186 pp., 10 pis.— Fernald, H. T. The
Digger Wasps of North America and the West Indies belonging to the Sub-
family Chlorionime. Proc. U. S. Nat. ^/ms., vol. 31, pp. 291-423, pis. 6-10.—
Garcia, F. European Grapes. N. Mex. Coll. Agric. and Mech. Arts, Agric.
Exp. Sta., bull. 58, 32 pp.— Gardner, N. L. Cytological Studies in Cyano-
phycea?. Univ. of Calif. Publ, hot., vol. 2, pp. 237-296, pis. 21-26.— Gil-
more, C. W. Notes on a Newly Mounted Skeleton of Merycoidodon, a Fossil
Mammal. Proc. U. S. Nat. Mus., vol. 31, pp. 513-514, pi. 12.— Herrera,
A. L. Invasion de gusanos en los estados del centro de la Republica. Com.
Parasitol. Agric, circ. 45, 14 pp.— Herrera, A. L. Destruccion de los
mosquitos en las habitaciones con el polvo de crisantema. Com. Parasitol.
Agric, circ. 48, 5 pp.— Hrdlicka, A. Anatomical Observations on a Collec-
tion of Orang Skulls from Western Borneo; with a Bibliography. Proc U. S.
Nat. Mus., vol. 31, pp. 539-568.— Inda, J. R. Una plaga de in.sectos llama-
dos " frailecillos" en el valle de Mexico. Com. Parasitol. Agric, circ. 46, 8 pp.

dnrttifopi' m. Hull. M iis, Ocmmir. de Monaco, no. 79, 6 pp , 1 pi.— Jordan,
1). S. AM) SxvhKii. .1 < ). A Hcviow of the Pceciliidse or KiUifishes of Japan.
Prnr. f. .V. \ut. vol. -.U. pp. 287-290.— Jordan, D. S., and St.\rks,

E. C. A Hcvi.-u ..f ihr I loinulci- and Soles of Japan. Proc U. S. Nat. Mus.,
vol. 31, pp. 101 JK,. .I.MuiAv, 1). S.. AM) Si AUKS, E. C. Notes on a Collec-
tion of Fi.sho. troin l>,)rt Arfhur. MMii.lnnia. Obtained by James Francis
Abbott. Prnr. ( . S. \<,t. Mus.. v.,1. A\ . pp. 1 6-.526.— JouBiN. L. De.scrip-
tion des nemerticns hathyprln^i.iues iat>tures au cours des demi^res cam-
pagnes du Prince de .Monaco (1<S90-19()5). Bull. Mus. Oceanogr. de Monaco,

Xo. 481]

Pl'BLICATIOXS Rh:CEIVEl)

59

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THE AMERICAN NATURALIST [Vol. XL!

Meteorites. Proc. Rochester Acad. Sci., vol. 4, pp. 225-231, pis. 23-25.—
Weller, S. The Geological Map of Illinois. III. Geol. Surv., bull. 1, 26 pp.,.
map.— Wheeler, H. J., and Hartwell, B. L. Commercial Fertilizers.
R. I. Agric. Exp. Sta., bull. 115, 114 pp.— Wheeler, H. J., Hartwell, B. L.,
Wessels, p. H., and Gray, J. P. Commercial Feeding-stuffs. R. I. Agric.
Exp. Sta., bull. 112, pp. 77-96.— Whitman, C. O. The Problem of the
Origin of Species. Congr. Arts and Sci., Universal Exposition, St. Louis,
1904, vol. 5, 18 pp.

Condor, vol. 8, no. 5.— Economic Geology, vol. 1, nos. 6, 7.— Eltka,
vol. 7, no. 45.— Indian School Journal, vol. 6, no. 10.— Journal op Geog-
raphy, vol. 5, no. 7.— Journal of Geology, vol. 14, no. 6.— Kentucky
Agricultural Experiment Station. Report on the Enforcement of the-
Pure Food Law. 197 pp.— Le Mois Scientifique, vol. 8, no. 10.— Missouri
Botanical Garden. Seventeenth Annual Report, 1906, 181 pp., pis.—
Museu Gceldi. Boletim, vol. 4, no. 4.— Natur.^: Novitates, vol. 28, nos.
12-15.— Nature-Study Review, vol. 2, no. 6.— La Nuova Notarisia,
ser. 17, Oct., 1906. — Rochester Academy of Science. Proceedings, pp.
231-344.— St. Louis Medical Review, vol. 54, nos. 9, 10, 12.— University
of Colorado Studies, vol. 3, no. 4.

(No. 480 was issued December 19, 1906)

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THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL
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CONTENTS

I. An automatic Aerating device for Aquaria . . . DE. LOXTIS MURBACH 61
n. The Flying-Fish Problem ..... LIEUT.- COL. C. D. DTTEN'rORD 65
ni. Cataloguing Museum Specimens .... PSOFESSOB L. B. WALTOH 77

IV. Some South American Rotifers JAMES MURRAY 97

V. Meristio Homologies in Vertebrates - J, S. ZINGSLEY 108

VI. On the Osteology of theTubinares DR. R. W. SHTIFELDT 109

Vn. Hotes and Literature : General Biology, Transmutation and Agriculture,
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Quaternary Remains of Man in Central Europe, Parian Races of the
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WILLIAM S. BAYLEY, Ph.D., Colby University, Waterville
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell University, Ithaca.
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA, M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University

CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM. Ph.D., Lake Forest University
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, Pittsburg
D. P. PENHALLOW, D.Sc, F.R.S.C., McGill University, Montreal
H. M. RICHARDS, S.D., Columbia University, New York
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ER^IN F. SMITH, S.D., U. S. Department of Agriculture, Washington
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington
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HENRY B. WARD, Ph.D., University of Nebraska, Lincoln
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THE

AMERICAN NATURALIST

Vol. XLI February, 1907 No.

AX AUTOMATIC AERATING DEMCT. EOR AQUARIA
LOUIS MURBACH

The use of the water blast as a means of aerating is well enough
known to need no more than passing mention. Its cost is prohi-
bitive for smaller schools and its use Hmited to lahonitoric^ where
noise would not be a disturbing factor. Furtlu'riiKuc it may be
desirable to have an aquarium under observation in dillVn iit rooms.
Some time ago a simple device was described A'r//.. noI. ;!s.

no. 453, 1904, pp. 655-661, 2 figs.) which, howrvcr. uvvv>^\vmv~^
the exchange of the water in the aquarium. 'I'hi- mi^ir involve
the loss of organisms if the flow were contitmous a> from a tap, or
it would necessitate lifting the water perio(hCallv.

These were some of the difHculties I eiicount(MV<l uhcn about a
year ago I wished to aerate some small a.|iiaria ci.maim'ng sea
water, in a class room where fresh water wa> availal)lr. but a flow
of sea water could not be had. .Vii oivh'iiary fihcr or varuuin
pump was fitted into a calcium-chloride jar about 4o cm. tall.
The accumulating air in the jar was carried through the -topper
hy a small tube to the aquarium. \ ball valve of paraffin held
against the lower opening of the jar by a lever and weight wa> to
regulate the outflow of water. This and siiiu'lar devices tri<-d.
failed to regulate the varying presstn-e in the su[)ply pipc<. and wa-
not satisfactory. If regulated for the day when more tap- were in
use, the pressure increased during the night, with few or no other
taps on, so that the fresh wattM- overflowed through the air tube
and diluted the sea water.

The above obstacles were entirely overcome in a devic(> that I
Jiit upon the past sttmmer at the Marine Riol.)gical Lal)oratory

61

62

THE AMERICAN NATURALIST [Vol. XLI

64

THE AMERICAN NATURALIST [Vol. XLI

pinch cocks until the desired flow is obtained in each, necessitating,
however, more attention than the simpler form. The main features
of this apparatus are: its automaticity, its noiseless action making
it suitable for the class room or laboratory table, its simplicity and
inexpensiveness.
Detroit. Mich.

THE FLYING-FISH PROBLEM'

LIEUT.-COLONEL C. D. DURNFORD

In a paper published in the Annals and Magazine of Natural
History for January, 1906, the impossibiUty, from a mechanical
point of view, of a flying-fish accomplishing sailing flight was
shown. The argument was based upon the fact that as a flying
animal the flying-fish is equipped with wings of a fractional sail-
ing value compared with those of a sailing bird. Also that if the
wings were many times larger, so as to bring the fish on an equality
with the bird in this respect, it could only sail with the bird's
limitations as regards direction of the wind, and with the bird's
frequent assistance from rowing flight. Also that if the figures
(which can be easily verified or, if wrong, refuted) are correctly
given in the article, the accepted aeroplane flight is miraculous,
unless a new law of Nature be discovered.

It is, then, perhaps advisable, if the present curious condition
of the question is to be understood, to examine how it has come
about.

The flying-fish problem is a very odd one in many ways, of which
the most striking is the unexplained power therein of the negative
to quench the positive. Throughout we find the aeroplanist's
"I cannot see the wing-movement" smothering a fairly equal
bulk of "I can, and have, and do see it."

Let us create a parallel instance, for a real parallel does not
perhaps exist : — Many people can see bullets in their flight.
Many others with equally good, or even better, sight cannot pick
up the flying bullets. Now if those who fail to see them said, and
if all books and papers on shooting supported them in so saying,

'This article was intended to appear simultaneously in the American
Naturalist and in the Annals and Magazine of Natural History^ut delays

publication here may be pardoned. — Editor.

65

66

THE AMERICAN NATURALIST [Vol. XLl

cannot see the bullets, therefore you, and all those who do see
them, do not see them," we should have a parallel to the current
odd mode of conducting the flying-fish problem.

It is in consequence of this supremacy of the negative that the
flying-fish problem has earned for itself the name of "eternal,"
for as soon as one new witness can see the flight, either another
new one fails to do so, or a reference is made to some ol)Scrver who
has formerly so failed; and this is equally satisfactory, for, in the
problem, even an old " I did not" is better than a new "I do."

It might naturally be supposefl that there nnist be an ovci-wliclm-
ing backing of probability, both mechanical an<l natiual, lo tlio
negative evidence in order to justify such dogucd (Uiiial lo the
affirmative of its common value. So far, h()\ve\er, from this
being the case, it is a second odd fact that l)ut one seemingly practi'
cal effort at |)roof has been made, and with this one exception
aeroplane flight rests wholly upon the flat negative.

system! in ' Annual Report of the Smithsonian ' I'ustitutiim,'
1904, p. ills, bv Dr. Tlu-o.lore (iill, an emphatic aeroplauist : —
"Mobiu. ,1S7,S. iss.H eontc-u.le.l that ' Mviuu-(i>h arr inrapahlr
of jl;rnnj hlu- italic, are hi.], f.>r the simple reason that the nu.seles
of \Uv pectoral fins are not lai-v en.-ti-h to b.'ar the weight of their

imuic'liateb i^-.,f. Whii.nau, a lii^h authoritv, <lenii>s i,. ac<-tu-acv.
In the .a.ne ariich- ^^v fin.I that this Matetuent is \ i^.,rou.siv
objected to bv (\ (). Whitman >lss()., uh,, nruv.l, -AJmilting
that in form, si/e, l.-noth, and structure the pe. loral fiirs of K.ro-
avfu.s are Ics. well adaj.ted to (light than th<- uings of most birds,
there is siiH ample room to beliexe, on anat.unical and pliN si„|, .gical

'I'hi. is a plain statemeiit moderately word.-d by a .list ingt^ishcd

Lsuetvd, as tho,.::'h In <•,m^incin^• armitneiu. U the ol.l irritating

to wliether flying-fish fly or d., not fly, and entirely ignoring the
new muscle as|)ect opened by Whitman.

No. 482] THE FLYING-FISH PROBLEM 67

Among the distinguislied naturalists tlnis rcfcrmi to in su})|)()t(
of Mobiuh's theory, Prof. Moseley, l.rinu ..f ili< •( li.dK ni^. r'
Exj)e(lition, and Mr. Boulen^rer are promincni ly nicnlidiir.l. I>,nt
Moseley, who cannot see the K.rnrnlii.s fla])i.ni<:'. can see ili,- Dac-
tylopterids doing so (p. 512): ilic [)()>sil.iliiy of u Inch a<t ixlcnicd
by Mobius from personal ohscrvalion as sironi^ly a> m ilic ca>c of
Exocootus! Whilst Boulenger merely (piolcd widid ot dtlicrs,
he himself retained, then as now, as he intornis nic, an open mind
U{)on the cpiestion.

It is surprising how largely this "general vcniici"" is ndhienced
by tlie researches of M(-)l)ins, the very Pn.fcssi.r whose solitary
so-called proof is (|nestione.j l.y WInltnan; so we will examine
more closely what he say> al)ont ihe muscles. '1 he (|uotanon is
continued from '"aloft ui the air,'"' al)ov(\

'"The pectoral iiuiscles ol birds (lepres^niii' tlieu' \vniu'> \\eii:li,
on an average, one sixth ot the total uciu'ht of the body, the [.ecioral
muscles of bats owe thirteenth, the nuiscles of the pecioial fni- of
fiying-fish only one ihu-t\ -second.'"

*If this proves anythin- whi.'li t.. the |.,n'iH.>e ii does n..t —
it may jm.ve that, a> flvin--fi>li have >..niewliat les> than half the
comparative muscl(> of bats, and i acconluii:' lo aei'oplanisis i eaiuiot,
for this reason, fh.theietore bat., whid. hav.^ M.nieuhat le-^ than
half the comparativt^ nurscle of birds, cannot tly.

Or. the other way about: - Hir.ls can fly. Kats. havuio- rather
less than half \hv coinpaialiNc nniMh- of bird^, <an iK . thuefoie

Those are reasonable <ledu( tious. but iherefoie tl\ u,-hsh ( an-
not fly" is an unreasonable uuc. ^

flying-fi.sh, if they can fly, cannot be cxpivted to llv like eitlu-r
hats or birds; ami, I may add, no one thinks or claims th.it tluy
do so fly.

But an even greater claim is made by a.-roplanisis. h is recog-
nized that there are two kuids of bM-.l-tli-hi . •'sailinu and •"low-
ing," the sailing beinu' urea tly the supei-ior form. >ailers ( an always

68

THE AMERICAN NATURALIST [Vol. XLI

to weight ratio. ^ Now flying-fish have a ratio of the lowest class
in comparison with birds (see 'Annals,' Jan. 1906, p. 162); yet
they are credited by aeroplanists with sailing of a higher form than
that of the best-equipped sailing birds — sailing, without even
occasional rowing assistance, at a slow speed, regardless of the
direction of the wind ! Such a feat — one utterly impossible for
an albatross,^ an eagle, a vulture, kings of flight — is given to
this last poor dabbler in the art upon persistently contradicted
negative evidence, two impossible parallels, and the one discredited

I have endeavored in the foregoing to show how observers have
been weighted and clogged by the unique system of handling an
admittedly difficult question — how a very able man, Prof. Mobius,
years ago undertook a research which required a very special knack
of eyesight in the observer. Probably the majority of men are
without this knack, and do not know it. Firmly believing what
I have endeavored to show must have been the false view pre-
sented to his retina, to be a true view, he wrote, with the cleverness
that belonged to him and the dogmatism of the believer, the text
of the faith which has guided and misguided scientists for over a
quarter of a century. His reputation was, and is, deservedly
great — so great that his word was practically law, and it came
about that if other scientists possessed the knack of sight and
differed from him so much the worse for them; they must be either
ignored, or explained away, any or no explanation being sufficient
for such a proper purpose. This is not a hard judgment. Any-
one, who is free from the superstition, on reading an ordinary
aeroplane article will recognize its justice.

1 Harting's formula — which governs this ratio
V weight in grammes,
in birds, is impugned by R. von Lendenfeld in the volume that we have been
quoting from (Ann. Rep. Smith. Inst. 1904, p. 129). The figures of his ex-

they strongly support Harting f^p= = 268, and not 4 03 as given by
Von Lendenfeld as the ratio of the partridge^.

^ Some notes by Prof. Moseley (" Notes by a Naturalist on the 'Challenger,"
p. 571, 1874) upon the small amount of true soaring performed even by the

No. 482] THE FLYING-FISH PROBLEM

Take a quite typical example of the common aeroplane blind-
fold acceptance from writer to writer of palpable impossibilities
as guiding facts. In the article that we have been quoting from
we may note the following (p. 500): "The best estimate has been
that an ordinary flight may extend from 30 to 50 yards in less than
twenty seconds." In order to get working figures we may call
"30 to 50 yards" 40 yards, and "less than twenty seconds" 15
seconds. This gives a rate of 5^ miles an hour!

Note this, you who watch the fish fleeing before a 14-knot

Such statements are the habit of the problem. Just in the
same way is it its recognized habit to quote, unquestioned, as "sail-
ing" parallels to the heavy small-winged fish, the |-oz. large-
winged swallow, and the parachute whose work is falling only;
or, again, to faithfully reproduce over and over again pictures of
impossible air-currents performing feats also impossible; or to
continue to ascribe the frantic efforts at flight of a fish fallen on
deck to natural spasms, although it is not credited with active use

of its wings either in air or sea; and so on. It is the way of the
problem, and no one is to blame.

Perhaps the odd unsuitability of the swallow comparison may
be brought more fully home by a sketch.

The ratio (Harting's formula) of a swallow (house-martin) is
4.2, and its wing-area 120 sq. cm. The flying-fish ratio is 2.6.
If we reduce the swallow to a 2.6 ratio, its wing-area becomes about
47 sq. cm.

This reduction to flying-fish ratio is shown by the shaded parts
of the sketch.

70

THE AMERICAN NATURALIST [Vol. XLI

Could anyone contend that a swallow could sail even in its pres-
ent poor and much-assisted way (for it is far from being a first-
class sailer) if the unshaded parts of the wing-areas were removed ?

Opinion is, however, undoubtedly changing. Many of the old
shibboleths are fast becoming discredited. The great distances
that the fish, under favoring conditions, fly clear of the water ^ —
the fact that they fly in calms as in winds — that they come on
board ships from lee and weather sides indifferently — that they
can and do turn in air ^ — that they often lose and often gain speed,
both from simple causes, on meeting a wave or on tail-dipi)ing —
that they can and do at times gain speed whilst still in air — that
they make for lights deliberately — that tliey rise and fall of set
pur[)Ose while in the air: all these and much more that has been
under the ban are being witnessed and certified to so incessantly
that soon only tlie high-|)riests of aero[)lane will be left contra-
dicting them.

F. ( r. Ailalo (' Natural Hist, of Australia,' Macmillan & Co.,
1S%) wiites- "I ha\e watched the^e beautiful creatines In tlic

hour and in all weathers, but after having closely watched

thousands of them through strong ghisses, I cannot give as ompliatic

72

THE AMERICAN NATURALIST [Vol. XLI

the fish could never arrive at a speed by which a very short aero-
plane flight could be attained even with their low ratio; but I do
say that such is not their common speed, and that in any case their
disregard of wind-direction disproves such flight.

Therefore another way must be looked for, and we are driven
back, perforce, to continuous wing-action, the manner of which
may be here examined as carefully as our information allows.

Premising that the flight varies greatly on different days and
under different conditions, the following is probably a fair descrip-
tion of their methods in an ordinary flight: —

1. The tail-impelled, visibly (to many) wing-assisted jump
from the water to a height where the wings can work freely.

2. The flight continued by an intensely rapid and labored
w^ing-movement — one easily mistaken for stillness, and usually
seen, if at all, as blur.

3. Short periods of slowing down of wing-speed, during which
the wing-movement becomes again visible. (These are the "vibra-
tion" periods, representing to aeroplanists loose wing-trailmg,
or dragging like a flapping flag — an impossibility; and, to Mr.
Adams, periods of wing-assistance — with limitations a possibility.)
These periods often precede a special spurt such as is re(|uired to
lift the fish over an oncoming wave.

4. Either sudden cessation of wing-mo vcukm it and consctiuont
immediate drop into the sea or a short slow down into visibility
(No. 3) previous to such drop.

It is to be noted that this vibration so often seen before the fish
enters the water is one of the many pointers to continuous wing-
movement, for such a time is a proper one for slowing down, but an
absurd one for renewal of wing-effort.

To return to Mr. Adams's paper. He notes, as have others,
the vibration of the wings as being in "an almost horizontal direc-
tion." This horizontal movement, if it exists, as is probable,
may afford, as I hope to show, a looked for key to the fish's action.

According to Pettigrew, it is a necessity of flight, where wing-
beats are in a more or less vertical direction, that the up-beat should
meet with little and the down-beat with much resistance from the
air. This is arranged for in the case of bats, birds, and certain
insects by means of special muscles and ligaments which automat-

No. 482]

73

ically flex the wing for or during the iijHstrokc, and extend it for
or during the down. (Pettigrew, 'Animal Locomotion,' Int.
Science Series, vol. vii. pp. 122, 182, 1!)4, cVc: IV.M. >

Marey ('Animal Mechanism,' p. 2();> c^e. : Int. Science Series,
1893) equally recognizes the necessity for a diminislied winir-area
in the up-stroke, but believes it to be <)l)tained in l)ird> tlnoui^Hi the
natural elasticity of the feathers, which enables tliein to return to
their ordinary position when the resistance of the air in the down-
stroke ceases to raise them.

The flying-fish's wing, as is known, is formed on quite a different
principle from that of a bird or bat. It opens and closes some-
what like a fan. A partial automatic closing of this fan at the foot
of the downward stroke in flight and opening at the top of the
rising stroke would both give the appearance of horizontal vibra-
tion when seen either from above or below, and would turn a some-
what difficult question of the mechanics of the flight into a very
simple one. Indeed we have here flying action on the same general
principle as that shown by Pettigrew and Marey to be necessarily
provided for in the case of bats and })inls. but the working details
of which are different and simpler, as becomes a simj)ler form of
wing.

Perhaps that is the explanation. There must, of course, be
some explanation, and that is not only the natural deduction from
the peculiar formation of the wing, but it also fits everything in.

The known (but indistinct) visibility of tlie larger rays of the
wings at times diu-ing flight points, perliaps. to a comparative
pause with wings full open before hegirning the down -stroke.
Such pause would give the open position, and with it the wing-
tracery prominence.

The form of these lishes' wings poiius to this fan-aciion rather
than to other known horizontal wing-actions of the nature of that
of certain insects — the connnon Hy. for instance : Marey, A-c vif.
pp. 204, 206).

The second quite recent and very important (.bservcr and writer
on this subject is convinced of the Hight-action. lie wrii(>s also
from personal observation, and is as free from proper mechanical
bias as from the improper follow-my-leader habit. One of his
remarks, "It is by no means impossible that flying-fish may soar,

74

THE A M ERIC A N NA T LIRA LIST [Vol. XLI

as eveji [my italics] birds do this." shows his mechanical freedom.
In a paper dated Oct. 2Sth, IDOf), Bri^ '(lalilee,' North Pacific
Ocean, Dr. J. Hohart Egbert, (^irnegie Kxpchtion, writes (' Forest
and Stream/ Jan. 27th, 190()): " Tliouo-li slill denied l)y some
observers, the power of propnlsion throiioh the air l)y means of
its fin-wings is generally accorded tlic flying-fish.' During months
at sea in the tropics the writer has ahnost daily watcluvl the flying-
fishes and studied their flight through the air. . . . The difficul-
ties of assiu-ing- oneself that the flying-fish moves its winu's during
its flight through the air are well under.!.. od, an<l aUo (he fact
lhat the^e diflicultie^ arr uvneralK r(Mn.)\<Ml wiicn opportuniix
is afhmlnl ..f ob.erxing th(^ flight of ccrlain of \hv lar-vr ^pvrw.
mider favorable conditions. That (lyino--fishes use their winus

can hardly be denied, since from the fo'c's'le head of a shij) plying

water almo.st mider the xcssel's bow. Tlils llappiiiu' motion of the

of the appendages .so rapid as lo be ainiosi bcvond human \ isnal
perception."

Quite so. That is thr to-be-expcct.-.l liiuht of an (exceptionally
low-ratio flyer having sp(>cial addc<l nalnral disai.ilities. Before
long it will l)e the accej.tcd one for flyinu-dsli.

More about the Pectoral Mii.srlrs.

Since writing the foregoing I have received a connnunication
from Prof. C. Stewart, F. R. S., Conservator of the Mnseinn of
the Royal College of Surgeons, who kindly gives me permission
to u.se the rcsuhs of a di.s.section ma.h- at the Mu>enni for the pm-
po.se of comparing the pectoral mu>eles of the (lying-fish with
those of a nearly related iK.n-flying (ish.

I quote from the letter of Mr. liurne, who made the dis-^eciion : -

>A little preniutiire. if Xutural Hi.storios and I'hu-yclopiEdiu.s are any indi-

No. 482]

THE FLYING-FISH PROBLEM

75

76

THE AMERICAN NATURALIST [Vol. XLI

ment was the best standard of size to take — better than length,
for instance. As a matter of fact, the fish were very much the
same length; the Exocoetus being rather the longer.

"The drawings, I think, explain themselves. The flying-fish
muscles were, as you see, considerably larger, both in area and in
thickness, than in Hemiramphus, and the same w^as the case with
the muscles on the deep surface of the fin. In their arrangement
they were much the same in both fish and the same as in other
bony fishes (the cod, for instance). The numbers on the surface
of the fins are the points where I took the thickness of the muscles
by plunging a needle into it and measuring the depth to which the
needle entered. You will notice the great length of the muscles
in Exocoetus: a long muscle means a proportionate length of con-
traction.

" . . . . there is a very marked difference in the size of the muscles
of these two fishes ....

" Believe me, yours faithfully,

R. H. BURNE

(Assistant in Museum).

The above tracing seems to give, roughly, about 44 times greater
bulk of muscle to the Exocoetus than to the Hemiramphus. With
this light it will not be out of place to requote and amplify the one
"proof," distinguishing the addition by itahcs: — "The pectoral
muscles of birds depressing their wings weigh on an average one
sixth the total weight of their body, the pectoral muscles of bats one

thirteenth, the muscles of the pectoral fins of flying-fish one

thirty-second," and the mutcles of a nearhj related non-flying fish
only one hundred and fifty-fourth.

As before, it does not prove that bats or flying-fish flap or do
not flap their wings, but it gives a different and, I hope, a proper
aspect to the figures which have done duty — of a kind — for so
many years.

CONTRIBUTIONS TO MUSEUM TECHNIQUE
I. CATALOGUING MUSEUM SPECIMENS'

L. B. WALTON

An essential feature in connection with a museum, is the main-
tenance of a careful record or history of the objects forming the
various collections, since a specimen deficient in data referring
to the locality, date and conditions under which it was obtained,
is practically valueless in comparison with one correctly catalogued.^

The inadequacy of the systems commonly employed, even in
prominent museums of America and Europe,' by which rarely
more than a number, name, and locality of uncertain value, are
more or less heterogeneously arranged in cumbersome and often
inaccessible volumes,* is apparent to any one who has attempted
to locate a desired speQmen, or when fortunate enough to ascer-
tain the location, to obtain concise information concerning it.
This condition of affairs is particularly obvious to the systematist
wishing to study the material belonging to a certain group or
from a definite area in a museum, for he may indeed be considered

2 1 have morely given t\x})ressi()n to the principle laid down by Goode in
his adniiialih' ]k\\h'v on nmsfuin a(hniiiist ration (Annual Report of the Mu-
seums AsMiciaiinti, IV'.-), al-o icpul .li-hr,l in the Annual Report of the Sniith-

77

78

THE AMERICAN NATURALIST [Vol. XLI

a fortunate individual if, after the loss of much time examining
the collections on exhibition and in storage, both catalogued and
uncatalogued, and in consulting the various volumes in which the
data are supposed to be kept, he obtains the data which he wishes.^

Consequently the following suggestions in respect to the cata-
loguing (often spoken of as 'registering' or 'recording") of
specimens have been brought together primarily with a view
toward facilitating the maintenance of such records in museums
of Natural History, although it is hoped that they may prove of
practical advantage in connection with other institutions of a sim-
ilar nature. The paper was outlined and partially written while
engaged in the rearrangement of certain collections in the American
Museum of Natural History, New York, during the summer of
1901. The completion, however, although a brief review was pub-
lished in the Ohio Naturalist for 1904, has been delayed in order
to make further inquiries concerning the systems of cataloguing
used in various museums, as well as for the purpose of profiting by a
more extended practical application of the method. This latter
result has been accompHshed in the cataloguing of specimens dur-
ing the last three years for a foundation of a small museum at
Kenyon College. It may be noted that very few changes from the
plan first proposed have been rendered necessary.

The literature relating to the subject of cataloguing museum
specimens is chiefly conspicuous by its absence, notwithstanding
the mass of information in regard to museums and museum admin-
istration which has been brought together in the Museum Journal
and a few other periodicals devoted to the interests of such insti-
tutions, and. in the papers by Meyer :00-03, Gratacap :02-03,

' In a vigorous article by Bather (How may Museums best retard the
Advance of Science, Annual Report of the Museums Association, p. 90-105,
1896) some of the difficulties of locating museum specimens are described as

certain specimeL^that had been described by Mr. de LorioL The various
curators whom I met at the Museum assisted me very willingly throughout
three days searching for these specimens, but they could not be found, and I

my friend, Professor Steinmann, who suggested that possibly the specimens

able expense and inconvenience I therefore returned to Strassburg, and sure
enough, there were the specimens carefully obscured."

No. 482] CATALOGUINC MI SFA M SI' l-j ■ I M i:\ S 79

Murray :04, etc. Meyer (p. 419) briefly outlines tlu' hiciIkmI used
in the Field Columbian Museum, while Murray (v. 1. p. L't, 1 1 some-
what naively suggests that "As a rule it is of ini]>oriaii( (' iliai the
exact locality from which each specimen has been ()l)taine(l
should be recorded .... This does not apply to archaeological
objects alone. . . .The date of finding or acquisition is often like-
wise of importance."

There are nevertheless a few papers which should be iiientiotied.

Hoyle, '91, described the cataloguing of specimens in the Man-
chester Museum and formulated a system of 'registraiioti ' in book
form, and of 'cataloguing' through the use of canN. His rei,ns-
tration catalogue corresponded to that designated in tlie succeed-
ing pages as The Department Catalogue. It consisted of four-
teen volumes bearing reference letters A-(^. be^imiinir \vith A-
Mammals, B-Aves, etc., and ending witli XMineraloi:y. and O-
Anthropology. Each volume contained space for speci-
mens and was ruled in perpendicular colunui> m) that sj)ace for
data concerning 'date,' 'name,' 'locality.' and 'remarks,' was
afforded. When a specimen arrived ar tlie nm^'um, the first
vacant number in the volume correspondinu' to tlie ^jroup to which
the specimen belonged, was affixed to it and the data concerning
it noted in the appropriate colmnn. After the specimen was thus
'registered' {i. e., our DepartmiMit ('atalouuei it was farther cata-
logued in what Hoyle describc^l a-^ the "('nrator> Catalogue"
{i. €., our Reference Catalogtie) by meairs of wliich an official
record of the contents of the museum arratiired accordin*; to a
natural classification, was maintaine(k This is verv similar to
that which I have terme.l The Referetuv Catalo-ue. It con-

80

THE AMERICAN NATURALIST

[Vol. XLI

representing systematic divisions, geographical distribution, type
specimens, etc., is at once manifest. Furthermore no space is
given for noting the authority for identification, date collected,
etc., name of collector, etc., for all of which data provision should
be made.

The "Curators Catalogue" may be criticized on this same
basis. Moreover in a catalogue, the chief purpose of which is
that of a reference or finding catalogue, there seems every reason
for arranging the cards in alphabetical order in preference to
classifying on a systematic basis. Hoyle, himself, in noting some
objections to the decimal system proposed by Petrie in Nature,
mentions the fact that "no speciaUst is ever satisfied with any other
specialist's work." Furthermore unless arranged according to
the alphabet as suggested under the Reference Catalogue, it would
be of no value to the public. The cards adopted should naturally
be of a standard size since odd sizes cannot be perfectly cut by
reason of the expensive machinery used. Ordinary 'guide cards'
would be much better than the 'genus' and 'family cards.'

Dorsey, '99, reviewed the method of cataloguing used in the
Field Columbian Museum of Chicago. As suggested in a preced-
ing footnote, this appears to be more or less of an heterogeneous
arrangement of cards, books, and manilla envelopes, which could
be much simplified.

Walton, :04, pubhshed a brief outline of the prese^it paper noting
the division into (a) The Accession Cataloune, il)) The I)e|)art-
ment Catalogue, and (c) The Reference Catal()<;ue, as well as
suggesting the general scope and methods of filino- the canls
employed in each.

Wray, :()o, called attention to the adoption of the card system
in the Perak ^luseum of the Federated Malay States, a result

procured,' 'How obtained,' ' Presi^tted by,' ' P)e(|neathe(l by,'
'Purchased from,' and 'Collected by.' Dnplicate cards were
made out, one set being filed numerically as a 'Register,' the
other according to the arrangement of the specimens in the museum

e when
Where

No. 482] CATALOGVIM: MI SI-IM Si'l-ri .][ i:\S 81

spomls to that which I have (h'sionatiM i the ' 1 ii incut Cala-
logue,' lacking the method of cross iiKh'xiiii: liy ( Npaii mciits,
marginal tabs, and colored cards (when <h'^ii;il.le The ;i ,")
inch cards used by Wray are too small, while ilie wiitin-- of two
sets for each specimen nearly doubles the clerical work involved
in the use of an Accession, I )e|)ai-tnieni . and Kefcrcnce catalogue
as noted in the following ])ages, since by the latter method a large
number of specimens are usually transcrihed on a sintjle dc})art-
ment and reference card. Space for certain valnahle data is like-
wise omitted by Wray. something unavoidable however with the
small card.

From the first it seemed evident that the card catalogue arranged
in unit cabinet sections would furnish the most satisfactory solu-
tion of the problem. The value of such a system had long ago
been recognized in connection with library and general l)usiness
methods, where it rapidly displaced lh(> bulky vohimes formerly

tages resulting from the tis(> (.f the card system are obvious since
(1) the retjtiired <lala are prcM-iUcd in a c<.inpact and ea>ily acces-
sible form; .2. the capacity i> unlimited. um'Icss ,vc.mb caii be

4 X (), and s, inches, and although other vizc-, could l)e made
and UM.,I. it is well t.> adopt one of thv.v. inasnuich as the regtilar

machinery Uscd is particniarly adapted for the three sizes. ' 'Vhv

82

THE AMERICAN NATURALIST [Vol. XLI

inch cards are too large and unwieldy. The 4X6 inch card,
however, is of sufficient size to contain all necessary data, without
being cumbersome in manipulation.

Card cabinets to contain the catalogues may be obtained in
various sizes, but by the adoption of the * unit ' card index section
containing six drawers adopted for the 4X6 in. card, future
units may be added as occasion demands, and the cabinet is thus
always complete.

Following a chronological order, the data which should be
rendered accessible in an adequately catalogued collection, can be
separated into three divisions. These are: (A) The Accession
Catalogue, containing a general record of all material received
by the museum. (B) The Department Catalogue, giving a com-
plete history of each specimen or group of specimens, (a single
species, acquired by each department. (C) The Reference Cata-
logue, having the names of all specimens belonging to each depart-
ment, arranged alphabetically so that the final disposition of any
desired specimen can at once be ascertained.

Of these, the Accession and Department catalogues are essen-
tial from a business as well as a scientific standpoint, while the
Reference catalogue, altliouuli not a necessary requisite, will be
found advantageous as a rt t'crcncc index to the specimens, particu-
larly in the larger mnscnnis. \\ iili tii<' exception of the one per-
taining to accessions, which should l)e in charge of the director of
the museum, each catalogue should be controlled by the head of
the particular department with which it deals.

While the records considered necessary vary more or less in

cerned, they can in general l)e redticed to the following tabular
form, covering the data which may l)e required in ]Museiuns of
Natural History.

A. Accession Catalogue (arranged numerically).

1. Accession number.

2. Date received.

3. Description.

4. How obtained.

a. Purchase (cost ).

b. Gift.

c. Exchange.

No. 482] CATALOGUING MUSEUM SPECIMENS 83

d. In trust.

e. Museum collectors.

5. From whom received.

6. Address.

7. Transportation number.

8. Collector.

9. Locality where collected.

10. Date when collected (approximate).

11. Correspondence filed under.

12. Remarks.

13. Date of entr}-.

B. Department Catalogue (arranged numerically)

1. Department number.

2. Accession number.

3. Original number.

4. Number of specimens.

5. Sex.

6. Stage of growth.

7. Scientific name.

8. Authority for identification.

9. Date of identification.

10. Ix)cality where collected.

11. Name of collector.

12. Correspondence.

13. Date when collected.

14. Character of specimens.

15. Remarks.

16. Date of entry.

C. Reference Catalogue (arranged alphabetically)

1. Name of specimen (common name and scientific
name, — genus, species, — listed on separate cards).

2. Department number.

3. Character of specimen.

4. Location.

a. On exhibition. Case No.

b. In storage. Drawer No.

5. Number of specimens.

The following suggestions have been found valuable in regard
to the data and their arrangement on the cards.

84

THE AMERICAN NATURALIST [Vol. XLI

A. ACCESSION CATALOGUE.

In this catalogue, all material ^ received or collected at a partic-
ular time from a particular source, (an accession), is placed under
a single accession number. Thus the catalogue will contain a
record of each group of specimens coming into the possession of
the different departments in the museums, and by means of a
series of cross references, consisting of tabs arranged as indicated

accompanymg

No. 482] CATALOGUING MUSEI M SI'EC I M ES S

85

by each department, whether they have been obtained by pur-
chase, gift, exchange, through museum collectors, or in trust,
and if by purchase, their cost, as well as the particular fund made
use of in connection with their acquisition.

The disposition of each item on the card should correspond to
its relative importance. In the following diagram (Fig. 1) a
convenient arrangement is suggested.

Classification by Departments. — A classification by departments
can be conveniently maintained by having tabs arranged on the
cards in as many different positions as there are departments.
Thus with ] incli tabs as in Fig. 1, eight departments may be

Accession number. — This should occupy a prominent place, pref-
er red ink.^ The iHiinbcrs should hr MMMallv arranovd in acconhmce
with the date of arrival of the accession, and at inKM'vals of one
hundred cards, a nunihcrcd niiidc card of a pariicular color t. (j.
dark blue) may he iirs,-rie.l. Where n.. previous catalo-ne of

commence with a innnher snfliciently iarp- (c. 7. lOOi n. allow
the eventual catalo.unini:- of former collections which ha\c come

logical as possil.l(^

numeral for the day of the month and a lioman mnneral for the
month, followed by the _v«>ar ir. 7., (l-IX is'.ls - September (i,

gin. At the end of every _vt>ar, a card can he in>erie.l, on the tab

THE AMERICAN NATURALIST [Vol. XLI

rial obtained by the museum during any particular period is at all
times readily ascertained.

Description of material. — The general nature of the consign-
ment should be indicated, (e. g. archeological material, mammal
skeletons, fishes) as well as the manner in which it is packed (num-
ber of packages, boxes, etc.). In this connection a record should
also be kept as to whether the accession is received as a ' purchase,'
'exchange,' 'gift,' 'in trust,' or through 'museum collectors.'
This can be readily accompUshed by having the above words
written on the card and placing a cross in the proper space at the
time of cataloguing. When procured by purchase, the price
should also be indicated.

From whom received. — The name and permanent address of
the person sending the specimens, is to be noted here.

Transportation number. — It is often convenient to have a rec-
ord of the number or numbers placed upon the consignment by
the transportation companies, particularly in the event of break-
age or loss of any of the contents of a package or box.

Name of collector. — Many collections are deficient in labels
bearing accurate information, consequently it is advisable to ascer-
tain the names of individuals concerned in collecting the speci-
mens, so that if desirable, further data may be obtained. The
address of the collector is to be noted, provided it differs from that
of the locality where the collection was made.

General locality. — When the collection is a small one from a
restricted locality, this can be readily indicated. If, however,
a large amount of material is represented, the principal region or
regions should be given.

Date when collected. — It is necessary to indicate merely the
approximate time.

Correspondence. — In order to readily refer to correspondence,
invoices, bills, and other memoranda relating to the accession, it
is well to indicate the initial name or number, together with the
year, under which they are filed.^

Remarks. — Under this heading can be noted the condition of

» Madeley :04 presents an elaborate arrangement for the classification of
oflSce papers in Musevnns based upon a provisional decimal system. It seems
unfortunate thut the standard decimal system (Dewey) was not utilized.

No. 482] CATALOGUING MUSEUM SPECIMENS 87

the specimens whether or not the collection contains any forms of
particular value (types, cotypes, etc.), as well as other general
information.

General suggestions. — In order to record small collections,
which may come directly to a department, blank cards may be
provided for those in charge, and upon the arrival of such an
accession, these should be immediately filled out and handed to
the person keeping the Accession Catalogue. Blank cards to be
similarly filled out and returned, can be sent to a person from whom
an accession deficient in data is received. The system of cross
references can be arranged to meet any demand. The method
employed as noted above, appears adequate for ordinary purposes.
Thus the name of each department is placed on a tab assigned to
a particular position, and when the cards are filed, the accessions
of a department will be indicated by the corresponding row of
tabs. A further subdivision which may be applied to each depart-
ment is in the use of colored cards. If for example the department
of anthropology, possesses three separate appropriations upon which
to draw for as many purposes, e. g. : (a) Explorations on the North
Pacific Coast, (b) The purchase of Michigan Antiquities, and
(c) Collections illustrating the life of the Aztecs; all accessions in
Anthropology of (o) obtained by purchase, or at the expense of
the museum from the one fund, can be placed on salmon colored
cards, while similarly all accessions of (6) and (c) obtained from
the corresponding appropriations can be placed on buff and blue
cards, respectively. Thus at any time the general condition of
the various funds of the department can be readily ascertained.
Geographical Distribution {e. g. nearctic, neotropical, etc. may be
represented in a similar manner.

Placing numerical guide cards at intervals of every hundred
cards, will greatly facilitate finding any desired accession number.
In a catalogue where the width of the tabs makes it possible to
have an area at the riglit fioiii wliich no tabs project, it is con-
venient to place the numerical lab as in Fi^^ 1.

Inasmuch as tlie majority of accessions cover a quantity of
specimens, such a catalouuc as tlie one described can be easily
maintained, and the advaniai^es which result through always
having correctly classified data accessible are an important item
in the making up of reports.

THE AMERICAN NATURALIST [\'ot.. XL!

B. DEPARTMENT CATALOGUE.

The department catalogue has the cards arranged numerically
in chronological order and should contain concise information
concerning each specimen, or group of specimens belonging to
the same species which were obtained at a definite time and place.
In the smaller museums the material may be grouped under depart-
ments of Zoology, Botany, Paljeontology, etc. as represented by

the Accession Catalogue (Fig. 1) each with its separate depart-
ment catalogue. In the larger museums, liowencr, it will often
be advisable for each department to have several sub-depart-
ments or group catalogues having the rank of (lej)artineiits. For
example the department of Zoology may maintain ( ataio-iies of
Vertebrate and Invertebrate Zoology, or of Pathological prepara-
tions, Neurological specimens, etc., or on a systematic ha-^is it
may have a catalogue for each phylum or branch of the animal
and plant kingdoms. The cross-reference classification by means.

No. 482] CATALOGUING MUSEUM SPECIMENS 89

of tah.s. liowcver. as rt'])re.sentecl in the department catalogue
(Fig. 2) w ill ii>iiallv !)(' siiflicient in the smaller museums.

Here I he ai i aiiueiiieiit of data will meet the needs of the average
department. Xear the middle of the upper margin of the card
should be placed the name of the particular dej)artni( tii to which
it refers, together with the name of the in.stitution. If the depart-
also appear, r. 7. Zoology Department ( alalouue. South Afri.an
Musenrn, Hir.ls.

Systematic cross reference classification by tabs. The elassili-
cation adopted will (ie[)end on the natm-e of the catalogue. If
half-inch tal)s are used on a G inch card twelve divisions are po.ssible
which in tlie zoological department cards above consi.st of 1.
Mammals, 2. Birds, Rej)tiles, 4. Amphibians, 5. Fishes, etc.
6. Tunicates, 7. lv-hitio(l«>rins. S. Articulates, 0. Mollusca, 10.
Vermes, 11. Coel(>nterates and Sponges, and ll>. Protozoa. For
certain reasons an arrangcni.Mit in the reverse order w(»uld be
more logical. In a l.oiani* a,l catalo-uc one could chooM' between
theol<lercla..ilicaiion<.f Ki.-hlcr. lss;;,uhcrca .omcwhai arbitrary
grouping gives n> the 1. .Vlgae, etc. 1'. Lichen^.;;. Hryojihytcs.

one of Knuler.' P.Kl l, w ith thirteen grouj)s and ;r)-4() classes. The
classification adopted in the other department catalogues, PalcTon-
tologv. Antliroj)olo!jv. etc., wdl m a similar manner represent to
a mo're or lc>s (>\lent the personal equation of the curator under
whose supervision they are maintained.

Geographical cross reference classification by colors. (ieogra|)h-

a particular color represent dcnnite area-. Sucli an arratige-

90

THE AMERICAN NATURALIST

[Vol. XLI

the state excluding the county, catalogued on buff colored cards
while other specimens from localities outside of the state would
be catalogued on salmon ^ colored cards. In the larger museums
where collections are made up of specimens from different parts
of the world, certain colors can be used to represent various regions,
(nearctic, neotropical, patearctic, etc.). Types, cotypes, etc.
could be catalogued on cards having the right half red, the left
half in accordance with the color representing the particular geo-
graphical distribution.

Department number. — A single department number will cover
a series of specimens of the same species, which have been obtained
at the same time in a particular locality. This method is more
satisfactory than assigning a number to each individual specimen
inasmuch as time would be lost by such a method and no particu-
lar benefits result. Should the occasion arise at a later period,
a separate number may be assigned to any specimen.

Accession number. — This should be indicated on the card, in
order that general information regarding the collection may be
obtained at any time. The accession number and department
number may be indicated in connection with the specimens as a
fraction {e. g. |§g ) whose numerator represents the accession num-
ber, and denominator the department number, or as a decimal
(294.896), or the accession number may be entirely omitted from
the specimens, since a reference to the department card will fur-
nish it when desired.

Original niunber.— This is the number which a specimen may
possess on its arrival. Often times it will be the field number
placed on it at the time when it was collected or it may refer to a
number assigned in a previous collection.

Number of Specimens. — This is essential in order to know the
amount of material in any collection. When duplicates are used
for exchange, the former number should be crossed out and the
new one substituted, while, at the same time, a reference number
referring to the exchange may be added.

Sex.— The sex can be designate.l 1)V \\w conviMitional si<,nis,
C?, 9, 0, representing, male, female, and luM-iiia))lin)(litf t'oniis.

No. 482] CATALOGUING MUSEUM SPECIMENS

91

Growth. — Embryo, young, adult. Measurements, weight, etc.

Scientific name.— In systematic work of this nature the generic
followed by the specific name must be used.

Authority for identification. — This is an important item which
is too -often omitted from the average museum catalogue. If a
specialist subsequently verifies a name previously given, this
should also be noted. In case the name is found incorrect a new
card is to be written.

Date of identification. — It is well to have this information avail-
able.

Locality where collected. — Too much care cannot be exercised
in accurately indicating the locality from which specimens are
obtained. It is safe to say that every museum has among its col-
lections material which would be of the utmost value, provide the
locality, even within a few hundred miles, could alone be ascer-
tained. Unfortunately in most cases of this kind, it is the collector
who is at fault. The cataloguer must rely on his data.

Name of collector.— Inasmuch as the ' personal equation ' must
be taken into consideration, the name of the collector is indispens-
able. Furthermore it often furnishes a clue to the history of a
specimen when all other means have failed.

Correspondence. — Letters, etc., pertaining to the particular
specimens can be indicated as suggested in the accession cata-
logue.

Date when collected. — This can be indicated as in the accession
catalogue.

Character of specimen.— The nature of a specimen, whether a
skeleton, an anatomical preparation, a mounted skin, etc., should
be given. If preserved in a special manner it is well to indicate
the formula, e. g. 5% formalin; 70% alcohol; killed and hard-
ened in chromosmic 3 hours, preserved in 95% alcohol, etc.
Explicit notes here will in the end well repay tlie time s[)ent in
making them. The back of the card will atrord a'lditional >\rM-v,
if needed.

Remarks. — This space is only to be filled out xvheii there is
something of particular importance to be noted concerning the
specimen, and of a nature which cannot be covered under the
other records.

92

THE AMERICAN NATURALIST [Vol. XLI

General suggestions. — The data, as well as their arrangement
on the cards, are naturally subject to various changes, in order to
conform to the requirements in different museums. It is well
to have a blank space for each item of information concerning
the specimen, although often " unnecessary, or even inadvisable,
provided there is reason to doubt its accuracy, to fill it out. As in
the Accession Catalogue numerical guides should be placed at
intervals of one hundred cards, while 'side locking cards' are
recommended.

C. REFERENCE CATAEOGUE.

The Reference Catalogue may with equal propriety be termed
a finding list, since its purpose is that of indicating the location of

NAME Cr4V>tobrancW6 altegkeaveusis i)aa<i

each specimen which belongs to the j)iirti( iilar (lc])iirtin(^iit in the
museum. The cards are arrange<l in al})liahetical ohUm-, both
the scientific name (generic followed by specific name in the case
of biological specimens), and the common name having a place
on separate cards, the latter, however, referring to the former,

No. 482] CATALOGUING MUSEUM SPECIMENS 93

(e. g. opossum, see Didelphys. Furthermore, the reference card
indicates the number of specimens of each species on exhibition,
or in storage, giving the number of the case or storage drawer in
which they are to be found.

A single card will usually contain the data concerning all mate-
rial belonging to a particular species, consequently the time involved
in maintaining a Reference Catalogue is an unimportant item, the
data (except location of specimens) being readily obtainable at
any time from the Department Catalogue.

The Reference Catalogue should be located in the principal
room containing the collections to which it refers, where it will
be readily accessible to each of the three classes of people for which
a museum primarily exists: (a) the specialist, (b) the amateur,
and (c) the general pubHc'

One method for arranging the data for a reference catalogue,
is shown below (Fig. 3).

Systematic Cross Reference Classification by means of Tabs.—
An excellent method which meets the usual rofniireinents, is that
of having the tabs arranged as in the l)('j);iriiii(Mit Catalogue.
Geographical cross reference by colors cannot l)c us(nl inasuiucli
as one card will often contain specimens from widely separated

Name of specimens. — Both the scientific name and the common
name should be given, the former on the card containing the data,
the latter on a separate card referring to the generic or specific
name of the particular species, {e. g. Brook Trout, see SalvcliJius
fontinalis, Pickerel, see E.sox, various species). By placing the
common name on cards having a particular color they may be
readily distinguished.

Department numbers. — Inasmuch as the department numbers
will be placed on all material, this will serve to establish the identity
of the specimen sought, and in ease further data is reipiired, the
corresponding number in the Dcpartincni Caialouiic can l)e con-
sulted.

Character of Specimens. — In alcohol, niounicd, skt'lcton. skin,

'See Bather, F. 1904. The Functions of a Museum; Ht-Survcy. Fop.
Sci. Mo., V. 64, p. 210-218.

94

THE AMERICAN NATURALIST [Vol. XL!

Exhibition, Storage, etc. — The location of a specimen is indi-
cated by the particular column under which it is placed. If on
exhibition, the number or letter of the case ^ will be given. Al-
coves or galleries may be designated by letters. If on storage,
the location will be similarly designated.^

Total number of specimens. — These columns will indicate the
total number of specimens of a given species ^ belonging to the
museum. If customary for the institution to make many exchanges
a balance column may be added, which will show the material
on hand as well as that exchanged.

The necessary steps incident to the cataloguing of a collection
which has been received may now be outlined as follows.

a. Catalogued as an Accession.

b. Placed in charge of a department.

c. Catalogued in a Department (^italooue and given a
department number.

d. Identified and labelled. I1)is data then added to the
department card.

e. Placed on exhibition or in storage.

f. Reference Catalogue filled out from data on department

The first three items should be attended to at once. A consitl-
erable interval will often elapse however before final desposition
of the specimen is made.

It would seem that only two general objections can be urged
against any system similar to the one propoM-d, namely; ( 1 ) The
plea that too much time will be occupied in ilie preparation of such
a catalogue, and (2) a certain inherent condition which precludes
the adoption of new ideas, ""['he only answer that need be given

No. 482] CATALOGUING MUSEUM SPECIMENS

95

to the former is that the space occupied by a specimen uiiworthv
of being j)roperly recorded, is more vahiable than the specimen
itself, while to the latter no reply is needed.

It is minecessary and often inadvisable to at once rcdiict' loinicr
catalogues to a card system. Incoming matori;il (mii \)v cai;!-
logued on the cards, and as the opportunity allows, ftoni liu^
previous records can be transferred to cards.

Conservatism * is a valuable factor in connection with all s(;ien-
tific work. It has its limitations however, and in order to make
definite progress in any direction, old methods must give place to
new ones — the fittest w^ill survive.

Ken VON COLLKGE, (iAMUIER, Ouio. Dcc. 1, 1900

^ Meyer, (:()() 01) in his (ixcellent revi(nv- of the niuseunis of the eastern

niuseurtjs in respect to the installation of the collections. In reply to this
criticism however it might well be suggested that to a certain extent at least
this lack of uniformity is an iiuHcatioii of healthy activity. It is not considered
necessary in this .•ou.ifry to rling to traditional ideas %xhich are too often

of dealing willi well known i)rol)lenis arc sought and evolved and if their

presidential address before the members of Sijrina \i ( I )cc. .{1 . H)():{ i cxprcsst-d
similar ideas regarding this tendency which he li:ul noted. •lii rniiur. m
Geniiany, even in England, the tradition of great ii.iines, the nistonis ot j;,-,.;,!

96

THE AMERICAN NATURALIST [\ ol. XLI

BIBLIOGRAPHY

Dorset, G. A.

'99. (The Cataloguing of Museum Specimens), American Anthropolo-
gist, n. s., 1, p. 473.
Meyer, A. B.

: 00-01. Ueber Museen des Ostens der vereinigten Staaten von Amerika.
R. Friedlander und Sohn, Berlin. (Also see translation in Report
of the U. S. National Museum for 1903, p. 311-808 with 40 plates).
HOYLE, W. E.

'91. The Registration and Cataloguing of Museum Specimens. An-
nual Report of The Museums Association, Cambridge, England.
Madeley, Chas.

: 04. The Classification of Office Papers. Mus. Jour., v. 4, no. 3, p.
73-95.
Murray, David.

:04. Maseums, Their History and their Use. 3 vols., J. MacLehose
and Sons, Glasgow, Scotland.
Walton, L. B.

: 04. The Cataloguing of Museum Collections. Ohio Naturalist,
v. 4, no. 3, p. 62.
Wray, L.

:05. A System for the Registration of the Contents of Museums.
Museums Journal, June, p. 407-412.

SOME SOUTH AMERICAN ROTIFERS

JAMES MURRAY

The undernoted rotifers were obtained from moss kindly sent
to me by Mr. N. D. F. Pearce, of Cambridge, England, in the
early summer of 1906. The moss was sent from British Guiana.
The locality from which it came was unknown, but it was some-
where in the interior.

A portion of the moss was still moist, but most of it had been
dried. The majority of the species were got from the dried moss.

As is usual when dried moss is examined after a lapse of some
time, most of the rotifers found belonged to the order Bdelloida.
Of this order 13 species were distinguished; 11 of the species were
already known, most of them being common and widely distributed
species. One, Callidina perforata,^ was only recently discovered
in India, and a very distinct variety occurred more abundantly
than the type. C. multispiriosa was represented by a variety,
probably of specific value. Two new species are here described.

Four species of the order Ploima were also found, — one Colurus,
two Monostyla, — and one Diglena. I was unable to determine
any of these.

Callidina angusticollis Murray > : O.")). - \'crv abimdaut. All the
examples hcloiii^tMl to thr type, or to a small variety. The Indian

C. perforata Murray. -Tlu' most ahimdam sju-cies in the col-
lection. The type ( Fio-. 1 ) was fairly plentiful, l)ut a variety,
described l)elo\v, was much more so.

C. p. var. amsricana var. iiov. (Figs. 2-3).— Case smaller than
in the type, length Klli ityj)e al)()ut 13() /<). Posterior process

Micr. Soc., 1906.

98

THE AMERICAN NATURALIST

[Vol. XLI

sharply marked off by abrupt constriction, not turned to dorsal
side as in the type, but in line with the axis of the case; — perfora-
tion towards ventral side (dorsal in type). Dorsal plicse of the
case not distinct, but an obscure tesselation or coarse stippling
instead. As in Indian examples, empty cases usually lacked the
ventral wall, as though some enemy had found this part vulnerable.

C. constricta Duj. ('41).— Plentiful.

0. aspera Bryce ('92). — A few examples.

C. habita Bryce ('94). — One example, living.

C. quadricomifera Milne ('85-'86). — One small hyaline example.

Ko. 482] SOUTH AMERICAN ROTIFERS

99

but the arrangement of the spines shows that it belongs t
species. This variety is also found in India and Africa.

(Fig.

-Long ante-
rior spines few, usually 4 on
each side, the 2d and 4th of
these much thicker than the
others. The lateral spine of
the anterior row on the central
segments of the trunk large
and very thick. Skin strongly
stippled or papillose.

Other smaller differences
from the type will be better
understood from the figure.
There was no variation from
this arrangement of spines in
all the examples seen. The
variety is much smaller than
the type. About 6 examples

Specific characters. — Small, 240 n long, hyaline or whitish,
with pale yellow stomach, food not moulded into pellets. Head
small, corona 40 wide, less than collar and about half diameter
of central trunk, discs touching, central process of upper Hp
single, truncate. Length of antenna \ diameter of neck. Jaws
18 /( long, teeth '2 1\ veiy thin. Foot 4-jointed, spurs narrow, taper-
ing, divergent; toes large and long, the two ventral put out and
drawn in. in the u>ual manner, wIhmi making the step, the dorsal
kept alway> extende.l an. I fornn-ng with tin- ^purs a tripod. Dorsal
skin Folds faint, few. lafral deeper. The striking peculiarity is
the tripod, whi.'h i> um'<|ue in the order. ( )therwis<> the animal
comes nearest ('. r/urnhrnj; .lanson. from which it is .li.tinguished
by the smaller head, closer discs, and tnmcate upper hp. Abun-
dant.

100

THE AMERICAN NATURALIST [Vol. XLI

C. speciosa n; sp. (Figs. 8-10)

Specific characters.— Yery small, 163 feeding to 238 it creeping.
Head very small, diameter of corona 26 /(, of prominent collar 38 a.
Food not moulded into pellets. Teeth 2/2. Anal segment with
lateral prominences. Foot 3-jointed, first joint with lateral proc-
esses, spurs small, tapering, divergent. Toes three. Dorsal
longitudinal and ventral transverse skinfolds forming symmetrical
pattern, which is constant. Length of antenna half diameter of
neck.

The most distinctive character is the pattern formed by the

skinfolds. Many species have a similar pattern formed by the
dorsal wrinkles, but no other species has the ventral surface so
ornate. Apart from this character it has no close resemblance

No. 482] SOUTH AMERICAN ROTIFERS

101

to any other species. Those which approach it in general form
and dorsal wrinkling have larger heads with separated discs.

Not abundant, about a dozen examples seen.

Rotifer longirostris (Janson) ('93).— Several examples of the type
were found, but none of the Indian varieties occurred.

Adineta graciUs Janson ('93).— Not plentiful.

A. vaga Davis ('73).— Rare.

LITERATURE

Brycf, D.

'92. On the Macrotrachelous Callidinae. Journ. Quekett Micr. Club,
ser. 2, vol. 5, p. 15.
Bryce, D.

'94. Further Notes on Macrotrachelous Callidinje. Journ. Quekett
Micr. Club, ser. 2, vol. 5, p. 436.
Davis, H.

'73. A New Callidina. Month. Micr. Journ., p. 201.

DUJARDIN, F.

'41. Zoophytes Infusoires, p. 658. Paris.
Janson, O.

'93. Rotatorien-Familie der Philodinaeen, Marburg.
Milne, W.

'85-'86. Defectiveness of the Eye-spot, etc. Proc. Phil. Soc. Glasgow,
vol. 17, p. 134.
Murray, J.

:05. A New Family, etc. Trans. Roy. Soc. Edinb., vol. 41, p. 367.
Thompson, P. G.

'92. Moss-haunting Rotifers. Science Gossip, p. 56.

MERISTIC HOMOLOGIES IN VERTEBRATES

J. S. KINGSLEY

One of tlie most difficult problems in vertebrate morphology
is to explain the serial homologies between the different groups.
In the lower segmented animals these difficulties, although they
exist, are far more simple and are far more easily explained. Thus
no one has any doubt that the tenth or the fifteenth somite of
Homarus is the exact equivalent of the serially homologous somite
of Cancer. Between the larger Arthropodan divisions the task
of comparisons of somites is possibly not so easy yet all attempts
at drawing homologies between, say, a hexapod, an arachnid and a
crustacean, are based upon the assumption of exact serial equiv-
alency. It is true that one author or aiiotlier has at times sug-
gested the possibility of intercalation or t^lision of a somite, but
these have been mere suggestions and liavc usually been discarded
in the discussions.

are forced to assume that the slioul.lcr -inllc and f.'iv liml.cf ttir
frog are the lionioh.oues of those of man. allhou-h their eoiuiee-

taken into account. In the ease of the i)elvie arch the numeri-
cal disparity of the corresponding somitt^s is even ureater, but in
either case the identity of structure of arcli and limb is so great
that doubt of homology is praerieally impossible. How then has
it come about that say the twelfth somite of the Am])hibian is not
homologuc of the twelfth but of more nearly the twentieth of
man?

In Gegenbaur's hypothesis that the oinlhvs are derived from
branchial arches and that thtvse have migrated backwards over

problems of the relations of -inile. f.I bo.lv srp.'ents. The back-
ward nuVration has been arreted at dillerent points in tli^- various

104 THE AMERICAN NATURALIST [Vol. XLI

groups. But this explanation will not suffice for other cases,
hence the probability that it is true for none.

In the frog as in all Ichthyopsida, there are but ten cranial
nerves, while in the mammals there are twelve. There is no
doubt that as far back as the tenth the nerves are exactly homol-
ogous in Amphibia and in the mammals. Relations to brain and
to points of distribution place this beyond question, but what
shall be said of the mammalian eleventh and twelfth? Are both
of these nerves from the post-cranial region which have been
transferred to the skull ? If so, does it not follow that the cranium
in the higher vertebrates is not the exact equivalent of that in the
lower? and that the differences have been brought about by the
transformation of cervical into occipital vertebr{3e. If this, in
turn, be so, are the occipital bones of the frog homologous with
those of the mammal ? Or are the basi-, ex- and supra-occipitals
of the one merely analogous of those of the other? Is Huxley's
argument for the derivation of the mammals from the Amphibia
because of the double occipital condyles in the two groups based
upon analogies rather than on true homologies ? Are the condyles
in Amphibia and Mammals not homologous but rather homo-
plastic formations?

Carrying this matter further back in the body, how are we to
explain that apparent shifting of the pelvis in such a form as
Necturus as described by Bumpus, Parker and others? Are
somites ten, twenty and the like exactly equivalent in the normal
and aberrant forms? And has there been an actual shifting of
the pelvic girdle from one somite to the next in some individual ?
Or has there been an actual intercalation of vertebrte, the one to
which the ilium is attached being constantly the same morpho-
logically if not serially? Or, lastly, have the limbs and their
arches arisen from a continuous fin fold and has every somite
which contributes to that fold the potentiality of limb formation
with all that this implies ?

To take another case. In Amphioxus there are a large number
of gill slits, a ninnber which is doubled during development by the
formation of the 'tongue bar.' Right behind the last gill slit
comes the entrance of the hepatic duct into the alimentary tract,
there thus being no oesophagus nor stomach intervening between

No. 482] M ERISTIC HOMOLOGIES IN VERTEBRATES 105

the pharynx and the Hver. Is this to be explained by saying that
in the vertebrates the posterior gill clefts of Amphioxus have closed
and that the space which they occupy has become converted into
stomach and oesophagus? In other words are these formations
of the vertebrate tube the homologues of a part of the gill region
of the acraniate ?

Then, too, what are to be done with cases of increased numbers
of gill slits; the Notidanids with six or seven, the Californian
Bdellostoma with its variable number, and Amphioxus itself?
This question is wholly apart from that which discusses the rela-
tions between metamerism and branchiomerism.

Numerous other similar questions will readily suggest them-
selves to all. There is no reason for enumerating them here.
The problem is, how are they to be explained. Must we find a
separate explanation for each or can we find some one principle
which will account for all?

This article is to be regarded in the light of a suggestion rather
than a full reply with demonstrations of validity. I have no
proof, other than analogies and the fact that the hypothesis here
presented answers all the demands of the problem, that the expla-
nation here advanced is the true one. It must be tested and the
tests are not easily made.

In the invertebrate segmented animals there is, at the beginning,
no metamerism. It appears later during growth, and in numbers
of forms it is found that the segmenting tissues are protkiced by
budding from groups of cells at the posterior end of the eiiihiyo.
These are most familiar in the annelid teloblast and are sr arccly
less well known in the Insects and Crustacea. Their number
varies between wide limits, but for the present purposes the most
important points concerning them, aside from their budding
capacities, are their position in a more or less plainly marked
transverse band and their situation at the extreme posterior limit
of the growing embryo. Extensive examination of the literature
has not shown similar budding cells in the Cuvierian group of
Articulata in other places than the tip of the growing embryo,
with the exceptions noted below.

It follows then that in these teloblasts and their equivalents are
the full potentialities of the future somites. From them arise

106

THE AMERICAN NATURALIST

[Vol XLI

all the cells which are utilized in every structure which is meta-
merically repeated, the material for the new somite not bein^
budded from any pre-existing somite, but always just in front of
the hinder end of the body.

This applies strictly to all cases which are known to me in the
arthropods as well as to most of the annelids; but in a few of the
latter group modifications occur in the process which have great
interest for us. As is well known in a number of annelids asex-
ual reproduction by transverse division occurs. At one or more
points in the body a new head may develop with the eyes, append-
ages, etc., characteristic of the anterior end of the worm, these
features arising from a somite which in its earlier stages is appar-
ently normal and like its fellows on either side. Then, just in
front of this new head the worm divides and two worms, each
with fewer somites than the original one, are produced and from
this time onward lead an independent existence.

Of these only the anterior worm need now be considered. After
the separation the segment which was just in front of the new head
of course becomes the terminal somite of the new worm. The
worm now increases in length and the new somites are formed by
material cut off from the terminal somite which thus must have
within it the equivalent of the teloblasts of the embryo.

From these facts it seems logically to follow that at least certain
somites in the body have the potentialities of forming material
for additional somites and must contain within tlicni the same
physi(.loui(;d possibilities as the original teloblasts from which
they arise. In other words, in the annelid before the heginning

was located at more than one point in the body, but it was not
exercised until after the asexual reproduction was well advanced.

In the case of the Naides the somites thus ])ro(Iuce(l are all
similar in character but in such instances as ProTnla, where heter-

by the formation of new somites which ditler in kind.

I'he ap{)lication of these facts to the variolr^ types of meristie
variation which occur in annelids n<-ed not be (hsi nss,.,| hc,.,.^ h^t
J think it is apparent that they will in ])art e\|)laiii some of them.
I do not mean to say that they reveal first causes but they do point

No. 482] M ERISTIC HOMOLOGIES IN VERTEBRATES 107

out the mechanism involved and may be used to reduce all to a
common rule.

In the same way the assumption that there are similar buddinji;
zones at various points in the vertebrate body will explain the
various conditions outUned in the statement of the problem. In
the vertebrates there is a continuous addition of new somites at
the posterior end of the body as in the arthropods and annelids,
implying the existence of the equivalents of teloblasts at the pos-
terior end. The assumption of budding zones at other points
will explain the other features noted. Such a zone in the occipital
region will allow us to explain the difference in the number of
cranial nerves in the mammals and in the Ichthyopsida and yet
allow us to accept the homology of the occipital bones throughout
the vertebrate series. The additional nerves are thus to be re-
garded not as transferred from the neck but as new or intercalated
structures. In the same way we may explain the varying number
of vertebrae in the different regions and allow at least one of the
pelvic vertebrae to be regarded as a fixed point and may be relieved
of any assumption of a shifting of the uinlles. It will also (>\|)laiii
many anomalies such as the attacluneiit of the two halves of tin-
pelvis to different vertebra^ and the incrcast'd nuiiiher of limibar
or thoracic vertebra^ in man.

This is to be regarded solely as an hypothesis. So far as I am

fact it is extremely prol>abIe that there is no >wh well .lefined /one
as is found in the band of teloblasts of the crnstacean. It is to be
regarded rather as a series of assiunptions. Uased in part upon
analogies, which, if true, would explain the questions with which
the present note began. The hypothesis is presented as a sug-
gestion to stimulate investigation and criticism upon an interesting
and difficult subject.

ON THE OSTEOLOGY OF THE TUBINARES.

II. W. SHUFELDT.
I. Historical.

Few of the groups of Birds have a more interesting literature
than this Suborder.

As early as 1827 M. L'Herminier placed the Tubinares together
in a family of birds (28th) and classified them upon the characters
of their sterna, assigning them to three sections; (1) the smaller
Petrels in which the xiphoidal end of the sternum was entire or
nearly so; (2) the Albatrosses, where it presented a shallow notch
upon either side of the carina; (3) those Petrels in which two
well-marked notches occurred on either side of the sternal keel.^

M. M. Hombron and Jacquinot in the year 1844, added some-
thing to our knowledge of the Tubinares,^ and they classified the
group upon the morphology of their palates, tongues, and beaks.
In one genus they placed the three genera Diomedea, Puffinus
and Priofinus, in another, the genus Prion, and finally, in their
third genus, — Procellaria. By them Pelecanoidt s \va> n inoved
from the Procellarida>, and placed in the AK a- near Alle, which
they considered its nearest relative (A. n/(/rir(i)i.s]. Viw years
later Gray and Mitchell (1849) divide the Procellarida- into the
Diomedeinse and the Procellariinc-e, and the last named into 5
genera (Prion, Pelecanoides, Thalassidroma, Procellaria, and
Puffinus), the group constituting the fourth family of their An.seres.'
In his Conspectus, Bonaparte divides the Procellaridre into the
Diomedeinse, Procellariina^, and the Halodrominae; the second

> Recherches sur I'appareil sternal des Oiseaux, pp. 79-81. v. iv. Paris,
1827.

^ Remarques sur quelques points de I'anatomie et de la. physiologic des
Procellarides, et essai d'une nouvelle classification des ces oiseaux, Compt.
Rend, de I'Acad. Sci. xviii, 1844, pp. 353-358.

^ The Genera of Birds, iii, pp. 646-650.

110

THE AMERICAN NATURALIST

[Vol. XLI

subfamily being subdivided into five lesser groups.^ But a few
years later (1864-66) this constitution was followed by the far
more accurate work of Coues, though that distinguished ornithol-
ogist complains of "having suffered not a little from imprudence
in believing Bonaparte/' whom to some extent he followed, but
upon the whole has given us a more natural classification of the
Tubinares.^

Both Bonaparte and Coues based their classification upon
the topographical anatomy of the birds of the suborder we are now
considering, but this was not the case with Eyton nor with Milne-
Edwards; nor with Huxley who followed them.^ All these
distinguished authors dealt more or less thoroughly with the
osteology of many of the Tubinares, as well as with such char-
acters as procellarine species presented externally. Eyton fig-
ured the bones of the skeleton of several varieties of Albatrosses,
and forms related to them, Milne-Edwards pointed out the
relations existing among Petrels, Gulls, and the Steganopodes ;
showing that the first two were more or less closely akin, and both
more remotely related to the last-named group of Birds. Huxley
in one of his groups of Schizognathous forms, the Cecomorphje,
in his celebrated paper, placed the Divers, the Auks, tlie (Julls,
and the Petrels in a group by theinsclvrs, and of the I- roct^llarida-
says that they "are aberrant forms, iticiiiiiiiii- towards the Cor-
morants and Pelicans among the 1 )esinouiiatha' " (/or. rif., p.
458).

Next of importance we find Professor Uciiihardt u\ 1S7;5, touch-
ing upon certain anatomical characters of Petrels, Albatrosses,
and Puffins, and presenting his classification of the Group, and
to his paper the reader is referred, inasmuch as his results are

' Conspectus generum avium, 1857, torn ii. pp. 184-206.

2 Coues, E. Critical Review of the Family Procellariidae. Proc. Acad.
Nat. Sci. Phila. pt. 1, (pp. 72-91); pt. 2, (pp. 116-144); pt. 3, (pp. 25-33);
pts. 4 and 5 (pp. 134-197). Parts 1 and 2 appeared in 1864, and the remaining

415-472.

Ill

likewise saw Garrod's studies of the Petrels appear, and finding
them 'holorhinal,' he parted them from the 'schizorhiiial ' Clulls
and related forms exhibiting a similar character. '

Other papers and works of minor taxonomic iniportaiuc (dii-
tinued to be put forth, when in 1882 appeared tlic \cry extensive
and meritorious work of Forbes dealing with the entire aiiatoniy
of many forms of the Tubinares, and a thorough study of their
probable affinities."'

Forbes divi.led the Tubinares into two famih-es. the ( hraniti.hr
and the Proccllarii.hc whicli last was sub.h'vi.lcl int.. the two
subfamih'cs 1 )i(.inc(lcina' and the Procclhiriina-. Osteology of
the Petrels and their allies fille.l a ].ronHnent ]^Uwv in this able
production, and 1 shall tmiuently have occasion in tlu^ ])resent
brief article to refer to it, es})ccially in instaiK c> wlu ic its author
had skeletons of .species which the writcM- has not been able to

Another classification is seen in that of Dr. Stcjneger whi<'li
was published in the Standard Xatnral llist.M-y .Ho>toni in isv,.
The following ..elected from Ins scheme will >how where he places
the M^il.inares:

Subclass \y. Snper-Order III. Order W. Sn].erfam. V.

I'hirhipidura^ ' Knornithes ' ( Vcomorpha- \ Pn.cellaroidea'.

In the Procellaroi.h.e are arrayed the three fanniies 1 )ioni<Mlida«,
Pr(.cellarii(he, and the Pelecanoidida^ Tlii> writer pla-o in hi.
.scheme the Tubinares wi<lely removed from the Stegano])odes,
which I believe to be a mistake, and a non-appreciation of the
morphological character, of the latter gn.np of Birds.

In his great work npon the anatomy and taxonomy of birds,
Fiirbringer makes th.> Proceliariiformes an " I ntenne.liate . Suborder '

112

THE AMERICAN NATURALIST [Vol. XLI

between his Orders Pelargonithes and Charadriornithes. He
considers the Procellariiformes to contain the Procellariae or
Tubinares to which group he gives the name of 'Gens.* The
Gens Procellarise according to him contains but the single family
— Procellariidse. Above the Procellariiformes in the Order
Pelargomithes we find the Gens Steganopodes.

In 1890 Mr. H. Seebohm in his "Classification of Birds," —
the "alternative scheme" makes an Order of the Tubinares,
placing them in his subclass Ciconiiformes, between the Stegan-
opodes and Impennes. Thus his third subclass of birds is ar-
ranged as follows: —

Subclass. Ordp:r.

Psittaci.

Raptor*

Pelecano-Herodiones

Tubinares.

Suborder.
14. Psittaci.
rl5. Striges.
16. Accipitres.
[ 17. Serpentarii.
fl8. Plataleae.
I 19. Herodiones.
120. Steganopode

21. Tubinares.

22. Impennes.

Professor Hans Gadow regards the T
light as they are by Fiirbringer, placing them as an Order Procel-
lariiformes, (9), between the orders Sphenisciformes (8) and
Ardeiformes (10), the first suborder of the latter being the
Steganopodes.^

The 'Procellariiformes' constitute Order XV of Dr. Sharpe's
classification, and it is subdivided into a suborder — Tubinares,
which latter is made to contain the three Families: (1) Diome-
deidte, (2) Procellariidae, and (3) Pelecanoidte. Of this author's
scheme, Order XIV contains the Sphenisciformes, and Order
XVI, the Alciformes.' This authority likewise widely separates
the Tubinares and the Steganopodes, the last being included in
his Order XXIII or the Pelecaniformes (loc. cit. p. 76). In

OSTEOLOGY OF THE mUXAUKS

1899 Dr. Sharpe changed this arrangement, entirely as will ho
by the following scheme which rej)resents 1 believe his li
opinions upon this subject.^ He now places the I'rocellariifoi
betw^een the Sphenisciformes and the Alciforuies.

^(XII).

Ilalocyptena. 1
Oceanodronia. l.'i

II Oceanitinae. \ Pelagodn
I Pealea.
[ Fregetta.

[ Bulweria.
f Ossifraga.

Ill Pelecanoidi.
IV Diomedi-i.

This scheme does not enumerate tlu
given by Dr. Sharpe in the Hand-Li si,
been discovered and described. TIutc

exhibiting at a glance their <

' Hand-List of Birds. Vol.

114

THE AMERICAN NATURALIST [Vol. XLI

Cope essentially agrees with Stejneger as given above, with the
exception that the Siiperfamilies of the latter are equal to the
families of the former. Thus Cope makes the Ce('()morj)ha'
contain the families Colymbidjv, lleliornitliidir, Alcida-, Laridie,
and Procellariidte.*

The writer of the present memoir added his own .snichcs to tlie
literature of this subject in a paper published in ISSl), which ap-
peared in the Proceedings of the United States National jVluseum,
it being, in its aim, more descriptive of material then in the col-
lections of that institution, rather than an atteni])t to classify the
Tubinares. In that \)i\pvr the skeleton of Orcauodroma jurcaia
is fully described and figured, also the skeletons of Fuhnarus
r/Iaciaiis an<l F. nnh/rrsll, \ru ligures being .levoted to the bones
of the latter .p(>( ie^.

Ocranodronia furrafa and Fuhnarus nuhjrrsii diiVer," and this
is foUovvcl by some notes on tlu- osteol.>gy of Vuj]iuus n irosfri.
and other mat(M-ial. Finally, a very (•omi)let(> acconnt is u'iven
of the skeleton of D/DiiKdcd allxilnis, it beinu- ilhistratcMl bv twebc

116 THE AMERICAN NATURALIST [Vol. XLI

is blunt and juts backwards and slightly outwards, being found
just at the point where the deeply sculptured supra-orbital gland-
ular depression terminates in front. Contrary to Forbes' state-
ment that "well-developed basiptervgoid facets are present in all

and ( 'vniochorca," ( j). I find tlicm liut ni(Umentary in this

.spccHiH'n of Pitjjiuus })())•( (dis ( Xo. 1777r). Smithsonian Collec-
tions), though thcv arc well-dcvclopcd and functional in a specimen
of Pvffinu.s creatopvs (Xo. 1S,773, Smithn. Coll.). In this last-

No. 482]

OSTEOLOGY OF THE TUBINARES

117

named species, too, the maxillo-palatines are well-anterior to the
ascending plates of the palatine; moreover, its vomer is notched
at its apex, and is not especially curved downward anteriorly.
These are three w^ell-marked differences in Pujfinus hormlis and
P. creatopus, and go to prove, what I have always held, tlint we
can never have too much material before as when coiupaiiiiir the
skulls or any other part of the anatomy of birds.

So far as my observation goes I find that Forbes's description
of the quadrate bone for the Tvbinares agrees with what I found
in other species of the group not examined or seen by him. But
my material does not bear him out so well in his description of the
foramen magnum of the Tubinares, and he says that that opening
"is more or less reniform, with the major axis transverse, in the
small species, whereas in the biggest it is oval, especially in Ossi-
fraga, with the long axis vertical. The moderately sized species
are here again intermediate in structure" (p. 417). Oi the two
shearwaters (Puffinus) before me, birds nearly of a size, and both
above the "small-sized species" of the group, it is found to be
oval in Pujfinus borealis, with its major axis \< iti(al. \\\\\\v in
Puffinus creatopus the foramen magnum is sulicin iilar with tin-
major axis transverse.

The mandible of Puffinus borealis has the articular mds some-
what massive, truncated posteriorly, with wry tier]) laiiial si(hvs
for its hinder half, and very shallow ones anteriorly. Apicahy it
is decurved, and there are lacking recurved angular ])r()eesses and
ramal vacuities. The articular ends are pneumatic, with the
facets for the quadrate, of course, the reverse in form to those
found on the last-named bone.

The distal elements of the greater cornua of tlie liyoideaii appa-
ratus are much flattened from above downward-, and, a- in the
Albatrosses, the parts anterior to the ba>ihranehial> are not per-
formed in bone. The first basibraneliial is siihcireular in form,

(Puffinusl

The sclerotic plates in an eye of horralis are >mall, and some-
birds.

Axial Skeleton: — In the skeleton of Pujfinus Ijorrali.s- at hand,

118

THE AMERICAN NATURALIST [Vol. XLI

I find twenty-one free vertebrae between the skull and sacrum.
Of these the thirteenth, fourteenth and fifteenth support a free
pair of ribs; they being quite rudimentary upon the first two, but
are long and slender on the fifteenth vertebra, and are without
unciform appendages. The following six vertebrae have ribs
that connect with the sternum by costal hsem apophysis. There
is also a pair of sacral or pelvic ribs, but their hsemapophyses
fail to reach the sternum, and their lower ends make extensive
articulation with the last pair of true costal ribs, at some distance
above the costal border of the sternum. The pelvis very much
resembles the pelvis of Rodger's Fulmar figured by me in the Pro-
ceedings of the U. S. National Museum (cited above), and there
are eight free caudal vertebrae plus a somew^hat elongated pygo-
style.

The costal border of the sternum is characteristically wide
from side to side, and the pits between the six facettes, unmarked
by pneumatic openings, are very^ shallow\ The sternums of these
shearwaters agree in their general characters with those of the
fulmars.

In P. horealis the xiphoidal extremity is doubly notched upon
either side of the sternal keel, and the form of the bone is tlicrv
symmetrical. This is not the case wdth the xiphoidal extrt'iiiily
of the sternum of my specimen of Puffinus creatopus. In it, not
only is the left side of the bone somewhat longer than the right,
but instead of showing the two usual notches of the right, it has
three, which appears to have been caused by a bifurcation of the
inner xiphoidal process. These inner xiphoidal processes in P.
ohscurus are wonderfully slender.

The shoulder-girdle is much like that of Daption capensis, and
in Figure 1 I present those parts in that species articulated in
situ with the sternum. This figure originally illustrated a paper
of mine which appeared in the Proceedings of the U. S. National
Museum for 1S87 (fig. 1, v. X. p. 379), where the skull is likewise
described in connection with other observations npon tlie osteology
of the Tubinarcs, and these should be read in .■,,niircii..n witli the

Ueturning to tlie shearwaters, I may say that the arrangement
of the })ones of the shoulder-girdle in some of them is as we find

No. 482]

OSTEOLOGY OF THE TllSlXARES

119

it in Daption, and tliis is the case with /^////////.s• nh.smru.^. The
sternum of the former, liowever, is noii-pneinnalic a coiKhtiim
not found in Puffinus.

Forbes in his work presents a careful and soiiK'what h'liuthy
description of the Pectoral arch in o;encraI for the T nJiliKin s.
and it agrees very closely with my own ol)servati()iis iipdn ihat

remarkable esi)ecially for the proiniiiciicr ii- jiittiii<r. pa])i]liform
ulnar crest, and (■oiisi)i(U(.ii> irianiinlar raihal crest. Its shafr
is quite straiu'ht, ami at its distal end, proximal, to the external

tached to thi.s l)y li<rament is an ossicle of n.me coiisi.lerahle si/c.

of the T: l)ent to an obtuse an-lc A ratlirr dcrp, wc1I-,I,.(iim.1
fossa exists innne<liately above tiie tub.Tch'. uhih- ihr

shallow. "In the ( )ceaiiili.he the huinefus i> (onspiciH.usly a
stouter and shorter bone, with its shaft evidently curved instead

120

THE AMERICAN NATURALIST [Vol. XLI

of being almost straight [as it is in Puffinus]. The epieondylar
process projects much less forwards, and is continued down by
an elevated ridge to the surface of the condyle itself." (Forbes,
p. 422).

Both radius and ulna in Puffinus are comparatively very slender
bones, the former, measuring 125 mm., is straight, and presents
a well-marked tendinal groove at its disto-superior aspect, over
the carpal enlargement. The ulna is likewise a very straight
bone in the shearwaters, with the elevations for the quill-butts of
the secondary remiges absent from the shaft. Its ends are but
very slightly enlarged, as they are in some birds.

The skeleton of the hand has a length almost equaling the length
of the radius. The terminal finger-points are long, slender, and
pointed distally. Claws are absent. The proximal phalanx of
index digit is very long and narrow; its blade not being fenestrated
as in the Laridse. Large and small shafts of the carpo-metacarpus
are rather close together and markedly straight. Above its prox-
imal end is a spindle-shaped, free ossicle of some considerable
size. Possibly it occurs in the tendon of the tensor patagii longus
close to its insertional extremity, but it exhibits no articular facette
for the wrist, as does the os prominens of the Owls and others.

The small phalanx of the medius digit is notably free, and
develops a tendinal tubercle upon its posterior border. Forbes
describes the pectoral limb as it exists in the ( )(raniti(ln\ in Ada-
compares the same as the skeh'ton of thi.-^ hmh is found io tlie
Diomedeina; {loc. cit. pp. 422, 423).

Puffinus borealis has a femur that in length hardly e<|uals half
that of the tibio-tarsus ; it is somewhat antero-{)osteriorly arched,
the convexity being along the anterior border. Its upper eii<l is
also antero-posteriorly flattened, with the trochanteriaii crest
about absent, and the pit for the ligamentum teres much scoojhmI
out. A small free patella exists. In the tibio-tarsus the strikirg
feature is the enormous development of its procneniinl < rest \\ \ih
a corresponding sub-suppression of the ec to( lu iiiial one. This
is even still more marked in Puffinus crralopns. whcrr upon the
posterior aspect of the common prominence, a well-nuuked,
transverse groove exists, apparently for tlie accommodation of the

No. 482] OSTEOLOGY OF THE TV BIN ARES 121

lower margin of the patella. The remaining characters of the
balance of the pelvic limb of Puffinus have already been correctly
described by Forbes, and consequently it will not be necessary to
reproduce his description in this place. He has also compared
those characters with those found in various other representatives
of this group of birds including Diomedea, Pelecanoides, the
Oceanitidae, and the Petrels, {loc. cit, pp. 424, 425.)

In examining the skeleton in the Oceanitidae I found among
other things that they lack in the skull the basipterygoid processes,
and that in them the uncinate bones, found in the skulls of other
Tubinares are also absent. The posterior margin of the xiphoidal
extremity of the sternum, is usually quite entire; and they have
but twenty-one cervico-dorsal vertebrae. These birds also possess,
in contradistinction to the Procellariidae, short and stout humeri,
a character which is also seen in the long bones of the fore-arm.

III. On the Taxonomy and Affinities of the Tubinares.

There is a combination of a few marked osteological characters
which will serve to distinguish any member of the present suborder
from any other existing avian group. The Tubinares all have
their skulls characterized by the presence of conspicuous supra-
orbital glandular depressions, which are large and generally deeply
sculpt.

They are likewise all holorhinal, as well as schizognathous
birds, wherein the vomer is usually of consi<lerable size, being
more or less broad, pointed anteriorly, and often depressed and
arched antero-posteriorly. Combined with these characters we
find that in them the hallux of pes to be either absent or else
rudimentary^ in that it is reduced to a single joint. Not more
than twenty-three cervico-dorsal vertebrae, nor less tlian twenty-
one are seen to exist. The sternum is short and broad, with its
posterior border either entire, or regularly 4-notched, or of an
asymmetrical pattern, or even jagged. The patella, when present,
is free and small, articulating high up on the posterior aspect of
the much-produced procnemial crest of the tibio-tarsus. The
sternal extremity of a coracoid is of remarkable width, being
nearly as wide there as the bone is long from summit to midpoint

122

THE AMERICAN NATTRALIST

[Vol. XLI

of base. The superior mandible of the skull is conspicuously
decurved apically, and very sharp-pointed; symphysis of mandible
also more or less decurved, and the articular ends of this bone,
truncated posteriorly.

When the skeleton of any bird has associated in it all the osteo-
logical characters here enumerated, they are sufficient to indicate
that the species belongs to the suborder Tubinares. These char-
acters are thoroughly diagnostic, and typical tubinarine forms
possess them in the avifauna in any part of the world.

I am of the opinion that the natural classification of the Tubi-
nares is as follows : —

Suborder. Families. Subfamilies.

Procellariidje.

2. PufRnida?.

Procellariinre.
Oceanitina^.
Puffininse.
Fulmariuje.

3. Pelecanoididie.
[ 4. Diomedeidse.

This arrangement does not include the extinct forms of this
suborder, but nevertheless the characters presented on the part
of these have been taken into consideration in connection with
taxonomical affinities.

When Mr. Forbes came to sum up his conclusions in regard to
this group of birds, at the close of his extensive paper, cited above,
he said that L'Herminier, A. Milne-Edwards, and Huxley have
all, in describing various points in the osteology of the Tubinares,
pointed out similarities of various kinds between their osseous
structure and that of various forms of Steganopodes, though they
still kept them close to the Larida\ Eyton, on the other hand,
places the various petrels he describes in tlie family ' Pelecanidte,'
and gulls forming a separate family liy themselves."

'Tkit no one will be prepared, I tliiiik, to dispute tliat the
Steganopodes are allied to the Ilerodiones. inchiding under that
name the Storks and Herons, with Sc<.pns only."

"Thus, on ost,.)l(>oieal grounds alone, there is >nfH<-ient gn»un(l

Herodines. And, in fact, neglecting the desmognathon.s structure
of the palate — the taxonomic value of which, per se, is becoming

No. 482] OSTEOLOGY OF THE TUBINARES

123

more and more dubious as our knowledge of the structure of birds
increases — there is little in the character assigned to the groups
PelargomorphsB and Dysporomorphce by Professor Huxley that
is not applicable to the general Petrel type." (Zoc. cit. p. 434.)

In this connection it is interesting to observe that the Tubinares
possess, in common with the Cathartidse, the Steganopodes, and
the CiconiidfE, a deep-keeled, broad and well-developed sternum ;
external osseous nares holorhinal; articular ends of mandible
posteriorly truncated; an evident tendency of the })a]atine bones
to unite with each other for their posterior moieties; powerfully
developed clavicles, which are strongly curved, — and these
osteological characters co-exist with other similarities to be found
in other parts of the morphological organizations of the respectix e
groups mentioned.

Structurally, the Cathartidse are of great interest, and the
anatomy of those peculiar terrestrial scavengers must be still better
known to us than it is, before w^e can hope to trace their probable
ancestry.

Remotely akin to the Steganopodes, the Falconida^, or more
generally, the Accipitres, also are linked with these more lowly
avian groups, — as are also the Ardeida\ through Scopus.

During the ages past, it is quite evident tliat hosts of intermediate
forms linking these families and oioiips luivc perished and become
extinct. This, taken in connection with \\\v very marked speciali-
zation of the remaining genera, goes far towards proving the great
antiquity of the entire group, and how vast that extinction of the
less specialized forms must have been.

My impression is that perhaps the Tubinares on the one hand
see their nearest relatives in the Steganopodes, in fact there can
now hardly be any doubt upon this point, — while upon the other
hand I am inclined to think that the penguins (Impennes) might
be with truth placed next below them, as Furbringer has done.
But such questions as these I will take up more thoroughly later
on, when I come, in another connection, to draw up my scheme
of classification for the Class Aves, and after I have paid further
attention to the osteology of other existing groups.

Note: — In closing this Memoir I would say that since it was
written there has appeared in the American Naturalist my con-

124

THE AMERICAN NATURALIST [Vol. XLI

tribution entitled "An Arrangement of the Families and the
Higher Groups of Birds (Vol. XXXVIII, Nos. 455-456 Nov-Dec.
1904, pp. 833-857), and, in so far as the taxonomy of the Tubi-
nares is concerned, it sustains what is set forth above; in other
words my opinion in the matter remains the same as it was six
years ago.

NOTES AND LITORAIURE

GENERAL BIOLOGY

Transmutation and Agriculture. — Mucli of the evidence ui)oii
which the evolution tlicory rests has been derived from the experi-
ments of practical breeders. It is doubtful, however, whether practi-
cal workers have ever greatly })rofited by the incorporation of the
results of their experience into general tiieories of evolution. The
present volume ^ seems to be intended as a general and popular reviev\
of the evidence which cultivated plants afford the student of the origin
of s{)ecies, rather than as a guide or handbook for those engaged in
plant breeding. Naturally many facts of interest to the breeder are
to be found in the discussions of the wide range of material treated,
but there is no attempt to formulate rules to be followe(l in any par-
ticular class of practical work or to emphasize the significance of any
particular theory of evolution for agriculturists. The arrangement of
the material under two main divisions, "Minor S|)eci(>s an<l Mutation,"
and "The Factors of Variation," iniglir sniXiXcM ih;ii th<- author is

the Lamarckian factors. Tlic source of inntcrin! is noi limited to the
results o])(aiiic(l hy connn<M-cia I l.reedcrs or aiirienltural experiment
stations, but recent e\[)erinieiita ! work of all kinds and especially that
of (le \'ries and his followers is (|iiite fully treated. In fact, the volume
furni.he. a railier i.UereMing index to recent literature bearing on the
evolution theory, it mu>t hv said, however, that it is not easy to grasp
the autiior's own point of view. The work gives somewhat the inii)res-
sion of a series of reviews, and while it is desirable that <-\oh.tionary
writings should contain less of theory and more of fact than lias fre-
(juently been the case, a work loses much in interest if it is not written
in support of definite theses which are kept constantly and clearly
in A'iew. Wanting, as it does, an obvious central purpose, the book
is not one of the kind to foinid a school and it will probably not in-
fluence evolutionary literature materially, but it does furnish a very
readable presentation of the results of much recent work and will
doubtless be of real service to many to wliom the more fundamental
works are quite inaccessible.

J. A. Harris

' Constantin, J. Le Transjormismc applique a ['Agriculture. Paris, F^lix
Alcan. 1906. 8vo, 300 pp., 105 figs.

125

126

THE AMERICAN NATURALIST [Vol. XL!

Form Analysis. — Slowly but surely the necessity of applying pre-
cise mathematical methods to the solution of many biological prob-
lems is becoming apparent to workers in both fields. The chief
application of mathematical methods has been in the study of varia-
tion and heredity, but the problems of leaf form, arrangement of leaves
on the stem, and the convolutions of the shells of gastropods may be
mentioned as having attracted the attention of niath(Mnatical workers.
In an address before the American Phil()S()j)hi( a] Society, Michelson *
emphasizes the importance of the ])r(»l)lcni> >>\' syinnictry and suggests
a classification of symmetrical and uiisyuimctrical forms.

J. A. Harris

GEOLOGY

River terraces at Brattleboro, Vt. Professor Fisher - Ims tesieil
the theory duit the river terraces of New England may he accounted
for by tiie behavior of meandering and swinging streams sh.wly
degrading previously aggraded vall.w> uiihout n.>cc.>an change in

No. 482] NOTES AND LITERATURE

the same time slowly degrading its previously a<i*i:r;i(l
encountered numerous rock barriers in its dowii-< uttin<r
controlluig the extent and character of the lateral swii
deterniinin<j the variety of terrac- pattern des.-rihcl.

The i>aiKT is al.un.lai.tlv ilhi.strat<.l l.v l>lnck diaura.
and bv maps and sections based on a careful s.n-vcv of

AXTHR()P()L()(;V

Quaternary Remains of Man in Central Europe.

Obermaier.^ The presence of man in central Knropc
nary no longer admits of doubt. The finds of an hcolo*:
tal human remains dating back to that period, ha\c I:
and well authenticated. They have, in fa< t, hvmmv m.

128

THE AMERICAN NATURALIST [Vol. XLI

No. 482] NOTES AND LITERATURE 129

Pagan Races of the Malay Peninsula ' TIh two handsonu' volumes
of over 1500 pages constitute unquestioiiahly the in(»t important con-
tribution to the knowledge of the less civilized [xoplo of -..utheastern
Asia. The work, according to the author. (|.. \\ \ rt M ,j. ) < laims to
belong to the sco|)<> of "descriptive ethnoirrapliy," l.nt iliis i> rather an

stood in this country. It i.s "essentially a c(»inpilaiion fr many

sources, but differs from most hooks of tliat kind, first, in heing ),ased
to a very large extent on materials hitherto impiihlishi'd. and accessil)le
only through private channels of information; and secondly, in having
been constructed with s])ecial knowledge of tlic .-ul.ject and in a critical
spirit." It is a work of "many facts, l„n few livp, ,t lice.," and should
be regarded not solely as a monogra])li on the trihcs de;dt with, "but
also as a necessary preliminary to a general scientilic survey of the races
of southern Indo-China and the Malay Peninsula" -which survey
is strongly advocated. The ohjectionahle term "})agan," used in the
title as a discriminative of races is jiistiHcd l)y the opinion that "the
point of religion (as between Mohammedan and non-Mohammedan)
was perhaps a better dividing line, on account of its definiteness,
than the vague, indefinite, and perhaps imdefinal)l(\ (piality of wild-
ness." The bulk of the book was written l.y Skeat. the attention of
Blagden being confined to languag.-.

The contents of the two volumes, I.esi.le. preface. !)il>li.)-rapliy, and
introduction, are, vol. I: Racial characters and aliinhies: \oi»vs on

in further St u.lio of the people, of the Malay Penins„h,'. wll as
that from the mainland further north and ih.' inlands to the s,>uth-

130

THE AMERICAN NATURALIST [Vol. XLI

Negrito, the Sakai, of suggested Dravidian ancestry, and the Jakun,
or aboriginal Malay. They dif¥er principally in head kmn, physiog-
nomy, and nature of the hair. The Semang are meso- to brachy-
cephalic, with woolly hair, and features approaching, in a number of
particulars, the negro; the Sakai are dolichocephalic, with wavy hair
and finer features; the Jakun are brachy cephalic, with straight hair
and with the features of the Malay in general. All are short in stature,
but the Semang are the smallest. In color the Semang are chocolate-
brown to black, the Sakai and Jakun ranging from brown to yellowish.
Both the Sakai and Jakun show numerous instances of admixture with
the Negrito.

The chapters on the foods and mode of life of the individual
tribes are valuable; but the diseases of the peo])le, their environment,
and especially their physiology are far from being treated adequately.
The total number of the 'pagan' aboriginies of the j\Ialay Peninsula
appears to be no more than 35,000 or 40,000.

For the mass of details concerning the habits, religion, folk-lore and
language of the tribes the reader must be referred to the original.

The book as a whole will not be found easy reading. This is partly
due to its plan, including several appendices, partly to the many native
names, and in some degree to the style of the authors. More tabula-
tion would have been of help. However, the work must be regarded

every .student of that region. For this purpose, luiwever, a more
copious in<lex, and page references instead of ti.e o<r:.sio,w,l "will l,e
found in another part of the work/' wouM have l.ee„ de^irahle.

The illustrations are not alwiivs sal isfaetory. Then- are a nuiiil)er
of photographs that show but little, and a few (r. r/. the "Kedah-
Raman," "Kedah," superior plane of the Semang skull, the "Semang
of Grit," the "Sakai at G. Kerbu," the "Group of Ulu Jelai Sakai")
which are wholly useless, being out of focus. It is not easy to see what
was the object of the authors or publishers in including these pictures

A. Hrdlicka

Growth of Parisian Children.' The paper presents the results of

' Tal.les ,lo cvuissanrv <l<-s ciifants I'ansiens ,le I a Hi ans. I'ar .\I.M. \ ariot
et Chaumet. Jiull. Mein. .Soc. d Anthrop. Paris, Vme 8er., Ml, No. 2, pp.
.51-65.

I^o. 482] NOTES AND LITERATURE

131

Parisian nurseries and schools. The series includes at least 100 sub-
jects of each sex for every year of life, which insures the value of the
averages. The study is the first of its nature made in France ; Godin's
well known observations were made on older individuals.

The results agree in the main with those of measurements of white
children in other countries.^ Up to the cud of their eleventh year the
girls are shorter than the boys; between their eleventh and twelfth
years they pass the boys in this regard, and continue^ taller until after
their fourteenth year, after which they are (Icfinitdv passed by the
boys. In weight the physiological excess of the f. male children
becomes marked even earlier and they (>xeeed die 1m)\>, ti(.ni the end
of the ninth to a little beyond their fifteeth vear.

A comparison of (Ih-m- ("lata uith th(»e ol.t'aiiie.l by Professor C. P.
Bowditch on liosion chihhvn shows that IxMween the ages of thirteen
and sixteen the Parisians shj^ditly exceed the Americans in height.
This can very likely be attributed to earlier puberty in the French
adolescents.

A. H.

Anthropometric data on the Norwegians. — Messrs. Daae report^ the

between 22 and 2:-; years of aiit>.

The data show that the average stature of the Norwegians of that
age is 172.1 cm. The tallest men are in the district of Jarlsberg-Larvik
(173.4 cm.), the shortest in the district of Finmarken (168.5 cm.).

103.00 to 100. It is' relatively shortest (102.2 to 100) in the Hergenhus-
Sud district, peopled by fisheruKMi wlio all the year around work with

Daae et le Dr. H. Daae. Bull. & M^m. Soc. <i Anthrep. Taris, \ n,r S. r.
VII, No. 3, 1906, pp. 158-164.

132

THE AMERICAN NATURALIST

[Vol. XLI

The population of Tripoli, according to the latest official data,*
amounts to 711,242. Among this are 16,670 Jews. The most south-
ern point at which the latter are found is Orfella. They live an ex-
tremely miserable life and in places suffer even partial slavery. They
do not emigrate because they know not where to go.

A. H.

ZOOLOGY

Dean's Chimaeroid Fishes ^ is one of the most strikingly illustrated
works yet issued by the Carnegie Institution. Any adequate sum-
mary of its contents is impossible here; all that can be attempted is
an enumeration of its contents. For several years Dr. Dean has
labored indefatigably in obtaining embryos of this group of rare
Selachians. The work is based on the eggs of the Pacific Chimcera
collei, the eggs of which were obtained from the gravid females and then
incubated in floating boxes, but unfortunately these often broke adrift
and about 150 eggs have been lost in this way.

After an introductory chapter on methods and the like Dr. Dean
first describes the appearance, habits, etc. , of the fish and then proceeds
to a study of the development. The egg-capsule is beautifully figured
and described in detail, this part of the work being made more valuable
by figures of the egg-capsules of other chimseroids, both recent and
fossil. The egg is fertilized before oviposition and Dr. Dean was
fortunate enough to get specimens showing various phases of the proc-
ess of fusion of the male and female pronuclei. Polyspemy is appar-
ently the usual condition. The segmentation is in general of the usual
Selachian discoidal type but is accompanied by a fragmentation of
the yolk. A single early stage of gastrulation is described in detail, the
striking feature being that the blastopore is not, as in other clasino-
branchs, at the edge of the blastoderm but inside its rim, a (ondition
which throws much light on gastrulation in other forms, eoneiusioiis
which are supported by two other stag€\s.

• M6hier de Mathuisieulx, L'Anthropologie, XVII, 1906, Nos. 1-2, pp. 237-

2 Dean, Bashford: Chimseroid Fishes and their development. Carnegie
Institution, Publication 32, Washington, 1906, pp. 194, 11 plates.

No. 482] NOTES AND LITERATURE

133

Of the stages after the closure of the medullary folds the accounts
are far less detailed than we could wish and there are many gaps in
the organogeny which remain to be filled but which cannot at present
be described on account of lack of material. Especially interesting
are the figures given of a reconstruction of the skull of a well advanced
embryo in which the pterygoquadrate bar is not completely fused
with the cranium. Other features of organogeny given are concerned
with (1) the integument and dentition in which embryos and larvae of
other chimseroids are considered and the conclusion is reached that
the dental plates represent fused denticles. (2) The skeleton which is
largely based on the work of Schauinsland. (3) The viscera. There
is, even in early stages, no continuous mesentery. A few words are
devoted to gut, gills and nephridial structures.

The third section, one of the most valuable of the work, is a discus-
sion of the fossil chimseroids. The existence of Silurian members of
the group is more than doubted, but, as shown by the Ptyctodonts,
they probably occurred in the Devonian. The definite knowledge of
the group began with the lower Jurassic, since which time numerous
undoubted chimseroids have occurred, the group attaining its maxi-
mum development in the cretaceous. Thvsv fossils and The stnictnrc
and embryology of the existing species arc invoked to show that the
chimffiroids are not a primitive group but are a ino(HHed and >|)eeiahz( d
development from forms more like the normal Selacliiaiis. An rwvu-
sive bibliography closes the volume.

.1. S. K.

Development of the Mammalian Lung. Flint (Am. Journ. Anat.
6, 1906) describes in a l(Mig ])aper the development of the lung and
associated structures in tlie pig Tlie anlage is asymmetrical, and its
origin, below the level of the gill pouches is an argument against any
phylogenetic connection between lungs and gill pouches. The develop-
ment of the bronchi is followed in detail and many variations noted, the
complete series including sixteen on one side and seventeen on the
other, a condition rarely occurring. iEby's conclusion that the pul-
monary artery differentiates two lung regions of different morphologi-
cal significance is not supported. The pulmonary veins arise as an
outgrowth from the undivided portion of the sinus venosus, the veins
to the right and left lungs developing by specialization in the capillary
plexus. In the eaiHer history the division of the respiratory ducts is
monopodial in ( haracter as in the lower pulmonate vertebrates and it
is only in the other stages that dichotomous division, characteristic

134

THE AMERICAN NATURALIST [Vol. XLI

of the mammals, sets in. The histogenesis and the development of
the lymphatic system are also traced. The early stages were studied
by Born reconstruction methods, the later by dissection and by corro-
sive preparations.

Half Hours with Fishes, Reptiles and Birds ^ is the second in the
series of books by C. F. Holder, designed as supplementary readers
for children in the grammar grades. The section devoted to birds
suffers from the same defects in the arrangement of material that were
pointed out in the review of the earlier volume (American Naturalist,
40, p. 140, 1906). The part dealing with fishes is full of interesting
information vividly presented.

R. H.

Notes. — In the Proceedings of the Indiana Academy of Science for
1905 (1906) Dennis and Petry give an interesting series of photographs
of the young of the turkey buzzard showing the changes in the plumage
from the tenth to the seventy-fourth day after hatching.

Zeleny (Proc. Acad. Sciences Indiana [for 1905] 1906) describes
the regeneration of an antenna-like appendage in the place of an
excised eye in the blind crayfish. The new organ has tlie ai)|)canince of
a functional tactile organ and the experiment has csiHcial interest in
that a functional organ has developed in place of the fnnctionhvss eye.

Martin describes (Proc. Indiana Acad. Sci. [for lOO.')] IDOii) a liandy
clamp by which the blades of 'safety razors' may lie used for xciion
cutting, thus materially reducing the cost, confusion, etc, of supplying-
section knives to large classes.

Madison Grant publishes some "Notes on Adirondack ?tlainmals"
in the Eighth and Ninth Report of the Forest Fisli and ( Janic ( oinmis-
sion of New York. The paper, which supplements Dr. Merriam's
well known work on the same region, is illustrated w ith some line half
tones, some taken in the forest, others in the New ^'ork Zoological

(\ W. .Johnson has eolleeted all the refereneo to the app.'arane.- an<l
<listril.ution of the Kn^rUsh ^Mrden snail, Ihlix hnrlrn.sls, in .\meriea
and is inclined t.> think (Nautilus, 20, p. 7:^,, m)i\) that it has not been

' Half Hours wit h I'ishes. Reptiles and Birds. By Charle.s Frederick Holder.
N. Y. American Book Company, pp. 255. Illustrated.

ISO. 482] NOTES AND LITERATURE

135

introduced by man within comparatively recent years nor by the
vikings" but is a much older inhabitant of this continent.

Ldnnberg (Arkiv for Zoologi, 3, 1906) discusses the systematic posi-
tion of the extinct Irish Elk. This is usually closely associated with
the common fallow deer. Lonnberg thinks that this association
rests almost exclusively upon the somewhat similar palmated antlers
but that in other and more ini])()rtant fciiturcs tlicre is more afliiiity
with the reindeer than with any other cervicorn, altliouo-h it pivsciiis
considerable specialization in its own Hue.

Froriep gives (Verhandl. An atom. Gesellschaft, XX, 1906) a detailed
comparison of the eyes of vertebrates and tunicates and concludes
that both are derivable from a common ancestral condition which is
closer to the optic pit of the vertebrate than to the eye of the ascidian
larva. Two weeks later comes the Anatomischer Anzeiger (xxix, p.
52() Nov. 24, 1906) in which Metcalf discusses the relation of the ver-
tebrate eye to that of Salpa suggested by Redikorzew, and holds that
the views of the latter are untenable but he says "It may not unlikely
be true that the condition with a single anterior enlarginncnt of the
central nerve tube is ancestral (cf. Amphioxus and tlic tunicate tad-
pole)."

BOTANY

The Journals: — Tlic Anirricdn Batanisf, September: — Saunders,
"Under Sierra Pines"; Bailey. "Tlw Leaf Alert or Drowsy " ; Dobbin,

^ylvaik-a "; Best, - Pfj/rhomlfrlum h ihrnjli-': Ilowe. "SoirH^ Addi-
tions to the Flora of'^Iiddiesex County, Mavs.'-; XayhT. '■Micro-
scopical Technique"; Merrill. " Lieh(-ii Notes no. K A Stu.ly o{
UmhiUrarin vrllni and T. s/Hvlnrhma ."

the Mis< ou Beach Plain"-; Slireve. "The I >eveloi.nHMit and Anatoniy
of Sarracenia"; ( )sterhout, 'TMiysiologically Balan.rd Solutions for
Plants"; Hasselbring, " The Appressoria of the Anthracnoses"; Frye,

136

THE AMERICAN NATURALIST

[Vol. XLI

" Nereocystis luetkeana"; Greenman, "Two New Species from North-
eastern America."

The Botanical Gazette, September:— Blakeslee, "Differentiation of
Sex in Thallus Gametophyte and Sporophyte"; Shantz, "A Study
of the Vegetation of the Mesa Region East of Pike's Peak: The
Bouteloua Formation — 11"; Kauffman, "Cortinarius as a Myco-
rhiza-producing Fungus"; Smith and Smith, "A New Fungus of
Economic Importance" {Pythiacystis ciiriophthora, — forming a
transition from Pythium to Phytophthora].

The Botanical Gazette, October :— Atkinson, "The Development
of Agaricus campestris" ; Crocker, "R6le of Seed Coats in Delayed
Germination"; J. D. Smith, " Undescribed Plants from Guatemala
and Other Central American Republics"; C. O. Smith, "A Bacterial
Disease of Oleander."

Bulletin of the Torrey Botanical Club, August: — Arthur and Kern,
"North American Species of Peridermium " ; MacKenzie, "Notes on
Carex — I"; Abrams, "Two New Southwestern Species of Pent-
stemon."

Bulletin of the Torrey Botanical Club, September:— Eaton, "Pteri-
dophytes Observed during three Excursions into Southern Florida";
Mathewson, "The Behavior of the Pollen-tube in Iloustonia cwrulea'' ;
House, " Studies in the North American ConvolvulacejB — II. The
Genus Operculina."

Bulletin of the Southern California Academy of Sciences, June: —
Hasse, "Contributions to the Lichen Flora of Southern California";
Parish, "Additions and Corrections," and "A Preliminary Synopsis
of the Southern California Cyperaceae — XII."

Journal of Mycology, July: — Kellerman, "Mycological Expedition
to Guatemala"; Charles, "Occurrence of Lasiodiplodia on Theobroma
cacao and Mangifera indica" ; Hedgcock and Spaulding, " A New
Method of Mounting Fungi Grown in Cultures for the Herbarium " ;
Peck, "A New Species of Galera"; Arthur, "Reasons for Desiring a
Better Classification of the Uredinales"; Morgan, "North American
SpcMM.s of T.cpiota. [I.] Descriptive Synopses of Morgan's North
Aiiicri( :iii Sp<Tii>> of M;ir:isiiiiti>"; ami " Synopsis to North American
Sprci*-, of llclioiiivco"; (ianctt, " Fi< l(l Notes on the Uredineae";
Kellernum, "Not.vs from Mycological Literature — XX."

Journal of the New York Botanical Garden, September: — Murrill,
"Further Remarks on a Serious Chestnut Disease"; Rusby, "Obser-

No. 482]

NOTES AND LITERATURE

137

vations in Economic Botany ]\Iade at Oscoda, ]\Iich." Gager, "Sym-

biosis in Gunnera mmiicata."

Journal of the New York Botanical Garden, October:— IMurrill,
"A Summer in Europe: Some Foreign Botanists and Botanical
Institutions."

The Plant World, August :— Fink, "The Gynsecocentric Theory
and the Sexes in Plants"; Rusby, "An Historical Sketch of the Devel-
opment of Botany in New York City" {concluded); Cook, "Tropical
Epiphytes."

Tlie Plant World, September:— Shreve, "The Hope Botanical
Gardens"; Gager, "Outline Study of Seeds and Seedlings"; Robin-
son, "The Filmy Ferns."

Rhodora, August: — Lamson-Scribner, "The Genus Sphenopholis " ;
Blanchard, "Some Maine Rubi. The Blackberries of the Kenne-
bunks and Wells — I"; Collins, "Notes on Algfe- VHI"; Fernald,
"Some New or Little Known Cyperaceae of Eastern North America."

Rhodora, September: nhuH-hard. "Some Maine Rubi. The
Blackberries of the KcMuul.unks aii.l Wells — H"; Fernald, "Some
New or Little Known (\ih-vavv:v ..f Hastern North America"
tinued); Knight, "A Now \ ari.ty o( Carr.r frispn-ma" ; Hill, "The
Perianth of Rynchosjxn-a m pillar, a var. h ris, t,r ; Knight, " lla-
benaria macrophylla in Maine."

Rhodora, October:— Collins, " Acro(ha>tium and Chantransia in
North America"; Robinson, 'The Nomenclature of the New England
Lauraceffi"; Fernald, "Some New or Little Known Cyperacete of
Eastern North America"; Robinson, " Filipendtda rubra, a new
Binomial."

The fourth annual volume of the Ifiternatwnal Catalogue of Scien-
tific Literature, M, Botany, is dated in July, 1906, and forms an octavo
of nearly 1000 pages.

Torreya, September: — Gager, "Tuber-Formation in Solanum
tuberosum in Daylight," Murrill, "A New Chestnut Disease" [Dia-
porthe parasitica]; Bailey, "A Newly Introduced Plant in Rhode
Island"; Hollick, "An Addition to the Flora of Block Island"; Rob-
bins, "Tubular Ray-Flowers in Gaillardia arisiaia"; Wilson, "Mv-
cological Notes from Indiana"; Harper, "A hitherto Unnoticed
Relation Between Viok pedata and Iris verua"; Bruckman, "Fasci-
ations in Arissema, Rudbeckia, and Viola."

138

THE AMERICAN NATURALIST

[Vol. XLI

Torret/a, October:— Harper, "Midwinter Observations in South-
eastern Mississippi and Eastern Louisiana"; Dowell, "Observations
on the Occurrence of Boott's Fern"; Farwell; " Note on the Identity
of Trillium obovatum Pursh"; MacKenzie, "Lespedrga .vmuhifn \n
New Jersey"; Gager, "Further Note on the Formation of Aerial
Tubers in Solanum."

Voh 7, part 4, of the current botanical series of T musacfions of the
Linnean Society of London is devoted to an account of Sutclifiia,
representing a new type of MeduUoseje from tin- li)\v( i- Coal ^bas-

Zoe, September:— Brandegee, "Plants of California," "New Species
of Mexican Plants Collected by Dr. C. A. Purpus," and "Plants of
Sinaloa."

The following papers of botanical interest occur in the recently
issued second volume of Proceedings of the American Breeders' Asso-
ciation:— Shamel, "Tobacco Breeding"; Montgomery, "The Corn
Plant as Affected by Rate of Planting"; Lyon, "Some Correlated
Characters in Wheat and Their Transmission " ; Ten Eyck, "Plant
Adaptation"; Freeman, "The Use of the Seed Plant in the Prevention
of Diseases in Wheat"; Ward, "Economic Value of Plant Breeding";
Westiratc. "A ^b■rho.l oF Bivc<lii,o- Strain of Alfalfa from a Single
In.livi.luar'; Wcl.hcr. " Convhiti*.., of ( 1,ara<-lcrs in Plant Htv<.lino " ;
Keyscr, 'A'ariation in Wheat IlvWrid.'"; Funk, "Practical Corn

son, "Laboratory^' Work in Pl!int Hivc.ling" T ( iauss. Breeding
Drought-Resistant Crops"; Hessey. "Croi. Iniproveuu-nt l.y Ltilizing
Wild Species"; Zavitz, "lireeding Cereals"; Hansen, "Breeding
Hardy Raspberries for the Northwest"; Carleton, "Fundamental
Requirements for Grain Breeding"; Hartley, "Value of Corn Pollen
from Suckers vs. from Main Stalks"; Stockdale, "Improvement of
Sugar Cane by Selection and Hybridization"; Havs, "American
Work in Breeding Plants and Animals"; llans<-n, " M.'tliods of Breed-
ing Hardy Fruits"; Williams. "Methods and Besults of llvl.ridi/.-
ing Fruits"; Keyser, "Metho.l^ in Whe;,t Breeding"; B,.;,cli, ''(irai.c
Bree(hng"; Fruwirtli, "Enclosing Single I'lanis, and its FtlVct on a
Large Number of Important Agricultural Specie-,"; Camp, "Br»MMling
Grapes"; Patten, "Results from Work in Breeding Hardy Fruits."

Young man, aged 26, excellent collector and setter of
Lepidoptera, can also make bird skins, desires a place
as assistant with museum, professional naturalist or private
collector. Is prepared to travel abroad. Write C, Box
4373 Willings, 125 Strand, London, W. C. England.

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VOL. XLI, NO. 483

MARCH, 1907

THE

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

I. studies on the Ophioglossacese PROF. D. H. CAMPBELL 189

II. Polygamy and other Modes of Mating among Birds DE. E. W. SHUyELDT 161
in. On the Wood Bails, genus Aramides, occurring north of Panama

OTJTEAM BANGS 117
IV. Notes and Literature : fi/o/o^/. The Reception of the Mutation Theory. Mon-
ument to Theodor Schwann, Fitch's Basis of Mind and Morals; Geology,
The Reconstruction of the Continents of Tertiary Times, The Mountains
of Cape Colony, Natural Mounds, Ancient Glacial Periods; Physiology.
Hough and Sedgwick's Physiology; Zoology. Guyer's Animal Microl-
ogy, MoUusca of lUinois and Michigan, A Monograph on Anurida;
Botany, A popular book on Canadian Wild Flowers, Notas, The Journals 189

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The American Naturalist

ASSOCIATE EDITORS
J. A. ALLEN, Ph.D., American Museum of Natural History, New York
E. A. ANDREWS, Ph.D., Johns Hopkins Universittj, Baltimore
WILLIAM S. BAYLEY, Ph.D., Colby University, WaterviUe
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell University, Ithaca.
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA, M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University
CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM, Ph.D., Cornell University
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, PUtsburg
D. P. PENHALLOW, D.Sc, F.R.S.C., McGill University, Montreal
H. M. RICHARDS, S.D., CoMia University, New York
W. E. RITTER, Ph.D., University of California, Berkeley
ERWIN F. SMITH, S.D., U. S. Department of AgricuUure, Washington
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington
W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis
HENRY B. WARD, Ph.D., Universiiy of Nebraska, Lincoln
WILLIAM M. WHEELER, Ph.D., American Museum of Natural History,
New York

facts and discoveries in Anthropology, General Biology, Zoology,
Botany, Paleontology, Geology and Physical Geography, and Miner-
alogy and Petrography. The contents each month will consist of
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THE

AMERICAN NATURALIST

Vol. XLI March, 1907 No. 483

STUDIES ON THE OPHIOGLOSSACE.^
DOUGLAS HOUGHTON CAMPBELL

The family of the Ophioglossacese comprises the three genera —
Ophioglossum, Botrychium and Helminthostachys, which are all
evidently related, but whose affinities with the other Pteridophytes
are not so clear, and there is a good deal of difference of opinion as
to where they should be placed. Most botanists agree that the
Ophioglossacese are related to the true ferns, but this view is not
universally accepted, although the results of the more recent in-
vestigations tend to strengthen this conclusion.

The most marked feature of the family is the peculiar fertile
leaf segment or spike; and the present paper is mainly concerned
with the question of the morphologic nature of this sporophyll.

During the past year the writer had an opportunity of collect-
ing a large amount of material of the 0{)hioglossacefe in Ceylon,
Singapore and .lava. 'V\\\> iiicliKled several species of Ophio-
glossum, one (if Hotrvchiiim, :ui<l the nionotypic Helmintho-
stachys,— so that it has been possil^lf To make a first-hand study
of all the genera belonging to the family. The following account
of the morphology of the leaf is ha-^cd nuiiiily \\\)on a study of
this material.

'riii: MnKnioi.ocv ..r thk Si-ohuimi yi.i..

fertile portion of the snike is more or le.ss extensively branched,

139

140

THE AMERICAN NATURALIST

[Vol. XLI

this being very marked in the larger species of Botrychium. The
two segments of the sporophyll may be almost entirely separate,
e. g., Ophioglossum bergianum, Botrychium ternatum, or the fertile
segment may be apparently an outgrowth of the base of the sterile
segment or from above its base.

The earlier views of the morphologic value of the fertile leaf
segment were strongly influenced by the prevailing theory that
the fertile portion was a secondary development of originally
sterile leaf tissue, and therefore must be homologized with some
portion of the sterile leaf. The belief more generally current
at present that the fertile structures of the sporophyll are older
than the sterile ones, inclines toward a different interpretation
of the real nature of the fertile segment.

Bower (Studies in the ^Morphology of Spore-producing Mem-
bers, II, Ophioglossacese. London, 1896) has given a very com-
plete account of the different theories that have been advanced
to explain the morphology of the fertile spike in the Ophioglos-
sacete, and we shall merely give here a brief summary of the more
important of these. ]\Iettenius (Fame des Bot. Garten zu Ixip-
zig. 1856, p. 119) regarded the two parts of the leaf as of equal
importance, but gives no data as to their method of origin, —
whether by the equal branching of a common primordium or
otherwise. Later writers, e. Holle (Bot. Zeit. 1875, p. 271)
and Goebel (Schenk's Handbuch, vol. 3, p. Ill) consider the
fertile spike as the equivalent of the fertile pinnse of such a fern
as Aneimia. The former considers the single median spike to
be the result of the coalescence of two lateral pinnie; the latter
as a single pinna which arises in a median position.

Bower himself has made the most complete study of the develop-
ment of the spore-bearing parts of the Ophioglossacete that has
ever been made. He concludes that the spike of Ophioglossum
is morphologically equivalent to the single sporangium of Lycopo-
dium. In this view he has the support of Strasburger (Bot.
Zeit., 1873) and Celakovsky (Pringsheim's Jahrb., 1SS4, vol. 14).
Bower has, however, more recently described a most remarkable
species of Ophioglossum (Ann. of Bot. 18, p. 205, 1904) (). .simp/r.r
Ridley, which makes possible another interpretation of the nature
of the spike, i. c, that it is a terminal and not a lateral organ.
The writer (Mosses & Ferns, 2d edit., p. 600) in view of the dis-

No. 483] STUDIES OF THE OPH lOCl.DSSACE.K 141

covery of this remarkable form, has ventured the hypothesis
that in O. 'pendulum the sporangiophore may also be terminal.
In order to make a thorough investigation of the question, the
collections of material already referred to were made and the
results of this study and the conclusions to be drawn from it are
given in the present paper.

THE GENERAL MORPHOLOGY OF THE 8P0R0PHYLL.

and thvvv is evi<lentlv very miicli confusion as to the species.

142

THE AMERICAN NATURALIST [Vol. XLI

Raciborski, who has published a list of Javanese Pteridophytes
(Die Pteridophyten der Flora von Buitenzorg, I.eiden, 1898) gives
only one terrestrial species, O. moluccanum Schlecht.; but it is
evident from the writer's collections that there are at least four spe-
cies belonging to Euophioglossum in western
Java and possibly more.

What seems to be the typical O. moluccanum
(fig. 1, A) is a species of moderate size. The
specimen shown has a sterile leaf somewhat
smaller than usual, but otherwise is typical.
One of the smaller forms of the same ( ?) spe-
cies is shown in fig. 1, B. In both of these
the sterile lamina is small, while the peduncle of
the spike is very long and not very much infe-
rior in thickness to petiole below the junction
of the spike and the sterile lamina. Most of
the other species of the section, e. g., O. vul-
gatum L., 0. mZi/ornzcww Prantl, 0. reticulatum
L., etc., agree in the main with 0. moluccanum,
and in none of these is there anything in the
external morphology of the adult sporophyll to
forbid the assumption that the sterile lamina
is a lateral appendage of the spike.

The second section of the genus, Ophio-
derma, comprises (). prndii/inii L., (). hifcrme-
dium Hook, and probably also (). simplex
Ridley. In the latter species (fig. 2), which
was discovered by Ridley in Sumatra, the fer-
tile leaf consists of a narrow basal part without
any lann"na, terminated bv a s])ike similar to
that in ().pn,dHl,iu,,'Au.\h wa> assumed to be
the nearest relative of this specie.^. There is,
Fig. 2.-l'la,u oi however, no |,e.hniele .levelope.l as is the case

arises from the lamina it>elf. i> continued into a >ort of thickened
mid-rib which is not developfMl above the in-ei'iion of the |.e(hnicle

No. 483] STUDIES OF THE OPH lOGLOSSACE.E

143

of the spike, and the latter may very well be interpreted eis the
apex of the leaf, the lamina being lateral and closely coherent with
its basal portion.

In all the species of Ophioglossum the growth of the basal part
of the young sporophyll is very much more active than that of the
lamina which remains relatively small, although the young spike
is conspicuous in the early stages. This is especially marked in 0.

'pendulum (Fig. 3). Tliis is the largest of the genus, and is a
striking epiphyte of moist tropics of the old world, extending,
however, to the llau aiujii 1 shun Is. The specimens figured were
collected in the botaiiicul u;u-<leii at Singapore.

In the youngest specimen siiown (Fig. 3, A), the thick fleshy

144

THE A M ERIC A N NA T URA LIST [Vol. XLl

leaf base terminates in a very small pointed lamina that is usually
bent over, suggesting the circinate vernation of the true ferns. In
most of the terrestrial species of Ophioglossum the young leaf is
folded straight in the bud. Under the arched hood formed by the
lamina is the young spike (*Sp.) which almost equals the lamina in
length.

Fig. 3, B, shows a somewhat older stage. The leaf has now
become somewhat flattened, but there is no clear demarkation
between the petiole and the small lamina. The fertile segment,
which shows as yet no differentiation of the peduncle and spike,
is conspicuous, and merges gradually into the thick petiole of the
leaf whose margins are more or less distinctly winged and pass
imperceptibly into the lamina above the insertion of the fertile
segment. The interpretation of the latter as terminal and the
sterile portion as a lateral appendage coherent with it would seem
entirely plausible. An interesting case is shown in Fig. 3, E,
where the lamina is almost entirely suppressed, and the terminal
character of the spike is very evident.

As the leaf develops there is a very great increase in size of the
lamina, which, in some of the largest individuals collected in Cey-
lon and Java, reached a length of one and one-half metres, or even
more. These large leaves usually have the lamina dichotomously
divided, and strikingly resemble the long (ir()<)j)iiig leaves of some
species of Platycerium. Nevertheless even in these huirer leaves
the segments are quite destitute of a mid-ril). This stops at the
base of the peduncle of the spike into which it is continued. The
spike in these large specimens is correspondingly large, and some-
times attains a length of 25 to 30 centimetres, with a breadth of
more than a centimeter (Fig. 3, I)).

Undoubtedly allied to O. pendulum is the rare O. intermedium
Hook. (Fig. 4). ^Miis is also perhaps the nearest ally of O. .^'im-
plex. In the ordinary form iVh^. 4, A, B) this is not nnhke a
small specimen of O. pendulnw, hnt it i> riiridly ni)rioht instead

leaf nnich >n,aller'an.l nlore >harply ..-paratrd from th.' petiole.
As in i). pendulum, however, the petioh- is pn.longed into the
peduncle of the spike with the same niid-rib like thickening,
caused by the coherence of the basal part of the peduncle with

i\o. 483] STUDIES OF THE OPHIOGLOSSACE.^

145

Even in the small number of specimens collected (the plant is
an extremely rare one) a number of very interesting variations
were found, some of which approximated quite closely the condi-
tion found in O. simplex. In these the lamina was greatly reduced,
and in one case (Fig. 4, E) formed merely the narrow wing along
the margin of the petiole and peduncle of the spike. In the other

cases the lamina was wider and its apex free, but even in these the
lamina was \ery small, and the terminal position of the spike
extremely evident (C, D).

In both 0. pendulum and 0. intermedium the spike is more
flattened than in the section Euophioglossum, and the central
sterile portion wider in proportion. Stomata are almost entirely
absent from the spike of O. pendulum, and the few that are occa-
sionally found are confined to the criitral part. In O. Infrniirdium
the stomata are more numerous than in (). piitdiduin, but much
less numerous than in 0. moluceatium, for example, where they
also occur upon the epidermis of the wall of the sporangium.

146 THE AMERICAN NATURALIST [Vol. XLI

The third section, Cheiroglossa, represented by the monotypic
0. palmahim L. of the American tropics differs from the others of
the genus in having, usually, several spikes which are not generally
borne in the median plane of the leaf, but are inserted near the
margin. Bower (loc. cit., figs. 116-117) has shown that there
may occasionally be a single spike which is then borne in the
same position as in 0. pendulum. He supposes that O. palmatum
has been derived from the form with a single median spike like
that of O. pendulum by branching of the spike, which not infre-
quently occurs in the latter species as well as in some others. The
separation of the originally connected spikes he assumes has been
the result of the great expansion of the lamina, which is much
broader in 0. palmatum than in any other species. Unfortunately
the developmental history of the sporophyll in 0. palmatum is
quite unknown.

The Young Sporophyll.
The differentiation of the two parts of the sporophyll takes

No. 483]

STUDIES OF THE OPHIOGLOSSACE.E

147

place at a very early period, and at this time the fertile spike is
already evident as a conspicuous protuberance on the adaxial
side of the leaf rudiment not far from its apex. Both divisions
of the young sporophyll terminate in an apical cell, and both
apparently grow in the same way.

Fig. 5, A, shows a nearly median section of a very young
sporophyll of 0. pendulum. This is a broadly conical body upon
whose inner (adaxial) face there is a slight prominence (Sp.) the
apex of the young spike. Fig. 5, B,

shows an older, but still very early ^ ^

stage, in which it is evident that the
spike rudiment extends completely to
the base of the young leaf , with which
it is adherent except at the extreme
tip. The apex of the young spike is
directed upward and its axis is almost
parallel with that of the sterile leaf
segment. From Bower's figures of
corresponding stages in 0. vulgatum
it is clear that a very similar condition
of things prevails in that species. In
such a stage as that shown in Fig 5,
B, the relation of the fertile and sterile
segments is not un]ik(> tliat of a stem
apex and leaf, and the conditicMi of
things here present would \ery well
lend itself to the intci pretation of a
terminal spike widi a sulitending ster-
ile lamina. At this stage the vascular
bundles are not yet difi'ereiitiated, and
the arrangement of these in the young
leaf still remains to be made out.

the temperate zones, both the fertile

and sterile segments of the leaf as is well known, except i

148

THE AMERICAN NATURALIST

[Vol. XLl

simple forms of B. simplex, are more or less extensively branched.
This is especially marked in such large species as B. virginianum
and B. lanuginosum.

The relation of the fertile and sterile periods is essentially the
same as in Ophioglossum, and there is the same variation in the
point of divergence of the two leaf segments. Thus in O. ohli-
■quum Muhl. the two are separated almost to the base. In 0. vir-
ginianum and O. lanuginosum (Fig. 6, B) the spike appears to arise
■close to the lamina of the leaf or even above its base. No material
was available for a critical study of this point in B. virginianum,
but in 0. lanuginosum Wall, where (see Engler & Prantl, loc.
cit., p. 471) it is stated that the spike arises from the base of the
sterile segment; even a casual examination will show that this is
more apparent than real (see Fig. 6, B). If the leaf be looked at
from in front it is very evident that the peduncle can be traced
for a long distance below the bases of the sterile leaf segments,
although only the anterior face is free, the inner face and sides
being completely adherent to the base of the sterile segments.

Helminthostachys.

A similar condition to that found in Botrychium lanuginosum
prevails in the third genus, Helminthostachys (Fig. 7), a mono-
typic genus of the Indo-Malayan region. This is much nearer to
Botrychium, in its general morphology, than it is to Ophioglossum,
although, in the character of both the prothallium and fertile
spike, it is to some extent intermediate in character between the
two genera.

In Helminthostachys the sterile segment, as in most species
of Botrychium, is ternately divided, and the anterior margins
of the stalks of the two lateral leaf segments are continued as
more or less conspicuous wings enclosing the adherent base of
the peduncle.

Distribution of the Vascular Bundles.

A careful study of distribution of the vascular bundles of the
leaf was made in most of the species that were available, to see

No. 483] STUDIES OF THE OPH lOGWSSACE.E 149

how far this harmonized with the theory of i\\v iciininal naiiire
of the fertile spike. The arrangement of tlie l)nn(ilcs lia> aiivady

been studied in the coninioiuT Kiir()[)eaii species, (). nibjafinn,
O. lusitanicuvi and B. lunaria. Bower has also investigated
this in O. hergianum, and more recently in 0. simplex, 0. pen-
dulum and O. palmatum (loc. cit. 1904). Of these forms the writer
has examined O. pendulum, and in addition to this a number of
other species which have not been hitherto studied.

In all of the species belonging to the section Euophioglossum
that have been examined, there is given off from the vascular
system of the rhi^conie a single leaf trace which divides at the base
of the leaf into two strands. Tliis is probably the case also in all
the forms associate.! with O. tNo/iiccanum (see Fig. 8). Accord-
ing to Praiitl. in (). I usitati iciini each of these two bundles gives
off a branch toward the a.hixial side of the petiole which unite and

150 THE AMHRICAX XATi h'ALIST [Vol. XLI

part of the petiole of Ophioglossum sp.,
X 25; A,B,at the base; C, higher up.

pass into the spike, the main
trunks passing upward into the
lamina. In the specimen shown
in Fig. 8, w^hich probably was not
the typical 0. moluccanum, while
the leaf trace divides into two
branches, as in 0. lusitanicum,
only one of these divided at the
base of the leaf, so that at a point
some distance above the base
there are only tliree bundles, two
of which are destined for the
spike. The single bundle which
is to supply the lamina is the
result of the division of one of the
two primary strands, the other half
of which forms one of the adaxial
bundles belonging to the spike.

O. MOLUCCANUM SCHLECilT.

A transverse section of the petiole in the typical O. moluccanum^
made some distance below the point of separation of the two
parts of the sporophyll (Fig. 9, A), shows four nearly equal vascular
bundles, of which one is on the outer (abaxial) side, the other
three on the adaxial side. As in all other species of Euophio-
glossum, these bundles are markedly collateral in structure. It
is probable that the central adaxial bundle is due to the branching
of one of the two adaxial bundles found near the base of the
petiole.

If a section be made just below the point where ihv two j)arts
of the leaf separate (Fig. 9, B), the three adaxial l.midlcs aiv still
recognizable, but the abaxial one has divided into several, which

taken a little higluM- up <(\ >l,ous plainly the Wa.es of the hv..
parts of the leaf. In tin- a-laxial paft. the pe.hinel.> of tin- spike,
the original tluve a-laxial l.nn.iles. are clearly evident, while in

No. 483]

SrVDIES OF THE OPHIOGI.OSSAClwE

L51

the further ramifications
of the abaxial bundles to
form the reticuhim of
veins in the leaf segment.
It is clear that in this
species three of the four
bundles of the petiole are
continued unbroken into
the spike, while only one
of these contributes to the
sterile leaf segment. This
would certainly tend to
confirm the view that the
spike is the principal part
of the leaf, and the lamina
is secondary.

The base of the spike
(Fig. 9, C, D) shows the
three bundles, but above
the base (E) these bundles
may branch, so that a sec-
tion higher up shows five
bundles. The ramifica-
tions of the veins of the
fertile part of the sj

Fig. 10 shows sections of a M'c.n.l toi-m of Ophioglossum,
collected at Biiiten:corg, cvidiMiiiy specifically distinct from 0.
molurrainnn. It wa. a plant of about \hv ^ainc ^hv, hut it .HflVml
both in ilic conlatr Mrrilr leaf and in the and ether ehar;e ters
of the spere.. It i. prol.abh- that Fi-.s.uhieh .hou. tiie extreme
lower put nf the petiole. aNo belong, to thi^ .per.i<... The l..uer
part of the petioh- ill ero.. Motion .hou . hut three l)undh- iuM. ad
of four, the nn'.hne a.hixial buiKHe l-ein- ahsent. In a >eetion
taken near the juneti..n of the spike and hiinina there were f..nr
abaxial l.tnidle." at.d li^e adaxial one.. It i. not e\a< tlv , lear

152

THE AMERICAN NATURALIST

[Vol. XLI

mary adaxial bundles, whose identity is not so clearly maintained
as in O. TnoliKcanum. In a section at the base of the lamina the

arrangement of the bundles is very much the same as in 0. moluc-
canum, and the three bundles of the spike are very similar. The
triple arrangement continues into the spike, and a section made
well above the base shows practically the same appearance.

O. CALIFORNICUM PrANTL.

0. calijomleum is a small
species from southern Cali-
fornia. In the anatomy of
the leaf it seems to follow
pretty closely the description
given by Prantl for O hmta-
nicum. A section of the petiole
(Fig. 11 , A i show^ four tuhixial
bumllcs, aii.l a >innl,. ahaxial

a medium sized specimen; Z), section of cle and lamina - Fi.U'. 1 1 , H) the
20. . , spike shows in some cases but

a single large bundle, evidently
formed by the coalescence of the adaxial bundles. There are five

No. 483] STUDIES OF THE OPHIOGLOSSACEAi 153

bundles belonging to the lamina, of which the posterior one is
apparently the original abaxial bundle, while the others are de-
rived from the two outer of the four adaxial bundles. A large
specimen which was examined showed three bundles in a trans-
verse section of the peduncle (Fig. 11, D).

Ophioderma.

Bower has shown that in 0. pendulum, 0. simplex and 0. pal-
matum there is not a single leaf trace, but the individual strands
of the petiole join the vascular system of the rhizome directly.
He also showed that the adaxial bundles which supply the spike
in the fertile leaf of 0. pendulum are quite absent from the petiole
of the sterile leaf, which in section shows no bundles at all on the
adaxial side. In the section Ophioderma the upper part only of
the peduncle is free, the lower

there are four adaxial biiiulles, '' ''

evidently the result of a bifurcation of tlu^ two w Inch arc mm'h lower
down. The three abaxial bundles remain uiu luinuvd c\('c|.t that
they are somewhat ^J^^'^T^^^^^^^^^ Ijnm.lcnuig

proicctiiiii' somcwliat froin the leaf and containing three bundles,
and the same mimbor occurs in the free portion of the pedun-
cle (Fig. 12, C & D).

154

THE AMERICAN NATURALIST

[Vol. XLI

The very much larger leaves of 0. pendulum show a corre-
spondingly larger number of vascular strands. Fig. 13, A to D,
shovi's sections through the petiole, base of lamina, and spike of a
medium size specimen. In the former eighteen bundles could be
seen, of which probably seven or eight are destined to supply the
spike. In the basal part of the lamina six or seven adaxial bundles
are plainly visible below the slightly projecting region which marks
the coherent portion of the peduncle. In both this species and
0. intermedium the free portion of the peduncle is comparatively
slender, and the number of
bundles less than in the broad-
er basal part. In the speci-
men figured there were three
bundles, of which the middle
one was evidently doubled,
and was clearly formed by the
coalescence of some of the
bundles before they left the
adherent part of the peduncle.
Higher up there were five
bundles arranged in a semi-
circle. The same arrange-
ment was found in the peduncle
of a larger specimen (Fig. 13,
E) taken from the spike which is shown in Fig. 3, D.

The complete absence of the adaxial strands in the petiole of
the sterile leaf, even at its base, is a strong confirmation of the view
suggested by both the older leaf and the younger stages that the
peduncle really extends to the extreme base of the petiole and is
joined directly to the rhizome.

BOTRYCHIUM.

The only species of liotrychium available for study was 0.
lanugimmtm Wall. ((.IIccKmI at Iloiton Plains in the uplands of
Ceylon. Tlie an aiigt-incnt <»f t!ic hiindles in the leaf of this species
agrees in the main with that of the otlier species that have been
studied (see Bitter, loc. cit., p. 458). The leaf trace divides into

Ko. 483] STUDIES OF THE OPHIOGLOSSACE.E

155

two at the base of the petiole, and these branches divide again
somewhat higher up (Figs. 14, A to C). Of the four bundles thus
formed, the two larger adaxial ones are those which supply the
spike, the smaller abaxial ones supplying the lamina. In larger

specimens of this species (Fig. 14, D), and the same is true in B.
virginianum, there may be a subsequent branching of some of the
bundles, so that a cross section of a stout petiole shows a larger
number of bundles, sometimes as many as ten.

Sections made at the junction of the spike and lamina (Fig. 14,
E) show anastomoses of some of the bundles which appear elon-
gated in section, but there seems to be no regular rule governing
the fusion of these. It is not quite clear whether any branches
are given off from the spike bundles ijito tlie laniiiia. hut this is
probably the case in regard to the two hit(>ral scpnciits of the
lamina. Within the peduncle of the >]>\kr in ihr lai-vr speci-
mens (Fig. 11, (;) th(> two original binidle.. are auain clearly
defined, but in some of the >niailer specimens these niav be com-

156

THE AMERICAN NATURALIST [Vol. XLI

Heoiinthostachys.

Farmer & Freeman (On the Structure and Affinities of Hel-
minthostachys zeylanica, Ann. of Bot. 17, p. 421, 1899) state that
in Helminthostachys there is, as in Euophioglossum and Botry-
chium, a single leaf trace which afterwards divides into several,
usually seven or eight, within the petiole. As we have already
seen, although the spike in Helminthostachys arises apparently
from the base of the lamina, in reality its origin is lower down, and
it may be traced for a long distance below the insertion of the
sterile segments.

In a section made near the base of the petiole, it appears almost
circular in outline with a ring of separate bundles. On the adaxial
side, however, there are two other bundles within the outer circle.
The number of bundles in the larger specimens collected by the
writer was decidedly greater than that given by Farmer & Freeman
(see Fig. 15, A). Higher
up the section is no longer
round, but slightly lobed,
indicating the bases of the
three branches of the ter-
nately divided lamina, and
on the adaxial side can be
plainly seen a fourth lobe,
which marks the position of
the spike. This is bound-
conspicuous bodies, the
sections of the wings that
extend down the petiole
from the lateral leaf lobes
7T ~' "• " ~= (Fig. 18.B&C). In this
region the separate bundles
of the basal part of the petiole are more or less coalescent, but the
two adaxial bundles remain separate and are those which later
extend into the spike. Still higher up the spike becomes more evi-
dent, and the two bundles belonging to it still more clearly separated.
In the free portion of the peduncle the two crescent shaped bundle

No. 483] STUDIES OF THE OPHIOGLOSSACE.E

157

sections are seen (Fig. 15, D), but it is evident that they are reall;^
composed of several coalescent bundles. A slight indication of
this can be seen also in the adherent basal portion of the peduncle.

CONCLUSIONS.

From a study of the distribution of the bundles in the leaf it is
evident that the bundles which supply the spike are not second-
arily given off from the main bundles of the petiole, but are them-
selves the adaxial bundles which can be traced
from the base of the petiole into the spike.
This would indicate that the spike is not a
secondary development upon the leaf, but is a
primary portion of it. From a study of the
earlier stages of the young sporophyll as well
as from the conditions shown in 0. simplex
and certain forms of 0. pendulum and 0.
intermedium, there seems to be little question
that the spike is really a terminal structure,
and tlie writer is inclined to believe that in all
i as(>s the spike may be regarded as the apex of
iht' leaf structure and the lamina as lateral with
regard to it. If this view be not accepted, it

of Mettenius, that the leaf is divided into two
equal branches.

In connection with the question of the termi-
°^ nal position of the sporophyll, the position of
the leaf in the embrv'o may be cited. In 0.
moluccanum — and the same is true in 0.
•'^ prdunruh.vnn .U^scrihe.l fifty years ago by
Z Mettcnins,- thc vonn.o- sporopliyte (Fig. 16)

strictly terminal organ. This (Mnl)ry() corroponds exactly to w hat
might be expected if the hypothesis advanced l)y the writer — tliat
Ophioglossum probably arose from some form resembling Antho-

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THE AMERICAN NATURALIST

[Vol. XLI

ceros — be true. This hypothesis assumes that, by the develop-
ment of a root from the lower part of the sporophyte and a
complete septation of the sporogenous tissue of the sporogonium
so that something resembling the spike of an Ophioglossum
resulted, there would be formed a plant not very unlike O. sim-
plex. We actually have in the embryo sporophyte of 0. moluc-
canum a plant which consists simply of leaf and root. Of course
the leaf is not sporogenous, but the ancestral forni must have
developed a sporogenous structure comparable to the spike before
the foliage leaf arose. The latter presumably was formed as a
lateral outgrowth of the sporogenous portion, as there seems to
be some evidence is the case in the young sporophyll of the living

The Affinities of O. intermedium Hooker.

Ophioglossum (Ophiodenna) intermedium Hook, is apparently
a very rare plant. It was originally described by Hooker from

No. 483] STUDIES OF THE OPJl lOGLOSSACE.E 159

material collected in Sarawak in Borneo. When the writer was
in Singapore inquiries were made at the botanical gardens as
to the possibility of obtaining material of this spe( i(>s, but it was
found that the original locality was lost, and the plant liail not
apparently been collected since it was first sent to 1 looker'.

The writer, however, found that tliis species had been collected
near Buitenzorg by Mr. J. J. Sinitli, of the herbarium of the gar-
den there. He was kind cnouiili lo accompany the writer to the
place where it had been (olleei(>d, and it was thus po.ssible to
obtain a fair anioinit of material uliieli was (>nough to show that
the plant is certainly quite distinct I'roni (). juudulion . of which
it has been supposed (Bitter, loe. eit., \). lb!)) that it was a mere
form, ])erhaps due to its terrestrial habit. In Hnitenzorg it grew
in a j)lantati()n of bamboo u.sually in the accumulation of
huinus and earth about the roots of the clumps of bamboo. It
is a small plant (see Fig. 4) and in its still' ti])right habit and nnich
longer peduncle presents a very diU'erent appearance from any
form of O. pendulum — althoniiii it is evident that it !>elon-^ lo

tuberous body a])parenlly deveioix'd a^ a root bud ■ Vv^. I, W. \\)

it approaches O. .r, u itli which it nun be pretiN .•k.^ely allied.

It <litrers. h<.w,>ver. in other respects than that <.f it. habit, from
O. pcinltihtiii The spores i !• ig. 17. !> ) are d(>cidt'dly smaller
than those of (). pnidnhim, aii<l the delicate reticulate markings
of the epispore (Fig. 17, K) are very ditlVrent from the markings
in the latter species.

POLYGAMY AND OTHER MODES OF MATING AMONG
BIRDS

R. W. SHUFELDT

For several years I have devoted much time to a study of the
phenomena of sex in vertebrates, comparing those of the lower
forms with the features presented by man. IMuch of the matter
thus obtained is now in the publisher's hands but some of it is
presented here.

The nature of man, .his customs, habits, and institutions, his
mental and physical characteristics cannot be fully and intelli-
gently understood unless all of our stock of accumulated facts are
studied in the light of w^hat we know to obtain along the corre-
sponding lines among all animals below man. That is to say, it
IS simply impossible to comprehend the morphology of man, unless
our studies of it are made comparative with our knowledge of the
anatomy of all other animals. So too with all else manifested
on the part of our species;— to get at the origin of all things in

to trace them dow^n through the x alc <>t' hvinu' forms hdow him.
It holds in our researches int.. the >vwurc cf x.ciety, ami it was
Letourneau who sai.l "Wlien once i( is estal.hshnl that man is a
mammal Wkr any other, ami only .h.tingni.lied from the animals
of this class l.y a greater cerebral development, all study of human
sociology nnist logically be jH-ecetled by a corresponding stialy
of animal sociology. Moreover, as >ociology finally depend^ on
biology, it will be necessary to seek in physiological conditions
themselves the origin of great sociological manifestations." '

162

THE AMERICAN NATURALIST [Vol. XLI

and derivation, I attempted to bring together what I knew of the
matter of mating among animals generally, carrying my investi-
gations into the various groups of fishes, reptiles, birds and mam-
mals. It is a very well known fact that with respect to our own
species, we meet in one part of the world or another, people who
practice every form of sexual relation, to say nothing of what is
met with along the lines of pervertism in such matters. Even in
the United States, we meet with any number of cases of marriage
devoid of all ceremony (anarchists); of free love; of monogamy;
of promiscuity; of polygamy and bigamy; of legalized concu-
binage (South Carolina); and of the divers unnatural relations
of the sexual perverts and inverts. Polyandry, that rare and
exceptional conjugal form, where the one wife has two or more
husbands, has never been instanced among us, so far as I am
aware. No such sexual association is met with among mammals
below man, and never among birds.

It is in this latter class of vertebrates that we meet with some
of the purest types of, as well as some of the most interesting
examples of the conjugal relation, and it is to a comparative con-
sideration of some of these that the present article will be devoted.

In reviewing the material for this purpose at hand, I have
drawn largely upon my own ornithological observations and
studies extending over a period of forty or more years. Then I
have consulted such works upon ornithology as I find in my
private library. With respect to the latter, I am obliged to con-
fess my surprise at the inadequacy of the accounts, and the marked
variance often exemplified in the statements of different authors
of recognized standing and reputation on the subject. Very few
books at my command pretend to make any comparisons between
the mating habits of })irds and the marriage customs of various
peoples, but there arc a iVw .

Beyond the matter of llic dillVivnt [.roccdiiivs of courtship in
the case of l)irds, tliciv arc no furtlicr (rrcinonials with them as
in tlic cast' of many, iiKh'cd, the niajority of the races of mankind.
So that, in the abstract, polynaniy in birds means exactly the
same thint;- as Imiium j>oly(iamy, and so on for monogamy, pro-
miscuity and other practices. Taken in the abstract, and barring
opinions to the contrary, many believe in the case of man, that

in prehistoric time, when lie was first differentiated
stock, he, wherever existing, was given over to ii
miscuity; that this was soon followed in many ic^i
form of polygamy, and polyandry where wonicii w vw
as promiscuity disappeared, and polygamy hc-canic f; ■
lent, some mode of monogamy ap|)earc(l. and this, a'
time is the form of marriage adopted In iicaily all ci
In other words these various customs havi' sliailcd iiif
— that is, in the main, promiscuity for ilic wild, pivlii^
followed l)y polygamy for ancient times, with moi

polygamy now o])enly followed in the I'nited State"
of the lowest existing races of the world are moiiogamc

These facts are thus briefly ])fesent('(l in that we i-
them with what occurs in the . lass of hirds. TIkv
one way, the lowest form, of exi^iiiii; l.inU dio,
mating be given over to pronn'scuity ; iIiom' higher
should be polygamous; and, hnally the mo-t ■
as the Passeres, be monogamous. 'This. ho\

while other, perhap., atv pn.mi>cn..u. .no l.inU
androus); atid .till oiIht. afroidiui: rxamplc. of
it Ls too, higher n,. in tl.r ..aic. jiim a. il i., a. l.rfo,
with the human spec io.

Tracing birds back thronuh ucolonic time a. boi
means of the niaI<Tial at hand thei'c i> no ()Mf>tioii

families of birds from vai'ions parts of (he world f\l

164

THE AMERICAN NATURALIST [Vol. XLI

way of characters) of their reptiHan relationships, are distinctly
more closely allied upon that account. ]Many taxonomers, how-
ever, have thought so; and have endeavored to show that all
existing true ostrich forms, the Kiwis, an(l tinanous are a sort of
modern affined struthious types. On the other hand a Kiwi
(Apters^x) is no nearer an ostrich, and an ostrich to a tinamou,
than a limpkin (Aramus) is to a bustard, and a bustard (Otis)
to a quail (Colinus). Therefore it need not surprise us, in view
of all that has been set forth above, that the various modes of mat-
ing of any of these birds should be entirely different, or that these
modes should fail to throw any light upon their affinities. For
a moment then let us see what some authors have to say in regard
to the mating of ostriches and their allies.

Professor Newton, quoting Lichtenstein, says: "Though some-
times assembling with Zebras or with some of the larger antelopes,
ostrichs commonly, and especially in the breeding season, live in
companies of not more than four or five, one of which is a cock
and the rest are hens. All the latter lay their eggs m one and the
same nest, a shallow pit scraped out l)y their feet, with the earth

more it would seem to uiianl their coiimion treasure from jackals
and small l)easts-of-prev than diiveflv lo lorward the process of
hatching, for that is otteii lett wliollv lo the suti." ^ From this
it is clear that the African Ostrich is a polygamous l)ird by nature.

The Rhea or South American ostrich (Rhea darwrni, amcricana,
etc.) IS also undoubtedly polygamous in nature, while the emeus
of Australia are said to l)e monogamous, though neither Newton
or rvciaft -av aiivtliing on this j)()int. Neither do they give us
aii\ iiitoniiatioii on this [)omt in regard to the cassowaries, birds

No. 483]

MAT IXC AMOM; lillUh^

165

more or less (•los<>ly allic-d to the (Mueus. Indeed, I am unable to
state whether a ca^-owarv is, l.y nature, polygamous or monoga-
mous. Their (^ggs have dcscrihed l)ut aj)])arent]y not their
mating habits. None of the above-named wrilcrs describe the
breeding habits of the kiwis i A p/rri/.r oin tii, iiiiinh//i and aiis-
trah's) and I am unable from jKM-sonal observation to state whether
they are by habit monogamous or polygamous , >ee Sir W aher
BuUer, Newton, Pyeraft, and other writers), 'idiese curious birds,
now being rapidly exterminated, are probably moiionaiiions. a^ Dr.
Claus says of them, "The kiwis are noctiu-nal \nv>U. w hich by day
remain concealed in holes in tlie eartli and n(> at night to seek
their food. They feed on insect-hir\ a- and worms, live in pairs,
and at the breeding time, which seems to come twice in the year,
they lay, in holes s('rai)e(i in the earth, a sirikinnly kirge egg,
which, according to some, is incubated l>y the femah\ and accord-
ing to others by the male and fenuile in turn.'" '

So far as I have been al)le to ascertain, the tinan)ous (("lyp-
turidje) are monogamous birds, while they a»ociate togetlier in
flocks during those times of the year when ihey arc not l)reeding.
Newton does not mention this in the " I )ictionaiy." and at this
writing I do not hapj)en to ha\c Hartlftt"s paper at hand (P. Z.

AH water birds of the main groups ai)pear to be monogamous
in the matter of their mating. There appear to be no excej)tions
to this rule to be met with among the several suborders of the
Pygopodes, Impennes, Tubitiar(vs, Steganojxxles, Lonijipemies,
Alcae, and the Chionides. .Vs we know, these oi',,ii|» ((.main
the divers, the penguins, the petrels, the p(-lican-> and \arious
allies, the gulls, and the ank tribe. Nearly all tlioe forms are
low in the scal(\ and in all w(> meet with near relatives among

166

THE AMERICAN NTURALIST

[Vol. XLI

plovers, the turnstones, the surf birds, the snipes, the phalaropes,
the avocets, and the ja9anas, the entire host being monogamous
by habit, with but one famous exception, namely, the truly polyg-
amous ruff (Machetes pugnax). The peculiar habits of courtship
and breeding practiced by this species have been well-described
by a number of continental naturalists/ Among the Limicolse
there "appears to be, among existing birds, but one other species
suspected of being a polygamist, and this is the double or solitary
snipe (Scolopax major) of Europe. Newton does not mention the
fact in the "Dictionary," but Darwin remarks in "The Descent
of Man," that "some of the above birds, — the black-cock, caper-
cailzie, pheasant-grouse, ruff, solitary snipe, and perhaps others,
are, as is believed, polygamists." (p. 406.) From all that I can
gather, it would seem that the question has not yet been decided.
Coming to the Cursone, the group contains but few species that
I know of, that have been suspected of being polygamists and
among these is the great bustard (Otifi tarda), — and with it
most of the evidence seems rather to point to the fact, that such
is the case. Whetlier any other representatives of this somewhat
numerous group (Europe, Africa, Asia, and Australia) are })olyg-
amous by habit, I am unable at present to say. However, the
birds called 'floricans' of India, closely allied species to the
bustards, are reported as practising ])(>lygainy. There seem to
be two known s[)ecies of these, -the Bengal { Si/ p/irol idr.^- hr>i-
fjalensis) and the lesser floriciui (N. auritin. 1 )iiriiii:- {tairinu'
season the two sexes live apart in groups, and in mating conic
together, and "when a male wishes to attract a temporary part-
ner, he does so by going through an elaborate series of perform-
ances."' It is possible that all the true bustards possess strong
inclinations in this direction, even if they are nor actually [polyg-
amists. Not so, however, with the stone curlew <K. n; j>d<tns^
a species I have relegated to the (/ursonc iiiiliouuli, 1 by no

' l.ydekker. R. The Roi/al \ot. lli^t. p. loS.

^iShufeldt, R. W. "All Arrangement of the Fmnilies and the Higher
Groups of Birds." The Amer. Nat., Vol. 38, Xov., Doc, 1904. pp. <S33 -857.

Xo. 483]

MATING AMONG BIRDS

167

Monogamous matings seem to be the rule with all the cranes
and rails, with their allies, near and remote.^

Probably no group of birds in the world's entire avifauna have
l)een more closely studied or had more w-ritteii about thcni than
the great gallinaceous group of fowls, includino' aiiionu' ilicin not
a few other such familiar birds as the turkeys, the uunica fowls,
quails, partridges, grouse, pheasants, and their various allies,
near and remote. Good and sufficient reasons there are for this,
as a very large number of them are, and have been, long domes-
ticated, as the chickens and turkeys. All of them constitute
game in every part of the world; while many of them are kept
in zoological gardens and private preserves, as the j)heasants
and others. None of the (Jalliformes, I believe, are polyandrous,
though many of the families are curiously divided up between
polygamy and monogamy, some being strong adherents of the
first-named ])ractice, while others, under no circumstances, depart

168

THE AMERICAN NATURALIST [Vol. XLI

reversed in the little common quail of the old world {Corturnix
communis), a well-known polygamous species, where the males
are both larger and handsomer than the females.

As to the Megapodes or brush turkeys (Megapodidje) of the
East Indies and Australia, none of the writers at hand state whether
they are polygamous or otherwise.

The habits of these birds are pretty well known, especially their
burying their eggs in immense mounds which they build, or con-
cealing them in sand-holes and burrows, in either situation they
hatch out by the sun and the heat of the fermenting vegetable
matter in the mounds. The young fly an hour after they are
hatched. Wallace describes several species of them in his "Malay
Archepelago," but does not state whether they are polygamous
or not, and neither Newton or Pycraft have anything to say upon
that point.*

Most ornithological writers lay it down as a rule that among
the Gallinre generally, where the cock bird is evidently larger than
the hen and its plumage is remarkably conspicuous, the hen,
being more or less plain in this particular, the species is polyga-
mous, whereas, when the sexes are nearly alike in point of size,
and but little difference in plumage, they are almost certain to be
monogamous in their mating. There are, however, a few excep-
tions to this rule.

Personally, [ liavc never stiulied the ciirassows and guans
{Cracidcc) in their native hannts, and therefore eainiot say, from
my own experience, anything in regard to their mating habits.
In this group, I take it, the curassows of South America are prob-
ably monogamous, as is likewise our Chachalaca (Ortalis v. mac-
calli), though in the case of the latter species, where the sexes
are nearly alike, few American ornithologists describe its court-
ship and niatino'. nctal.ly Hendire. Cones, Ui-lgway (Alannal),
and others, while continental writers rarelv refer to' it. Neither
Audnhon or WiKo.i ever saw \hr bird.

Finally, the >ul.ord.'r (iaiiin.-e i. >een to contain live very ele-

>Loc. cit. Art, ■■.M.-a|HHl,.- l>t. ii, p. r,:VJ. and I'v.ial!. ■ l.ivin- Animals
of the World" Lot.d. p. 411.^ One writer states that several hen niesu[)()des

No. 483]

MATING AMONG BIRDS

169

gant families of birds, representatives of which, in more or fewer
species, are found in all parts of the world. These are the pheas-
ants (Phasianidfe), the grouse (Tetraonidse) , the American Par-
tridges (Odontophoridae), the Guinea fowl (Numididcie), and the
Turkeys (Meleagridai). Great is the wealth of species in the most
of these several families, and while some of them are polygamous,
others are strictly monogamous, and the habits of any of tlicm
may be changed through domestication, and tluy soinctiines
infringe upon, or even break, some of the rules oi\cn in tore-
going paragraphs. Included in their ranks are all of our eounnon
domesticated gallinaceous fowl, and occasionally the habits of
some of these are very remarkable.

Very much do I regret that I cannot give more space to this
group as it is both an interesting as well as an inij)ortaiit one;
moreover, authors are by no means unanimous in their opuiKuis
in regard to the modes of mating, and in the case of xtnie species
we have apparently no data at all. Considerable part of the
literature has been carefully looked up by me. No one sct ins to
question but what such species as the capercailie and black urouse
of northern Europe are polygamous. Pheasants an<l their near

but in this I may be wron.y-. The wild tnrkens of North Ain(>riea
are also polyganiists. thoiio-h it is said that the oM males i^-enerally
have a favorite hen.while the other females he favors are but his
concubines. Peacocks are polygamous but the \arioiis species
of Guinea fowl are eminently monogamous. When the latter
are domesticated, however, as vast numbers of them are, I have
personally known a male Guinea fowl to take charije of six or
seven hei'is. and the latter would all lav the usual nund.er of e,u-s
and brino- forth their youno-. From all I can -ather, it ha. been

monogamous. This .eems toVe the ca^' too. with tlie l.ir.l> we
call quail (American partridnvs: ( ),lo.itoi.liorid,c i. ihou::!, 1 am

our typical o-n.use iCa'nada. dusky, Fra.d<lin-s a>.d others!,' the
sage cock, howev(>r, i.> polygamous i ( Vntrocercus

Audubon, wluxse life-histories of our game birds are so thorough-

170

THE AMERICAN NATURALIST

[Vol. XLI

ly unsatisfactory^ in his account of the mating of the pinnated
grouse (Tympanuchus) gives one the impression that he beheves
the bird to be monogamous, while in his account of the ruffed
grouse (Bonasa) he states in referring to the latter species, that
"The males have the liberty of promiscuous concubinage, although
not to such an extent as those of the pinnated grouse."^ Bendire,
on the other hand when describing the habits of the ruffed grouse
(B. umhellus) says, "By many persons the ruffed grouse is consid-
ered polygamous, and while I can not actually disprove that
assertion, I doubt it very much."^

Again authors are at variance in their opinions with respect
to the several species of the sharp-tailed grouse (Pediocsetes)
^nd E. T. Seton, quoted by Bendire, says of the prairie sharp-
tailed grouse in describing the remarkable dance of the males, "Its

erratic character can hardly be questioned The whole affair

bears a close resemblance to the mancKuvring of the European
ruff, and from this and other reasons I am inclined to suspect
the sharp-tail of polygamy."^

The curious hoatzin of tropical South America (( )pisth()comus)
in a way related to the (iallina?, is said to be polygamous, but as
yet we stand quite in ignorance of some of the habits of this inter-
esting form in nature.

Sand-grouse f Syrrliaptcs) and their kin 1 believe are nionoga-
moiis, and I tlo not at tins writiiin- i-ccall any species ol' wild pigeon
(Colinnl.it'oniiesi that has anv other fofni of mating in the breeding

No. 483]

MA TJXG A MOXG BIRDS

171

nogamous, and I believe tlie entire swan, ooose aiul <lnok tribe
(Anseriformes) are, — at least in natnre, although there may be
exceptions to this that I either do not recall for the moment, or
have not come to my notice. When domesticated, however,
ducks may become highly polygamous, and it is a well-known
fact that in this state it is not difficult to cross various species and
rear interesting hybrids. Cases of this character are reported
by Darwin, who states with respect to birds that "In several
groups I have not been able to discover whether the species are
polygamous or monogamous."^

"Very peculiar fancies," says Letourneau, "sometimes arise
in the brains of certain birds. Thus we see birds of distinct spe-
cies pairing, and this even in a wild state. These illegitimate
unions have been observed between geese and barnacle geese,
and between black grouse and pheasants," and further, when
(juoting Hewitt from Darwin as to how a ( i t 11 1

duck threw over the male of own species and deliberately courted
a male pintail that had been placed in tlie water with lier, mated,
and would have nothing further to do with the nialhird. he says
"that conjugal fidelity docs not always resist a strong;- ini|>res>ion
arising from a chance encounter; that novelty has a disturbing
effect; and, finally, that inditrerencc and coldness can rarely hold
out against the jxTsistcnt ad\anccs of one who loves anlcntly
enough not to yield to discouragenicnt. Dante ha- already made
this last reflection in his celebrated line -

'Amor cha iniH' aniato aniar penlona.'

To (jiiote Dante a pwpn.s of the illicit amours of a j)intail and
a wikl duck may shock the learned, but the aptness of the (pL.ta-

human organisms." '

Polygamy is not practiced, so far a-^ 1 am aware, by any of the
flamingoes (rha'in'copt<"ri i, or rc[)rc>cniati\c> of ilic crane— fork
assemblage (Herodiono or the diurnal Kaptorc- including
all the vultures (Ac(i|)itrcs

'Loc. cit. pp. '219. -J IS.

172

THE AMERICAN NATURALIST [Vol. XLI

the owls (Striges), the Caprimulgine forms (Caprimulgiformes).
None of the Coraciee (rollers, etc.) I believe are polygamous, or
the kingfishers (Halcyoniformes), or the Bucerotes, or representa-
tives of such suborders as the Upupse, the Meropes, the Momoti,
or the Todi, but when we come to the humming-birds (Trochili)
some authorities still seem to be in doubt, and no less a distinguished
ornithologist than Mr. Salvin told Darwin that he was "led to
believe that humming-birds are polygamous," ^ but, the present
writer by no means entertains any such an opinion.

Comparatively speaking, very little is known of the courtships
and matings of the Jacamariformes (jacamars and puff-birds)
and the Trogoniformes (trogons), but I believe none of them to
be polygamous in their habits, although if found to be so it would
in no way surprise me, on account of the relations of the latter to
the cuckoos.

When I say this I do not mean to imply that any of the cuckoos
are strictly polygamous, and no writer seems to be perfectly cer-
tain on that point. What the mating habits of the touracos (Muso-
phagidie) is like, I am, at this writing unable to say, but it is very
interesting and important for us to know. Those who have had
opportunity to study them have, as in so many instances in or-
nithological history, overlooked all this. The literature upon the
nidification of the cuckoos (Cuculida') would make many volumes
so it is quite unnecessary' to dwell upon it here. Their depositing
their eggs in the nest of other birds is simply parasitism, and for
all I know to the contrar}^ the European cuckoo may be the
veriest polygamist in the world's avifauna, and the same is true
of others of his kin that follow the same practice. It is not likely
that these birds are monogamous, it being far more probable that
they follow some form of promiscuity, or where there is a scarcity
of mal(>s, even ])()lyiin(lrv ? All these remarks likewise apply
to oiir (ou hirds ( Molotlini-. i of the rasxTiformrs, birds which I
am <|iiit(' sure from pcisoiial Dbscrvatioii may he cither polygamous,

eastern form, the ruby-throat, nor in any of those I have had the opportunity
to study in the west.

No. 483]

MAr/.\(; AMOXG BIRDS

173

it is only through the force of circumstances that \)\ViU aic ever
the latter, as some seem to contend.

Some of the breeding habits of Cuckoos in vai ions part^ of the
world are truly remarkable, as witness tlio^e of thi^ country
(Crotophaga, Geococcyx and ( 'ocevzn-^ i. It wonM appear, from
what we know of its habits, that our Ani> may be strictly i)olyg-
amous (Crotophaga), inasmucii as several females of this species
all lay their eggs in the same nest, — but even so, they may be the
mates of different males.

There are some wonderfully interesting questions that arise,
when we come to study the courtships, mating, and m'dification
of the cuckoos, cuckoo-like birds, and the cowbirds, and especially
when we apply this knowledge, in a comparative way, with the
customs followed by our own sj)ecies. Space, or rather its limi-
tations, will not admit of my diM-ns.iu- any mk Ii matter here.

that will take fully into .-ouMdlTat ion> all >u<'li !,ne>ti..ns, and
where suflicient data is av;ulal.le. end.-avor to throw >ome liirht

174

THE AMKRIi J .V AM 77 RMJST [Vol. XLl

Cypseliformes, and the Eurylsemiformes, although I know of no
species or family among these several groups that are not strictly
monogamous by nature, while they may differ very widely in
their habits of nidification. Uiifortuimtely. we sti]| know very
little about the life-histories of the iyiv-l)ii(l^ of Aii>trah'a (Me-
nura), and some naturalists believe tiieni to l)e polyginious.
Again, Darwin quoting Lesson says "that l)irds of paradise, so
remarkable for their sexual differences, are polygamous, but
Mr. Wallace doubts whether he had sufficient evidence."^

In closing this article it is well to note that what T have set forth
in it has probably long been known to the niajorily of oeneral
and observing naturalists of each veneration, bnt not >o to the
average reading pul)lic, and, unfortiniatcly lutt to a i;i<-at many
people to whom the knowle.jov would be of con^idt^rabh- interest
if not of })ositive vahie.

It is clearly shown that birds, as a Chis. anion- \-erU'brates, in

more easily ..xphiinc. L That thr s-iti>fa< tioii of the ..'xnal instinct
and the (■(|iiaiiy imperative demand, on the part of nature, that
the species he j)erp<'tuated, if po>>il,h', i. the e-^-ential part of the

iar with general biolooy, and the i>a.t an.l pn-ent life histories of
anhnals on tlii> plaiHl, woid.l for an instant claim that any of these
mating habits in bird> ^^■vvv of a criminal nature. It i> oidy the

No. 483]

MATING AMONG BIRDS

175

ignorant, the superstitious and narrow-minded who entertain such
views. We have plenty of storks, black grouse, and even Euro-
pean cuckoos and American cow-birds among our own species,
but the significance of all this, and its biological importance to
our kind, I shall endeavor to point out in another connection

ON THE WOOD RAILS, GENUS ARAIVIIDES, OCCUR-
RING NORTH OF PANAMA

OUTRAM BANGS

For many years I have been gathering all specimens I could
of the splendid, great Wood Rails of the genus Aramides with the
hope of some day monographing the group. Unfortunately I
have as yet been unable to bring together sufficient material from
South America to attempt to include in review the forms of that
country. I now have, however, a complete set of the species and
subspecies of Middle America from Panama north to the northern
limit of the genus in southern ^Mexico. A critical study of this
material together with a number of skins kindly lent me by the
United States National :\Iuseum, the American IMuseum of
Natural History, and the Bureau of Biological Survey of AVashing-
ton, which include the types of Aramides flumheicollis Zeledon,
A. axillaris Lawr. and A. albiventris Lawr. has induced me to
publish now a short synopsis of the forms of Aramides occurring
north of Panama.

My views expressed in the following pages will be found to differ
a little from those of recent authors, such as Sharpe in Yoi. XXIII
Catalogue of Birds in British Museum 1894 and Biologia Centrali-
Americana, Aves, 3, 1897-1904, and I describe as new one form
from ]\Iexico, allowing to the region here treated three species
and two additional subspecies.

In all species of Aramides the sexes are alike in color and there
are but slight individual or seasonal differences, apart from those
caused of the w^holly mechanical processes of fading and wear.
Some species have a juvenile plumage, still worn when the bird is
nearly full grown, that is quite different in color from tiie livery
of the adults — A. axillaris and its allies. Other species, appar-
ently (I have seen but one young individnal of .1. alhlrnifris
phnnhrlroUis, and none at all of the other subspecies ni alhlrrnfris

tinctly difi'erent in color from that of tiie adults.

177

178

THE AMERICAN NATURALIST [Vol. XLI

If the specimens examined by me are correctly sexed, there
is also no average difference in size between the sexes in any of
the species or subspecies. All, however, vary much in size indi-
vidually, in fact to a deorcc 1 am ^^•h()lly unable to account for.

KIN(; NOKTIT OF PaXAMA

A. Sexes alike in color.

1. Under wing-coverts banded black and white

A. axillaris Lawr.

1. Under wing-coverts banded l)lack and cinnamon-rufous or

2. Back of head, between gray forehead and gray neck not
distinctly chestnut, but grayish-brown or l)rownisli-gray

A. cajanca (Miill.).

2. Back of head distinctly chestnut 3.

3. Back concolor, olive 4.

3. Back not concolor; olivaceous-tawny anteriorly, olive pos-
teriorly . . . A. alhivcniris flumhei colli s (Zelcdon).

4. General color paler; light colored crcsc(Mit around the bhu-k
belly j)atch, ver\' wide, white

4. General color darker; light colored crescent around the
black belly patch, narrow, fulvous

A. alhiventris mexicarms nobis.

. 1 ramidrs a.rll/ari.s- I .awr. Troc. IMiii. Acad. p. 107, bSGS. Sharpe
Cat. Bird^ Hr. Mii^ 23, p r.C. n\n\. (Vnt. Am., Aves vol.

3, p. 31S, pi. LXW ll, is'.i: I'lOl.

Typk Uo<m.itv. IJarniM.nu-lla, ( '< .lonil.ia. Type, now No.
45655, Ameri<-aii Mu>fu f Nat. Hist., New ^'oI■k. exainii.ed.

It iia> been .upp«,MMl that tluMV ua. a bivak in tli.- ranuv of
this species and that it did not occur in southern (Vntral America
south of Honduras. There is, however, in the Cndcrwood collec-
tion, lately purchased by John E. Thayer Ivscp. a young example

No. 483]

WOOD RAILS NORTH OF PANAMA

179

of A. axillaris, nearly full grown but with ihv niulcr j)aits still
brownish slate-color, from Costa Rica. Uiifortiiiiatcly the label
bears nothing more definite than "Costa Rica." While certainly
very rare in southern Central America, 1 still believe .J . axillaris
has a continuous range. It is a rare species in northern South
America, and seems to be nowhere so plentiful as in the region
lying between southern Mexico and Honduras. At all events I
can detect no difference between northern and southern speci-

Characters. Size small; bill short; under wing-coverts
banded blackish and white; neck and head, except throat, ru-
fous-chestnut; a conspicuous gray patch occupying upper inter-
scapular region and lower hind neck.

Color, Adult plumage. Throat white; head, neck and
breast bright rufous-chestnut; upper interscapular region and
lower hind neck gray (about slate gray); back, wing-coverts
and wings except primaries and secondaries, olive; rmnp i)r()wn-
ish black; tail and upper and under tall-coverts, black; belly
and thighs slate-color; ])riuiaries hazel; secondaries also hazel
but duller, more dusky toward ti})s; under wini; covert> and
axillars banded black and white; bend of wing an<l tips of axilla rs
usually banded black and ha/el; "tarsus vermilion; b<>ak un'«'ii,
basal portion yellow; iris brown."'

Young differ from adults in having the neck and muler parts
dull slate-color, and the characteristic gray patch on lower hind
neck and upper back less distinct though still evident.

180 THE AMERICAN NATURALIST [Vol. XLI

105554 yg. British Guiana.
16375> yg. Costa Rica.

168. 53. 51. 39.
156. 50. 51. 40.5

Remarks. A. axillaris is a small species with a short bill,
related to A. mangle (Spix) of Brazil of which it is the northern
representative. Judged by the few specimens I have been able
to examine I should think the two were specifically distinct.

A. axillaris is very different from any species occurring in the
same region with it, being at once distinguished by its small size,
black and white under wing-coverts and red-brown neck. It
does not appear to be subject to any geographic variation —
southern and northern examples being, so far as I can see, quite

Aramides cajanea (Miiller).

Fulica cajanea Miill., Syst. Nat. Suppl., p. 119, 1776, based on
Daubent. PI. Eul. pi. 352.

Rallus chiricote Vieill., N. Diet, d' Hist. Nat., 28, p. 551, 1789.

Aramides cayanea Sharpe, Cat. Birds Br. Mus. Vol. 23, pp. 57-
58, 1894.

Aramides cayanea subsp. A. Aramides chiricote Sharpe, Cat.
Birds Br. Mus. Vol. 23, pp. 58-59, 1894.

Aramides chiricote Biol. Cent. Am. Aves, Vol. 3, p. 318, 1897-
1904.

Type Locality: Cayenne.

Geoc/rapkic Distrihution: Tropical America in general from
northern Brazil nortli tlirough Panama and Chiri(iui to the Pacific

allies ()ccii|)y an intciiiKHliatc ])()siti()n in the genus, being much
larger tluiii .1. (txillari.s and allied species, but inferior to the
gigantic .1. i/jxicalid (Nicill.i of southern South America); bill

head gray, di.lln-, browner on oeejpnt; i.aek coneolor, olive;
breast, coneolor bright, deep, I'eddish tawnv.

gray (nearest slate-gray) this color sometimes extending a little
> Coll. of E. A. & O. Bangs, Boston.

No. 483]

WOOD RAILS NORTH OF PANAMA

181

onto mantle, darkest on forehead and palest on <•
put darker, duller, often brownish — grayish-hn
gray or grayish-olive; back and wings, exoej)! ))rii
ondaries greenish-olive; primaries and srcoiulanc
dusky olive toward tips, the outer sccoiKlaiitvs
edges, and inner secondaries mostly dlivc; nun
suffused posteriorly with deep reddish olive; iipj
and tail black; breast deep reddish tawny, \ar
tone (with season or age?), sometimes toward h
toward orange-rufous; belly and under tail covert
slate color; under wing coverts and axillars ban
hazel; "tarsus poppy-red; terminal part of bill g
yellow; obital ring red; iris red, sooti after (ha

15408
16536
9942^
9943
17940'
148191
7060^
7650

182

THE AMERICAN NATURALIST

[Vol. XLI

Remarks. After very careful comparison of a large number
of specimens I fail altogether to make out a subspecies, chiricote.
I can find no constant differences whatever between skins from
Brazil and Surinam on the one hand and the most northern exam-
ples from Chiriqui and Costa Rica on the other. Indeed Sharpe
in Catalogue of Birds in the British Museum (Vol. 23, pp. 57-59)
does not assign any well defined geographic distribution to the
two subspecies he recognizes. The patch of a duller color on the
occiput varies considerably in shade — with season I think, that
is with the condition of the plumage, whether fresh or much worn.
Different examples from Panama and Costa Rica differ quite as
much in respect to the shade of color and distinctness of this mark-
ing as do any two that can be picked out from the northern and
southern parts of the range of the species.

Inhabiting the Pearl Islands in the Bay of Panama is a slightly
paler and slightly smaller race of this rail. Tlie four examples
taken there by Mr. Brown cannot quite be mutchcd l)y continental
specimens, but the differences are too slioht and in this genus of
two unimportant a nature to base a subspecies uixm.

There appears, however, to be in Bra/il a well nuirked sub-
species, the exact range of wliicli I am tliroii^'li want of suflicieiit
material unable to define. Skin "f" of Sliarpe's list in ( atalo.une
of Birds, belongs to this form (see footnote, p. .')S i and there is one
skin in the National Mnsenm, No. lMIlM from fSt. Catliarines ?)
Brazil collected by Lemuel Wells, that appears to at;-ree exactly
with Sharpe's Rio de Boraxudo specimen, differing from J. caja-
nea in being mostly gray above, the gray of the upper neck per-
vading the entire mantle, the wing coverts alone being olive and
these paler and decidedly more grayish olive than in .1. cajanea;
the rufous color of under parts, as pointed out by Sharpe in his
specimen too, is also paler. This bird is not (Uillhnila rn/irrps
S]>i.\, which, judged by the plate, is true A. cajdiira, an<l un-
.loubtedly rep'resents a' vali<l form.

Another peculiar individual is a verv old skin in the Nalional
Mn.emn. no. LltOT, lab.'led liueno. Avre>, .] . K. 'r..wn<end
In color this example agrees with true .1. rajanra except in having
the rump nearly wholly dark reddish olive.' It i>, h(.w(>ver, very
1 icl 1 nth proportionally shorter tar>us and bill, the wing^.

Xo. 483] WOOD h'MLS SORTU OF /M.V.I. l/.l

183

considerably worn at that, measuring, 200, tail S('», t;ir>ii~> 7.1.
culmen 55. It may represent still another suhsjxH ic^.

Thus while in the northern part of its ran^v Iin.ih noitlicin
lirazil nortli, this rail does not vary to any extent with ocouraphic
areas, there seem to he in southern Soutli America si'veral uco-
oTaphie forms.

Aramidcs cajanea is another very (hstiiict sjx'cies. nearly allied

distinguished by its shorter, thicker hill, and duller color of the
occipital region, which in J. alhi roiiri.s is always hri-i-ht chestnut.

■ ArAMIDES ALBIVKXTKIS AIJUVKNTRIS Lawr.

Aramidcs cdbhrntrh Lawr. Troc. Phila. Acad., ji. 2:]4, 1867.

Aramidrs raifonra, sul)sp. B. Araninirs alhirrnfris Sharpe, Cat.
Birds. Br. Mus. \o\. 23, 1S04, pp. olMlO.

Aramklcsalbivnifr>\'\Vio\. ( ent. Am., Aves, A ol. :!. p. :nO, 1807-
1904.

Type Locality; Britisli Honduras, 'lype, now Ameri-
can Museum of Natural History, exann"iied.

GEOCRAniic DisTHiin tion: British Honduras and Vucaian,

Ci1aha( TKHs: About tlie size of .1. rajaiira or sli-luly iai-er;
bill lonoer and more slender than in diat spe<MV>; all the <'oior.
pale; a la rue. conspicuous patch of briu'ht chestnut ext(Midin,-

wing covert, 'han.led Mack and pale hazel.

Color: Thn.at dull white, this color extending well down
under surface of neck; on the occiput, exteudin- backward to

cl.esttmt; rest ..f neck and head -ray iabout Cray No. (1 I>t Bid-

interscapulars as \\ell ochraceous-rufous more or less mixed witli
olive, this marking usually very conspicuous, thou*:li ne\er form-
ing a complete mantle across back as in the soutlu>rn snbsjiecit^s
plumbeicollis; primaries and secondaries, light, bright hazel;

184

THE AMERICAN NATURALIST

[Vol. XLI

rump black, somewhat dusky olive anteriorly; upper tail coverts
and tail black; breast pale tawny-ochraceous becoming ochra-
ceous-buff posteriorly; a wide crescent shaped marking of white
or sometimes cream-buff, around upper part of black belly patch;
belly and under tail coverts black; thighs slate color; under
wing coverts and axillars banded black and pale hazel, the tips
of the feathers sometimes buff.

British Honduras.
Belize, British Hondu

130327 — " 177. 58. 80. 65.

148192 — " 184. 62. 77. —

15246^ c?ad. Yucatan, Rio Lagartos 177. 63.5 74. 62.5

33668^ — Guatemala, Chiapam 187. 62. 78. 66.

42777 — Central Guatemala 176. 58. 75.5 66.

Remarks. Typical A. alhwentris occurs only, so far as I know,
in the coast region of British Honduras and Yucatan, and in its
very pale coloration parallels other bird forms of the same region
such as the clapper rail, lately named Rallus pallidus by Nelson.

Two specimens from Guatemala I refer here, though they are
intermediates, between A. alhirnitri>f alhirnitri.'i and A. alhirnifri.s'

Yucatan and British Honduras {'xaiiiplcs, the other from (liiaj)ain
on the Pacific coast being more like the Mexican bird. To the
northward true A. alhiventris is replaced by a darker form with
less distinct and more fulvous crescentic marking on the belly,
that occupies southern Mexico and that I have named below as a
new subs|)eci<>s. Farther south in Central .Vnierica .1. alhlrmfrh
is represent<-<l l,v u form .1. pi innhrin,!! is .,nit(- .HtlVrcnt

like it in gcncial tiiat 1 have no hesitation in regarding the south-
ern form as a snl)^[)('( i«v^ latlier than a segregate species.

^CoU. AiuericHU Muscnun of Xutural History.
'Coll. Unitod States Xatioual Museum.
3 Coll. E. A. and O. Bangs.

No. 483]

WOOD RAILS NORTH OF J'AXAMA

185

I find no indication of intergradation between A. cajanca and
A. alhiventris and must regard them as distinct species. The
much longer more slender bill of A. albiventris and tlie conspic-
uous chestnut patch on the back of the head, always serve to dis-
tinguish it in all its subspecies from A. cajaiiea.

Type: from Bikmki Veru ("ruz. Mexico, a.lult 2, no.

2281 Coll. of K. A. and ( ). Han.i^s. Colhvtcd Jimr 4. 1001, by
A. E. Colburn and P. W. Sliufcldt.

Geographic DisriniU TioN : Soutlicrn Mexico, in States of
Vera Cruz, Tabasco, Oaxaca and (diiapas, north to Hidalgo (one
skin from Orizava no. 2!I2;)1, V. S. Xat. Mns.) and on the coast
at least to Tampico.

Characters: Very simiUir to true J. aU)i rnitris, hnt pale

narrower and less distinct, strong buff in color, not white or cream
buff; all the colors darker — gray of head and neck, greenish
olive of back, and tawny of breast; much less siitl'nsed with
ochraceous or taw^ny on scapulars and outer interscaj)ulars thongli
this marking is often indicated; throat less purely white, more
grayish and this marking more confined, extending less onto
under surface of neck.

228P Type $ ad. Mexico, Vera Cruz, Biiena Vista. 177.
2280 cS'ad. " " " ISC.

1415362 9 ad. Mexico, Vera Cruz, Tlacotalpam. 17'.».

141537 (5^ ad.
141539 9 ad.

' Coll. of E. A. and O. Bangs.

'Coll. of Bureau of Biological Survey, Washington.
^ Coll. of U. S. National Museum.

186

THE A M ERIC A N NA T URA LIST \Yor.. XLI

Remarks: Aramidcs alhiveniris mexicamis is the northern
representative of this group of the genus, occupying the southern
tier of states of the Republic of ^lexico and southeastward passing
gradually into true J. alhiroitris of ^'ucatan and British Honduras.
Though well chaiactcrivuMl sul»])e('ifically it is in general much
like true A. alhirvuiris. It can, however, always be told from that
form by the characters pointed out al)ove.

Aramidp:s AiJin i.M Kis tumbkicollis (Zeledon)

Aramides plumbeicollis Zeledon Anales. Mus. Nac. Costa Rica,
1, p. 131, 1887. Biol. Cent. Am. Aves, Vol. 3, p. 320, 1897-1904.

Type Locality : Jimenez, Costa Rica. Type now no. 1 1 3603,
U. S. National Museum, examined.

Geographic Distribution: Costa Rica, specimens exam-
ined from Jimenez, Carrillo, and Cariblanco de Sarapiqui, north
at least to Segovia River, Honduras.

Characters: Similar to A. alhiveniris mexicanus, but slightly
smaller; bill actually shorter, though relatively of about the same
length; differing in color principally in having a comj^lete mantle
across upper back of olivaceons-tawny — the back thus bicolor,
olivaceous — tawny anteriorly, greenish olive posteriorly; breast
rather darker than in the other two snbsjjecirs more nearly as in
A. cajanca; crescentic marking made by paler feathers around
black belly patch, when present, narrow and buff in color (in two
skins, one from Carrillo and one from Cariblanco de Sarapiqui
this marking shows ver\' distinctly; in the type and one other
skin from type locality it is l)arely indicated).

$ ad. Costa Ku

' Coll. of E. A.

No. 483] WOOD RAILS NORTH OF PANAMA 187

Remarks: I feel confident that I am right in placing this bird
among the subspecies of A. albiventris, rather than to allow it
specific rank. In all essential points — the long slender bill and
chestnut color of the occiput and crown it agrees with A. albiven-
tris albiventris and A. albiventris mexicanm. The brown mantle
strikes one at first as a very strong point of difference, but this
is in reality only a difference of degree, many northern skins
showing a very decided approach to it, though it is in them never
quite complete all across the back as it invariably is in the Costa
Rican bird. The southern form is also somewhat smaller and
darker in color below than either of the other two races, but every
indication, in my opinion, points to its being a representative
geographic form — subspecies — of the A. albiventris type.

The specimen from Segovia River, Honduras, unquestionably
belongs here, as first pointed out by Richmond (Proc. U. S. Nat.
Mus. 16, p. 528, 1894). It is young, and as it happens is the
only young example of any of these rails, except .1. axillaris,
that I have seen. The feathers of the underparts, especially
the belly, are more flulfy than in the adults and in color it
differs in the belly (black in the adult phnnagc) being bhick only
at the base of the feathers which arc externally tipped and suf-
fused with the tawny color of the breast and in llie rump, also
clothed in fluffy feathers, being ,l(-<i.le,lly paler and browner.
Though badly shot in the back and neck with many feathers
from these parts lacking, the complett^ mantle of olivaceous-
tawny is plainly to be seen. The bill is not full grown and is
very immature in appearance. Judging from this skin it appears
that the species of Aramides of this group do not have a brownish
gray breasted juvenile pliunage as does A. axillari.^ and its allies.

NOTES AND LITERATURE

BIOLOGY

The Reception of the Mutation Theory.— Wheri the first Lieferung
of "Die Mutationstheorie " appeared in 1901 a frequent question was
whether the work would be made available for a larger audience
by the preparation of an English translation. That the interest
of Americans in this subject is very real was soon evidenced by an
invitation extended to Professor de Vries to deliver a series of lectures
at the University of California. A second edition of the thick volume
containing these published lectures was necessary in a few months.
Besides a French translation of the American lectures we now wel-
come an attractive German edition by Klebahn.^

Species and Varieties was reviewed in the pages of this journal
(Am. Nat. 39:747-751, 1905) and it seems unnecessary to discuss
the scope or contents of the work. The translator had the benefit
of the corrections prepared by Professor de Vries so that the transla-
tion is comparable with the second American edition. An especially
commendable feature of the present volume is a fine series of over
fifty illustrations. These are drawn in part from the larger work of
the author, in part from his unpublished drawings or photographs,
and in part from living material or other sources.

It must be gratifying to all serious students of evolution to see the
widespread interest in these works. Whether or not they admit
the general applicability of de Vries's theory, they must at least
realize that after long years of marking time students of evolution
have at last begun to march. No one should scorn the results of
comparative studies, but their limitations should always be kept
clearly in mind. The spirit of experimental work is in the air and
let us hope that there will be no turning back because of difficulties
encountered in the way. Just here a word of warning may not be
out of place. In experimental physiology and morphology it is con-
sidered essential that the factors involved and the results secured
be quantitatively expressed. In ecology and evolution the impor-

Ins Deutche iibertragen von H. Klebahn. Berlin. Gebruder Borntraeger,
1906. Q. xii + 530 pp.

189

190

THE AMERICAN NATURALIST [Voi>. XLl

tance of quantitative methods is just as great. While de Vries was
making the now celebrated experiments upon which his theory is
based Pearson and his associates were developing the methods of
quantitative investigation in variation and heredity. It will be unfor-
tunate indeed if present day workers neglect this new and power-
ful instrument of research. But with a proper combination of ex-
perimental and biometric methods it should be possible to gain a
very precise knowledge of the processes involved in species formation.

J. A. Harris.

A Monument to Theodor Schwann. — Theodor Schwann was born
at Neuss on the Rhine, December 7, 1810. On the centennial of
that date it is proposed to unveil a monument to his memory in his
native town. A considerable sum is already in hand and a committee
representing all countries has issued an apjx-al for subscriptions for
the memorial. As is well known, he with Schlcidcn, placed the cell-
theory on a substantial basis sixty-five years ago; wliile his later work
was almost equally valuable though not so startling in character. He
became an authority on fermentation, decomposition, digestion and
spontaneous generation, and, not least, was the discoverer of pepsin.
A monument to his associate has been erected in Jena \vhi](> his master
Johannus Miiller has a bronze memorial in his native town, Coblcnz.
Contributions may be sent direct to the 'Stiuhix he Sparka»c, .\cu>-;
am Rhein, Germany' marked 'Schwanndenknial ' or prol)al)ly to the
American members of the Committee, Prof. C. S. Alinot of Boston
and Prof. R. Ramsay Wright of Toronto.

Fitch's Basis of Mind and Morals.* — This book is a brief exposi-
tion of the principles of evolution as stated by Darwin and S])iMicer,
together with a discussion of the evolution of mind ami of the nat-
ural code of ethics. The point of view of the book is ]»1h iionienal-
istic; the style is simple, clear and direct. For those who have
thought seriously about the problems of evolution the work has little
value; for those who wish to be stimulated to such thought it may
prove profitable.

The atithor contends that there siiould be a natural co.le of ethics.

that it should be the rciilt of niaii"> knowlcluc of natural causes and

> Fitch, M. H. The Physical Jiasis of Mintl and Morals. Chicago, Charles
H. Kerr and Company. 1906. 266 pp.

No. 483]

NOTES AND LITER AT V RE

191

effects. "But I repeat, he says that until iiicii coiik- to coniprcliciKl
a natural cause for every uaiural cfVect they slidiiltl he coiitn.llcd
in their attitude toward environnieut, incltidiiio- tli,ir lin.ilicr men,
by some code that will have the proper effect, howwcr liascd that
code may be." (p. 255.)

GEOLOGY.

The reconstruction of the Continents of Tertiary times is the
topic discussed in a |)aper by \Y. 1). Mattlunv.' l'siii<: the evidence
furnished by the distribution of fossil and recent :Maninials, he tries
to reconstruct the outlines of the old land-masses, and illustrates
his results by seven niai)s, which represent the geographical conditions
of the earth's surface in Postcrctaceous time (immediately after the
close of the Cretaceous), in the middle Eocene, in the middle Oligo-
cene, in the Miocene, in tlie Pliocene, in the early Pleistocene, and
in recent times.

This pajicr inuloubtedly marks an important progress in this branch
of resoarcii. ('..tnparing it with tiic hist att.Miipt to reconstruct the
oldconti.KMits. nia.l(> I.v Orli.iaMU in I'.KL' (I'r. Aincr. IMiilos. Soc. 41),
we srr that hr.v onix iuo inai.. unv -ixrn, tnr the h.urr and for the

Mltthcu'. ,na|...'ahh.),ii:h throii.- lor the lou.-r Trnian mnv-ponds

Tertia^^ tn'ihat nf ih<' Mio.vnr. Hut ' coiiipK't.- amvnncni raiiliot
be expected, considcrlD.u- the (>Ntrcmc diniciiitics with which stich
investigations an^ {•oiiiicctnl. Indeed, it is rather stirprising that

192

THE AMERICAN NATURALIST [Vol. XLI

chiefly in Tertiary times, and the agreement of the zoogeographical
facts with paleontology and geology tends to show, that these recon-
structions of the old continents are not merely wild speculations.
However, the results cannot yet be accepted as final, and although
some of the major features of old geography must be regarded as well
established, much remains to be done in detail. Chief of all, additional
groups of animals should be studied, and an attempt should be made
to correlate the results obtained by them with those of Ortmann and
Matthew; and further, attention should also be paid to the Mesozoic,
and if possible, Paleozoic times. It is to be hoped that, for instance,
the distribution of certain molluscs and lower vertebrates may fur-
nish evidence with regard to these ages, although it is only natural
that this task will be much more difficult, since the facts are very
scanty, and their meaning is largely obscured by the changes in
subsequent times.

A. E. O.

The Mountains of Cape Colony. — In the Cape Colony, southern
Africa, are ranges of folded mountains very similar to the Alleghenies
of the eastern United States. During the summer of 1905 Professor
Davis had an opportunity to study the Cape Colony ranges while a
guest of the British Association for the Advancement of Science dur-

geologists and geogra pliers, l)ecimse of the striking similarity, in
practically all essential features, of the two widely separated moun-
tain groups, whether compared as to structure, the relation of folded
areas to undisturbed plateaus, the erosion history and development
of drainage adjustments, or the control exerted by the physiographic
features upon transportation, etc. In climate, however, a marked
contrast between the two localities exists. The paper is illustrated
by a number of drawings and photographs.

D. W. J.

Natural Mounds.-— During the hist two years a luimlier of j)apers

14, 708-717,

No. 483] NOTES AND LITERATURE

193

Mr. Campbell figures and briefly describes tlie mounds, reviews the
various theories of origin, some ten in number, uliich have been
advanced by various writers, and concludes from his own st\uiies
of the subject that the mounds have been built up by niits or small
rodents, more probably by ants. A bibliograpiiy \hv Mihjcri is
appended to the paper.

Ancient Glacial Periods.— During recent years the evidences of
repeated glacial periods during ancient (^a-olopcal time have been
accumulating so rapidly that whereas much doubt \\ as cast upon the
earlier reports of such glaciation, it is no longer possible for the unpre-
judiced student to doubt the conclusions which the evidence forces
upon us. The famous Dwyka glacial formation of South Africa is
now well known, and its equivalent in India, the Talchir. I. C.
White and David White have recently rcn< h.d ihr conclusion, inde-
pendently, that the equivalent of these I'cnnian or Pen no-Carbon-
iferous glacial deposits occurs in south(M-n Brazil in w hat is called the
Orleans conglomerate dlarial deposits in Au-lialia arc reported
from both the I^'rniian and the Cambrian or older l.cl.. A. P.
Coleman has recently reported evi.lcn.c of a lower Ilnn-nian ice a-e
in Canada. Mr. Schwarz^ discusses three ulacial p. rio.l. in Sonth
Africa, those in addition to the Pennian Dwyka In ini: nu»t j.robalily,
according to the author, of Devonian and Archaean aire. The relation
of the glacial beds to other members of the ovneral ^tratiirraj^liic series
is pointed out, and the evidence of the glacial origin considered. It
is this last point which in every case is critical. The fart that a large
number of reports of ancient glaciation are being piiMi-hed does ,i,,t
strengthen the evidence in favor of ancient glaciation ni any ]iartienlar
case. Each reported instance must be critically exammed as to the
value of the evidence supporting it.

194

THE AMERICAN NATURALIST

[Vol. XLI

PHYSIOLOGY

Hough and Sedgwick's Physiology.*— "The authors of this work
beheve that extensive and fundamental changes must be made in the
elementary teaching of physiology, hygiene, and sanitation, if these
subjects are ever to occupy in the curriculum of education the place
which their intrinsic importance requires." This sentence from the
Preface to this new book by two well-known professors of biology is
the key-note to its importance, for their intention in this respect cer-
tainly has been fulfilled. Not only the students of high schools,
academies, and colleges actually need to know the facts and principles
set forth here, but so also does the long-graduated 'average man' if
he would live well. Especially is it one more step toAvards the recog-
nition educational theory is certainly about to make, that in education
every part of a boy's body one is educating at the same time and in
the most real manner also the capability of his whole mind.

The book is divided into two nearly equal parts: 'Physiology,'
and 'The Hygiene of the Human Mechanism and the Sanitation
of its Surroundings,' respectively. The latter half is subdivided into
accounts of personal hygiene, domestic hygiene, and public hygiene
and sanitation, with an important introductory chapter in addition.

The matter of the first part of the book is better than its arrange-
ment in chapters, for the nervous system is placed last and the mus-
cular mechanism early in the list. For the learner the much more
preferable order is just the reverse, it being certainly difficult really to
understand any one of the great organic functions until the coordi-
nating purpose of the nervous system is mastered. One deplores
too the omission of at least a brief discussion of protoplasm in general
as an introduction to its differentiated natures.

A far more serious omission (but one morv easily defctisibltO is
that of the basal principles of rcpnxhiction. WIhmi all is said, at

» The Human Mechanism: its Physiologj' and Hygiene and the Sanitation
of its Surroundings/' Theodore Hough and William T. SedgAvick. Boston,
Ginn & Company, [1907]. Pp. ix + 564. Illustrated.

No. 483]

\()T/':s AM) Ln\

195

not intended for grammar schools nor for the first years of the liiirh
school even, but for schools whose students might soon aspin- to he
husbands and wives.

The chapter on muscular activity is uniquely fine in iis cHm ns>ion
of the necessity for physical exercise, and in combination with |.tvvioiis
chapters on muscle-function and neural coordination ahno>t meets
the insistent demand pedagogy is beginning to make for bodily skill
as a basis for learning. One misses, perhaps, an adequate descrip-
tion of the kinesthetic mechanism for muscular control, as well as
sufficient information as to habit and the emotional reactions. On
the other hand, 'rhythmic segmentation' is allowed far more promi-
nence than the doubts as to its existence warrant.

The hygienic portion of the book is rieh in clear and precise infor-
mation of really great importance to everyone. .Moreover it is set
forth in a manner as scientific and up-to-date as could be desired.
Could an enlarged wall-copy of figure UC ("A domestic well badly
situated in a farmyard"), be distributed broadcast by the state boards
of health, our city hospitals would soon cease to be over-filled with
typhoid patients in October and our farin-hoiises would \)v less sad-
dened by cholera infantum in the summer.

The account of personal hygiene is at once cniinently praciii al and
entirely scientific — a needful combination .s(>ld()ni attained. More-
over it is more complete than is common in text-books of t!ii< s(»n.
It seems as if too little emphasis perhaps, were placed on tlic impor-
tance of moisture in the air of dwellings, this need heino- met hy ( .m-
tinually open windows. It is the throat-speciali.sts who hot realize
the general lack of moisture in the atmosphere of onr hoii-c-. hut
there are of course other reasons (such as that moist warm air fcel>
warmer than does dry warm air) which are important in the theory of
ventilation.

Few but physioloo:ists familiar with the re(iiiircd falsiti.'. as i..
alcohol and tobacco which reek in certain states, esjiecially \\est\\ard,
will reahze how excellenr is the dix iis^ion of tlioe wry important
topics in this book. The facts arc clearly stated and the principl. s
laid down,— their dan,ircr> in ovcrn>c any Mnd(>nt in a school
for normal persons may certainly sec and he warned hy for himself.

The 147 often fatnil'iar illn-I rat ions of the work are adc(,nate and
for the most part well execntcd.

196

THE AMERICAN NATURALIST [Vol. XLI

ZOOLOGY

Guyer's Animal Micrology,^ though burdened with a horrible
name, is one of the best and most practical works upon microscopic
technique with which we are acquainted, ranking, in this respect with
Bohm and Oppel's well-known "Taschenbuch" which, by the way,
is not referred to in the list of works cited on p. vi.

The especial merit of the work lies in its great practicability. It-
does not burden the beginner with a large number of alternatives;
but starts him 'at once with a f(>\\ reagents of almost universal avail-
ability and sets him at work with his specimens. Only when these
have been carried through and converted into slides are other methods
and other objects considered.

In the Appendix are given an account of the microscope and its
accessories, a list of further tried and proved reagents and a table of
tissues and organs with methods of preparation which will doubtless
prove of value to instructors as well as to students. The list which
is given embraces over 250 objects and is more than ample to illustrate
any practicable course in normal histology. The final chapter of the
Appendix deals with methods preparatory to microscopic preparation
and study of a series of animals which are frequently used in the
Zoological Laboratory.

Omissions of what we would like to see in such a work are few. We
have found no mention of Cox's Golgi method which presents certain
advantages over the silver impregnation; the Golgi method for dis-
tinguishing bile capillaries is not referred to, nor is the value of Lyons
blue for differentiating cartilage. The method of rolling wax plates
for reconstruction, credited to Ruber (p. 128), has been in use for
many years. In the ' Memoranda' on p. 30 it is stated that material
which is to be kept indefinitely should be put in tightly stoppered
bottles, but there is no hint as to the injurious effects of the extracts
of cork and that some other method of closure should be adopted.
But why find any more fault with such a useful and excellent work?

J. S. K.

. F. Guyer, Chicago, Ur

No. 483]

NOTES AND LITERATURE

197

MoUusca of Illinois and Michigan.— F. C. Baker has recently
catalo^iied^ the :Mo]hisca of IHinois, enumerating in all 332 species of
which 91 are terrestrial and 240 are aquatic (the figures are the
author's, the cliscre})ancy is not explained). Tlic I'nionidac number
89. The list gives localities with considerable detail; no new species
are described. Bryant W alker's catalogue of the terrestrial Pulmonata
of Michigan 2 is more claborale, jriving descriptions and in most cases
figures, with an onthnc ol the svnoiivmy of the 79 species recorded
from the state. In ilic Intiixhictioii. besides general notes on dis-
tribution, adequate di ir( i loiis ;irc o;ivcu for the collection and prep-

A monograph on Anurida. Those in charge of the I.iverj>ool

of short M(>inoirs on the morphology, life history, and oecology of
various typical animals and ])lants found in that region. The thir-
teenth of the series appears in volume 20 of the Proceedings and
Transactions of the societv. It ck^als with the interesting (\)]lem-
bolan, Anurida an.l is I>v'a. I). Imms. The habits nii.l structure
arc dcMnlH.I t,,,ni onom.il obMn.iiion- ih. ... . ouni ih( .|.N<l.,p-

BOTANY

A popular book on Canadian wild flowers.— In 188.", the venerable
Mrs. (-athcrinc I'arr Traill was among the leaders in popularizing a
knowledov of Vnu-rican wil.l flowers In- pres,>ntinir th.-m unteehiii-

198

THE AMERICAN NATURALIST

[Vol. XLI

successfully followed in this countrv by iiuuiy and excellent works
of the suine character. A new and revised edition of her book has
now made its appearance.^ Like the original, it has passed through
the hands of Macoun and Fletcher, for the determination of the plants
included; it should stimulate in many people of the present day that
love for plants and their Avays w^hich comes through knowing what
they are,, and toward which the first edition did such good service two
decades ago.

W. T.

Notes. — An interesting and appreciative sketch of de Vries, by a
former assistant, Henri Hus, has been separately issued from The

A handsomely printed volume of botanical studies presented to
Kjellman on his 60th birthday has been distributed by the University
library of Upsala.

A detailed account of the history of natural science in the Aberdeen
Universities has been reprinted by Professor Trail from "Aberdeen
University Studies."

Semon's terminology, "equally applicable to the movements of a
plant or the thoughts of a man, " is used by Francis Darwin in a lecture
on associated stimuli, printed in The New Phytologist of November

A lecture on "Mendelism and Microscopy" is published by Scour-
field in the Journal of the Quekett Microscopical Club of November.

The viability of old seeds has been tested recently by Becquerel,
as reported in the Comptes Rendus of June 25 last, and abstracted
in the Gardeners Chronicle of November 24.

A concrete presentation of the results of local ecological study of
the modern sort is afforded by Woodhead's Huddersfield paper occupy-
ing no. 261 of the Journal of the Linncan Society, Botany.

Strasburger contributes an illustrated paper on the thickening of

ivissenschaftliche Botou ik.

'Traill, Mrs. C. P. Studies of Plant Lifi; in ('Hiuuhi. Toronto, Willimn
Briggs, 1906. 8vo. xvii + 227 pp., with 8 reproductions in natural colors
and 12 half-tone engravings, from drawings by Mrs. Agnes D. Chamberlin.

No. 483] NOTES AND LITERATURE 199

Habit illustrations of a number of the economic plants of West
Africa occupy Heft 5, Vicrte Reihe of Karsten and Schenck's "Vege-
tationsbilder."

in the . I rh-ir jOr Hnhui ik of Stockholm.

A morphological and anatomical study of CraiiofJnis- (nncricanus
and C. ovaius is published by Holm in TItr Amrrlnni ,h,uri„il of
Science for December.

An extensive segregation of the components of liliiis (jlabra is
effected by Greene in the Proceedings of the WashitujUm Avmlrmii of
Sciences of December 18th.

Agnes Chase ]iu1)lishcs on Paniccfc in the Proceedings of the Bio-
logical Society of Wo.s'h inf/tmi of Deconiber 8.

An interesting study of die I'>uolciioid genus Dunaliella is published
by Teodoresco in the limu Cnirrale de Botaniqve of Se])tcmber 15.

/'//A/////// >h Banjanuw is said by Raffill, in The dardmrr/ C hrnn iric

An illustrated monograph of Kav(Micli:i is publislicd hv Di.tcl in
vol. 20, Abt. 2, HclY :\ of the lirihrjfe /mi, IJolanix licn ( •.Miliall.lalt.

A practical account of the fungous .liM-i^cs of tulips an.! tlicir
treatmcnl is contril.ulc.l hy l\lcl)ahn to ( la rh n jlnni of November 1.

A small text book of funu-i. including nunphologv, physiology,
pathology , (■la.>iti<-aii..n, ctr., i.y Ma-cc has Iummi i-ucd from the

i|)liic skct. li of Mitten, with portrait, is published in The

200

THE AMERICAN NATURALIST [Vol. XLl

An excellent, conservatively handled, local flora, of a very rich region,
is that of the State of Washington, by Professor Piper, recently pub-
lished as vol. 11 of Contributions from the U. S. National Herbarium,
an illustrated volume of 637 pages.

In contrast with the highly diversified flora of Washington, is the
homogeneous flora of the Altamaha grit region of the coastal plain of
Georgia, to which is devoted a volume of 357 pages, by R. M. Harper,
forming vol. 17, part 1 of the Annals of the New York Academy of
Sciences. In this region, comprising about 11000 square miles, only
814 species or varieties of vascular plants are recognized, and 75 of
these are weeds. Mr. Harper's study has been carried out on the lines
of ecological analysis with special reference to geographic distribution,
and his paper is illustrated by a map and 28 half-tone plates which
form one of the best series of such illustrations yet published.

Habit illustrations of antarctic vegetation are given by Skottsberg
in Reihe 4, Heft 3-4 of Karsten and Schenck's Vegetationsbilder.

Further "Contributions to Canadian Botany" are being published
by IVIacoun in current numbers of The Ottawa Naturalist.

The official precedings of the International Botanical Congress
held at Vienna in 1905 have recently been issued from the Fischer
press of Jena, in the form of a quarto brochure of vi + 262 pages:
the scientific papers presented before the Congress form a similar
quarto of vi-|-446 pages, freely illustrated, and published by the

A polyglot code of the rules of botanical nomenclature adopted by
the 1905 International Botanical Congress of Vienna, has been sepa-
rately issued from the Fischer press of Jena. The pamphlet is indis-
pensable for every phanerogamic herbarium. One of the most debated
acts of the Congress was the adoption of a list of several hundred
generic names which were considered so thoroughly established as to
be exempted from supersession by earlier names which have failed to
come into general use. This list is included in the pamphlet.

Professor Bray's " Wgc^tatioii ..f the S,.tol Country in Texas,"
elsewhere published, is also printed in vol. 7 of the Transactions of
the Texas Academy of Sciences.

No. 483]

NOTES AND LITERATURE

201

For Juliana and Orthopterygium, Mr. Hemsley proposes a new
Order, Julianacefe, to go between Juglandacese and Cupuliferae,—
in The J ournal of Botany for November.

Brand describes and figures under the name Trijolium pratense
foliosum, a glabrous clover recently introduced into American culti-
vation from Orel, Russia. {Bulletin no. 95, Bureau of Plant Industry,
U. S. Department of Agriculture).

A considerable number of new species of the orchid genus Acovi-
dium are described by Ames in the Proceedings of the Biological
Society of Washington of September 25.

The Department of Agriculture in India has begun the publication
of an important series of botanical memoirs, from the Agricultural
Research Institute at Pusa. The three numbers thus far received
refer to "Fungus Diseases of Sugar Cane in Bengal," "The Hau-
storium of Santalum album," and "Indian Wheat Rusts." A fourth
paper, on "Gossypium ohtusifolium," and a fifth, "An Account of
the Genus P}i;hium and some Ch\i;ridiace8e," are also announced.

An account of Cratfegus, as richly represented in the vicinty of
Albany, has been separately issued by Sargent and Peck from Builetin
105 of the New York State Museum.

A colored plate of Rihr.^ crunifutii is given in Curtis s> Bnfanind
Magazine for November.

The first issue of The Bulletin of the Picfon Academy Scientific
Association contains an account of the Myxomycetes of Pictou County,
Nova Scotia, by C. L. Moore.

Huber publishes a synopsis of 18 recognized species of Hevea in
vol. 4, no. 4, of the Boletim do Museu Goeldi, of Para.

The fondness of cats for Act in id ia poly ga ma is re-recorded by Fair-
child in Science of October V^.

Several new Cuban grasses are .Icscrihc.l l)y Hackel in the first
Informs Anual de la F.sfarlihi Cmfrnl Agrn„6mira dr Cnha, issued in

volume oF Thr Pln/lppinr Jnurna/ of Srirn<l 'r.uUains a Ii>t of known
Philippine fun.u-i, by Hieker.

Adams, in The Irish Xnturalist for November, notes that a molil
of fermenting hay thrives at an induced temperature as high as 135.5°

202

THE AMERICAN NATURALIST [Vol. XLl

Magnus has separately issued from vol. 21 of the Naturwissen-
^chafiliche Rundschau an account of the destructive mushroom para-
site, Mycogone perniciosa.

An exhaustive account of a Sclerotinia-rot of apples is given by
Molz in the Centralblatt Jilr Bakteriologic, etc., Abteilung 2, of October
27.

A study of the influence of selected yeasts upon fermentation, with
reference to cider making, by Moncure, Davidson and Ellett, forms
Bulletin 160 of the Virginia Agricultural Experiment Station.

The Ustilaginales of North America are revised by Clinton in the
recently issued vol. 7, part 1, of "North American Flora," under the
editorship of Professors Underwood and Britton.

A descriptive account of the economic plants of the world and of
their commercial uses, by Freeman and Chandler, is being issued in
fortnightly parts by Pitman and Sons, of London, under the title
"The World's Commercial Products."

Brief descriptions, with 3-color illustrations, of the most noxious
weeds or "proclaimed plants" of Victoria are being published by
Ewart and Tovey in The Journal of the Depaiimrui of Agrlnilture of
Victoria.

An illustrated account of the seed of red clover, and its impurities,
by Brown and Hillman, forms Farmers' BuUdin no. JOO, llic U. S.
Department of Agriculture.

Laubert gives an account of Ambrosia arfemtxiafolid as a (Jcrman
weed in vol. 35, no. 5, of Landwirtschaftlichc Jahrbiichcr.

Stockberger gives an economic account of Spigelia marilandica and
its surrogates in Bulletin 100, part 5, of the Bureau of Plant Industry,
U. S. Department of Agriculture.

A portrait of I>ord Avebury forms the frontispiece to Nature Notes
for October.

being published by Paris in current numbers ..!' tlu- l!rn,r ] ln>i imk
de VAlgerie.

"Date varieties and date culture in Tunis" is the title of Uulhiin
no. 92 of the Bureau of Plant Industry, IJ. S. Department of Agri-
culture, by Kearney.

No. 483] NOTES AND LITERATURE

203

An illustrated practical guide to judging and selecting corn is given
by Shoesmith in Bulletin no. 139 of the Kansas Agricultural Experi-
ment Station.

An economic account of the cultivation of Agave caninla in the
Philippines is given by Edwards in Farmers' Bulletin no. 13 of the
Insular Bureau of Agriculture.

preparation in Africa, in Der Tropenpflanzrr for October.

Chemical studies of Althusa, Grindelia and Pittosporum, by Power
and Tut ill, have recently been distributed as papers from the "Well-
come Research Laboratories of London.

An interesting account of the use of tree bark etc. for bread making
is given by Dillingham in the recently issued vol. 3, part 5, of the
Bulletin of the Busseij Institution of Harvard University.

Some good root-habit photographs of Ficus are reproduced in
Arboriculture, for October.

Biffen analyzes Mendel's laws of inheritance with reference to
wheat breeding, and the inheritance of sterility in barley, in the recently
issued Cambridge volume of reprints from vol. 1 of the Journal of
Agricultural Science.

A second edition of De Vries' "Species and Varieties: their Origin
by Mutation," corrected and revised under the editorship of Dr.
MacDougal, has been issued by The Open Court Publishing Com-
pany. The frontispiece is an excellent but somewhat informal por-
trait of the author, at work.

Further evidence of the germicidal effects of copper is given, in
official orthography, by Kellerman and Beckwith in Bulletin no. 100,
part 7, of the Bureau of Plant Industry, U. S. Department of Agri-
Livingston publishes an important study of the relation of desert
plants to soil moisture and to evaporation as Publication no. 50 of The
Carnegie Institution of Washington.

A paper on the effect of tension ii])on the developnient of inechanical
tissues in plants, by Ball, is contaiiuMl in vol. 7 of tin- Transartinns of
the Texas Academy of Srirnrr.

From a study of the strength of the bands which Thyridopteryx
fastens about twigs, the results of which are published in vol. 17 of the

204

THE AMERICAN NATURALIST [Vol. XLI

Report of the Missouri Botanical Garden, von Schrenk concludes that
the radial force of twig gro\\i;h may equal a pressure of 40 or more
atmospheres.

A biographic sketch of C. B. Clarke, with portrait, appears in the
November Journal of Botany.

A short account of the :McKinley or Dinkey grove of big-trees is
given by Guthrie in Forestry and Irrigation for October.

The Journals. — Botanical Gazette, November: — Chamberlain,
"The Ovule and Female Gametophyte of Dioon"; Brooks, "Tem-
perature and Toxic Action"; Cook, "The Embryogeny of some Cuban
Nymphseacese."

The Fern Bulletin, October:— Fellows, "The Fern Flora of Maine' ;
Gilbert, "Polypodium vulgare var. alato- multifidum, var. nov."; Clute,
"The Genus Oleandra"; Negley, "Where Florida Ferns Grow";
Palmer, Asplenium ebenoides in Chester Valley, Pa."; Ferriss, "On
Cultivating our Ferns " ; Clute, " Rare Forms of Ferns, — I " ; Squires,
"A New Station for Selaginella douglasii" ; Puffer, "The Rusty
Woodsia in Cultivation."

Torreya, December: — Harper, "Some Hitherto Undescribed Out-
crops of Altamaha Grit and their Vegetation"; Berry, "Leaf Rafts
and Fossil Leaves" ; Sheldon, "A Rare Uromyces."

Rhodora, November: — Hitchcock, "Notes on Grasses"; Blanchard,
"Some Maine Rubi. The Blackberries of the Kennebunks and Wells
—HI"; Fernald, "Twelve Additions to the Flora of Rhode Island";
Leavitt, "Regeneration in the Leaf of Aristolochia sipho" ; Fernald,
" Potamogeton spathceformis a probable Hybrid in Mystic Pond."

Torreya, November: — Howe, "Some Photographs of the Silk
Cotton Tree {Ceiba pentandra), with Remarks on the Early Records
of its Occurrence in America"; Hill, "A Mississippi Aletris and
Some Associated Plants"; Shafer, ''Hibiscus oculiroseits" ; :Murrill,
"How Bresadola Became a Mycologist"; Burnham, "A New Species
of Monotropsis " ; Blanchard, "A New Dwarf Blackberry."

Journal of Mycology, November: — Long, "Notes on New or Rare
Species of Ravenelia"; Atkinson, "A New Entoloma from Central
Ohio"; Kellerman, "Fungi Selecti Guatemalenses Exsiccati, Decade
1" [label data]; Morgan, "North American Species of Lepiota"
(continued); Kellerman, "Index to North American Mycology"
(continued).

No. 483] NOTES AND LITERATURE

205

The Ohw Naturalist, November: — Schaffner, Mabel, "The
Embryology of the Shepherd's Purse"; Hambleton, "Key to the
Families of Ohio Lichens"; McCleery, "Pubescence and other
External Peculiarities of Ohio Plants."

Tfie Plant World, October:— Arthur, "The Paired Seeds of Cockle-
bur"; Tullsen, "The Probable Origin of Key-Fruits"; Parsons,
"Children's Gardens and Their Value to Teachers of Botany and
Nature Study"; Blumer, "Wild Fruits and Shrubs of the Priest
River Valley"; Taylor, "The Germination of the Morning Glory."

The Bryologist, November: — Fink, "Further Notes on Cladonias
— VIII " ; Hagen, " A Study of Tetraplodon australis " ; Bailey, "Van-
couver Island Bryology — I"; Lorenz, "Notes on the IVIosses of
WaterviUe, N. H."; Haynes, "Ten Lophozias"; Collins, "Notes
on Polytrichum commune."

Bulletin of the Torrey Botanical Club, October:— Arthur, "New
Species of Uredinese — V"; Harper, "Notes on the Distribution of
some Alabama Plants"; Piper, "Notes on Calochortus."

Journal of the New York Botanical Garden, November: — Britton,
"Recent Explorations in Jamaica"; Underwood, "Report on the
Condition of the Tropical Laboratory"; Taylor, "Collecting in the
Mountains West of Santiago, Cuba."

Journal of Mycology, September: — Kellerman, "A New Plowrightia
from Guatemala"; Arthur, "A New Classification of the Uredineae";
Bain and Essary, "A New Anthracnose of Alfalfa and Red Clover";
Atkinson, "Two New Species belonging to Naucoria and Stropharia";
Morgan, "North American Species of Lepiota (continued)"; Hedg-
cock, "Some Wood-Staining Fungi from Various Localities in the
United States"; Kellerman, "Notes from Mycological Literature —
XXI," and "Index to North American Mycology (continued)."

Of Mr. Elmer's Leaflets on Philippine Botany the following articles
have been issued :— Elmer, "Philippine Rubiaceae," "A Fascicle of
Benguet Figs," "Additional New Species of Rubiaceae," and "Pan-
dans of East Leyte"; and Copeland, "A New Polypodium and Two
Varieties."

{No. 482 was issued February 16, 1907).

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VOL. XII, NO. 484

APEIL, 1907

THE

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

The Geographic Distribution of Closely Belated Species

DR. ROBEBT GREENLEAF LEAVITT
The Coincident Distribution of Belated Species of Pelagic OrganiBms as Illus-
trated by the Chaetognatha . . PBOF. CHARLES ATWOOD KOFOID
The Attached Young of the Crayfish Cambarus clarkii and Cambarus

and Mutualism ; Geology, Relative Geological Importance c
nental, Littoral, and Marine Sedimentation, Obser%'ations in South
Africa, Geology of the Big Horn Mountains, A Glacial Lake in Tibet ;
Zoology. A Statue of Lamarck, Gardiner's Maldive and Laccadive
Archipelagoes, Kolhnann's Atlas of Human Embryology, The System-
atic Position of the Tubinares, Beebe's The Bird, The Conus Arteriosus
in Teleosts, Does half of an Ascidian Egg give rise to a whole Lar\'a ?
Digestive processes in CoUembola, Fresh Water Amphipods of North
America, Some Problematic Worms; Botany. Sukkulente Euphorbien 276

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THE

AMERICAN NATURALIST

Vol. XLI April, 1907 No. 484

THE GEOGRAPHIC DISTRIBUTION OF CLOSELY
RELATED SPECIES.^

BY ROBERT GREENLEAF LEAVITT.

The botanical researches of the members of the New England
Botanical Club are largely taxonomic and floristic. With some
of us this is vocation, with others avocation. The majority per-
haps pursue the study of plants in the field and make collections
of them in herbaria for their own personal satisfaction. Floristic
studies may properly be an end in themselves, whether followed
as a business or only for recreation. In the latter case they
need no further justification than the fine and pure pleasure they
afford to those who love them for themselves. But the results
of these studies, for whatever conscious motive pursued, may have
an application and a destination far beyond our private aims.
Collections of specimens and reports of distribution recorded
in accessible journals by well-informed non-professional as well as
professional botanists, may help materially in answering some of
the largest questions of biological science. In this paper I hope
to make it clear that refined taxonomy and most thorough-going
plant geography may have a direct relation to the enormously
diflScult problem of evolution.

Organic geography has, indeed, already served the cause of
evolution, — in aiding to secure general acceptation of the Descent
Theory. Darwin and Wallace, drawing upon the works of tax-
onomers, were able to point to features in the distribution of

• A paper read before the New England Botanical Club at the meeting of
Feb. 1, 1907. Published as Contribution from the Ames Botanical Laboratory,

208

THE AMERICAN NATURALIST

[Vol. XLI

species which support the notion of common descent. Plants
and animals, they said, occur upon the surface of the globe just
as if they had originated by evolution, and in a manner unintel-
ligible on the assumption of special creation. Species are uni-
versally found in the neighborhood of other species which they
resemble; or to put this generalization in evolutionary phrase,
species arise in geographic proximity to the species from which
they may be supposed to have sprung. The geographic evidence
was an important part of the testimony accumulated by Darwin
('59), to which he gives two of the fifteen chapters of the "Origin
of Species." Wallace ('55) had already published an essay
arguing for the evolutionary conception of organic history, the
main thesis being this: "Every species has come into existence
coincident both in time and space with a pre-existing closely
allied species." Thus the evolutionist has been under deep
obligation to the taxonomer from the beginning.

The obligation is likely to be much increased with the lapse
of time. I do not agree with D. H. Scott, that the determination
of the actual course of descent is the ultimate, or chief, object
of the scientific systematist.' The fact of evolution being ad-
mitted, and the course of evolution lia\iiiu- been ascertained^
there still remains the question, " Hy what methods have new
forms emerged from old ones?" — a subject not less interesting
or important than the others, from any point of view. It schmus
to me, furthermore, that the final i^oal of phyioncon-raphy is not
reached in the reconstruction of the continents and islands of
former epochs, and the reviving of ancient states and changes
of climate, through the study of the history of the vegetation of
the earth ; nor is its purpose satisfied in teaching us through

fariously adapted to tiieir environments. Hioloiiieally considered,

matic botany and zoology and with experimental morphology
in composing the solid basis of an a(h (|uate th(H)ry of evolution.

'The present Position of Palseozoie Botimy. Lot^v^ I'mir.v.Mi-. liei Motan-
icjp, 1: 139 (1907).

No. 484] CLOSELY RELATED SPECIES

209

The finely discriminative work of modern taxonomers, much
as it confuses and discourages students of other aspects of bio-
logic science, is necessary for several reasons but especially is
it necessary from the point of view of the evolution problem.
Doubtless systematic and experimental work will be more fre-
quently cooperative henceforth, and such studies as those of
Alexis Jordan, de Bary, Rosen and Wittrock will be repeated
with many of the so-called polymorphic plant groups by students
of the greater problem.

This view is apparently opposed to that lately expressed by a
prominent worker in experimental evolution, who seems to deny
this applicability. "The underlying fault," he says, "consists
in the fact that taxonomic and geographic methods are not in
themselves, or conjointly, adequate for the analysis, or solution,
of genetic problems. The inventor did not reach the solution
of the problem of the construction of a typesetting machine by
studying the structure of the printed page, but by actual experi-
mentation with mechanisms, using printed pages only as a record
of his success. Likewise no amount of consideration of fossils,
herbarium specimens, dried skins, skulls, or fish in alcohol may

eration, although from such historical data the general trend
or direction of succession may be traced." — MacDougal (:06,
p. 4).

Nevertheless, it may be shown that, while such studies are
not in themselves adequate to the solution of genetic problems
they have a very high corrective and evaluatory worth.

Geographic studies foiuided on an exact taxonomy have a
corrective function. It is axiomatic that no theory having its
origin in experiment can be acccj)tc(l if it seems to be in funda-
mental (lisconl with what wc know of the present disposition
of the organic world. For example, the theory of MutJition as
developed by de Vries cannot be acci^ptnl for the animal king-
dom, if, as seems to certain zoolo^M'sts, it is irreconcilable with
the facts of the distribution of animals. And even if an hypoth-
esis is not positively excluded by the facts, it may be weakened
or practically nullified by comparison with large bodies of facts

210 THE AMERICAN NATURALIST [Vol. XLI

gathered broadly; so that we may fairly ask the experimental
school to admit that results, however well proved for the condi-
tions established by the experimenter, ought to be assigned little
worth if they find only a trivial correspondence in nature at large.
We make the same demand of the physiologist with respect to
such a phenomenon as geotropism, for instance. The extended
study of this form of irritability has its justification only in the
fact that plants in nature so widely show the effects of geotropism
in their forms; the value of the experimental results is great
because the phenomenon is manifestly widespread in free nature,
being observable in the erect attitude of countless main axes
in field and forest, in the fixed angles of side stems, the vertical
descent of tap-roots, etc., etc. In like manner a true theory of
specific origins should find strong confirmation in the study of
the broadest aspects of plant and animal life. Every grand
agent of specific modification should leave its distinctive mark
upon the character of life as a whole, and if we rightly apprehend
the nature of the agent we may expect to be able to distinguish
its special mark or effect when we know plants and animals
thoroughly. I think that it will appear from considerations
which I now bring before you that the distribution of species
must have peculiarities corresponding to the particular class of
evolutionary forces which have been at work. If this be so, suit-
able studies in geographic taxonomy must possess high evalua-
tory worth when we wish to estimate theories of evolution.

The Effects of Different Evolutionary Agencies upon
Specific Distribution

Let us examine the necessary effects of the chief supposed
evolutionary agencies upon the character of specific distriluition;
and first contrast Natural Selection in I)ar\vin\ stricici- sense
with Mutation, in this regard. Natural Selection works within
specific limits. Its materials are the small, or individual, varia-
tions within the species. By the accumulation of tiiese variations
as they occur from generation to generation new characters are
built up. The change in a species is slow and the whole species
within a given competitive area moves along together. When

No. 484]

( lObhl^i RI I MED .SPEC IIS

we consider that sufficient change has occurred to w{
epithet 'new,' as apphed to the conchtion of tlie ^n-ouj
that the new species has risen upon stepping stoiKvs o
self, since the survival of the fittest has had its convc
extinction of the unht — that is, the 'old' spec ics
the given area only a single new species is louml rc[)
vanished old one. For any given area of conijuMiiiDii
forming effect of Natural Selection then, is moiiolvpic.
( 06, ch I ) cleaih states the truth that Natiii il -iUi
out isolation effects monotvpic evolution, and oiiK hv i
isolating factors of some kind resuhs m poh i \ pic <
Nageli's earher exposition of the monoiN pic ctlcct ot
Selection was explicit (Naueh. 7;)).

Ontheothei li iiid Mm ition I.k ik. rh( .puu^ ind
rily at least, must o-iv(^ a polvtvpic aspect to the urtttip
specific area. I'lic pari'iit species is conteinjxiraiieoiis
new spcdes to uludi ii ^n(^ iim 1 lu lu u ui.l iIh <
side In- side for a linuv wiilicut n(,,oTapliic iM.laiion and i

AMthont Lolninll „l IMX MMI llll^ 1> iIm pillllin <

Subseeiiienilv c,)nipetiti..ii niav leave onlv .nie ..f tli.

to f^o^\nll^ (.1 l.Mchim tiiiu i phx moIcuk d iM»]iti.m
the imin(<liU( k Milt ..t Mm itu.n l',iit is , mh tlu 1
of Mutation iim>l certainlv he tlie allocation of r\o<vh.
species, or kinds, in tli(> same area wiiliont anv M)rn

If ^^e compau ( )i ihojuu ^i. utm^ uihK i .md
enMronment, XMtli \ itin d ^(Uction on til. <'i . hind
tion on the other, we sei^ that ( )rtlio<:-eiiesi> must in ma
agree with Natural Seleciion rather than with Mutation
cerns the distrihulion of us ])rodncts - >pecies. 1 hroi
single region of unitoini < , oloi,i< .1 . Inr u t. i ilu ( tlu i ot
mental moiildmo. so-called, upon a u-i\eii ori^amc siock

geographic district, as tliev alwavs are when the surtaee i.

212

THE AMERICAN NATURALIST

[Vol. XLl

merits in the district; or, if interbreeding suffices to reduce the
diversity in some degree, at least several kinds perferring different
hal)itats. While in the broader geographic sense this effect
would be polyty})ic, in that any geographic district nnght have
several different closely allied types, each type would fit a partic-
ular set of conditions; there would be definite allotment and
topographical separation of the derivative species, and each
ecological field would present a monotypic aspect. A distribu-
tion quite distinct from that due to recent Mutation would be

Thus while the geography of species may or may not be deci-
sive as between the evolutionarv theories known by the names
Natural Selection and Orthogenesis, both those modes are dis-
tinguished from IMutatioii in the unincdiaK' cllects winch they
have upon distribution. >Sn(h sjx'cific (hstnbntion as Moritz
Wagner asserted to be ninvcrsal or almost nnivcrsal. li it could
he pro\ed, would be piadualK t ital tc the MiMation Theory
regarded as a general explanation ot .sjx'cilic evolution.'

the competence of geographic evidence. Indeed the\ appear
at times to recognize the propriety of the appeal to nam re; 1 )e
^^les refers to Draba, Viola ami similar gM.ii|.. iiid Ma< Doii-al
in the paper alreadv cited alludes to stiidi.>s of [.lant distribution
and addiKo tin (an ot <I.>mK idand Opiintia. in tlu Vn/ona
<leMMt llu <li^tinun>h(d /o<.lo^i.t ^^\^u u(UitK ...iitned to
pass upon the merits of the Mutation Theorv, (evidently without

' let Danviii (Oris. Hp., eh. IV.) in his theory of Divergence of Chanxcter,

Xo. 484] CLOSELY RELATED SPECIES

213

much knowledge of De Vries's work and apparently after a read-
ing of only the popular lectures published in this country/ was
at least thus far right, that he searched for indications of muta-
tion in the distribution of animals. The evidences should be
found even in museums, providing the museums are representative,
and providing mutation is a sufficient explanation of the origin of

The Necessity of Isolation: ]Mexdeliax IxnKHiTAX( e

Before coming to an examination of the facts as thcv an> repre-
sented by writers, it will be well to consider for a nioinciii a the-
oretical side of the subject, namely the siii>])(.s(m1 nrcrssiiii of
isolation as a factor in the evolutionary process, (irantiiiu that
new forms may appear upon the scene by Muiaiioii. wliai is to
become of them? How can ]\Iutation be said to oriuiiiate new
species — that is, stable groups — if through iiiterbreeihiiij tlie
mutants are at once swallowed up by the parent species with \\ hieh
they grow commingled V The opponents of the Miitaiioii Theory
hold that the isolation which the experimetiter practico in his
garden by means of paper bags, etc., is lackin-' in natnre and
that this difference between the garden and free natnre vitiates
the exi)erinuMits.

(h'senssed in the writings of Romanes and (inl'iek: the hilter has

from the theoretical standpoint and upon the ha>is ot" the very
limited knowledge of heredity of a few years ai^o. The eon( lu-
sion of these authors i.s that some kind of M-r(-i:ati..n or iM.hnion
is necessary for the ^necess of a new race. W hih- W a-iier 's!))

214

THE AMERICAN NATURALIST

[Vol. XLI

difference of breeding time, difference of local habitat, and phy-
siological properties precluding inter-breeding. The clear dis-
cussions of Romanes and Gulick have rendered superfluous
much in recent disputes on Isolation and Evolution.

Lately new conceptions in the theory of heredity have materi-
ally changed the conditions of the argument. Experiment has
shown that new characters may not be immediately swamped
by promiscuous breeding, but may on the contrary, in the fusion
of new and old races, predominate in full force over old characters
which they sometimes have the power of entirely subduing.^
While this result is very suggestive, too little is as yet positively
known to make an extended discussion at all profitable. Those
who are inclined to argue the matter may well take caution from
Davenport's opinion on the integrity of unit characters. "While
admitting, thus, the reality of unit characters, the further study
of the evidence of liybridization in poultry has led me away from
the conception that they are rigid and ininuitiil)]e as atoms are,
which may l)e combined and recombined in various ways and
always come out of the process in their j)ristine ])urity. This
is by no means the case. Very frequently, if not always, the
character tliat has been once crossed has been affected l)y its
opy)osite with which it was mated and whose place it lias taken
in the liybrid. It may be extracted therefrom to use in a new
combination, but it will be found to be altered. This we hav('
seen to be true for almost every characteristic sufficiently sludiod
— for the comb form, the nostril form, cerebral hernia, crest,
muff, tail length, vulture hock, foot feathering, foot color, ear
lobe, and both general and special plumage color. Everywhere
unit characters arc changed ])y hyl)ridizing.

"How does this fact 'bear i.u the ri\al theorii- of evolution?

It has an imporiant lu
the statements ,>f <lr\
of organisms arc built
from another,' and 'T
as between the moleci

1 Besides the Mendelum results s(>e also dc Vries (: 03, 2, p. 396 et seq.) on
the crossing of mutants with the parent species.

No. 484] CLOSELY RELATED SPECIES

215

comb is a different unit, but they are not sharply separable. Crest
and no crest are units, but they run into each other in hybridizing.
Unit characters may show transitions, and, if so, they may have
originated gradually, so far as I see. It does not follow that
they must have originated gradually" — Davenport (:06, p. 80).

Castle and Forbes's results with guinea pigs indicate the
same modifiability of unit characters. These authors (:0C, p.
13) say: "From the foregoing observations it is clear that, while
the long-haired and short-haired conditions are sharply alterna-
tive to each other in heredity, the gametes formed by cross-breds
are not in all cases pure. Frequently they consist of a blend
or a mixture of the two alternative conditions, constituting in
effect a new condition intermediate ])etween the other two. A
study of other characters alternative in iKMCility yields roults
somewhat similar,

"Albinism is, in heredity, the most sliarply alu'riintivc uf char-
acters, yet cross-ln-eedino- hvUxwu albin.. and pi-nuMiicd ouinra-
pigs may modify tlie cliavacler both of the albino racr and of die
pigmented one. This modification may take on a variety of
forms, as has elsewhere been pointed onl i( "a>tle, :().") i. It may
result in the production of mosaics ( piunuMiteil aniinai> spotted
with white), or of albinos with a modified peripheral pigmenta-
tion, or of albinos visibly like their ancestors hnt transnn'ttini;- a
different set of latent characters. A,n-ain, die ron-h or ro>etted
coat of certain races of n-uinea-pius is sharply ahernaiive to smooth
coat, yet cross-breeding of ron,uii with smooth races may indnce
curious modifications of the ronuli character or prodnce smooth
individuals bearing the merest trace of the roni:h cliaracter.

"All these facts are in harmony with the hypothesis, for which
there is strong evidence on the cytolouical side, that each sepa-
rately heritable character is represented by a differeiiT strnctnral
element in the germ (egg or spermatozoon!. In fertilization
the paternal and maternal representatives of a chaiacter become
more or less closely united, this nnion persistin-- ihroii-h all

sexual elements. At that time the paternal and maternal repre-
sentatives of a character separate from each other and pa.ss into
different cells.

216

THE AMERICAN NATURALIST

[Vol. XLI

"But the paternal and maternal representatives of a character
may in the meantime have exercised on each other a considerable
influence. In the case of some characters, as ear-length in rab-
bits (Castle, :05a), they completely blend and intermingle, so
that a new character is produced strictly intermediate between
the conditions found in the respective parents.

''In other cases the modification may be slight, as if the pater-
nal and maternal representatives of a character had been scarcely
more than approximated. Sometimes in cases of alternative
inheritance no influence of the cross is observable in certain of
the 'extracted' individuals, but if any considerable number of
individuals is examined, others will be found in which the cross-
breeding manifests its influence. From this we conclude that
gametic purity is not absolute, even in sharply alternative inheri-

These are very interesting qualifications of the Mendelian
principle of gametic purity. They suggest that new characters
might be swamped by repeated crossing, unless they were of
such overwhelming importance that they quickly won out in the
struggle for existence, to the immediate extinction of the bearers
of the older alternative characters. However, discussion may
here well wait upon further discovery.

But this may be said: If characters are gradually in()(lifiaV)le,
time becomes a necessary element in experinu iiTs on evolution and
possibly long periods of time may be needed for the demonstra-
tion of certain slow natural processes. For the present we may
well hesitate to accept the conclusion that Mutation is the sole
and only possible mode of evolution. Refreshing as the new
method of research is, in the midst of oceans of tiresome specula-
tions, and most valuable and even absolutely indispensable as
the i-e>ults already arc, the latter arc certainly small compared
to the hulk of our iuii(»rancc rcH-arding inorphogenetic processes.

and from infatuatio,. with ncw"i<lcas on the other, will look for
sonic iudci.cn.h'nl nutans of estimating the probable significance
of the new theories, it is the chief object of this paper to suggest
that such estimation may be rested upon the evidence of organic
geography when the evidence is available in sufficient body.

No. 484] CLOSELY RELATED SPECIES 217

If we were right in what was said above about the specific
effects of different modes of evolution upon (hstribution, the
first question to be asl^ed of the geographer is this: Are species
universally so distributed that each one occupies a region of its
own, or a habitat of its own; so that even the nearest related
species are strictly separated in sj)a('c. either in the broad geo-
graphic sense, or at least topograpliicallv /

It is to be noted that the inquiry lias iwo >t<'i)>. or >iauv- The
first relates to the distribution of organisms in thr br.,a.Icr x-iisc.

identical districts, or have largely coincident ranuv-. in many
cases. The adherents of the Mutation Theory exi.ect to lind a
considerable proportion of sucii instances, ("ertain of ii^ oppo-
nents have believed that the advance of the theory might 1 .l.x ked
on this first level. But if their efforts fail hvvv tluy are i)repare(l
to fall back upon the second line of <lefenee. 'n.e >.(<.nd stage

in relation to ecological condition^, and a>k> whether anv of the
allocated forms — if some are found cxi^i >i<le ly --idc with-
out even local .segregation. Di.sciples of de \'rie> expeei thai
instances will occur in such number> as lo s;iti>fy the ihMnaud-
of their theorv; while the opposit(> pariv thinks ilmi practically
no in.stances will be discovered giving counienance to tl.,- idea
of :\Iutaiion. Tluy expeei that all caM-. ..f -eneral geoo-raphie

tion, alfording practical is,,lation: and so hop.' to withstand th(>
final a.s.sault of the Mutationi>|s.

The application of the botanical eviiK-ncc j)resented in thi-
paper is to the first stage of the iiupiiry. .^omc of it is manifotis
applicable to the second stage also.

Specific 1 )isTh'i iu i i<>\ i\ tiik Am.mai. Kin<;i).»m

218

THE AMERICAN NATURALIST

[Vol. XLI

ance of Darwin's Origin of Species, first as an advocate of the
theory of Natural Selection, but shortly as its opponent. Through
twenty years of controversy he insisted upon the inadequacy of
Natural Selection, and as the prime factor in the diversification
of species sought to substitute Spatial Separation and to estab-
lish his own Law of IVIigration and Colony-formation. He sup-
posed a new species to arise by the migration or escape of a single
individual or of a pair from the domain of the old species into
new territory, where in geographic isolation and freedom from
the influence of the old stock a new race might be founded. The
divergence of the race from the old type he supposed to result
(Wagner, '89, pp. 286-295, 401) (1) from the individual peculiari-
ties of the parental pair or individual, which peculiarities in the
absence of the normalizing influence of interbreeding with the
whole body of the old stock would necessarily become accen-
tuated; and, (2) from the new environment. His theoretical
view^s, which throughout are questionable, are of less consequence
than the facts which he adduced in their support; the facts indeed
upon which he first formed these views. Wagner himself was a
traveler, observer, and collector in several i)arts of the wc)rld and
continually recurs in his writing to his ex])erience ni the field with
regard to endemic, narrowly r(

tant local varieties

elming numbers.

ill classes of animals and to some
extent also from plants. He represents specific distribution as
having a strictly mosaic or chain-like character. Everywhere we
find vicarious species and local races in separate habitats. Ihe
facts are presented at great length and with careful detad, and
seem to form a consistent body of knowledge, which impresses one
as being pregnant with a rational principle of wide import.

Mr. C. H. Merriam about a year ago addressed the zoological
section of the American Association for the Advancement of
Science on the topic, "Is Muta
the higher vertebr

(Merri

arguments

and conclusions ]

long, to my mind, too largely in the eonjeel uval
while this author's grasp upon the real character o t e iu>
work and upon his theory seems comparatively feeble, the da

No. 484] CLOSELY RELATED SPECIES

219

of distribution brought forward for several groups of mammals
are valuable. Dr. Merriam considers the geographic relations
of certain American rats, chipmunks, and ground squirrels; and
refers besides to other groups. His representation of specific
distribution agrees with that held by Wagner, with a qualification.
Merriam shows that the mammals in question occupy distinct
areas with very little exception, but that the areas often overlap,
and that the overlaps are likely to constitute narrow transition
zones characterized by the presence of intergrades. Actual phy-
sical barriers are often wanting.

President David Starr Jordan has also discussed the Mutation
question from the standpoint of organic geography and assembled
from his own experience and that of others a considerable body
of evidence regarding birds, while he himself speaks for fishes
(Jordan, :05). His paper, which appeared in Science a little
more than a year ago, contains some extraordinarily sweeping
assertions. He says: . . . .Moritz Wagner (1868) first made
it clear that geographical isolation (niumliche Sonderung) was a
factor or condition in the formation of every species, race, or
tribe of animal or plant we know on the face of the earth." The
principles set forth by Wagner ''have never been confuted,^
scarcely even attacked, so far as the present writer remembers,
but in the literature of the present day they have been almost
universally ignored." The question is much discussed whether
minute variations may serve to establish a new species in the
presence of a parent species, or whether wide fluctuation or muta-
tion may do so. "In theory either of these conditions might
exist. In fact both of them are virtually unknown. In nature
a closely related distinct species is not often found quite side by
side with the old. It is simply next to it, geographically or geolog-
ically speaking, and the degree of distinction almost always bears
a relation to the importance or the permanence of the barrier
separating the supposed new stock from the parent stock." "The
contention is not that species are occasionally associated with

' See the works of Darwin (72), Romanes, Weismann (72), and Niigeli cited
in the Bibhography. Weismann's paper relying upon the case of Planorbis
multiformis in the Steinheim chalk should be considered in connection with
Hyatt's Memoir on the same form ('80).

220

THE AMERICAN NATURALIST [Vol. XLI

physical barriers, which determine their range, and which have
been factors in their formation. It may be claimed that such
conditions are virtually universal. In a few cases, a species ranges
widely over the earth, showing little change in varying conditions
and little susceptibility to the results of isolation. In other cases,
there is some possibility that sahations, or suddenly appearing
characters, may give rise to a new species within the territory
already occupied by the parent form. But these cases are so
rare that in ornithology, mammalogy, herpetology, conchology
and entomology, they are treated as negligible quantities. In
the distribution of fishes the same rules hold good, but as the
material for study is relatively far less extensive and less perfectly
preserved than with birds and insects, we have correspondingly
less certainty as to the actual traits of species and subspecies,
and the actual relation of these to the intervening barriers."

President Jordan summarizes the distribution of species in a
law, as follows: "Given any species in any region, the nearest
related species is not likely to be found in the same region nor
in a remote region, but in a neighboring district separated from
the first l)y a barrier of some sort." That the intent of the law
involves both animal and vegetable kingdoms seems clear from

President Jordan says that his conclusions, much as they differ
from a priori judgments or the results of experiment, are the
unavoidable outcome of the study of distribution, and that they
are as a matter of fact "accepted as self-evident by every com-
petent student of species or of the geographical distribution of

Taking the facts of animal geography as they appear in these
several essays, typical of a larger number which might be cited,
we may say that as a whole they militate against the operation
of Mutation in any wide sense in the animal kingdom. This
conclusion is not prompted by the attitude of certain of the zoolo-
gists mentioned, who seem to have ma<le l)iif a curxny study of

No. 484] CLOSELY RELATED SPECIES

221

the ^Mutation Theory, but is drawn from the geographic evidence.
It is, however, true that the evidence is rather scanty. ]More-
over there are some exceptions to the general law of distribution,
and if these exceptions should, upon further research become
very numerous, the prejudicial force of the law would be much
diminished. But from the evidence at hand we may infer the
very general truth that animal species are distril)iit(Ml according
to Jordan's law of geographic isolation; that when exceptions
occur, the exceptional species are taken over into some other
category of isolation. The nearly universal patch-work char-
acter of specific chorolog}' — as at present depicted in the works
of zoologists — strongly suggests the gradual spreading out of
individuals over the surface of the earth, their settlement here
and there in isolated districts or topographically (h'stincr stations,
where shielded from promiscuous intercrossing they hav(> under-
gone transformations, which have been different in the (htlVrcnt
areas; transformations wliieh, advancing l)y whatt'ver force> or
conditions, whetlier tho>e of Xatiu'al SehM tif)n or of ortlion-cne-^is,

history is that which forms itself in the imagination of most students
of animal geography and has appealed most strongly to me as I
have reviewed the literature of the subject.

The Distribution of Plants.

Turning now to the vegetable kingdom we find, first, that
there have been few or no exhaustive (vssays (U'ah'ng with the
question of specific (hstrihniion in n'htiion to the theorv of evohi-
tion. In the second j)hice, it may be >ai(l at once that when
botanists have tm-ned their attention in thi> (h'rection their \ iews
generally do not coincide with tlio.->e of the zoologists as to the
nature of the facts.

NiigeH (73) opposed Wagner in a paper of uhich the purport
is succinctly ex|)resse(] in the iith\ "Die gt^scllschaftHche Mntstch-
ung neuer Spezies," the social oriyln of new species. This

acter of specific distribution, to the study of which he gave much
time in the field for several years, lie calls particular attention

222 THE AMERICAN NATURALIST [Vol. XLI

to the association of species of plants and their varieties upon the
same ground, and states that when one form replaces another in
consequence of change of ecological conditions within the same
district, the replacing form is not related to the other in the closest
grade of affinity, but in some degree more remote. He clearly
recognizes the intimate relation of distributional studies to the
question of evolution.

There is a little bit of evidence from Wallace (:00, p. 391).
He says he made inquiries of two experienced English botanists
to find whether well-defined varieties occupy areas to the exclusion
of the type and do not occupy the area or only a very small one
with the type. Only one such case was found in England. Wal-
lace's conclusion is that such varieties of plants occupying consider-
able areas to the exclusion of the type are not common.

Asa Gray ('59, p. 193) expressed the following opinion:
"Whether capable of scientific explanation or not it is certain that
related species of phienogamous plants are commonly associated
in the same region or are found in comparatively approximate
areas, however large, of similar climate."

The case of Draba verna L., is most interesting. As is well
known, about two hundred distinct species, or at least kinds, of
Draba have been distinguished within the limits of the original
I^inna^an species Draba verna. These numerous forms were
studied in cultivation by A. Jordan, and later by De Bary and
F. Rosen. They are found to come true to seed, and for this
reason are by these authorities spoken of as species. Their geo-
graphic distribution is discussed by both Jordan ('73) and Rosen
('89, p. 613). The conclusion is that as a rule the forms which
resemble each other most are found in the same stations. The
joint occurrence of next related species is indeed a fact which
])articularly impressed both of these writers. Rosen thinks that
it is very unlikely that these closely related species originated
separately and by chance came to be associated in the fashion in
which they are now found. Such an explanation might serve, he
says, if one or two cases only were to be explained ; but it becomes
absurd when we consider that the concomitance of next related
forms is wide-spread. Rosen ends his account of this group of
Draba species with a very clear statement of the mutative origin

^s^o. 484] CLOSELY RELATED SPECIES

223

which he is obliged to assign to these forms; without, of course,
using the terms of the Mutation Theory, which he partially antici-
pates by several years. "The Erophila [Draba] species owe
their existence to the free variation of their forefathers. This
consists not in a mere heightening or further development of single
characters, but variation fashions new characters and combines
old characters in new ways. Therefore the forms which arise
from species do not intergrade."

He says that, while Selection plays no part in the origin of these
forms it operates upon them after they appear. And of the laws
which must control this sort of variation he speaks as follows:
"Variation is not blind, vaguely working in all directions, but is
obviously determined by laws unknown to us: for we are obliged
to assume that the same or similar combinations of next-related
forms have arisen in different places. But what can these laws
be?" It is most interesting and significant that Rosen is led to
these de Vriesian conclusions through florisfic and geographic
studies.

The following excerpt from A. Jordan (73, p. 4) has so direct
a bearing upon our argument that I give it entire : " Ayant observe
dans leurs stations diverses, pendant plus de trente ann^es, une
foule de veg^taux de toutes les families et de toutes les categories,
des plantes annuelles ou vivaces, bulbeuses ou aquatiques, des
arbres ou des arbustes, j'ai pu constater presque partout que
lorsqu'un type linneen, vraiment indigene dans une contree, y
dtait commun a ce point qu'on pouvait le citer parmi les plantes
caractdristiques de la vegetation d'une certaine etendue du terri-
toire, ce type y etait presque toujours represente par des formes
diverses, plus ou moins nombreuses, croissant en sociSte et pele-
melc [ital. mine]. L'observateur superficiel, qui parcourt le terrain,
n'est frappe que des ressemblances de ces diverses formes; il
n'aper9oit pas leurs differences, ou, n'y attachant aucune impor-
tance, il ne s'arr^te pas a les consid^rer attentivement; il croit
n'avoir affaire qu'^ un type unique, susceptible de quelques
modifications accidentelles et sans valeur. Tandis que celui
qui observe avec attention pent aisement se convaincre, sur les
lieux, que ces modifications apparentes se retrouvent sur des
individus divers, tous parfaitement semblables entre eux. Si,

224 THE AMERICAN NATURALIST [Vol. XLI

pour pouvoir continuer et completer son observation, il arrache
des pieds vivants de ehacune des formes qu'il a pu distinguer
et les replante ensuite, dans un meme lieu, afin de les suivre dans
tous leurs developpements, il se convaincra bient6t qu'elles pre-
sentent des differences appreciables, dans tous leurs organes
S'il seme leurs graines, il les verra se reproduire avec une parfaite
identity de caracteres.

"Voila le fait que j'ai pu constater moi-m^me mille fois, que
j'ai fait constater dans les lieux que je ne pouvais visiter, en France,
en Corse et en Alg^rie ou ailleurs, par divers botanists qui m'ont
envoye soit des graines, soit des pieds vivants de formes nom-
breuses, recueillis dans les memes station et appartenant aux
memes types linneens. Je ne dis pas que les plants communes
soient toutes egalement et partout diversifiees. II y a, sous ce
rapport, de grandes differences entre elles. Je dis seulement
que le cas oil elles presentent diver ses formes croissant en societe
est le cas le plus ordinaire [ital. mine], et je crois que ce fait pa-
raitra clair, patent, indiscutable, a quiconque prendra la peine
de le verifier s^rieusement."

In the literature of this subject, as far as I have read it,
essentially the only writers who insist on the isolation of nearly
related kinds of plants are the zoologists. Their assertions
are not, however, supported l)y evidence from the vegetable
realm.

I have examined the distribution of North American Orchi-
dacese from the standpoint of this paper. Furthermore, I have
consulted with several specialists in different groups as occasion
offered. Several members of this club have given me information
with permission to publish it along with the evidence gathered
by myself. I may take the groups in sequence.

For Alga?, Mr. F. S. Collins speaks as follows in regard to their
general distribution and in particular the distribution of nearest
related species: "As regards fresh water algae, it almost seems
as if geographical limitations did not exist. Of course this is
not entirely true, but the area of distribution in the case of the
great majority of fresh water algse is vastly greater than in the

No. 484]

CLOSELY RELATED SPECIES

225

case of most flowering plants.^ The limitations seem to be those
of temperature, exposure, character of attachment, and to a less
degree, geological characters. Take ihv oumius \;nKlieria, for
instance. The last serious work is bv ( ioiz ; ;i >{\u\\ of tlie species
of Vaucheria in the neighborhood of Hast l. Switzerland. There
are 12 species there; 8 of them occur in Knijiand, G in the New
England States, 7 in California. Only three other fresh water
species are recorded for North America ; one is a P^uropean s})ecies,
found in the West Indies but not elsewliere so far on this continent;
the other two are from California. Now these two species, grow-
ing together, belong to the same subgenus, and I know of no
described species that I should say belonged in between them.
Take the genus Spirogv-ra. The best book on this is that of
Petit, Spirog}'res des Environs de Paris. He includes 37 species;
of these 34 have been found in North America. We have also
five other species; three of tliem are European, though not found
about Paris; the two others are from (ireenlaiid and Florida
respectively. It is mucli the same with all the fresli water alga-;
of the very inc<)iis|)icii()ns species, the records from (Hstant stations
are not so ahun(hint. bm tliat is laru'ely because the>e minute
forms have been little studied ont>ide of Knrope.

greater. It would seem strange that marine alga- on the two
sides of the Atlantic, should dillVr nmch more than the fresh
water a1g;v of the two coiuinents. hut snch is the fact. Still the
resemblances are nmch greater than with flowering plants.
And there are many instances where clos(>ly allied species or
varieties have practically the same range. I will give a few such
pairs, and in each case there seeni> to be no species or variety
anywhere else that would stand between the two in question.

"Cladostephus rnilcillafus and C. spon;/insus have the same
range, in temperate waters on both sides of the Atlamic. They
are the only species of the g(Mnis in that range. Fiicus (drntaftis
and F. evanescms have their hea.hiuarters in high arctic regions,

' Compare Alph. Dc C:inaollr. ( in.i,naphie Botanique, 1, p. 499: "Nous

d'autant plus petite que la classc dont elks font par k a une organisation plus
complete, plus dcveloppee, ou, selon Vexpression usU6e, plus parfaUe."

226

THE AMERICAN NATURALIST [Vol. XLI

extending to Great Britain, New Jersey and California. Myrio-
irichia filiformis and M. clavceformis have practically the same
range as the Cladostephus species. Ralfsia borneti and R. ver-
rucosa have a slightly more northern range on both sides of the
Atlantic. Phyllophora hrodioei and P. memhrayiifolia range from
France and New Jersey to Norway and Labrador. P olysiphonia
violacea and P. fibrillosa from Virginia to IMaine, from the Med-
iterranean to Scotland. P. harveyi and P. olneyi are American
species, or possibly varieties of the same species; they have the
same range as P. violacea and P. fibrillosa; wherever I found
one, I should expect to find the other.

"I could keep on for some time in this way, but will give only
one more instance; that is a group of species in the genus Anti-
thamnion, one of the most beautiful of the red algse. On the
European coast A. plumula ranges from Morocco to Great Britain;
A. cruciata about the same; .4. floccosa from the English channel
to high arctic regions; A. horeale from the Faroes north; A,
pylaiswi from Norway north. On the American coast A. plumula
and A. cniciatum range from New Jersey to Cape Cod; A. floc-
cosum and A. horeale from Cape Cod to Greenland; A. pylaiswi
from Long Island Sound to Greenland; A. americanum from
New Jersey to Portland, Maine. On our Pacific coast A. floc-
cosum ranges from California to Alaska; yl. horeale from south-
ern Alaska to high arctic regions; A. pylaiscei from Washington
north. Now these are all so closely allied that Rosenvinge some
time ago proposed to unite them all under the older name, yl.
plumida. He is a man with a strong tendency toward uniting,
it is true, and has since concluded that A. cruciatum, and possibly
A. floccosum are distinct; but at any rate, this shows how closely
allied they are.

"Some things about algje seem very much like supporting the
mutation theory; when the same species occurs in widely distant

Dr. Evans, while lacking the opportunity to ])ay very extended
attention to the subject, has given me the iollowinw' indication
of the distributional conditions in Hepatica\ "One of the l)est
examples of a cognate pair of species is Ijpfolrjrunca cllipfirn

No. 484] CLOSELY RELATED SPECIES

227

and L. exocellata. The first of these species is very widely dis-
tributed in tropical America, growing on the upper surface of
thick and glossy leaves. The second species is less abundant
but nearly always occurs mixed with the first.

"Among northern species Lophozia barbata and L. lyoni are
closely related and often grow together, although each retains
its distinctive characteristics. The same is true of Gymnomitrium
concinnatum and the much rarer G. corolloides; of Sphenolobu^
exsectus and S. exsectcBformis; of Anthoceros levis and A. puncfains.

"As a group of related species I might mention the ventricosa-
group of the genus Lophozia. This contains about half a dozen
closely related species, most of which are circumpolar in their
distribution. L. ventricosa, L. alpestris, L. porphyrolema, L.
longidens and L. confertifolia are all known from New England,
being most abundant in mountainous regions. Of these L. por-
phyroleiwa and L. longidem grow on rotten logs, and the others
on moist rocks, although L. longidens is equally at honu' on eitlier
substratum. Although I have no definite data that tlicsc species
actually grow mixed in North America, their ranp-s cdiiu iile to a

been at all accufately .studied', ali.l I feel sm-e tliat further study
would considerably lengthen the short list I have given."

Mr. A. A. Eaton has given me several examples from Equisetum
and Isoetes. "Equisetum jiuviatile is circumboreal in its distri-
bution. No third form stands between this and E. palustre, yet
the latter has practically the same range. E. scirpoides and E.
variegatvm arti a cognate pair, and yet both have in general the
same range throughout the northern part of both hemispheres.
E. variegatum has a variety, E. variegatum jessupi, distinguished
by anatomical characters. Its range, Vermont to Minnesota,
is quite covered by that of the species. E. hrvigaium has a near
relation, without an intermediate, in E. hiemale intermedium y.
and this on the other side is next to tlu> variety afme. The last
of the trio is wide-spread in northern North America and overlies
the other varieties, which also essentially coincide in their ranges.
E. arvense is found in Europe, Asia, N. America to Virginia and

228

THE AMERICAN NATURALIST [Vol. XLl

southern California. Its near of kin (without intermediate), E.
telmateia, is found with it (broadly speaking) in Europe and
California. Starting with E. pratense a next-related species is
E. sylvaticum. The former belongs to northern Europe, Siberia,
Alaska, Canada, the Rocky Mountains, Labrador, and south-
ward to Massachusetts and New Jersey. The other is circum-
boreal, covers the range of the first and with us goes somewhat
further south to Virginia.

"In Isoetes we find the following coincident ranges of close
relatives. Isoetes tuckermanni is foiiiul (|iiite plentifully in New
England and completely overlies the range of its varieties harveyi
and horealis. I. Engelmanni is found plentifully throughout the
region east of the Appalachian range, from New Hampshire and
Vermont to Pennsylvania, extending sparingly to (Jeorgia. It
overlies the ranges of its varieties carol in iana, foiitaua and valida.
I. canadensis is found from Pennsylvania to ]Maine and Quebec,
appearing again in British Columbia. Its next of kin in the genus
is /. engelmanni, whose range for the most part it covers, and the
two species are not rarely found commingled in the same pond.
/. holandrri is found from Wyoming to (^ilifornia and Washing-
ton. Its next of kin would apjx'ar to be /. piigniwa of the Mono
Lake region of California, and the two .-.pecics were found by
members of the King Ivxpcdition in contiguous areas. It may
be supposed that /. puijnuni is an abnormal form of /. holandiTi
and hence not competent in this relation, but the next of kin of
bolanderi is /. echinospora var. hrainiii, which overlies the range
of bolanderi, but is widely distributed otherwiM- in North America."

I have inquired of President lirainerd al)out the conditions in
Viola, and particularly whether pairs of closely related species
are found within the same range>. He an-wers: "Many ])airs

No. 484]

CLOSELY RELATED SPECIES

229

To Professor Charles Sprague Sargent I am indebted for inter-
esting information as to the distribution of North American
Crataegus. As is well known, numerous species have bc<'n (hs-
tinguished within the last few years, of which sonic five Inmdrcd
have been named. These species are readily and unniistakahlv
recognized by special students of the genus, by means of Horal
characters such as number of stamens, color of anthers, form of
inflorescence, etc.; by fruit characters, configuration of nutlet,
time of blooming and fruiting, character of foliage, veining, pres-
ence or absence of hairs, etc.; traits which appear to be constant
and reliable as shown by extended observation in the field and
by cultures of seedlings carried on now for a number of years at
the Arnold Arboretum. In these cultures, the sowings from the
several species result in crops of seedlings of remarkable uni-
formity within the limits of each species, and in the instances in
which the seedlings have flowered and fruited, of notable con-
formity to parental type. This result must certainly diminish
the scepticism with which the proposal of such a vast number of
species within the one genus has rather naturally been met in some
quarters.

In answer to the question whether the ncanv-r n lated species
are separated, as the law of 1). S, Jonhin and of Wagner w^ould
require. Professor Sargent re|)li('s in the m-uative.

In the genus as it is represented in North America several
groups are distinguished, which hi part (•()rresj)<)nd to the spec-ies
of the older writers, and whicii may lie reathly recognized by
anyone with a little attention- such are Crns-oalh, Punctata-,
iEstivales, Tenuifolite, Pruinosae, Intri( ata', llaht hata", Anoinahe,
Molles, Tomentosse, etc. These groups arc in p'neral fairly
well restricted to particular geograpliic x'ction^. For e.\anij)le,
the Tenuifolia^ the largest group in the northeast, do not cxtt'iid
west of tlie Mississippi river, or go southwani except alonu' tlie
mountains. The Flava^ are found only in the ^onthca^t. The

along the mountains southward to the end of the Alle-rhanies,
northward into Vermont, and westward through New ^'ork and
Ontario to southern Michigan, within which distributional area they
mingle with all the other northern groups. In some cases a group

230

THE AMERICAN NATURALIST

[Vol. XLI

predominates in a region, in other regions several groups are
nearly equally represented. Within each group, divisions can
be made; but in the case of these divisions geographic sepa-
ration does not obtain, since species of all the divisions of a group
are Hkely to occur in any part of the general territory proper to
the whole group. Regarding the ultimate units, or species, those
which are most closely allied are likely to be found promiscuously
associated in the same district and without the semblance of isola-
tion. For example species of the Pruinosse or of the Intricatse
with 10, or with 20 stamens, or with rose-colored, or with yellow
anthers are found growing within a few feet of one another, and
may cover common districts of several hundred square miles.
In these cases, while it is the number of stamens or color of anther
which first attracts attention, other specific characters exist which
adequately distinguish the species. As an example of promiscuous
association, the vicinity of Albany may be pointed out, where
the five species of Intricatse heretofore found in New York state
grow in a small area. In Ontario we find twenty-five species of
Tomentosse, many of them growing very close together. In the
distinctly southern group Microcarpse we find the two species,
C. apiifolia and C. spathulata, growing over the same areas, while
the third and more distantly related species, C. cordata has a some-
what more northern range. In general, the reverse of Jordan's
law would more nearly represent the distribution of American
species of Crataegus.

Coming now to Orchidacea?, I may say that I adopted the line
of examination suggested by the form of Jordan's law; that is,
I looked for pairs of kinds. I say kinds instead of species in-
tentionally. The main problem should not be confused by the
difficulty of agreeing upon a definition of species. What the evo-
lutionist has to account for is not the definitions of systematists,
but the multipHcity of hereditary types; he has to explain the
antithesis between the uniformity which heredity seems at first
to promise, and the diversity which actually prevails among
organic things. A definition of species is demanded in taxonomy,
but is somewhat less necessary in studies like the present. We
do not require that the foims be related in some particular taxo-
nomic sense; but only that they have different hereditary charac-

No. 484]

CLOSELY RELATED SPECIES

231

ters. In order to avoid complications I have used the word kind
to designate such different types, instead of the words species,
variety, etc., which have restricted technical senses.

I have sought for closely related pairs of kinds so made up that
in each case no third kind stands between the members of the
pair in resemblance. Such pairs I may call immediately cognate
pairs, or for short, cognate pairs. A pair may consist of two
species, two varieties, two subspecies, a species and a subspecies,
a species and a variety, etc. It is assumed that such cognate
pairs represent recent forkings of the phylogenetic tree; and that
if we could collect all such cognate pairs in the vegetable kingdom
we should have a representation of all the youngest forkings.
Evidently their distribution would be very illuminating, for the
youngest branches are on the average the least disturbed geo-
graphically, and the distribution of the members of these pairs
would represent as accurately as we could ever discover it, the
position of things at the moiiient when forking takes place. That
is, we should liave a ucouiaphic cliart, more or less distorted it
is true, of the origin of kinds. If the members of the pairs are
universally, in the vegetable kingdom, separated from each other,
then — as already explained — Mutation is excluded as a true
cause of diversification of hereditary types in plants. For among
several forms of isolation to which Mutation may conceivably
give rise, and which are, therefore, not inconsistent with the mu-
tational assumption, the one form of isolation to which it could
never give rise is geographic isolation.

I repeat that I have examined only the broad geographical
aspect of distribution and not at all the topographical, for which
exact data are wanting. Let the reader recall the two stages of
this general inquiry: my evidence belongs to the first of these.
I present the following facts as a contribution towards an answer
to the question. Is Mutation instantly excluded from a place among
the considerable powers in evolution, by the broad aspects of specific
distribution in plants? I have taken only one step. But this
may be of some little importance, especially in view of the asser-
tions concerning the distribution of plants which have been made,
and in view of the lack of even broadly geogi-aphical statistics.

232

THE AMERICAN NATURALIST [Vol. XLI

Evidence from the Family Orchidace.e in North America

The American Habenarias have been given careful study in
our laboratory. In addition to our own collections, those from
several of the largest herbaria in the United States have been
brought together. The species have been delimited with minute
attention and then the distribution of each species, represented
by the large amount of material assembled, has been recorded.
Thus exceptionally full and reliable returns have been secured,
which are available for the present paper.

H. ciliaris R. Br., and H. blephariglottis Torr., are a pair of
perfectly distinct, yet extremely similar species. While instantly
distinguishable in the field by their colors — the flowers of the
former being yellow or orange, those of H. blephariglottis pure
white — the dried specimens are separated only upon close inspec-
tion. The best distinguishing character is then the degree of
fimbriation of the lip, which is considerably greater in H. ciliaris
than in //. blepharigloiiis. No third species stands between
them. They are spread together through the eastern United
States. H. ciliaris is found in Massachusetts, Connecticut, New
York, Ontario, Michigan, New Jersey, Pennsylvania, Ohio,
Indiana, Delaware, Maryland, District of Columbia, Virginia,
Kentucky, North Carolina, Tennessee, ^Missouri, Arkansas, S.
Carolina, Georgia, Florida, Alabama, ^Mississippi, Louisiana,
Texas. H. blephariglottis is found in Newfoundland, Nova
Scotia, New Brunswick, all New England, New York, Ontario,
Michigan, New Jersey, Pennsylvania, Ohio, Virginia and North
Carolina; and if we include the southern form which may possibly
be distiuguislied, the range is extended to South Carolina, Georgia,
Alal.atiia, and Mi..i^.i].|.i. Whether we allow that the southern
form is distinct (m- not i> iininaienal, since it does not stand between
//. bh'pharigloffi.s and JI. rl/iaris, and the latter species covers the
range of both the northern and the southern forms of the other.

//. cristata R. Br., H. chapmanii Ames, and //. ciliaris R.
Br., form a group of very closely related kinds. //. cristata is
like a very small H. ciliaris, with a broader and un-clawed lip,
petals oblong or somewhat obovate instead of linear, and a spur

No . 484] CLOSEL V EEL A TED SPEC ' I ES

233

shorter than the ovary instead of longer. //. cIki juiidinl is iiuer-
mediate between the others in perhaps every resj)e( t, and fact,
with the absence of any new character of its own, makes this
species appear very hke a hybrid. Its apparently local occur-
rence is in favor of hyl)ridity. But allowing it to be independent,
it forms a pair with //. cristata on one side, and with //. oUiaris
on the other. On the other hand, removing llic ])Ianfs now
grouped as H. chapmanii, because of suspected hybrid origin,
we have left a very close pair in the two supposed j)a rental tyj)es.
On any disposition of the matter, the geographical ranges of the
three kinds are found to coincide widely. The range of H.
ciliaris, as above shown, extends from New England to middle
Florida and Texas, and inland to Michigan, Missouri and Arkan-
sas. That of cristaia includes all the Atlantic states from
New Jersey to Louisiana, with Pennsylvania, 'JV-nnessee and
Arkansas added. Specimens of 7/. chapmanii have been seen
only from northern Florida.

Hahenaria psycodcs (Jiay. and //. jii/.hriafa R. Br. are a very
close pair, with no interniediary. They are with some difficulty
distinguished, yet statistical studies that I made upon them some
time since convinced me that authors, including the most reliable
authorities, are right in considering them specifically distinct.
No one character can be relied upon to separate them in\ ariably
but all characters of each species fluctuate, so that any given
part in one may run into the form characteristic of the other s])ecies.
The l>alance of characters, however, is almost always decisive.
The geographic ranges are very largely the same. Both are
fomid in Newfonn.lland. Nova S.-otia. N<-w Brunswick, (Quebec,
all New Kngland, Neu York, New .le,-ey, Pennsylvania, and
North (/arolina. II. psi/nHlrs vxwiuU further we.i. and //.
fimhriata a little further M)uth. a-, represented in the .•olhutinns

H. pvramcrna Gray, has for it> prol.abl> n.-an^i rehnise //.
fimhriata — or possibly //. p.^ijcadrs ■ - \\\\\\ no species hciueen.
The three species mentioned, with //. h iH'ophaa , form a group of
close affinity. While JI. jx rama na is more w idely (hstributed
westward and southward (111., Mo.. Ala.), and //. jimhriata much
further northward, thev occupy extensive territory together; viz.,

234

THE AMERICAN NATURALIST [Vol. XLI

Pennsylvania, West Virginia, North Carolina, and Tennessee.
If H. psi/codes is substituted for H. fimbriata in the comparison,
the geographic result has the same influence on the discussion.

H. orbiculaia Torr. and H. macrophylla Goldie are so close
that the plants of the two kinds have long been accepted by col-
lectors and described by authors as of one species. The differ-
ences are at first sight slight, but are apparently constant and
sufficient for distinction. The former species has a much greater
range, which completely covers that of the latter. II. orbiculaia
extends from Labrador and Newfoundland westward through
Michigan, and Minnesota, to British Columbia and Washington;
and southward through New England, New York, and Pennsyl-
vania to South Carolina and Tennessee. It is found in every
district where H. macrophylla is found; viz., Newfoundland,
New Brunswick, Ontario, Michigan, New England, and New
York. The status of these two species is discussed by Ames in
Rhodora for January, 1906, with illustrations of the flowers.

The diflBcult genus Spiranthes has lately been thoroughly
studied by Ames, who has given the results in Orchidaceje, Fasc.
I, pp. 113-154. The abundance of material examined may be
seen from the citations of specimens in the detailed statement of
the distribution of each species.

*S. cernua Rich, has for nearest allies, first, the variety (which
some authors regard as a species), S. cernua var. ochroleiica Ames,
and secondly the species S. odorata Lindl. S. cernua may be paired
with either of them. 8. cernua and S. cernua orchroleuca in the
dried state can be separated with certainty by no macroscopic
character. They may be distinguished by the seeds, however,
S. cernua being polyembryonic.^ Unfruited specimens being
indistinguishable in the dry state, the exact distribution of each
form may not be very precisely defined, but Rydberg in Britton's
Manual gives the range of var. ochroleuca as from New Hampshire
and Massachusetts to Pennsylvania and North Carolina. I
myself have identified, as being unmistakably typical S. cernua^
specimens from Massachusetts, Ontario, Iowa, and Georgia.

^ See my notes on the embryology of the two forms in Rhodora 2, p. 227
(1900) and 3, p. 61 (1901). In S. cernua embryo formation takes place without
poUination.

No. 484]

CLOSELY RELATED SPECIES

235

Thus the range of the species overHes that of the variety. Geo-
graphic isolation is wanting.

When we compare S. cernua with *S. odorata we find again a
very strong Hkeness. S. odorata is usually much larger in all
parts than the former. The length of the scape relative to
that of the leaves is greater in S. odorata, and its leaves are less
strictly radical. Those not expert in the genus Spiranthes would
often distinguish the two species with difficulty. They might be
regarded as elementary species in de Vries's sense. S. odorata
has been found in Virginia, Georgia, Florida, Alabama, Louisiana,
and Texas, and its range thus coincides widely with that of S.

S. romanzojjiana Cham, and S. porrifolia Lindl. are very closely
related species, which no other species approaches. The former
is by very much the more widely dispersed, since it crosses the
continent, while S. porrifolia is confined — according to speci-
mens seen — within the states of Washington, Oregon, and
CaHfornia. S. romanzoffiana is represented in our records by
many specimens from these same states, and there is therefore no
general geographic separation in this case.

S. laciniata Ames and S. vernalis Engelm. and Gray are ex-
tremely similar but distinct species. The former is confined to
Georgia, Florida, Alabama, Louisiana and Texas. S. vernalis
occurs in all these states, but reaches far beyond tliis area.

S. beckii Lindl. and S. gracilis Beck are an iniiiu-diarely counate
pair of near affinity. The former grows in the Atlantic states
from Massachusetts to Texas. S. gracilis covers the same range,
but is also to be found further north and further inland. There
is no geogi-aphic isolation.

Cijpriprdium puhescens Willd. and C. parvi forum Salisb. have
had attention at tliis laboratory for several years, observations
having Ixcn made in the field and in the herbarium, and coUec-

Measurements indicate that tliere are two pronounced tendencies
as regards size of flower. In life, the plants generally have an
appearance of thstinetness, and most field naturalists whose
opinions have been asked, have maintained that the two kinds are
specifically different. The manuals treat them so. Yet they

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THE AMERICAN NATURALIST

[Vol. XLI

occasiouaily intergrade and perhaps can be regarded merely as
subspecies. I have found them growing together in closest
proximity. We have specimens of C. farvifloruin from Ontario,
British Columbia, New England, New York, New Jersey, Penn-
sylvania, Ohio, Indiana, Michigan, Wisconsin, Washington;
and of C. pubescens from New England, New York, Pennsylvania,
Ohio, Illinois, Wisconsin and Minnesota. The manuals extend
the reported occurrence of both plants to Georgia. Thus the
ranges of these two very closely allied kinds coincide over a very
wide extent of territory. No other form in the world stands
between them.

The genus Calopogon is confined to the eastern half of the
United States (if we except the occurrence of C. pulchellus in
Cuba), and comprises four close species and a variety, or five
species. These species all come together and overlap in Florida.
One, C. pulchellus R. Br., ranges from Newfoundland to Florida,
Cuba, and Missouri, and geographically includes all the rest. C.
pallidus Chapm. ranges from North Carolina to Florida and
Alabama; C. parviforus Lindl. from North Carolina to Florida;
C. multiforus Lindl. is confined to Florida. Here, therefore, are
several pairs of cognate species not geographically separated.

Pogonia vcriiciUata Nutt. and P. affinis Austin make a pair
of very nearly related yet distinct species. The former extends
from New England to Florida and west to Wisconsin and quite
surrounds the other, a very rare species occurring sporadically
in Vermont, Massachusetts, Connecticut, New York and New
Jersey. Here again geographic isolation is wanting.

The conclusion from this examination of North American
Orchidacese is that cognate pairs of kinds w^ith uniform or widely
coincident ranges are too numerous to leave any force at all in
Jordan's law in its broad sense as regards this family in our flora.
If one member of each pair w^as derived from the other member,
or both were derived from a parent species, then, as jar aft the
geographic evidence goes, the new species may have originated
in the same district with the old one; i. e. without geographic isola-

No. 484]

CLOSELY RELATED SPECIES

237

CoxcixDiXG Remarks.

In concluding this paper I may make some remarks of a general
character touching the whole prohlem.

First, we note that zoologists and botanists are rather distinctly
opposed to each other in their views of the actual state of specific
distribution. The suggestion is offered that zoologists may best
discover the condition and interpret its meaning among animals,
and botanists among plants. In no case is it safe to reason deduc-
tively from one kingdom to the other. In the factors affecting
their evolution plants and animals differ vastly.

Secondly, in seeking for the laws of specific distribution we
should first take the facts as we find them. We should acrree
to consider that in the absence of explicit evideucc to tli<> coiin-ary,
kinds now found in coincident ranges have been so situated from
the beginning. In any given cases this assumption may or may
not represent the truth, but we have no right to postulate move-
ments in the past, of which there is no certain evidence, in order
to save a preconceived theory. We may call sucli hyj)oth(>tical
migrations into being, in a strictly hmitcd number of ( mm's. \ipon
a reconsideration, if from a first cxaininaiioM ot the unuio.hlied
facts some law emerges so strong and com])ulsory that the few
exceptional instances must somehow be brought into Jiccord with
it.

In the third place, if I may express my personal iniy)ression of
the matter with regard to plants, it seems to me that tlie sttidy of
specific distribution in the vegetable kingdom is not hkely to be
unfavorable to Mutation, regarded as a method, but ])erhaps
not the sole method, of evolution. It is true that in examining
the distribution of species of plants, one encounters an effect
which seems to be connected with geographical (h>iant'e. \Veoftt>n
find that a species of wide distribution exhil)its shghtly (htferent
phases in different divisions of its range. These phases are some-
times too subtle for definition and pass into one another by degrees,
yet are evident to students of particular groups. Such cases do
not look like the work of Mutation. They exemplify that which,
to conceal ignorance of causes, may be termed a geographic effect.

238

THE AMERICAN NATURALIST

[Vol. XLI

But this aside, the indications seem to me to be that a good many
instances sustaining the notion of mutative origin will be found
among plants. It is not to be expected that the number of such
cases will be relatively large. On the assumption of periodic
Mutation as the origin of species, with competition between
associated mutants and the survival of those mutants best fitted
for existence in the original habitat, and the spread of mutants
with new capacities into areas or habitats not open to the parental
species, we should expect to find as a rule a single species occupy-
ing a given territory or ecological footing, and related species in
separate, neighboring areas or habitats; though it is evident
that mutants instantly endowed by Mutation with physiological
or chronal isolation might continue to exist side by side with the
parental species or with sister mutants if there were no active
vegetative competition between the associated stocks. As a
matter of fact, in many species of plants competition for sub-
sistence between individuals of the same parentage is practically
absent. Unification of congenital mutants may be brought about
by continued interbreeding. This would eventually destroy the
geographic evidence of Mutation in any given case. But in such
amalgamation the effects of Mutation may not be destroyed ; for
new characters may during amalgamation be perpetuated in full
force. It is single characters, rather than constellations of charac-
ters, with which the Mutation Theory is primarily concerned
The number of cases of association of closely related species
resembling recent mutants, in proportion to the number of cases
of geographic or topographic segregation of closely related species
would depend upon the balance betv^^een the activity of Mutation
on the one hand and the operation of the forces tending to isolate
or to amalgamate the products of Mutation on the other. If
mutative periods are far apart in most species — and stabihty
of the organic world may preclude great frequence — while the
segregating or amalgamating powers are constantly at work,
then the occurrence of the social condition indicative of Mutation
may be expected to be relatively infrequent.

In order to use geographical evidence effectively against the
Mutation Theory, its opponents must show that the social con-
dition of closely related forms is, to use President Jordan's words.

No. 484]

CLOSELY RELATED SPECIES

239

"virtually unknown." In the vegetable kingdom this is likely
to be an arduous task. The indications are that the adherents
of Mutation will be able to bring forward enougli cases of social
distribution to render phytogeogra})hic woaj)ons useless in the
attack upon this Theory.

LITERATURE

Castle, W. E., am) Foubks, Alexanokr.

1906. Heredity of Hair-length in Cuinea-pigs and its hearing ...i die
theory of pure gametes. Published by the ( ai iu^iiif bistituiion
of Washmgton, D. C.
Darwin, Charles.

1859. The Ongm ot hp. U( ^ .h.pt,,. 11 „ I IJ (..n-Mphi, .1
Distribution.

1872. TheOiiginof .sp(<i<s ()tl. I .litiu,, J, ipiu 1 ,^ i .i i^iti.ht.^
to Isolation a.ul to DivergennM.f Clmrartrr.

1906. Inheritance in Poultry. Publishr,! l.v tlir ( •;.r,u"-ir liisiituiion
of Washington. D. V.
De Vriks, If.

1905. l.xolmioiK racial and habitudmal. Published by the Carnegie

240

THE AMERICAN NATURALIST [Vol. XLI

MacDougal, D. T.

1906. Discontinuous variations in pedigree cultures. Pop. Sci.
Monthly, Sept., 1906.
Merriam, C. Hart.

1906. Is Mutation a factor in the evolution of the higher vertebrates?
Proc. A. A. A. S., New Orleans, 1906, p. 383. Also Science, N. S.,
23, p. 241 (1906).
NXgeli, C.

1873. Die gesellschaftUche Entstehung neuer Spezies. Sitzungsb.
d. math.-phys. Klasse d. k. b. Akad. Munchen, 3, p. 305 (1872-3).
Romanes, G. J.

1886. Physiological Selection. Jour. Linn. Soc, Zool., 19, p. 337.

- 1906. Darwin and after Darwin. Vol. 3, Isolation and physiological
selection. Opinions on isolation [with clues to valuable data of
distribution]. The appendices contain important matter.

Rosen, F.

1889. Systematische und biologische Beobachtungen iiber Erophila
[Draba] verna. Bot. Zeit., 47, pp. 565, 581, 597, 613.
Wagner, Moritz.

1889. Die Entstehung der Arten durch raumliche Sonderung. Gesam-
melte Aufsatze. Basel. This edition contains the following pa-
pers of importance: Die Darwin'sche Theorie ujid das Migra-
tionsgesetz der Organismen. Leipzig, 1S6S. — UcbcM- (1(mi Eiiifluss
der geographischen Isolirung und Koloniciit)iMuiijj: ;nit die morph-
ologischenVeranderung der Organism. :\IiiiHli.ii, ls7(), Sitzungsb.
d. k. bayer. Akad. Wiss. 2 July (1870).— I'rln-y die llnisichung
der Arten durch Absonderung. Kosnios. lite. I, L', :!. ilssO). —
Darwinistiche Streitfragen [5 pajHTs]. Ko-mu^ ! IssJ. issh.
Wallace, Alfred Russell.

1855. On the law which has regulated th.' int mdurt iu,, nf nrw species.
Annals and Magazine of \at. History, Septeml.er. is.V.. Also
in his Contributions to Xatural Sel..'tion. [.o„.lnn, is;:., p. .).

1900. Studies, scientific and >o, ial, 2, p.
Weismann, August.

1872. Ueber den Einfhi.^ d<T ls(,lirun- auf die Anluldnn- Leipzig,

THE COINCIDENT DISTRIBUTION OF RELATED
SPECIES OF PELAGIC ORGANISMS AS ILLUS-
TRATED BY THE CHJi:TOGNATHA.
CHARLES ATWOOD KOFOID

No small part of the diversification of the organic world has
taken place in the open sea. Whether we accept the view that the
littoral and abyssal faunas are derivatives of the pelagic, or regard
the latter as secondarily derived along many lines from the organ-
isms of the shore and bottom, the fact remains that many groups
have undergone great diversification both in the specific and in
higher categories in the pelagic habitat. Illustrations of this
process are to be found in the diatoms, the Protozoa (notably
the Foraminifera, Radiolaria, Dinoflagellata and Tintinnoina),
in the Scyphomedusa^, Siphonophora, and Ctenophora, Ostra-
coda, Schizopoda, Amphipoda, Decapoda, Heteropoda, Ptero-
poda, Cephalopoda, and Tunicata and certain families of fishes.
The Nemertini, Aimelida, Rotifera. Holothuroidca and the llcni-

are exclusively marine and ])e]agic, and their aflinitirs arc with
the more primitive types of invertebrates. It seems })r()l)able
that their entire evolution, or at least their generic and si)ecific
differentiation has taken place in the marine habitat. Their
present distribution is therefore of prime interest because of its
bearing on the relation of isolation to the origin and ))reservati()n
of species.

Barriers are far less in evidence in the environment of the j)ehigic
fauna than in that of the shore or of the land. A t\nv instances in
limited regions along the margins of great ocean currents as, for
example, along the edges of the Gulf Stream or in liorizontally
stratified waters, there are abrupt transitions in temperature,
but in the n)ain t]w changes in temperature, ilhnninution, density,
and substances in solution or suspension, are so gra<hial that
zoological })rovinces are delimited with difficulty and mainly in
terms of tempcratiu-e, on the high seas away from the infiuence of

242

THE AMERICAN NATURALIST [Vol. XLI

shore conditions. In a large and somewhat vague way isotherms
and isothermobaths constitute the barriers of the sea. Many,
and in some groups, most of the pelagic species are wide-ranging,
found in most seas, through a greater or less range of temperature.
The pelagic fauna has thus a considerable cosmopolitan element
and part of the differences which result in the contrasted poverty
and richness of pelagic fauna are due to changes in the numbers
of individuals and in the proportionate representation of the
various components, as much as, or even more, than to restrictions
in the distribution of species. In so far as the species of any group
of related organisms establish themselves throughout a wide,
coincident or overlapping range, in like degree isolation becomes
problematical as a factor in the origin of new or preservation of
old species.

Our knowledge of the horizontal and vertical distribution of
pelagic organisms is lamentably incomplete and partial, and no
less so of the Chsetognatha than of other groups. Fowler (:06)
calls attention to the fact that he finds no published record of a
single species of that group between 160° E and 80° W, nearly the
whole of the Pacific Ocean! Unfortunately no report was pub-
lished on the Chaetognatha of the Challenger Expedition and
the results of later surveys have not yet appeared. We find,
however, an excellent summary of the known distribution in
Fowler's (:06) report on the 'Siboga' collections, based largely
on his Biscayan investigations, Fowler (:05), and the wwk of
Doncaster (:03) on the jVIaldive and Laccadive fauna, of Aida
('97) on that of Japanese waters, of Steinhaus ('0()) and Strodt-
mann ('92) on collections from the Atlantic, and of various re-
corders in the lists of the Conseil permanent pour V exploration
de la Mer, from the waters of Northern Europe. The data thus
assembled by one whose critical knowledge of the species has
enabled him to sift out svnonyms and eliminate probable errors,
are far from being adecjuate to give a complete or satisfactory pre-
sentation of the distribution of Ch^tognatha in the seas named.
They are, nevertheless, of sufficient fulness to afford a basis for
the consideration of the extent to which isolation of species prevails
in this typical pelagic group of organisms and to mark out clearly
the necessity for additional data on vertical distribution and
breeding seasons for a critical and final analysis of the problem.

Ko. 484]

DISTRIBUTION OF CH.ETOGNATHS

243

It is the purpose of the present note to call attention to the
important contributions which investigators of pelagic h'fe might
make to the discussion of this phase of the prol)lenis of evohition
especially since monographers of pelagic groups are best qualified
to judge of the degrees of affinity between the species of the genus
and can determine whether the most closely related ones have a
coincident or contiguous distribution. It is exceedingly desirable
that future expeditions investigating the life of the high seas be
equipped for a fuller analysis of the details of vertical distribution
and that data on breeding seasons of pelagic species be included
in monographs whenever available.

Genus Krohnia

This genus includes three species, K. hamata, K. siibtilis, and
A', pacifica. The first are two oceanic species of wide distribution,
the last an Indo-Austral species of surface neritic distribution.
The horizontal area of distribution of the first two species is largely
coincident, A', hamata being known to extend to higher latitudes
(81° N., 52° S.) than A. subtllis (Oif X., 2\)° S.). As might be
expected from its temperature relations, A', hamata is recorded
from lower levels in the tropics than is A. .suhtllis. Data on this
point are not very complete as K. suf^fllis is not an abundant
species. The closing net catches of the Plankton Expedition
indicate a maximum depth of 1500 m. for A. hamata and S5() m.
for A', subtilis. The two occur together between 300 and 500 m.
(37° N). The extent to which the vertical distribution of the
two species overlaps cannot be deterniined from tlie available
data. Fowler (:05) shows that tlie size of the individual of A.
hamata increases with the depth in the Biscayan region. The

and large ones with occasional small ones 1h>I.uv that level. The
sexual conditicm at different levels was not noted. The |)()ssil)ility
of overlapping distribution is certainly present l)ui contiguous
distribution is by no means e.\( lude<l.

Krohnia hamata is foinid in the mesoplaiikton of the Indo-
Austral region, where A', pacifica is also found, but in surface
waters exclusively. These two sj)ecies were thus contiguous

244

THE AMERICAN NATURALIST

[Vol. XLI

rather than coincident in their distribution. There is thus little
conclusive evidence of coincident distribution in the few species
of Krohnia.

Genus Spadella

The case of the two species of Spadella, S. cephaloptera and
S. draco the area of distribution of the latter, which is a wide one,
includes that of the former which is a neritic species from the
northwestern coasts of Europe and the Mediterranean. They are
both surface forms and their distribution is of the coincident type.

Genus Sagiti'a

The genus Sagitta as revised by Fowler (:06) includes twenty-
one species. Their general horizontal and vertical distribution
is shown in the accompanying table taken from Fowler's (:06)

J

I

1

1

I

1

1

1
1

1

1
1

1
1

I

1

1

i

i

1

i

t

i

t

t

t

t

i

\

i

:

(Siboga) report. S. bipunctata is omitted by him from the Indo-
Austral region in his text because of the uncertainty of its identifica-
tion since it is quite similar to the young of several other species
in the list. Of the twenty-one species, eleven, including S. bipunc^

No. 484] DISTRIBUTION OF CH.ETOGNATHS

245

lata, occur in the Atlantic, ten in the Indo-Austral, eight in Jap-
anese waters, and two in the siibantarctic, in the e[)iplankton. In
the mesoplankton of the i\.tlantic eight species are found, and
three in the Indo-Austral. In the hiru'cr ocoirrapliical rcLn'ons
of the Atlantic we find coincidently in ilic cpiplankton, in tlu-
Arctic, three species, in the subarctic five, in the north KMupcratc,
eight, in the tropical, five, in the soutii temperate, four; in the

TEMPERATURE IN DEGREES CENTIGRADE

246

THE A M ERICA X NA T 1 1L\ LIST [Vol. XLI

20° to 32° — 11

This, in conjunction with the fact that some of the low tempera-
ture species belong to the mesoplankton of the tropics, indicates
that the center of radiation of the (;enus has been in the tropics,
or that specific differentiation has been relatively more rapid in
that region than at the lower tenij)eratnres toward the poles.

These broader outlines of the distril)uti()n of the species of
Sagitta are suggestive of a considerable degree of coincidence of
distribution of species, it may be of closely related ones, and
prompts to a closer analysis of their relationships and distribution.

The determination of degrees of relationship among species of a
genus is a matter of inference from structural details for whose

guided by the selection of characters on whicli chissification is
based, l>y experience in deahng with the specific analysis of the
material, and subjectively, by the conception of sjx'cies which one

modified structures, some clue t.^ the distance of their removal
from the i)arent stock, or from each other. On the other hand
the elementary >pecie> arising by nuUation from <Knnthrra Immirvk-
iana mav be regarde.l a> geneti<-ally e<|nally related t.> each other
or to the parent .to<-k, but if we bas,> oiu' judgment of the degrees
of the relationship which they exhibit solely on the structural
characters whi. li di.tinguidi them, wc wonM be for.vd to conclude
that there wa. considerable disparity of relationship among them.
The n.ntation tlicry admit, a wid.^r latitn.i.' in <'sti,naling the
relati<.n.hip of .pccic. than docs tlu- unmodiiic.l Darwinian point

We have, however, in Sagitta onlv the ronit, and n..t the process
of si>ecific .litrerentiati<.n with which to deal, and are therefore

No. 484] DISTh'lllCTfoX OF (11 .KT< HiXATHH 247

forced to depend >M\ stiuctiiral tv.s(>iiil)lances for the

determination of sjxM'ilic rrlationsliips.

The s])e('ies of ^auiiia aiv distinouislicd, anion^ other less
quantitatively ('N])Itssc(1 cliaracters, by (1) size, (2) ratio of tail to
total leno'ili. (.{) miinlHT of jaws, (4) number of anterior and (o)
posterior tivtli. An analysis of Fowler's (:0()) specific diagnoses
reveals three t,n-ou])s of related s])ecies within wiiich couplets of
most closely related species may be noted.

The first of these, the srrratodoitata otouj), includes five species:
S. serratodentata, a eurythennal cosmopolitan s|)t>eie> with little
tendency to sink to deeper waters in tlie tropics; .S. Ixdoli, a
neritic surface species from Indo-Austral waters; N. /criKv and
<S. rohuMa, neritic and surface species from the Malay and Maldive
Archipelagos; and 8. siboc/w, taken only in hauls from dee{) water
in the Malay Archipelago.

The following table of cjuantitative characters of the species
taken from Fowler's records serves rather to indicate their close
reseml)lance than to dilferentiate them. Other characters such
as proportions, form of the (yes and teeth, assist in dia^mosis.

sibogce 9-20 21-83

ferox l()-2() 29-36 r,-G

248

THE AMERICAN NATURALIST

[XoL. XLI

age of coincidence 80%. Of the 65 collections 55 were made at
the surface. Four of the five species thus have a coincident dis-
tribution, including the mostly closely related couplet ferox-
rohusin and the very closely related S. hedoti and S. serratodentata.
The hedoii-sibogos couplet appear to have a contiguous distribution
in the upper and lower levels, respectively, in this region.

A second group of species which show considerable resemblances^
to each other are S. hexaptera, an oceanic, stenohyaline, eury-
bathic, and eurythermal form; S. enflata, a warm water form of
wide distribution in the e})ij)lankton of warm-temperate and
tropical seas; and S. pidchra, a neritic surface form from the
Malay and Maldive Arcliipelagos.

The accompanying table indicates the relationships of the
three species of the hexaptera group as suggested by the quantita-
tive characters.

hexaptera 15-70 20-25 6-8 8-4 2-7

pulchra 9-22 18-27 5-7 6-9 10-15

euflata 22-26 16-22 7-9 7-10 12-17

An examination of the 'Siboga' lists shows that the three species
occur together irs 24 catches, two in 26, and but a single one in 26,
the {)er(rentage of (.'oincident occurrence l)eing ()(>%. The most
closely related couplet in this group is pulchra-enfiata, the former
a neritic, the latter an oceanic si)ecies. 'I'liese two occur together
in the Maldi\^s and also in the 'Siboga' collections, where *S.
enflata is one of the most abundant species. It is found in every
one of the 34 collections in which S. pulchra appears. Of the 34
coincident occurrences 29 are in surface collections. Those three
related species have here a coincident distribution and N. Ju.ra pfera

A third grony) of related species includes N. hi putivfafa , and two
(•()nj)lets of most closely relate<l species, [areata- pldiictoii Is and
iHujh'cfa -re(/idari.s. I'nblislu'd records indicate that tli(> first
named species is a cosmoj)(>litan one of wide range. 1 )ifliculti(vs
attend its specific determination so that Fowler is of the opinion
that it is possibly only an Atlantic neritic form not occurring in
Indo-Pacific waters.

No. 484] DISTRIBUTION OF ( U.KK u.S.\ TIlS 249

The members of the first couplet, /urco/r/ i>!(iiic/>>n is. a iv Atlantic
species, the former of wide distribution, '>\ X. to 7 S., in the
epiplankton of colder waters (17°) and the nicsoplankton of the
tropics. The latter occurs only in the epiplankton of the tro|)ics.
This couplet of most closely related species has a contiguous
rather than a coincident distribution. The distribution of both,
however, is overlapped by that of the very closely related S. hi-
punctata. The degrees of relationship as suggested by quantita-
tive characters may be inferred from the accompanying table.

regularis 4.5-7 28-40 5-7 2-4 4-6

The members of the second couplet of most closely related
species, S. regularw and SI. ncglecta, are both surface neritic forms
of the Malay Archipelai^o and Japanese waters. .S'. rrr/u/an's is
neritic also about tlie MaMives and ir may !)<■ that 1 )<>iicaster (:03)
overlooked the very similar N. iiKjlicfn in the eoUeetions from
these waters. The distribution of these t\v.. most cIom Iv related
species is thus widely overlapping, if not in(h-ed eoiiK ident.

The distribution of pelagic organisms, a- ilhisrrated by the
Chtetognatha thus affords several probable in>taiices ,,f the isola-
tion of the members of couplets of most closely related ^iieeies hy
isotherms or isothermobaths. This isolation is similar in many
of its aspects to that so often foimd hetweeti terrestrial speci.^s.
It may well be that isolation has heen an essential factor in the
differentiation of the members of these couplets. l-:\en more
general, however, in the pelagic world and amonu- the species of

We have now no evidencr of 'ditrere.uial s.-as,,,',.. temper '.lures,
or levels at which breedin- mi-ht occur in thes,> closely related
species. Should these ditferentials ultimately prove to be absent

250

THE AMERICAN NATURALIST [Vol. XLI

we would be forced to conclude that isolation has had no part in
the origin, differentiation, and continuance of these related species.

In Dagitta bipunctata Miss Stevens ( :03) has described a method
of close fertilization. As yet we have no light on the extent of its
occurrence in other species where the presence of enlarged seminal
vesicles and external male parts affords suggestive though not con-
clusive evidence of external and presumably of cross fertilization.
Should all species of Chsetognatha prove ultimately to have close
fertilization we would have in this a most effective means of isola-

The apparently wide-spread phenomenon of coincident dis-
tribution of related species among pelagic organisms appears to
cast some doubt upon the universality of the operation of isolation
in the evolution of species as originally maintained by IVIoritz
Wagner ('68) and recently revived by President Jordan (:05).

The contrast here afforded also raises the question whether the
two types of 'species' really belong fundamentally to the same
category or not. Are those with contiguous distribution, and also
many of the geographical species and subspecies of land verte-
brates, of a standing exactly equivalent to that of those having a
coincident distribution? Are, for example, S. furrata and S.
planctonis merely the extremes of an environmental series begin-
ning in the warm surface waters and ending in deep waters of
lower temperature? In other words are they the result, in part
at least, of the pressure of the environment ? A statistical study
of the distribution and variation of such a pelagic couplet and a
comparison with a similar study of a couplet having a coincident
distribution would be most instructive in indicating whether or
not any distinction exists between 'isolation-environmental' spe-
cies on the one hand and 'selection-nuitation (?)' species on
the other. Are intermediate forms (Hjiially absent in both types of

the Llivi.huds of the two types? Ab..vr all will tl„- in.livi.luals oi

ronnuM.ts are trimsp..>(Ml ? An.l linally will tlir ,>,Hvi,«> with coin-
cident <listributi()n exliibit any greater specific ..tal)ility ^un<ler

Investigators of pelagic organisms have been morphologists so

No. 484] DISTRIBUTIOX (>F ( ■/IJ:T()( ,.\ ATI/ S

251

generally, rather than primarilv svstcinatists, that ilu- lH'ariii<^ of
the data of the geographical (h'strihutioti of the ofixaiiisins with
which they have been dealino-, upon the l)roa<lcr jjiohlcms of
evolution has been somewhat iicolccftMh It is on>atiy to l)c hoped
that the life of the sea, primitive, ancient, (Hversified as it is, may
yet shed some light upon the problems which this brief paper can
do little more than suggest.
Zoological Laboratory

University of Califorxia
Dec. 15, 1906

BIBLIOGRAPHY

AlDA, T.

'97. ChcTtogiiaths of Misaki Harl)or. Annot. Zool. Jap., Vol. I,
pp. 13-21. Tab. III.

DONCASTER, L.

:03. Chjptognatha, with a note on the variation distribution
of the group. Fauna and Goog. of the M;il(h\v and l,;i(c;idi\ e
Archipelago. Vol. I. pp. 209-21S, pi. t. xt tiLnn. - Mf, U).
FOWLKH. (i. H.

Alonogr. XXI, 85 pp., 3 Pis., 6 Charts
Jordan, J). S.

:05. The origin of species through isol;
XX n. pp. .-)4.-)-:)fi2.

Vvhvr (he Darwinsche Theorie in Be/.ug auf die ge(.grai)liis("he
\'erbreitung der Organisnieii. Sitzber. d. bayer. Akad. d. Wiss.
Munchen, Bd. 1, pp. 359-395.

THE ATTACHED YOUNG OF THE CRAYFISH CAM-
BARUS CLARKII AND CAMBARUS DIOGENES

E. A. ANDREWS.

A REMARKABLE fact in the life history of the crayfish is that the
young associiate with the mother for many days after leaving the
egg, being at first firmly fastened to her and later going back to her
for protection until finally quite independent.

As pointed out in The American Naiuralist, :\rarch, 1904,
Cambarus affinis molts twice while fast to the mother and leaves
her only in the third stage. Some facts as to the character of this
incipient family life in an American Astacus from Oregon will be
given in another communication. The object of the present note
is to describe the association of parent and offspring in two more
species of Cambarus and to compare this with what is found in
C. ajfinis and in Astacus. The illustrations are all of ('. darkii.

The young of C. clarkii were obtained from eggs laid in con-
finement by adults shipped from New Orleans, November IS,
1904; some 18 out of 61 surviving the journey. Two of these
active, prawn-like and brilliant red crayfish, one male and one
female lived in a shallow sink of warmish water during the
w^inter and by ]\Iarch 25, 1905, the single female lunl the abdominal
basket full of many very small and vt>ry dark-colored eggs. These
eggs were already in the stage H of ReicluMil)acli but differed from
that in having the abdomen larger. Facli egg was about 1\ mm.
in diameter and partook of the exceptionally vivid coloring of tiie
adult, the large oil-like yolk drops being wine colored instead of
yellow as in Cambarus ajjiiii.s.

When re<-eived in November, tlnvc of the fctnalo examined
had only minute yellow eggs in tlie ovaries and no s,>crni in the
ainnili. while the males had small testes but yet mature .perm in
the va>a defcirntia. It would thus appear that the seas.m of

earh- spring, l.u't thi.-> can be deteriuiued onlv l.v (.bservations in
thefiehl.

254

THE AM EH IC AN NATURALIST [A^ol. XLI

By April 17 the eggs had become coated over with a dark deposit,
but the embrv^o within was far advanced and easily escaped when
pressure caused the egg case to spring open. With Zeiss 2. A. it
was evident that the embryo was clothed in a loose cuticle, or cast
off shell, which loosely invested the tips of the first and second
antennae, the chelae, the walking-legs, the abdomen and thorax as
well as the ends of the gills when torn out of the gill chamber.

These embryos were now essentially the same as when they
hatched three days later. The eyes were almost sessile and with
the pigment restricted to a narrow (lescetit and this pigment
reflected yellow light but appeartMl black by ti-ansmitted light.
The yolk was still a large dark mass of sa(l(II<"-l)ag shape. The
tips of the fourth and fifth legs were strangely bent back like
hooks while the tips of the claws of the chelae did not as yet seem
to be recurved. All over the body the extremely dark crimson
pigment cells again emphasized the agreement of embryo and
parent in intensity of coloration.

But the detail of anatomy of the telson was the most important
character for understanding the subsequent attachment of the
young to the parent. The abdomen ended in a simple, flat,
rounded telson that bore a row of simple spines along its posterior
edge as seen with 2. D. in figure 3. The spines were fourteen or
fifteen on each side symmetrically placed right and left, and a
group of seven or eight of them on each sifle, near the median
plane, seemed to push off the loose cuticle, which on the middle
plane, was close to the body. The spines, or better, papillae,
were highly refractive and clear except that some showed granules
and some vacuoles in their homogeneous contents. Some of them
had small protrusions at the tips as if paste-like material had
extruded from within.

arched <.ln-, mvx and J-cnicd o,-oJn tojjcthcr. On the aninml's
left tlic >pinrs 7 and 1 1 wen- -rown toovthcr at their tips while S

ami th.- >anH- was true of and 12. 'with hiuhrr [.owrr, I nun.
4.45 conip. oc, tnfts of fine threa-ls, or hbrils w<t.- x-en .hvrrpn*:
from the tips of many s])ines to pass, posttMiorly, often beyond
the tips of other spines. Some of the.se threads passed ont to the

No. 484]

ATTACHED YOUNG OF CRAYFISH

255

loosened cuticle and seemed fastened to it. On the ri^ht of the
specimen fine wavy lines sujrgested secreted films rather than fibrils.

As will he seen later these specialized s})ines are ^^landular
stnictnrcs (hat make (he tcl.soii adhere firmlv (o the cast-off cuticle
and th.i.s make pn^iM,- di.' -U'lsun thr,'ad""cf the hatching hirva.

The .mail uuml.er ..f .pine. -n.uped. touerher Nxilh the fact
thai the wUnu of eadier eml.rxonie .(a-e. i> in<-i.ed en the middh-
nf the posterior ed^e Mi^-ct. tliat the.e 7 or S .pine, max he
eomparal)le to the 7 nr S .pine, .ecu on ca-h .ide of the inei.ed
telson of the lobster cinl)r_vo (Fig. 72; Herrick; The American
Lobster) before it molts at the time of hatchini^ and is in a stage
which Herrick compared to a protozoea, or other early larva.
On this basis a very remote ancestral state has been retained to
the extent that its spines have been applied to the new use of
attaching the larva to its cast cuticle.

Before speaking of the hatching larva' it must be recalled that
all crayfish eggs arc fastened to the ])leoj)ods of the mother by a
hardcnino- mas. whose oriuii. is somewhat in dispute. In' C.
clad-ii the pieopod. of the mother were so transhicent that the
transverse striation of the imiseh-s was seen through the exoskeleton
and with 2. 1). the gland cells that arc supj)osed to take part in
fixing the eggs to the pleopods were seen as polygonal areas of
secretion droplets separated by clear lines.

All over the bases of the pleopods these areas were massed
together but the terminal part had them arranged in transverse
bands that crossed the anterior face and extended into the sides
but left the posterior face without glands. ( )n the bands anteriorly

the glands to the surface and near these were .ome short, sharp
setie w^hich occurred a.uaiii at th(- tips of the ph-opods.

While it is possibK- that thes(- .harp scta« aet in priekin- lUc
egg. and liberating an a Ihe.ixe material a. .'laimtvl in Wiiliam.on
for crab's egg., and that tlu^ u'lan.l. of the ph-opod. have nothinu"
1<) <lo uith fa.teidn- the e--.,\et tiii. .eem. ^rr^ improl,al>ie. a.
d.een-> are fa.tentMl to a lai-e'nia.. of material Minilar to the e-g
(•ii.e a-Kl .talk an<l uhieh bin. Is all the l.)!.- plumos<> .eia> together

256

THE AMERICAN NATURALIST [Vol. XLI

When fastened to tlie mother each egg was in a remarkably
elastic case wliich had a rongli, dirty outside layer and a clear
inner layer containing the same microscopic drof)lets seen in C.
affinis and similar to the droplets coming from the pleopod glands.
Each case was continued on one side as a long stalk that in turn
was continuous with the hardened secretion binding together the
plumose sette along the edges of the pleopods. The stalk was
hollow though flat and wide and was a continuation of the dirty
outer layer of the egg case, separating from the inner layer on one
side to form a large hollow^ bell or tent.

Between the egg and the egg case was a variable amount of
coaguhnn showing fibrils in it.

By April 17th some of the larvie had hatclied while otliers were
not yet out of the egg cases. The young, figure 1, had the usual
embryonic look of crayfish at hatching; a huge swelling of the
head region owning to the presence of nuicli yolk there; a weak
development of the locomotor part of the head-thorax so that the
five pairs of weak legs all arose posterior to the iriiddle of the head-
thorax; a weak, down-bent abdomen of little use in locomotion;
eyes almost sessile and of little size or perfection. These larva;
were transparent and showed the heart beating rapidly and the
scaphognathites rapidly baling the water out of the gill chaml)ers.
'Hie dark area in figure 7 represents the dark red yolk mass;
and the scattered dots, the aborescent pigment cells that w^ere
thickly scattered over the head-thorax and abdomen with but few
upon the third maxillij)ed, ba^e of antenna, three basal segments
of the antennule and some few segments of the periopods.

Normally the larva- remained n])oii the mother and did not
move aboiU, and when jHiiled olV and put on the bottom of the
<nsh thev couhl iK.t Mand up but eonhl progress by lying upon
the side and llappino. the alnlonuM..

At halc.hin,-. figure 1, the young w.-re .o weak they would
have dropped to ilw bottom but for the " telsou-lhread " whicii
i. the cast otr ••uticle pulled ,Mit into a thread or band and fastened
at one end to the WUm of the larxa by the ^p.vial uU>u .pines
deMTibcd above and at the other end to the in^idr uf the rn.v.
A. th(- egg ease still tvinaiii> faM by it> >taik to the mother the

No. 484] ATTACHED YOUNG OF CRAYFISH 257

and obtain a hold by them to the egg stalk or to parts of the material
covering the plumes of the pleopods.

In hatching the larva escapes not only from the egg case but
from its loose cuticle and this cuticle, where it covers the abdomen,
is pulled inside out, but leaves the telson spines fast as before to
the inside of the cuticle over the tip of the telson. The cuticle is
so strong that larvjB may be picked up by the telson thread and
their weight does not break it even when hanging in the air.

The attachment of the cast cuticle to the inside of the egg case
seems to be an indirect one; apparently the larval cast cuticle is in
some way fast to the egg membrane and that in turn adherent to
the inner of the two layers that makes the egg case, but this was
not definitely seen. In many eggs the embryo when young lies
upon the side of the egg near the stalk and we suspect some re-
lation between the region of fertilization and of stalk formation.
Later, when the embrj^o hatches, it goes out back foremost through
a crack in the case opposite to the stalk. In the old embryos the
tip of the telson is carried forward to near the eyes and not far
from the stalk of the egg case and in that same region of the egg is
found the connection of egg case to embrv'onic cuticle. Possibly
there may be some common factor, as gravitation, that determines
at fertilization the position of the embr)'o, the place for formation
of the egg stalk and the connection of egg case and larval cuticle.

The ])art plav(>d by the sj)ecial telson spines in hohling the larva
fast to the Iclsoii tliread is shown in figures 2 and I, which show
how the wrinkled telson thread is connected to fihrilhir material,
fastened to and interlocked with the long, curved and arched
spines. In y)assing from the condition shown in figure 3 to that in
figure 4 the cuticle over the abdomen has been pulled off and
turned inside out and is now fn>e from the hirva except where
held by the Tiial.Tial fiiriii>h.Ml by the u-landnlar >},!,„...

Whni the ycunn- -vt h..ld ..f thr nicthrr |.lcn(...ds with^ their

fastened to th.- eg- va.v.

In thiv lii-M lar\al ^Uvj^c ihr.r y..nM- rraxfi^h unv about 4\ mm.
long from tip of telson to a j)oiut between the eves where the

258

THE AMERICAN NATURALIST [Vol. XLT

The accompanying camera sketches from specimens hardened
in Worcester's liquid show the generally imperfect state of the
appendages of the first larva, which lived for a few days an inert
embryo-like existence fastened to the mother and not eating but
only rapidly aerating and circulating its blood as the yolk was
being transformed. The first antenna, figure S, has only four
segments in its exopodite and iti its (Midojxxlite and agrees with
most of the other appendages in being devoid of set;e. This bare-
ness of the appendages of tiie first larval stage was first pointed
out in the English Astacus by Huxley and seems connnon to all
crayfish larvae in their first stage. In place of set* there are but a
few spinules at the tips of the first antenna and on the basal seg-
ment there is a small ear-pit; but as yet the entire appendage
would seem of no use as a sense organ.

The second antenna, figure 9, has only 24 segments in the slender
part of its filament, beyond the three large broad segments, and the
exopodite scale bears a blunt process and a row of few, sharp
spines. The tubercle upon which the nephridial canal opens is,
as in all young crayfish, proportionally very large.

The mandible, figure 10, has a smooth edge with no teeth and
is probably not used. The first maxilla also, figure 11, is very
simple and probably of no use.

The second maxilla, on the other hand, figure 12, bears the
large scaphognathite which is very active in removing water from
the gill chamber. The seta? along the edges of tlie scaphognathite,
though represented in the figure as smooth, were in reality, under
2. I)., set with five side l)ranches so that in this only activelv mov-
ing appendage the setje are present as plumes that would seem to
be of use in striking against the water and m making the ap[)endage
fit more closeK nito tin p,.^,^. h idm- out of tlu gill (hambei

The three maNihipcd-. li-nro i:;. I I. lo. are stranuvlv lacking

No. 484] .4 TTA CHED YO I Xd OF ( 'RA YFISH

259

lobes and setvc aid respiration in niakino; tlie inlet water more
free from dirt.

The chela% figure 10, are long and strong but as yet not sj)ecial-
ized as cutting organs. The tips of the claws are i-ecm-ved as
Huxley first found them to be in the P'nglish Astacus so that once
shut upon a penetrable mass they could scarcely be loosened by
the larva, figure 17. By means of these lockinu" lips tli(> young
become fastened to the egg stalks and lo ilic liardciicd secretion
on the mother's pleopod seta^ so that th(y |)rol»;il)ly remain fixed
in one s{)ot all the time they live in the first stage. Tlie simple,
acicular seta" seen along the edge of the chiw, figure 17, may pos-
sibly aid in tactual reflexes to enable the hu-\ a to shut its claw on
suitable substances.

The next two pairs of legs are very like the cheiic, l)ut more
slender, short and weak.

The fourth leg, figure IS, with no claw, has its arthrobranchs

The fifth leg has no nilU al all aLoeiate,! ^\ith if! tlie pU-uro-
brancii of Astaeus l>ein- absent n..t oi,K in ilie a^luh ("an. hams

deternii.ied by Faxon, in ('. rustirus. The hrancliial fornuihi'is
thus the sanu> in the larva' as in the adults.

On the abdomen the appendages have the incoinpleteness of all
crayfish larva'. The first pair are not begim and the sixth {)air
are forming under the exoskeleton within the base of the lelson.
The other four pairs are vety small and apj)arently (piite useless

abdomen and with the endopodite more aincriov and the exopodite

the larger and both endopodite and exopodite ai-e \eiy simple and

The telson, figure 'J. is a simple, (-lonuaicd, llai plate showing
within its elear'sul.siai,.-e ra liaiin- lines ni,lin- at (he nuirginal

spines ;„m1 aU,, thr oudine^ of tlu' Ion- exopodiles of the sixth

pleopods Ixin- alon- on ea( li ^de <.f tlie ,eetum and anus. On
the veiural side, figure 1 . ihe l.as,- ,.f the telson is <|niie protui)erant
over the part of tlu> enclosed [)h-op()d that will i)e the exopodite.

260

THE AMERICAN NATURALIST [\^ol. XLI

In larvae that have been in the first stage a few days and are
about ready to molt it is obvious that the radiating lines in the
posterior part of the telson are the glands secreting the setse which
will replace the marginal spines at the next molt. In a prepared
section of the posterior part of the telson of such a larva, figure 5,
the old cuticle is separated from the epidermis by a space across
which the tips of the forming setae pass toward the hollow bases
of the old spines. Each old spine has a new seta beneath it but
as there are also other setae the second larvae will have more setae
than the first had spines; the long plumes, however, figure 7, are
slightly fewer in the second larvae than the spines in the first.

Each developing plumose seta seems a flat plate ending in a
fine central thread and with its edges frayed out in short fine
threads. The base of each is deep within the epidermal ingrowth
that forms the secreting gland. Each gland seems a row or rod of
cells, indicated by large nuclei in a common protoplasm in which
no cell walls were seen. The longitudinally striated base of the
plumose seta forms the axis of the rod of cells. The space between
the radiating glands was in part occupied by blood, staining,
like the setae, yellow, while the nuclei were red in borax carmine
and orange G.

Similar, but less developed rods of cells were also seen in sections
of the internal buds of the sixth pleopods where they were forming^
plumose setae that projected into a bag surrounding the pleopod.

By April 24th, when some of the larvae had begun to molt, it
was evident that something was abnormal, as some larvae in both
the first and the second stages fell away from the mother and
died. The mother also died, April 28th. The hatching was pro-
longed more than is probably normal so that many first and sec-
ond stage larvae were found side by side for a few days. Some
of the young in the second stage remained with thv morluT for a
few days but made excursions away from her and apiiii returned

is not certain that this w:is ncniuil in C. vlarkii. While upon the
mother these youno- held firmly with their ehehe, hut they let go
when the dead female was lifted out of the water. When upon
the bottom of a dish they were able to stand up and walk feebly,
and after a day, they swam backwards on their sides by flapping

No. 484]

ATTACHED YOUNG OF CRAYFISH

261

the alxloinen. They tended to climb over one another and one
hrld so fast to a dead fellow that it could be shaken loose only
wilh (liffK iillN . 'rhey also climbed up on to the dead female an<l
on to a piece of ("aiiton flannel where they held fast i)y their chela?
for a time and then ,-oi down and >\vam actively if .iisturbed by a
l.ipette.

Thoiiuli the h-.rva in tlie x'cond >ia.uv may thus away from
the mother it donbtlcss n^tnrns <'ven into the' third sta,uv a^' Faxon
records findin- upon the abdomen of nniseum spe<'ini,MH hirv;e
with characters evidently of the third sta^^-.

The second stage young, figure (>. wtM-e still so tran>hiceiit that
in the abdomen the digestive tract and the ventral gant^dia couhl
be plainly seen.

In r. affltn: it was noticed that the young, in i>a-ing from the
Mvond to tlie thir.l >tage. was sn.pended from it> < a.t cuticle by an
anal thread which bound its anus to that of the ca^t ..ff cuticle
an<l as the < law> ..f that cutich- still hehl fast to th.- mother the

able to take hold again with its uL chiws. ' In ('. riarkli the same

that die.l just after molting. In these there was a long thread
that issue.i from the anus and. pa.>ing .iown through tlie hollow
cast olf cuticle of the ab<lom<-n. was fastene<l at the b..ttom of it to
the flat telson. By the strain of the anal thread the ca.t otf ab-
dominal cuticle had l)een telescoped; tlie old telson being dragij;(>d
up against the collapsing rings of the cast cuticle. A> in ( \ a//7///>
this anal thread was only the cuti<'ular iim'ng of the intestine not
cast off eiuirely at the >ame time with the external cuticle and thus
serviceable in "bin.ling the larva to it> old .lu^ll. If thi> tardine.. in
casting the lining of the iiu<-stine i> normal in ('. c/url:!/' it would

upon the mother, which is prollblv tUr ra^v even if it ha^ .ome

In .•oiUrasting the s^.-ond >tag.', ligure f., with the tir.t, linure 1.
we hud an increasr in >i/,e, tin- bo.ly being now .V. mm. long with

body and in xhv perfection of the limbs. While tile head-diorax

262

THE AMERICA N NA TI RA LIST [\'ol. XLI

still contains much yolk it is less swollen and more elon^^ated while
the alxlomen is relatively larijer and it is more useful as its telson
hears a fringe of setiie.

The limbs are changed but little, yet they now hear some setx
though these are too small to show nnder low nuignifications,
figure (i. 'J'he rostrum is still triangular, hnt sharp, and though it
is still l>entdoN\n hctw(>('n tlu^ (wc. it can he M-ni from a .lorsal
view an<l aho fro.n a .i<lr n ieu , lignrc (i. ulim- it. i)aM- i. ^i.ihle
near the lycs which are now dcci<ledly stalked.

The first antenna had six segments in its exojxxlite and in its
endopodite and the former bore five sense setje, three on the
termiiuU and two on the fifth segment. The ear was a wide open
cavity with three or four finely barbed seta^ along its external

The second antenna now had a long spine and a row of 19 or 20
plumose seta' on its scale and its filament contained .'U segments
some of which seemed to he dividing.

The mandible edge was now no longer smooth hut ha<l six or
seven teeth on its free edge and three above the j>alpus.

^rhe spines at the lips of the chehe, fig.nv L>(). >too<l at about
right angle, and were but slightly recurved, .\long ea-li edge of

narrow blade, {iuMur L'o' which k'mkI.mI to be cracked or .trialed
a<-n... it. lenglh. The.e <-u)ting or ra.pin- .pine, are a .peciali-

Xo. 4S4]

ATTACHED YOUNG OF CRAYFISH

263

In the figure the dotted lines represent tlie enclosed pleopods
and in them the radiatinn; setie ulands in wliicli are fonnino- the

Zeiss 2. I), these^crlands were lono- tubes from each of which pro-
jected a phnne.the tip of whi<-h turne.l to one side in the space
between the edn-e ,>f tlic plcopod and the enveloping- sac. In the
same way the posterior end of the telson showed lono- tubular
glands forniing a set of phnno>e set;c to rej)lace those already
present. The tip of (-i< li new ])hune j)roiected slightly from its
gland into the hollow ba.e of the existing plume, which' would be

All these seta' seen in formation in long tubes are richly barhexl

later at the next molt. ' ^

Only some five or six of these specimens of (\ rlarkii stirvived
to change into a third stage, April 29 to ^lay 1st. but these agreed
with all known crayfish of the third stage in tun ing a complete
tail-fan, with both telson and wid(>ly (>xpauded >ixth i)leopods
together forming a very laig-e area for n>>iMaiire to the water and

Though these few individual. >eenied weak thev'lx.th walked
and swam easily. The color had now become a darker fle.di-color
from the crowding of red pigment c<>lls, but the area about the
stomach was lighter and on each side of the stomach there was
a small, narrow, dark band representing the yolk.

As above stated it is ])robable that in luiture the larva- in the
third stage remain with the mother for a time, and then gradually
become entirely indej)eudent.

Faxon. April N.
females withoiu .
Tallx.t (•o.,Marv

264

THE AMERICAN NATURALIST [Vol. XLI

The young were hatching upon six of these females May 22n(l
and just before this an examination of the embryos showed a
dehcate loose cuticle over each tip of the chela, over the abdomen,
and over the body, and an egg opened in strong sugar solution,
and then put into water showed a cuticle swelling up all over the
antennse, the chelaj and the abdomen. But when carefully dis-
sected it seemed that this cuticle was not a case over each append-
age but rather that it was a large bag over the thorax, a side pouch
over the abdomen, a large side pouch over all the pereiopods and
a side pouch over all the gills. Probably, however, there are two
thin membranes, an outer vitelline membrane of irregular form
when stretched over the protuberant regions of the animal and an
inner, real cast-off cuticle, that goes over each appendage; for
some dissections showed the embryo inside a delicate spherical
bag fastened to the inside of the egg case, and observations upon
the hatching larvae seemed to show them drawing out the limbs
from separate envelopes.

At the end of the telson there were groups of spines fastened to
the cuticle by refractive fibrillar coagulum. On each side a grouj)
of six spines arched over and connected very mueli as in C. clarkii,
figure 3, and here the cuticle was thrust ofi' further, wliilc on the
middle line it was close to the telson.

In one individual the actual hatching lasted forty-five niiinites;
the egg case cracked open opposite to the stalk and the embryo
slowly "oozed" out back forward. During this process some
movements of the legs were seen as well as a rhythmic pulsation
of the lateral lobe of the liver lying close to the yolk mass on each
side of the body, and swaj'ing movements of the yolk mass. This
tube was filled with yellow liquid for ten or twelve seconds and
then grew narrow and white for about two seconds and again filled.
It seemed as if tlie tube weiv contractile itself, but the yellow liquid

In either case the rhythmic filling would seem useful in ai<ling in
digestion of the yolk, uhich was the only available food so far.
Shoiikl it prove that the a<li:lt liver also rhythmically fills and
empties it would be an interesting addition to the anatomical and
physiological evidence advanced by H. Jordan (Pfluger's Archiv,
1904,) to show that the iiver' is the chief organ for absorption
as well as secretion.

No. 484]

ATTACHED YOUNG OF CRAYFISH

265

As soon as out of the egg case tlic larvn lu'^aii to kick its legs
and in a few minutes the scaplioiiualliitc slowlv iiiovi'il, stopped
and began again, finally establishing a ra])i(l rliytlini. On adding
carmine, the currents made by the scaphognathite were visible
and its movement seemed comparable to a scooping motion of a
hollowed hand, the fingers downward, thus forcing the water
through the dorsal part of the respiratory passage as the fingers
closed the lower part and tlu-ii rising up to close the upper part
and prevent a back set of water into the passage way.

Once out of the ego- cax- \hv hirva was still fastened to it by a
telson thread coii-^i-tiiiL;- of a -hovt ^wId'j: from tlie tel^oii spines to a
large cruniplc<l ma- that MTincd a cast otV cntich' and lay just
within the gaping egg case an<l \va> fasrcnc.l to it. in^'.h'. hy the
intervention of an expanded membrane which may possibly have
been the old vitelline membrane. This mend)rane wa^ t)omid to
the inside of the egg case by a few short fit)rils over a round area
smaller than the base of the egg stalk and often near it. Thus
suspended the larva moved its legs weakly and now and then shut
its claws and violendy flapped its abdomen without breaking
loose from the telson thread. Soon the larvje became fast by their
claws to the egg stalks or to the material on the plumose seta? of
the mother's pleopods.

In this first stage the larva' remained cowered down close to the
pleopods and were so firmly fastened to the mother In' their claws
that they did not break loose when the pleopod was thrown into
Worcester's li(jtiid, though they jerked their legs and ])owerfnlly
and violently Happed their abdomens. Those left locked to the
pleopods of the mother lived three to four days and then molted
into a secon.l stage, May 2f>.

They were very large. :, to (i mm. long when strct.'he.l ont and
4i mm. as they lay with the weak ahdomen carrie.l forward imder
the thorax an<l w<M-e very attractive ol)ject- hecan-e of the swaying
of the dark red and gol.len yolk nia>s, the contraction^ and change
in color of the lohes of the liver N|>read like the (iiiM-ei'- of a hand
deep in over the back, and of the liery. ruby-red. neuroiidike.
branching pigment cells >pangled over a hody so translucent as to
show the white blood corpuscles hurried along the vessels over the
red yolk, along the sinus at the edge of the (•ara})aee and out and
in through the legs an<l antemia> like shtitdes.

266

THE AMERICAN NATURALIST [Vol. XLI

Camera lucida sketches of the first larva of C. diogenes showed
it larger than even the second stage of C. clarkii but in simplicity
and proportions essentially like the first stage. As usual in hatch-
ing crayfish the appendages were almost all bare of setse; the eyes
were nearly sessile; the rostrum a small triangle close to the body
and between the eyes. The yolk far forward in the head-thorax
distended that region and left the region for the gills and pereiopods
of less extent.

In the first antennae there were four segments in the larger, club-
like exopodite and also in the slender, smaller endopodite and
there were no sensory setse.

The second antennse were carried curved backward and down-
ward but not close against the thorax as in C. affinis and each had
only spines upon its scale and 35 segments on the slender part of
the filament.

The mandible had no teeth but its edge was very slightly waved
where the epidermal cells seemed about to secrete slight thicken-
ings.

The scaphognathite used as a baling organ also was exceptional
amongst the appendages in bearing plumose setae which formed
a row along the edge and were longer and more easily seen than in
C. clarkii.

The gills were larger and with more side filaments than in C.
clarkii but were suddenly reduced upon the fourth pereiopod so
that the anterior arthrobranch had but a few filaments and the
posterior none. On the last thoracic somite there were no gills,
as is the case in all the young of Cambarus thus far studied.

The four pairs of pleopods had the endopodites slightly longer
than the exopodites and the entire appendage was very much
longer than in C. clarkii and with evident spines on both tips.

The telson with its enclosed pleopods was very much like that
of C. clarkii and bore on its posterior edge the same kind of spines,
about 14 on each side, six of which wore specialised gland ducts
archc.l over and ioincd u^ivtUvv and l.<.un.] to the tfl^..n strino-

stage and a day befon
extended from their gla

Xo. 484] ATTACHED YOrX(; OF CRAYFISH

Each new seta luitl its lateral l.arh. cIom.i' a|.|)iv.M.,
axis. Moreover the new cnticlc .•Men.lc.l" inwani to

tul.e. eii.lin-al.rnptly, Mn.n-lv nvall'.l thr likr iiil.r
alonu- the seta' of the eardi worm, rrol.al.l.v ai n
sleeves heeoine everted and m. all-.u ihr .ndd'm rMr
setae to a length e(jiial to the length of tho^c tuho ;■
length of setpe already lyinu' between the old aii<l new <

Molted into a second stau'*- tlie yoimu' i1 loiji'ins \
long, H wide and 2 dec|) and had anteiuia' ."> nun. I(»ii<:

next molt — some hve days. May 2(lth to th

upon the mother's pleopods, hut were not so hrinly fix
as thev fell off when put into Worcester's liquid.

While in most respects the larva was essentially like
larva of C. clarkii a number of differences were noted.

The rostrum was le.ss l)ent down than in C. darkii a
and pointed and visible from the side as its tip exteii<l<'(
the eyes; its sides moreover were not straight as in ('
arched so that something of the adult character
expressed.

The first anti una \va~- xcx concave on the upper sid-

externaTly by u few small spines an. I one very iin|)erf
seta. Beneath these spines the piuinoM' MMa' .)t' the
were seen in formation. The exojHi.lite hoi'e
three on the sixth and two on th<' fifth an.l on the foii
The exopodite. an.l en.lop<..lit(- uere ea.'h .lixi.ie.l

268

THE AMERICAN NATURALIST [Vol. XLI

A dissection of one of these second larvae revealed a mass of
membranous material and both simple and plumose setse in the
intestine suggesting that these larvae may eat the egg cases and
setse from the mother's pleopods.

The creature was still translucent enough to show the ventral
ganglia through the exoskeleton of the abdomen and was dotted
over with pigment cells of stellate form, which when expanded were
light red and when contracted very dark, while deeper in were
diffuse and indefinite blue cells. In the antennae and legs as well
as in the antennal artery the corpuscles were going outward rapidly
and returning somewhat more slowly in wider vessels.

Two days before molting into the third stage the new inner
cuticle was already formed and the new setse projected into the
bases of the old. The yolk had become reduced to a small dark
remnant on either side and even to the naked eye the gastroliths
were conspicuous as two pink-white opaque areas, one on each
side of the stomach enveloped in a clear glassy coat.

The third stage began by June 1st and had the adult character
of a tail-fan made of the telson and the fully expanded sixth pleo-
pods all fringed with perfect plumose setae. These larvae were
10^ to 11 mm. long, 3 wide and 2h deep and expanded the tail-fan
about 4 mm. while the antennae were 6 mm. long.

These third stage larvae when recently molted were still some-
what translucent and of a faint pink color with red-tipped claws
and though the stomach was plainly visible the gastroliths were
lacking on the outside. But within the actively moving stomach
was a brownish liquid containing white particles or in some cases
whole gastroliths moved about actively. In some cases the intes-
tine contained colored material in its anterior part.

The specific gravity of the larvte had so changed tliat they now
floated in Worcester's liquid though the first and sf'cond stages
sank; they were also less resistant to this fluid and died more
quickly than when vounger.

When the iarvH- luid chanovd int.. th<- third Mauv it wa. noted
that the six mothers no hniger had egg cases an.! ea.t .Miti. le> upon
their pleopods and as their fa>ees containe.l part.> of phnn.^e set;e
of adult size it may be that they aided in cleaninu' ..tf their pleojwds
though there is some evidence that the second hirva- may eat off

No. 484] ATTACHED YOUNG OF CRAYFISH 269

that material and Souheiran stated that the yoiin^ of an Astacus
ate the egg cases and larval skins.

The third stage larvie stayed near the mother some ten days or
more, often, when disturbed, climhiiiu- on oih- aiioili(M- and crowil-
ing under and upon the motlici'. hut aftci' iliai iluy wciv (|iiitc
independent and seemed to hiwr no associntioii wiili die moilicr
though kept in the same small a<|iiariiim.

Walking and swimmino- the yoimu' sought food o\(t the KoUom
of the aquarium and in a day cleaned otV all the l.rown d.>|.o>it
from a spray of Myriophyllmn and when another piece was uiven
them ravenously set ahout teai'in,u' oil' and eatino- the hactei'ial
slime and algal growths. When given Chai-a they >ei;'.ed nii
internode with their month j)ai-ts and ])tished it with iheir feet
somewhat as a dog gnaws a bone, but when j)ieees of internode
were cut off for them they seized them by one end and walked
about suc;king the contents out. Such a larva holding its head
high and supporting a stick longer than its body, held by its
mouth parts straight out in front of it, ludicrously suggested the
enjoyment of a huge stick of candy. Animal food in the shape of a
dead comrade was eagerly sei:;ed and {Milled to ])ieees and a small
earthworm was eaten tip m a few lioiirs.

Living thus, at a temperature of .v") the yonnu' were very

water. After two weeks some molted without nmeh change
<.f si/e but by duly 3rd some were 13 to b") mm. long and the only
survivor. July l.")th, 18 or 19 mm. long.

From the above account it appears that the voting of (Uimharns
clarkii and (Uimbarus diogcnc.s associate with the jtarent in the
first and second stages and in part of the third and this sort of
family life is aided both by special recurved tip^ on tlu' chehe and
by a peculiar teUcn threa i; an. I a-^ thi^ i^ irue aU.. in C. nvmis a.

gen.'ral fact Inr all ^pr, ir^ the^c tu.. umna Mnn .Al.r all ilieM>
ci-ayhsh show in the voun- .tniclural .•liara.-irr. and !.al.il> that
make them unfit for "free life like that (.f their niarine relative..

\vith the mother with whom they live as in a kind of elementary
family.

270

THE AMERICAN NATURALIST [Vol. XLI

In this departure from ancestral conditions C. affinis has gone
farther than Astaciis in the following respects. In the first stage
and in the second stage the telson is more reduced and both pairs
of antennae are more simple and to some extent this is also true in
the other species of Cambarus here described. Thus in tlie first
stage, Astacus has 50 to 66 spines along both the posterior and
lateral edges of the telson while the three species of Cambarus
have spines only upon the posterior edge and they are less than 30.
Astacus also has in the first stage five segments in endopodite and
exopodite of the first antenna and .")() in the filament of the second
antemia while the three species of ( anilfarn. have but four in the
first case and 2.) to '.^o in the last.

In the first larval stage the three species of Cambarus thus
agree amongst themselves and depart from Astacus in the direction
of simplicity which is presumed to be a secondary reduction in
connection with protected life upon the mother.

In the second larval stage C. affims alone has spines merely
and no plumose seta? upon its telson and is thus most remote from
fitness for the active life of its ancestors. In the second stage
Astacus is most like a free form in having its telson fringed with
much more perfect and ninneron. j)luin()se seta; than are found in
C. clarkii or (\ <li,>;/nirs. In Asta-us also the first antenna has its
ear-pit well overarched by a row of plninose seta* but in C. darkii
there are only 3 or 4 plumes, in ('. (Hixji'tics bnt one i)lunie and in

second antenna has .")4 s(«gnient>, in ('. (liot/cnrs about 40, in C.
clarkii 34 and in ('. alJiii/.s :v.). in ('. (ijjinis alone is there a reten-
tion of simple >|)ine> Mich a> occur in the first larval stage, so
that the scale of the second antiMnia here still bears no plumes.

Thus in the second .ta.u'e C. is most removed from Astacus

but C. diogrNrs and f. r/arkli .lepart h's> from the anceMral Asta-

No. 484]

ATTACHED YOUNG OF CRAYFISH

271

From consideration of the larval lif'(> \v(> come to the same
general conclusion as that generally drawn from study of adult
anatomy and geographical distribution, namely that ("amharus
is a more highly evolved form than Astaeus and that (\ ajjiiiis is
one of the higher, more specialized forms of the genus.

As to the relative position of (\ clarki't and ('. <//'o(/rnr.s there is,
however, doubt and (liserej)en(y-. The aihilt characters seem to
leave no doubt that C. clarkii is nmcli tlie more primitive, h-ss
specialized and more like Astaeus of the two. But in the adjust-
ment of the larva to family life (\ dUxirnrs would seem to have
progressed less far tlian (\ clarkii, at least in the first stage C.
diogenes has more segments in its second antenna and in the second
stage more. sense seta- in the first antenna as well as more segments
in the second antenna. ( )n the other hand (\ clarkii would be
more primitive in liaviiig more seta- over the ear-])it and if in
nature the yotmg actually get loose from the mother in the second
stage they would be more like A>tacu-^.

be paid to the first three larval stages as ai<ls in determining the
relative positions of the species and their j)robable derivation
from ancestral forms.

From a thorough study of larvai of many species and from experi-
ments in cross breeding some idea might be got as to tlie nature
of the causes that seem to be leading some of the more evolved
crayfishes to develop further that association of parent and off-
spring which forms in the crayfish a simple stage in family life.
December 20th, 1906

NOTES AND LITEliATURE

JUOLOGY

Beebe's Log of the Sun.' Merely to turn over the ])afres of this
beautiful book dispels all desire for (•aj)tious criticisin. 'I'he pub-
lishers have done everythintr to ])resent ir to the public in the best
shape while the fifty-two full ])a(ie plates in color by Walter Kin^^ Stone
and the numerous ti-xi li<inrcs from photographs and from wash and
charcoal drawino-. iii;ike ihc work a delight unto the eye. Each of
the fifty-two weeks ot the year has its chapter; in some eases chosen
with a full a])preciation of tlu> fitness of things, in others placed in
position because one week would do as well as another. Some of
these chapters have previously ai)pearcd, without illustration, in
other places, while others were prei)ared exj)ressly for this volume.
Naturally the birds attract the most attention, with the mammals a
close second, but other chapters deal witli reptiles. II^Ik .aiid insects,
while the invertebrates of the sea are not iieulcctrd and even those
marvels of crystallograi)liy, the snow flakes, have their all..tted space.

The text itself is writt.Mi in an ea>y. u-raeeful mainief with a full
appreciation of the wonder^ of tiaimv and with the most s\ inj)athetic
spirit. Here and there, perhaps, a statement i- exairii-eiated or, may

they shall not be detailed here. \{vm\ the luMpk. look upon the living

world about you — sea, shore - plain or tureM with the oj)en eyes

of the author and you will .see the mai vels lie lia> >een and a myriad
others of which he tells you nothing.

Laloy's Parasitism and Mutualism. -- 1 )r. Laloy devotes an intro-
ductory chapter of his recent work- to a eonsid(M-ation of th(> various
reciprocal relations between living things of which seri<\s para.sitisni
and mutuali.sm are the opposite extremes. I'ollowing this th(> first
part deals with para.sitism under S(>ven ehapUM- headings: geni'ralities,

'The Log of the Sun; a chronicle of Nature s Year. By C. AVilliuni
Beebe. New York, Henrj- Holt & Co., 1906, pp. xii + .345, $6.00.

th<;caire de I'Acad^mie de M^decine. Preface de M. A. Giard, professeur j\
la Sorbonne. 1 vol. Bibliotheque Scientifique iiiternationale; 82 text fig-
ures. 6 fr. F^lix Alcan, ^diteur, Paris 1906.

275

276

THE AMERICAN NATURALIST

[Vol. XLI

plant parasitism, plants parasitic on animals, animals parasitic on
plants, animal parasitism, the role of parasites in pathology, and
finally parasitism in the evolution of species. This last chapter
presents in striking fashion an opinion previously advanced by this
author regarding embryonic and sexual parasitism.

In the second part, devoted to mutualism, are grouped under separate
chapters discussions of social life among plants, mutualism between
plants and animals, social life among animals, and mimicry. Under
these headings are discussed many interesting questions of an unusual
sort. The author has selected instances of an illustrative tyj)e and
presents them clearly and attractively.

The scope of the work is uncommonly large, einhrac iuo- ;,s ii does
both plants and animals and scant 300 pages are narrow limits in
which to present such discussions in a form to escape criticism. To
a zoologist it appears as if on the whole too great space had been given
to the plant side and yet this may be distorted perspective on the part
of the reviewer. The figures deal almost exclusively with plants and
insects, with the former largely in the majority.

In many respects the work hardly represents present knowledge
on the subjects discussed. Thus, in speaking of the hookworm, to
which the author devotes a considerable' section, the statements tiiat
this parasite sucks blood is jx'rliaps excusable, thouirli in WHY.', Loo^^

upon and quoted, as well a> <(Mifinncd, since then. However to
outline the life history with the larva cnevMed in a roisiant envelope
and infection takinn- place by the inoulli astoiu.hiiin' i„ view of th--

No. 484] NOTES AND JJTERATURE

277

278

TIIK AMERICAX XATURALIST [Vol. XLI

indiciitioiis of tlic coiitiiioiital orio-in of a o-ivon formation. Applying
the results of his studies to specilic portions of the geologieal column,
the author concludes that certain important formations, heretofore gen-
erally referred to a marine origin, are most probabh' continental
depasits.

Professor Barrell's j)a])er is an important contrihution to a series
of studies which urr rcsultiiiii- in a vcrv nianif<-st niovcniciit away

' 1). \V. J.

Observations in South Africa. — Professor W. .M. Davi-, |)rcscnts^
l)is visit to the Colonies of South Africa in ihc snniincr of MlOo.

Hric.l. and th.- pn.l'.l.'nis lo l,c r.l.idnvd l.ricflv .lalcd. The next
tucntx paov. arcrnainlv c.mrvnM.l uith a .tndv..f ihc (ape Colonv
ran-.- .•nnH<lrivd uith ^pr.ial n-ard „. ,|,nr n'MMuLlance to the

of its origin, the evidence being weighed with a desire to discriminate

leveling without baselevcling as the other. ( )thcr problems of interest,
MK-h a- the ori-in of l\u- ziu"-/.-!- -or-v l.rlnu the \-i< toria falls of the

I). W. J.

Geology of the Big Horn Mountains.- The results of five seasons'
field work in the Pig Horn Mountains of Wyoming and Montana

17, i.i>. :!77-4.W, lOOG.
nil Taper No. 51, 1906, 12<S pp.

Xo. 484]

NOTES AND LITERATURE

279

;.re einl)0(li(>(l ii. tliis .splcn.li.lly illiisi nin-.l ivport l.y ^Fr. N. II. Darton.

end, while iiiinnr tlrxuiv. wnd f.-iiill. ...■. nr. 'I'lic .avn.Tnl uvolo.i^-i.-
history of the rctrion i. tnicc.l. an.l in .-cik lu.i.,ii xhv miiicnil ivxMirc.-s.
wate/supply and tinihev are doscrihcd. Sonif y.-ais a-n Mr. F. E.
Matthes prepared an unusually valuahlc coniour map nf ilic central
portion of the range and discussed the fcatiiiv due lo ohn sculpture
Mr. Darton's report gives a comprehensive aeeount of the general
geology, adding much to our knowledge of this inierestin<r region.

and presents in this paper' an aeconnt of a lake which .seems to owe
its origin to glacial erosion, and uhi. l, elosely re-Mni^les th.- famous
valley lakes of Switzerland. A-ronling ,o p.vxiou. ol-MTsers ihe
lake has been fonued bv the danuiiiiiu- of an old outlet l>v fans spread
out acro.ss the valley "by tributary stream- but Mr. llm.tington
presents pretty clear evidence that the basin is terminated by a rock
lip rising well above the present level of the lake, and of course nuieh
farther above the lake bottom, th(^ lak(^ being 1 12 fe(^t deej) aeeordiiig

ings and photographs.

280

THE AMERICAN NATURALIST \Yoj.. XLI

ZOOLOGY

A Statue of Lamarck — As yet there is no memorial to this emi-
nent naturaHst but now it is proposed to erect one in tlie Jardin des
Plantes in Paris. The matter is in charge of a committee of the
Museum d'llistorie Naturelle in Paris; subscriptions may be sent
to Professor Joubin, the secretary, i)o, Rue de Buffon, Paris, France.

Gardiner's Maldive and Laccadive Archipelagoes/ parts of which
have been noticed in these pages as they have appeared, has now
been comj)leted. The whole makes two quarto volumes of 1079
l)ages and 100 plates. In this concluding part is an account of the
Myriapoda collected by R. I. Pocock enumerating eight species, and
some supplementary remarks upon geogray)hieal distribution and

Kollmann's Atlas of Human Embryology.' 'I liis is, as its name
iTnpli<'s, an atlas of developnient . TIkm-c is no true text, merely
descriptions of the three liinidred and forty fiunres which are intended
to illustrate tli(> features of hunian eiubrvolou-y. 'I'hcse figures,

from 'u!i.h'.lrauing> are uith feu eveptinn^ e\eellenl, uhil.'lhn.e

made by the /inc process are usuall^ more eni.le. h. a feu . a.e.
other animals than man have been called upon to supply the illus-
trations. Thus the (>arly phases of the niainnialiaii onuns are based

Bonnet's work upot. the <loi: and S<-lenka^ up«>" =M><- <-alled
in to illustrate other .-arly features; while < hiek and (ish furnish illus-

' The Faun;,
edited by J. ^t i
1906. 3s, 6(1.

^ Handatlas

No. 484] NOTES AND LITERATURE

281

trations of monstrosities, and the development of the skull is intro-
duced by Schauinsland's figures of Callorhynchus, and Stohr's of the
salmon.

While the work has consi(l(M-al)lc \n\uv for the inctlical >tu(l('iit
in that the illustrations suj)])l("iiitMit thosc^ of ilic usual text l)o<)k,
the morphologist finds the volume less a(la|)t( <l io lii> needs. ( )ne
might wish figures showing the early slao-es of the vertebral eolmmi,
more details regarding the development of the lower jaw. better
illustrations of the embryonic adnexa, and some eonneetion l)etw<'en
the figures of the head cavities and the definitiv eve miix les which
develop from them. Three figures illustrate the development of
the diaphragm. In two only the sej)ttnn transver>nni is >liown;
in the third the diaphragm has nearly its delinitive eondliion, but
there is nothing to show the origin of the 'pleural portion.'

The Systematic Position of the Tubinares. In a recent nunituT
of this journal (41, p. Ill, 1907), Dr. Sluifeldt in the hi.storieal intro-
duction to his paper 'On the osteology of the Tubinares,' has this to
say about my treatment of these birds in the Standard Natural His-
tory, vol. IV, pp. 84, seq., (Boston, 1885) : "This writer places in his
scheme the Tubinares widely removed from the Steganopodes, which
I believe to be a mistake, and a non-appreciation of the morphological
characters of the latter o-roup of Birds."

If the main object ..f the l)ird vohnn.' of the Stan.lard Xatnral

have been a ditVerent one. That 1 fully indicat.'d th<-ir i>roper place
and also fully api)r.-<'ialed tln-ir ' morph.)logical characters' will be

"The arrangeinerit may n..t be ivgar.le.r a! final, however, for there
are reasons to suspect that it will In- necessary. <mv long, to divide the
schizognathous swimmers into three or<lers. Kretmopo.les for the

282

THE AMERWAN NATURALIST

[Vol. XLI

And finally, on page 85, I again emphasized the true position of
these birds by reiterating that it is "rather probable that the Tubi-
nares should be placed in the neighborhood of the Steganopodes and
Herodii."

Professor F irl nng r i I last review of this subject (Jcnn.
Zeitschr Xaturvv., 36, pp. 644-646, 1902), does full justice to the
subject as follows: "Stejneger-Cope ('85/'89) follow Iluxlev in
the rather unfortunate establishment of the Cecomorphjie, but Stej-
neger mentions particularly that the Tubinares perhaps are lietter
regarded as a special ordo with nearer relation to the Steganopodes
and Herodii. . . .On the strength of later coiisidcratioti^ 1 >till adhere
essentially to the opinion expressed l)y ine in IsSS, hnt I am iiiehned
to place their relationship to the < 'ieoniiroruio nioiv in the foreground
and that to the Laro-Liinieol;e nior ■ in th ■ backgronnd than then ....

On the other hand, I eaiinot follow th<.-.c author. wIk, argue for
placing them too far from Laro-Limicohc," the group called

Beebe's The Bird^ in tlie American Nature Series is easily one

thisepocliof bin! books has pro.hieed. it marks, v.e hope, the l.egin-

thos.- who n..u' ke..p bin! lists as a pastinie will take up the serious

titles of >ome of which are as follows,-- The I'ramework of^lie Bird,
The Skull, 'i'he Foo.l of Birds, 'Vhr Sens. s. B<-aks an<l Bills, The
Eggs of Birds, etc.

Mr. Beebe is curator of birds in the Xew York Zoological Park.
His position lias enablc.l him to observe at clo.se range tiie hal>its of a
great variety of binls, and also to (h.scover the needs of an in.piiring
public. .Mr. Be(>be is, however, nnich more than a keeper of animals;
he is a trained scientist and a skilful lecturer. He has succe(>dcd in
this book in arranging a large amomit of accurate information clearly

the n>adcr's infrcst.

' Beebe, C. William. The Kud. It I nn uidlucTDn. Xew York,
Henry Holt & Co. x + 496 pp. 371 figures.

No. 484]

NOTES AM) LITERATURE

283

in the author's mind. The lirsi cliaptrr, tlicivtoir, pivscius ilic
essential fa(•t^ ^^hi(■ll pahi<(.niol..K> . ..in nl>nt.>. to out know UmI^ti^ of
the bird .uhI t liroiiiihoiit the book there are fre(|uent ami illuniinatinjr
references to lioiiiolooics or analogies in the kindred classes. The
delieate balance ol Nature and the complex interrelations of all
organic lih; are well illustrated in the chaj)ter on food.

Where a large number of forms are (Hscussed it is dilficult to avoid
the appearance of a mere catalogue of compiled facis. Kvaiis Dic-
tionary of Birds is a noticeable example of work of tins kind. Mr.
Beebe has avoided this danger 1)V a liap])y introduction troin time to
time of bits of personal ob.servation, or by enlarging on some excep-
tionally interesting habit or structure. The reference to a flamingo
observed by Mr. Beebe, weeping from terror because a condor was
playfully "galloping" around it, illustrates also the author's happy
choice of words.

The suggestion of problems to the solution of which careful obsen-
ers can bring assistance, the frequent references to Nature's evasions
of our pet theories, and the conservative position taken on dis])iited
points, begets in the reader a stn.ncr ami deserved tVeliiii: ot confi-
dence that Mr. Beebe possesses together with Ins power ot pici nres(|ii.'

scientist. Mr. Beebe is evidently a strong believer m M-\iial M'ledion.
but he puts forth (p. 318) an interesting suggestion that the dispiav
of the male bird instead of affecting the jesthetic sense of the female

tion.s on" color chanlvs due directlv to .-nvironment or food. White-
throated .,,arro^^. and uood thru^he. turne.l ahn-.^r Mack when

Thouii-li primarilv intemle<l for tin- instruction of amateurs, Mr.
Beebe'. ixu.k i. one that nmII at om e ^^\^^ an honorable place in the

school shotiM b.' without it. It will b.^ tin'' hope of all who use this
manual, that Mr. Beel)e will follow it by a similar treatment of the
intelligence of birds.

R. H.

284

THE AMERICAN NATURALIST [\^ol. XLI

The Conus Arteriosus in Teleosts. — One of the characters which
have been relied upon to distinguish Ganoids from Teleosts has been
the presence in all (xanoids and, with the exception of Butirinus,
its absence from all of the other group. H. 1). Senior now shoAvs^
that the tarpon of our southern waters has a conus with two rows

Does half of an Ascidian Egg give rise to a whole Larva? — In

reply to criticisms of Dricscli, Conklin n-tuni. t.. tlii. (jucstion which
was' discussed in his earlier i)apers and maintains (Archiv f. Entwick-
lin.gsnKvliai.ik, 21, 19()(.)) the general correctness of his former account.
The hair l)lastomere cleaves as if it were still part of the entire egg;
(•(u-rcspoiidingly the resulting gastruUe are half gastruhe and are in
no wise l)ilaterally symmetrical and the anlagen of muscles and
mesenchyman unilateral in position. Similarly the larvfe uj) to the
time of metamorphosis are half larvie, having only the parts belonging
to one side — right or left — represented. They "are such as would
result if a fully formed larva were cut in the median plane and the
cut edges of each half then came together, the dorsal and ventral
mid-lines joining. These results follow from the early differentia-
Digestive processes in Collembola. Dr. .1. W. 1m)1s()iii and Miss

TonHH-rn,. .nd m h.,' '( ■o'iKmmI. 'la TlL <|rnv the rxis,.,uv of
Malpio-hian tnlxil.- in thcr inM-ci. :,nd .u.lr th,- lad that thry molt

tion of the intier half ..f the intestinal epithelium, the (legcnerate
portion being ca^t out soon after the molt, .•arrvino- with it a part of
the nuclei wiiieli aiv n-plaee,l mit.»tieally from those- which persist.
In this degeneratinu- mas> is containeil xxlic urate as well as gre-
garine so that this is an exen-torv process. Adult specimens molt
everA- six or <-ii:ht day>: the cast skin i. dev.Mired.

Fresh Water Amphipods of North America. Miss Ada Weekel

No. 484]

NOTES AND LITERATURE

285

286

THE AMERICAN NATURALIST [Vol. XLI

doptera found in New England. The second portion (over half)
is given to a key to the caterpillars of all but the smaller Lepidoptera
of the same region.

J. F. McClendon has described four new species of ^lyzostoma
(Proc. U. S. Nat. Mus., 32, 1907, obtained from the collection of
crinoids in the National ^Museum.

BOTANY

Sukkulente Euphorbien.^ —This is the first of a series of illustrated
handbooks of succulent plants designed by the author to meet the
demand that the scattered literature should be brought together in a
form accessible to the many cultivators of this group of plants. The
aim is to give both a scientific classification and cultural hints on
those species now in cultivation and this work has been admirably
done, both in the text and in the numerous half-tone illustrations.

One hundred and nine species and eight varieties of Euphorbias
are treated, nine species and two varieties being new to science and
three species have been renamed. The species listed are for the most
part natives of Africa, though a few are from the adjacent islands and
from India, and three are American. A good working key is given
to the twelve sections under wliich the ■>|>;-( ies are arranged and the
sections again are each provided with good comprehensive keys so
that a species may be readily determined. The descriptions are full
and clear, supplemented by full synonymy and by additional notes on
habitats and comparisons between species, ending in a short note on
the culture required for the species. Following the treatment of spe-
cies a chapter is devoted to the general culture required for this group
of succulents and the text ends in a full index To thv literature bearing
on the group.

In view of the fact that several new species apjiear in the piibliea-
tion it may be well to state that, though the title page bears the .late
of 1907, copies have been distributed in December I'.MIii.

1 Berger, Alwin. Sukkulente Euphorbien. Stuttgart, 1907. 12 mo. v -f
135 pp. 33 Abb.

{No. 4S3 was issued March 29, 1907)

The first of a series of Colored
Plates of the ^ ^ ^ ^ ^

THRUSHES

OF

NORTH AMERICA

By FUERTES & HORSFALL,

was published in BIRD^LORE for
February. ^ ^ ^

The series will be concluded this year

20 CENTS A COPY. $1.00 A YEAR

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vol. XLI, NO. 485

MAY, 1907

THE

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

I. Tli« Functions of the Spiracle of the Skate . . HEBBEST W. SAND 287

II. A Critical and Statistical Study of the Determination of Sex, particularly in

Human Offspring F. H. PIKE 303

III. Chubs' Nests ALFRED W. G. WILSON 323

17. Notes and Literature : General Biology; Mendelism— The problem of age,

growth and death.— The hypothesis of mimicry. Zoology; Palms and
Soles. — Literature of Ichthyology. — Nettling hairs of the brown-tail
moth.— Divided eyes of insects.— Notes. Botany; Winter rest.—

Notes 329

V. Correspondence: The Flying Fish Problem 347

BOSTON, U. S. A.
GINN & COMPANY, PUBLISHERS
29 BEACON STREET

Chicago London, W. 0.

378-888 Wabash Arenue 9 it. Martin's Btrci

The American Naturalist

EDITOR

FREDERIC T. LEWIS, M. D., Harvard Medical School, Boston, Mass.
ASSOCIATE EDITORS
J. A. ALLEN, Ph.D., American Museum of Natural History, New York
E. A. ANDREWS, Ph.D., Johns Hophins UniversUy, BaUimore
WILLIAM S. BAYLEY, Ph.D., Colby University, WatervUle
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell UniversUy, Ithaca
WILLIAM M. DAVIS, M.E., Harvard UniversUy, Cambridge
ALES HRDLICKA. M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford UniversUy
CHARLES A. KOFOID, Ph.D., UniversUy of California, Berkeley
J. G. NEEDHAM, Ph.D., Cornell UniversUy, Ithaca
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, PUlsburg
D. p. PENHALLOW, D.Sc, F.R.S.C, McGUl UniversUy, Montreal
H. M. RICHARDS, S.D., Columbia UniversUy, New York
W. E. RITTER, Ph.D., University of California, Berkeley
ERWIN F. smith, S.D., U. S. Department of Agriculture, Washington.
LEONHARD STEJNEGER, LL.D., SmUhsonian InstUution, Washington
W. TRELEASE, S.D., Missoun Botanical Garden, St. Louis
HENRY B. WARD, Ph.D., UniversUy of Nebraska, Lincoln
WILLIAM M. WEffiELER, Ph.D., American Museum of Natural History,
Neva York

The American Naturalist is an illustrated monthly magazine
of Natural History, and will aim to present to its readers the leading
facts and discoveries in General Biology, Anthropology, Zoology,
Botany, Paleontology, and the several branches of Geology.

The contents each month will consist of leading original articles
containing accounts and discussions of new discoveries, reports of
scientific expeditions, biographical notices of distinguished naturalists,
or critical summaries of progress in some line; in addition to these
there will be briefer articles on various points of interest, editorial
comments on scientific questions of the day, and critical reviews
of recent literature.

All naturalwts who have anything interesting to say are requested
to send their contributions; candidates for the higher scientific degrees
are invited to contribute concise summaries of the special literature
pertaining to their chosen topics. The editors will endeavor to select
for publication only that which is of true scientific value and at the
same tune so written as to be intelligible, instructive, and interesting
to the general scientific reader.

All manuscripts, books for review, exchanges, etc., should be
sent to Frederic T. Lewis, M. D., Harvard Medical School, Boston,
Mass.

All business communications should be sent to Messrs. (iixx
& Co-MPAXY, 29 Beacon St., Bcston, Mass.

Annual lubicriptlon, $4.00. net, in advance. Sinele copies. S5 centi

GINN & COMPANY, Publishers

THE

AMERICAN NATURALIST

Vol. XLI Mai/, 1907 No. 485

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF
THE MUSEUM OF COMPARATIVE ZOOLOGY AT HAR\-ARD
COLLEGE. E. L MARK, Director. No. 189.

THE FUNCTIONS OF THE SPIRACLE OF THE SKATE

HERBERT W. RAND

I\ tlie latter part of September, 1004, I spent a few days at the
^^oo(^s Hole laboratory of the United Mat(\s Hureau of Fisheries
for the purpose of making a stiidv oi cci iaui blood vessels of the
skate. At that late season the Fisliencs laboraioi-v had abandoned
Its hsh traps and I was supplied with material tiirough kindness
ot officials of the IMarine Biologieal Laboratory, wliich was still
maintaining a trap in A ineyard Sound. One afternoon this trap
was hauled and some seven or eight common skates {Raja cnnacca)
were taken. The skates were thrown with nnnieroiis other fish
into the bottom of a skiff which was towrd back to the laboratory
by the steam launch, — a distance ot about a niilc. Arrived at the
laboratory, I picked out the skates and threw theni into a lar<>:e
shallow tank for the pmiH.M ot ua^hni. t.oni th( n. th( .and .nd
debris ^\hich had beconn tituhHl to ih.m ui tiaii^t In Mt ^^
of the fact that the fisli had bciMi out ot the water nrarlv an hour
and had been .ub,Mt(d to no ^u^ ( ..(tul tu .ttn(nt it did not
occurtoniebnt thit th(N ^^<r. d. id o, it h im b. x ond tht po.^-
bilm of iCMMiii, \^^ uu .n> ot a hoM 1 tnnud upon thun a

time, that feeble resj^iratorv motions weiv ni j)fouiv>s. .\s I
continued to play the water over the lish the resj)iratorv nioiions
became stronger. Shortlv one skate slid over the opening ot the

288

THE AMERICAN NATURALIST

[Vol. XLI

outlet of the tank, closing it, and in a few moments a half inch of
water had accumulated over the bottom of the tank. Thereupon
the skates set up an energetic spouting of water from the spiracles,
— a mode of behavior which had never before come to my notice.
At frequent intervals a large stream of water was ejected from each
spiracle, rising vertically to a height of one or two inches. (The
fish were of uniform size, — about a foot in width across the pectoral
fins.) The animals were not submerged, it should be remembered,
but were less than half covered with water, most of the dorsal
surface, including the spiracular region, being well out. The
vigor and frequency of the spouting and the fact that so many
skates were doing it at the same time produced an effect striking
enough to compel attention. No doubt this behavior has been
observed previously by others. A "Spritzloch" is certainly a
spout-hole. But I could recall having met only the briefest
reference to the use of the elasmobranch spiracle in respiration,
so I postponed the fate of some of the skates and placed them in an
aquarium suppHed with running sea water, with ;i v'ww to watching
their respiratory movements. During the nexi tVw days I observed
the fish as I could, but other work had ])rc( ('(lcii( (\ so ihat i was
unable to carry on any systematic study of tlicir behavior. How-
ever, my impromptu experiments broiioht to li<;lit one or two
facts which seem to me worthy of iiu niion.

As must be well known, the modified first visceral cleft (si)iracle)
serves in the skate chiefly as an iiicurrciit opctiino- tor the rcsjjira-
tory stream. So far as this function i.> concerned, as poiiucd out
by Garman (74), the spiracle is probably of greater importance
in the rays than in the sharks, owing to the fact that the rays, for
the most part, lie flat upon the bottom of the sea, and this habit
places the mouth at a disadvantage as an incurrent respiratory
opening, while in the perpetually roving sharks such is not the case.
These facts are very likely connected with the fact that the spiracles
occur as large openings in all the rays while in many of the sharks
they are either very small or completely close.!.

Many writers make the statement that water may pass either

notes that, whereas the sting-rays have in the spiracular passage a
valvular fold preventing outflow, in the common skate no such

No. 4,S5]

THE FUNCTIONS OF THE SPIRACLE

289

structure is present, so that water may pass either way. Dumeril
('65-70, tome 1, p. 210) states that water usually enters the mouth
through the spiracle, but less frefjuently passes in the reverse
direction.

While at rest on the bottom of an a<|iiai-iinii, the >katc sli^iitly
elevates the head above the surface of tlic I.oKom in tli(> maiiner
described in Brehm's Thierleben (Breiini, 7!), ]>. .isD. wliicli may
well be quoted : "Abweichend von andercii HocU'niisclu'ii lie^jen
sie mit dem Vordertheile ihres Leibes niemals fc.st aiif. sondern
stiitzen sich so auf ihre Brustflossen, dass in dcr Mitte ciii I lolilraum

mit Wasscr zu versoro-en. offnen sie ihre AtluMul.K h.^r. indcni sie
dcii Xolhen zuriickziehcn, fallen die Kiemensackr. M-hlies.eu die
Athemlocher und treiben das verbrauehte Wasser .luicli die
Kiemenspalten nach aussen." Acconiinii- to my observations the
skate takes in water not only by the spiracle but also tliroui^li the
mouth, although considerably more water ent(>rs through the
spiracle than through the mouth.

When fully open the external aperture of the spiracle in the
common skate is nearly elliptical in outline, but the curvature of
its anterior margin is much greater than tiiat of its posterior margin.
The anterior lip of the opening bears tiie rmhinentary gill and
the closing of the spiracle is effected mainly i)y the contraction of
this gilled lip, while the posterior lip, being nearly straight when
relaxed, contracts but little.

pendidiun-like regularity. Dm'ing one of the prolonged resting
periods of llu^ fish, the interval between successive openings is
longer than when the fish is active, and the spiraclt' is not opened
wide,— indcvd. the o])ening may be only a narrow slit. During
more active rt\s[)iration the anterior lip of the spiracle moves back
ami forth with a (piick .lecisive motion an.l the spiracle is opened
to its ittmost width. .Vs the spiracular valve opens, the l.rancliial

tightly closed. At the same iini(> that the spiracle is open, the
moiuh also is opened more or less and a certain (piantity of water
enters. I satisfied myself as to the inward current at the mouth

290

THE AMERICAN NATURALIST [Vol. XLI

by watching the movement of sohd particles suspended in the
water in the vicinity of the mouth. Much the greater volume of
water, however, appears to enter through the spiracle.^ During
an expiration the spiracle is shut, while the mouth tends to close
but does not close tightly. The mouth action was always a little
sluggish as compared with the action of the spiracle, especially
in opening. As nearly as I could determine, mouth and spiracle
closed together, but the opening of the mouth was slightly later
than that of the spiracle. As spiracle and mouth close, the bran-
chial region is contracted and the water contained in the gill
chambers is forced out through the gill clefts.

That water does not flow out through the mouth as well as
through the gill clefts during an expiration is probably due to the
action of a well developed respiratory valve similar to those de-
scribed for teleosts by Dahlgren ('99). The dorsal flap of the
valve (Figures 1 and 3, vlv. d.) is a conspicuous bilobed fold of the
oral membrane, while the ventral or mandibular flap — a less
extensive fold — is broadest in the median region of the lower jaw
and becomes much narrower towards the sides of the mouth.
Judging from the relative widths and the positions of the two parts
of the valve, it appears that the prevention of outflow through the
mouth must depend mainly upon the action of the dorsal flap.
Garman ('74) mentions only the dorsal one of these two folds.

The elevation of the forward end of the fish above the surface
on which it rests would seem to facilitate the respiratory process.

flat-bottomed glass vessel. The vessel was placed upon a higlf table so^that
one end projected some distance beyond the edge of the table. The fish was
induced to lie with its head in the overhanging part of the vessel. I found that
an object held just underneath the mouth could be seen directly through the
head of the fish by looking, at the proper angle, into the spiracle as it opened,
and, similarly, an object held just above the spiracle could be seen by looking
upward into the mouth as it opened. This was sufficient proof that mouth
and spiracle were open at the same time. In order to see the dorsal and ven-

total reflection from the siirfinc of the wntcr H;i\in>: tlir \\;itci- :ii ;i ccrtMiu
depth and looking upward Irom uinlcrn(';ii h the ii\ crluiiiiriiii: xc-^rl at ]\\<\

sufficient clearness, while at tli<" same time 1 had a direct view ot the \ciitral
surface of the head.

292

THE AMERICAN NATURALIST [\''ol. XLI

I watched the respiration of skates in aquaria supplied with running
water, observing the fish at times when they had not been dis-
turbed in any way for several hours. At such times the rate of
respiration was always slow, — usually from 22 to 30 inspirations
per minute. At fairly regular but long intervals there occurred
a break in the regular alternation of inspiration and expiration.
This break was brought about in the following way. Immediately
after an inspiration (and therefore in a period ordinarily marked
by a contraction of the pharynx with closed spiracle and open
gill clefts) the spiracle remained open and the gill clefts remained
tightly closed while a particularly vigorous contraction of the
pharynx caused the contained water to be ejected forcibly from
the spiracle. It is apparently by muscular action that the gills
are kept closed during the spouting, since the pressure of tlie
water in the gill chambers would tend to force open the extei nal
valves. During the spouting the mouth was open, as it is (huing
inspiration, and some water escaped from it, but very little as
compared with the amount ejected from the spiracle. It is doubt-
less due to the respiratory valve that the outflow from the mouth is
not greater. The contraction which caused the spouting was
immediately followed by an expansion of the pharynx, thv spiracle
still remaining open and the gill clefts closed, and respiration
then proceeded in the usual way. In animals which had Ih cii at
rest for several hours, the rate of rcs|)iratioii l)cin<^ \\\vu at its
lowest, the spouting occurred at iiitcr\al> of five to ten iiiimites.

Having found that spouting is a feature i)t' normal respiration
in a resting fish, I next sought to discover what part the spouting
plays in the respiratory process. With this end in view, I observed
the fish under other conditions than rest.

Effrds of Exercise.— The rate of respiration in a fish varies
with the degrec^ of activity. To induce rapid respiration 1 caused
the fi^h U, take exen i.e. This was etfectively .lone l,y gnisping
and 1 lli.th I t fill I I thetaih Th<' iuo>t violent ellofts

(Ij A skate had been un<listnrl)<-.l over ni.-lit in an acpiariuin
supplied with running sea water. When (ir>t observed in the

No. 485] THE FUNCTIONS OF THE SPIRACLE

293

morning the fish was at rest, the respiration being verv slow and
the spouting infrequent, as described above for the resting condi-
tion. I have no record of the precise rates in this case. Tlie
fish was then exercised and immediately removed to a shallow
tray of water for easier observation. The rate of respiration was
markedly increased, rising to 47 inspirations per minute, and a
spouting occurred on the average after every nineteenth insj)ira-
tion, that is, a little oftener than twice a minute. TK\> avnagc
was obtained by counting the number of inspirations within a
period covered by eleven successive spouts. The actual number
of inspirations between two successive spouts varied from 15
to 23.

In a similar case the rate of respiration while at rest was 22
inspirations per minute, with spouting at intervals of several
minutes. After exercise the rate of respiration was 39.5 per
minute, with a spouting after ever}' seventeenth inspiration, or at
the rate of 2.3 spouts per minute.

(2) A skate which had been undisturbed, so far as 1 know, for
two days was found resting quietly against the side of the a(|nannin.
The rate of respiration and the frequency of spouting were deter-
mined. Then the fish was exercised vigorously for five miiuites,
after which it was given five minutes to become quiet so that
observations could be made. Following are the results of the
experiment.

Inspirations Inspirations Spoutings

After exercise 47.5 67 0.71

Increase 58% .500%

(3) A skate which had been under experiment was allowed
to rest for about an hour. At the end of that time the rates of
respiration and spouting were determined. Then durin^r tl)(> next
half hour the fish was subjected to some annoyance l>y irritation

294

THE AMERICAN NATURALIST [Vol. XLI

of the spiracle and neighboring parts (see page 299). After these
experiments the fish was exercised vigorously for a minute or so,
after which the rates were again observed. Following are the
results of the experiment.

The high respiratory rate (47) iiiiiiuMliatcly after the hour's
rest a|)parently means that the fish ha<l not recovered from tiie
effects of th(^ e\})eriiiieius which preceded that horn*, a rate as high
as 57 having been iiuhiced in (he course of these t^xperiments.

(4) In a skate immechately after exercise, conditions were as
shown in the following table.

Inspirations Inspiriitions Spoutings

This skate was then
small vessel of water, w
At the end of the three h

No. 485] THE FUNCTIONS OF THE SPIRACLE

295

(c) with (b) 26% 340%

111 this experiment the rates of respiration and spouting are in-
fluenced by two factors, exercise and the quality of the water,
and the effects of these two factors can not be separated in the
resuhs. The experiment is cited because it shows strikingly,
and in accord with other experiments, that, as the rate of respira-
tion rises and falls, the rate of spouting likewise rises and falls,
but in much greater proportion.

(5) Another observation shows the effect of quiescence. A
skate immediately after exercise breathed 40 times per minute
and spouted twice per minute. After three hours'
(during the first hour of which the fish was extremely re
the frequency of breathing had decreased 44%, while the frc.
of spouting had decreased 08%.

In several other experiments similar to those just drx rili
same general results w^ere obtained. Fish which had Ixrii
quietly for several hours were found to breatlir tV..m l'l' m .JC
per minute, while the spouting occurred ai iiii« rval> of -
minutes. After vigorous exercise the fre(}ucii(y of l.iv
was always increased to a rate between 40 and OO per iiiinn
the spouting occurred once per minute or oftener. Thii^,
the rate of respiration becomes more rapid as the resuh of
following a period of rest, the frequency of s])()utiiig is inc
also, bid in much greater proportion. A veiy rouuli avt rairi'
all of the observations taken tou'ctluT, shows that, when-
rate of respiration is increased about lOO',, the rate of -.p^
is increased at least 500%.

With quiescence, the rates of respiration and s])ontinu-
towards the low resting rates, bui the >j)oniinu- rate falls of
lively much more rapidly than [\\v vn\v of bn-athinu-.

Effects of Partial Asphyxiation.— Is th.^ fr.Miuen.y of sp.
aflfected by partial asj)liyxiation / The behavior of the lisli
first brought into the laboratory suggests this tpiestion. Tl
lowing experiments were made.

296

THE AMERICAN NATURALIST

[Vol. XLI

(1) A skate was put into a rectangular glass vessel measuring
about 12 by 18 inches, containing sea water to the depth of about
3 inches. The fish was allowed to become quiet and then was left
undisturbed for two hours, during which time a copious stream
of water was flowing into the vessel. At the end of this period
the animal was found resting quietly, respiration being at the rate
of 22 per minute, while spouting occurred at very irregular intervals
averaging about 1^ minutes.

The stream of running water was now shut off and the fish was
left in the vessel without change of water for about three hours.
During the earlier part of this time there were alternate periods
of quiet and unrest. In one of the periods of quiet, the respiration
was slow and the spiracle was only slightly opened. But after a
minute or two of these resting conditions, respiration became
markedly quickened, the spiracle being opened wide at each
inspiration, and shortly the fish raised its head and began to swim
about, usually trying to swim up the low vertical side of the aqua-
rium so that the head was thrust out of the water. This activity
lasted usually less than a minute, after which the fish dropped to
the bottom of the aquarium and became quiet, the respiration at
once slowing down to the normal resting rate. Sometimes the
performance was varied in that the quickened respiration which
marked the close of an interval of rest was followed, not by the
swimming activity, but by a vigorous spouting, after which slow
respiration was resumed. At still other times the period of unrest
was marked by both the swimming and the spouting. Occasion-
ally the spouting occurred also in the resting intervals.

During the second hour after the incurrent stream of water
was shut off the alternate periods of rest and unrest continued.
The rate of respiration, however, gradually increased, reaching
a maximum at the end of the second hour when the fish was
breathing 59 times per minute and spouting about once per min-
ute. Respiration was equally rapid during rest and unrest. The
activity was often much more violent than in the first hour of the

in the third hour of the <-\[)('riiii('iii thr rate of respiration
diniinisluMl with increasing rapidity. Following is the record
(the rimning water having been shut off at l.()0 P. M.).

No. 485] THE FUNCTIONS OF THE

297

3.00 P. M. Rate of respiration oO prv minntc

3.20 " " " " ru "

3.45 " " " " .-,()

4.00 " " " 10 "

The spouting continiKMl ;it tlic rate of alioiit once per minute.
The resting periods wviv coiisidcraMy ](iiin(.r than in the j)re-
ceding hours and the activity was less violent. The fi>li cvidcntlv
was becoming sluggish. Returninu- ;i( to oI).s(tv(" tlic

I found the respiration obviously mucli slower and rapidlv dimin-
ishing in frequency. Before I could determine the rate the
respiratory motions suddenly became very irregular and spasmodic
and then the action of the spiracle abruptly stopped. I waited,
perhaps half a minute, and then, fearing a premature end to the
experiment, T turned into the afjiiarium a stream of water, wasln"n(:
it about the head of the fisli. Witliin a nn'nute feeble an<l -low
respiratory movements began, slmr/h/ jollnim/ hi/ four r'Kjorous
spoutingsin rapid succcssiiw. Iie-piration (juickly Ix'came >tr()nL^er
and its rate increased ra])idly, reach inn- is per niimu*' at 4 . L'2 o'clock.
The rate of spouthig, at the same time, was 1..') per mimite. an
increase of about 50^:; over the rate at l.OO o'clock.

At 4.29 the fish was taken out of the water and left lyin- on the
table top. For several minutes it strni^^^led viuonm^ly, hut at
the end of eight minutes the respiratory motions had ceased and

limp. The heart, however, was lu-atin- strongly. TlH-n ' the
animal was put into well aerated s.-a water. At first no sign of
retiu-m-ug activity a])i)eare.l. The spiracle was wide open and
motionless. I therefore began kneading the gills and flirected a
stream of water into the spiracle. Almost immediately very weak,
slow and irregular spiracular motions began, and in the course
of two minutes regular res])iratory movements were in ])r()gress,
although still weak and very slow. The s])iracl<> <lid not close

This was not regarde.l as sp..uting. The action of the >|,iracle
rap-'dly <puVkene<l and siren-rhened, and about four minutes

spouting wlii<-h was then occurring fre<|uently. ^I'he count was

298

THE AMERICAN NATURALIST [Vol. XLI

the spouting occurred five times, while for the entire period of
three minutes there were, on the average, four spoutings per
minute. At the end of the three minutes the rate of respiration
was found to be 41 per minute.

(2) Following is the record of another experiment.
10.00 A. M. A skate was removed from the water.
11.15 A. M. Feeble respiratory motions of gill chambers
and spiracles still in progress at the rate of 28 per minute.
The spiracle is continuously wide open, its anterior Hp
contracting very slightly at each expiratory movement.
The moutli is contiiiuously shut.

11.20 A. M. The skate is put into well aerated sea water.

11.21 A. INI. The spiracular action is stronger and weak
mouth action begins.

11.25 A. M. The spiracle closes completely at each expira-

These experiments, then, so far
conditions of gradual approach t(
is left in a small volume of unchn
increasing restlessness attended b^
greater frequency of spouting,
there were, earlv in the experiinei

No. 485] THE FUNCTIONS OF THE sriUACI.E 299

Later in tlic (>x|.erinu'i
high, with IV('(|ii('nt s\)>
At the near a])i)n)ac
ually diminishes, l)Ui
frequency than under i

slow, but during the (
spouting oeeurred with
in one minute. Wilhi

gradually fell, as the nite

with much greater fre(ni(Mic
Spouting Induced hii 7V.
vations led me to trv'tlie t
in the vici.n-ty of t'hr cm
margin of the spiniclc \\;i>
rod or with a stiir hit of
diately a spouting from 1..
stimulation, or persistent iii
in a vigorous spouting fron
had once been provoked
repetition of the stiiniil.itid
spouting. But after an iin
renewed stimulation nsiiall\
One skate was especiall;
much more prom])tly aixi
animal was exjjerinicnied \

only. Thisone-sided.p<.ntnii:ua>i»
and then from the other, in fairl_\ ra|.
stimulation of the spiracles aherna

300

THE AMERICAN NATURALIST [Vol. XLI

was squirted from the spiracle with such energy as to rise through
an inch of water and some seven or eight inches vertically upward
into the air. Frequently the stimulation was followed, not only
by the spouting, but by a sudden dash to another part of the tank,
as if to get away from the annoyance.

Tactile stimulation of the skin in the region of the eye also
usually caused spouting. A gentle touch uj^oii the outer corneal
surface of the eyeball almost invariably provoked a particularly
vigorous spouting from the corresponding .s))ira( le. Indeed, stinni-
lation of the cornea w^as found to be a more certain way of ]>r()-
voking spouting than stimulation of the spiracle itself. The
response was always immediate and definite and ui nearly every
instance unilateral.

I tried also the introduction of solid materials of one sort and
another into the gill chambers. I first tried sand, allowing a little
to sift into the spiracle when it opened for an inspiration. Some-
times a spouting resulted, but equally often, even though a con-
siderable quantity of sand was introduced, no response whatever
followed.

P^xperimenting in a similar way with another fish, T found in the
aqiiarimn some shre<ls of filmy substance of doul>tful nature.
Tlu-v appeared like hits of slou-he.l-otf .kin. It well exemplllies
the impiompni < 1, u .< in ..t dl -.1 di. m (\punnuus ilial, makino-

hand, I caused some of this doubtful filmy substance to he sucked
into the spiracle at an inspiration, invariably material of this
sort was promptly expelled by spouting. Often one or two in-
spirations intervened between the one by which the foreiun material
w^as drawn in and the spouting by w^hich it was expelled. I sually
the spouting occurred from both spiracles at once,— rai-ely from
only the one at which the foreign material was introduced. The
material was always ejected by the same sjiiracle at which it
entered.

In the one-sided spouting the action of the unstinuilated spiracle
appeared to be uninterrupted. The stinnilated spiracle simply
remained open during one closing of the other.

Summarizing the foregoing accoutu, it apj)ears that the spiracle
of the common skate serves chiefly as an in-take for the respiratory

No. 485] THE FUNCTIONS OF THE SPIRACLE 301

stream, but at somewhat regular intervals the stream is reversed
and an expiration takes place via the spiracle, which thereby
becomes a spout-hole. With quickened respiration due to exer-
cise, the spouting occurs much more frequently than in the resting
fish. Also, when a skate is confined in a small volume of water
which is not changed, respiration is quickened and spouting occurs
much oftener than under normal resting conditions. Wliether
in this case the higher rate of resj)ii'ati()n is due dircctlv to the con-
dition of the water, or to the activily vaumhI l)y tiie unfavorable
quality of the water, I am unable to say. :\rKtMidi ick (7'.t ) states
that, in the presence of an insufficient supply of oxygen tlie fish
"breathes hurriedly." Finally, spouting occurs with t'X(•e»!^c
frequency in skates which are just beginning to recover fioni an
advanced stage of asphyxiation. What, in view of these fatts,
is the probable r6le of the spouting, so far as it is a resj)irat()i v act ?
May it not be roughly analogous to " taking a deep bn atli r An
occasional reversal of the respiratory stream may sei\t' to < !ear
out the gill chambers, resulting in a more nearly complete ehanue
of water in them. The ureater frequency of the sjxiniinL;- when
respiration is (luiekentMl. by whatever cause, and its exce^sivt^
frequency in recovery from asi)hyxia indicate, I tliink. thai it has
some importance in the way of increasing the ellicitMuy of the
respiratory process.

Spouting in response to tactile stinuiiation in ihe vicinity of the
spiracle indicates that the fish may, uiKh-r natural comhtions. em-
ploy the spout-hole as a means of expelHiiL;' foi'eiuii soHd materials
from the gill chambers, or of dislodging objects from the surface
of the body in the region of the spiraeh^s and (>yes. The bt^havior

the sea-bottom havt> a habit of settling lh(Mn>elves into the sand

apparently, be very likely to sift into the spiracles, and one might
suppose that sand would be particularly irritating. Hut in my
experiments the skates were indill'eriMU to the introduction of con-
siderable quantities of sand, while soft iilmy materials were
promptly spouted out. On fiu-ther consideraiion. it occurred to
me that sand, being a finely divi(le<l substance, would easily wa>li
out through the gill clefts, whereas, being heavy, it could not so

302

THE AMERICAN NATURALIST [Vol. XLI

readily be forced up through the spiracles. But the larger frag-
ments of soft material (such as bits of sea-weed) are likely to be
caught on the gill-rakers, tending to clog the branchial passages,
and could best be dislodged and expelled by a reversal of the cur-

The prompt, vigorous, and almost unfailing response to a touch
upon the cornea suggests that the fish regularly employs spouting
as a means of keeping the eyes unobstructed. The external
opening of the spiracle is so near the eye that a stream spurted
from the spiracle would readily wash away foreign objects which
settle upon the eye.

Regarding the spiracle as one of a series of visceral clefts which
were primitively similar in structural relations and in function, it
is evident that, serving as it does such a diversity of uses, it has
come to differ from the more posterior visceral clefts quite as
markedly in its function as in its structural conditions.

tup.ltoc;raptiy

79 IliiMhlMi. 1>mM s 1)„ IimIh Uip/.g x\i+i2() pp ,

'99. ihf M.iMll.irv und Mandibular Breathing \alves of Telcost
!-i>h< s. Zool. P,ull., Vol. 2, pp. 117-124, 3 figs, in text.

'65-70. Histoiro Naturolle dos Poissons. Paris. Tome 1, 720 pp.,

'74 On the Sk itcs (P ij r) of the 1 lutein Coast of the United utiles
Proc lio^tonboc .Nit IIi',t , ^ol 17, pp 170-181, 1 fig m text
['Kendhtck, J. G.

79 On the R<.piritor% Mcnemcnts of I i.he^ Joinn \n it tnd
Physiol., Vol. 14, pp. 461-466, pi. 28.

A CRITICAL AXD STATISTJCAT^ STUDY OF THE
DETERMIXA'llOX OF SKX. I ARTICL L.\RLY
L\ IIl'MAX ()lTSl'inX(;.'

F. H. PIKi:.
I. Introductiox.

Cu£not ('99) and Strasburger (:00) summarized the evidence
in favor of the heredity of sex in animals and plants, respectively.
Rauber (:00) in the same year as Strasburger, declared for the
heredity of sex in man.

Bateson in 1902 suggested that the ISIendelian law might apply
to the heredity of sex. Castle (:03), accepting Cuenot's and
Strasburger's views without question, formulated an hypothesis
to account for the heredity of sex in accordance with ^Mendel's
law. Weldon (:01) had already shown that ^Mendel's original
results with cotyledon color in peas differed from the theoretical
numbers by something less than the limits of error. At the time
Castle's theory ap])eare(], it occurred to nic to gatlicr statistics
of births in order to (letcriiiiiic in a similar way the probaliility
that the actual inimhcrs of male and tVinalc l)irtlis would he the
numbers deinaiulcd by the hypothesis.

II. Rkview of Previous Work.

The idea that the sex of the offspring could be influenced by
changing the environment of the parents or of the very young
embryo has long been current, "^^ing's ('S3) experiments on
tadpoles, in which he was apparently able to control the sex by

' This study was begun under the direction ..f Pn.f.-M.r C. !I. luarmnunn
of the Department of Zoologj' of Indiana rniv.M>ity an<l \Na^ roinplrted in
the Hull Physiological Laboratory- of tlu- I nivcrMiy ui ( hi.a-o, Ti,r author
desires to express his obligations „. th. lualtl. .mI,,-..,-. an.l n-i^tra.-who
have supplied hi.n with statisti..; ^t.. hi^ ..„ll,.a.ni.. u, tb. Hull Lalx,nUo:y for

3oa

304

THE AMERICAN NATURALIST [Vol. XLI

changing the nutrition, have been cited as a demonstration of
this point. Statistics of human births have been judged in such a
way as to lend some support to this view. A good review of this
aspect of the question has been given by Geddes and Thompson
(:01). The vaHdity of such conclusions has been well discussed
by Newcomb (:04), who has made a statistical inquiry into the
probable causes of sex in the human subject. Newcomb concludes
that the causes of sex are beyond voluntary control.

Cuenot repeated Yung's experiments on tadpoles with contrary
results. Eggs from the same mother, but of different layings,
gave a more constant proportion of males to females than Yung
obtained. The ratio of males to females in the young (54.85%
females) did not differ materially from the ratio (61.5% females)
existing among the metamorphosed tadpoles found in a state of
nature in the vicinity of Nancy. Born ('81) found 52 per cent
of females in the metamorphosed tadpoles near Breslau. Gries-
heim ('81) found 63.63 per cent of females in young Rana tempo-
raria in the vicinity of Bonn. Pfluger ('81) found 64.5 per cent of
females in the same vicinity, and 86.8 per cent near Utrecht. The
percentage of old females in the latter vicinity he found to be 51.2.
Pfluger concluded that the sex was determined in the egg. From
his results on tadpoles and other animals, Cuenot hkewise con-
cluded that sex was not influenced by the conditions of develop-
ment. He decided further that there was a certain sexual ratio
common to the frogs of any particular vicinity, and that this ratio
might vary among frogs of different locahties. In view of the
comparatively small number of frogs observed, the last conclusion
may possibly be open to question.

Von Malsen (:06) and Issakowitsch (:06), the former for the
worm Dinophilis apatris and the latter for Daphnia, have recently
reaflfirmed the statement that an abundance of food and a low
temperature cause a greater number of eggs to develop into females,
while a higher temperature and a scarcity of food result in the
development of a greater number of males. The food supply,
according to them, is the main factor in this process, nnd the
temperature acts only indirectly by influencing the imtriiioii.
It is to be remembered, however, that in experiments dealini^ with
a whole animal, it is difficult to exclude all causes except food and
temperature.

No. 485]

THE DETERMIXA TIO.X OF Si:X

305

Strasburger (:00) made main- cxpcrimnits with (lid-emus j)l:ints-
growing them on various kinds of soil and imdcr various condi-

tions, in the attempt to

modify tlie

sexual ratio,

. Tiio followir

resuhs with Melandrinm

albnm may 1

l)e cited as an

example:

lABLE

Kind of i^oil

Females

/' '

410

562

Unfertilized garden soil

235

282

Fertilized field soil

384

479

124.4

Unfertilized field soil

254

307

120.8

Sand

321

411

128.0

Totals

1604

2041

Mean 127.2

Thus sexual ratios for groups of plants grown under the most
diverse nutritive conditions did not differ greatly from the mean.
Strasburger concluded that an arbitrary determination of sex in
dioecious phanerogams has never been accomplished, and he is
inclined to apply this conclusion to all plants.

Rauber (:00) studied statistically the distribution of sex in man.
He found everywhere an excess of male births, but this early excess
in the young was changed, because of the greater mortality of
the males,^ to an excess of females in later life, and particularly
in old age. He showed that the sexual ratio for Europe was 1000
female to 1060 male births, and that this mean ratio was fairly
constant in the different parts of Europe. Reasoning on the basis
that, if sex was determined by environment, the great diversity
of external conditions in tlie different parts of Europe should
cause a considerable difference in the sexual ratios for the differ-
ent countries, he concluded that sex is hereditary in man.

x'Vccording to Rauber, there is uonnally an excess of female
births in horses, sheep and certain other donic.^tic animal-.

Punnett (:04a) has made a statistical stndy of the disirihntion

306

THE AMERICAN NATURALIST [Vol. XLI

of male and female births in London, in order to determine whether
or not the sexual ratio is affected by the nutrition of the parents.
For this purpose he divides London society into three groups,
following Rowntree's (:02) division of the society of York. These
groups are (1) the servant keeping class, (2) the artisan class in
which the family earnings are in excess of 26 shillings a week,
and (3) the laboring class in which the family earnings fall below
26 shillings a week. Rowntree found that, compared with a
standard dietary containing 125 grams of proteid and possessing
a total energy content of 3500 calories, the first group has a dietary
containing more food than is necessary for the maintenance of
health; that the second class has, in general, a sufficient diet,
although the family must practice strict economy in order to pro-
cure it; the third class is, as a rule, seriously underfed, the average
deficiency in proteids amounting to as much as 29 per cent. Assum-
ing that these considerations apply to I^ondon as well as to York,
Punnett finds that there is either no effect upon the sexual ratio
which can be attributed to parental nutrition, or, at most only a
very small effect. He finds also that the statement of Diising as
to the greater proportion of males among the first born children
is supported by the statistics of the English lying-in hospitals.
Furthermore, mothers whose first birth occurs between the ages
of nineteen and twenty-three years bear a larger proportion of
males at this birth than mothers whose first birth occurs either
earlier or later in life.

IIL Statistical Data.

On the fundamental errors in the statistics. — Rauber (:00) has
considered the errors in even the best statistics, and only a
brief discussion of them will be given here. In order to compute
the exact sexual ratio, it is ntH-essarv to obtain statistics of all
births, both premature and full term, livinu' (»r still born. The

homologous or duplicate twins, (lc\ eloped tVoin a sinulc ovum,
and invariably of the same sex, should be coiintcil as a single
birth. The author has at hand no snfhcicnt data n|)on which to
base an idea of the ina^-nitndc of the error which niiuht be uitro-

No. 4S5]

THE DETERMINATION OF SEX

307

duced by counting such twins as two births. Of the eighteen
cases of twins and triplets considered by Wilder (:04), twelve
pairs of duplicate twins were females. If such a large propordon
of all duplicate twins should be females, the error introduced
would be considerable, and the preponderance of male births
increased. The number of cases given, however, is too small to
warrant drawing conclusions as to the relative frequency of male
and female duplicate twins.

The sexual ratio. — The sexual ratios for eleven European
countries, as they existed sometime during the latter part of the
nineteenth century, have been taken from the tw enty-eighth annual
report of the Massachusetts State Board of Health through the
courtesy of the late Dr. Samuel W. Abbott. The figure for Eng-
land from 1628 to 1642 is that given by Lexis ('92). The ratio
for the United States was computed from 2,021,955 births —
1,038,432 males and 983,523 females — the statistics for which
were furnished by the health officers of the various states liaving
reliable statistics of births.

German Empire (1 871-1 S80) 1062

Switzerland 1063

Austria 1067

Italy 1071

France 1063

Belgium . . . . • 1058

Holland 1063

Denmark 1058

England (modern times. Living births only) . 1038

England (1628-1642) 1068

Sweden 1060

Norway 1061

Massachusetts (1876-1896) sdll births included . 1066

Massachusetts (1856-1875) living births only . . 1055

United States 1056

Mean of all ratios 1060

308

THE AMERICAN NATURALIST

[\'oL. XLI

A considerable increase in the sexual ratio occurs when still
births are included. The sexual ratio for the city of Chicago,
based upon all births reported in the years 1898 to 1902 inclusive,
(141,233), is 1065. During this period 4828 males and 3554
females were prematurely or still-born. If these premature and
still-births are deducted from the total number reported, the
number of males to 1000 females is 1035. It becomes necessary,
then, to decide whether or not still-births shall be counted. Still-
births must be reported under penalty. Since the living children
are the ones voluntarily reported, their record is not complete;
but we may suppose that the parents are as apt to report a birth
of one sex as of the other. The statistics of living births, there-
fore, in localities where birth registration is not compulsory,
probably approach more nearly to the true ratio than the ratio
based upon both still- and living births. If birth registration is
compulsory, the ratio should be computed on a basis of all births
reported. The ratio for Chicago computed on the basis of living
births only is very nearly the same as the ratio for England. The
ratio for Massachusetts, computed from living births only, is less
than that based upon both living and still births. It is probable,
therefore, that the incompleteness of the statistics is the most
serious source of error.

The constancy of the sexual ratio.— An examination of the sta-
tistics shows a remarkable constancy of the sexual ratio in all
parts of Europe and in the United States, for a period ranging
from 1856 in Massachusetts, through 1871-1880 in the German
Empire, to the year 1902 in Chicago. During these years and in
the various countries, there were periods of war and peace, of
famine and plenty, beside a great variety of racial and climatic
conditions. Yet the greatest variation from the mean, exclusive
of England, is only eleven in 1000 — a difference of one per cent.

In the same country for a period of years, the ratio is approxi-
mately constant. As an example, we may take the statistics for
England (Tal)le III) during the twelve years from 1888 to 1899,
inclusive.

No. 485] THE DETERMINATION OF SEX

309

Table III.

(Taken from Ses

sional I aj)ti.s (

. t le ouse o

.OK s)

Year

Total Births

Mal,.<

1899

928646

4731 72

.];,;, 4 74

!();>(,

1898

923265

468920

1032

1897

921693

469180

452513

1037

1896

915331

465660

449671

1035

1895

922291

468886

453405

1034

1894

91457^

453016

437273

1036

1892

897957

456622

441335

1034-5

1891

914157

465660

448497

1038

1890

869937

442070

427867

1033

879868

447172

423696

1033

888

885944

451218

434726

1037

10,864,950

5,527,287

5,336,663

1036

It wi

11 be seen that the ratio for any

one year does not differ by

than four in one

i thousand fror

n the mean calculateti from

than ten million

births. As a

further example,

the sexual

in Massachusetts, based upon li-^

,'ing births only, for the years

1856 to 1875 inclusive

'. is 1059. The

• ratio for the yei

ars 1876 to

1896 inclusive, based upon a considerably greater number of births,
is 1053. The mean for the two periods is 1055. The sexual ratio
for the period in which the Civil War occurred differed by approxi-
mately one half of one per cent from the later period of peace,
and by less than one half of one per cent from the mean of the
two periods. If external conditions exerted any effect upon the
parents in such a way as to change the sex of tlie offspring, the
change due to such influences was not greater than one in two
hundred.

Social, political and material conditions in i-aiglaml (iiuing ilie
for the two periods — KHIS to 10(10, and lO.'Kl to lOOO re-])ecti\ ely

310

THE AMERICAN NATURALIST [Vol. XLI

of Chicago computed upon the living births only for a given
period, and upon all births for the same period. It does not,
therefore, appear necessary to assume with Strasburger that the
sexual ratio for England has changed to any considerable extent
in two hundred years.

The effect of a war upon the sexual ratio.— It has long been a
current belief that more males were born in a period following a
war than in a similar period of peace. Newcomb considers this
statement unworthy of serious consideration. It may be said
that, so far as the United States is concerned, such statements are
based upon an insufficient number of births, and that the statistics
are for the most part worthless. I have many letters from state
health officers to the effect that there are now no reliable statistics
of births in their respective states.

The sexual ratio independent of external conditions. — In view
of the remarkable constancy of the sexual ratio under diverse
social, poUtical and material conditions and for long periods of
time in different races, it seems incredible that the determination
of sex should be dependent upon external conditions.

If the sex of the offspring is independent of external conditions,
what is the determining factor? Two general explanations are
open. There is first the possibility that sex is determined by a
series of accidents, as Newcomb suggests, and second, the possi-
biUty that sex is hereditary.

The possibility that sex is determined by a series of accidents. —
Newcomb likens the sex of a child to a particle floating on a
stream of water. In the early part of its course the stream is
single, but an obstacle divides it into two at the lower part. A
particle entering the stream at the upper part may pass on either
side of the obstacle, the exact course depending upon a multitude
of accidental causes up to a certain point, after which its course
on one side of the barrier or the other is fixed. So with an ovum.
In its early development, there is the possibility of developing into
either a male or a female, the sex depending upon a series of acci-

Newcomb showed from statistics that the i)r(.l.ahility that twins
will be of the same sex is .77, and the pro})ahiiity that tlun- will be
of opposite sexes is .23. It is impossible to tell froni Newcomb's

No. 4So]

THE 1)I-:TI:NM IXATIOX OF SEX

311

tables whether duplicate twins wvw fxcliidcd, as thcv slioiild have

been excluded, the probability iliat ..nliiiarv twins woiiM l)e of
the same sex nii<^ht have bccMi even less than .77; for diij)licate

Sex determined before the first cleavage of the ovum. ].v\ us
now examine into the bearing of these con.sideratioiis upon New-
comb's hypothesis. To continue his simile, two j)arti( lcs stariiiiij;
together will have a greater chance of remaining touctlier and
passing on the same side of the barrier than two particles somewhat
removed from each other. Himilarly. two ova (lcveloj)ing together

offspring of the same sex than two ova developing at ditrerent
times, but neither the two particles nor the two ova iiivariablv
follow the same course. If any series of accidents acting upon the
ovum after fertilization is to determine the sex of the twins, it is
incredible that it should always ])r()duce the same re>uh in lioth.
Since, however, duplicate twins ai-e always of the same <e\, this
view becomes untenable, and we must limit the action of a series
of accidents to the ])eriod j)receding and j)ossibly including fertili-
zation. The conclu.sion that at or inunediately after fertilization,
the sex of the offspring is determined once for all seems inevitable.
The effect upon the ovum of any series of accidents must cease
before the first cleavage is accomplished.

Is sex determined by either parent alone?— Having concluded
that the sex of the offspring is determined at or before the time
of fertihzation, we may inquire further whether the sex of the
offspring may not be determined by the ovtiin alone, or by the
spermatozoon alone. First, the ovum may have the potentiality
of developing into either a male or a female embryo. Dnring
maturation the chromatic material neces.sary for the (iev(>loj)inent
of an embryo of one sex is cast off in the polar bodiis. and that
necessary for an embryo of the other sex is retained. The sperma-
tozoon thus plays a purely asexual role. This hypothesis ])o>tH-
lates a qualitative reduction of the chromatin in maturation.
According to another variety of this hypothesis the ovmn assumes
the asexual role, and the sex of the embryo i> .letermined solely
by the spermatozoon. Since all spermatozoa do not [)roduce

312

THE AMERICAN NATURALIST

[^'oL. XLI

embryos of the same sex, there must have been, at some period in
the development of the spermatozoon, a quahtative reduction of
the chromosomes, those necessary for a male going into one
spermatozoon, and those necessary for a female into another.
If we accept this hypothesis, we must show why a constant and
unequal proportion of all ova or of all spermatozoa have chromo-
somes, e. g. the accessory chromosome, which will produce an
embryo of a certain sex. As an alternative hypothesis we may
suppose that both ovum and spermatozoon play a sexual r6le,
and that the sex of the embryo, in common with other character-
istics, is determined by both sexual elements. This view, as I
shall show subsequently, is the more probable.

If we cannot explain the cause of sex by postulating a series
of accidents of unknown nature occurring after fertilization, can
we explain it on the second possibility, — heredity? And if so,
which of the two great laws of heredity are applicable to the case ?
The first question I shall answer in the affirmative, and proceed
to the discussion of the second.

The application of Mendel's law.— On the basis of Mendel's
law we must suppose that each ovum has equal chances of develop-
ing into a male or into a female embryo. Given two thousand
ova, chosen at random, the chances are even that a thousand of
them will develop into males and one thousand into females. We
might reasonably expect also that in some groups we would find
an excess of males, and in others an excess of females, but the
mean of all groups would be 1000 each of males and females.

Punnett (:04b) in order to test Bateson's suggestion, attempted
an enumeration of the sexes in Carcinus moenas. He found an
excess of females in groups of individuals of the same size, but
this excess decreased in groups of younger individuals and there
were indications of an approximately equal distribution of the
sexes at the time of hatching. The exact proportion of the sexes
at the time of hatching could not, however, be determined.

Mcintosh ('04,) from a study of the Norway l()]).ster. concluded
that the young were hatched in about equal |)r(»j)()rti(ni> nf tlie
sexes, but was not able to determine the exact pioportioii.

Taking the English statistics given in Table III as a basis, we
may compute the probability that the actual distribution of males

No. 485]

THE DETERMINATION OF SEX

313

and females would be obtained bv such a random choice. Instead
of the theoretical distribution of 1000 males and 1000 females,
the actual numbers of males and females in 2000 births are 1017.6
and 982.4 respectively, or, for convenience in calculation, 1018
males and 982 females. The probability that, in choosing at
random, we should obtain such a distribution is .60984 X 10"^"°.
The probabihty that in eleven siuh choices, we would alwavs
obtain the same (Iistiil.ini..n is infinitesimal. The probabihty
that, in every case where \\\v l.iiihs are numerous enough to be
representative of the actual eondiiions, we siiould alwavs obtain
practically the same distribution is ])raetieally zero. There is
about the same probability that Mendel's law UMs for all these

A single concrete example taken from organic cheniistrv will
serve to emphasize this point and perhaps to make this mathemati-
cal abstraction clearer. In the transformation of aeetaldehvdc to
lactic acid by the addidon of hydrocyanic acid, saponification and
oxidation, each of two isomeric forms of lactic acid is, on the
theory of probability, equally likely to be jyrodnced. 'J'he two
forms differ in optical activity, one being dextro-rotatory and the
other laevo-rotatory. Kx])eriinenlally, it is found that the two
forms are actually ])roduced in exactly e(|ual amounts, and the
mixture of the two is o|.tically inactive. A variation of from
three to seven per cent from the theoretical yield w(.uld he fatal
to the theory of probability. In u'eneral. in the synthesis of organic
bodies in which two isomeric forms are in.ssif)|e ;ind tlieoreticaliv
equally probable, the experimental results agree much more <'loselv
with the theory than do the statistical results of hmnan })irths.

Neither ovum nor spermatozoon play asexual roles. — It is evi-

314

THE AMERICAN NATURALIST

[Vol. XLI

who maintain the truth of such a hypothesis to explain the
nature of this unknown mechanism.

In the case of the accessory chromosome (McClung, :02), the
statement is made that it occurs in one half of the spermatozoa of
Orthoptera and Hemiptera. If we are to suppose that the acces-
sory chromosome acts as a sex determinant, and that sex characters
are to be treated as if they were Mendelian alternates (Wilson,
'07), we should find a sexual ratio equal to unity or differing from
unity by an extremely small per cent. We cannot, however,
account for the determination of sex in the human subject on any
basis of an equal division of spermatozoa into male and female
producing sperms, unless we suppose, as Wilson concedes for the
sake of argument, that sex may be modified by external condi-
tions. The statistical evidence is strongly against this alternative.
If it can be shown that the accessory chromosome occurs in the
spermatozoa of a species in the same proportion as the sex to which
it gives rise occurs in the young of that species, the statistical
evidence in its favor will be increased. At present, there is no
such evidence in its favor, as we do not know the exact sexual
ratio of the species in which the accessory chromosome has been
observed.

The strongest evidence known to the author in favor of the
Mendelian theory of dominance in the determination of sex is
that cited by Harper (:07) in regard to plants. That the stamens
should develop and the pistil be suppressed in the fungus-infected
female plants of the campion is strongly suggestive of the recessive-
ness of the stamens under ordinary conditions.

The application of Galton's law.— To explain the remarkable
constancy of the sexual ratio by Galton's law, we have only to

is inherited equally from the paternal and nuiternal ancestry;
and to explain the preponderance of males in the itrcscnT uciit ia-
tion, we assume that in this ancestry for fiveor six u;(Mi(M;iri<iiis back,
there has been a preponderance of males, lii this wr arc justi-
fied since there is direct statistical evidence that, for more than
two hundred years, there has been an excess of male birtlis in
England. Accepting the statistics as hein<;- reasonably accm-ate,
the accordance with Galton's law of ancestral inheritance is much
closer than with Mendel's law.

Xo. 485] THE DETERMINATION OF SEX

315

IV. The Biological Significance of the Sexual Ratio.

Rauber, apparently taking the view that an excess of females is
the normal condition, explains the present preponderance of male
births in man by supposing that those tribes or families which,
in primitive times, had the greatest proportion of males would
possess a certain advantage in warfare and thus be enabled to
overpower those in which there was a larger proportion of females.
The male preponderance, once estabhshed, would be perpetuated
by heredity. This ingenious explanation, does not, however,
account for the excess of females among the domestic animals.
What the sexual ratio was in primitive man we have no means of
knowing. Neither do we know what the sexual ratio was in horses
and sheep before they were domesticated. One would expect
that the sexual ratio in wild animals would de})end somewhat
upon the mating and breeding habits of a species. In those species
of birds, in which one male mates with one female for a season or
for Hfe, we might expect that the sexual ratio would be nearly
unity, the excess of one sex or the other depending on which one
was exposed to the greater dangers and had the less chance of
growing to maturity. In herds of wild horses, cattle and ])i.s()ii,
there are many females to one male. All but the .stn)ii<:(\sr nialf>
are killed off by the others and the number of adult males is thu>
kept down to 'the needs of the herd. Those iii.livi.hials which

than males, and it is iM)>sil)le, in fact nioiv than pi-obablc. that man
has unconsciously, by selection in breeding, increased the pro-

No one sexual ratio may be taken as the standard. — If sex is

herclitary, we ini-lit reasonal)ly expect that the relative mnnbers
of male and female births in any species would be those which,

at the period Of sexual' niatm-ity of its individuals the irrlatest
advantage in the >trn,uol,. for exist^uv so far as the pnMlu.-tion of

enj.uMhe maxiinuni reproductive power, and this cnn'diiiun would
be fulfilled when there were no superfluous, sexually mattu-e males
or females.

316

THE AMERICAN NATURALIST [Vol. XLI

Let us suppose that a species possesses the maximum reproduc-
tive power when there are x males to n females, the relative magni-
tudes of X and n depending upon the breeding habits of the species.
Let us suppose also that a males and b females die before reaching
sexual maturity. The number of males born will therefore be
a + X, and the number of females b + n. The sexual ratio will
be , J or k , X 1000 if we wish to express the number of
male births to 1000 female births.

In a monogamous species, such as the American robin, the repro-
ductive power of the species would be at a maximum when there
were equal numbers of sexually mature males and females. If
the males are more likely to be killed off than the females, a would
be greater than b, and the sexual ratio would be greater than unity.
Surplus males or females would die off without reproducing. In
a polygamous species, such as the ox, it is not necessary that there
be equal numbers of sexually mature males and females to give
the species its maximum reproductive power, and a + x might
well be less than b + n. The sexual ratio would in this instance
be less than unity. The relative proportions of the sexes in any
species may, therefore, be looked upon as one of the physiological
adaptations of the species, determined by the conditions of its
existence.

V. The Experimental Point of Attack.
If sex is inherited according to Galton's law it should be possible,
by suitable selection of the parents, to establish a strain of animals
or plants in which males or females occur with any desired degree
of frequency compatible with perpetuation of the species. The
practical benefits of the favorable results of such an experiment
to the dairy and grazing interests of the country would be difficult
to estimate. Castle has recently ])ublislifil some experiuuMits on
the effects of inbreeding, cross-brmiinu', and selection upon the
fertility of flies (Drosophila) in wliieh lie foniid no marke.l elinnge

No. 485]

THE DETERMINATION OF SEX

317

often be of different sexes while duplicate twins are invariably of
the same sex if nutritive or any other conditions outside of the
ovum itself are responsible for the sex of the offspring. It is per-
haps conceivable that, in the case of ordinary twins, the placental
circulation may be more highly developed, and the nutrition con-
sequently better, for one twin than for the other, or that some
peculiar local characteristic of the uterine wiill inay aflVct one
twin more than the other. The probability of any mk Ii dissimi-
larity of conditions in the case of dupHcate twins, \v!ui(> tli(^ same
placenta supplies both w^ith nutriment, and any local peculiarity
of the uterine wall affects them equally, is very small. It must
therefore be admitted that ordinary twins may be subjected to
more diverse conditions during development than duplicate twins,
and it is conceivable that the latter might sometimes be of opposite
sexes if we could vary the conditions during development. This
would manifestly be a matter of great difficulty in mammals, but
a simpler method of attack is open.

Roux ('85) and others have shown that the indivi.lual blasto-
meres of a frog's egg will, when separated from tlic otluM s. develop
into complete embryos. Such embryos are presmnahly compara-
ble in all respects to duphcate twins, and if by any means we might
cause two blastomeres from the same ovum of any animal nor-
mally reproducing sexually to develop into embryos of opposite
sexes, we w^ould have a demonstration that sex was not determined
at the time of fertilization of the egg. Failure to produce from
the same egg two embrj^os of opposite sex would be evidence that
w^e have, at present, no known means of changing the sex of the
embryo after fertilization of the egg. It is incumbent upon those
who maintain that sex is determined by the environment to show
that two embryos of opposite sexes can be produced from the
same ovum. The experimental solution of the ])rol.lem of the
causes which influence the sex of the offsprinu". as well as the
significance of sex itself, is to be souirhr in the >im])h> ceU whose

It is obvious, also, that the problem of sex determination is t)ut a
particular phase of the much wider problem of the extent to which
the ovum may be modified by a change in the external environ-
ment. Furthermore, if we acquire experimental data on the deter-

S18

THE AMERICAN NATURALIST [Vol. XLI

mination of sex, we will at the same time acquire experimental
data on the question of a period of sexual indifference in the devel-
opment of the individual. If sex is determined, as appears prob-
able from the statistical data, at the time of fertiUzation, it is
difficult to conceive of a period of real sexual indifference in the
histor}' of the individual. But if we can influence the sex of an
individual after fertilization of the ovum, we will at the same time
demonstrate a period of sexual indifference in development.

The bearing of artificial parthenogenesis on the problem of sex.
— In a personal communication to the writer, Dr. Woelfel has
suggested that if, by any means other than fertilization by a sperma-
tozoon, we are able to cause an ovum of an animal which normally
reproduces bisexually to develop to sexual maturity, we will have a
demonstration that one parent plays a purely asexual role in the
production of sex. This conclusion, however, does not follow
necessarily. Moreover no individual arising by artificial partheno-
genesis has as yet grown to sexual maturity. Whether this failure
of normal development is due to improper nutrition of the young
or to a lack of some essential detail in fertilization cannot, perhaps,
be stated at present. One is inclined to regard the production of
a sexually mature individual, whicli may in its turn reproduce, and
the transmission of certain heivditaiy characteristics to the off-
spring as two essential details of inuli/ation. Until these phe-
nomena have been imitated by artificial means, one is loath to

by artificial parthenogenesis (Loeb :06). The study to artificial
parthenogenesis may have an important bearing upon the deter-
mination of sex, but the true significance of the work already done
is not apparent.

VI. Summary and Coxcu sioxs.

The statistical study of the distribution of sex in man sliows
that there is a sHght but constant excess of male Ijirtlis. The
greater mortality of the males leads to a preponderance of females
in old age.

There are certain unavoidable errors in the statistics, the two
most serious being (1) incompleteness, and (2) disregard of dupli-

No. 485]

THE DETERMINATION OF SEX

319

cate twins. It is not probable, however, that these errors are in
such a direction as would change the sexual proportion if we
could get absolutely correct statistics.

The sexual ratio is remarkably constant in widely different
localities and at widely different times; in a given locality the ratio
is not altered by the varying social and material conditions of the
parents, as indicated by statistics.

The study of duplicate twins shows that if sex is determined
by a series of accidental causes, such causes cannot be operative
after the fertilization and first segmentation of the ovum.

The logical conclusion from the statistical data is that sex is
hereditary. Mendel's law does not apply. The constancy of the
sexual ratio for more than two hundred years may best be ex-
plained by supposing that sex follows Galton's law of ancestral
inheritance.

If sex is hereditary, we may explain the significance of the sexual
ratio on the basis of natural selection by supposing that the pro-
portion of the sexes in any species is such as will give that species
the maximum reproductive power at the time of sexual maturity
of its individual members. The sexual ratio may be expected to
vary for different species, depending upon the mating and breed-
ing habits of any particular species. The sexual proportion may
be considered as one of the physiological adaptations of a species.

The conclusions drawn from statistical data should be tested
experimentally. There are two experimental points of attack:
(1) Breeding experiments to determine whether the sexual pro-
portion can be altered by selection. (2) Experiments on the
separate blastomeres from one ovum to determine whether two
embryos of different sexes can be reared from the same egg, and
whether there is a period of sexual indifference in the development
of an individual.

320

THE AMERICAN NATURALIST [Vol. XL!

BIBLIOGRAPHY.

Only a partial bibliography is given here. Cuenot and Busing ('84>
give all the literature for animals up to 1899. Strasburger and Gregory
give the botanical literature. A less complete review is given by Loeb
(:06).

Bateson, W. and Saunders, E. R.

:02. Experimental studies in the physiology of heredity. Reports
to the Evolution Committee, No. 1, London.
Born, G.

'81. Experimentelle Untersuchungen iiber die Entstehung der Ge-
schlechtsunterschiede. Breslauer artzUche Zeitschrift, 3, p. 25.
Castle, W. E.

:03. The Heredity of Sex. Bull. Mus. Comp. ZooL, 40, pp. 189-218.
Castle, W. E., Carpenter, F. W., Clark, A. H., Mast, S. 0., and Bar-
rows, W. M.

:06. The effects of Inbreeding, Cross-breeding and Selection upon,
the Fertility and Variability of Drosophila. Proc. Amer. Acad.
Arts and Sciences, 41, pp. 729-786.

Cu^NOT, L.

'99. Sur la determination du sexe chez les animaux. Bull, scientif.
de la France et de la Belgique, 32, pp. 462-535.
DtisiNO, C.

'84. Die ReguHerung des Geschlechtsverhaltnisses bei der Vermehrung
der Menschen, Thieren und Pflanzen. Jenaische Zeitsoh. f.
Naturwissenschaft. 17, p. 593.
DusiNG, C.

'85. Der Experimentelle Prufung der Theorie von der ReguHerung
der Geschlechtsverhaltnisses. Idem, 19, supp. Heft 2, p. 108.
Geddes, Patrick, and Thompson, J. Arthur.

: 01. The Evolution of Sex, Chapter IV, 2d. ed. London.
Gregory, R. P.

:04. Some observations on the determination of sex in plants. Proc.
Cambridge Phil. Soc, 12, pp. 430-440.
Griesheim, a.

'81. Ueber die Zahlenverhaltnisse der Geschlechter bei Rana fusca.
Arch. f. d. ges. Physiol., 26, p. 237.
Harper, R. V.

:07. Sex Determining Factors in Plants. Science, N. S. 25, pp. 379-

:06. Geschlechtsbestimmende Ursachen bei den Daphniden. Arch-

No. 485]

THE DETERMINATION OF SEX

321

Ceboreiien und der Gestorbenei
iwissenschaften, 3, p. 816: new editic
1900, 4, p. 177, cited by Strasburger (:00).

:06. The dynamics of living matter. N. Y. and London. Chapte
IX and X.
McClung, C. E.

:02. The accessory chromosome — sex determinant? Biol. Bui
3, pp. 43-84.
VON Malsen, H.

influsse und Eibildung bei Dinophilv

number and arrangement of the male
the proportion of the sexes in the Nor-
ibridge Philos. Soc, 12, pp. 441-444.

Pfllger, E.

'81. Einige Beobachtungen zur Frage uber die das geschlechta-
bestimmendon Ursachen. Arch. f. d. ges. Physiol. 26, p. 243.

'82a. Hat die Concentration des Samens einer Einfluss auf das Gesch-
lecht? Idem, 29, p. 1. (Has no effect.)

'82b. IJebor die das gi^schlcchtsbestimmende Ursachen und die ges-
chlecht.sverhaltnisse der Frosche. Idem, 29, p. 13.
PUXXETT, R. C.

:04a. On nutrition and sex determination in man. Proc. Cambridge

Phil. Soc, 12, pp. 262-276.
:04b. Note on the proportion of the sexes in Carcinus mainas. Idem,
pp. 293-296.
Rauber, a.

:00. Der Ueberschuss an Knabengeburten und seine biologische
Bedeutung. Leipzig. (Abstract by R. F. Fruchs in Biol. Cen-
tralbl., 21, p. 833, 1901).

:02, Poverty, a study of town life. Second edition.
Roux, W.

'86. Ueber die Bestimmung der Hauptrichtungen des Frosch-embryos
im Ei, und uber die erste Teilung des Froscheies. Bresla'uer
iirtzliche Zeitsch. [The literature is given by Wilson (1900)1.
Strasburger, E.

:00. Versuche mit diocischen Pfianzen in Rucksicht auf Geschlechts-

322

THE AMERICAN NATURALIST [Vol. XLI

verteilung; Biol. Centralbl., 20, pp. 657-65, 689-98, 721-31,

753-85.
Weldon, W. F. R.

:01-:02. Mendel's Laws of Alternative Inheritance in Peas. Biome-

trika, 1, pp. 228-254.
Wilder, H. H.

:04. Duplicate Twins and Double Monsters. Amer. Journ. of Anat.,
3, p. 387.
Wilson, E. B.

:00. The Cell in Development and Inheritance. 2nd ed. New York,
p. 408 et seq.

:07. Sex determination in relation to fertilization and parthenogenesis.
Science. N. fe. 25, pp. 376-379.
Yung, Emile.

'83. Contributions a I'histoire de I'influence des milieu physico-

chimiques sur les Hres vivants. Arch, de Zool. experimental,

2e serie, 1, p. 31.
'85. Influence des variations du milieu physico-chimiques sur le

d^veloppement des animaux. Arch, des Sciences Phys. et Natur-

elles, 14, p. 502.

CHUBS' NESTS

ALFRED W. G. WILSON

of the numerous small streams tribiitarv lo tlic rpper ( )ua\va
River, the passing voyageur cannot fail in liavini^ liis attnition
drawn to curious conical piles of coarse gravel and j)el)ble.s which
occur along the river shores. Locally these piles of stones are
called "Chubs' Nests." The following notes are published in
the hope that they may prove of interest to American Naturalists.

The accompanying plates will give a general idea f)f the shape
and character of these heaps of atones. They are ci.nical in form,
with a circular or oval base. The vnhinie of the travel of which

they are built will var\^ from a g I sized wlieelharrow load to

about a cart load. The individual pebbles vary in size; the great
majority would readily pass through a two inch ring. In a few
cases oblong pieces of schist al)out three inches in length were
noted but their cross section would not be more than one square
inch. The largest pebbles used in the construction of the lieaps
would weigh at least half a pound each; mo>t of the pebbles would
weigh less than four ounces each. Th(> rock material from which
the pebbles have been derived is often (juite different from the
rock of the immediate vicinity, showing that the pebbles have been
transported some distance to their present resting place. In a
number of instances it was foimd that the interior of the heap
consists almost entirely of small pebbles less than an inch in the
maximum dimension, the larger ones forming only an oiuer laver
over the whole cone.

The dim<>nsions of two of these heaps of stones were as follows: —
Xo. I. Ha.e, h'ligth (;..-) feet, width o feet at one end (left of figure
1 and 1 fet>t at the other, height 21 in. Ik-, angular slope of the
side of the cone about 4S° t.. the vrli. al. This pile was built of
mixed pebbles, chiefly granite and mIum. See figtire l.i

No. 2. Base, nearly circular and foiu' t'eet in diameter, height
22 inches, angular slope 49° 4.V. ( Sei> figu re 2. )

323

324

THE AMERICAN NATURALIST [Vol. XLI

In a very large number of cases examined the stones were found
to be piled quite loosely so that the slightest jar set them sliding
down into a position of more stable equlibrium.

Along the larger streams and rivers the heaps are usually found
in small bays off the main stream or on bars and ridges on the
sides of the main channel, in quiet but never in dead water. In
some places near the watersheds they occur in midstream, and
occasionally they are sufficiently numerous to hinder and partly
obstruct canoe navigation, where the water is shallow and th^

Fig. 1.

In the early spring when the waters are high and usually more
or less turbid the cones are not in sight; but as the waters recede
the apices of the cones gradually appear above the surface and
late in the season the water may have receded so that the whole
cone together with the bar on which it was built comes into view.
The tops of the "nests" shown in figures 2 and 3 were fully five
feet above the surface of the water when the pictures were taken
in August. In early June the water was probably six feet higher
and the tops would have been under at least a foot of water.

As to the origin of these curious heaps of pebbles, the Indians

No. 485]

325

and Bushmen all attribute them to small fish — called Chub by
the whites and Awadosi (stone carriers) by the Indians.^ Per-
sonally I have made numerous inquiries but I have not been able
to find any one who will say he has actually seen the fish at work,
still they all insist that it is the fish who make them. A careful
examination of over one hundred heaps, scattered along a line of
gravel more than three hundred miles in length has convinced the
writer that the cones are of animal origin, that the materials have
been assembled by some intelligent agent, not by stream action.

These heaps of stones are said to be built in the early spring
and are presumably used for spawning purposes. They are
always in places where the water is smooth but still flowing.
Except in the very beds of the rivers of this north country, pebble
and gravel beds and bars are not found. The shores of the
streams are almost universally clay. At liiuh water tlic rivers
expand and invade the woods so tliat. as one ot' iny canoemen
expressed it "The pike go into the l)nsli to Innit."' In midstream
the water is usually flowing veiy swiftly at higli water and along

' Bell. Robert. Recent Explorations to the South of Hudson Bay. The
Geographical Journal, July, 1897. p. 16.

326

THE AMERICAN NATURALIST [Vol. XLI

the shores the ground is covered with logs and bushes. Ground
suitable for spawning covered by a moderate depth of water is
rare. In nearly every case where the nests were seen the bottom
consisted either of large boulders and cobbles, or of soft materials
and sand with a certain admixture of partly decayed logs and
lower types of plant life, chiefly algse. On this bottom the conical
heaps of stones were built up. It seems not unnatural to suppose
that they serve the dual purpose of offering a clean gravel surface
for the deposition of the eggs, and at the same time raise these
eggs nearer the surface of the water and thus into a zone of more

light and warmth tlian if they wcrv dcpositc'd dirc(;tly upon the
bottom.

The fish which are said to be the arcliitccts of curious
nests vary in size up to about 18 inches in Iciiuih. .iiid in weight
up to about two pounds or a Httle over, 'i'hcir vcntial nspoct is
white, the dorsal dark gr;iy-])lack, and the broad sides me silver
white. The cycloid scales mv li.ruv and dii.-k, ;iiid di<- t.ody is
about three times as dccj) wide. The tliinl pliitr ^hows a
specimen about 14 iiichrs in Icnuih which was ca])inrcd and laid
upon the nest before making the j)i('turc. rresident David S.
Jordan to whom these data have been submitted considers that

CHUBS' NESTS

327

the fish is "prol)ably the Silver Chub or Fall Fish, Snnnnfilu.s-
corporalis Mitchell." It may be interesting to sportsnuMi to know
that the fish rise readily to the fly, occasionally can be cau<;ht witli
a troll, and are easily captured with an ordinary hook baited with a
piece of bacon rind. The flesh is coarse and the bones are few
and large, reminding one of mullet.

In 1844 Chubs' nests were found in the Magalloway River,
Maine, by Dr. Jeffries Wyman. He described them to the Boston
Society of Natural History (Proceedings, Vol. 1, p. 196) as " mounds
of pebbles, two or three feet in diameter, which he was told were
heaped up by a fish called the Chub, at its breeding season, and
that its eggs were deposited among the stones." He referred to a
similar habit attributed to the lamprey eel and remarked that he
was not aware of any other instance of the kind.* Dr. Robert
Bell, in the report of his explorations referred to above, has pub-
lished a figure of a characteristic nest. He states that a varying
number of chubs work together in building a mound, bringing
the stones in ilicir months, one at a time, from far and near.

In consi.lciinu- the relative sizes of the pebbles and the fish that
move them, it nnist be remembered that under water the weight
of the stones will be from one quarter to one third less than the
weight in air. In the cases of the larger heaps of stones it is often
found that there is an area greater than the base of the eone over
which the stones are scattered. In one case we found what ap-
peared to be the base of an old cone and the inference seems to be
that in the rebuilding every spring they repair the old nests, shift
them at times, and utilize materials from abandoned nests to
construct new ones or to enlarge the old. The larger nests are
probably the work of several seasons.
Montreal, January, 1907

1 The nests of the lamprey are " gravel filled pockets. " "The central part
is usually 15 to 20 cms. deeper than the edges, so that the whole is dish-Uke
in appearance; at the lower edge there i.s always a pile of stones." The
stone carrying habit of the lamprey has been described by S. H. Gage, by
Dean and Sumner, and by Young and Cole {Anm-ican Naturalist, 1900, vol.

care^among fresh water fishes (Rep^of the Smiihsonian Inst., 1905, pp. 402-
531) Theodore Gill does not include either the lamprey or the chub, the

NOTES AND LITERATURE

GENERAL BIOLOGY

MendeUsm.* — In a well printed booklet of eighty-five 4 X 5^ inch
pages, R. C. Punnett of Cambridge, England, has presented an ad-
mirably clear and concise account of Mendelism. After reviewing
the simple and fundamental experiments of the Abbot of Brunn, the
writer describes the more recent discoveries to which they have led,
and in conclusion shows them to be of the highest practical and
scientific importance. Although the mendelian principles of heredity
are well known in America through the publications of Castle, Daven-
port, and others, a brief review of them as presented by Punnett may
still be of interest.

It is found by experiment that when a certain pure bred tall variety
of plant is crossed vrith a dwarf, the resulting hybrid contains both
the factors for tallness and shortness. If A represents the tall factor
of one parent and a the short factor of the other the hybrid which
contains both is Aa. It is not of medium height, but is like its tall
parent. A character such as tallness in ])eas which is retained by
the hybrid is called dominant: one like dwart'iu ss which is latent in
the hybrid is named recessive.

When hybrids Aa are bred together, they pnxhicf in the next gen-
eration 25% of pure tall forms, AA; 50* , of tall hyl)ri(ls. Aa; and
25% of dwarfs, aa. The famiHar fornuila may he written thus:

1st generation Aa
2nd generation AA 2 Aa aa

The tall hybrids Aa, and the pure tall plants A A, are indistinguish-
able except by further breeding. Then it appears that one in every
three contains only the factors for tallness. Such plants, like the
dwarfs, breed as true as if derived from an unbroken ancestry of pure

It is not al\vay> the caM" that the hybrid resembles one of its parents.

•Punnett. H, ('. M>n.i.li.^m. Second Edition. Cambridge, MacMillan
and Bowes. 1907. IGnio. vii + 85 pp.

329

THE AMERICAN NATURALIST [Vol. XLI

The blue Andalusian fowl is a race which in breeding produces 25%
of black offspring, 50% of blue, and 25% of white splashed ^\ith
black. It is evident from these proportions that the blue race desired
by fanciers is essentially mongrel, and can never be made to breed
true. The black race and the splashed whites remain true when each
is mated with its own kind, but when crossed they produce the blue
Andalusian.

The fixed proportion of pure and mongrel forms in the offspring
of hybrids may be readily explained. The factors A and a, derived
by the hybrid from its parents respectively, are transmitted through
its germ cells in equal abundance. The factors become segregated,
so that one half of the germ cells contains only A, and the other half
a. In the process of fertilization an A will unite with a as often as
Avith A ; and an a will join A as often as a. Thus there will be 2
Aa for each aa and AA.

When two different inheritable factors occur in each parent the
number of combinations in the offspring is much greater. Mendel
found, with peas, that the height of the plant (tall or dw^arf) and the
color of the seeds (green or yellow) were transmitted independently
of one another. A and B may represent respectively the factors for
tallness and greenness which are dominant; a and b the factors for
shortness and yellowness which are recessive. If a tall green-seeded
plant AB, is crossed with a dwarf yellow ab, tall green-seeded hybrids
containing the factors AaBb result. Every germ cell of such a hybrid
contains one factor for height and one for color; tlu^y are e(|ually
distributed in the four possible combinations AB, Ah, ciB, and ab.
When such a group of germ cells fertihzes a .similar (jjioup, the follow-
ing combinations are to be expected:

AABB AAbb aaBB aabb

2AA Bb 2Aa bb 2aa Bb

2AaBB

4AaBb

Thus among sixteen individuals nine contain both dominant factors
and in the case of the peas are tall green-seeded plants. Three con-
tain only the dominant A, and are tall yellow-seeded forms; three
contain only the dominant B and are green-seedcnl dwarfs. ( )nc
contains neither dominant and is a yellow-seeded dwarf. Tlii. ratio,
9:3:3:1 Mendel verified by experiment.

The sweet pea known as the 'Painted Lady' has a bright pink color
due to its sap, and this is dominant over the absence of such sap color

No. 485]

NOTES AXD UrEUArVRK

331

in which case the flcwcr is white. In ilic civmn sum p.., ili<'
sap color, the tint hc\\\\i (hie to pitriiicnicd clirniiKii.hiM-. The

A similar roiill has h.vn nh.rrvr.l in l.rr.Mlin- lnui>. 'rhr ruM'
comb of tlic W yan.lollr ivpc ami the pea (•(.nil) of llic Indian -anir
are both .Innnnant ovrr ihr sinuir ,-unil. of the [..■u-liorn type. WIum,
a rose cnil. i> < ro.M.,l ^^\^\^ a p« a conil. a new typ,' n >ull>. .Ics. rihe.l
as the " wainni - r.unl.. It ivM inMcs that of tlie Malav bnr,l. When
such hybrids aiv bn .l toovtlirr fnnr types of comb appear in the next

of 9:3:3:1.

ism. When a gravis cros.sed witli an albino, -ray hybrid^ iv.uh uhiel,
produce young in the proportion of ^nays, ;; blaek>, and 1 white>.
The factors involved are ]>i<rinentation. .1. dominant over albinism,
a; and grayness, />'. d.nninani over blackness. The four white
animals which appear identical include iln-ee forms, namely aa BB,
2 aa Bb, and aa hh, all of which lack the pij,nnetitati()n factor >4, From
this it appears that the wild i,nay color consists of a factor for pigmen-
tation and another for irniyne^. Uy loss of the former a white rabbit
results, and by loss of the latter, a l)lack one. In the offspring of
such a white and l)lack, r< n rsimi occurs to the original gray form.
Similarly white pea l)lo><oin^ may .^n h contain one of the two factors
for pigmentation, and by cro»in<r snch whites, reversion to a wild
colored type has been obser\cd. Tims reversion ha> been dcline(]

in the course of jihylo^'enetic development.

From these and many other observations, the author concludes
that no hoiticnlinrali>t <an j)ropose to rai.se a tall pea from a dwarf
bv a proce-,^ of mamirinir, nor by selecting minute fluctuations, but
only by obtainin.i: new airirregations of unit <'haracters throng], br.-ed-

' In Mr. Pulinett''s'book no refenMue i. ma.lc to 'mixed inheritan.e'
whereby the parental chara< icrs arc blended in the offspring. It nm-t
be remembered, however, that ral)bii.-> with ears of medium li'iigth are

332

THE AMERICAN NATURALIST [Vol. XLI

crossing of some tall and dwarf plants produces those of intermediate
height. The omission of such limitations may cause a student to
believe that Mendelism is the universal law of inheritance.

F. T. L.

The Problem of Age, Growth, and Death.— In a series of six
public lectures Professor Charles S. Minot has made known the results
of his studies, now in progress, concerning the essential nature of
senescence. Rejecting such criteria of old age as a halting gait or
arterio-sclerosis, which pertain chiefly to man, he has sought those
features which apply as well to the aged frog or fish, and even to still
lower forms. Such characteristics are found in the decreasing rate
of cell division, the increase of protoplasm at the expense of the
nucleus, and the progressive differentiation of the protoplasm. Old
age is therefore essentially a cytomorphic phase.

The rate of cell division is expressed by the "mitotic index" which
is the average number of mitotic figures found, in sections, among a
thousand nuclei. The mitotic index falls from 18 to 13 in rabbit
embryos of 7^ and 13 days respectively. Drawings, on the same scale,
of nuclei of the various tissues in rabbit embryos of 7^ to 16^ days
show a striking reduction in the actual size of the nucleus, except in
the case of the nervous tissue. Even there, in relation to the proto-
plasmic mass, the nucleus may be relatively small.

The rate of growth begins to decline before birth, and this rate of
decline rapidly decreases until old age, when growth is at its niiiiimuni.

held to apply to man, both in physical development as shown by
statistics of weight, and in mental development as determined by
psychologists. During the first months after birth, progress in
acquiring concepts of time, space, the ego, and the external world is
more rapid than in later years. As with weight, the rate of decline
is most abrupt at the outset, becoming gradual as age advances.

The study leads to the paradoxical conclusion that the changes of
senescence are most marked in the years of infancy, for the popular
idea of maximum efficiency as the mark of maturity is set aside.
The embryo in adding an ounce to its wcioht is rated as advancing
more rapidly than the child in gaining a poumi ; the inx . t which leaps
many times its own length would be regarded as more successful in
jumping than the mammal which can far outdistance it.

No. 485] NOTES AND LITERATURE

333

Because of the rapid early decline in the rate of development
Professor Minot believes that the age of college entrance should be
lowered, and that professional studies should be entered upon at a
younger age. A final publication of these researches, which have
extended through many years, is in preparation.

F, T. L.

The Hypothesis of Mimicry.— Dr. Franz Werner of the University
of Vienna is a skeptical critic of the Mimikrylehre which he regards
as due to a rather crude anthropomorphic point of view (Biol. Cen-
tralhl, 27, pp. 174-185). He considers first the non-poisonous
snakes which are supposed to have acquired a protective resem-
blance to the poisonous forms of other genera inhabiting the same
locality. Since no snake-eating animal is known which makes a dis-
tinction between poisonous and non-poisonous forms, the latter can-
not be protected by the similarity of pattern. ^Moreover in some cases
there is reason to believe that the non-poisonous snake is the older
type: and that the venomous Elaps or Vipcra is the "imitator." The
stingless insects which deceptively resemble bees and wasps fare no
better than the harmless snakes, for stinging forms arc "not in tin-
least protected from their natural enemies; they fall a prey to many
birds as well as to lizards, frogs, toads, and spiders." Finn is cited
in evidence that the poisonous Danais is as eagerly devoured in India
by lizards of the genus Calotes as are its mimics. Poisonous forms
which often exhibit bright warning colors "to signaUze their unpalata-
bility to enemies in good season" are not secure.

The similar patterns and colors of various snakes in a given locality
may be due to similar climatic conditions and food supply, the pig-
ments involved being physiological by-products. Color photography
is invoked to account for the correspondence in color l)(>twecn an
animal and its habitat. A physiolopcal rather than a tclcoloirical
explanation is desired. In otiicr words, it is believed that >iiiiihir
causes produce both the forms wlucli nunnc and thoM' which are
imitated, and that there is no other rehition hetwcvn ih(> two. Krom
the reports of field observers the number of in-tanco of eti'ciivc
mimicry has been so reduced that "as good as nothin>: remains.'"
Dr. Werner believes that man alone has been seriously deceived.

334

THE AMERICAN NATURALIST

[Vol. XLI

ZOOLOGY

Palms and Soles. — Dr. Schlaginhaufen of Dresden has written a
brief description of the palms and soles of man and the apes, based
upon a hterature of one hundred and fourteen publications.^ The
volar surface of the hand and fingers, and the plantar surface of the
foot and toes are thickly covered with slender ridges, the cristae cutis,
separated from one another by depressions, the sulci cutis. Along
the summit of a ridge, a row of SAveat glands opens. A primitive stage
in the formation of the ridges is seen in the Prosimiae, which have
small round elevations (insulae primariae) surrounding single sweat
pores. Besides these primary islands there are larger elliptical forms
on which several sweat pores may be arranged in a circle or ellipse,
surrounding a central depression. These lenticular islands are due
to the coalescence of primary islands radially arranged. A crista
is formed by the coalescence of a linear series. The minutiae of the
cristae, upon which personal identification depends, consist in the
branches of the ridges, which may end blindly or anastomose; in
detached ridges; and in the ridge patterns. The two principal
patterns are the more or less concentric tactile figures, and tli(^ Y shaped
groups called triradii.

Besides the bas-relief of cristae, palms and soles pn sc nt the liiiiii
relief of tactile cushions, toruli tactiles. For each extrtMuity tlifit- are
typically five digital cushions at the tips of the firigers or toes; four
interdigitnl riis/n'ons near the metacarpo- or metatarso-phalangeal
joints; and two or tlin r proximal cushions, — a tibial and an elongated
fibular, or a radial and two ulnar, one behind the other. This ar-
rangement is typical for pentadactylous mammals and the cushions
are well developed in marsupials, rodents, the insectivora and pri-
mates. Often the interdigital cushions fuse, as in the cat, and that
between the thumb and fingers may be suppressed. Secondary
cushions are not infrequent — such as a central cushion found in
Cebus — but none occur in the anthropoid apes or in man. Cushions
are accumulations of connective tissue and are not to be confounded

» Schlaginhaufen, O. t)ber das Lri.tcnivliri .l. r lIolilha.Ml- un.i I-u^^^.>hl('n-
Flache der Halbaffen, Affen und Mcnx hctira-M n. /• /,/,/-// ,/. A, ml. >,. luitir..
vol. 15, pp. 628-662. Since writing this n-vicw, thv editor luts n-<riv..d the
announcement of the following book. Kidd, \N'. Th srn.sr of touch in mam-
mals and birds wUh special reference to the papillary ridtjes. London, A. and
C. Black, 1907. 8vo., 174 figs. 5s.

336

THE AMERICAN NATURALIST

[Vol. XLl

Otohime, a new genus of gurnards (0. hemisticta) is described in
the same proceedings by Jordan and Starks, from Japan.

In the same Proceedings, Professor John O. Snyder gives a review
of the Mullidse or Surmullets of Japan.

In the Bulletin of the Bureau of Fisheries, vol. XXV, 1905 (issued
1906), are several important papers on the fish-fauna of our island
possessions.

The "Fishes of Samoa" by Jordan and Scale contains a list of the
species collected on the American island of Tutuila and the German
island of Upolu by David Starr Jordan and Vernon Lyman Kellogg
in 1902. About 500 species were obtained, 92 of them new to science.
Most of the latter are small fishes taken through the use of poison
(chloride of lime) in the pools of the reefs. The reef fauna of the
islands of Samoa is remarkable for the number of brilliantly colored
species. In this paper are twenty-six colored plates of the most strik-
ingly colored of these small reef-fishes, noted since the days of Captain
Cook. These plates are from water color sketches by Kako Morita.
The origin and purpose of these brilliant hues of coral-reef fishes is
one of the most difficult problems in evolution. It is to be noted that
these colors are not confined to any one family, but that more than a
dozen families of fishes participate in them.

With this paper is a check list of all the species, 1704 in number,
now known from the region called Oceania, which includes Hawaii,
Polynesia, Micronesia, and Melanesia. In all this region the fauna
is essentially continuous, except as regards Hawaii. In this separated
island group, the genera remain the same as in Polynesia, but the
species as a rule are difl'erent. This difference is clearly due to the
operation of isolation and segregation.

In the same paper is a valuable discussion of the Samoan names of
fishes, and tlu> root-words ctmiposing them, by Mr. W. E. Safford.

Aliii()>r siiiiiili.iiicniis with tills paper, but apparently with a few
weeks priority, is a iiieinoir "Zur Fischfauna der Samoa-Inseln" by
Dr. Franz Steindachiier, in the "Shzungsberichte der Kaiserliche
Akademie" (1906) in Vienna. Dr. Steindachner describes the spe-
cies of fishes collected at Apia in Upolu, by Dr. Rechinger. This
collection contains 120 species, of which 20 are new. Only one of
the new species is contained in the series described by Jordan and
Seaie. This is .SV//ar/V/.v rrrhlmjrri Steindachner. calle.l Snlarlas
garmani by Jordan and^S.v.lc.^ A ncvv /vm///r,-m, is added

In the next volume of the Bulletin of the Bureau of Fisheries,

No. 485] NOTES AND LITERATURE

337

(XXVI), Jordan and Seale discuss the "Fishes of the Islands of Luzon
and Panay" as represented in a collection made in 1900 by Dr. George
A. Lung, Surgeon in the United States Navy. Dr. [.iiiig ()l)taincd
at Manila and Iloilo, 249 s])ccic>.. <,f ^^h\rU ci-ht.v.i :nv .I.^m tIIxmI ms
new. One of these species, Uhitint/ohins /iiik// proves idcMiiicil wiih
an older species Rhhwcjohiu.s u,hnln.„s (F.u-skal), and P, trusrirl, .s^

'VUv saiiK- ri. h faniK. is discnssc.l in a similar paper which imnie-
dial.-ly lollous il.c ..thcr In the same Hnlletin, "Fishes of the Philippine
Islands" In Kvennann and Scale. This treats of the collection exhib-

by Mr. Charles J. Pierson, formerly of Stanford University. In this

One of these, Platophrijft palad, should have been referred to the
genus Pscudorhombus of Bleeker, of which the Americ-an genus
Cencylopsetta seems to be a synonym. Tliree other species of Pi^cu-
dorhombus are by some slip of ilie pen referred to Plniophrys. It

It should lead to a general descriptive cataloirnc of iIk- vast iisli-iauna
of the Au.stralian contin(Mit.

In the Bulletin of the Museum ..f ( 'omparativ /u.,l.>iry. vol. L,
1906, Dr. Charles K. KaMman dcM ril..- mnnrruu. ^hark'. t.vth and

THE AMERICAN NATURALIST

[Vol. XLI

Lahille describes a remarkable new genus of maekerel-like- fishes
from Argentina under the name of Chocnogastcr holmhenji. The
dorsal and anal fins are provided with finlets; llie mouth is very large,
and the body is covered with large scales. An alHed fish is described
by Dr. Lahille from Port L^'ttelton, New Zealand, under the name
of Lepidothynnus huttoni. Both of these are regarded, probably
correctly, as related to Gasterochisma mdampuft of New Zealand.
Figures of all three of these species are given by Lahille.

In Volume III, of IMarine Investigations of South Africa (1905),
Dr. J. D. F. Gilchrist, Government Biologist of Cape Colony, describes
seventeen new species of fishes found in rather deep water off the
Cape of Good Hope. Several of these are most interesting additions
to our knowledge of fish-forms.

The Biennial Report of the State Board of Fisli Cennnissioners
of California for lOOC. contains u.eful account. ..f ilu- in.ui of Cali-
fornia, those of the Sierras bv Dr. B. W. K\vnn;irni, tlic others by
Dr. D. S. Jordan. The report is edited by ( liailc- A. N'o-clsaiig.

One of the most valuable monographs of ;i siiiiilc tvpc of fishes is
the magnificent paper entitled, "Chimteroid V\>\u's .iiid Thrir Devel-
opment," by Bashford Dean, published by the Cnriicixic Institution.

This paper treats especially of the anatomy and dcx .1. .pnicnt of
the Cahfomia Chinncra called Hal-fisli or I'lli pliant-fisli, Cliiniwra
coUifi, as studied in tlu- Hopkin^ -ca-nlr I.alioraton at I'acilic Grove
in California. The- paper eontains a r.'.-ord of tlie oilier living and

a liiolily ni..dified and .pe. ia li/e< I ..iWiooI from llie o-n.np of primitive

Dr. (;. \. Bouiengcr continnes 'lii^ paj.ers .m tli<' tivsli water fishes
of Africa with a memoir on the fislie> of Lake Tan,i:an\ ika.

In the Proceedings of the l\oyn\ Aca.leiny ..f Amsterdam, Professor

No. 485] NOTES AND LITERATURE

339

Mazatlan, is the same as Eleotris picta from farther south. Dorm itafor
latifrons, of the Pacific slope, is regarded as different from Dormitator
maculaius of the Alhuitic. The name, Chmmphnrus banana, is used
mstead ot ilic doiilii tiilly idciil ifiable C/ionophorus (or Awaous)

dium inultipinirlalum is a new s[)eeies from Oaxaca. Excellent
figures are given of many of the species. In the Annals and Maga-
zine of Natural History, XVIII, 1906, Mr. llegan has numerous
papers on fishes. In "Descriptions of Some New Sharks in the
British Museum Collection," the Japanese Orectolobus is separated
from O. barbatus, as Orectolobus japonicus, and the Japanese Monk-
fish as Squatina nebidosa. This had, however, been earlier named
Squatina japonica by Bleeker. In another paper in the Proc. Zool.
Soc. London for V.){)(\, Mr. Regan discusses the classification of the
sharks and ray^. pmposini: a new classification.

The t'ollnv.ini: is KcKuirs arrangement of the families;
Subclass Sriarhr,,

Series 1. Trcmafopnca,

Order 1. Pleuroptcrijcfii (extinct)
Families, Cladoselach idw.

Order 2. Acnnthodli (extinct)

Orcctolobida;,
Sri/lliorliinidce,

340

THE AMERICAN NATURALIST [Vol. XLI

(HeterodontidcB)
SqualidoB,

\ SquatinidcB.

Suborder 2, Hypotrema
Di\dsion, Narcobatoidei
Family, Torpedinidce
Division, Batoidei

Families, Rhinobatidce,
Raiidce,
Dasybatidce.
vSeries II. Chasmatopnea,
Order, Ilolocephali,
Family, Pyctodontidce (extinct)
Squaloraiidce (extinct)
Myriacanthidoe (extinct)
Chimccridae

The principal feature of this arrangement is the grouping together
of the Cestraciont and Squaloid sharks as a division corresponding
to the Galeoidea. Except for the reduction of some families to a
lower rank, and a few changes in names of groups, this corresponds
fairly with that adopted by recent American writers.

Under Diagnoses of New Central American Fresh Water Fishes,
Mr. Regan describes Rivulus flabellicauda, from Costa Rica, Rivulus
godmanm, from Guatemala, Pacilia salvatoris, from San Salvador,
Xiphophorus strigatus, from Vera Cruz and Oaxaca, X. brevis, from
Honduras, Agonostomus macracanihus and A. snlvini, from Guate-

In the Anatomischer Anzeiger, Dr. l ine I )aliltj;reii describes the
anatomy of the electric organs on the top of the head in the Electric
stargazer, Astroscopiis y-gra;cum.

These very interesting organs constitute a new type of electric
organs, quite different from those of the torpedoes and other electric

In the Proccpdi.ics cf the Aca.lcmy of Natural Sciences of Phila-
delphia, Henry \V. Fnu Irr .h-. nl..-s ( •r>,trupnw„s <,ahln as a new species
from San Donii.i-.,, ami ('. hrrim/l, fror.i Siirina.n. He gives a list
of the cold-blooded vcrtcl.ral.^s ohlainr.! about tlx- Florida Keys.
Eighty-six species of fislic^ arc ivi-ordcd, one of tliciu n-o-arded as
new. This is Con^m ?///// //.V inonm'. w hii li sccius to 1 Ii(m\ riicr a s|)c-
cies of Darfyln.tropii.s, not evidently dilfcrrut from / )acti/l(,.sr(>pii.s' tri-
digifatvs, found by him at Key W(>st.

No. 485] NOTES AND LITERATURE

341

In a paper on "Rare or little known Scombroids, No. Mr,
Fowler proposes the new subgenus Pampanoa ior 1 raclnnofiis (//a urns,

Mr. l-owlcr 111 I his jiiid oilier papers adopts the generic names of

'riie.se names, ni liis view, become available, because Walbaum in 1792
reprinted them all with their diagnoses, although not adopting them
or in any way reinforcing them. In the judgment of most writers,
a name published before Ivinnanis does not accpiire validitv bv a reprint

should be made.

sosteus or Lepidosteus, Brauia will replace Ahrami.s, and (>lanni.s
Lichia.

A review of various genera of Soutli Amencan Characins is given
bv Mr. Fowler, as also series of useful notes on fishes of Pennsylvania.

In the Bulletin of George ^\ ashington Ijniversity, vol. I, 19()6, Dr.
Theodore Gill tells "the remarkable story of a Greek fish, the Glanis,"
{Parasdums arhstotelis). This species was known to Anstotle, but
modern authors have, with a few exceptions, overlooked its existence.

In the Smithsonian Report toi !<)()) l)r Theodou Gill giM^ an
interesting review ot our knowliM !<:■(■ ot Parental ( are amoiii: 1-resh
Water Fishes." The literature of ihi. Mil.jeei i> fiillv di^eusM'.l.

In the Zoologischer Anzeiirer, Hr. L. l?eii:- (li>( ii»e> ilie fish, -,
of Lake Baikal and those of the Aniiir Uamu. He eon.iders Cnth,-

are also listed by the same author.

In the Bulletin of the Vcadunu ImptiiaU (U^ Niiiuis, 1. Btig
di.scusses the lampreys of the Ru.ssian Empire.

The species of Lampeira or nver lam]m^v in this vast region he
reduces to two, L. fluviatiUs and L. planen. To the former he refers
Imnpihn a,n,<t i>\ \la.ka, / jnpnn i< a iA } m I unuhhahntni

ind / ot th. ] iMMii I nil.<l Nit.s I In 1 ..I d. lu iiimmIh.ii

In the Pro.-, /ool. ,N,e. Lond.>n, Prof. W. H. Penliaii, and \V. .1.

342

THE AMERICAN NATURALIST

[Vol. XLr

In the 24th Annual Report of the Fishery Board for Scotland, Dr.
H. C. Williamson describes the small cod-fish, Gadus mimiius and
Gadus esmarki, and records two cases of hermaphroditism in the
common cod-fish.

The fourth part of the Fishes of Japan by Otaki, Fujita and Higur-
ashi appears with descriptions in English and Japanese and with
excellent colored figures of the common 'Tai,' the "national fish" of
Japan, {Pagrus major), oi the Ayu {Plecoglosmus altivelis), next to-
the American Eulachon, the finest of all food-fishes, and other spe-

In the Zoological Series of the Field Columbian Museum, Dr. T.
H. Bean publishes a catalogue of the Fishes of Bermuda. 261 species
are recorded, many of the more rare forms being figured. The new
species, previously described in the Proceedings of the Biological
Society of Washington, vol. XIX, for 1906, are the following: Hippo-
campus hrunneus, ( = H. hudsonius Jordan & Evermann, not of
DeKay), Holocentrus meeki, Eupomacentrus chrysus, Iridio decoratus,
Iridio meyeri, Iridio microstomus, Cryptotomus crassiceps, Mona-
canthus tuckeri, Rhinogobius mowbrayi, Lahrisomus lentiginosus,

In the series of Occasional Papers of the Bcrnicc Pauahi ^riiscuni
at Honolulu Alvin Scale gives a list of "Fislu-s of the South Pncific"
collected by him in the Marquesas, Tahiti. Solonuin Islaiids, and
elsewhere in the South Seas. Numerous ww s|K ( i( s arc dcsc rilxMl,
and illustrated in not very satisfactory fashion ])v j)h()t(>(rraphs

In the same series, William A. Bryan describes a few new or rare
fishes from Honolulu.

In the Records of the Australian Museum, VI, 1906, Edgar R.
Waite gives descriptions of Australian and Tasmanian fishes, and
studies in Australian Sharks, with photographs of the egg cases of

In the Proceedings of the Biological Society of Washington, Hugh
'SI. Smith and Alvin Scale describe a number of species from the

III tlic Hiillctiii of the Michigan Fish Commission, No. 8, Mr. Ellis
L. Micliacl (•atal..<^s the fislics of :\Iichigan, with rcfcretur to all

Nettling Hairs of the Brown-tail Moth.' It is well known that

1 Tyzzer, E. E. The pathology of the brown-tail moth dennatitis. Journ..
of Med. Res., vol. 16, pp. 43-64.

No. 485] NOTES AND LITERATURE 343^

certain barbed hairs from caterpillars of the brown-tail moth, when
applied to the skin, may cause a severe inflammation. Dr. Tyzzer
has found that these hairs occur over "two velvety brown spots which
appear on the dorsal aspect of the fifth and sixth segments after the
first molt." Similar spots are found after each succeeding molt up
to the last two spring molts, when they appear on all segments from
the fifth to the twelfth inclusive. At this time they occur also in
relation with the lateral tubercles of the same segments, so that the
caterpillar becomes much more poisonous than in its young stages.

elevations upon the caterpillar; the barhs. which at iiiK-rvals tend to
encircle the hair, point outward. If these hair>. which arc ca>ily
detached, are rubbed upon the skin they work their way inward,
pointed end foremost. It was supposed that the irritation which
followed was purely mechanical. Dr. Tyzzer has demonstrated a
chemical poison in the following manner. If the hairs are placed in
a drop of blood between a slide and cover glass, a modification of
the red corpuscles takes place at the apex of the hair. There the
rouleaux are broken up; the corpuscles shrink and become at first
spiny, and then spherical. That this is not a physical phenomenon
is shown by substituting hairs of similar shape from the tussock moth,
when no reaction occurs. It is believed that a poisonous substance is
emitted from the apex of the hair, although no pore is visible. If the
hair is broken the reaction occurs about the fracture, but otherwise only
at the pointed extremity. The poisonous substance is not destroyed
by baking the hairs for one hour at 110° C, but is destroyed at 115°.
In the latter case the hairs produce no dermatitis when applied to the
skin, and no reaction in the drop of blood. The poison is insoluble
in alcohol, acetone, chloroform, ether, acetic acid, and dilute liydio-
chloric acid. It appears, however, to dissolve in distilled water at (10°
C, and a further chemical study is in progress.

In regard to animal coloration it may be noted tliat tlie cater-
pillars of the tussock moth, said to present 'waniiiiir c()h)rs,' have
non-poisonous hairs; those of the lo moth, with a iiriH ii 'protective
coloration' are somewhat poisonous; and the poisonoiH hrown-tail
caterpillars have neither a warning nor a [>roteetive color. All three
forms, moreover, are eaten by birds.

Divided Eyes of Insects.— G. D. Shafer has studied the <livided
eyes in certain Odonata and Diptera ' and has followed tlu' late sta-es

344

THE AMERICAN NATURALIST [\^ol. XLI

of their development in two species. The modifications are introduced
in the nymph stage and are almost complete in the subimago, though
the eyes rapidly increase in size at the time of the final molt. Shafer
thinks that the two divisions of the eye are for \dsion in different kind
of light the regions with larger elements and less dense pigmentation
being available in twilight or in the darker hours.

Notes. — Dr. Lawrence E. Griffin has published in the Missouri
Valley College Quarterly Bulletin, (6, No. 4, 1907) a handy guide to
the dissection of the dogfish (Acanthias & Galius). Copies may be
had from the author at Marshall, Missouri, at 25 cents each.

BOTANY

Winter Rest.— In a very comprehensive series of experiments,^
in one of which as many as 283 species were used. Dr. Waher L.
Howard, of Columbia, ]\Io., has studied the effect of increase of
temperature, narcotics, lack of light, and dehydrating agents upon
plants in the resting condition. He comes to the conclusion that
the resting period is due to external influences, which also determine
its duration and intensity. It may be interrupted by the use of the
above mentioned agents. Though the results they produce are identi-
cal, their action is different. An am})le citation of literature enhances
the value of this paper.

Henri Hus.

Notes. — A quarto of 340 pages, devoted to a revision of the genus
Lcjjidiuiii ])>• Tliclhmg, has been separately issued from vol. 41 of
the Xrw )h>,hsr}rnjtni der AUgem. Schweizerischen Gesellschajt
/. (/. (icsnmini Xatiirwlsscnschaften, as a contribution from the Zurich
Botanical ^Vluseuu).

Some of the difficulties of cactus study are ])ointed out by Griffiths
and Hare in an economic leaflet issued as Bnllrtin 7W. iO,>, part 1, of
tlie Buivnii (.f Plant Industry. 1". S. Dcpiirtincnt of AgricuUure.

( )l,.(Tv;. liens on SaiTa. rni;i aiv | .ul .lislic( 1 l,y Macfarlane in The
Journal oj Hnhinf/ tor .lanuary.

Pflanzen. Inaugural-dissertation, Halle^ 1906. pp.111.

No. 485] NOTES AND LITERATURE 345

Opuntia pmilla as a Cape weed is discussed by Nobbs in The
Agricultural Journal of the Cape of Good Hope for December.

Illustrations of the celebrated cypress of Tule are given in Forest
Leaves for December.

Notes on rare ferns about Media, Pa., and especially Asplenium
ehenoicles, are given by PaluKT in vol. 2, no. 1 of the Proceedings of

From notes in Xatiire of I)ecem})er 13 and January 10, it appears
a subject of debate whethel- Spharotlieca mors-ura: is a new pest in
England or one of 30 years' standing.

Berghs gives an account of the nuclear phenomena of Spirogyra
in vol. 23, fascicle 1 of La Cellule.

Cruchet publishes on Labiate rusts in the Centralblaft fur Bak-
teriologw &c., Abteilung II., of Dec. 28.

The biology of tlie sand areas of Illinois is the subject of vol. 7,
article 7, of the Bulleim of the Illinois State Laboratori/ of Natural
History, by Hart and (ileason.

A short readable expr^.ition of liis views on evolution and mutation
is given by l)e^'ri(^s in Thr Munisf tor January.

An illustrated l.aiuH.onk <.|- -'V\^v Microscopy of Vegetable Foods"
with special reference to ihc> detection of adulteration and the diagnosis
of mixtures, by Winton and Moeller, has recently l)een issued by
John Wilev and Sons of New \'ork and Chajunan and Hall of London.

The flora of the Cuban 'Sierra Maestra' is considered in a forestry
studv rei)orted by Fernow and ^Faylor in the Forestry Quarterly of
December.

Tobacco-culture experiments, bv Hunger, occupy jiart 3 of the cur-
rent vohune of Archives du Musee Teyler.

(Hiaynle (Parthenium) rubber is the subject of a statistical note

346

THE AMERICAN NATURALIST

[Vol. XLI

Vol. 5, no. 16 of the Bulletin of the New York Botanical Garden
forms a general descriptive guide to the grounds, buildings and
collections.

The Report of the Michigan Academy of Science, vol. 8, contains
the following papers of botanical interest : — Kauffman, ' Unreported
Michigan Fungi. . . . Beal, 'A Study of Rudheckia hirta,' and 'Some
Botanical Errors Found in Agricultural and Botanical Text-Books.'
Dandeno, 'A Stimulus to the Production of Cellulose and Starch,'
'A Fungus Disease of Greenhouse Lettuce,' and 'The Aerating Sys-
tems of Plant Tissues'; Pennington, 'Plant Distribution at Mud
Lake'; Smith, 'Some Notes on Nodules'; and Sackett, 'The Asso-
ciation of Psevdomonas radicicola with Bacillus ramosus.'

W. T.

CORRESPONDENCE

Editor of the American Naturalist:

The Flying Fish problem, discussed by Lieut. Col. C. D. Durnford
in the American Naturalist for February (page (15), seems to be now
reduced to a (juestion of keenness of eyesight . I )o the wings or pectoral
fins of this fish in flight move so swiftly that the motion cannot be seen ?

The initial start of the fish ,.n Icavin.^i- 1 h(> water is^clearly due to

initial leap from the water, or wlieii hy skimniiiig along the surface
the tail touches the water, the n iiio-s arc seen to be in Vi\])n\ vibration.
When the tail frr.' fn.n, ilu- \NatiM-. ilir u in-^ niv outspread fan-
fashion and seem 1.. 1... lu-M finnh aiMlni n.M uiihuut Mbration, to be
folded when the (i-.h drop^ into the water. It lake-< Mroiig muscles
to hold the wings taut; we may admit that tlie fish has these; it would
take stronger muscles to cause the fish to move through the flapping
of the wings.

The problem is this: Does the fish fla)^ its fins? In the view of
Col. Durnford it does. In liis \\v\\ ilic vibrations are so rapid that
to most observers they cannot v\rr\)\ at \ \\r bciiiiiniiig or end

of the flight, when the tail is in \ hr water.

In the view of others, tiie wiiios ar.' not flai.jHMl al all. When the

347

348

THE A M ERIC AN NA T URA LIST [Vol. XLI

be in a state of rapid vibration, but this is apparent only, due to the
resistance of the air to the motions of the animal. While the tail is
in the water, the ventrals are folded. ^Yhen the action of the tail
ceases, the pectorals and ventrals are spread and held at rest. They
are not used as wings, but act rather as parachutes to hold the body
in the air. When the fish begins to fall, the tail touches the water,
when its motion begins again, and with it the apparent motion of the
pectorals. It is thus enabled to resume its flight, which it finishes
with a splash. W' hile in the air it resembles a large dragon-fly. The
motion is very swift, at first in a straight line, but later deflected into
a curve. The motion has no relation to the direction of the wind.
When a vessel is passing through a school of these fishes, they spring
up before it, moving in all directions, as grasshoppers in a meadow."
Very truly yours,

David Starr Jordan

February 23, 1907

PUBLICATIONS RECEIVED.

350

THE AMERICAN NATURALIST [Vol. XLI

Bull, de rinst. Ocearwg. de Monaco, no. 92, 24 pp., 19 figs., 4 pis.— Juday, C.
O.stracoda of the San Diego region. II. Littoral forms. Univ. of Cal. Publ.,
Zool., vol. 3, pp. 135-150, pis. 18-20.— Juday, C. Cladocera of the San Diego
region. Univ. of Cal. Publ., Zool, vol. 3, pp. 157-158, 1 fig.— Millspaugh,
C. F. Flora of the sand keys of Florida. Field Columbian Mus., hot. ser.,
vol. 2, pub. 118, pp. 191-243, 19 maps.— Moore, H. F. Survey of oyster
bottoms in Matagorda Bay, Texas. Bureau of Fisheries, doc. 610, 86 pp., 13

Wyoming Exp. Sta., bull. 71, 39 pp.,^11 figs.— Nowell, H. T. Duty of water
on field pease. Wyoming Exp. Sta., bull. 72, 16 pp., 4 figs.— Schaeffer, C.

at Brownsville, Texas, and in the Hi^Tchuca Mts., Arizona. Brooklyn Inst,
of Arts and Sci., vol. 1, pp. 291-306.— Schlaginhaufen, O. Uber das
Leistenrelief der Hohlhand und Fusssohlen-Flachen der Halbaffen, Affen, und
Menschenrassen. Ergebn. d. Anat. u. Entw., vol. 15, pp. 628-662, 14 figs.—
Schlaginhaufen, O. Beschreibung und Handhabung von Rudolf Martins
diagraphentechnischen Apparaten. Korrespondi-nzbl. d. Deutsch. Gesellsch.
f. Anthr. Ethn. u. Urgeschichte, vol. 38, pp. 1-6, 4 figs.— Schlaginhaufen,
O. Ein Canalis craniopharyngeus persistens an einem Menschenschadel und
sein Vorkommen bei den Anthropoiden. Anat. Am., vol. 30, pp. 1-8, 5 figs.
— Shafer, G. D. Histology and development of the divided eyes of certain
in.sects. Proc. Washington Acad. Sci., vol. 8, pp. 459-486, pis. 24-27.—
Springfield Musei.m of Natural History. Bird migration; dates of
arrival of birds within ten miles of Springfield, Mass., from 1901 to 1906.—
Vles F. Sur I'existence de la Mye dans la Mediterranee. Bull, de I'lnst.
Ocf-anog. de Monaco, no. 94, 2 pp.— Wilcox, W. A. The commercial fisheries
of the Pacific Coast States in 1904. Bureau of Fisheries, doc. 612, 74 pp.

Boletin de la Sociedad Aragonesa de Ciencias Naturales, vol. 6,
no. 1. Bulletin biologique, Dorpat, Russia, no. 1. Bulletin of the
Charleston Museum, vol. 3, no. 2. Second Report of the Wellcome
Research Laboratories at the Gordon Memorial College Khartoum.

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NATURALIST

A MONTHLY JOURNAL
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CONTENTS

I. A Graphic Method of Correlating Fish Environment and Distribution

ALBEET H. WRIGHT 361

II. The Uicrogametophyte of the Podocarpineae

E. C. JEFFBEY and M. A. CHEYSLEE 355

III. The Problem of Color Vision JOHN M. DANE 365

IV. The Breeding Habits of Amblystoma punctatum Linn. BEETEAM G. SMITH 381

V. The Staff-tree, Ce/astrus scandens, as a Former Food Supply of Starving

Indians FEANE T. DILLINGHAM 391

VI. Notes and Literature : Genera/ Biology; The spirit of nature study.— Hetcro-

genesis. Zoology: Anatomical terminology. — The blending and over-
lap of instincts in birds. — Variation of scutellation in the garter
snakes. — A method for removing the gelatinous coats of eggs. — The
star-nosed mole on Long Island, N. Y. — Notes. Botany; Cytology
and mutation. — Variation and differentiation. —Cotton. — Notes.
Geology; The elements of geologj' . . . • 396

VII. Publications Eeceived 411

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E. A. ANDREWS, Ph.D., Johns Hopkins University, BaUimore
WILLIAM S. BAYLEY, Ph.D., Co»y University, WaterviUe
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell University, Ithaca
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA., M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University
CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM, Ph.D., Cornell University, Ithaca
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, Pittsburg
D. p. PENHALLOW, D.Sc., F.R.S.C., McGHJl Univer»Uy, Montreal
H. M. RICHARDS, S.D., Columbia University, New York
W. E. RITTER, Ph.D., UniversUy of California, Berkeley
ERWIN F. SMITH, S.D., U. S. Department of AgricuUure, Washington.
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington.
W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis
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WILLIAM M. WHEELER, Ph.D., American Museum of Natural History,
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THE

AMERICAN NATURALIST

Vol. XLI June, 1907 No. 48G

A GRAnilC MF/rilOl) OF ( ()iniHLA'riX(J FISH
EXVIKONMFN r AN D I )IS'rin HI ''ll< )\

albi:rt hazfx wrigiit

It is some years since ornitli()lof?ists saw the advantages of a
graphic means of rej)resentiiig com])lex bird waves and their
coincident relation to i)liysical conditions. In ichtliyologv a
schematic method whereby fish (hstribution and cnxironnient
can be correlated is not less valuable.

The study of a stream and its fishes inv()lv(-s the consideration

ma.ss of data, that one is mijH'ilc.l ro a(i(.|)t scuu" uraphic method
to make results appear (|nickl\- and clcarK.

stic.un indiKlnii, i \ i-utx ..t'[H-^'''l» ...ndition. I lu fiLi < on-

th. mouth ot th( Muaui. and the (oiu.pondinil N»ni<al on the
right of the chart is its source. The dotted verticals numbered
below (1-S) mark mile points. Beneath the "Misc. Data" space,
these mile lines are not dotted but continuous.

The hea\7' horizontal oj)posite eacli species indicates the ranjxe
of that species in the stream. \\ hcivvcr the hori/oiital is broken,
it indicates the occasional oeciirrence ot die species. ANheiiever
a species gains entrance to a --ti'eam troiii two or mon> pomt> an
arrow tip at the end of each ot its raiiuv hues iii.licate- the (hrectioii
ot Its migration; e. 7., in our hvpotheiie -iivani the carp. ( i/pnniis

T\w continuous horizontal above tin- fiiNi specie, in the li.t.
upu.entsth( .intake ot tlu uatu 1 lu boitom i. .ho^n bx ,!„
(UiMd hue labele.1 "Bottom (.t Mm am llu asna^t dtptli
351

352

THE AMERICAN NATURALIST [Vol. XLI

at* any given point is, therefore, the vertical between these two
lines, read from the scale at the extreme right of the chart.

The continuous horizontal immediately below the last species
enumerated, represents the altitude of the mouth above sea-level.
The profile line indicates the drop in the stream. The approxi-
mate altitude of any given point along the stream is shown by the
vertical between the two above mentioned lines, and read from
the scale at the right.

The continuous horizontal in "Valley Cross Sections" repre-
sents the stream, on either side of which is shown a section of the
countr}' for one and a quarter miles. In these cross sections at
every mile point the geologic formation can be indicated.

The current, width and bottom data are self explanatory. In
the "Miscellaneous Data," bridges, marshy regions, dams, etc.,
are represented so far as possible, by the conventional signs
employed by the U. S. Geological Survey, thus:

Bridges (A) e.g., below the one mile mark, at the

two mile mark, etc.

Woodlands (B) " at the one mile mark.

Swamp (C) " between the one and two mile

Important Tributary (D) represented by a forked wavy line.

The position of the tributary line
in the upper or lower part of the
"]Misc. Data" space indicates
that the tributary enters the stream
from the right or left side respec-
tively.

" at the 1.25 mile mark, (Enters

from the left side.)
" at the 2.6 mile mark. (Enters

at the 5.8 mile mark,
from the right side.)

354

THE AMERICAN NATURALIST

[Vol. XLI

■(I)

c. g.,

at the 7.25 mile mark. (Enters
from the left side.)
at the 7.5 mile mark.

Takiiif? the common bullhead, Amemrus nebulosus, as an
exam})le, one is able to read from the chart, concerning its distri-

of the stream, gradually decreasing in abundance at the end of the
second mile. Throughout the middle course where rock or
gravel bottoms and swift water or rapids occur this species is
absent. In the upper course where the current and bottom are
influenced by the dam, located 5.5 miles from the mouth, it re-
appears. In the latter iiistMiicc, its |)i'('s(Mice so near the head-
waters is due to canal ovci liows at the 7.5 mile point. It seldom
frequents water less than 4 feet (ieej). In both ranges, the drop
in the stream is slight, so that the current is just perceptible at
the mouth, and imperceptible at the dam. At these two points
the stream's width is respectively 27 and 32 feet. In both places
a muddy bottom obtains. In the lower course the stream lies
on a delta formation while in the upper course the underlying
stratum is glacial drift.

The Johnny darter, Boleosoma nigrum, occurs at the source
of the creek, due to a contribution at floodtime from another
stream across the divide, the two sources being on the same level
and continuous at some seasons. ^I'he falls on either side of the
4 imMc ]»()iiit would pircludc its reaching the source from the mouth.

deductions might be drawn. Should it seem desirable to make
the work more intensive, to restrict it to a limited portion of a

bution, etc., the following:

THE MICROGAMETOPHYTE OF THE PODO-
CARPINE.E^

E. C. JEFFREY AND M. A. CHRYSLER

Although at the present time the views in regard to the rela-
tionships of the Coniferales depend very largely on the study of
their gametophytic or sexual generation, our knowledge in regartl
to the gametophyte of the coniferous families is often very meager.
The two families concerning which information is actually most
needed are the Podocarpineae and the Araucarineae, exotics con-
fined chiefly to the southern hemisphere. There is a prospect that
our ignorance in regard to the Araucarineae will soon be less dense
than it is at present, a consummation devoutly to be desired on
account of the prevailing views, which make them the most ancient
of the Coniferales. It is proposed in the present article to describe
certain features of the male sexual generation of the Podocarpineae
observed in material which we owe to the kindness of Dr. Cockayne
of Christchurch, New Zealand, and Dr. Treub, Director of the
Royal Gardens at Buitenzorg, Java. To both of these we tender

made it possible for us to study souw of tlu- Aiistiala-iaii i;tMU'ra
of the Podocarpineae. The material ar our (lis[)o>al wa- fixed
in formaline or alcohol and conse(|uently leaves soiiiethinu to be
desired in the preservation of cytological details. A> we shall
however confine ourselves to the gross features of inielear structure
which do not suffer seriously by the methods of pre>ervation de-
scribed, this will not be a serious disadvautaue.

The first species to be consideretl is Pmhtcarpus pol if.sfdchi/a ,
material of which we owe to Dr. Treiih. Director of the 15otanic
Gardens at Buitenzorg. The male cones in our pos-^ession are
in various stages of anthesis; but some of them >ii(.\v (juiie young
anthers or microsporophylls in the ui)|)er n-oion of tiu> axi-. This
feature has made it possible for us to follow step l>y >te|> the
development of the male gametopli_vt<- u[) to tlie time of the ^lied-

' Contributions from the Phanerogiunic I.a!)r.rat<.i i.'< ut Harvard I ni verity.
No. 8.

355

THE AMERICAN NATURALIST [Vol. XLI

<lin^ of the ])oll(>n or inicrospores, in spite of the fact that the
iiiatenal represents a single collection. A figure 1, represents
the first mitosis in tiie microspore, which it will be observed is
well advanced towai-d completion. The state of preservation of
this material is remarkable in view of the fact that it was fixed
m strong alcohol. In /; fignre 1, is to be seen the first prothallial
cell fnlly formed and lying over against the Uj)per or poKicrior side
of the microspore, lieneath it, is the residual mu lens surrounded
by vacuolated protoplasm. In c figure 1, is to b<> seen the mitosis
which precedes the formation of the second prothalhal cell. In d

figure 1, the second prothallial cell is complete and lies against the
first. At about thi^ time the .trough thickened posterior ^^all of
the microspore, which s(^ems to be a peculiar f(>atin'e of podocar-
pineous pollen, becomes markedly sculptured as is shown in (L In
a figure 2, is shown a still later phase where the s(»-( alle(l ncuera-
tive cell has become added to the prothallial cells, which he on the
posterior wall of the microspore; it arises from iuioilier (hvision of
the residual nucleus. The contents of the pnlh n ui.uu .u thi> .t.me
resemble in detail the conditions to Ik- tound ui the aluetuicdus
microspore before the prothallial cells have begun to degenerate.

No. 486]

PODOCARPINE.E

357

In Podocarpus however tliere is no atrophv of the protliallial
riidnnents at this stage, but tliev undergo further chanoes of a sur-
prising character, comparable onlv to those recently described bv
Thomson^ in the genus Araucaria. In h figure 2, a later stage of
development is shown, in which each of the prothallial cells has
undergone transverse or anticlinal division. Division generally
takes place first m the outu piothalhal cell hing next the ^^all

of the microspore and subsequently in the second prothallial
cell. Contrary to the statements of Coker' in regard to P. conacm,
where a similar but less well marked condition has been descril)ed
as a probable abnormality due to artificial conditions, anticlinal
divisions of the prothallial cells are not initiated l)v direct divi-
sion of the nucleus but by true mitosis. In tlu' (vlls <lfn\c.l tiom

358

Tin: AMKinr.W \ ATI ' h'A LIST [Vol. XLI

the second protluilluil cell in b figure 2, the nuclei are still in the
spireme condition. Occasionally anticlinal divisions occur in the
generative cell as in the prothalhal rudiments. One such case is
re{)resented in c figure 2, which is an obvious and clear mitosis.
I sually however in P. 'polystachya such divisions of the generative
cell do not occur, although they are exceedingly common in some
ol the other species which we have had the opportunity of study-
ing. At about this time the prothallial cells lose their walls; and
their nuclei, floating freely in the cavity of the inicros{)ore, are no

longer (mkIom-iI liy ( ytoplasmic liodit^s. The nuclei, however,
persist inilflinitfly and j)a^s out as a swarm into the pollen tube.

spore at this >tauc, tlic ivsidiml or tiilx" mich-us can l)(> distinguished

cell, or the central cell dcrivcil from it in ca-c it has undergone

No. 486]

PODOCARPINE.E

359

anticlinal divisions previous to being set free from the prothallial
complex, always retains its protoplasmic body as is generally the
case in other Gymnosperms, and thus cannot be confused with
any of the other contents of the microspore in tlie condition which
immediately precedes anthesis.

In Podocarpus jerruginea from New Zt alaiid. iiiat<M-ial of which
we owe to the kindness of Dr. Cockayne, the carHer stages are
not so well represented as in the species described above, but so
far as they have been followed they present no essential deviation
from the course of events in P. polystachya. A figure 3, represents
the abietineous stage of development in this species. The preser-
vation is even less good than that of the Podocarpus already de-
scribed, and the protoplasm has shrunken from the microspore
membrane. In b figure 3, is shown a fully developed grain, in
which only one of the prothallial cells has undergone division.
The generative cell in this case is also free from divisions, although
it has rounded off and is almost ready to be set free from the
cavity of the microspore. C figure 3, presents a tangential view of
the first prothallial rudiment, which in this case has undergone
two anticlinal divisions, so that four cells have resulted. D figure
3, presents a longitudinal section through the air chambers and
shows anticlinal divisions in both of the prothallial cells. E figure
3, shows a similar condition" in the prothallial rudiments; but in
this case there are two lateral derivatives of the generative cell.
The latter are very small in size compared with the central cell of
the generative complex and with the derivatives of the prothallial

As is represented in figures 3 and 4, starch is commonly found
in the pollen grains, especially in the younger stages, tliougli its
presence is by no means constant. A similar feature has been
noticed by Coker in the article cited above.

n figure 4, shows the ganietophytic <levelo|.n.rnt in a probably
mature microsix.re of l)arr;/,!iuin Hi^hrlll!;, another n-proenta-
tive of the Fodocarpincie. The material in this cax- proved to be

360

THE AMERICAN NATURALIST H'ol. XLI

ven^ badly preserved. Dacrydiiim is distinguished from Podo-
carpus by the transverse striations of the thickened posterior wall
of its microspore. In the species of Dacrydium which we have
examined more than two prothallial cells are present, but the
derivatives of the prothallial rudiments do not seem to be as
numerous as they are in Podocarpus, where there may apparently
be as many as eight present (P. ferruginea).

Through the kindness of Dr. Cockaviie ha\e had the op-
portunity of comparing the microganiet()])liyTic (lc\el<)pment of
Podocarpus and Dacrydium with that ])reseiit(Ml in Agathis,
probably the more ancient of the two living genera of the Arau-

natelv verv much shrunken, possiblv on^accolmt of the small
amount of alcohol m which it was preserved, but this fortunatelv
does not interfere with the understanding of the general conditions
present in the microgametophvte. In h figure 5. is shown an
apparentlv mature microspore, ^^e cannot however speak with
certaintv on this point, since none of the microsporophvlls in our
possession have shed their pollen. It is to l)e noted that the
conditions present in this figure closelv reseml)le those depicted
in h figure 2. and d figure 3. In other words there are sub-
M(|utnt uituhnil(h\i lonsprt < nt m tlu two pi .tli ilU il in Imunts
which an <ru..idb 1 u<l dmn tlu n ik u. tlu Vbittnu . 1
figure rcscinl>h's ch.sdv and dilVcrs in tlic fact that onlv one
of the piothilliil I ihncM 1 . 1( n< hM 1< 1 In < figuie 5,
1^ ^ho^^n a tuu 1 ti i! \ < \ t i t tl < i . tl illi il ( t lU Ihere

PODOCARPINEM

361

It is apparent from the foregoing paragraphs that in two genera
of the Podocarpinejie there are unusually numerous prothallial
cells present in the microspore, which are derived by the subsecjuent
anticlinal divisions of the two primitive prothallial cells. T\vaX
these features are perfectly normal ones in the Podocarpinca' i>
made clear by the fact that all our material is from plants grown
in their native habitat and presumably under natural condition^.

These features are fiinluT parallchMl by x\w coiiditions presented
by the microspore of tlic araiu aiiaii uoiuis Aijathis. The question
here arises if we arc lo i-coanl tlic rich j)r(itliallial (Midowment of
the Podocarpinca- as tlir iviciilioii cf a tVaimv possessed by tlie
ancestral ( "(.tiifcral<- <.r a^ ;, ivccut < nio-ciicii.' adaptation, which

tions found in the Gymnosperms in uvii.Tal, particiihuly the
more ancient of those still living. In the primitive /(.idogaiuous

362

THE A M ERIC AN NA T URA LIST [A^ol. XLI

Cycads and Ginkgo there are one or two prothallial cells present.
The generative cell undergoes only a tangential or periclinal
division in connection with the formation of the stalk cell and
antheridial cell. The antheridial cell in both the Cycadales and
Ginkgoales gives rise to two spermatocytes, the mother cells of
antherozoids. In the Abietinete, which we know from the
evidence of the fossil remains extend very far back geologically
in forms allied to Pinus, there are two evanescent prothallial cells
present in the mature microgametophyte, and a generative cell
which as in the zoidogamous Gymnosperms gives rise to stalk and
antheridial cells by periclinal division. The antheridial cell in turn
gives rise to two cells which are to be regarded as the homologues
of the two spermatocytes of the Cycadales and Ginkgoales. In
the Araucarineae, so far as our knowledge goes, there are formed at
first two prothallial cells, which may subsequently undergo more or
less numerous anticlinal and possibly also periclinal divisions.
The final history of the generative cell is obscure, but it is to be
inferred from the brief summary of Thomson (loc. cit.) that the
antheridial cell of the Araucarinete does not divide into two as in
the AbietinctT and the ancient zoidogamous Gymnosperms. In
the Araucarine£e there is a further remarkable feature in that the
pollen grain does not reach the micropyle of the ovule as in the
other Coniferales and all other known Gymnosperms living or
fossil; but is deposited on some part of the ovuliferous scale or
megasporophyll (on the 'ligule' in Araucaria) thence sending a
pollen tube down to the ovule, in a manner analogous to that
obtaining in the Angiosperms. Thomson, adopting the prevailing
hypothesis that the Araucarinese are the most primitive Coniferales,
designates this peculiar mode of fertilization as primitive or ' pro-
tosiphonogamic'

This view presents some difficulties, for if the quasi-angiosperm-
ous method of fertilization found in the Araucarinese is ' primitive '
it is difficult to see why such a method is entirely absent in the
older g),'mnospermous series, the Pteridospermje, Cordaitales
and Ginkgoales, or being ancestral for the Coniferales is entirely
lost in the coniferous families other than the Araucarineae, which
have moreover a method of pollination resembling closely that
of the older Gymnosperms in that the microspores are received
through the micropyle. The reported presence of only a single

A'o. 486]

PODOCARPlXK.i:

sperm-cell in the Araucarineje sup{)lies another ariruineiit a<;aiii.st
their being more primitive than the other Coniferales. Their su-
perior antiquity further does not rest on any sound paljeonto-
logical Imsis, for so competent an authority as Schenk (Zittel's
Handbuch) remarks that if more abundant and more ancient
geological occurrence were to be considered as a criterion (.f anti-
quity, the Araucarineie must yield place to the 'ra\o(lin( a'. It
appears not unlikely, especially in view of observations made l)y
one of us on Mesozoic Coniferales, shortly to be published, that
the ' protosiphonogamic ' method of fertilization which is the inter-
esting discovery of ]\Ir. Thomson, is correlated with the prolifera-
tion of the prothallial cells in the Araucarineje, since the greater
length of pollen tube, in the absence of any special conductive
tissue such as is found in the Aii>:iosj)erni.s, calls for a greater
development of prothallial tissue. The failure of the pollen to
reach the micropyle, on the other hand, may have been due to the
unfavorable inihieiu t' of drought upon the fluid secretion which in
other ( oiiifcrs Hoats tlie pollen to the micropyle.

Turning from the Araucarineje to the Podocarpinea\ we find
\(^ry similar conditions in regard to the prothallial j)rolifi'ration-.
The plan of prothallial development here as in the Al)i(>tinra' and
Araucarineiv involves two prothallial cells, but as in the Arau-

tion. That this i> the n-ue view <.f the matter is rcn.lclvd more
prol.al.lr by the fact that even the gcn.>rativc cell may be albvt.T

jrrnnj-lu nuA P \larr;idloidrs, <\vsvv\W^ above, ^'bhere i'. <.er-
tainly no reason from our knowle.lge of the older and zoi.lo-

364

THE AMERICAN NATURALIST

[Vol. XLI

very remotely connected with the Abietinese. Their pecuhar pro-
thalHal developments represent an apparently cenogenetic super-
addition to the primitive type of coniferous microgametophyte
found in the Abietinese. If this view be taken of the position
of the Podocarpineae, it may well be extended to the Araucari-
neae which present a similar microgametophytic development,
although it would take us too far afield and would involve the
discussion of yet unpublished data in regard to living and fossil
Coniferales, to defend that proposition in the present connection.

Summary

1. The Podocarpineae as represented by the genera Podocarpus
and Dacrydium are characterized by a proliferation of the two
original prothallial cells through more or less numerous anticlinal
divisions.

2. The anticlinal proliferation of the prothallial cells in some
cases is accompanied by a similar proliferation of the generative
cell, an abnormality which appears to have been described in no
other Gymnosperms.

3. Similar proliferation of the two original prothallial cells
has been observed in the araucarian genus Agathis.

4. The proliferation of the two prothallial cells in the Podocar-
pineae and Araucarineae and the proliferation of the generative
cell in certain species of Podocarpus, cannot be regarded as a
primitive feature.

5. The ground plan of microgametophytic development found
in the Podocarpineae and Araucarinea? points to their derivation
from an ancestral stock allied to the Abietineae.

6. Since the Podocarpineae and Araucarineae present many
features of similarity in general habit, in geographical distribution,
in the organization of their megasporophylls, and the development
of their microgametophytes, it seems not improbable that they
are somewhat more nearly allied than has been supposed.

THE PROBLEM OF COLOR VISION

JOHN M. DAXE

The problem of color vision is one of the most intricate which
the biologist is asked to solve. The following paragraphs are
intended to indicate the several methods which are being employed
for its solution, together with some of the results thus far obtained.
The anatomy of color vision will be considered first; then in turn
its physiology and its development; and finally, the abnormal
conditions of color blindness, together with the theories of normal
vision to which they have given rise.

Anatomy. The mechanism of color vision is lodged in the rod
and the cone cells. A ray of light, after passing through the
lens of the eye and its vitreous body, penetrates several layers of
the retina, thus arriving at the proximal ends of the elongated
rod and cone cells. These cells are arranged in a single row.
The light traverses the length of the cells to their distal ends which
it stimulates. The rod and cone cells project against a single
layer of heavily pigmented cells, the stratum pigmenti retinae
(Fig. 1, S. P.). These have non-retractile processes which are

the f<)nn of'donljlttMllVn-vs!^^^ .ranules. niiurat.-s into th'ese
processes when the eye is illinninated ; in ilie dark it i> withdrawn
into the cell body.

Every rod cell consists of a rod, a rod fiber, and a nucleus,
arranged as shown in Fig. 1, A. A rod, which is from 40 to 50 //
long and L5 to 2 ri in diameter, consists of a doubly refractive,
lustrous oiifcr scqment, and a singly refractive, finely granular
nnur .se<jmn,f. In serum or dilute osniie aeid the outer segment
breaks into a series of regular transverse discs which are lu'lieved
to indicate a stratified structtu-e in the living rods, \ isual pin-ple
is a pigment which occurs only in the outer segments of the rods.

365

366

THE AMERICAN NATURALIST [Vol. XLI

It bleaches rapidly in the light, but (unless the pigmented stratum
has been removed experimentally) it is soon restored in the dark.
Light thus appears to incite chemical processes in the outer seg-
ments of the rods. The inner segments are sometimes described
as having a longitudinally fibrillar structure in their outer portions.
The opposite ends pass rather abruptly into the very slender rod
fibers. Each fiber somewhere in its course expands to enclose
the nucleus, and finally terminates in a pyriform enlargement.
The nucleus in preserved specimens may have its chromatin
arranged in a few broad transverse bands.

Every cone cell consists of a cone, a cone fiber, and a nucleus.
The cones like the rods are divisible into outer and inner segments.
The outer segment is usually shorter than that of the rod (12 fi)
and tapers somewhat to its rounded extremity. It never contams
visual purple, but otherwise, as for example in breaking into trans-
verse discs, it resembles the outer segment of the rod. The inner
cone segment bulges like the body of a flask. It is divided into
an outer, longitudinally fibrillar, ellipsoid portion, and an inner
contractile myoid portion. The non-contractile ellipsoid is said
to become strongly eosinophilic in the dark. Because of the myoid
substance the cones, unlike the rods, may alter their length. The

ntractility is said to be less i

, and less i

the latter than in some amphibia and fishes where the myoid
segment is reported to shorten from .Id ." to The nuclei are

found in a mass of protoplasm near the l)ase of the cone; beyond
the nucleus the protoplasm forms a cone fiber which is thicker
than that of a rod and which ends in a branched and expanded

The stimuli received bv the outer segments of the rods and
cones are transmitted through their fibers to the nerve cells of the
retina, and thence to the brain. A single retinal nerve cell receives
the stimuli from several rods and cones.

Since rods and cones are believed to have different relations to
the perception of color their distribution in man and other aninuUs
should l)e significant. In the peripheral portion of the human

where vision is nH»t aciiic, ](><1> and tones are equally abundant,

Xo. 486]

THE PROBLEM OF COLOR VISION

367

and in the fovea itself only cones are found. These cones, how-
ever, are strikingly rod-like in form, and greatly exceed the rods
in length (Fig. 1, B). Slender cones are also found in the thick-
ened area centralis which in many mammals replaces the human
fovea.

In the ape, horse, pig, cow, sheep, and dog the rods and cones
are similar to those of man. In rodents which avoid the light
the cones are "very small and hard to detect since their inner
segments scarcely differ from those of the rods, from wliich they
may be distinguished by their much shorter outer segment. M.

nu. r.f. Rod, z.s.... as. S,P.

Schultze at first (piostioned the existence of cones in the mouse,
guinea pig, mole, hedgehog, and bat. Tiie cat undoubtedly has
cones but they are small, slender, and except in the area, infre-
cpient."^ Birds have a single or double fovea, like that of man.
Cones are small but very numerous, and in their inner segments

various sliadcs of yt'Uow. urceu or red. l'n'>\iui;il)ly these drops

36S

THE A M ERIC A N NA T URA LIST [Vol. XLI

believed that reptiles have only cones. Tn fishes and amphibia,
l)()th rods and cones occur; in some sharks, rays, and eels, however,
the cones so resemble rods that they may i)e overlooked. Whether
or not deep sea fishes are without cones is apparently unknown.
In the various (jroups of aninuils the rods and the cones each
present modifications of structure, with which as yet physiological

Physiology. The physiology of color vision is the study of the
functions of the rod and the cone cells. In passing from a bright
to a very dim illumination one ex|)crienccs a momentary blindness;
after becoming accustomed lo tlic darkness, a modified form of
vision is regained. In this iirl/ii/hl risimi the fovea is far less
sensitive to light than the more peiiplit-rai parts of the retina.
Moreover all objects appear in shades of gray. The spectrum is
bright but colorless, and its brightest part has shifted from the
yellow portion toward the blue. Von Kries has explained these
facts by assuming that the cones are the agents of day vision, and
the rods of twilight vision.' Cones, exclusively, occur in the fovea
where day vision is most acute; and rods predominate where
twilight vision is at its best. The fluctuations in the visual purple
of the rods show that they respond to the varying intensities of
dim light, and tliis purple is known to desintegrate most rapidly

or not the bleached t'dds are active in day'^vi.ion has not been

It is probable that all cones do not respond to color stimuli.
In the periplieral })orti()n of the retina there is a partially color-
one another; and the outermost portion of th(< retina is always
totally color blind. Since cones occur in tliese areas they also
must'be color blind. From these con.si.h-ration^ it is reasonably

No. 486]

fairly be questioned whether the lower animals are capable of
color vision. The biological importance of this problem is very
great, since prevalent theories of the development of the colors
of flowers, and the bright plumage of male birds, assume a color
perception in insects and female birds essentially like that in man.
To learn what a bee actually sees has been thought impossible
since it re(|uircs that otic should possess the nervous system of

Tiierc is a laruv litcralun^ (Icaliiiu' witli the distinctions which
the lower aiiiiuals make Ix'twccti vai-ioii^ colors, but the factor of
iiit(Mi>ity or briulitiic» lias seitloiii been satisfactorily eliminated.
The troui fishei'iiiaii is eonfidcMit that one lisli, at least, discrimi-
nates eoloiN w ith |)recisi(.ii. ( 'aret'iil (^xperiiiients with the chub,

to (eliminate briu'litiu-ss, indicate that the chub distinguishes red

370

THE AMERICAN NATURAEIST

[Vol. XLI

ceive colors." Two colors of equal l)rightness are distinguished
better than two grays of equal brightness; and though the bright-
nesses are the same, colors may be distinguished from grays. '

In the dancing mouse, however, the cones of wliich are at least
very rod-like, Yerkes has recently found that color vision is ex-
tremely poor. There is some evidence of (hscrimination of red
and green, and of red and blue, but none whatever of l)lue and
green. A]){)arcntly such visual guidance as is received results
from dif^'ercnccs in brightness. The mouse discriminates blacks
grays, and whites.^

Because of the inherent difhculties in the investigation of color
vision in the lower animals, comprehensive results liave not
yet been obtained, but the newer methods |)romise notable
discoveries.

Development. Since color vision is a complex differentiation,
it might be expected that in the course of development, an individ-
ual should successively pass through the simpler stages by which
it was acquired. Anatomically it has been shown that the retinal
layers first become distinct at the center of the retinal cup, and
that the differentiation of tlie retinal cells decreases from the
center toward the periphery. In the chick it is said that the cone
nuclei may be identified at an earlier stage than the rod nuclei,^
but it is not generallv recognized that one t'onn of visnal cell ])re-
cedes the other.

The development of color vision has hvvu theon^tieallv consid-
ered l,y Mr^. l.iidd Franklin.' Il.-r theory assnnies that tli,> color-

phoio-ehemieai substance called the gray substance, which is com-

These gray molecules, which persist in their {)rimitive state only
in the rods, upon disas.sociation furnish us with the gray sensa-

No. 480]

THE PROBLEM OF COLOR VISION

371

tions. In the cones the gray molecules have undergone a devel-
opment such that a certain portion only of the molecule becomes
disassociated by the action of light of a given color.

Tlie ditt'erentiation of the primitive gray molecule is supposed
to have taken place in three stages (Fig. 2). The first stage is

" " ' other the sensation of red.

If the rcil and u'rccii liToupini^s are disassociated together the

372

THE AMERICAN NATURALIST [Vol. XLI

vision the red end of the spectrum is lost, and the green-bhie por-
tion is its brightest part, he considers that the photo-chemical
substance of the rods is attuned only to the green-blue light, which
is perceived as colorless. Later this photo-chemical substance be-
comes sensitized in two stages, first to include the green-yellow,
and then the yellow-red, which however are still perceived as
colorless light. Thus a gray molecule like that of ]\Irs. Franklin's
first stage is constructed. It occurs in the color blind peri{)heral
cones. The formation of color-reacting groupings in the partly
sensitized gray molecule leads, according to Schenck, to those
forms of human vision in which the red end of the spectrum is
shortened.

Observations upon the color perception of young children do
not support these developmental theories. Holden and Bosse ^
tested two hundred children by placing before them square pieces
of colored paper attached tt) a gray background of similar bright-
ness. If the child made an effort to grasp the scpuire, its color
must have been perceived. It was found that the average child
would react to all colors by the tciilli month, the red end of the
spectrum causing response a little earlier than the violet end.
When ribbons of six spectral colors were i>laee(l before children
of from seven to twenty-four months, red was selected first; orange
or yellow second an.l lliinl; an.l -reen, blue and violet last of all.
Naoel^' .h,n\e<l his child „t' tu entx -ei-ht nu.i.t h. each of the spectral

him their names. Ued an.! -reen were learned .-asilv, l>nt l)lne
^^a. acquired uilh -.vater dillicuilN than anx other <-ol'or, in<.lnd-
ing violet, (ireen, violet, and red were preternMl; l.iack, yellow,
white, gray, and blue ha<l secon.lary rank. ( )ther exf.eriments
with the color perception of children have givtMi ditVerent r<>snlt-.
It is clear, however, that children are not known to pass from a
color l)lind stage, through one of yellow-bhu/ vision, to a discriini-

Xo. 486]

THE PROBLEM OF COLOR VISION

373

which any of the colors is unknown; and the notion derived from
studying the color terms and references in ancient literature,
that man in historic times had a deficient color sense, is not sub-
stantiated. It may be that as in children, the red portion of the
spectrum was preferred to the blue, but even this is not e>tal)-

All the colors which are normally perceived
may be ])ro(luce(l by combinations of the spectral red, green, and
blue. Normal \ ision is therefore trichromatic. Sometimes in
trichromatic vision the red end of the spectrum is shortened; in
other cases a nu'xture of red and green, which to normal persons
appears pure yellow, may seem tinged with red or green. Thus
there are variations in trichromatic vision, (ireater abnormalities
may take the form of dichromatic and monochromatic vision. The
latter is a rare pathological condition in which all colors are per-
ceived as shades of one; vision therefore is essentially colorless
(achromatic), the images obtained being comparable with photo-
graphs. In dichromatic vision color perception is so limited that
all of the shades perceived may be ma'le by combining two (.f the
spectral colors red, green, and blue; blindnes. to the third of the>e
colors may be partial or com|)lete. Tlu^ ordinary color blindness

hereditary. It may cause so little trouble as to pa^s undetected
until the age of seventy. All attempts to overconie the color
blindness by educating the color sense in various ways, have failed.

Since dichromatic color blindness plays so large a part in the
theories of normal vision, a portion of Dr. Pole's description of his
own case is here inserted. He says,' "In the fir-^t place we see
white and black and their intermediate gray. i)rovided they are
free from allov with other colors, precisely as others do. (Such
statements are conHrme.l by those who are color blin.l in one .-ve,
the other being normal.) Secondly there are two c.)lor^. namely
yellow and bhie, which also if unalloved we se,-. so far as can be

374

THE A MEHH WX .V.l TURALIST [Vol. XLI

colors of which we have any sensation. It may naturally be asked :
Do we not see objects of other colors such as roses, grass, violets,
oranges, and so on ? The answer is that we do see all these things
but that they do not give us the color sensation correctly belonging
to them; their colors appear to us as varieties of the other color
sensations which we are able to receive. Take for example the
color red. A soldier's coat or a stick of sealing wax conveys
to me a very positive sensation of color, by which I am perfectly
able to identify, in a great number of instances, bodies of this hue.
But when I examine more closely what I really see, I am obliged
to conclude that it is simply a modification of one of my other
sensations, namely yellow. It is in fact a yellow shaded with
black or gray, a darkened yellow or yellow brown."

Dichromatic vision occurs in three forms, in two of which red
and green are not differentiated from one another. The three
forms are named protanopia, deuteranopia, and tritanopia respec-
tively. In protanopia the red end of the spectrum is shortened;
that is, a portion which to the normal person is red, appears black.
The remainder of the red, the orange, the yellow, and the green
appear as successively lighter shades of yellow which, toward the
blue, becomes gray or white. This white shades into bhie which
deepens toward the violet end of the s])e( tnim. \n deuteranopia,
which is the normal condition of a jHTipheral zone of the retina,
the red of the spectrum is not shortened. Red, orange, yellow
and green appear as lighter shades of one color, called red or
yellow, and shade into a white or gray band which is a little nearer
the red end of the spectrum than the corresponding band of protan-
opia. Blue is perceived normally. Tritanopia is a rare form in
which yellow and blue are not recognized. The spectrum presents
red and green portions, separated by a white band in place of the
yellow. A dark green is seen in place of blue and the violet end

No. 486] THE PROBLEM OF COLOR VISION 375

up by blue light. As shown in the figure, orange is a mixed sensa-
tion due to the simultaneous partial destruction of the red-green
and the yellow-blue substances. Yellowish green and greenish
blue are likewise mixtures, and violet is supposed to combine the
partial construction of the yellow-blue with the destruction of
the red-green, the latter being indicated by the broken line. There
are four pure sensations, red, yellow, green, and blue. Color
blindness may be due to the absence or deficiency of the red-green
substance (protanopia and deuteranopia, the two forms being
varieties of a single type), or to lack of the yel]ow-l)lue substance
(tritanopia). Ilering further considered that there was a white-
black substance, built up in darkness to give rise to the sensation
K-g yh w

376

THE AMERICAN NATURALIST

[Vol. XLI

cists recognize that only three are necessary. Accordingly the
physicist Young proposed a simpler theory antedating that of
Hering. It was advocated by Helmholtz, and is generally known
as the Young-Helmholtz theory.

According to the Young-Helmholtz theory there are three photo-
chemical substances, red, green, and blue respectively, which are
stimulated by the various rays of the spectrum as shown in figure
4. Absence of stimulation produces black, and the simultaneous
disassociation of all three yields white. Protanopia is interpreted
as red blindness, due to deficiency of the red perceiving substance.
Deuteranopia is green blindness, and tritanopia is blue blindness.
Since it would appear that the })ercepti()n of white must be lost

with the disa])pearan('e of one of the three elements, the theoi
has been variously modified, in j)n)t;nio[)ia The red and the gree
substances may We so iiltered that each fe^poiiiU hoth to red an
green light iFieki, or the vvi\ and the ui-eeii siihstances may I
imperfectly segregated, as assumed by Mrs. Franklin's theor
The close relation between the red and green substances is shov,
in Koenigs presentation of the Young-Helmholtz theory (Fig. 5
The absence of either would give rise to somewhat similar cond
tioiis. siieli as occur in protanopia and deuteranopia. The figui
iraheates that in triehroiuatie vision, the colors from vellow i

No. 486] THE PROBLEM OF COLOR VISION 377

intersection of the blue with the green and red curves respectively.
In the absence of the blue substance, the white band is near the
yellow. This accords with the observations upon the color blind.
The absence of the green substance would not shorten the spec-
trum, but the lack of the red or blue would cut off their respective
ends. All of these features are equally well explained if, instead
of the absence of one of the three substances, such a modification
of its reaction is assumed as would be illustrated by a lateral
shifting of its curve in the diaoram. Thus in red blindnf>s the
red curve is shifted to cover more closely tiie territory of the ureen;
in green blindness the green is shifted toward the red; and in blue
blindness the blue and green curves are brought together. Thus

in the color blind all three substances are pres
form. Since this modified Young-Helmholtz
well with observations on color blindness, it is o
as the most satisfactory explanation of color \
An interesting attempt has been made by 1
theory into relation with structural elements
believes that thcvisual cells of invertebrates a
a fibrillation which is transverse to the direct
light waves, and that the tendency of the v
cones to separate into lran>verse dix - is evi
structure. Many hundreds of >nch fil)rils n

378

THE AMERICAN NATURALIST [X'ol. XLI

to act as 'conductors or resonators,' a fact which would not exckide
chemical changes resulting in fatigue. The long fibrils respond
to the red end of the spectrum and the short ones to the blue. In
Mjual length and only monochromatic vision
; their varying length allows a range of
)n in the form or dimensions of
the cones would bring about cor-
responding changes in vision.
The increased length of the
cones at the fovea provides for
a greater power of color discrim-
ination. If the base of a cone
were absent or cylindrical it would
be red blind.

This theory is illustrated in fig-
ure 6. On the right is the dia-

tlie latter radiate from an axial
lilament, the existence of which
has been discussed and denied
by other investigators. The fi-
brils in the right half of the cone
are drawn as responding to red,
yellowish green, and violet light;
the Young-Helmholtz curves are
shown on the left. In nonpolar-
ized light all of the fibrils in a

spond uniformly, but in polarized
liiiht only such are effected as are
I the left of the figure. Thus the
,,hmu-.l. The correctness of this

It will be noted that
theories the mechanism
a single cone. The lie

No. 486]

THE PROBLEM OF COLOR VISION

theory, although the three substances could exist in a single cone,
each is declared to exist in a cone hy itself. This is considered
to be strongly in favor of the vah(ht_v of llic ^'ouno-I lehnholtz
theory. Since physiologists find no instance in wliich different
sorts of impulses are conveyed over a given nerve fiber, it is believed
tliat a single cone fiber can transmit only one .sort of color M-nsition.
The stimuli of the red, green, and blue cones respcc ti\cly arc
supposed to be gathered l)y .separate nerve cells of the retina, and

only il.l. otvcn. and blue sei'isations respectivel v" I1ie nn'xing of
the MMi^..ti..n.. ^i^inn rj.r to the perception of sha.lo and tints, is
tiuM-efore acconiplisiied in the brain and not in the cones. In an

perception of the colors of stars. The image of the star- is so

that one cone perceives its color is invalidated by the fad that the

rapid .succe.s.sion upon .several coiit>> which may unite in uivinu" the
color perception. Those who bcucve in the s|.,vihc cueruv of the
rod and cone fibers dismiss at oikv several .if the iheori.'. of color
vision. It must be nMnenibered. however, that the separation of
the cones into forms respoudinn- to re.l. blue, and oivm light,
with three corre.sponding sets of nerve cells and fillers to convey
these separate stimuli to the brain, does not rest upon anatomical
evidence.

Physiological Laboratory
Harvard University

THE BPiKKDIXC IIAHirs OF AMHIA'STOMA PiWC-
TATl'M LI NX '

382

THE AMERICAN NATURALIST [Vol. XLI

stituting the base must be strongly adhesive when fresh, for the
spermatophore is firmly attached to the object on which it is de-
posited. The cap is usually hemispherical in form, with the con-
vex surface upward; but the material of which it consists often
runs down the side of the stalk, or is found projecting in downy
tufts like the cotton from an open cotton-boll. In many cases
the caps have a frayed appearance, as if they had been disturbed ;
in occasional specimens the cap of spermatozoa is partly or almost
wholly absent. The appearance in the latter case is like that of
a spermatophore of Triton viridescens from which I have seen the
ball of spermatozoa taken up into the cloaca of a female. The
dimensions of the complete spermatophore are about as follows :

Height 6-8 mm.

Breadth of base G-8 mm.

Diameter of stalk near top. . .2.5-3 mm.

" " cap 3-4 mm.

As compared with some spermatophores of Triton viridescens
obtained from specimens in captivity, these under discussion are
slightly taller, with a smaller base and a stalk of much larger
diameter, surmounted by a larger mass of spermatozoa. The
spermatophore of Triton viridescens has a broad flattened base
from the center of which rises a distinctly conical stalk tapering
to a very slender spine, at the top of which aitaclicd a small ball
of spermatozoa; the spermatophores attnhntc<| lo Aniblystoma

When found on April 9 and 10 the spermatophores were all in
Kood <()ti(htion, with some slight differences in the freshness of
tlieir appearance. In two or three days they became infested
with fungus, disintegrated quite rapidly, and in a week very few
few of them could be found. Had new ones been deposited in the
interval, they could readily have been distinguished from the
old ones; but no more spermatophores were deposited. Hence
it is scarcely possible that the period during which spermatophores
are deposited lasts longer than two or three days.

I'he spermatophores shown in the figure had been attacked by
fuuirus and were beginning to disintegrate when photographed.
The hasc is thcn'fon" no lonucr clear, but on the contrary the

No. 486]

HABITS OF AM BLY STOMA

383

Ident{ficatio7i. In order to identify the spermatophores, search
was made for the parent animals. This resulted in the capture
on April 11, of three specimens of Amhlysioma puncfatum Linn,
which were found eml)edded in rotten wood under a stump at the
edge of the water of one of the ponds where the spermatophores
were numerous. From two of these specimens a few drops of
seminal fluid, containing an abundance of spermatozoa, were
obtained hy stripping; from the third, which proved to he a female,

spermato})h()rcs and similarly treated. In >tni(tutv, >i/.c. ami
staining reactions the two were iilentieal.

Another .species, J. tl(jrl)ntiit (ireeii. alx) (iccuin in the vicinity

384

THE AMERICAN NATURALIST [Vol. XLI

of Ann Arbor, and a single example was taken on April 9, in a
field several hundred yards distant from the nearest pond where
spermatophores were found ; but the eggs of the two species are
easily distinguishable, and in the case of A. pundatum were iden-
tified by means of eggs laid in the laboratory. With the exception
of one bunch of eggs of A. tigrinum, all the eggs found in the pond
where spermatophores were observed, were those of A. pundatum.
With the single exception above noted, the two species have not
been known to breed in the same ponds in the vicinity of Ann

B. The Spermatozoa. The spermatozoon of Amhlystoma punc-
tatum is extremely long and slender. The head stains well with
Delafields' haemotoxylin, the middle-piece less deeply. The tail-
piece is bordered on one side by a very delicate undulating mem-
brane. Some of the dimensions are as follows:

Length of acrosome 20 /x

"head 106

" " middle-piece 14 /x

" tail-piece 480 /x

Total length 620 //

The spermatozoon resembles in size and form that of Triton
viridescens, with which it was compared, but the latter has a
middle-piece twice as long, and a more conspicuous undulating

As compared with the spermatozoon of Crytohranchus allcr/hr-
niensis (Smith '06) the sperm of Amhlij.stoma pundatum is nearly
three times as long, with a proportionally much longer middle-
piece; the entire structure is much more slender and thread-like.

In freshly mounted seminal fluid the spermatozoa were seen in
active motion. They tend to cling together parallel to each other
to form bundles or ringlets, revolving with a circular motion;
when so clustered tliey retain their vitality much longer than when

It -nHhially hwumv, .l..uer until with a InVI, nia-nification it is

if form across the entire fi.'M <.f the niicn.sc.pc. Tl^. un.lulatino-
membrane <loes not wind alx.ut the >liaft as in ('ryi.t..l)ran<hus,
but continues on one side of it. When dead, the sperms are usually

HABITS OF AMBLYSTOMA

found much convoluted, indicating a greater degree of flexibility
than is the case with stouter spermatozoa like those of Crypto-

Experinients were performed to determine the length of time
the spennatophores would retain their vitality in water, hence the
interval within which they would have to be taken up by the female.
In all the spermatophores examined the spermatozoa were motion-
less; but since the examination was not made until the evening
of April 10, probably the spermatophores had been in the water
for many hours. The effect of the cloacal secretion of the living
female was then tried, to see if it would revive these spermatozoa;
no such result was produced. Freshly obtained seminal fluid
mounted in water retained its vitality for many hours; but as this
experiment was not performed until April 18, only a small amount
of seminal fluid could be obtained, and in this the sperms were not
m a vigorous condition. It fresh seminal fluid were taken in the
proper season and mounted in quantities to correspond with that
deposited in a sperniatophore, it might retain its vitalitv much
longer. I'he viscous lupiid in which the spermatozoa occur does

V fHsliK ,]q>„s,u,l sp(rnuitophoie ot J nfon um/oa//s ^^as

tlu bill n\ .,,(,!.,,!.. . . uti. t. i..d .| .If in.l (xun! I 1 hi
tl.(> n.icn.s.-niK-. K\rvvu Unuvs fn.i.i th. tiui, tlu .[x nnatopliore

tophore thui ^n^\n^ ^^nuU\ In Ktiuud l..n.( i tli ui r^^(l^< lioui

C. The Eggs. Those of A. punrfafiim have hvvu dcscnlu'd
and hgurul In ( laik (SO) Uu < uith th<n iii.hM.h ,1
gelatinous envelopes, occur m conipact buii( li(>s, MirioiiiHU'd hv
axerx tlmkidh mass Thetntiu Miiutint i> u.ii ,lh ot w omI
sh.ipt <)tltnn«ail\ is laigt n.u . h t Hu .>t J f ,/> nam

ai( , ii.h lu^.i ih... lh(.M ..t I In/, mum ind du nninh . ot
cgn-s „, a bmich IS ii.iiallv n-iva1.T.

Vt th( tiiiu ot i\u <I.M< XMN ot du .pu.n.tnphoH ^n^ U^^
biMKhont (ould In tuuu.l Ih. iiini.lKi uhK iih k , <d

for a week; at the eiitl ot that tune eggs were iound in early seg-

386

THE AMERICAN XATURALIST [Vol. XLI

mentation stages, showing that thev had been quite recently laid.
The egg-laying season follows immediately after the deposition of
spermatophores, and lasts six or seven days. Nearly every bunch
of eggs found on April 10 was close to a group of spermatophores.

On April 16, in the pond where 25 groups of spermatophores
had been counted nearly a week before, about 55 bunches of eggs
were found. Of these, many bunches were deposited in groups of
two to four, probably by the same female. The number of aggre-
gations of eggs very nearly equalled the number of groups of
spermatophores.

D. The Adults. Secondary Sexwil Characteristics. During
the breeding season, at least, the cloacal region of the male is quite
prominent; that of the single female examined was much less
swollen, and the orifice was smaller. The cloaca of the male is
lined with fine parallel papillated ridges, extending inward for a
few millimeters; between these ridges are deep grooves, lined with
cilia whose beat is outward. These ridges and grooves were not
found in the single female examined. According to Kingsbury
('95) the female Amblystoma, as well as the male, has cilia in the
cloaca but the tract is less extensive. The urogenital sinus of
the male is larger than that of the female, probably to hokl a con-
siderable supply of seminal fluid preliminary to the deposition of a
spermatophore. No secondary sexual cliaracters to indicate the
clasping of the female by the male were found.

II. Discussion.

Andrews ('97) described the structure and distribution of some
spermatophores which he attributed to A nibli/sfonia piiiirtnfvm, but
without positive identification. He states tliat tiiesc spermato-
phores were more slender and higher than those of Triton virid-

No. 486] HABITS OF AMBLYSTOMA

On account of the late season at which my investigation was
begun, no direct observations of the process of fertilization were
possible. Clark (79) says of some specimens of A. pundatum
m conhneinent: "The males showed no inclination to clasp the
females, but (juietly deposited quite large masses of an apparently
rather thick liquid, opaque white, on the bottom of the dish in
which they were kept. Upon examination this was found to con-
sist of spermatozoa moving actively in a licjuid." The manner in
which the spermatozoa reached the eggs was not observed.

Fertilization is undoubtedly internal. Of this the evidence
adduced by Kingsbury ('95), and the presence of spermatozoa
in the cloaca of the female as described above, furnish sufficient
proof. It remains to consider how the transfer of spermatozoa
is effected by the spermatophores.

The number of spermatophores is evidently very much greater
than the number of females; and unless there exists an enormous
disproportion between the sexes, each male must deposit a large
number of spermatophores. Their abundance and the manner
of their distribution, render it a very easy matter for the female
to find enough of them for purposes of fertilization. In some
portions of the pond it would seem scarcely possible for a female
to move about in the water for any length of time \\ itiiout l)rnshing
against some of these spermatophores; hence tlu>re is the possi-
bility of finding them bv chance contact.

In the ca^-s nf thos,-' rrodela in whic-h, as in Triton r/r/./r.™
(Jonlan -HI and Ilihon an.l Ax.)lotl ((iasco M) the

number of spermatophores <lep.)sited In- a single male is small,
particular safeguards are needed in order to facilitate their delivery
to the cloaca of the female. In tlies(> forms the physiological
necessity which re(|uires the co-operation of the female in order
that spermatophores mav be deposited insures the presence of the
female at the right time; su!.st>(|nent reactions safeguard the

the spermatophore. The complicated behavior of the adults in
these cases finds its biological significance not only in the increasing
certainty of the process, but in a corresponding economy in the

388

THE AMERICAN NATURALIST

[Vol. XLI

number of spermatophores that must be depositerl. "With Amhly-
stoma pundatum, on account of the very large innnber of sperma-
tophores, there is the probabiHty of a simpler mode of behavior,
and the spermatophores may be found lar<jely by chance. The

Axolotl there is evidently no clasping- of the female by the male,

"Vhv result of the ex])erimental work on the vitality of the seminal
fluid in water indicates that the s])ermatophore is not necessarily
taken uj) by the female immediately after it is deposited; probably
it is capable of effecting fertilization after exposure to the water for

On account of the shortness of the breeding season, the sperma-
tozoa can be retained in the cloaca of the female for only a few
days at most, before fertili.-.ation is effected. The position with
respect to the spermatophores, of the earlier eggs fouiul, .suggests
that in some cases the eggs are deposited inunediately after the
spermatophores are picked up.

The extreme flexibility of the sperm is doubtless correlated with
the process of internal fertilization. In Cryptobranchus, in which
fertilization is external (Smith '07), the egg envelopes nuist be
penetrated after a brief exposure to the hardening elVect of the
water, and a much more riuid spermatozoon is nM|uir(Ml.

In the evolution of terrestrial from a.|natie vert<4.rate \Wv, a
transition from external to internal fertilization takes place.
External fertilization is not ada[)ted to terrestrial conditions,
hence in the land-living vertebrates it occurs only in some of the
forms that revert to the water during the breeding season — r. e. in
the Amphibia. Internal fertilization is an ada{)tation to terres-
trial life in the sense that it is a condition aiU(>ce(lent to that life,
not a result brought about by it; it may occur in purely aquatic
vertebrates, as in the Elasmobranchs and a few Teleosts. Internal
fertilization by means of spermatophores is a method still adapted
to aquatic rather than to terrestrial conditions. It is a method
intermediate between external fertilization on the one hand aiul
internal fertilization withont spermat()])li()res on th«' other. \'iewe(l
in the light of tli(> linhil. of the ]w^hvv vertel.rale.. the oecnrrence

Ko. 486] HABITS OF AMBLYSTOMA

water represents an advance upon the habit of external fertihza-
tion, and a stage in the evolution of habits that are to make possible
the invasion and permanent occupation of the land.

Internal fertilization also finds a biological significance in the
fact that in the course of its development there is gradually effected
an economy in the amount of seminal fluid required for fertiliza-
tion. This factor may account for the persistent development of
the habit under aquatic conditions, where external fertilization
is still possible; the incidental result is a preparation for terrestrial
life.

In existing Amphibia we may find illustrations of various stages
in this evolution of the breeding habits correlated with a transition
from the water to the land. In Cryptobranchus, one of the lowest
of the Urodela, leading an aquatic life and showing only in its
methods of respiration and locomotion an advance to\\'ard terres-
trial conditions, external fertilization takes place. This is evi-
dently the primitive condition for the Urodela. In Amblystoma,
a urodele living partly upon the land but returning to the water
to breed, we see developed the peculiar habit of fertilization by
means of sperm atophores — a mode of internal fertilization favored
by aquatic conditions. In Triton viridrsrrns an economy of
seminal fluid tiirough a reduction in the nuinl)cr of sperm;itoi)h(>re.s

Savi., Molgc a.spcra Duges and GlossoUga lla(jcnLdlvn Latastc.
according to Bedriaga ( '82 and '95) the male emits spcruiatopliores
while still clasping the female; in Triton iorosus Escli. (Ritter '99)
it is probable that a very similar process occurs; in none of these
cases, with the possible exception of Molge aspera, is there direct
cloacal contact. Finally in the Apoda (the Sarasins '87-'93;
Brauer '97) we find the establishment of a method of internal
fertilization bv direct c-loacal contact, thus fulfilling the require-
ments for continnons residence ii})on the land.

390

THE AMERICAN NATURALIST [Vol. XLI

THE STAFF-TREE, CELASTRUS SCANDEXS, AS
A FORMER FOOD SUPPLY OF
STARVING INDIANS

FRAXK T. DILLINGHAM

In many kinds of hard and horny seeds there is present, as a
reserve material, a carbohydrate which upon hydrolysis yields
mannose (a simple sugar closely related to glucose). This carbo-
hydrate has been named mannan. It is one of the hemi-celluloses,
a group of substances closely resembling in appearance the true
celluloses, but easily resolved into simpler carbohydrates by the
hydrolytic action of enzymes or of dilute acids. There is no lack
of evidence that mannan which occurs abundantly in the so-called
vegetable ivory, Phytelephas macrocarpa, and in the seeds of many
other palms, as well as in the wood of coniferous trees, is in spite
of its hardness, fit food for camels, neat cattle, sheep, and various
rodents. This is illustrated in the girdling of pine trees by mice,
as recorded by Thoreau in "Walden."^ He says:— "There

inches in <lianirtrr, ul!ich lia<l been unnued by mice tl.r ,.re\inns

deep, and thev uo.v cbliool to uiix :i hn-c proportion of pine
bark with their other diet. The^e tree^ wore aliNc an<l apparently
flourishing at mid-summer, and many of them had grown a foot,
though completely girdled; but after another winter such were
without exception dead. It is remarkable that a single mouse
should thus be allowed a wliole pine tree I'or its dinner, gnawing
round instead of up and down it; but perliaps i( i.> ntM("->an ni
order to thin these tn^N. which are wont to urow up densely.

It is known that the root of a .lai)anesf j)huit, ( 'oiioplialliis
komijaku, rich in mannan is used as human tood, and tlie (piestion
may fairly be asked whether the former n.e of bark brea.l by the
inhabitants of Scandinavia miglu nor ha\e been deix-ndent up<m
the mannan in the bark. After di.cu^Mn- thi. matter in the

'Walden, p. 300. Jas. R. Osgood & Co. B<.>f(.n. IMH

392

THE AMERICAN NATURALIST

[Vol. XLI

Bulletin of the Bussey Institution (1906, Vol. 3, pp. 120-128),
the writer learned that some tribes of North American Indians in
times of extreme dearth were accustomed to keep body and soul
together by boiling and eating the bark of the Staff-tree, Celastrus
scandens. The Staff-tree is also called the staff-vine; false,
climbing or shrubby bittersweet; wax-work, fever-twig, yellow-
root, climbing orange-root and Jacob's ladder.

Radisson, wintering near the outlet of Lake Superior about
the year 1G58, found the Indians suffering greatly from starvation
He writes:'— "Those that have any life seeketh out for roots,
which could not be done without great difficulty, the earth being
frozen 2 or 3 feet deep, and the snow 5 or 6 above it. The greatest
subsistence that we can have is of rind tree which grows like ivy
about the trees; but to swallow it, we cut the stick some 2 foot
long, tying it in fagot, and boil it, and when it boils one hour or
two the rind or skin comes off with ease, we take and dry it in
the smoke and then reduce it into powder betwixt two grain stones,
and putting the kettle with the same water upon the fire, we make
it a kind of broth which nourishes us, but become thirstier and
drier than the wood we ate."

In the Report of the U. S. Commissioner of Agriculture for 1870,
(p. 422), there is the following statement:— "The Chippewa
Indians use as food the tender brandies of the Stuff tree (Celastrus
scandens). This climbing shrub, the hoi.s- rdor.s of the French,
or twisted wood, is sometimes called hhirr sweet. It has a thick
bark and is sweetish and palatable when boiled."

In view of the above statements, specimens of both the bark and
the wood of the Staft-tree were tested for mannan. On the
grounds of the Bussey Institution, on Jan. 24th, 1907, branches of
the Staff-tree were cut in pieces about one foot in length. Both
the inner and outer bark were removed together, no attempt
being made to separate them. The outer bark was thin, but the
inner bark was thick and fleshy. The material was carefully
<lii<(l, ground to a fine meal, and a weighed (|uantity of it was
l)()iltMj with dilute hydrochloric acid for three hoiu-s. A small
portion of the liijuor thus obtained was neutralized with sodium
hydroxide and examined for mannose by the addition of a few

' Voyages of P. E. Radisson. p. 204, Prince Society Edition, Boston, 1885.

No. 486] THE STAFF-TREE AS A FOOD SUPPLY

393

drops of ph.enylhydrazine acetate. No niannose hydrazone crys-
tals formed at this ])oint. 'J'he remainder of the hquor, after l)e-
ing similarly neutralized, was evai)orate(l to dryness; the residue
was treated with a small .|nantity of water; and the concentrated
li(luor thus ohiaincd was tested for nianiiosc hy addini: a few drops
of phenylliy.h-a;:inc .•.<rtatc. With the i.i.l" of tl.c microscope,
the formation of characteristic crystals of mannose hydrazone
was observed.' The wood proper (including the pith) was re-
duced to a fine meal and then treated in precisely the same manner
as was the l)ark.

From these tests it apj)ears that unlike the bark of most decidu-
ous trees, that of the Staff-tree contains an abundance of mannan.
The bark of the Staff-tree, moreover, contains a larger quantity
of mannan than does the wood proper.

To confirm Radisson's statement as to the effect of boiling, a
few branches of the Staff-tree were boiled with water for about
one hour. At the end of this time the bark was found to peel off
with great ease. It was seen to be thick, pulpy, and very mucilagi-
nous, and it had a rather agreeable taste.

It is evident from these experiments that a part, at least, of the
physiological value of the bark of the Staff-tree may be justly
attributed to the presence of mamian.

NOTES AND LITERATURE

GENERAL BIOLOGY

The Spirit of Nature Study.' — Nature may be approached in a
very unscientific spirit. Thus Emerson was led to dedicate to the
Bofanist the folloAving quatrain:

Do thou of the Ages ask
What me the Hours will bring.

What the hours brought he so ex])resse(l that tli(^ Rhodora has become
a universal type of Ix^tanic bt^aiity. In a different spirit the New
England botanists named their join-nal Rhodora, for they profess
to have been uninfluenced by Emerson's familiar lines; they sought
a characteristic local plant with a short name which would commend
itself to bibliographers. The spirit of nature studv, accorduig to Dr.
Bigelow's interpretation, combines the seiuuncnt.il .irid the scientific,
with its emphasis upon the former. This ;ij)|H'ars iii such iHi\ ice as, —
"Take frequent rambles into the countrv; nsxxmu' wuh natural

rcnicmbranccs ot them. . . . Mil)sr(|nciu vcars ot trouble cannot oblit-
('rat(> the clianiicd niij)rcssi(.ns.' - At ilic iic.M stopping place there
will !)(■ no taircr huidscaiH-. imr inoiv i.caimtiil skies, no statelier
tn... nioi.- loNon. m,,,.,,..^ ik.i l.ii^ln. i tluuers; more cheerful

Therefore nuuv time sh.ml.l I.e uivcn t.. natinv study in the schools,
and many educ-ators niv (pi.u.d to ilii> vWvvx. School children should
be taken to the couiitrv and should have |)laiits and animals at home;
rabbits and gounls arc parlicularlv rccnimufnded since the former
are reasonably small and the latitM- urow iipwai'd 'where land is cheap.'
College methods ot instruction slioiild not he extended to el(>nientarv
schools. Of the sixte(-ii half-tones uliich illiistrat. tlu hook tuehe
are photographs of hovs and irirls out m

396

THE AMERICAN NATURALIST [Vol. XLI

as ornithological specimens, nor wild flowers as material for herbaria.
Dr. Bigelow's expeditions are not for 'what one can get.' It is ap-
parent that within the present century the destruction of such irre-
placable plants and animals as remain will not be tolerated, either for
sport, for food, or for amateur collections. Since nature study in the
schools should save the swallows' banks from the small boy and
protect native plants from bouquet gatherers, it may prove of great
value to the community. Tliis, however, is not strongly presented in
Dr. Bigelow's book, which iiichides a photograph of eleven women
gathering bunches of violets, and recounts, as one of the author's
pleasurable rcnuiusctMKvs, the bloody death of a woodchuck.

Dr. Bigelow's t\venty-thir(^ informal essays are enlivened by many
quotations and anecdotes; their author ai)i)rc( iat<>s th(> "fun of being
a naturalist" and his good natured lunnor is all at the expense of the
"bug-hunter's" critics. He is at home with hoys and girls for whom

in St. Xirho/d.s; hilt the book here considered is addressed to parents
Heterogenesis. 'The idea that eggs of one species may give rise

' Bastian, H. C. The evolution of life. Reviewed in \ature May 2 1907
vol. 76, p. 1.

Son & Co., 1907. "Svo, ix+ 103 pp., 5 figs.

398

THE AMERICAN NATURALIST [Vol. XLI

the dorsal ixincreas acrrssort/ pannra.s\ the bias of human anatomy is
apparent. In this rcsprct Professor Wilder's rejected principle ought
not to be abandoned. 'J'lie (iernian committee has adopted thirty-
four of the forty terms sanctioned by the American Neurological
Association; in ten of these, however, nouns which had been dropped
as superfluous are retained. In general, the committee declined to
introduce new terms, to combine nouns and adjectives, or to eliminate
syllables or letters for brevity. It followed Professor Wilder's advice
in preferring descriptive to personal names, definitely retaining only
two of the latter, Wolffian and MueUerian. Intestinal glands, parotid
duct, splenic nodule, and renal corpuscle replace glands of Lieber-
kuehn, Stenson's duct, and Malpighian corpuscle, the last term having
been applied to radically different parts of the spleen and kidney.

After a trial of more than ten years this anatomical nomenclature
adopted at Basle, and known consequently as the BNA, has become

This

• Barker of Jol

i Hopkins

use whatever English efjuivalent he desires for the ol
Students are strongly advised, however, to use t
terms as English words. The Latin terms are tli
ones." We agree with Profcs.sor Barker that "th(
decision to adopt these terms is reached, the better

: and Overlap of Instincts in Birds.'

Ko. 486]

NOTES AND LITERATURE

including (1) spring migration; (2) courtship and mating; (3) nest
building; (4) egg laying and incubation; (5) care of young; and (6)
fall migration. Some birds, like the robin and blue bird, pass through
two or three reproductive cycles before the fall migration. Tlie fish
hawks and eagles which repair their oki nests in the autumn do not
act in "anticipation of spring" but exhibit a recurrence ot" ilie iu',stin<j:
instinct, due to beginning a new cycle which is never finished, ^'oimi:
birds may be abandoncnl in the fall when the migratory impulse »>v(m--
laps the parental instinct. jiduit r()l)in lias been seen to offer a

string to its fully gr()\\ n youno-. and try to cram it down the throat of
a fledgling. Later the old bird llcw with the string into a tree. This

the cowbird's egg, which it does so j)erfectly. It indicnics rather that
the reproductive cycle has been broken 1)\ tear, and a new one i>
begun, in these rare cases the old nest beinjr n taiufd a- a site n» hinld
upon. The herring gull also will bury it. u hm n. . ^. Ir ha- 1.. vu
interrupted through iVur.

followed in th,>.f(Mn-.Kal..f ( onll.al^l(iv.■^^■nI■..l(>oyl,van.■xIra(.^linarv
/J//^^/,»v all,<ih,nH„s,^ ' It Mate, that "afn-r iIh' .'l-^ arc drj-o^nd
they arc nsually unardcd for a lime by the male, who fi^'his and (h'lv,-

himself eats some of the eggs, bnt on' accouiU of the sKmn.-.. of hi^

is onlv guarding his own food supply: the origin of tlie l)roodiiig
habit in this case .seems to he the feeding habit." If .)ne d..nbts that

Journ. of Comp. Neur., 1907, vol. 27. pp. ](»7 1!>S.

400

THE AMERICAN NATURALIST

[Vol. XLI

A Preliminary Note on the Variation of Scutellation in the Garter
Snakes.— Three years ago the writer began an investigation into the
relationships of the different races of garter snakes (Thannophis) in
an attempt to determine the laws involved in the differentiation of
the genus. The results of this work are being included in a mono-
graph of the genus, but as it will be several months before this work
can be completed it has been thought best to publish a brief outline
of some of the conclusions.

In the progress of this investigation it was seen very early that before
a serious attempt could be made to determine the affinities of the
different races, the significance of the variations in scale arrangement
or scutellation must be determined. This was attempted with the
following results:

(1) The number of dorsal scale rows on an individual snake
decreases posteriorly by the elimination of certain rows, and
the series eliminated are always the same for snakes with the
same number of rows, as for example T. sirtalis and T. saurita.

(2) The rows dropped posteriorly in individual snakes are those
which have entirely disappeared in races with a fewer number
of scale rows.

(3) The reduction in the number of dorsal scale rows is generally
accompanied by a reduction in the number of labial, ventral,
and subcaudal scales (gastrosteges and urosteges).

(4) There is considerable evidence that the reduction in scutella-
tion is directly or indirectly associated with a diminution in size.

The general reduction in scutellation described above is exhibited
by each of the several (natural ?) groups into which the genus can be
di\ided. These groups all occur together only in northern Mexico,
which may be considered the center of origin for the genus. The
races that occur in this region all exhibit the maximum scutellation
for their respective groups, the dwarfing in size and scutellation
taking place at points away from the center of origin. The dis-
covery of these methods of variation in the different series of scales
has been an indispensable aid in determining the affinities of the dif-

A Simple Method for removing the Gelatinous Coats of Eggs.—

of amphibian eggs tlie writer liir iij)i>ii a -iinph' jind ra{)id method of
freeing them from their irclannous eiiv( l()iH>. While, because of its

402

THE AMERICAN NATURALIST [Vol. XLI

dellids with which group (or the Histriodrilids) the author is indined
to place it. The internal structure was not studied.

The Sijstematw Position of TricJtoplax. Exer since its discovery
Trichoplax has been one of the zoological problems, and now Thilo
Krumbach of Brcslau offers evidence^ to show that it may be the
planula of the hydroid Eleutheria. His proof is not conclusive but is
based upon the histological similarities between the ])]aiiula and
Trichoplax, and upon the fad that Triclioitlax :i|.p("are.l suddenly in

He suggests .mIm. lh..t M..nti.vlli. Tnptup/nr n ptan^- l.clnno. to
Eleutheria clapamli.

Caesar Bottger report- r,lii,nh, plmladi jnnm^ Irom the North
Frisian Islands, and (luou . .,1m. ii. pn-cncv tn.ni the Kast Frisian
Islands. It has previonsly only Ikh'ii known from the Atlantic coast

'Zool. Anzeiger, 31, p. 268, 1907.
»ZooI. Anzeiger, 31. p. 291. 1907.
*Zool. Anzeiger, 31, p. 296, 1907.

No. 486] NOTES AND LITERATURE

403

BOTANY

Cytology and Mutation. - Immediately after tlie rediscoven of
Memlers law and \hv |.ii!)licati()ii of DeVries's great work on mnta'tion,

organization of the germ cells. The most recent contribution to ih(>
literature of this subject is a paper by Gates^ on (Enothem L(im<irrkiaiin
and O. lata.

The author finds that the regular abortion of the pollen iii (Enofhcm

growth of iis lining (the tapetum) as described by Pohl, but to some
other agency the nature of wliich is not yet explained. Pollen (level-

1 Pearl, R. Variation and DifYerentiation in Ceratophyllu.n. Carnegie
Institution of Washington, 1907, Publ. 58, 136 pp., 2G figs.. 2 pi.

406 THE AMERICAN NATURALIST [Vol. XLI

Notes and illustrations concerning Robinia Neo-Mexicana are pub-
lished by Phillips in Forestry and Irrigation for February.

An illustrated economic account of Nyssa aquatica, by von Schrenk,
has been reprinted from the "Silver anniversary edition" of The
Southern Luml>ennan.

Vaccinium Dohhini is the name proposed by Bumham in Jlie
American Botanist of Fel)ruary for a New York relative of V. vacillans.

A revision of Spilanthes, by A. H. Moore, constituting no. 33 of the
new series of "Contributions from the Gray Herbarium of Harvard
University," is published as vol. 42, no. 20 of the Proceedings of the
American Academy of Arts and Sciences.

A paper on Citharexylum, by Greenman, forms Publication 117 oi
the Field Columbian Museum.

On Pringle's Santa Catalina Mountain material of 1881, Dode
bases a new Juglans ela^opijn n in tlic Ihillcfin de VHerbier Boissier
of February 28.

An economic account of the walnut in Oregon is published by
Lewis in Bulletin no. 92 of the Agricultural Experiment Station of that
State.

A new Californian oak, Qucrcus Pricei, is described by Sudworth
in Forestry and Irrigation for March.

Several new aloids ami otlicr succulents are described by Berger in

urns zu Berlin.

Agave desert i is fion.v.l in detail in Icones Selectee Horti Thenrnsi.s;
vol. 6, fasc. 1.

A series of notes on Ahietinea-, by Hickel, are appearing in the
Bulletin de la Snriitr I)n,drnhn,i^inr <le Frmwe.

Cardotand Tlu'riot rcju-rt on a collection of C,:", Ala>kan ino.ses in
vol. 2, no. 13 of the rnirrr.ih, njCnIijnrnia Puhlimtinn.. Bntan,,.

No. 486] NOTES AND LITERATURE

407

An enumeration of the fungi collected by Simmons on the second
Norwegian Polar expedition, by Rostrup, was published in no. 9 of
the Report on the Expedition shortly before the death of the author,
which occurred in January.

Several quite distinct puff balls and ])lialli)i(ls of Arircntiiia urc
described and figured by Spegazzini in a \y.\\)rv ivcuily disi lihutcd
from vol. IG of the Anales del Miiseo Nacioinil </r liiinm. Ai,r.s'.

A flora of Central Europe, with text cuts and colored i)lat('s, by
Hegi and Dunzinger, is being issued in 70 monthly parts from the
Lehmann Press of Munich.

With vol. 3, fasc. 7, issued in December, Coste's "Flore Descriptive
et Illustr^e de la France" etc. was brought to a conclusion, the final
signatures dealing with Pteridophytes.

An ecological systematic account of the flora of Columbia, r^Iissonri,
by F. P. Daniels, forms vol. ], no. 2 of the Scientific Scries of Tlir
Unirer^ify of Mis',oun Studies. TyUr n.u .p.-. i... or ^ari.-n... ..,,.1

species, of Pteridophytes and \'22 genera, with l().)'.i sprcu-., of
Spermatophytes.

A general biological study of the -and areas of Illinois, Wy Hart and
Gleason, forms vol. 7, article 7 of the nii/hlni nj the H/nmi. Sfatr
Laboratory of Nnfitral in.sfon/.

The distribution and a.lai.tation ot ihc N^^-tation of Texas are
discussed bv Brav in lixl/rhji im. S.' (Seieniilic Serie. n... 10) of the
University of Texas.

A studv of the flora (»f the Sand K<>v> of Florida, l.v M illsi)an<:h.
forms Puhlirntlun //\ of the Field C.hnnl.ian MuxMini.

408

THE AMERICAN NATURALIST [Vol. XLI

Studies on the pollination of AVisconsin flowers are being published
l)y Gracnieher in current numbers of the BuUeiin of the Wimnisin
Natural Hhtory Society.

Von Iherincr contributes an illustrated account of the myrmeco-
philous ( '(•(Topias to recent iniinbcrs of Engler's Botanische Jahrbucher.

Disscniitiation by tlir aid of ants is the subject of a well illustrated
memoir ly S.-niaii. 1(m-, forniinc:' vol. 41, no. 7 of the K. Svenska Veten-
sknpsnk,ulrwir>,s IIa,nl/inr,ar.

IVtcii, i^ p'ul.li-^l.r.l. uiili'illHMr.aions, in vol. 3, part 2 of the Anna'h

A long list of plants known to contain i)nissic acid is separately

Studies on tiie influence of spectral colors on the sporulation of
Saecharomyces are reported by Purvis and Warwick in vol. U, part
1 of the Proceedings of the Camhridcje Philosophical Society.

The root-knees of Somieratia arc well figured in the Annual Report
of the Director of Forest ni uj I he Philippine Islands for the Period
July, lOOr, to June SO, imn;.

A rope-like tumor of Hetulo pnpiilijniid is described and figured
by Penhallow in a separate from noI. 12 of the Transactions of the
lioyal Societi/ of Canada.

No. 486] NOTES AND LITERATURE

A series of "Forest Planting Leaflets," each dealincj with a sii
.species, is being jjublished as Circular.^ of the Forest Sen-ice of
United States Department of Agricuhure.

Studies of the wood of Javau trees, l.v Moll i.nd Janssoniiis,
being published by the lirill P.vss „f Leiden.

A second edhion of ih<' useful " Ke^ t.. rh.- (icnera ..f W.mhK VL
in Winter," by Wiegan.! and F uNwordiv. ha> been issued l.v
authors, whose address is Itha. a, X. \ .

Ilanbury is given in The CmhL^- Chmnu-h of March lOth.

A portrait of IL N. Ridley i. ^^Um in l^njura/ l.ije for January.

An appreciative notice of Mar-hall Ward, by the late Director
Kew Gardens, appears in Tlie Xnr l'/ii/f(>/<u/i.si of January 31.

Fascicle 4 of de Wildenian's " Knnineration .les Plantes Recolt '
par Emile Laurent," issued in Februar
bi..graphic sketch of Laurent.

Further articles on Burbank an<l his work, by DeVries, appear
the Bt'ohnps-rhes Crnlrafhiatf for September, The Open Court
Noventb.-r, an. I TheCeufun, Maqa-Jtir for March.

W. T.

rrEOLOGY.

The Elements of Geology.' - Professor Xorton of (

410

THE AMERICAN NATURALIST

[Vol. XLI

111 tlio order indicated. A hnal chapter in this part ot the book dis-
cusses off-sliore and deep-sea deposits. Under the heading "Internal
deological Agencies" the following chapters appear: Movements
ot the Earth's Crust, Earthquakes, Volcanoes, Underground Struc-
tures of Igneous Origin, Metamorphism and Mineral Veins. Histori-
cal Geology is treated in the usual manner, the principal systems and
some of their characteristic fossils being described in order, beginning
with the Pre-Cambrian. Special emphasis is laid upon the evolution
ot the North American continent and the evolution of life upon the
planet.

It is probable that manv will doubt the wisdom ot dropping out
structural geologv as a special subject and treating it onlv m connec-
tion with geological processes. There are difhculties in the way of
such a treatment, one being the danger that the elementary student
will not discriminate sufhcientlv between the process, the structures
diu to tht j)rocess, and tht tructures which mereh afiect the operation
of the process, all of which are treated under a single tit e. In the

different tactors involved, although in places a stronger (liscnininatioii

PUBLICATIONS RECEIVED

412

THE AMERICAN NATURALIST [Vol. XLI

{No. m was issued May 21, 1907)

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VOL. XLI, NO. 487

THE

JULY, 1907

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

ennial. Eemarksby CHARLES W. ELIOT 413

jf a New Species of Telenomus with observations on its Habits

nd Life His

ffiOREILL 417

III. The Development of Pinnate Leaves . . . • • FREDEBIC T. LEWIS 431

IV. Contributions to the Pleistocene Flora of Canada . D. P- PENHALLOW 443
V. Significant Results of a Decade's Study of the Tunicata. WILLIAM E. BITTER 453

VI. Notes and Literature : General Biology ; The A|assiz Centenmal.— llie pre-

Stone-gatherSg fishes. Botany: The search for mutations.- Biology

in the Journal of Agricultural Science. -^otes 461

Vn. PubUcations Received

BOSTON, U. S. A.
GINN & COMPANY, PUBLISHERS
29 BEACON STREET

Mew York Chicago

The American Naturalist

FREDERIC T. LEWIS, M. D., Harvard Medical School, Boston, Mass.
ASSOCIATE EDITORS
J. A. ALLEN, Ph.D., American Museum of Natural History, New York
E. A. ANDREWS, Ph.D., Johns Hopkins University, Baltimore
WILLIAM S. BAYLEY, Ph.D., Colby University, WatervUle
DOUGLAS H. CAAIPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., Cornell University, Ithaca
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA, M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University
CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM, Ph.D., Cornell University. Ithaca
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, Pittsburg
D. P. PENHALLOW, D.Sc, F.R.S.C., McGUl University, Montreal
H. M. RICHARDS, S.D., Columbia University, New York
W. E. RITTER, Ph.D., University of California, Berkeley
ERWIN F. SMITH, S.D., U. S. Department of Agriculture, Washington.
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington
W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis
HENRY B. WARD, Ph.D., University of Nebraska, Lincoln
WILLIAM xM. WHEELER, Ph.D., American Museum of Natural History
New York

The American Naturalist is an illustrated monthly magazine
of Natural History, and will aim to present to its readers the leading
facts and discoveries in General Biology, Anthropology, Zoology,
Botany, Paleontology, and the several branches of Geology.

The contents each month will consist of leading original articles
containing accounts and discussions of new discoveries, reports of
scientific expeditions, biographical notices of distinguished naturalists,
or critical summaries of progress in some line; in addition to these
there will be briefer articles on various points of interest, editorial
comments on scientific questions of the day, and critical reviews
of recent literature.

All naturalists who have anything interesting to say are requested
to send their contributions; candidates for the higher scientific degrees
are invited to contribute concise siunmaries of the special literature
pertaining to their chosen topics. The editors will endeavor to select
for publication only that which is of true scientific value and at the
same time so written as to be intelli^ble, instructive, and interesting
to the general scientific reader.

All manuscripts, books for review, exchanges, etc., should be
sent to Frederic T. Lewis, M. D., Harvard Medical School, Boston,

GINN & COMPANY, Publishers

THE

AMERICAN NATURALIST

Vol. XLI Juhj, 1907 No. 487

AGASSTZ ( EXTKXXIAl .

414 THE AMERICAN NATURALIST [Vol. XLI

teacher and expositor. He expounded clearly and sympathetically
before any audience the fundamental principles of his science, and
gave examples illustrating the principles with both hands, and
with shining, smiling face. He was just that, — a teacher by
nature, an enthusiastic, earnest, moving teacher.

As Professor Gray has just said, he came into this Puritan
society like a warm glow into a chilly room. He was a revolution-
ary spirit in Harvard College, an exception to all our rules. He
welcomed special students, for instance, who could not possibly
pass the examinations for admission to Harvard College. He
kept them for years in his laboratory', training them in his obser-
vational method, — quite a new introduction among us. Many
of our best people disapproved of that method! The son of one
of our most distinguished surgeons submitted himself to the teaeii-
ing of Agassiz in the crude zoological laboratoiy, and received
several trilobites upon which he was expected to spend weeks, —
examining them, seeing what he could discover in them, and mak-
ing a record of his discoveries. He was kept at this sort of work
for weeks without a book, and without plates. \\v was to mak(>
his own plates. At last the son .les(ril)e<l this (.n-.v.s to the

was at bottom a naturalist, lik.^ .'very phy.icini, or sui-eon. and

AikI llien, what a new kind of professor Agassiz was in this old
t,,\Mi: He hail none of the regular habits of the traditional Har-

when" to ..noke in the Coll.-v Yard wa> a -fave ollVnce. He
never wrnt to rhnreh. Sin,(h.v wa> his day of rest, but he did tiot
take it in the New l-^iLilaiid fa-hion. His mode of lecturing waa

No. 487]

AGASSIZ CENTENNIAL

415

unexampled among us. His conception of the duty of a jUDfi ^-^s(»|•
to investigate, to discover, to collect, w^e had only noiiccd faimly
in a few exceptional American teachers. Those nictliods lia I hccii
introduced in small measure among us; but those wvvv the \mmc
ideas of Agassiz as a professor and a teacher.

There were but two pitiful little collections in the })()sscssioii of
the University when Agassiz first came here ,— a collection of
minerals, imperfect, small, and never properly arranged, and the
beginnings of a botanic garden and herbarium. The idea of mak-
ing great collections of natural history objects hardly existed
among us; \\v had hardly aspired to such collections.

And then, he rais(Ml such astonishing sums of money for these
new subjects of zoology and geology. A good deal of jealousy

other departments lonu' (established in ( 'anibridgc for the traditional

uncle Afr. George Ticknor's, hearing- this jealoiiM- expressed by
one of Professor Aga.>i/^ (•.>ilea-iie. i„ Harvard I niver.itN. Hm
Mr. Ticknor said. -- " 1 ).Hri l>e alarmed; Aga^.i/ uiil -et more
monev out of the ( 'onnn..nwealth of Ma.s.ehu.ett> for hi. .ubje, t.
than any of you have <lreanted of oviiin-. than anv of you eould

416

THE A M ERICA X XA TCRALIST [\ ()T,. XLI

nol.le tVuMi.l, a.Kl w.^ >av with Lono-tVllow at the Satiinlav (1nl),—
We miss liiin -ivatly, I'.ut we rejoice in his eoiniiiu- hack to us in

i)KS(M{iP'ri()X OF A m:\v spkciks of tflfxomus

www ()HSFU\ ATIONS OX FIS IIAHITS AXl)

FiFi' IIls'^()Ii^■

418

V/Zr WLHK 1\ ^ \UR\L1ST XTJ

liaich; ihev were presumably destroyed hy y)arasites which failed
to mature or to emerge, for from the four non-parasitised egg
hatches 155 bug nymphs hatched from 15G eggs. In the case of
<«\(M-y parasitized pentatomid egg batch collected at Barstow,
'Fexas, or at Tlahualilo, Mexico, no nymphs hatched, showing
that in the majority of cases comi)lete destruction results when-
e\-er an egg batch of a host species is discovered by its tiny
enemies. Of 41 batches collected, 30 or SS% were parasitized
In Tilcnomus ashnuadi Tin-, probabh represents more dosely

the individual eggs. Although the nuiltiplication of tlie host
species ap{)ears to be effectively checked by these beneficial insects
by midsunnner, the y)entatomid bugs affected have already had an

No. 487]

A NEW SPECIES OF TELENOMUS

419

Description and Rfxords of Breeding and Collection.

This parasite having been pronounced a new species of the
genus Telenomus by Dr. W. H. Ashmead, the writer takes pleasure
in dedicating it to this eminent authority on the parasitic Ily-
menoptera who has described more than 500 North Anieric an
representatives of the family Proctotry{)ida\

Telenomus asJmeadi, Sp. 9 (Fig. 1, h). Length 1. OS- 1.15
mm. Black with fine pale pul)escencc.

Head.— Width, .50 mm., scarcely wider than thorax, marked
with impressed reticulations; mandibles black. Antemuv clothed
with pale pubescence, dark brown to blackish in color, except
pedicel the color of which gradually changes to light brown at
distal end. Proportionate dimensions of antenna! segments (Fig.
2, B) are as follows:^

Scajje Pedicel Segments of Flagellum

: : 1 2 S 4 6 6 7 8 9 1-9
Length 105 2S 80 18 12 11 17 15 15 16 20 124
Greatest width 10 12 11 11 13 15 18 18 17 16 13

Thora.r. — "Width .55 mm., length .48 mm. Dorsum marked
with impressed rctu illations, pubescent; mesoscutum rather
roughly longitudinally grooved posteriorly; scutellum smooth
and shining with a few pale hairs arising from minute j)unctures.
I>egs clothed with moderately dense pale pubescence; coxa>
black; trochanters, tibire and tarsa^ light brown by reflected
light, yellowish brown by transmitted light; tarsal segments
successively darker to the last; femora dark brown or brownish
black by reflected light, dark brown by transmitted light. Pro-
portionate length of segments ot hind tars;r not mcliKhng tarsal
daws, as follows:" I, ,\ !, ,. hvahnc. iridescent;

venation brownish; length ot tore wini: I nmi.. urcatcst width,
.46 mm.

Abdomen. — Lengtli .5.)2 nini.. wichh .51s nmi.; basal two
thirds of first segment striate above; ba>al two tlnids ot second

No. 487]

A NEW SPECIES OF TKI.KXOMV

423

No attempt was made to ascertain the nature of the tropism or
sense which leads to the (hscovery of host eggs by the adult para-
sites, but it a[){)cars tliat the attraction is as ^^reat for eni])ty e^^^'
shells of TfufaNfa rusiafm as for unlialchc.l i-u- of F.usrhistns
servus in a suitable Ma.irc of developiiu-iit I'nr Mi<-c....fiil para^iti^n.

men the pentatonii.l e^m. batch i. UnmA, tlic adult parable
carefully examines it with her antenna>. if the cjr^^. arc sati.sfac-
tory slie sets to work industriously and oviposits in one egg after
the other. The process of oviposition in a single host egg recpiires
from two and one cpiarter to three minutes. The body of the
female during this o})eration is held rigidly by the legs in a position
nearly perpendicular to the siirface of the host egg at the point of
introduction of the ovi{)()sitor. The puncture may be made
through the egg cap or top end of the egg, or it may be through
the side of the egg. The latter is more frequently tlie case with
eggs located on the outside of the batch.

Before leaving the egg the female scrapes it for a few seconds
with the tip of the ovipositor, usually moving it around the point
of insertion making a nearly complete circle, then reversing and
with a shorter radius passing aroimd to or beyond the starting
point, then perhaps reversing again and with a still shorter radius
making a nearly complete circle. Sometimes a much more irregu-
lar figure is traced but it is always curved for the most part, and
so far as observed the direction of the movement is abruptly
reversed from one to three times. It is difficult to imagine any
useful purpose of this instinctive act except to mark the para-
sitized egg so that it can be detected as unsuitable for further
attack liy parasites of its own and j)robably other species.

Egg lai/iiif/ capacifii. The largest number of pentatomid eggs

fenude of the species here coiisidcM-cd is 27. The total number
of eggs which a female para-ilc may deposit a])pcars not to be

424

THE AMKRK'AX XAirh'MJST

[Vol. XLI

NiMUKk oi I'l.M AToMii) Eggs Parasitizki) bv Single Speci-
Mi:\s OK Tdcnomus ashmeadL

Julv 17
Sept. IG

; hand which bear (

De\liopmfnt\i PunoD P\Kvsnis \.\
Period of Eggs of the Most m>

Sept. -MUM. 1. 10 A. ^r.-l !>. m. " " 14

In the case of tlie hrst, second and fifth records the exact time
of the beginning of oviposition is <;i\('n to (he nearest liour. In
the remaining records no note was made a> lo tlie exact time at
whicli ovi[)ositu>n began. J he iasi \\\v records, ahhougli made
at Ddh. hM-^ i(t<i n.^iMMUHi, Ml liu.i u,d | ni.it. o.in.ualK
ttnii, H,,.|..u I . Mm Ji.Mi 11, ( ill., IK. 1, pnind ni th. ucid

426

THE AMERICAN NATURALIST [Voi XLI

changed color only slightly if at all. Two days later the eggs
which had changed but little up to the time of the previous exami-
nation were as dark as the rest and indistinguishable from them
as far as appearance was concerned. Parasites developed to
maturity in each of these ten eggs and the first one was noted to
have emerged at noon on October 16th.

Development in infertile host eggs. It is the writer's observation
that shriveling of the eggs of pentatomid bugs indicates infertility
although in some species, as in that of the Harlequin Cabbage
Bug {Murgantia histrionica), a slight shrinking normally occurs
just before hatching. With this as guide for the experiment, a
batch of ten eggs was selected, which had been deposited by a
specimen of Enschistus servus which previously had deposited a
batch of infertile eggs. Four of these eggs were reserved as con-
trols and a female parasite was given access to the remaining six;
after having made the usual examination she was observed to
begin oviposition. The four control eggs shrivelled in the course
of a few days, but the six eggs into which the parasite had ovi-
posited became dark in color and to all ai)pcarances promised to
produce adult parasites, ^one appeared however nor did shrink-
ing occur, and several weeks later when the eggs were oi)ened
their contents was found to consist of a very dark colored vitelline
membrane toi^otliM uidi a ^,,,..11 shrivelled blackish mass on one
side, Avhich vva. nun < o.i.. ..l.l< as insect remains. A si,nilar
condition was found oc( asionally in parasitized eggs lu'lieved to
be fertile and belonging to a batch from which many adult para-
sites appeared. The failure to produce adult parasites from the
eggs used in the above experiment is therefore not positive evi-
dence that this was due to infertility of the host eggs, ^^he experi-
ment shows however in a fairly conclusive manner that adult
females of the species of proctotrypid here considered will readily
parasitize infertile pentatomid eggs, and that the resulting larval
parasites will develop suffu initl v to cause the host eggs to take on
the characteristic .-oK.r ot para^itiznl fertile eggs.

Farthniognu s,. ,nu/ it^ ,<l 'I »v to si x of offspring. No absolute
proof of paitlunoon.etH d(Nelopinent of the eggs of TiUmmus
ashrtyadi was obtained, but the (ontiilnitarv e\i<len(e from the
few breeding experiments undertaken furnishes a good basi.s for
the supposition that parthenogenetic development occurs and

No. 487]

A NEW SPECIES OF TELENOMUS

427

moreover has a marked influence on the sex of the offspring. The
following diagram shows the history of the offspring of a single
female, which was one of a lot of 120 females and 19 males bred
from eggs of pentatomids collected at Barstow, September 12,
1905:

9 (probably fertilized)

9 9 (females probably fertilized)

????? 9 9 99 9 99 9^ (probably not

, ^ , , . , , ^ , . , . fertilized)

12j>c?,19 4d^c?,6? 206^(5^,19 3c?d^
The specimens recorded in the diagram by a " V escaped or
were otherwise lost so that their sex is unknown. The last genera-
tion included eighteen specimens which are not indicated in the
diagram, since they came from eggs laid before the individuals
of the preceding generation had been separated fi-oni one another.
Of these eighteen, fourteen were males, and tlie others escaped
before their sex had been determined. Two females bred to
maturity in tiie laboratory imder conditions which allowed of
less doubt concerning their supposed infertility, oviposited in eggs
of Fentatoma li(/ata and produced 27 and 21 adult parasites respec-
tively; of these 17 and 15 respectively were males and the remain-
der escaped before their sex was determined. To summarize: —
The offspring of the probably fertile females numbered 10 females,

1 male, and 5 undetermined; the offspring of the probal)ly infertile
females numbered 93 males, 2 females,' and 20 uiidetcrniitied.

tiona/and probably the stn.iiuv>( cviM.mkv Tii;ii tlir tVrtiHtv <„•
infertility of the eggs of the parasite (ictcriiiinc^ to a utcat extent
the sex of the offspring. Sixteen females and S males were br(><!
from a batch of eggs of Pentatoma liqaia colNn'ted in a cotton held ai
Tlahualilo, in July, 1905. Another batch yielde.l 2() femalr. and

2 males. Pentatomid eggs collected at liarstow in the months
of August and Sej)tember, 1905. j)f()ducc(l 125 t'cninics and I'l
males. In all, of aduh parasites bred from pentatomiil eiios
collected in the fiehl. Hi] or s5', uere femah'., and 15', umlv-^.

No. 487]

A NEW SPECIES OF TELENOMUS

429

during their larval existtMice to nHiniiv no food for carrvini: on,
at least to a certain extent, tlieir rei)ro(luctive fun.-tion^. It is
not unhkelv. however, that their loiinvvity and rej)ro(hietive
capacity is increased by such food as they nn'^ht obtain under
natural conditions.

Sr.MMAKY AXl) CoXCLrSIONS.

1. A species of the proctotrypid genus Telenomus, helicAcd
to ])e new, is desci-ihed under the name Tclenonms ashmcadi.

2. The species, although originally bred from the eggs of
pentatomid bugs of the genus Pentatoma, readily attacks the eggs
of species of the genera Thyanta and Euschistiis and such eggs
ordinarily produce adult j)arasites differing in size from the parent
in direct correspondence with the size of the host eggs.

3. The develo[)ing ])arasite invaiiably occupies a fixed position
in relation to the embryo of th(> host, and emerges from the egg
through the end from which the bug nyini)h normally hatches.

of pentatomid egi^s at >onie (h>tanee; four specimens w(mv as

:.. 15efwe'en two and three minutes are re.piired for ..viposition,
after uhieh th.- >nrfaee of the hoM egg is marke.l bv the oxipositor
in a characteristic manner, presumably for aidi.ig in its sub.se-
(juent detection as unsuitable for attack by other parasites.

6. The maximum number of pentatomid eggs known to have
been successfully parasitized l)y a single specimen of Telenomus
ashmeadi is 27; but there is evidence that this number may be
greatly exceeded.

430

THE AMERICAN NATURALIST

[Vol.. XLI

9. Pentatomid eggs with translucent membranes containing
developing nymphs of a pale color undergo a characteristic darken-
ing as a result of the parasitism. Other eggs witli more opaque
membranes and dark colored developing nymphs do not exhibit
characteristic; changes in external appearance.

JO. Females of Telenomiis ashmeadi show no hesitation in
ovipositing in infertile pentatomid eggs, and such eggs when
parasitized do not show the shrinkage which is usual in infertile
eggs. In the case of Emchistus servus they undergo tlie changes
in color characteristic of the parasitized fertile eggs of this species.
Although no adults have thus far been bred from parasitized in-
fertile eggs, the development of the parasite is at least partial, and
the observations here recorded furnish only slight evidence that
complete development in infertile host eggs is impossible.

11. Sex of the offspring seems to be controlled to a great extent,
if not al)solutely, by fertilization. In nature where the chance
for a female parent to be fertilized is at a maximum the female
sex greatly predominates, whereas under laboratory conditions
w^hich artificially reduce the chances for mating of the adults,
the male sex predominates in an even greater proportion.

12. In confinement in the laboratory, adult life under ordinary
temperature conditions lasts but a few days. The maxinium
period recorded during the month of July is tlnvc days, dining
September four days, and during October eiulii days.

1.3. The a.lulls of l\lnwmu.s ashmeadi }ia\c net been obserxed
to k'vd an.l ap])an'ntlv this is not necessary for carrying on repro-
ductive fiiiKtion-..

WASHINcnOV, I). C.

BIBLIOGRAPHY.

AsiniKAi), W. TT.

1893. A inniinoniph nf thr North Ai.KM'icnn Proctotrypid^. U. S.

X„f. .l/'/N.. l)ull. I.-..

Mm,;.,\n. II. a.
MoFUilLL. A. W.

g 1907. The Mexican conchiicla in wcsici n Texas in 190.'). U. S. Dep.
of Agric, Bur. of Ent., bull. 64, pt. 1.

THK I)KVkl()p:\ikx r of pinnate leaves

432

THE AMERICAN XATFRALIST [Vol. XLI

purely ar})itrary, hut it iu.licates a possible uio<le of dexelopmeut.
The lower pair of leaflets m h is close to the stipules; in r nn.l d

sively orcater. Tu r oue stii)ule is enlari^ed aud leaf-like at its tip,
havitiu- a coarsely serrate upi)er uiargin and contaiuint; a mid-rib;
ill / ihcrc is a complete leaflet close to the sti])ule. 0, h and i
show an imperfectly developed ])air of leaflets in relation with the
stipules. ./ and /.: have a perfect pair close to the sti})ules, and in
/ and m this third pair is more distally placed since the i)etiole is
lon<rer. lluis the series su/rgests that the third pair of leaflets
is tleveloped from the outer porti(m of the stipules. Forms sliow-
hvr the similar additi(m of a fourth pair of leaflets may cnsil\ he
obtained, to<;cther with those which present the fiist and second
pairs in relation with the stipules. In the leaf a there is an un-
paired fourth leaflet on the rifjht, V)ut the sti|)ule on the left is

From the examination of mature rose leaves it a[)pears, there-
fore, that leaflets are adde.l from the stipules. Tt would be in-
ferred that the basal leaflet^ are the ia^t to dcNclop, but that the
stipules arise before the first pair ..f lat.Tal Icafiris. lu the rose

434

THE AMi:in<\\\ SM l hWLlST

[Xni.. XLI

prefers akn.petal). Niiux-li wlu. I.elieve.l that all leaves -row
near their apices, stu<li(Ml particularly the leaf development in
Aral lit --/nno.s-a L. uliich i> of the Lasifu^jal type/ At the distal
ends (.f its chief sul.divisioiis there are lobed leaflets like those of
the sumac. Steinheil" who believed that leaf-growth was generally
basal, considered that compound leaves were an exception in that
their outermost leaflets were the youngest. TrecuF recognized
both the basifugal type of comjKnmd leaf and the hasipefal which
would inchule both the blackberry and rose. He distinguished
also a mixed and a paraUel type. These are discussed and rear-
ranged by Eichler^ (])p. 1()-21). In addition to basifugal and
basipetal he recogni'/es six types, namely di\-crg(^iir, com ci-geiit,
simultaneous, ternary, cyclical, and parallel (but the last is not
in Trecul's sense). In the divergent form, the Icallcts develop

No. 4S7]

435

In Eichler's tables Rhus fi/pltlna is with the hasiFuc^al leaves;
"Rosa canina, tomcniosa, arvensis, etc." are \\\t\\ the basipetal.

436

Tin: AMi:i,'lc.\.\ XATl h'AI.IST [Vol. XLI

rose and calls them basipetal (p. 525). There are two serious
objections to grouping the rose leaf with that of the blackberry or
potentilla. Neither the terminal nor the basal leaflets of the rose
are ever lobed to produce new leaflets as in the blackberry; and
in the blackberry the stipules are not involved in leaflet production
as in the rose. If the term basipetal is to be retained for the
blackberry and potentilla type, the leaf formation in the rose may
be described as stipular.

Embryonic Leaves .— The interpretation of series of leaves such
as those shown in Figures 1-3 depends upon the study of their em-
bryonic development, for they are mature leaves and can never add
to their lobes or leaflets. The number of these parts is determined
before the leaf expands.

According to Eichler (loc. cit.) a leaf may begin in two ways.
In some cases, immediately below the tip of the axis, there arises
simultaneously in all its parts, a wall-like proliferation of cambial
tissue corresponding to the entire insertion of the future leaf to-
gether with its stipules, if it is to have any. In other cases, beneath
the growing tip of the axis a low papilla or conical proliferation
appears, which quickly spreads laterally so that more and more
of the circumference of the stem is involved in the leaf formation.
This spreading ends before the leaf begins to hv snbdivided nito
lobes. In one of these two ways the pn monl uil Iraj i> tonn.Ml,
from which (and never from tlx- stem) all i)arts of rlie matuiv leaf
develop. They are not pushed out from the stem. The pri-
mordial leaf forms from the stem; after that, all growth of the
leaf is onlv an elaboration of the primordial leaf.

After its fornuition, the primordial leaf begins to difterentiate a.
stationary basal zone, whieii is concerned only with the formation
one which ]>roduces the petiole
,vavs follows blade formation;
1 before the stalk l)egins. In

438

THE AMERICAN NArURALIST [\'<)L. XLl

development of a leaf to this point is characterised by sej?-
ion of its parts, accompanied hy relatively little increase
ime. ^I^liis is its cmhn/oiuc sfcu/e. It is followed hy the
/ r.rprnis-hn in which there is a .u'reat incn^ise in A-olume
t The :id(hti<)n of lolx's or h-aflefs. The rccoo-nition of these

below the growing-
length it presents

stipnle appears almost sinuilta
T leaflet, before any of the lov
nate close above the stipules."
eaf presents lower lol)es for the

of leaflets, and an upper lobe

toward the apex (the lowest leaflet is marked //). An arrest of

440

THE AMERICAN NATURALIST

[Vol. XLI

a{)proxi

matelv inferred from the variations m mature leave

udy of the mature leaves is therefore sif,niificant, but it ought
to 1)6 confirmed by embryological examinations.

Evolutionary Significance — A knowledge of leaf development
is of great importance to the paleontologist, and in a paj)er entitled
"Localized Stages in Development in Plants and Animals" Jack-
son presents a study of mature leaves from a geological and evolu-
tionary point of view.^ His conclusions are stated to be m the
direct line and the natural outcome of Hyatt's principles of develop-
ment. Professor Jackson's work upon leaves was continued by
Cushman who published three papers on localized stages m this
journal.^ The leaves at either end oi a biandi whidi tnmin.itis
in a flower are 'well known to be simj)l('r than those alon- Us mid-
dle portion. Simple leaves are expected luai iht bud MaUs,
the sepals, and the cotyledons. M. llihnre ((itioted by liecul)
sought to explain this arrangement by iiutini\e conditions, the
simple leaves in the young plant or near tlie (lower are (hie to ucak-

The eaiK kaNts aie said to upi
forms; and the su((essi\e lea\e^
most complex forms winch the

similar to those of MHing plants, ti.eii uiiiix aiciits au to .u souo
in the adults of ancestral groups.

^I^hp nature of ancestral forms is for the geologist to decide;

CONTRIBUTIONS TO THE PLEISTOCENE FLORA
OF CANADA

D. P. ri;MIALL()\V

Eaely in the present vear I nvcivcl from Professor A. P. Cole-
man of Toronto T'niyersity, a vrry line colK.ction of leaves from

branch which it was possible to identify with accuracy. Thc^c
specimens prove to be important since they serve to conhriii in
rather striking ways, conclusions already reached through previous
studies of the Don material, and they furthermore aflford addi-
tional evidence bearing upon the preglacial existence of types
now unknown in the living state. It is therefore thought desirabl(>
to place on record such facts as are revealed bv a studv of tliis
collection.

^riie last prt'vious record of the Don plants was inade by me in

cduv of the Plri>ro,vnc of Virginia and Marvland. and the evidence
they brin- forward oces to .sh..w that .'ssenTially the .aine flora

Acer pleistocenicum Penh. — Tlii^ s[)ccics appear- for the fourth

time in colh-ction. from tlie Don Valley, and in the prcM-nt iir.tance
it forms a large p.-nvnta-e of the (Mitirc material. Most of the

One of the best of tlie>e is reprodnee<l 1um-(> on a (limini>!ied s,.;,le
(Fig. 1), as it is nu.re complete than that employed f.>r tlie original

444

THE AMERICAN NATURALIST

[Vol. XLI

same. With the possibilities arising from more extended com-
parison of material representing a wider range of variations, the
conviction l)ecomes stronger that the opinion so stated is a tenable

Acer torontoniensis n. sp.— The Don collection embraces a
number of specimens, some of them fairly perfect, representing
a species of maple altogether iniknown, either in the fossil or the
living state. This leaf appears to present two principal variations
which depend in part upon the relative depths of the principal
sinuses and the character of the minor lobes or teeth, but chiefly
upon the fact that in one form the base of tlie leaf is only slightly
if atalllobed. while in the (.1 her case two large lobes extend down-
ward from tiic iii-criii)ii of ilic blade mi the petiole and enclose
the latter. 'i\v() i)rin( ij)al veins extend from the base of the mid-
rib to the corresponding principallobe.>, and two subordhiate veins
of varving prominence extend diagonally downward from near
the same point, into the two minor and variable lobes which form
the base of the leaf blade. From this description, as also from
the two specimens shown in Fig. 2 it will be seen that this leaf
belongs to the same group with our common hard maples. Com-
parison with these latter also shows that its nearest representative
among existing species is the common rock or sugar maple, Acer
.s«rr/mrm?m Wang. Comparing the upper fossil of Fig. 2 with
one of the more ordinary types of leaf of the sugar maple, it ai)j)ears
that the chief points of difference are to be found in the form of
the sinuses and in the character of the large teeth or smaller lobes.
If again we compare the lower fossil leaf in Fig. 2 with the cor-
responding type of leaf of the sugar maple, the resemblance be-
<-omes nnK-h' stronger bv reason of the similar basal lobes, which
]ri\e unfortunatelv been much broken away in the fossil. The
dillVrences noted are such as might well result from changes inci-
<lent to natural development, whereby the more simple tends in
the direction of the more compound, and ul.en to this there are
joined the actual resemblances, the\ >uu-e>i a \ er> nitmiate rela-
tion between the existing sugar maple and the t'o-sil, of such a
1 ... <l,.,t tl..- jailer ina\ i)e tlie ancestral form

Ko. 4S7] I'LKISTOCKM: I I.OIIA OF ( .\\M>.\ 445

446

THE [UhRK IV \ \HR\LLSI [\ o\ \Ll

inconspicuous, or prominent and extending
enclose tlie petiole as in Acer saccharinuni
l)r„ad and shallow; the terminal lobe with
'th; the lateral lobes with one or two large
de; the teeth acute, rarely somewhat acumi-

rved the occurrence of maple
1. but it is at present impos-
ot the recognized leaves

Carya alba \utl-— Although uevvr abundant, the leaves oi the
hickory have been noted in three former collections from the Don.
Their form and \enation are so characteristic as to leave little
room for doubt as to their true character.

Rollick ('00, 221, 222) now records the occurrence of three

■ from the Talbot and two from the Sunderland For-

of Marvland. Only one of these is specifically recogniz-
able, and to this the name Hicona pseudo-glabra, Ilolhck, is

Cercis canadensis I..— The red-bud, an altogether new constitu-
ent of the Don flora, is represented in the present collection l)y a

Madura aurantiaca

No. 4S7] FLKJsrnci-Xi: Fl.Dh'A OFC.WADA 447

448

THE A M ERICA X NA TUHALIST [\or.. XLI

anv ])revi(Mi^ collection from the Don. tlie leaves contained in the
im'sent one are sudiciently characteristic lo make the determiu-

Platanus occidentalis L- ( )ne small and imperfect leaf shows

represented, this tret^ lia> neverthele>> been louml in a ])r(>\i()ns
collection representino' two localities, and it is a recoiini/ed con-
stituent of tlie Don Hora.

IloUick ('()(>, 231, 232) ha. ^hown die occnrrence of P. acrroidr,
Goepp., together with another lar-e leaved bnt nnnamed si)ecies,
in the Sumlerland Formation of Maryland.

Populus grandidentata Michx. This w(>ll known hnt sparingly
represented species is a well recognized clement of the Don flora,

Prunus sp.— The genus is represented in the present instance hy
a single drupe of an oblong form. A similar but somewhat slu)rt(M-
fruit has been found on one previous occasion.

Quercus alba I..— The white oak is represented in the present
collection by fragmented but well characterized leaves. This
species has been observed previously in only one collection. The
specimens obtaiiu.! from (^.aol Hill were so imperfect as to make
the determination open to some (|nesti()n, but the present material
is suffieientlv pcrfc<i to vvmovv all <lonbt.

Other oak leave, arc also (Mni>raced in the 1906 collection, but
the fragments are too incomplete^ to justify reference to a particu-
lar species. It is <iuite probable that they may represent the
white oak, l)ut this cannot l)e stated with any degree of certainty
since the Don flora embraces no less than seven recognized species,

the second time it has been fountl.

No. 4S7]

449

452

Tllh: ami: UK \\ XATCHALIST [Vol. XLI

to Gray ('80, II, 147) tlie two species now occupy an unstable
position of such character that "a Httle further drying of the
climate would precipitate their doom."

The evidence aflonh^d U^ i!ie PNm.kh ciu dn. ..f Tnumio is
therefore in accord, in tins respect, with that tuniished hv certain
Pleistocene deposits at Khmra, Ni'w ^ork, and hv the concltisions
elsewhere stated with resi)ect to the recession of hequoia, Taxo-
diinn, and probably also Pseudotsuga, from the present (rreat
Plains region of Saskatchewan and Alberta ('04, 04-05).

McChLL UxivEKsrrv

MONTHKAL

•02. Sih-a of North America. Ill & IX. 1902.

Till-: SK. Ml- KANT IM-Sll/l'S OF A DKCADK'S

s^^I)^ ok tiik tcnk ata

WILLIAM K. i;LLii;i!

Morphology; 4, Knil.ry.iK.uy ; IMiysido-y.

1. Taxonomy and Affinities. -'I'lirrc an- x-vcral (jucsiions of
general interest thai iiatui'ally arix' iiiidfi' thi- head. Tlie Tuni-
cata behig a coni])arativcly >iiiall (■la>s of animals, does the prog-
ress made in gettin-' lioM of lie iirw kiii<l> iinHcaif that we are
approaching conij)l('iciics> in this (hrcction":' 'l\\r class hcing as
thoroughly pelagic at mic extreme of it> liahilat, a^ tlioroiighly
littoral at another, ami as thoroughly ahys-al ai ^lill another as

of number of kinds upon environment ^ 1- ilie pi-oMre-> ot knowl-
edge bringing out auythinu- concln-^ive as to the ureaier >ucces-. of

Herdman's "Revised ( "lassilicalion of the 'I'nnicala,"' pul.li.hed
in 1891, contains a total of .".iN >pecie.. Thi^ list supposedly
includes all the species known at that time. By a reasonably
careful enumeration, those (lescril)ed since that year nutnher 521,
making a total of UH»!) s|)ecies now known. Tlu- better explored
portions of the sea, such as the Atlantic about the Hritish Islands,
the coasts of Continnual l-nn.pe. and the Me.lite.auPean. hase
yielded verv feu of the neu nuv.. prol.al.b not m-.re than half a

water, lilerdman and .-luitcr : the M-a> traversed l.v the Sibou'a
h\pedition Hnit.-r ; the Cex l,>n region llerdn,an . th. southern
-ouih \me,i.an rei^ioi, Mi.hebui : the .fapan. ...a^l. , Oka
an.l llaitm.-vei : the I'a, ilic North Vnieii, an u-iou Ultter);

456

Tin-: AMKh'lC.W \.\ TrRALlST [Vol. XLI

pleura and am al)le so far to testify to tlie correctness of (rold-
sehmidt's observations. On the other hand Metcalf finds some
eviden(^e of metamerism in the rapheal nerve of the Molguli(Ue,
Perhaps these facts sh'trhtly support Perrier's view that the aikilt
sedentary simple ascidian is nearest to the vertebrate ancestor
and that consequently Aj)p(Mi<licularia is a modified ascidian larva.

Seeli^rer and .Metcalf doubt that there is a true homol()(iy between
the tunicate and vertebrate hypophysis. Xeither docvs Seeliirer
believe the vertebrate thyroid to be related uviici k ally to the iinii-
cate endostyle.

Lar<rcly from the character of the stiu'niaia and the ahseiKc of
an epi<..rdimn, Julin hold, a ^bMlite,ann."an .pe- ie> < alle.l In him

A. K. intra-'la.. kin.hi|,.. perhap. nnd.inu' of oreuler uvneral
interest iia-. come m liuht (hiin)i: the decade than that ( )ctacnenms
is an ascidian i)roper and has nothinu- to do with the Salpidte.
Metcalf and Hitter have made this positive.

The ])olvphvletic character of the eompoun.l ascidians is now

458

THE 1 MERK W \\1LR\J 1ST

[Vol. XLI

be mentioned the tendency to multiplication of ganglionic out-
growths observed by Metcalt in fcalpa.

4. Embryology, — Knowledge of tunicate development has prog-
ressed in numerous ways during the decade. According to my
judgment two of these are particularly signihcant to the general
biologist. They pertain to the verv^ early embryonic life and to
the multiplication of branchial stigmata. Among the investiga-
tions of the early embryo those by Conklin easily hold hrst place.
To the embryologist, one set of facts brought out by Conkhn
stands with special prominence in the midst of many that are
important. These relate to the cjuestion of organization in the
unsegmented egg. Conklin's figures seem to furnish strong sup-
port for his statement that " it is doubtful whether any other case
of cytoplasmic localization hitherto reported is more remarkable
than that which has been described in the preceding pages for the
ascidian egg."

Important as are these particular truths of ascidian develop-
ment emphasized by Conklin's observations, there are other as-
pects of his work which appeal particularly to the general zoologist.
In a resume of what is known about cytoplasmic localization
the author savs: — "The annelids do not approach the chor-

No. 487]

STUDY OF THE TUNIC AT A

459

my own part, sj)eakin<i; as a .syst('ii»atist, I am fully ])iv])are(l to
accept the full consequences of just this outcome. Furthermore
I see something of the reach of such consequences.

The fact about the development of branchial stigmata which
seems to me second in significance only to cytoplasmic localization,
is that the origin of the epithelium of new stigmata is always
dependent on that of preceding ones. The general question of
how repetitive parts arise is basal. These results on the l)rancliial
stigmata of ascidians seem to be decisive so far. Sclys-Long-
champs, Dumas and Julin are specially to be mentioned in these
investigations.

time ago about the origin and fate of the follicle cclis of ihc ascidiaii

decade, thanks to Floderus.\iancroft, 'Fodarl. and KorotnefV.

( )rinin of tlu> heart from the ectoderm is affirmed by Salensky.
lie treats this along with other instances of ontogenic origin of
as( i(han parts in deiiance of rigid germ layer tenets, under the

has rect>ived no thorough investigation (hninu- the .lecade.

5. Physiology.— Studies of the degeneration and rejiive

460

THE AMERICAN XATURALIST [Vol. XLI

croft that the ectodermal ampiillse in the test substance of the
Botrvlhis colony are rhythmically contractile and have a regular
circulatory office. There is coordinated activity among large
numbers of ampulla? in the same colony and this seems to be
without the intermediation of the nervous system.

The unique form of tunicate heart action has continued to
attract attention. The most comprehensive study of it is by L.
Schultze on Salpa. "The heart of Salpa is an example of purely
muscular self-regulation in a highly coordinated mechanism of
motion." This sentence summarizes Schultze's positive results.
Neither cerebral influence nor blood pressure play a causal })art
in the phenomena. Of similar general import Bancroft and
Esterly find proof of polarization through its own activity in the
heart of Ciona. On the other hand ]\Iagnus, Hunter and Froh-
lich bring forward rather strong evidence of the dependence of
heart action in Ciona on ganglia and to some extent on the brain.

NOTKS AM) TJTERATURE

Brooks, Killers ;

Commemorative ineetiiijjs were held ii
Hall, Cornell llniversitv, Professor Bui
address on "Wluit we owe to Agassiz," a
in the Cornell Era (vol. 39, pj). 441-44()).

To the younger generation of scientists t
personality of Agassiz belongs essentia
children arc still led by an inborn love of i
ponds for strange creatures, and to hriii
watering trough. Sometimes they are 1
medicine where parental warnings fail t(i
the presumably lucrative bounds. Ocea

on the rnoui
Agassiz^saic

THE AMERICAN NATURALIST [Vol. XLI

conditions were believed to be natural. At twilight the salamanders
come out from the mossy logs, and the male having found a mate
crawls under her body, working his way forward. His front legs are
swung around those of the female and he strokes the under side of
her head with his nose. After a time a spermatophore is deposited
on the moss. It is a pyramidal structure 8-10 mm. high and 4-6
mm. wide, sharply pointed with the apex upward. After its deposi-
tion the male swings his body 90 degrees to one side, but retains his
grasp, and his head remains beneath that of the female. The cloaca
of the female is thus brought over the spermatophore which is taken
up before the pair se{)arates. Mating occurs from July to September

The eggs are fertilized and in the following spring or early summer
the young arc laid. A new set of eggs is then mature and ready for
fertilization.

F. T. L.

Stone-gathering Fishes. — In the Amcnrtni Xaturalitt ior May
(pages .:;_>;3 327 of the current volume), Dr. AUred W.G. \Mlson has
given important information and excellent illustrations of so-called
"Chubs' Nest Although, as stated in a foot-note, no account of
these nests is included in my article on "Parental care among fresh
water fishes," it is recorded (p. 436) that several American c\'[)niii(ls
"also take care of their eggs, especially the lb)nic<l Dace [SriHof/liis
atromaculatus), the Black-headed DaJe (Pnm p/m/, ^ ,nn,n,ln.), .,n.l
the Stone-roller (Cam/>o.s'tow,a anomalum.)." 1 did not giw aii\- tiir-

isfactory, and Professor Jacob Jleighard had informed me that he
would soon publish an account of the habits of these fishes.

The evidence as to the exact species that heaps the stones in question
needs confirmation. Dr. C. C. Abbott, in "A naturalist's rambles
about home" (1S84, p. 408) po iti\e]y declares that Semoiilus cor-pornlis
(called by him bullarvs) "ditt'ers materially from the birds and even

spring. Once the eggs are laid upon their bed of sand, all care as to
their future vanishes"; he says nothing about the oviposition of
Scmolilm afwmarulatu.s noticed on page 409 and called by him S.

No. 487]

NOTES AND LITERATURE

The question thus arises whether the fish which prepared the chubs'
nests observed by Dr. Wilson was Seniofilus corporalis^ or *S. atroma-
culatus; the former has been declared not to be a nest-maker and the
latter is known to be such.

Our own common catfish does indeed carry "stones aimj/ from its
nest" but the following instance of the contrary liahit lias I)cei! de-
scribed in my article (p. 453). The Australian catfisli ( . I riu.s- (iiislm/is)
according to Professor Kichard Semon resorts to flat, sandy, and
stony parts of the river under a rapidly passing cnrn iif to spawn.
"When depositing its eggs and building its nest tiie fish oocs to work
in the following way. It begins by pre])aring a bedding about half
a yard in area, consisting of gravel and small pebbles among which

venting them fronrbeing washed away by the stream, or carried' off
by water birds .... or hy marauding littie fishes. The material for
this defensive structure i. derived frotn the aboxe nHMiti..ne<l rin-
(surrounding the n(\st) wliich iliereby becomes devoid of all siones
and gleams brightly in its smooth garb of wliite siind. It is wonder-
ful to observe the accuracy of ihe fish's handiwork and the perfcM t

circle described by the ring. So far as I could see the fish moved tlic
bigger stones by pushing them along widi iis tail, '['lie whole all'air

fied by hlin as Hh'nnrhllni.^ alnmasu.^.

Theodore Gill

472

THE AMERICAN NATURALIST [Vol. XLI

Carnegie Mus., 2 : 343-524, 1906) studied the crawli.hcs ..f Pennsyl-
vania and states that "anything that looks like a 'nmraiion" in de
Vries's sense is entirely unknown." Closely allied species eiilier have
distinct geographical distributions or if found in one locality they pre-
fer different habitats.

W. D. Tower is of the opinion that "the evolution of the genus
Leptinotarsa and of animals in general has been continuous and
direct, developing new species in migrating races by direct ivs]X)nse
to the conditions of existence" {Carnecjie Inst., I'lil.l. 4S). \\v states
that "there is not at present evidence to show the origin of any heritable
variations in the soma"; and that "in these beetles we can get new
permanent variations by stimulating the germ cells and in no other
way." Such an inheritable character he produced artificially by sub-
jecting adult beetles to abnormal conditions of temperature, moisture,
and barometric pressure. The eggs produced and developing under
these conditions give rise to new forms which breed true even under
normal conditions. But the parent beetles when restored to normal
conditions produce offspring of the original type. The new forms are
therefore believed to be due to influences of environmental conditions
on the germ plasm. F. E. I.utz of the Cold Spring Harbor Station
has reviewed Dr. Tower's work from the mutationist's point of view,

No. 487]

473

Ohio, and he uotvd that oc. asioiially, as at Cabhi John Bridge, Md.,
in 1900, sports are relalivel.v v.mv abundant. . . .If 1 had been so fortu-

eonehisions, and h^^c .U|.|>..r1cd the inut:Ui(>n tlieorynmst

loyally...."

Aifjner-Ahafi (Ann. // /.v/.-.V,//. M .\nf. limn/., 1: Isl .-,;^1,1900)

Porritt {Rep. Brit. Ass., 7i\: :Ui-y-XV2, 1907) ha.s (riven a detailed
aecount of the increa.se of melanism in Yorkshire U'{)i(h)])tera. The
author confines liimself strictly to recording the facts, some of which
seem to support the mutation theory although others do not.

In all tliese ca.ses it is olrserved that zoologists are cautious about

Biology in the Journal of Agricultural Science. .Mtlionuh the n al

474

THE AMEBIC AX XATLRALIST [Vol. XLI

Pl'B LT ( Wn ( ) XS R K C K I \' K I )

476

THE AMERICAN NATURALIST

[Vol. XLI

(No. 486 was issued June 22, 1907).

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VOL. XLI, NO. 488

AUGUST. 1907

THE

AMERICAN

NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

Pag,

I. Observations on the Natural History of Diving Beetles

JAMES G. NEEDHAM AND HELEN V. WILLIAMSON 477
II. Habits of the Short-tailed Shrew, Blarina brevicauda 'Say)

A. FRANKLIN SHULL 496

The capilaliznlion of sj)ocific luinies. Anthropology; Handbook of
Amerirmi Indiana north of Mexico.— tiamc-^ of the North Aniericnn
Indian,s. Zoology: Oogenesis in insects.— Parthenogenesis of BaciUus

logical literature. Botany; The fungi of termite nests.-The longleaf
pine. — Purple -producing bacteria. Geo%/; Rate of recession of
Niagara Falls 528

BOSTON, U. S. A.
GINN & COMPANY, PUBLISHERS
29 BEACON STREET

Entered at the Po$i-Omce, Boston. Mats., as Second-Class Mail Matt

The American Naturalist

FREDERIC T. LEWIS, M. D., Harvard Medical School, Boston, Mass.
ASSOCIATE EDITORS
J. A. ALLEN, Ph.D., American Museum of Natural History, New York
E. A. ANDREWS, Ph.D., Johns Hopkins University, Baltimore
WILLIAM S. BAYLEY, Ph.D., Colby UniversUy, Watervtile
DOUGLAS H. CAMPBELL, Ph.D., Stanford University
J. H. COMSTOCK, S.B., ComeU UniversUy, Ithaca
WILLIAM M. DAVIS, M.E., Harvard University, Cambridge
ALES HRDLICKA, M.D., U. S. National Museum, Washington
D. S. JORDAN, LL.D., Stanford University
CHARLES A. KOFOID, Ph.D., University of California, Berkeley
J. G. NEEDHAM, Ph.D., Cornell UniversUy, Ithaca
ARNOLD E. ORTMANN, Ph.D., Carnegie Museum, Pittsburg
D. P. PENHALLOW, D.Sc, F.R.S.C., McGiU University, Montreal
H. M. RICHARDS, S.D., Columbia University, New York
W. E. RITTER, Ph.D., University of California, Berkeley
ERWIN F. SMITH, S.D., U. S. Department of Agriculture, Washington.
LEONHARD STEJNEGER, LL.D., SmUhsonian InstUution, Washington
W. TRELEASE, S.D., Missouri Botanical Garden, St. Louis
HENRY B. WARD, Ph.D., University of Nebraska, Lincoln
WILLIAM M. WHEELER, Ph.D., American Museum of Natural History

New York

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THE

AMERICAN NATURALIST

Vol. XLI August, 1907 No. 488

OBSEin'ATIOXS OX THE NATURAL HISTORY OF
DIVING BEETLES

JAMES G. X1:E])HAM AXD IIKLl' X V. A\ ILLTAMS( )X

Our predacious diviner l)eede? of the family Dytix idie are
fairly well known as niuseuiii species, hm the >tudy of their life
histories and hahits ha- hceii -inuularly lu'uicctcd. A nuniher
of our genera and a few of oui- >pe( ie> occur ai>(> in Knroi)c, and
the natural history of M)ine of theM' ha^ l.een Mu.lied th.-re; hut

Therefore the foll-.^in.- oh.ervat i.Mi. on the hahits and adapta-
tions of the uronp may serve to direct attention to an iniworked
but interesting- field.

Dvtiscida-areveryc-omitionat Lak<. Forot, an.l are verv acce->-
ible in a catnpn> pond that lie. almo>t mnh-r the uin<lows of ,!,c

iuthor for that purpose with eia^-es for a nmnl.<-r of year--. TW
material incidentally aecuinulate<l in that work, coml.ined^ with

author during the academic year 1 <.)().')-('), will con>titute this paper.
The campus pond locally known as the "(iyni Po!id'"i from

made by damming a short, sj)rin<:'-ftMl branch of one of the ra\ ine>
that bound the campus. It has betMi in exi-KMice for many yeais.
and conditions in it are now qtiite natural. It is some sixty meters
long and about half as wide, an.l if attains a .l.>pth of four and a

end it hecom.'. diallolv, and is lilled with a <lenM- an.l clear growth

478

THE AMERICAN NATURALIST

[Vol. XL!

of cat-tails (Typlia). Tl
anywhere, but the hollovA
with oak leaves from the surrounding forest trees.

It is in the typha beds of the upper end of the pond, extending
from the shore outward into water of about a meter in depth, tliat
the diving l)eetles are commonly found. These beds cover an
area of hut little more than a square decameter, but in them we
have found twenty-nine species of Dytiscidie, as follows —
*Laccophilus viaculosus (iermar JI i/dwponis modcsfus Aube

LaccophUu^ ptsclaius Aubc Jh/droporus dichrous Melsli.

Larrnphilns- pro.viniNs Say Ih/hlus nmlusus Aube

*/)'/Wr.v,s'//.v larusf!-is Sjiy Afjdhu.s .suhfiiwatus Sharp

liidrssus jla r/rollls Lee. Ai/dhiis d i.sinlrf/ratus Cr.

Bidrs.sii.s ajjiin's Say lihanliis iiofaius Fabr.

H'a lamhiis i>injuali.s Fabi-. ( ,i)id>r1,'s .sridptUis Harr.

(Wlamhii.^ pin>rtafiis Say lli'idafirus piccuH IvCC.

('aJa,nhus disnar ' I'rilins .v, ml.s'ulcatu.s Aube

Calambu.s imprc.s.w-pinirlafNs Sell.

Deronedes caiascopium Say
*IIydroporus undulaivs Say
Xo other Dytiscida- have beiMi foi
that are occasionally cast up on tli
that w^e have picked up from ilicdr

Distribution by Size and Depth of
the above list tliat arc niark«Ml w

The Activities of the Adult Beetles. — Thei'e is
difterence m the s^^nnlnlno ^n^^^^n> of the^e heeth

CUnster piol^hh iiiaiuh M the hicrhest effiticiu.
The^long heautifullv fri.iired hiiul le^s are moved :
the flattened and fiinuv.! tnrsi nn<l the hhide-like 1-

scantily fniiired tcct dI m)1iic d the lower in
little nKxHfied in ih.-ir uunum fr-in ih.-.t .
pro.hirt ni.hM.hnlK l.iit httl. t.-^ult m I

No. 488]

DIVING BEETLES

483

484

THF 1 MI RIl [\ \ iJiR 1/ /^i [\oL XI I

coadaptation. This joining is so perfect m the higher forms, such as Cvbi
that if after the prothorax has been detached from the after-bodv an atf e
be made to replace it in its natural position, this is ver\' easily effected:

A (limimslu'd ir.si.sfancr to f/ir irafer has been brought about in
manv wavs,^ notahlv hv the roinuhiig of the contours of the body
especially at the neck and shoulders, so that it assumes a l)oat
shaped formf; by the depression of the evesf; by the loss of hairt
and sculpture; by reversal of theantennie; by recession of the fore
and middle legs into the concavities beneatli the thorax at the sides;
and 1)V the flattennig ol tlic hind legs m the horizontal planef.

brouy-ht about in nianv wavs. the seven most striknig'of which
art i> tollous — th( flittcnin. dounan.Isold(ini^ta>toftlu hind
co\. I u i /> / / f th( \. mill .lilt u< oMht iiHi i>r(Muim,

ment of braces at the loints to tiirihcr hunt motion to one plane,
making the leg more rigid and oar-hke; the shortenmg ot the prox-
imal joints of tlie leg f ; the lengthening ot the joints ot the tarsusf
accompanied l)v the flattening ot these joints and occasionally
ot tlie tibial sj)urs as well; the development of swimming fringes

No. 488]

DIVING BEETLES

in the thin lateral margins of the tarsus; the recurvatun^
tarsi to a more dorsal position, in with the itiotion ot the
of gravity of the body t; and finallv th(> loss ot (he hind chi

The modifications having to do with the takiiiu' and
of air are mn. h le.> ohxu.u. ...n^i.t in th. Ipi^

of the malum. ot tht (Kti.. ..h..idN mdiiionMlto, th.ii <oin,
and strenudu.un^ tun. t.om whi.h tnui to inak. ., ^^^u
air-compartiiicnt : and in the sh:i-hi luo.hlication ot the ii

in the enlaruTincnt ot the hnnhno'-i ahdoininal -piraclc- to

atus of terrestrial beedes has been cvidcntlv tairlv a(l«'(|uat
the main problem has been that ot i;(-ttiiiu' through tiic \\at(
sufficient ease and speed to capture picv and to t-^capc t

The Larvae Studied.— Five sptH-ics ot hirvir ot DvtiM-
kept under observation. Unfortunatclv but one ot thtv-c
porus undulatus Say) was reared to the a.hih l»fctlr. 1 he
are here named tentativelv. it Ix-mu' possible to make a suppv
as to the genera to whicii thi^v belong:-, ba>ed on tl
of the Gvm pond, on their size, and on then' nk
Emopean form. The kn...t k.na Mnonnnnd
^^ele tho.e of Ihn.di. Th, .e an ot iIh .muou^

book. We found them in Mav in -real number., fee.h
Corethra piipie in the deep narrow straits ol open water 1>«
standing aquatics, but thev are >o well known and the\- r
such quantities of live food (hulv, that we.hd not aiienipi t
them. Of still more snakv form aii.l with an e.piahv -o.hI
opment of s^\lnllmlu tiuu. . on h and the .i.lc ot n
abdominal sciiinents is the ai^ile hir\ a ot Aeihii.. Ir i> an e

486

Tlir: AM 1:1: K AX .\ ATI L' A LIST [Vol. XLI

rarabid.T A\e obtained luinu'rous .pedinen. m the fall of 1905
^vhen our ca»-es were first started, and these furnished our early
exi)erience. 'Hie hrst lot collected, kei)t over nicrht in a small
vessel, ate each other; in the niorninfr hut one remained. The
second lot, kept over night in a large vess(^l with pleiitv of proper
food, did exactly the same. Then we made a screen cage with
separate compartments, set it m an afiuanuin and i)iit our third
lot into it, one larva in each compartment. These then chmhed

No. 488]

489

lar\a of Ac

indicated in the diaur
pli^lied l>y xen .li-h

zontally once or twice, and finally n>f lo the --iii tacc, lail uj)\\
as just described. This species swims \cry rapidly. twi>tiii<i'
turning its long slender body like a snake. AVhen di-sturlx

darts witli a peculiar indescribable motion of the whole body
away from tlie i>oint of disturbance. Sometimes it niak(\s just
one quick dodge, and sometimes it goes through a series of wrig-
gling movements so swiftly executed that the eye cannot follow
them. This dodging feat must be of great advantage in avoiding

The larva of ITydroporu. i Fiir. 4^ has ouU a scanty <leN,>lop-

490

TIIK .l.l//;A'/r.l.V SATl h'M.lsr [Vol. XLI

(FijT. 7 D). Placed in a deep vessel ..f cit-ar walrr tlie Hydro-
porus larvjie spend mucli time .swiiniiiiiiu' nlxnu in this manner,
very rarely risin(^ to the surl'act\ Dutinu' vaiioiis periods of

half a dozen specimens to^-etlier in a lariiv hcakcr, nor one of them
rose to the surface. \Vlien kept in a shallow .lish of water with
plant fragments, they sjKMit nnich lim(> crawling- alnnit on the
bottom, creeping beneath dead leaves, or hiding in the hollows of
the typha stems.

Suppln»nital dr.srrlplions oj hlfhrrto >n,L-m>irn larra^ of D,jfi.nihr
I. Coptotomus interrogatus I'nhr. (supposition). L.mio|1i of larva:

492

Tin: A MKh'/CAX XATL h'Al.lST [Vol. XLI

HABITS OF THE SHOIIT-TAU.EI) SHREW, BLARIXA
BREVICAUDA (SAY) '

496

THE AM i:in( .\.\T( RA list [Vol. xli

was passed through a spool and bent down at the ends in tlie form
of an inverted U. The sharpened ends of tlie wire were tlirust
into the ground. The spool was held in such a position that a
thread unwound from it could easily i>ass into the burrow. On
the reel thus formed were wovnid scvimhI yards of carpet thread,
to the end of w4iich a snail was ti(>d l)y means of a hole pierced
through its shell just back of ihc lip. The shell was then placed
near the opening of the burrow. The thread was marked at
intervals so that it would be possible, without lirst (indin<; the shell,
to determine how much had been reeled olV. At the next visit to
the heaps the thread was found extending into the burrow for
about a foot. The shell was still fast to the sirinu'. but had been
broken open and the snail was gone.

Now that I knew where to look for the animal, 1 bcuaii to set
traps. At one place there were two laru'c heaps of shells about a
meter apart, each near a bin-row desccndinu abinptiv nito the
ground. Between these was a well worn path in the snow at tlie
surface of the ground. Into this path a steel wire tiap was sunk
by digging out a bit of the earth, so that the trig«rer of ilie traj) was
on a level with the bottom of the trail. No bait was used. At

hrrvwauda. Manv of the snails lia<l been rein.)\v<l. sliowino- either
that the animal had f.>r some time escaped the trap, or that another
shrew luuhaniMl on the w<.,k attu tlu h,M had b<-en < aptmed

Mv work wa> tluM. ord.^re.l a<v<H-dinu' to the following plan:
(], diMoxn a. man^ luap^ of M.aib a. possible in d.llerent
situations, an<l to record minutely the chan-es in location of the
shells aboNC ground; rl) \. M,on a^ the tio>t had thawed out of
the ground, to e\ca\ate the bunow. an.l - an h toi n(M>. To
capture in the meantime one or more .^hiew> and eontna- them in
the laboratorv; and iA^ To mak. xaiioM^ p^v < holoiiu al Mudies
in the laboratory and in the field.

All mx Ih Id obM n.Mion^ w, 1. mad( ii -n < 1 1 .u amp, a tiact
of snails were found, though search was made for them at other

498

THE A M ERIC AX XA TURALIST [\'..t.. XLI

500

THE AMERICAN XATURALIST [Yoi.. XLI

ture of the night, rather than the maximum or the average. The
Fahrenheit scale is retained as given in the weather records.
The snails killed in formalin which were placed at the burrows in
this area are not included in the counts.

in diameter, and the minimum nightly temperature, for a period of

°F Occupied Empty To

Apr.

As in Table I it is seen that the number of empty shells was
never diminished, showing that once the empty shells were brought
to the surface they were not ordinarily moved again. Throughout
the seven weeks there is a steady increase in the number of empty
shells. If the last two columns be compared, it is observed that
on ten of the sixteen days the change in terni>cr:itnrc and tlie
change in the number of shells are of oj)p()sitc sign, - - that is,
with a rise in temperature shells are remoxcti and \uv \vv>:\.
These ten days include all the most marked tenipcianiiv clianges,
namely, those on Feb. 20, Feb. 27, Mar. S, and Mar. 27. >iniilar
to these are the changes for Mar. 1 and Apr. :i The ( (.nspiciions
exceptions are Feb. 24 and Mar. 31, when, though the temperature

No. 488]

HABITS OF Tin: Sinn-:

changes are marked, the change in the number of shells is of the
same sign as the temperature change. Further, with the one
exception of ^lar. 6, all the considerable changes in the number of
shells occurred at times when the temperature changes were of
opposite sign. On this date there seemed to have been a marked
removal of shells into the burrows, while at the same time the
temperature had fallen. However, five days had elapsed since
the last preceding visit. In this time there had not been a steady
decline of the temperature; but the temperature had risen 8°
between Mar. 1 and Mar. 3, and then fallen 12° from Mar. 3 to
Mar. 6. Had I observed the shells on Mar. 3, the number of
shells might have been much smaller than for ^Far. 1, and then
increased to ^lar. 6. This seems especially probable since some
of the individual heaps showed an increase on Mar. 6, and others
a decrease. The decreasing effect of the rise of temperature
prevailed.

Nests axd Burrows

The Burrows. — Tlio record of the snails was closed Apr. 7.
Though a tVw occupied shells were still above ground, the weather
was then so warm that several of the snails were found crawling
about. Records of their transportation were untrustworthy after
that time, and were discontinued. By this time the frost was in
large measure out of the ground, and excavation of the burrows
was begun. Two methods were employed. AVhere the ground
was not very wet, flour was sometimes blmvn into tlic bunou s ^^ ith
a small hand bellows. The burrow Avas then ciuvfnlly opciicii as
far as the walls were whitened, and more flour was l,i.,un in. If
the ground were wet, the flour so-.n becamr uun^wuvA and lo.t
much of its whiteness. A more Mu.r^^f.il i.uMhn.l ua^ m ,,a^s a
rather stiff rubber tube into the burrow to keep it open whilf the

502

THE AM ERIC AX XATUHALIST

into the ground, so that it could not l)e said with ee
even the undergrourul burrows were those of the shrew
I have found underground nests used by ^Microtus, no
ing the emphasis which Rhoads (1903, p. 100) places o
nient that the nests of this species are built ''(it fhr su,-/
position of any nests that might be found could not tli
as a safe criterion. Some of the burrows iuid h-. ai» of

Microtus never ii-^e<l >iiails for t'ood. '\\> deteiiiiine
two meadow volo weiv coiiliiied in iron ca-e^ in ihe
Each was given a vessel of water, and ('(pial cai'c \\a- la
each cage clean and dry. One Microtus \\a> ui\en

No. 488]

HABITS OF THE SHREW

503

only water and snails. In each case the one confined with snails
died in less than 48 hours, the other remained in uood condition.
From these experiments I have concluded that all linrrows with
snail shells in or beside them were at one time used In tlu> shrew.
Taking this as the only criterion at first, I found other features
later which distinguished the burrows of the two animal-.

The burrows used by Blarina were usually iV) t.) ;5() nun. in diam-
eter. Those at the surface were exactly like tho.e of ^Microtus.
running in zigzag fashion imder weeds and grasses, often pushing
the latter aside, sometimes crushing them down, crossing and re-

504

THE AMERICAN XATURALIST [Vou XLI

the surface, rising as abruptly at one end as they had descended
at the other. The two openings of a single burrow were never
found closer together than a meter, and they were occasionally
four meters apart. This fact gives another means of distinguish-
ing the burrows of Blarina from those of ^licrotus. As far as I
have observed, the openings of an individual underground burrow
of Microtus were never as far apart as a meter, usually not more
than 35 or 40 cm. Microtus burrows, moreover, were not found
to extend as deep into the soil as those of Blarina.

Nests and the Burrows Near Them.— Nests are found along the
course of the burrows. In digging out the burrows some sixteen
nests were unearthed. Some of these were along ditch banks where
the groundwater level was lower than elsewhere. All the others
were in small (>levations such as mounds where celery had been
burinl or l.ills thrown uy) by roots of fallen trees. The' nests were
t'ouiui at depths of 15 to 40 cm. They were
, \ \ . 12 to 15 cm. in diameter, and slightly de-
^ ^'-^^ ^ pressed from the spherical form. They were
BiarlnalirciicM usually made of grass, sedge, and leaves of

nettle, goldenrod or ash, arranged in the form
of a hollow ball, the shell of which was 1 to 3 cm. thick. One
was composed entirely of hair which mi(Toscopic examination
showed to be that of the meadow vole. When plant materials
were used, the plants furnishing them were invariably found
inniiediately adjoining the nest. If grass was near the nest, it
was used almost to the exclusion of other leaves. Coarse mate-
rials were used without being shredded or torn into smaller
pieces. This constitutes an easy distinction between the nests
of Blarina and such nests of Microtus as are constructed of any-
thing larger than grass. In all the nests of Microtus wliich I
observed, coarse materials were torn apart; sedge leaves 6 to 8
mm. wide were shredded into three or four strips, and corn blades

than about 2 cm. Wlien the same kinds of material were used in

Xo. 488]

HABITS OF THE SHREW

505

singly. Tlic excrement was m'vmsh l.lack when fresh, sliglitly
brownisli when (h'v. It was voided in spindk-shaped portions
23 to 30 mm. Ion-, coikMl in xarion. ^^a^ a. in Fi-. 4. Very difi'er-
ent is tlie excrement of Microtiis, wliicii is found in black or brown

^Usually two, tln-ee, or four burrows radiated from the nest. At

an ordinarv bun-ow with a rubber tube until the burrow seemed
to come io a blind end; tlie tube could be pushed no further in

then excavated. When 1 had ai)proached within about 35 cm.
of the place where the Hrsi l.urrow had lu-en abandoned, tlie second
burrow also was closed. A third burrow ai>proachiu,(^ the same
spot wa. next duo- open, and it likexNi.e ended blin<lly. A spade
was set in at the i)oint toward which the three burrou s conxerj^ed,

bodv of a meadow vole near by sliowed that the nest w as then being
used, and was not a relic of'the preceding >ear. Careful search
all round the nest failed to rexeal a burrow lea.lin- to it. ^Fhe soft,
loose soil was then carefullv dug up to a distance of 40 cm. around
the ne.st and 20 cn,. l>elo^^ it at.<l exMmined as it was throNvn out
to disccncM- the shicw. None was foun<l, m, the soil was thrown
backan.l stamped d.>u n. \t n,> next \isit a l.urn.w opened to the

where the shrew had escape<l. It had e\ Identlx been present when
1 dug up the nest, l)ut had esca[)ed my spade. In this ca.-^e 1 have
concluded that the shrew was obliged to force its way through
the mass of loose soil for a distance of al)out 40 c-m. every time it

506

THE AMKRH 'AX SATURALIST [Vol. XLI

the burrows. Sometimes they were thrust into the soil at the sides
of the burrow in groups of 2 to 10. At other places short branches
led either downward or laterally from the main burrows, and then
expanded into chambers filled with shells, mingled with loose soil.
Such chambers contained in some cases as many as 80 shells.
Generally, all the shells were em])ty, l)ut one such chamber con-
tained m shells, of which :A were still occupied the last week in

beneath ti.e main burrow, the branch burrow leading to it was
spiral in form, like a winding staircase. ( )ne of these is illustrated
in Fig. 2. In two instances empty shells, broken exactly like those
found elsewhere, were found iir-^ide nests which, from tlu'ir com-
position of slu-(Ml(K-d material and position at the surface of the
ground, must have been Mi.rotus nests.

Method of Burrowing. — The method of burrowing was observed
and experimented on in the laboratory, where a shrew captured
by hand was kept for some time. When it was first confined,
loose bhick soil was place.l in the cage to a depth of about 10 cm.
Into this soft soil the shrew at once thrust its nose, and by violent

through the >oil like a wedue. Xo difficulty was experienced in
burrowing 20 or ;',() cm. in a minute. 'Vhv UKnements underground
wen- evi.h-nce.l bv the moveuH-nt of the soil at the surface; but
no ridge was forined above the burrow. When clods were en-
countered, they were rea.lily moved, even if fifteen or twenty times

opening to the surfa;-e at seven or eight points was worked out.
The aggregate length <.f burrow was not known, as 1 did not dig
out the soil, l)Ut it include<l the whole cage whirh measured ;;.-> l.y
4S cm.

the end of a week there was an aggregate of 35 cm. of Imrrow

No. 488]

HABITS OF THE SHREW

507

with three openmgs. An extension of 40 cm. had Keen made from
one of the former openings. 'VhU ^andy M>il liad not tu'en packed.
Hence the difficuhy in hnrrowinu in it mn>t liavc l,cen dne to its
weight, not to its hai-dness. In neither the l.h.ck nor >andy soil
did the shrew loosen the soil with its teetli as k'eimicott ( I SoT. j).
94) has conjectured Neither then nor at any other time (hn-ing
the confinement of tlie shrew did I observe any marked surface

Food.

50S

THE A M ERIC AN NA TURA LIST [Vol. XLI

covered cage in which earth was placed to a depth of about 10 cm.
AVlien practicable, live food was furnished. Among the various
foods tried were meadow voles and house mice {Mus musculus),
Mav beetles (l.achnosterna) and their grubs, moth lar\ae, other
insects and pupae, earthworms, snails, sowbugs, carrots, crackers,
roots of grasses and other plants. None of the last three articles
were ever touched as food. If any article proved especially ac-
ceptable to the shrew, that food was I'ui-iiishcd exclusively for
several days, and the quantity consuiiicd \vii> noic.l. From these
figures the average per day was coinpuicd. 'i'lic result in each
of tlie foods thus tested is given in Table HI.

Other articles of food were furnished at other times, and some
proved favorites; but owing to the difficulty in securing the food
no quantitative data were secured. Other insects, such as various
ground beetles, giant water l)ugs (Benacus), and Hydrophilus
friangalari.^, were furnished. All were eaten, but the ground
beetles were tlie favorite. Other larvae of insects besides Lach-
nosterna were readily taken, even the "woolly bear" of Pyrrharctia
isahella. Sowbugs were eagerly devoured. When live food was
not to be had, l^eef was furnished, and was eaten readily. I made

Xo. 488]

HABITS OF THE SIfRKW

509

anise oil, and the specimens taken still had fragments of the nut

Method of Capturing Food.— On several occasions I witnessed
the capture of prey. In the case of the voles and the mice, the
attack was essentially the same as described l)y Merriam (1S8(3,
pp. 106-168) and Morden (1883, p. 283). The hoii^e mouse,
being very agile, was not taken in the open, but only when it eiUered
the shrew's burrows. I observed this twice. The clumsy vole,
on the other hand, was pursued above ground, cornered in the cage,
and caught. In each case the shrew seized the animal's ear in its
teeth. After the shrew had been dragged around the cage until its
victim was almost exhausted, it quickly loosed its hold on the
ear, seized the head in the parietal region, and pierced the skull
with its teeth. In two cases the prey w^as dragged part way into
a burrow after it had been killed. In the third case it was eaten
at once at the surface. The brain and cranium were eaten first,
then the neck and shoulders. The skin was closely cleaned and
rolled back till the tail was reached. The snout, legs, skin, and
tail were left.

Some difficulty was cxp(M'it'nced in making observations on the
eating of snails. When beef ov mice were furnished, snails were
not touched. Finally, when all other foods were excluded, snails
put ill tho cage of a morning were devoured before the following

ing by this means that the sluvw would (^at snails at that time of
year (early in June). I starved it for a .hiy. then in order to keep
it at the surface put it into a caur with sandy soil, and gave it a
few snails. The snaiK N\<'re large and their ^lu'lK uere har.l.
The shrew put its lower jaw into the aperture in an attem])t to
reach the snail. Once its forefoot was thrust in. Failiiiu' to get
the snail in this way, it set its teeth across the omcr turn of the
shell and tried to break it. This it failed to <io in my pn MMiee,
but later the same shells were found broken. It -eeiii-- from these
observations that in the ca^t of laru'c >hells, iireakinu' i- a last

510

THE AMERICAN NATURALIST

[Vol. XLI

being mostly small and immature shells of the same species as
those on the left. These small shells were of course much more
fragile than the mature ones.

From the way in which the attempt to get the snails was begun,
it appears that when the shells are not broken the snails are dragged
out through the aperture. I did not see this done. To determine
whether it could be successfully accomplished, I seized an extended
snail with a heavy forceps and pulled upon it strongly. With
a steady pull the attachments to the shell slowly yielded and the
snail was removed almost entire.

Psychology.

I have described reels which were set at several of the burrows
to determine whether the animal that was moving the snails occu-
pied the burrows. The same reels were used to determine how
the occupied shells were distinguished from the empty ones. The
possible means that suggested themselves were the weight of the
snail, and the senses of touch, sight, and smell.

Muscular Sense. — To learn whether weight was the criterion,
an empty shell was stuffed with sandy soil till it was about as
heavy as an occupied one. This, with an empty shell and an
occupied one, was placed near one of the burrows. Each shell
was tied to a reel, and all were placed at equally accessible points.
The occupied shell was drawn into the burrow at the time of the
first decided rise of temperature, while the other two were left
indefinitely. The experiment was repeated, but the occupied
shell was so placed that the shrew would have to go round the
empty and stuffed shells and under the reel in order to get it.
The occupied shell was again removed and the other two left.
The experiment was twice repeated at another burrow, with, the
same results. Evidently weight of shell is not the determining
feature. It seemed possible that the center of gravity might not
be at the same point in a <^tuffed shell as in an occupied one, and
that the shrew could detect this difference. Therefore the position
of the center of gravity in ;i ^tnllV-l and an occupied shell was
determined by balaneinu' on a knifr e<lg(> and by suspension;
it was found to be the same in the two shells.

No. 488]

HABITS OF THE SHREW

511

Tactile Sense.— It might be supposed that the shrew would
reach into a shell with its feet and feel whetlu r the Miail was there.
I found later that the tactile sense was acute. Whvw tlie sluvw
was running at full speed in its cage and came n|)oi) an ohstacle,
it invariably stopped short before touchiiiu- it except with it<
vibrissae. The most common obstacle was it> wnter disli, which
was frequently moved about to different places iu the cage. I am
not certain that I ever observed the shrew rim against the water
dish even immediately after it had been moved. I have seen the
shrew run past masses of such favorite food as earthw^orms without

notu u.o tlu ni. but ^^hcu a u.mtu m it. uii -Im- t..u< hed the ta. tde
hairs, the slnvw turned at once au.l seized it. To Irani uiirther
the tactile mum ua. UMd w .luuuuu. iIh pi. m um nt a Miail in
its shell, I stufled theap. iiur..nr mnm iI ....upiMi -Ik 11^ uuli .oil
firmly, so that the snails wciv out ot M-lit and ivacli. 1 he cxtrrual
appearance of one of thcM' >\\v\\< was im a liuuian (.l.srr\-cr prtviM-Iv
like that of a stuffed on. iiuh ii u a^ la Id ii|> u. ilu h_l,i llu u

512 Tin: AMi:UirAX XATI-RM.IST [V.n. XLT

alon^ witli stuffed and empty sliells, at several biirrows. The
occupied shells were acrani removed, to die exclusion of the others.

Sight.— Blarina has not been accredited with acute vision,
the principal function of its eyes beinp;, as IVIerriam (1886, p.
]().">) has supposed, to distinguish light from shadow. To deter-
mine thi> point at first hand, various te^t.> were made on the shrew
in the laboratorv. Objects varying in size from a lead pencil to a
l)ook ^^■cIv waved ix'fore the shrew, first at a distance of a foot or
more. No notice wa^ taken of them. The distance was gradually
shortened unlil tli(^ ()l))(>cts almost touched the shrew's vibrissae,
but still the animal was not disturbed. Once when a cigar box
was thrust violctitlv toward [ho shrew, the latter shrank back
and inunediatcly turned to face the ol)iect. Thinking that the
response might have l)een due to an cuirent>, I mo\ed various
objects, as cards, boxes, or books, toward or ]mi>\ th(> >luv\\ m an
oblique position so as to produce currents. The -hi(>\\ ui\aruibly
noticed these although its head was sometime^ tin-ncd a\\a\ from
the object. I then blew lightly ui)on the animal and it turned
towani me and chattered vehemently. I have concluded that, in
the above case where notice was taken of the cio-ar 1)0X, the response

were nuKhM.n the opposite ^id.- of tlu- . nu<- >o a^ to thi
o^crtheshrew,theauimal^^a.a^on..Mli.tuHH.d. If;
as a book, were used to cast the shallow, the >hn-
hurried into one of its l>urrows. >i.uht, then, » an
to distinguish occupied from cnipn oi- >tutlcd >iuiil >
where a human observer could not di>( ( iti a diih
remaining possible means ot di-iinuui-hinu ilicm i> 1
Smell. — When mice or brcf were phicctl in the ( <i
almost invariably came out of it> l>uiro\\> in ;i -h
rarely did so when tlu li.l w;i^ nicrclx iai>ed and io\\«
other objects, as the water di-h. \\<"ic put ni. In tli

No. 4S8]

HABITS OF THE SHUEW

513

514

THE AMERICAX XATURALIST [Vol. XLI

in April. Both days were rather cool, though the sun was shin-
ing brightly. Still more eonclusivf, at least in regard to heat,
Avas the behavior of the shrew when l)rought out to be {)hoto-
graphed in a dish lined with white paper. The animal was exposed
to direct sunlight when the temperature was about 30° C. It
tried at every point to get under the paper lining of the dish, while
its breathing rapidly increased. After some 8 minutes of exposure
it was evidently overcome by heat, and after dancing wildly about
a short time on all fours, lay motionless. Long continued bathing
with cold water was necessary to restore it. It is evident that times
of daylight and even ordinary sunnner heat are not selected by
the shrew for its greatest activity. On the other hand, c\eii if
there were no direct evidence of daylight activity, the (■a])tui-c of
shrews by hawks (Fisher, bS03) shows that th(< animal, occasion-
ally come out upon the surface by day.

Summary of TRixdi'vL Hksi lts.

1. Blarina hrcvicauda preys upon various snails of the genus

2. These snails are hoarded, and arc in giMieral moved to the
surface of the ground as the tcmpcratnir fall> iuid into the l)nrn.ws

3. Empty shells which are brought to ih.' <\\v\-m-v arc not moved
back into the burrow.. The basi. of .li.iinciion L.-turrn empty

combined with the weight.

4. P'mpty shells not left at the surface arc stored about the
nests, along the burrows, or in special chambers.

5. Other principal foods are voles, mice, insects, and earth-
worms. Vegetable foods, except nuts, are not employed.

0. ^rhe burrows of Warina hrvrivamh are similar to those of
Mirrutn^- j,nni<,ilnnn<„.. l>m may b(> < !iM in^iui^hed 1a the I'ollou-

a. 'ilH> rim. of Ularina. wIkmi m h ler-roim. I. open t<. th.^ surface

No. 488]

HABITS OF THE SHREW

515

6. The nests of Blarina are always underground; those of
Microtus are more usually at the surface.

c. Blarina uses all its nesting materials unaltered; ^Microtus
shreds or tears coarse material.

d. The excrement of Blarina is greenish black, coiled spindle-
shaped, about 25 mm. long; that of :Microtus is black or brown,
spindle-shaped, 5 to 8 mm. long.

7. The smell, hearing, and tactile sense of Blarina are acute;
its sight serves merely to distinguish light from shadow.

Discussion.

The short-tailed shrew is easily recognized. It differs from other
shrews by its large size, having a total length of 120-124 mm., by
its short tail (23 mm.), and relatively small feet (hind foot, 15 mm.).
From the common mole and Brewer's mole, it is distinguished by
its smaller size, and by the absence of digging forefeet; from the
starnoscd mole it is further separable by the al)sence of tentacles
around the snout.

In the field, the work of Blarina is readily distinguishable from
that of either the common or .iavuo^ed mole })y the smaller burrows,
and the absence of humps of ranh ^^h^u■h air m, cliaractcristically
heaped up by both moles. A i'lirthor <li>iin( iinii i> tlu' rul-r ot'
earth over the burrows of tlic iiK.h's, csix'cially the ci.imiuHi in.)!i>.
Blarina does not make such a Ti<lu<\ at Ica-t in >ntr ground.

It has been noted that the riui> usvA l.y lilarina at the surface of
the ground are preciselv like lli...,' ,.f Microtus l,ut that the
burrows as a whole differ in several ropect^. The most interesting
of these differences concerns the material of which the ne>t> are
com{)osed. Shredding or tearing it into i)ie(e.> would ])erliaps
make the nest more comfortable and the slu-ew is fully capahle .^f
thus altering its material. But the shrew is canu\or(.u> and
]Microtus is a rodent. To the latter, with its gnauin- inci^.i-,
accustomed to dividing and tearing roots of gra>>e^ and liir t..irk
of trees, the shredding of nesting material is a natural pio.v.^.

The collecting „f empty ^hell. around the n.-r of ih<- ^luvw

516

THE AMERICAN NATURALIST [Vol. XLI

of the vole, rejected parts of its food. Microtus nests are commonly
made of the husks, leaves, and silk of the corn, or of the chaff
and leaves of the wheat which it devours. It is easy to conceive
that in this way the nesting habit of the shrews also originated.
If this be true, the use of grass, leaves, and sedge, now so common
among the shrews, must be a secondary modification, since these
articles are not rejected food materials.

The fact that in the laboratory the shrew did not make any
defined runs at the surface, suggests that it may not make any in
the field. If this is true, the runs which it occupies were probably
made by IMicrotus. They may have been entered in pursuit of
game, and when the original owners were captured, their burrows
were appropriated. The finding of broken snail shells in Microtus
nests seems to support this view, since Microtus does not eat snails.
The shells must have been carried thither on a foraging expedition,
and devoured in the nest of the vole. To what extent the runs used
by Blarina have been appropriated by it has not been determined.

Many of the shells found around the nests of Blarina, in under-
ground chambers, and in the burrows, were shown by the numbers
painted upon them to be those which were previously heaped at
the surface. The snails, therefore, were being hoarded, and
used gradually. Bachman (1837, p. 370) mentions that beetles
are hoarded by shrews of the genus Sorex, and Merriam (1886, p.
169) thinks it probable that Blarina stores food. Dahl (1891)
has found masses of earthworms, having their anterior segments
injured, in tlie burrows of the European mole; but Adams (1903,
p. 14) thinks they merely fell in and could not get out,— he does
not explain the injur}' of the anterior segments. There is no
mention of hoarding among shrews on as large a scale as this of the
snails seems to be. It has been noted that the snails were carried
out on top of the ground in considerable numbers when the
temperature fell markedly, and were taken l)ack in equally
large numbers wlien there wa^ a iii;ukc<] ri^c in temperature.
The snails seem to be kept in the coUk-st [.lace avaikable. In
cold weather this is above ground; in wartn wciirhcr, in the bur-

No. 488]

HABITS OF THE SHREW

517

early in April when it was warm enougli for them to crawL This
may have been due to the fact tliat my first shrew was (•ai)tiire(l
at this set of burrows, so that only one shrew was left to devonr
the snails originally intended for two. liodies of Microtiis wcie
hoarded but were not transferred to the surface. I'liis a^ain
indicates that the cold storage serves to keep the snails ininiohilc
rather than to prevent decomposition.

Too little has heretofore been known of the short-tailed shrew
to make an estimate of its economic importance practicable.
Stomach examinations are almost wanting, my own work including
but two. However, from data concerning the quantities of food
in laboratory and field, I have attempted an estimate of the eco-
nomic importance of Blarina.

Three principal elements determine the economic value of a
species, namely its range, its abundance, and the character and
quantity of its food. Of the range of Blarina, Rhoads says (1903,
p. 192): "Atlantic Ocean to Nebraska and Manitoba; Quebec
to Virginia." This is practically the northeastern quarter of the
United States. Of its abundance, the same author says (p. lOiii:
"This species stands preeminent above all others of our nianiniais
in its combined abundance and universality of distrilxiiion in all
conceivable situations. Not a place have I trapped over in tlic
two states but what it Avas amoni:- tlie first species to he canuht.
It is found in our deepest, coKU-st niouinain ravines, on the stormy,
barren mountain top, in the hanl^s and valleys of low tidewater
streams an(] maritime nuirshes, and delights in roving from the
cool sphagntnn bogs of the N. J. cedar swamps where tlie temper-
ature may be Inflow (10° to the hot sand barrens of the adjoining
fields whli a niid-.lav heat of 110.° Forest and plain, sand and
clay, ban-en or fruitful field, back w..ods or door yard, heat and
cold, wet and dry. day an.l night, have common charms for this
cosmopolite." It is (Hfhcult to conceive of the shrew in some of
these sittiations after having observed hs almost futile atteinj.ts

Yet numerous records atte->t it^ presence in the-e ^ituation-^.

different mannnals foun<l in the pellets of owls to (leterniine tlie
relative abundance of the aniinaN. .^hrcws necessarily came far

518

THE AMERICAN NATURALIST [Vol. XLI

down the list, because few were captured; and he found that
Blarina farva is more abundant than B. brevicauda. It seems to
me that the small number captured is due to the fact that they are
underground most of the time, rather than to their rarity. From
my own observations, assuming that a pair was present at each
nest that was being used, as I found to be the case in two instances,
there were at least two pairs to the acre over the region studied.
This number should be easily maintained for, according to Rhoads
(1903, p. 195), they produce four to six young at a litter, and
breed the year round.

The quantity of food eaten in a month has been estimated as
follows: From Table III was computed the quantity of each item
which would have been consumed in a month had that article
alone been furnished. For example, one month's rations of voles
alone would be 20; of house mice, 30; of adult May beedes, 450;
and so on. It has already been stated that 130 snails were eaten
by one shrew between Mar. 1 and Apr. 31. However, since the
moving of the snails by the shrew had practically ceased by Apr, 7,
it seems probable that the snails were eaten in a litde over one
month. Moreover such other foods as insects, earthworms, and
voles were available at the same time, so that the snail diet was not
the total. It seems reasonable to assume that 120 snails alone
would make one month's rations, since that is more nearly the
ecjuivalent of 20 voles.

The distribution of the dietary amoiiL^ tlic diflVrent articles is
largely a matter of judgment, and in Taljlc I\' the (|uantities are
based on the relative abundanct> of iho varions items in the swamp
region studied. For example, voles were abundant, and have
been allowed to constitute 40% of the diet. Earthworms, on the
other hand, were comparatively rare in tlie peat of the swamp,
and have been allowed but 5%- The table of course rei)rest'nts
only a sort of average for the year. Snails are evidently caren in
much greater nundxM's during several months of the winter, when
the hiseet .Het i^ neee>.arily limited. The snail di<-t i. pn.bahly

shells, all rnht'i:/r<' tlnim.lr.. id.out ihr oprmnn. .,f Mn.-dl !.urn.\s~-
in high land near Lexington. Ky. 'J1ii> wa. probably the work

No. 488]

HABITS OF THE SHREW

519

of Blarina. In other situations than the j)eaty swamp, the earth-
worm diet is probably greater than I have estimated. In pro-
portion as other foods not liere included are empkned, the
quantities in the table will be diminished.

Table IV. Showing estimated quantities of various staple foods devoured

Meadow voles (or equivalent in mice)
Adult insects (of the size of Lachnosteri

Estimating the number of shrews as I have done at four per
acre, it appears that the number of meadow voles devoured bv
them on a farm of 100 acres in a year is 100 X 4 X 12 X 8 =?,Sm.
Since this number can scarcely be supplied, the capacity of the
shrews for keeping the voles in check is not strained. Where
this quantity of voles can not be found, either other foods nnist
be eaten in equivalent amounts, or the shrew is capable of sub-
sisting on shorter rations, or the estimated four shrews ]X'r acre

a year than the f;ivnuM- himself. The shrew even compare;
favorably, IVoin iUc (•(OM..iiiic standpoint, with the common owls
Montgomery i ls!)!i i t'xaiiiiiu'd the pellets of four long-eared owl;
for a peri...l of tuo inonTh^. and found that these birds had de
vonred ;U7 .mall niamn.aK. mosilv Microtus. This is an average
of l:! per mo.ith for each owh " Blarina devours 20 voles pei

520

THE AMERICAN NATURALIST [Vol. XLI

of owl capture shrews but the species of shrew is not stated. Mont-
gomery (1899) found the skulls of shrews in the pellets of the long-
eared and the short-eared owl. The number of shrews taken,
however, is relatively small. For example, Fisher (1893, p. 53)
found in 562 stomachs of the red-tailed hawk 45 specimens of
shrews. Of these one third were short-tailed shrews, taken in
10 individual stomachs. In 39 stomachs of the barn owl (p. 139)
5 specimens of shrews were found, among which was Blarina.
Montgomery (1899, p. 566-567) found that out of 347 skulls of
mammals taken from the pellets of the long-eared owl, only one
belonged to Blarina. These figures show that the item of shrews
does not count very heavily against the hawks and owls in esti-
mating the economic value of these birds.

The subject of bird enemies of the shrew recalls the disturb-
ance produced in the hil)oratory by tlie fluttering of the pigeon's
wings. The sound was probably recognized as a familiar one by
the shrew. This accounts for the fact that the animal never
became oblivious to this particular sound.

Surface fl90(), ])p. 155, 160, 189, 197) has found shrews in the
stomaclis of four spe("ies of snake, though in small munbers. In
at least one case lie was al)le to identify the specimen as Blarina.
Rhoads (1^K)3) and Stone and Cram (1902) state that small mam-
mals are ca{)tured l)y foxes, minks, weasels, and skunks. In
several instances tliey mention shrews among the ninnber, but in
no case is specific mention made of Blarina brevicauda. Dicker-
son (1907, p. 356) records that three specimens of Blarina brevi-
cauda were found dead in the fallen nest of a red squirrel. She
believed them to have been killed and stored there by tlie white-
footed mouse. This mouse is well known to utilize des(M't('(l nests,
among others that of the red squirrel, but whether it kills shrews
is doubtful. It apj)ears to me more probable that the shrews had
been killed by hirger l)ea>ts of ])rey and rejected, possibly on ac-
count of their odor (Rhoads. IIM).;. ]». |".);5; Stone and Cram,
1902, p. b^2). and had then Ix'cn picked np \>y the w^hitefooted
mouM". Tlii. ni..n.e is said by Stone and Cram (1902, p. 132)

IV-Hii l.ii.l ciienn'es the shrew can escape to its burrows. From
tho^e enemies that can pursue it in its burrows, some other means of

No. 4SS]

HABITS OF THE SHREW

521

escape must be employed; perhaps it pushes out into the loose
soil. The instance of the obliquely descending burrow at one nest
suggests the "bolt run" by which the European mole is said to
escape when its fortress is attacked (Adams, 1903, p. 13). This
burrow, however, was probably not a back door escape, since it
ended blindly and the shrew did not enter it at this time of attack.

The short-tailed shrew is so well protected from its enemies that
no animals appear to depend upon it for food. It is abundant
and widely distributed. In security it devours such quantities
of voles and insects that its economic importance is considerable;
and since, unlike the other common shrew, Sorex 'pcrsonatus, it is
almost exclusively carnivorous, there is little to detract from its

LITERATURE CITED

iiACHMAN, J.

1837. Journ. of the AauL <>/ X,iL Sri.
Kennicott, Robt.

1857. Tlie Qun.h'uiHMls (,f Illinois :

■.of Phihuhlphia. Vol. 7, pt.2.
„J oj I'aU uts for lSo7. Agricul-

MoKDKX, .1.

1883. Canuilion Spnrtsman n,ul Xntundist. Vol. 3, p. 2S3.
Mekuiam. C. Jl.

1886. The Mammals of the Adirondack Region. Henrv Holt. 316 pp.
Dahl, Fr.

1891. Die Xahrungsvon-iitho Jcs Mavihvurfs. Zool. Am. Bd. 14,
Fisher, A. K.

1893. Tho ll;.\vk< nn.l OwN of thr rnifd StaK- in tlirir R.'lation to
Agriculluiv. r. .^. Ihpt. Aarir.. Div. of ( )niith. and Mammal.,
Bull. Xo. -A.
^roNTCa.MKHv. T. II.. .Ik.

1899. 01)s('rvati(>ns on Ow U wiih I'aM iculMr Rrizard to their Feeding

522

THE AMERICAN NATURA LIST [Vol. XLI

Stone, W., and Cram, W. E.

1902. American Animals. Doubleday, Page and Co. xxiii + 318 pp.
Adams, L. E.

1903. A Contribution to our Knowledge of the ^Mole (Talpa europtea).
Mem. and Proc. of the Manchester Lit. and Phil. Soc. Vol. 47,

1903. Mammals of Peimsylvania and Xew Jersey. Philadelphia.
Pubhshed privately. 266 pp.
Surface, H. A.

1906. The Serpents of Pennsylvania. Pennsylvania State Dept. of
Agric, Monthly Bull, of the Div. of Zool., vol. 4, nos. 4 and 5.

DiCKERSON, Mary C.

1907. The Pageant of Nature. Country Life in America. Vol. 11,

NOTES AND LITERATURE

GENERAL BIOLOGY

The Philosophical Problem of Life.— Dr. Venvoni, Professor of
Physiology at Gottiiigen, has recently piibHshed a lecture upon the
investigation of life, delivered before the society of political science
at Berlin.^ At the outset he states that the search for a cause in
biology is unfruitful and unscientific. "There is no process in the
world which is determined by a single cause. Every process is always
dependent upon a number of other processes and it is unjustifiably

arbitrary to select one of these and to account it the first cause

A scientific investigator can only establish the several conditions
which are necessary for the occurrence of a process. If these are
known, the process is accounted for, — explained. The process is
nothing more than the expression of the sum of the concomitant con-
ditions. The conception of cause becomes therefore superfluous and
worthless."

Accordingly one must regard as superficial such affirmations as
that an insect is colored green because it is thereby protected, or
that a mammalian embryo has gill clefts because its ancestors did.

From the study of the conditions of life Professor Venvorn concludes
that, — "To produce life artificially we must know completely all
the elements of the fixing sub.tiuur We nui.t knoxx the n^iatixe
amounts. We must undtM-staJKl their an-angement in the <rll l.odv.
If Axe (oiiM ( ..nM.iu I h a .^ ni h.lf.lh.m all thr * o,„l.tiun. ot lit. .
theartlhnal («11 xxonl.l at on.* lu. It xx ouM . . na.nU l.e.Mnnuh

and transmit its qualities — but the prospect of producing life is
a complete Uto[)ia. We have not learned to approach the complex

conditions involved in a living organism The chemical fabric of

a cell should first be so understood that it could be imagined a- a givat
machine shop, in which the mechanism of life could be observed bv
wandering among the atoms as among wheels and c^limlrr.-.

Consciousness also is held to be a product of these < (.n.liii.ms. It.
according to DuRois-Kevniond, we could brink' t(.«r« tlier ai mu e aii.l
in their proper relations all the aioiiis i.t wliieli ( .esar \\a> e<.in|HiM d

» Verworn, M. Di, Krj,.
45 pp. IMk. 80Pf.

524

THE AMERICAN NATURALIST

[Vol. XLI

when he crossed the Rubicon, Ave should have reconstructed Caesar,
body and soul. The artificial Csesar would have the same sensations,
aspirations, and ideas as his predecessor at the Rubicon. Both con-
sciousness and life, therefore, are the expression of definite conditions to
determine which is the object of scientific investigation.

Professor Verworn here ascribes as a cause of consciousness an
unknown arrangement of atoms. A more conservative opinion has
been expressed by an American biologist, as follows,^ — " The work
of physiologists has been so devoted to the physical and chemical
phenomena of life that the conviction is widespread that all vital
phenomena are capable of a physical explanation .... Let us give up
the ineffectual struggle to discover the essential nature of consciousness
until we can renew it with much larger resources of knowledge."

In regarding the construction of a living cell as a complete Utopia,
Professor Yenvorn differs from Professors Le Dantec and Cressou,
The former writes,^ — "Our knowledge of colloids is still so recent
and rudimentary, that we ought not to expect to see the making of a
cell accomplished soon; but it will come some day by careful analysis,
permitting a rational synthesis .... The scientific world today is so
prepared for the discovery that the premature announcement of spon-
taneous generation in gelatine submitted to the action of radium
surprised no one. . . .It is not necessary for an enlightened mind to see
protoplasm made to be convinced of rlic al)sen(e of imy essential
difference, — -any real discontinuity, hciwvcii liviiio; and dead matter."

Professor Cresson,^ after quoting Hudiiicr that "doiibth^ss some

"Such a hope is at least sonicwliat reasonable and prolmble." \Vhen,

known to be realized at ]m>s(Mn. and that tlie conditions in the past
when life arose are eriuall> unknown, one is inclined to accept Professor

It is unnecessary to refer further to Dr. Le Hantec's volume, which
was ]mblishc>d some months ago in English, and has been fre(|uently
reviewed. Dr. Troson's more nncnt volume is a simple introduction
to naturali.stic philo.soph\ . The autlior describes the development

'Minot, C. S. The pn.Mrn, m <nn~. i.HiMiess in its ijiological aspects.

No. 488]

NOTES AND LITERATURE

525

of natural science and its conflict with the "old geocentric and anthro-

pocentric philosophy which seduced and satisfied our ancestors

Science has descended upon this philosophy like a tempest and nothing
is left. The earth is not the center of creation. Man is not an excep-
tion in the universe. The adaptation between living things and their
environment is explained by evolutionar}^ principles without supposing
an intelligent creator." In the preface, philosophy is said to be a
matter of temperament. "For some, naturalism is the final word of
true metaphysics; for others, it is devoid of all truth." In this way,
perhaps, the author acknowledges, that there are many who see in
evolutionary principles the manifestation of an intelligent creator;
and who find in man, though one animal among many, much that is
exceptional. It is stated by Professor Cresson that naturalistic
philosophy is not science, though suggested by it. The determination
of the conditions of life, as described by Verworn, is science itself.

F. T. L.

The Capitalization of Specific Names.— It is agreed that the name
of a genus shall always begin with a capital letter and that the specific
name shall usually begin with a small letter. Zoologists are inclined
to begin specific names invariably with small letters, but botanists
employ capitals for a varietv of purposes as shown in the following
examples:

Zoological Names. Botanical Xinncs.

a. Slita cauadrn.v\s^ ,hn,rus Cnmulmsh

c. a a Jo pacha iliri folia Li/thrnm II n^sopijolia

d. Hn-nomis isabcUac Rosa Beatricis
Wliat(>vcr ivasons exist for beginning these botanical names with

capitals ii[)])ly with equal force to the zoological names; and the advan-
tages of the invariable rule for lower case letters are no greater in zool-
ogy than in botany. ^Moreover, as expressed l)y the \'ierma Congress
of botanists,— ^' The i^rinciples and forms of n()nionclature should

order to determine u])()n a niiifonu i.raciirc tur tlic Natnntlisi (in

526

THE AMERICAN NATURALIST [\^ol. XLI

"Recommendations for improving the nomenclature in future."
Among these is § C. "Specific names should ahcays be written with a
small initial letter, even when derived from persons or places, and
generic names should always be written with a capital."

1865. The British Association code was revised, and although the
rule for small letters had been very generally adopted, the section relat-
ing to it was omitted. The revised code stated that " It is not a matter
of great importance and may be safely left to naturalists to deal with
as they see fit."

1881. The Societe Zoologique de France stated,— " Every one
agrees that the name of the genus should be written first and begin
with a capital letter. For the specific names, there is also unanimity if
they are common nouns or adjectives, — a small letter is used. Should
proper nouns and adjectives be treated in the same way ? Some per-
sons adopt and recommend the practice. Your committee considers
that the question is of ver\' minor importance. It believes that it
conforms to the most generally established usage in recommending
the capital, which is not inconvenient, and may in fact, in certain
cases, be a useful distinctive mark."

1881. The rules of the International Geological Congress at
Boulogne, in regard to the nomenclature of species, merely state that
"the rules of Latin orthography are to be followed."

188G. In the code of nomenclature a(lo|)tc(l hy tlic Ain(>ncan
Ornithologists' Union, Canon VHI states that "proix-r tunnes of
species, and of subspecies or 'varieties,' are s\\\)i\v wonls, simple or
compoimd, .... written with a small initial letter. "

framed a code containing Art. 13.— "While specific jfnd substantive
names derived from names of per.M.ns may be writtc^n a capital

HMi."). At \'ieniia the 1 iiieniational Botanical Congress adopted
th<' fnll.Avir.^r nrnnn,„„<lnt,nn. Chap. III., Sect. 4. Recommenda-
tions, \. "Sp.^.-ili.- n;,nie> Ix'trin with a small letter except those

1«»()7. The lu.i.H'nrlature <-onnni.^inn u\ the H..t:.t.ieal Club of the

No. 488]

NOTES AND LITERATURE

528

THE AMERICAN NATURALIST [Vol. XLI

The opinions of the botanists who are associate editors of the
Naturalist, upon the capitahzation question are as follows:

a. "I shall be yevy glad to follow the practice of lower case spelling
for specific names in conformity with zoological usage."

b. "I am not verv' positive in my opinions of right and Avrong on
the capitalization question. I am going to try to follow the Vienna
practice as consistently as possible. The zoological decapitalization
has some valid arguments against it. In its favor is the fact that no
knowledge is required on the part of those who adopt it, other than
that the specific name chosen is to be used. I should suppose that
for the Naturalist a uniform practice for the different departments of
biology would be adopted, and the line of least resistance would be
imiform decapitalization. Any proof-reader is then competent to

c. "I have no decided opinion on the subject. ISIy instinct is to
use capitals for adjectives derived from proper nouns, as it somehow
does not look right to me to see them spelled with small letters. I have
no objection to offer if it seems best to adopt the uniform rule of small
letters for specific names."

d. "Botanists should follow the international code. Personally
I come near it, that is, I write names derived from persons with capi-
tals, e. g., Goldoni Lcwi.si, and names derived from other proper nouns
with small letters, e. g., Goldoni perwsylranica. American scientific
men (some of them) seem never satisfied to do things in nomenclature
the wav the rest of the world does. Really we ought to write G.
Lnrisi^ and G. Penns,,l ranim following the genius cf the Latin lan-
guage."

In order to know wlirtluT L;ilin usage had any luN.ring upon the

I)r. A. A. Howard, Professor of Latin at Ilananl ( ■..Ucge, wlio wrote,
— "There are no ancient rules whatever tor th<' use ..f capital letters
in Classical Latin. Our earliest uianuseripis aiv wriiUMi t lirougliout
in capitals, and so are all inscriptions. When the uiiiuiMule letters

530

THE AMERICAN NATURALIST [\' ol. XLI

the Bureau of Eiluiology, besides to tlie editor, i.s a coiupilatioii ot hrief
but comprehensive, simply worded and well illustrated, authoritative
articles, Avhieh represent the sul>stanee of our actual knowledge of
the Indian. A further perfection and po.v>,ibly e\ten^ion of the subject
matter will be attended to in future editions.

The work will prove in general a satisfactory reference book on the
North American aborigines, and a valuable handbook on the subject
in higher schools and colleges. It has, with its othcM- merits, the dis-
tinction of being the first work of its natmv in existence. The l)ib-
liography, though mostly restricted to synonymy, according to the
original plans of the work, is neviM-t li.-lcss ample and v/ill facilitate the

Wig\UauVhU'^ ... tl,l. uork a,.. Mi.> VMu-y,

diamberlain, Tewke., Kroeber, (.atMh.-t, ( yn^, TLoma.. Ih'^^.■tt,
Boas, Cnslm,- CnKill... l[,.,lg... llnlli.ka. Hough. \)ov..y Mas..,,,
M..(;,>i,v. ^h.olH■^. Sv,.,non, l)ix..n. Culin. Mattl.rus, lh>^^itt,

No. 4<SS]

NOTES AXD LITERATURE

531

532

THE AMERICAN NA T URALIST- [Vol. XLI

ZOOLOGY

Oogenesis in Insects. — It is a much debated question whether
the sex or germ cells are set apart at the outset of embryonic
development or arise later by modification of certain of the somatic
or body cells. The continuation of Marshall's studies on the anatomy
and embrj'ology of the wasp Polistes pallipes ^ treats of the early
history of the cellular elements of the ovary. The author finds that
in the embryos and very early larvie, each undifferentiated ovary is a
sync;yi:ium with a number of nuclei similar in structure. In the course
of development oocytes, primitive nurse-cells, and follicular epithelial
cells are developed from the undifferentiated cells of the distal end of
the egg tube. In a similar study of a Phryganid ^ he found that the
first differentiation had taken place in a fairly old larva. At this
stage the cells may be either "1st, undifferentiated or, 2d, passing
through the first stages in the development which is to result in the
further differentiation of oocytes or nurse-cells. Cells of the first
group may either remain unchanged and become the epithelial cells
or they may pass through the same stages as tliose of group two."
Thus ilarshall believes that the sex cells arise late and have a common
origin w-ith certain other cells in the ovary.

These results agree essentially with those of the earlier workers,
notably Korschelt, 'SG, on the history of the giM-m cells of inse.-rs. btit
are in'sharp contrast to the results of Ilcyiiioiis Lcc-iillon '0(1 01,
and many other recent workers \\ ho r,,iiirii.l tliai ;lir -mii cfll- ;iiv iu
origin perfectlv distinct from the lolliciilar cpitlu'liuin.

^ ^ * W. A. IUlky.

Parthenogenesis of Bacillus rossii. -The theory that each body cell
contained both male and female constituents, and that the egg cell in
becoming mature gave off its male elements in the second polar body
has aiso^'been much discusMMl. Tliis idea was su[)ported by finding
that the second polar body was not given off froTu certain eggs which

No. 488] NOTES AND LITERATURE 533-

developed parthenogenetically. According to Baehr/ the walking
stick Bacillus rossii must be added to the list of parthenogenetic species
in the development of which the second polar body is formed, and the
first divides in two. There is no evidence of their functioning further
for they apparently degenerate and disappear.

Contrary to a generally accepted belief that parthenogenesis in this,
species quickly leads to degeneration, the author reared perfectly
healthy females from at least the ninth parthenogenetic generation.
Apparently only females are produced,— it is a case of normal thely-
tokv.

W. A. R.

Phagocytosis. — By means of a clever technique ^Nlcrcier^ has been
able to throw new light upon the much debated question as to tlie
nature of the phagocytes in the batrachians and the insects. On
injecting sterilized, powdered carmine before the beginning of meta-
morphosis he found that it was taken up by the leucoc}ies and that
leucoc\-tes thus marked were yet capable of phagocj-tosis. Through
this method he was able to demonstrate beyond a doubt the active
participation of the leucocytes in the degeneration of the muscle
fibers. In the case of the batrachians the muscles exhibited signs of
degeneration at the time that the leucocA-tes entered but in the case of
the fly r'alliphora siicli signs were not to be detected microscopically.
The fil)(M- l)( (oin(>s bn.kcn up into sarcolytes which are engulfed by the
phago. vtcs. riiriv is no such phenomenon as the formation of
myoclasts and cons(Hiucnt autophagoc\-tosis. The author was able
to demonstrate with ecjual clearness the active participation of the
leucocytes in the destruction of the fat body of Calliphora and to dis-
tinguish them from the so-called " pseudonuclei " of Berlese.

W. A. R.

Histolysis in Queen Ants.— Janet ^ has studied in queen ants, the
degeneration of the wing muscles, which begins xery soon after the

' Baehr, W. B. v. '07. Uber die Zahl der Richtungsk(>rper in partheno-
genetisch sich entwickelnden Eiern von Bacillus rossii. Zool. Jahrb. Aiiat.
xxiv pp. 174-192. PI. 16.

2 Mercier, L. '06. Les processus yihasooA-taires iiondant la metamorphose

pp. 1-151, pis. 1-4.

Chez les reines des Fourmis. C. ll. Ai ad. :^ci. Paris, ixliv, 1907, pp. 393-196.

534

THE AMERICAN NA TURALIST [Vol. XLI

nuptial flight. This histolysis docs not l.co-in siinullatu-ously or
advance with equal rai)i(lity in all of these intiscles and hence anioug
fasicles apparently hitaet niay he found those in which the degener-

states that throughout the process there is no phagocytosis, or ingesting
of solid particles by leucocytes. The wing muscles are finally com-
pletely replaced bv adipocytes which, he belieyes. arise from leucocytes.

W. A. R.

Notes on Entomological Literature.— The (irmi P'Kpnoif of
Lomstidcr. PodLipolsky ' has studied both the chemical and the
spectroscoi)ic i.rciiMniitles ..f the green pigment extracted from the
wings of Locushi rir'niissivti,. lie was able to separate a yellow and
a green pigment compU lcly ])arallel to, if not identical with, the xau-
thophyll and the chloroi)hyllan of i)lant-green. The paper is yery
suggestive as regards methods.

W. A. R.

Innrr M , fa niurphosis of the Trirhopfrra.— ^Slm'h as the caddis
flic, have I.e. II studied from the biological and the systematic view
point. ( (iiii|):iiaiiv<-ly little is known regarding their histologic struc-

W. A. R.

Starving out thr Cod/inr/ Moth .— Vm\or this caption Fabian (iarcia
of the New Mexico Agricultural Experiment Station issues a call to
fruit growers to exterminate the codling moth in a single season!
The late frosts of last April l.^ft little pome fruit in the territory: if

No. 488]

NOTES AXD LITERATURE

535

'Handlirsch, A. Die T-.-ile In^^kt*
Formen. Leipzig. Willi. i:iigeliimnn.

536

THE AMERICAN NATURALIST [Vol. XLI

Berleses Entomology.— Fascicles 21-22 (pp. 585-648) of Berlese's
magnificent work ^ are just at hand. They conclude the discussion
of the nervous system and begin that of the organs of special sense.
Like the preceding fascicles these are not mere compilations but are
rich in new facts for the student of insect morphology.

W. A. R.

BOTANY

The Fungi of Termite Nests. \Ve are accustomed to think of
Belt's classic observations on the leaf cutting ants of South America
as the beginning of our knowledge of the relationships between ants
and fungi, but Fetch ^ assures us that Sweathman in 1781, nearly a cen-
tur>' before Belt's discoveries, stated that in tropical x\frica some species
of termites had chambers in their nests in which grew a kind of fungus
used by the ants as food. Although the "fungus gardens" of the true
ants of tropical America have been quite fully described, we have had
until the present time no comprehensive treatment of the similar habits
in the termites of the Eastern Hemisphere. Fetch brings together

scattered observations on this sul)j( ( t.

Ceylon does not afford such variety of form and size of tcrinit(>s as
Australia and Africa, but the nests of Trrmrs mimunni and T. obsruri-
ceps, the onlv two species which Fetch studied, are abundant every-
where except in the highest districts. The ant hills, roughly conical
in form, are only about six feet high. Their upper portion is con-
tinued into one or more hollow conical structures called chimneys.
The form of the nests varies greatly; they may slope gradually to the
top of the chimneys, they may branch into several chimneys or they
mav have a solid apex and bear the chimneys at the side. They are
built of earth and grains of sand brought up from the interior of the
nest and cemented together by a secretion of the termites. A large
portion of every nest is underground. In the early stages of develop-
ment the presence of a nest is usually indicated by three or four chim-

coU uomo vol. 1. Milan. Societd Editrice Libraria.

2 Fetch, T. The Fungi of Certain Termite Nests. Ann. Roy. Bot. Gard.
Peradeniya, 3: 185-270, pi. 5-21. 1906.

No. 488]

NOTES AND LITERATURE

537

neys 10 -20 cm. high, surrounded by the scattered earth brought up
in excavating the underground chambers. In fact in some cases the
nest is entirely under ground and the chimneys are wanting. Reasons

Internally the nest is composed of numerous chambers roughly
oval in sha])e, 5-25 cm. in diameter and .5-15 cm. in height, connected
by numerous galleries sometimes as much as 1 cm. in diameter but
generally only large enough to permit of the passage of two or three
insects at once. Similar galleries connect the chambers with the chim-
ney. For a discussion of the purposes of this structure the original
paper must be consulted. Some idea of the extent of the uiidergroinid
system of these nests may be gained from experinuMiis which retell
made; in one case water was run in for two hours iVoin a i)ii>e (leli\( r-
ing 15 gallons per mmute but this was quite fruitless lar as Hlling
the opening was concerned.

The chambers, except the royal cell, are generally nearly filled
with a structure designated as the comb. This is a grayish or brown-
ish mass, traversed in all directions l)y a labyrinth of anastomosing
galleries, and closely ivsenibliiig in general ap])ea ranee a coarse l)aiii
sponge. The combs lie free in the eliaiiib.M's, l(\iviii«r a rh-ar space of

has disappeared. The fact that the same substance is found in the
intestines of the workers and soldiers, taken in connection with the
regularity of formation of the comb from the small pellets, shows that
this is made up of the excreta of the tennit(vs. It ^\ ill tlins be seen that
the comb itself is not of fungus oriu-in.

The surface of the coinl) is uiwii a -Tavi-h or ulaiicoiis appearance

538

THE AMERICAN NATURALIST

[Vol. XLI

Some have suggested that this fungus is one which is found in the
neighborhood of the nests on decaying wood and that it is introduced
into the nest accidentally by the termites, but in an extensive investi-
gation of the fungi of Ceylon in which large quantities of dead wood
passed through his hands, Fetch was never able to find any form at
all similar to that in the nests.

Occasionally an agaric also develops from the comb. This species
is the chief edible form of Ceylon and so generally is it esteemed that
it is difficult to obtain perfect s])eeimens, for the natives who collect
them for food do not secure the long stipe intact and unfortunately
they do not overlook many examples. This fungus has never been
foiiiid oTowiiig from th<' li'ill ilsrir l)iit is always proihuv.l frotn the
undergrouiKl porlioiis of ihc nest. Tlie conih I'roiu wliich it develops

specimens w(>r(> found to grow from combs nearer the surface. The
connection of the agaric with the hyphte described above has not been
demonstrated. Efforts to germinate the spores or to grow the sphere-
producing mycelium froni the tissue of the agaric have proven unsuc-
cessful. It is not improbable, however, that they are stages of the
same species. At first the agaric forms brownish-white, soiiiewliat

No. 488]

NOTES AND LITERATURE

but probably only in wet weather. In this species a number of stipes
may develop from the same comb.

If a piece of fresh comb be removed from the nest and placed under
a bell jar the spheres will decay if the insects have been removed but
both spheres and external hyphtie will be eaten if the termites remain.
In the course of two or three days after the surface of the comb has
been freed from these, small jrroups of erect hyphic, indistinguishable
from those which give rise to the agaric. l)\ir app;irciitlv derived from

thin structures resembling the conidinl f(»riii> of \yl;iri;i. retell liiis

concliKles that it^' is probal.ly A'. „i;irijls. 'n.- tennire. eat this too
as it develops. After coiUimied rain A", nlr/ri/x-.s grows from deserted

Besides these forms, Mucor, Tlianmi.liui.i, ( ■ei.lK.Usporiuni. a.ul
Peziza sometimes grow on ccmihs removed from the ik-i.. >itiee none
of these are found in the nests, though some of them are e;i[>;kl.le of
growing underground, it seems probable that the iiise.K "uee,! out"
undesirable fungi as they develop.

dcfiiiitely"^proNed that tluw foftn the foo-l of tl.^ in^.t^. Tl. ■ luo
species stiKlie.l prefer fungi, or woo.l which ha^ ben aitackd by
fmigi. \Vh. ther a (litlVrence in food is a faclof in I he .litr.'n'iuialioii

is composed of sphere, of .wollen Veils uhieh in .letail reM'inble\ he

spheres of the frniilc ne>ts and the " Kohlrabihatifehen " of the
leaf-cutting ants invotiirated by Mollerare parts of a normal mycelium
and that their form has Ix-eii little, if at all. inodiiied by the insects.

J. Aktiiuh Hakkis.

The Longleaf Pine. - Schwarz's The Lrmr/kaf Pi»r ' is an attractive
little volnm.>. describing in a oooiilar stvle the silvics of Pin us palusfri.-

540

THE AMERICAN NATURALIST

[Vol. XLI

The longleaf pine is characteristic of the so called Southern Pine
Forest, and occurs principally in a belt some 125 miles broad, from
Virginia south and west along the coast to within a short distance of
the Mississippi River, and in southeastern Texas. The chief type
is that of a pure forest. Owing to various destructive causes, these
forests are largely in groups of different ages. A second, mixed type
is found farther inland, and is largely determined by differences in the
composition of the soil. Here the longleaf pines occur on the hiUtops
while farther down, on the richer or damper slopes are the oaks,
hickories, and other deciduous species, with shortleaf and loblolly

The natural course of evolution of the longleaf pine forest and its
method of reproduction are briefly sketched. The species is intolerant
of shade and requires direct overhead light, since the dense terminal
clusters of leaves shade the buds from side light.

The chief danger to which the southern forests are subject, is doubt-
less fire, hence this is treated at considerable length. The fires in
longleaf pine forests are exclusively surface fires, which not only
des'troy the young seedlings in the grass, but injure the butts of the
older trees, causing often considerable damage. The frequency of
fires, set either accidentally or purposely for burning over grass lands,
makes imperative the employment of rangers and the construction of
fire lanes about commercial forests. As a rule, seedlings of one or
two years' growth are destroyed by surface fires, but older plants
usuaily escape total destruction by virtue of their thick bark and the
dense head of long needles that not only protect the terminal bud but
form a miniature fire screen by hanging down about the short stem
to the ground. Frequent fires will, hoAvever, kill even these older
seedlings, to say nothing of their destructive action on the humus.

The future silvicultural treatment of these forests is considered in
Chapter 8. The forest must be perpetuated as well as exploited.
Cutting to a diameter limit of 16 inches has been recommended.
In some cases, a method of clear cutting with reserve trees left for
seeding the cut over area will probably be found good. The aim of
future management will also be partly to bring these ton >rs info a
more uniform condition instead of their present un ai ii n oulariiy.

Although more extended tables as to rates of -lowih aii<l \.)liinies
might have been added, this little book will no (l<.ul.i MTvr it> purpose
in helping the linnberniaii and the general r.'ader to a \n \\i-r niider-
standing of the proper study and ireanneni of our MHirlieru jMne
.1 Allen.

542

THE A M ERIC A N NA TURA LIST [Vol. XLI

recediiiff upstn-ain, that the gorge below the falls is the result of this
recession, and that it would be possible, by sufficiently accurate obser-
vations, to determine the rate of recession. He then discusses the data
upon which computations of the rate of recession must be based, con-
sisting of surveys of the crest-line of the falls made in 1842, 1875, 1886,
1890, and in 1905; and camera-lucida sketches made in 1827. After
considering the relative accuracy of the different surveys and sketches,
and platting the results together, the author concludes that a gradual
recession of the Horseshoe Falls is demonstrated, while a much slower
rate of recession is indicated for the American Falls. These changes
are strikingly represented by contrasted photographs and sketches
made from the same view-point, but many years apart.

Conc(;rning quantitative results of the study, the author points out
that the available data may be treated in a variety of ways, and made
to yield widely di wrgcM.t' nvsults. The lack of' harmony is dii<^ in

must Ik- taken into account in estimatiiiir su. li time. A >l,(.rt iv,..>i(
by \\. Carvel Hall on the latest survey ,.f llic civ^f lin<- ..f (he li.lU
appended to the paper.

A sprinkling of "reformed" (one i> templed to >ay deronned ")
spelling throughout the paper occ asiounlly distraets the reader's
attention from the matter itself to the maiuu-r in which it is i)re.setited.

I). VV, JOIIN.SON.

544

THE AMERICAN NATURA LIST [Vol. XLI

Basaltische Ergussgesteine vom Tepler Hochland. Arch. f. naturwiss. Land-
e,,durchf(rrschung v. Bohmen, 1905, vol. 13, no. 3, 72 pp., 2 pis.— Ruttnkr, F.
Die Mikroflora der Prager Wasserleitung. Arch. /. naturwiss. Landesdurch-
forschung v. Bohmen, 1906, vol. 13, no. 4, 46 pp., 6 figs.— Smith, E. H. The
blossom end rot of tomatoes. Mass. Agric. Exp. Sta., bull. 3, 19 pp.— Stej-
NEGER, L. A new Gerrhonotine lizard from Costa Rica. Proc. U. S. Nat.
Mus., vol. 32, pp. 505-506.— Stejneger, L. A new salamander from Nica-
ragua, Proc. U. S. Nat. Mtis., vol. 32, pp. 46.5-466. - Wakd, H. B. Icono-

skuU of Brac-huiK'henius. with ..hscrvnt i..ns „ii I he
saurs. Proc. U. S. Nut. Mu.^., vol. 32, pp. 447-4!
K. Trachytische und Andesitische Ergussgesteiii

pp., 1 pi.— Woodruff, F. M. The birds of the Chicago
of Set., Nat. Hist. Sur., bull. 6, 221 pp., 11 pis.

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VOL. XLI, NO. 489

SEPTEMBER, 1907

THE

AMERICAN
NATURALIST

A MONTHLY JOURNAL
DEVOTED TO THE NATURAL SCIENCES
IN THEIR WIDEST SENSE

CONTENTS

I. The structure of Cilia, especially in Gastropods . LEONAB

II. The Poison Glands of Noturus and Schilbeodes . . HUGH DANIEL REED 6S3

III. The Structure of the Silk Glands of Apanfehs glomeratus L.

EOBEBT MATHESON and A. G. EUGGLE8 667

IV. The Kest of the Kelp Fish CHAKLES F. HOLDEB 687

V. Notes and Literature: General Bio fogy; Specific chanicters in curly enibiTy-os.

births.— Recent public;ition.s concerning the structure of insects.—
Stridulation rhj-thm of crickets.— Notes. Botany; Plant geography. ~

VI. Publications Received

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ALES HRDLICKA, M.D., U. S. National Museum, Washington.
D. S. JORDAN, LL.D., Stanford University

CHARLES A. KOFOID, Ph.D., University of California, Berkeley,
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ERWIN F. SMITH, S.D., U. S. Department of AgricuUure, Washington.
LEONHARD STEJNEGER, LL.D., Smithsonian Institution, Washington
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THE

AMERICAN NATURALIST

Vol. XLI Sepiemher, 1907 No. 489

THE STRUCTURE OF CILIA, E.sPECI VEL^ I\
GASTROPODS

LI'OXAin) W. WILLIAMS

the liope that thev niav coiiinhute to tlie solution of the (iiHicult
prolilem of the .structure ot ciha.

"\^hlle lookuig over fresh niaruie plankton tnun NarrauaiiM'tt
Bav, we came upon an unidentified hnr connnon larva ot a pnuo-

546

NATURALIST [Vol. XLI

contraction of the protoplasm above the cilium carries its base up-
ward, and cramps the cihum against the overhanging edge of the
groove until the lower portion of the cilium is bent into an S-shaped
curve (Fig. 1, c and d). The contraction continuing, the cilium is
forced past the edge of the groove and flies out and back with a
very rapid stroke — the efl'ective stroke of the cilium (Fig. 1, e).
It is carried by the force of its stroke far beyond its position of
rest to which its elasticity brings it back in position for another
stroke. There seems no doubt that these cilia are elastic rods
(Fig. 2, g) which are moved by the contraction of the protoplasm at
their bases. The cuticula around the base of the cilium rises and
falls with the movement of the cilium as though it formed a plate
into which the cilium is set. Consequently it appears that the
contractile fibrillae of the protoplasm are inserted in the cuticula
and not directly in the base of the cilium. We do not know what
part the basal body plays in this movement but we believe that
it forms a pivot upon which the cilium turns somewhat as an
echinoderm spine turns upon its base.

The cramping and subsequent escape of the cilium account for
the rapidity and force of the effective stroke and also explain the well
known fact that the cilia of rotifers and veligers always seem to move
only in the opposite direction to that which is necessary, since the
effective stroke is too rapi<l to he visible. We do not believe that
the groove is a commoi\ ciliary structure, — on the contrary it is
probably present only in cilia like those mentioned, whose effective
stroke is invisible. However, we should call attention to the pits,
collars, and ridges at the bases of the flagella and cilia of Protozoa,
Porifera, and spermatozoa to which as yet no function has !)een

No. 4S9]

THE STRUCTUh'H OF (ILIA

others who consider that the isolated cihum is ciipahle
but hicks onk the necessary stimuhis.

^riiat the core or axis of the cihnni is alwav
iT-nprol)able bv the tact that many psemloi)cdia and tl
of the Siictoria, which are admittedly homolouou. with c
haye a central canal or a fluid core (Imo-. 2, m. Ah
experiments of Zacharias ^ who caused the s[)ennatu/(.
phemus to produce slender ciha-hke pseudopodia by
them in a 5 per cent, solution of sodium phosphate, ;
wazek ^ who saw the retracting protoplasm of an mjuu-d
of Siphonaea brijopsis produce in h\e minutes (iha wh
the rate of 40 strokes a minute, and who ' also toiind in CI
ciha appearing as small processes ^^inch in 8 nnnu
halt then normal si/e and beat 10 time, m 20 stcon.K
ceitdin ciha must consist of but slightK modihul i
These experiments also indicate that a solid or a pern
i« not always formed, for in the first two cases the cilia w-
dt.tuncd Ttis(h 11 tint 1 tnbi (ontiiniii^ i fluid
escape either becausi- ol tlu ( t il tui;;(»i oi iHiau i (

tube ^Mll a(t piidMh

550

THE AMERICAN NATURALIST [Vol. XLI

fications by Engelmann/ Piitter, Parker and Gunvitsch.^ Ac-
cording to this theory, the ciliuni consists of an axial snpport and a
contra(.'tile protoplasmic sheath, 'i'he natnre ol* the axis has heen
less the subject of discussion than that of the slu^ath which Mngel-
mann regards as fibrillar, Piitter as j)r()toplasni with temporary
fibrillar arrangements, and (iurwitsch as ])n)t()plasm of changing
surface tension. As stated above, \'(M-W()rn thinks that the ciiium
of ctenophores is formed of two cohmms of contractiU' protoplasm
whose differential contraction moves the ciiium, and Kiigelmann
seems to lean toward this view.

Less generally accepted theories are those of Benda and Schiifer.
Benda ' believes that the ciiium is passive and is operated by a
mechanism at its l)ase, but the cases cited al)()ve of the movement of
cilia entirely separated from the basal body and the cytoplasm,
and the failure with few exceptions to find such a mechanism,
make this view unacceptable. The velar cilia above described,
the existence in some cells of the liypobasal layer wliich seems to

Vcruorn's obM'rvatioii> n},oii (•t.Mioj)horc cilia Mign,-r. however,
that ahhoiigh this theory will not apply to all cilia, it is the only
theory which explains the action and stnictun> of certain cilia.
^Schafer"" re-ard. the cilinm a. an ciaMic lube (Fig. L'. r., one

forced can>iiig tli<- ciiium to Lend o\cr touani it. Ic- ria^ic ^idr.
This the..rv is plausible and the stnictniv is nuvhanically

tozoan flagella. It seems, however, that the action of the .nctorian
tentacles which are evauinated and invaginated like the finger of a
glove can only be explained by this tlieory.

No. 4S9] THE STRU

The generally accepted tin

of essentiaiK tlic ^-auw \vnv
toplasinic sheath which end-
Primitively the sheath ( Fiu'. '
not marked oil' stnicturallv
of the ectopla^n. .-eeondai
into contractii.'aiHl imneoiitr;
areslKHM. in the l\.llnu i,m <.\-
(Fig. 2, g) of velar cili;. a
confined to the hase of
reversible cilia (Fig. 2. / ! tin
two opposite bands w hich in
sides of the supporting axis,
more strongly than the other {
but certain organic and inor<

THE POISON GLANDS OF NOTURUS AND
SCHILBEODKS

554

THE AMERICAN NATURALIST [Vol. XLI

this is added a quotation from Cope, — "From it [the pore] may
frefjuently be (h-a\vn a sohd gelatinous style ending in a tripod,
each limb of which is dichotomously divided into short branches of
regular length." Jordan and Evermann ('90) make a similar
statement and add, — "The sting from the pectoral spine is very
painful, rescnd)ling the sting of a bee, but worse." Again, Jordan
('04) writes the following footnote,— "The wounds produced by
the sting of their sliarp pectoral spines are excessively painful. In
tlie axil is usu^illy a ])ore, probably the opening of a duct from a
poison gland. Tliis matter deserves investigation." Finally
('05) he writes, — "In two genera, Noturus and Schilbeodes, a
poison gland exists at the base of the pectoral spine, and the wound
gives a sharp pain like the sting of a hornet and almost exactly
like the sting of a scorpion-fish."

Boulenger ('04) does not consider this axillary sac a poison
gland. In this connection he says, — "I think this condition of
things has nothing to do with a poison organ, and is merely a
rej)etiti()n of what is observed in loaches and in the characinid
Xencx liarax, where 1 have found a gelatinous substance filling the
short (hu t by wlii. li the membrane of the air bladder is placed in
coniinunication with the skin and the sensorv organ of the lateral
line."

found in >exeral <hlV.Mvnt ,un.up. ..f lid..'., (iln.thcr , •('.',»> has
deMTibe<l a poi.on apparatus in T halus.ophnjuc rrtlrulaia u hich
inflicts a wound followed by poisonous .symptoms. At the ba.se
of the dorsal and opercular spines in this species he found a sac
connected with a canal passing through the whole length of the
s{Hne and oi)ening through a slit at its di.stal extremity. 11ms the

innocnuu-^ M.I.Manrv and tlu-reforc I ha-.r n<.t lu-italcl li> <i.-ig-

555

spine caused the h. jet a foot or m..re tVom the o]>eiiiii,-' of

the fishes were (Hsphiyed in the markets. Sehim'.h i'7oi and

the dorsal and opercular spines. There are mimerous referem ( s
to the poisonous nature of the weever fishes, Imt the two author^
mentioned are the only ones, so far as I know, who have des( rihed
the structure of the glands.

Wallace ('93) in the toa.lfish, Opsanus iHaimrhus^ inn. .Hs-
covered glands in connection with pores in the axilla, on the ilorsal
portion of the operculum and on the siu-face of the pectoral fin.
This author does not consider them as poison u'lands althonnli
the spines of this species o'ive a "slio-|,tiy painful stin.i:- and the
-lands apparentlv are of the same type as th<.se <.f the weever
fishes dcscrihed l.y Sclnnidt and Tarker. and tlu.s,. of the mad
toms to I.e described presently.

With the ahove facts and views in mind and a d, mh' to know
more of the action and structure ..f the supposed p.-ison -land^.
all the known species' <.f Xoturus an.l .-chiilu'o.Ies. except N.
junrhris, have heeii examined. The stin- will he .ies. ril.e.I first,
and then the structure of the ulan.ls.

The Sting.— The sthii; of the ma.l toms has l.een descrihe.l as

556

THE AMERK 'AX XA TURALIST [Vol. XLI

Evermann (MS.) describes the pain as, "A very stinging sensa-
tion, more like that which would result from a severe nettle sting.'
This describes precisely the majority of stings, for in handling
live specimens ordinarily only the tip of the spine enters the flesh.
The mechanical injury is so slight that it frequently is impossible
to locate the wound except for the stinging sensation. From an
ordinary sting, such as is received in handling the live fishes, the
pain continues from one to several hours, depending probably
upon the amount of poison entering the wound. Both in sensation
and duration these wounds differ from those made by the prick
or puncture of a sharply pointer! instrninent. The swelling is
hardly perceptible,* except in cases of \t rv scmuh' punctures, in
which event the flesh about the woun*! l)!'( ()nu's distiuctly swollen

and sliglitly discolored. Similar results, but more marked, are
j)r()(hi(rd l)y introducing a portion of a fresh gland underneath

The Poison Glands. — All the species of Xoturus and Scliilbeodes
possess axillary glands which open througli a poif^ situatt'd in the
axilla just below the post humeral pr()('(vss i Fiu. 1, Tiie
position of the pore with reference to the l.a^c of the (in and the

the ,.oiv i'. nioiv or I-., ^iii-likc and i^ Mtuatrd that uhcn the
fin i- addu.lrd ilic .].i!u- n,-. riih-i- dircrtlv a'-n... or parallel with

No. 4S9]

plished (luriiio- aililiidioii l>iu
metliod of aiK.iiilini: the >}nuv
is 0})eiH'(l widely when the tin i
the lii).s of the poiv. if aiiytlm
abduction of the fin. In a])}
lips of the pore are contiinied
folds with a jrroove between tli
a safe conduit from the pore to

The t^hinds are pear-sha])ed,
with tiie apex toward the axil-
lary pore (Fig. 2, p. g. and p.
o.). In most species examinee I ,
the gland is inclined l)ackwards
and lies just beneath the skin
and entad of the posthnmeral
process (Fig. 2, h). Jt is sur-

gland was o nun.

The glands are divi.led into
three main lobes, each of wliicli
i.^ub(H^id(>dintolo],nk-. Tlu-
lumen of the gland i> extremely
narrow (^xcept near the pore

558

'/'///•; AMEliK ■ 1 .V SATURALIST [Vol. XLI

senting the lumens of further su))divisions. I have fre(iuently
found globular masses of the secretion filling and tiepending
from the pore (Fig. 2, s) but have been unable to withdraw one
with the branches intact. The secreting cells are granular and
very large. They range between 80 and 200 microns in their
greatest diameter, and vary in shape due to compression with
neighboring cells. Most of the cells contain two large nuclei. As
a rule the two nuclei are separated by a short space, but fre-
quently they are found very close together or apparently joined.
This may indicate recent division as Parker has suggested in eon-

560

THE AMKRICAX X.\ TCRA LIST [Vol. XL.

same will ])r()i)al)lv he foiiud tni(> of N. noclunnix for the dorsal
.pine nland i. heie ^^ell .lex<'l<.pe<l n> the i,ectoral. The
presence of spine ojan.U nun he detennined h^ ( arefull^ slit-
tino- tlie overl_vin.ir skin and serapiny- tlie s})ine with a needle.
It' the j,dand is present, the serapinos when stained and
mounted show the tyjiieal (^laiid strncture almost as clearly as
sections. Sometimes the strncture can he made out with a
dissecting? lens without either staininu" or scraping the tissue from
the spine. In the fresh condition the olands are of a translucent
jelly-like consistency and ap})earance. 'J'here is no difference in
structure between the axillary and s|)ine glands.

It appears from the figures and descriptions of Schmidt and
Parker that the poison organs of the mad toms and the ^^ee^er
fishes are identical in structure. In the latter group the glands
arc found only in connection with the opercular and dorsal spines
which they surround in precisely the same maimer as do the i)e( toral
and dorsal glands of Schilheodes gyrinm and ^S. noclunni.^.

Differential stains fail to reveal^ the presence of aiiv imiscular

cells and force their secretions to the exterior. 'i1ic same is true
accoiNliiiir to Schmi.lt and Parker of the weevcr fishes. Tlie
latter ( 'SSi wrings. "No special muscles are present in connection
with the ghindv 1 am iin liiu>d to think that in the discharge of

that "Along the ventral side of the upper gland are found a few
hmidles of the extensor muscle of the gill cover but they could
hardly })ro(luce any ])ressure on the gland, and moreover, no
organ, adaj)ted for active ejection of the secretion, is found."

released, for in no case lia^ a natural opening been found in the
cells, and the secretion, when >tained are found to contain rup-
tured and rao-..-ed (elU in main .4' ^^lu<h the nucleu. can still be
ma<le out. FrcpientK -min|.n<>d cell, aiv found lloatina in the
secretions alon-- with the ruptniv.l ones, rsnallv these are small
cells, prohahirimii.atiiie one. uhi.h lune !>.<>„ n.i.i n^^a^ uith
the others.

The relation of the fin and body nnisclc to tlu- -l.ind i. >uch

562

THE .1 .1/ EUK '. 1 .V .V. 1 TURALIST [Vol. XLI,

the axillary pore would most likely be poorly provided with poison
except for the presence of a gland opening at the very end. It
seems plausible, therefore, that the absence of serrae is correlated
with the presence of glands developed about the spines, as in
Schilheodcs f/i/n')nis and N. nocfunius and as probably will be found
in aS. lepfacanf/iiis ' when an uninjured individual is examined.

The Origin of the (Hand Cells. — Cutaneous glands are generally
to be regarded as invaginations or proliferations of the epidermis,
certain cells of which become transformed into the secreting cells.
Wallace ('93) has shown that in the axillary glands of the toadfish
the clavate cells of the epidermis become the secreting cells, whereas

Pg. c.c. p.c. sp.

the smaller cells become tiie supporting elements. Similarly, in
writing of the weever fishes, Schmidt says,— "After a comparison
of the contents of the gland sac and the adjacent epidermis it
seems clear that in the gland tissue the secreting cells replace the
clavate cells and that the ordinary epidermal cells are gradually
transformed into the supporting plexus." Parker calls attention
to the lack of well defined ducts in the glands of the weever fish.
He says,— "There can be little doubt tiuit the gland is developed
as an epidermic involution the whole of which gives rise to secre-

» The only specimen available was a very small one. the spines of which
had been denuded of all tissue whatsoever.

Xo. 4S<)]

POISOX GLAXDS

503

tory cells, so that there is no marked dinVrentiation into inland and
duet."

The glands in Notunis and N'hilheodcs arc likcuisi^ invagina-
tions of the epidermis. Tluy are surrounded hy ;i slicaih of corium
pushed inward l)y the dc\ clopinu" i:ian(L A> sliowu in Fio;. 2,
d and /, the sheath is tliiinu'r and <l(Miscr than the nv^i ot' llic corium.

is a layer of pigment cells ( Fig. 2, j). c. i wliicli is frc(|uciuly fouml
to extend about the gland hetwciMi the sheath and thi> i;lan(l ti>sue,
thus occupying the same relative position as in tiu^ skin. The
arrangement of the pigment layer in relation to the spine glands
shows that they are invaginated from near the tip of the spine.
Near the tip, the pigment layer (Fig. 5, p. c.) turns upon itself and
passes over the gland to its base, toward the root of the sj)ine.
Thus the glaitd 'is invested bv two lavers of pigment cells with
corium l)etween tliem (Fig. T), d). 'I'he ehivatc .rll> <.f the skin

the two. The other rv\U .,f ,h' epidernu. a-^ ihc Ja-id i> ap-
proached, graduallv a.Muue an irregular diape. then bccnnun-
more elongated they a,v hnally tranMonn..! it, to th.- extremely

toadfUh. hi the '^Uiwl^ of the hatter ilie e].id.M-n.a! character of
the c-lU i. harely io.t. Thu> the .Hme celU. uhich in the mad

are perfech' diMincl in the -iaud> of the t.-adnd.. a.id proj.Ht into
the u'huid cavity.

564

THE AMERICAN NATURALIST [Vol. XLI

3. In addition to the axillan' glands Schilheodcs giirinus and
S. nocturnus possess glands developed about tlu^ pectoial and
dorsal spines. These are of the same tyj)e and structure as the
axillary glands. The end of such a glaud-l)earing si)ine projects
slightly through a slit in the epidermis.

4. Spine glands are not found in those species which possess
well develo])ed serrae upon the spines.

5. Schi/lxodrs Icpfacanfhus, because of its close relation to S.

the axillary glands.

G. A study of the mature glands tends to justify the following

a. The glands are of epidermal origin; those in the axilla
invaginate from the pore, and those in the spines from
the slits near the tips of the spines.

b. The gland sheath is modified corium.

c. The clavate cells of the skin become the secreting cells
of the gland.

d. The ordinary epidermal cells become elongated, forming
the supporting network of the secreting cells.

e. The glands of the mad toms are essentially like those

of tlu> weever fishes.

566

THE AMERICAN NATURALIST

[Vol. XLI

Jordan, D. S. and Evermaxx, B. W.

'96. The Fishes of North and Middle America. U. S. Nat. Mus.,
1896, bull. 47, p. 145.
Kent, W. Saville.

'83. British Marine & Fresh Water Fishes. Fisheries Exhibition
Handbook, 1883, p. 29. London.
Parker, W. X.

'88. On the Poison Organs of Trachinus, Proc. Zool. Soc. London,
1888, p. 359.
Sacchi, Marie.

'95. Atti. Soc. Ligust., 1S95, vol. 6, p. 89.
ScHAriDT, F. T.

'75. Om Fjarsingens Stik og Giftredskaber. Nord. Med. Arkiv.,
1875, vol. 6, nv. 2.
Tybring, Oscar.

'86. Poisonous Fishes. (Translated from the Danish hv Hermann
Jacobson). Bull. U. S. F. C, 18SG, vol. 6, p. 14S.
Wallace, Loiise B.

'93. The Struf'ture and Development of the Axillary Glands of Batra-
ehus. Jo),nL Morph., 1893, vol. 8, p. 563, pi. 27.
Wiedersiieim, R.

'06. Vergleichende Anatomie der Wirbelthiere. 1906, Jena.
Wright, R. R.

THE STRUCTURE OF THE SILK GLANDS OF
APAX TELES GLOMERATUS L.^

ROBERT MATHESOX AND A. G. lU'CCLES.

Apanfrlrs (jhmcmius is a hyraenopteroiis social j»araHU- ..f the
larvae of Finis rapac, the eommon eahhaue worm. The adult
females deposit at each oviposition from fifte(Mi to thirty-five vii^zs
in the young larvae of Pieris. The parasite- on hat. hiuii. feed tipon
the lymph and fatty tissue of their host and oiow very rapidly,
becoming full grown at about tlie end of the larval life of the
caterpillar. They then penetrate through the skin of th(Mr iiost
and, while emerging, spin their characteristic sulphur-yellow co-
coons. The silk glands, as seen in sections of the mature larvae,
are enormously developed. Althotigli the silk glands of lepidop-
terous and trichopterous larvae ha\e been the objects ot detailed
study by Helm, Gilson, and others, very little is known concerning
these glands in the Hymenoptera. As regards histological struc-
ture the only works of importance arc those of ( liolodkovsky,
his student Pikel, and Bor.Uts; an.l. excepting the latter who
gives a brief discussion of ihc-^v glan.i> in the acnh-ate Ilymenop-

larvae of various Tenthre.linidae. Theref.>re at the suguv-^tion
of Professor Rilev we were led to invotigate more fully the >ilk
glands of Apanteles.

The work was carried on in the Knto.nologi. al Laboratory of
Cornell Universitv. We wish to extend onr thank, to Profe»ors
Comstock. Riley" and :Mac( Hllivray, for their constant aid and
advice.

Anatomical Disposition of the Silk Glands.— The silk glands
of Apanfdc.s qhmcrafus arise near the base of the lal>inm and
extend through the body cavity to the antepenultimate segment
of the abdomen. Li the abdominal region of marnre larvae they
consist of two pairs of thin-walled, tnuch convohite.l. cylindrical
tubes (PI. 1, fig. 3) which completely surround the alimentary

568

Till': AMERICAN NATURALIST [Vol. XLI

canal. Each pair of tubes unites in the first abdominal segment
to form a connnon thoracic division. These connnon tubes, extend-
ing forwards with many convolutions in the thorax, turn ventrad
just behind the developing heafl and passing on each side of the
sub-oesophageal ganglion, end in short ducts. These ducts unite
in the labial region to form the press which occupies more than
half of the common duct.

In young larvae just hatched, and for several days later, the
glands show no convolutions whatever. They lie as straight
tubes, two on each side of the alimentary canal and extend caudad
to the antepenultimate segment (PI. 1, fig. 1). jNIoreover the
structure is the same throughout their entire length, no regional
differences occurring. Their walls are thick and their lumina
very small. Later they become much convoluted, and their
lumina are greatly distended, till in the mature larvae at time of
emergence from the host, the abdominal division has practically
ceased to secrete, becoming simply a reservoir for the already

The silk nlaiids may be divided into two general divisions; 1..
Secretory. l>. ( \,M.lii("ting.

The Secretory Division. — The secreting division nuiy be con-
venienllv divided into two portions, abdominal and tli/.racic

Thr ah,lomim,( pnrtiun (•<.... |)riM- that part ot' tlic uland .'xt<M.d-
ing ca.idad hum the point of j,i.,ctinv of the ulan.lular tubes in
tiie first abdominal segment.

In the freshly hatched and y(.unu' larvae this portion consists
of two pairs of straight glanduhir tubes, one pair situated on eaeh
side of the alimentary catuil iV\. IJi-. 1). On .>aeli side th(> tuhes

No. 489]

SILK ^/„l .\7>,

ture in the larvae of Ji<>ml>i/.r
stating that the prochiciii*;- pi
fine resistent cuticiila, in wli
various' thicknesses, unit(Ml to
or oblique trabeculae. He (h
are closed by a structureless 1

As the larvae feed and un
and fourth days, eoin?nenees

dular tubes become more and
of emergence of the parasite fn
fill and greatly distend the peri
brought out in the longitudinal
respectively, PI. 1 , figs. 2 and 4,

Along witii the great inerea
goes a corresponding inereas
become larger, but their radial
gradually becomes distended
at the time of the sj)iiniing of th

PI. 2, figs. 7 and S.

very thin hner
the enormous i.
short larval life,
tmlinal sections.

cat is thi,^

practically the whole body eav

570

THE A M ERIC A N NA TURALIST [Vol XLI

In ApanieJcs ghmcraius, owing to the thinness of the glandular
walls, it was inipossihle to secure tangential sections which would
give surface views of the nuclei. As the nuclei do not stain deeply
at this stage, they ai-e rather difficult to differentiate, although we
secured faiiiy good toults by staining with Grenadier's borax
carmine. :i> .liowii in figure 34.

The (yt()i)la><ni is dense, granular, and vacuolated, especially
during the latter part of the larval life when the glands are at the
height of their activity (PI. 1, figs. 10 and 12). Through the
cytoplasm run trabeculae, extending in many cases from the exter-
nal border to the inner margin of the cell. These trabeculae
appear as fine radiating lines, but later, with the thiiming of the
glandular walls, they disappear.

Gilson, '90, performed some interesting experiments in order
to determine the method of secretion. He ligated the entire liv-
ing larvae, disposing the ligatures in two pairs, the two threads of
each pair being close together. He then divided the larva into
three sections by cutting between each pair of ligatures. Treat-
ing the cut surfaces with mercuric chloride and collodion he secured
living isolated portions of the c-aterpillar, in each of which the
silk glands, es])ecially near the ligatures, continued to secrete.
1.1 .u<-h isohited portions hefouud vacuoles present in thecvtoplasm
un.l even in the iniel.'i of the >ilk ulan.l>. 'VUv>v vacuoh'. he e.,n-
Mdered ,he .ilk ^.'eretion. lie did not .Ueeeed in e.tal.lishing
whether they h.y Ix'twe.^n the radiating iral.ecnhi,^ or not. In
the case^ of Apanfrlrs ,,l.>mrratu.^ the condition which ( dlson sought

the secrcte<l product is used till at the time of emergence from the
host. Xmnerotis vacuoles are present in the cytoplasm, becom-
ing most abundant dining the time of the greatest glandular activ-
ity. The contents of these vacuoles remain unstained bv anv of

st;)in. «l a^ i^ iioted hilcr. Wlu'thcr the presence of tlie.e vaeiioh's

sists of but two^ecn-ting tulx'.. lying one on .'aeh ^i.h' ..f the ali-
mentary canal (I'l. 1, iig. 1 i. Each is formeil by tlie union of the

Xo. 489] SILK (iLAXDS OF M'AXTI-LL

two tubes of the al)(h)iniiial (livi>i()ii in ilie fir>i al)
and extends cephaUid in ilic >!i<'rt <hict wliicli he
of the sub-oesophageal oanylion. 'I'lic dmracic
divided into three well defined parts,- the 1st, or i
the 2iid, or middle thoracic; and the ;ird, or y

The 3rd, or posterior thoracic, division during
larval life consists of an almost straight cxlin.lri
it becomes nnich comolnted and it^ ^^al^ i)e<-om
in every way it markedly resembles the abdomin;
cytoplasm is denselv grainilar, deeply staining, a
lated, especially near the {)eri))liery of the cells (PI.
16).

The 2ml, or middle thoracic dixi.ion. is 'piite si-
lt extends from the beginning of the second tho"
the first division. The cells of tliis portioii of tl
greater radial diameter than in any other parr
"gres" or '*gum," so prominent in the Ist di\is
and in some places difficult to distinunish. The ct
ized by a faintly staining, loosely granular cytoph
the periphen-of the cells, is much vacnolated
20).

The 1st, or anterior thoracic divisi<.n.
It extends from the bciiiiutiim of the lirsi ihoraci
duct. This portion of the uiand !> chaiactevi
layer of "gres" or "ixnm" adhering ( h
the secreting cells (IM. 1, i'vj;. -'■'); pi.

572

TV RAJ AST [Vol. XLI

There is no indication of a vestige of these glands, such as Gilson
found in Limnoyhilus rhombicus, one of the Trichoptera.

The Conducting Division. — The conducting portion of the silk
glands is Y-shaped, with a median stem and branches pointing
caudad; each branch joins the thoracic portion on its respective
side of the body. The press commences at the juncture of the
two branches. The entire conducting portion is very short, being
wholly confined to the labium. In cross section the branches are
seen to be composed of a number of cells surrounding a small lumen
(PI. 3, fig. 25). The nuclei are oval to rounded in shape — never
branched. Posteriorly the cells are colunmar and contain elongated
nuclei, but anteriorly the cells become fiattened (PI. 3, fig. 30).
There is thus formed an enlarged lumen at the anterior end of
each branch of the conducting tube. Also by the increased radial
diameter of the posterior cells the amount of "gres" or "gum"
that can pass forward is regulated (PI. 3, fig. 30).

The cuticular lining of these branches forms chitinous folds
or ridges which are not perfect spirals but appear as incomplete
rings. rPl. 3, figs. 25, 29 and 30, in).

The Press.— Although the internal disposition of the silk glands
\\a. faniilar to the earlier anutoTui^ts, nothing was known regard-

lonnati.ui .4' th.' ^ilk tliivad h u..^ not till thr iiPp,!n..nt unik.
nfCilM.,, and Hlaiu-ihar iIh- iniiiui.- --iriu miv tlii. <.ruan in Leo-
defined.

No. 489]

SILK (,!.AM}S OF A l'A\T/-f.l-:s

573

Berlese, '06, denies the pres(>n(e of a press in the silk glands of
the larvae of Hymenoptera, Diinera, and ( "oleoplera. He figures
a sagittal section of the head of the lai-\a of X iilotoma rosae hut
does not represent muscles as j)n'sciit in the ri^uion of the con-
ducting tuhe. On the other lunid the pn>>cnre of a press in liviiicn-
opterous larvae has heen recordc<l l.y l-ckst.Mn, 'HO, in Ij/da
pratcnsi,, and IMkel, in Loplnirus ph,l. 15., d. of workers
figure this oruan as preset hut uixc no defn.ite <l(lails n-anlinu
its structure. Tikcl states that in structure it is siuiihu- to that
descrihed hy Tichornirow for Bntulnix wnri.

Sfrurfmr'offin /'/v..s.- In the (asr of Apanhh. nhmuraiu.s the
press is highly developed, it couuneuces at the union of (he two

than half'of die conunon duct. Dorsally the press is' <.otu'ave.
traxersrd hy a lono-im.hiud fuirou into ^^\^n■\^ pass dorsal pair

fi-s. 1'7, J!»an(l;!(). The Neutral surface is ( on\.-\ , Fius. JTand.'^O);
its ((>IIs are son.cuhat elonualed and s(-< ,vtc the thicker <-hitinous
layer of tlu^ conunon .hict. The dorsal uium h-s consist ot nuuicr-

of M-N e.al (ihcts. Thc^ are l.u aled as sh<,v n in c ross tion in Fi..
27. 'V\w^ are insrrte.i on opposite si<les of the press and. passing
almost direcllv veutrad. have their origin on the veiUral ho.ly wall,
being attached <lirectly to the chitiu. .

'i"he huuen of the press, as seen in ( ross se( tion i Tiu. 27 is hoise-
shoe-like in form. When the uuiscies are n>ia\ed this lumen is pM< -
ticalh <'losed, thus prcNcntinu- the furthei (-ress nf the thiead. The

dexelope.! on the \entral si.Ie. This 'ehiliuocs kn.M' is dire, th
continuous u i,h that of the <lucts lu Fi.. 27 the dorsal poniou of
this iaver appeals thi<k.l.ut this is due to theol.li.inu> of ihesMti.Mi

574

THE AMERICAN NATURALIST [Vol. XLI

and it is not till about three days after hatching that the furrow
and attached muscles become clearly differentiated. Being func-
tional for such a short time its complete development does not
take place till late in the larval life. It is not till shortly before
emerging from its host that the press becomes completely developed
as it is not necessary that it should be fitted for spinning during the
entire lifetime of the larvae but only for the very short time occupied
in the buihhng of its cocoon.

In srructure the press differs from that described for Lepidop-
ttM'a in that the lateral pair of muscles is not present, and from
tliar of the Trichoptera in that each dorsal muscle is single and not
divided into two distinct bundles as figured by Gilson.

Fuiiciions of the Press. — The functions of the press in the Lepi-
doptera liave been carefully worked out by Gilson and Blanc and
there is no doubt that the functions of this structure in the Hymen-
optera are similar. These functions may be summed up as fol-
lows—

1. The press modifies and regulates the form and diameter of
the two threads.

2. It regulates the layer of "gres" or "gum" whicli surrounds
these threads.

3. By the relaxation of the musdes tlie chitinoiis walls on
accomit .a' their elasticity, contract and hold the thread innnovahle

( Jilson attributes to the press another function, that of forcinu' the
thread to the exterior when by accidtMit or voluntarily the thread
is broken in the spinneret. This is deniecl by HIaiic who holds
that the contraction of the .muscles of the press, distending its
lumen to the fullest extent, together with the pressure upon the
contents of the gland due to the elasticity of its walls, and the general
muscular contractions of the body, serve to force the thread to the
exterior when l)roken in the spinneret or even in the conducting
tube.

No. 489] SILK (U.AXDS OF M'.WTFJ.ES

575

tion of how the thread is first forced to the exterior and also how it
is extruded when broken.

It is not necessary for us to describe the mechanism of spinnini;
as that has been fully done by Blanc and Gilson for the Lepidop-
tera and their descriptions serve equally well for the liv-UKMioptcrous
larvae. The method of spinning the cocoon in A panfrh.s filnimm-
tus has been well described and figured by Reaumur.

The Spinneret.— From the i)ress a sliort chitiuous tube leads to
the .spinneret .^p. Fig.. lN. 2\) and :5(> . The lunu'ii of this tube
gradually widens as it a|)pr()aches the ('xierior Kiu'^. L*!i and 'AO).

press occupies .4 mm. The spinneret is situated just beneath the
buccal cavity and consists of two small chitinous projections
directed cephalo-dorsad (Fig. 28, sp.).

Functions of the Different Parts of the Silk Gl&nd.—Ahdotm'nal
Dirision. A. previously pointed out the abdominal division of
the silk glands consists, i'n the young larva(^of tN\o pairs <,f straight
cylindrical tubes lyinu' on each si.le of tin- alinientaiy eanal. As
the larvae urow these parts beconie active, their hiniina become
gradually di<tend(>il with the s(>ert>ted i>roduet till, in the mature
forms, the ulanduiar walls are so thin as to warrant the assertion
that tlieyhaNepraetieally ceased to sccivte and are nicrelv rcscrxoirs
for the accumulated product.

'Fhe (|uestion as to whether the "gres" <.r - ^^luu" is scvreted
by one j.artieular reo-ion of the gland and the silk by another is still
unsettled, (iilson in 1S!»0 and auain in l v.»4 came to th(> conclu-
sion that both are secreted throughout the gland; and that the
silk, properly speaking, is the result of a selection etl'ected in rhe
layer of secreted pn.duct lyin- next the internal fa<'e of tin- cells.
AVhether this proccs. ,)f schn-tion is a ehemi<-al phenomenon or

a sru.ly o'f -'everai series of sections he conclud'es that the outer or

spe.-ial afHm-tv for colorinu' a-ents" Neither ..f Ihes.- conclusions is
absolute since he. lid nor (ind then, to hohl I nie in all eases.
Hlanc. •SI), p. L'[. stams that "The silk sc

576

THE AMEBIC AN NATURALIST

[Vol. XLI

formed in this region. This substance is the 'gres.'" He con-
siders (pp. 27-28) the "gres" as nothing more than the peripheral
layer of silk oxidized in the reservoir, the oxidation being due to
the presence of a large number of tracheae in this region.

In the abdominal portion of the silk glands of Apanteles glomera-
ius the secretion appears as a hyaline, faintly staining product.
Fixation in Flemming's solution and staining with iron haematoxy-
hii gives it a greenish color, the peripheral portion often being
l)Uu k. In tlie anterior part of the abdominal portion the peripheral
hiyer is always stained black. Safranin colors the entire secretion
salmon color, the peripheral layer always more deeply, especially
in the anterior part of the abdominal portion. Mayer's acid
haemalum and eosin do not color it at all.

Judging from the staining properties of the secretion the conclu-
sion that there is a differentiation between the peripheral layer and
the central column appears justified. ^Yhether this j)eripheral layer
is the " gres " or not is a question which we do not pretend to answer.

Thoracic Division. This pf)rtion of the gland remains activelv
secreting much longer than the abdominal division. The char-
acteristic appearance of an actively secreting ])orti.)n is shown in PI.
2, fig. H;. Thv peripheral area often ai)pear.^ k>ss dceplv stained
and nnmerons vacnoh-s are always present.

Tlie character of the sc.-retion' in tiie posterior thoracic <hvision
apjK-ars similar to that of the al.doniina! portion except that the

are generally more nnmerons. It also takes tlu same chai'acter-
istic colorations.

The anterior and middle thoracic portiojis secrete a somewhat
(liferent ])roduct. The cliaracter of the cells of the middle portion
wonid indicate that snch is the case here at least. Fixation in
Fleming's solution and staining with iron haematoxylin gives to
this division a vcrv cliaractcristic n[)])earance. The cvto])lasm
i> filled uith rather large ronn.lc.l l.ia.-k grannies thns ea.'ilv dilfer-
.ntiating this portion from the other two thora-ic .livi>ion>. The

In .iniilariy tn.ated gland, many identical black grannies are s.^en
near th(> interna! >nrfa<r of the cell, jnst within the internal mem-

the first thoracic division I PI. 1, figs. 10 and 12j.

Xo. 489]

SILK GLANDS OF A PAN TELES

577

The dense secretion ooverino- the inner snrface of the 1st thoracic
portion indicates that tliis is its own peculiar product and not that
of the fohovvinfr (Hvision. T]n> secretion appears closely striate
in a rmhal manner. Acid liaenialuni and eosin do not color it;
iron haeniatoxylin, safranin, and (irenacher's borax carmine stain
it but sli^ditly. The central colinnn of silk, however, is more
densely stained with iron haeniatoxylin and safranin.

As the glands of Fhilippi are not present it is only speculation
to suppose that the secretion of the second thoracic portion may be
of a nature similar to that of these jjlands in the Lepidoptera. The
function of the product of the olands of Philippi is not definitely
known. Nearly all workers dill'er in their conclusions in regard
to this question. The conclusion of Blanc, 'Ul, and Berlese, '()(>,
based upon the experiments of Robinet, '30, seems the most logical.
These authors consider its finiction that of lul)ricatin<: the threaii
which is to pass through the i)ress.

The fact that the thread, in Apan1rJi\^ (jlomrmitts, begins lo take
on its definite form at the posterior end of the 1st thoracic ])ortion
might indicate that the secretion of the 2nd division had a coagu-
lating effect upon the silk and "gres."

SUMMARY

1. The silk glands of Apanirirs- rjlomemlu.^ differ from those
in the Lepidoptera and Tric-hopt.'ra in that then- are four tubo
in the abdtmiinal region. TluMr histological structure is similar
to that of Lepidoptera and TricliojJtera but difiers markedly from
that described for the tentliredinid larvae.

2. In immature larvae the epithelial cells of the whole produc-
ing region are actively secreting. Ximierous ^■acuoles are present
in these secreting cells, especially near the periphery.

^ T,i nlon.I-; (Iv.-,! in Fl , M 1 M .1 i 1 1 < .'s solutiou aud staiucd with iron

578

THE AMERICAN NATURALIST

[Vol. XLI

distended and nearly fills the entire body cavity. It is probable
that this portion now acts merely as a reservoir and that its cells
have ceased secreting.

5. The glands of Philippi are absent and it is probable that
the second thoracic portion performs the functions of these glands.

6. The press is well developed. It differs from that of the
I.epidoptera in that the lateral pair of muscles is absent, and from
that of the Trichoptera in that there is^a single pair of dorsal
muscles rather than two distinct pairs.

7. The product of the gland is a double thread as in the Lepidop-
tera and Trichoptera.

CoRXELL University
Ithaca, N. Y.

BIBLIOGRAPHY

Berlese, Antonio.

1907. Gli Insetti, loro orgaiiizzazione, sviluppo, abitudini e rapporti
coiruomo. 1907, vol. 1, pp. 521-523.

BORDAS, L.

1895. Appareil glandulaire drs Myineno[)teiv<. .1////. Sri. Nat. ZooL,

1895, vol. 19, pp. r_'-2().
Carriere and Bi-RGER.

1897. Die EiitwickluMgs.uv^rhichic dcr M.-iunhi.^iH' (Chalicodoma

muraria Fal.r.) iiii Ki. Nora Aria Anul. I.,n/,. Car., 1897,

CHOLoDKnVSKV, X .

1895. i:nrn,H.,t.,nnsrh.. Mis.Tll.'n, I. Vvhrv Aw Spinnariism ,ler

1901. I'cb.T Ln Spiiniapparar drv Ly.la-Larvn. A/l>/r»,. Znl. /.

1893. ( In Cviulouri, ;,! I )itlrirnr,.^ ii, Homologous Organs. 63rd Kept ^
Ilrit. As... A'h-. .V/., lMt;5, pp. 81.3-816.

No. 489]

SILK GLANDS OF APANTELES

579

No. 489] SILK GLANDS OF APANTELES 581

Fig. 28, pi. 3.— Longitudinal section of the press, showing the position of,
the dorsal and ventral muscles, s. L, silk thread passing through the

Fig. 29, pi. 3.— Same as Fig. 28, but showing the attachment of the muscles
directly to the chitin of the conducting tube. X 267.5.

Fig. 30, pi. 3.— Longitudinal section of the press, of one of the conductmg
branches, and of the beginning of the 1st thoracic portion, a., point
of union with the conducting branch of the opposite side. X 2(j7.n.

PLATE 1

PLATE 2

PLATE 3

THE NEST OF THE KELP FISH

CHARLES F. HOLDER

588

THE AMERICAN NATURALIST [Vol. XLI

She frequently pushed her way through the clump of weed but more
often passed around it, the silken tenacious cord binding it together
in a globular or oval mass about the size of a hen's egg. The
entire nest shown in the photograph was formed in about two hours,
the fish dropping to the bottom of the tank after each effort and lying
there for ten or twenty minutes.

The accompanying photograph of probably the first nest of
Heterostichus to be recorded was made under my direction by

NOTES AND LITERATURE

GEXERAL BIOLOGY

590

THE AMERICAN NATURALIST [Vol. XLI

other vertebrates. . . .is highly instructive and disclost-.s to the thought-
ful person deeper and weightier secrets than cire to he found in the
so-called "revelations" of all the religions of the eartli. Compare
attentively the successive stages of the chick, i)ig, rabbit, and man
shown in the accompanying figure. In the first stage (the upper row),
in which the head with the five cerebral vesicles and the gill arches are
clearly marked out but the limbs arc still wholly absent, the embryos
of all vertebrates from fishes to man differ from one another either

which the limbs have begun to develop, distinctions between the
embryos of lower and higher vertebrates have begun to appear; yet
the human embryo even now is scarcely to be distinguished from those
of the higher mammals. . . .These are facts the significance of which
cannot be overestimated."*

As drawings of embryos, the well known figures of Haeckel here
reproduced are totally valueless. The front limbs have been trans-
ferred to the neck, and the characteristic features by which any one
familiar with embryos can distinguish a pig from a rabbit have been
wholly overlooked. Although Parker declared that 'one diagram
would represent all,' his figure of the embryo mole could not possibly
be mistaken for a f)ig. Moreover tiie pig at this stage could be dis-
tinguished from the rabbit or man by its pancreas alon(^ The com-

The ^,pherical yAk ..t tlic Ik n'> vo., the elongated vesicle of the sheep
(that of the piii iH'iiig (juite as I.mg but not so slender), the round
smootii vesicle ot the lahbit and the villous human vesicle are radically
(liHeietit troiii e;i( li other. Since these membranous structures are

ot the embr\<) they must be regarded as expressions of differences

Haerkel. ]]. Ant hr()i)ogenie. 3rd ed. Leipzig, W. Engelmann, 1877,

THE AMERICAN NATURALIST [Vol. XLT

Montgomery states that two species are as distinct in the egg-cell stage
as in any later one, "no matter whether the differences are as percepti-
ble or not."^ Such a statement, however, evades the question whether
or not embryos of related species can actually be distinguished from

The four suborders of rodents, represented by the squiriel, mouse,
guinea pig, and rabbit respectively, according to Lee may be distin-
guished at very early stages. His studies do not enable him as yet to
recognize the genera of one suborder, — namely the gophers, prairie
dogs, squirrels, and cliijimunks — until the embryos are far advanced.
Differences in tunicate eggs of closely related genera have, hoAvever,
been recorded by Conklin, and ^McClung can distinguish several
species in one genus of grasshojjpers by the chromosomes of their
germ cells.

F. T. L.

ZOOLOGY

New Text Books of Zoology.— The most iiiii)(>rtaiu service thiit
biology can render to students is to train their rcaxm and ilicir power
of observation, and to free them from a too dccj) rcvcn-iice for authority.
This service can also be p('rforine<l l)y tlie other natural .sciences,
physics and chemistry, which an- coniiiionly nichidcd in the cnrricuhi
of high schools but which rccitiire expensive apparatus Ix-yond the
reach of many schools. Biology, however, can be profital)ly taught
with so slight an equipment that every school can aflonl to underiak(>
to teach botany or zoology, or both. The teaching- of biolouy lias
often failed to yield the results that educators liave expecte.l. This
is so because teachers too often yield to the temptation to tell ilie
students the facts and theories which they ought to learn by their o\\ n
efforts, instead of teaching them how to discover, to class ity, and to
draw proper conclusions.

Professor Gletm W. Ib-rrick. of the Mississippi Agricultural College
has endeavored to meet tlie needs of the high school of limit. -d means
whose students will, as a whole, have no further instruction in bioh.^^v.

3\o. 489] NOTES AND LITERATURE

tiling more than twenty species of vertebrates and invertebrates,
together with questions, tlic majority of which appear to be answered
by the context, the remainder by the companion text book.^ Tins
method scams to us unwise and the book is apparently less useful than
Kingsley's Elements of Compamiive Zoology (2nd. ed., Henry Holt
-and Co., 1904). The latter is a cheaj>er book which gives facts that
are otherwise inaccessible to the student and asks questions which
can be answered witliout appeal to costly apparatus, about easily
obtained and inexpensive animals. The answers, the student must
gather while learning both to find and arrange facts and to draw right
conclusions. A more extensive work is that of Linville and Kellv;
their Text Book in Geneial Zooloqij and Guide jo- Lahomfonj and Field
J\ oik VI Zoolofji/ (Ginn and Co , 1906) gi%e facts and suggestions,
especially tor the reading of original articles, together with necessary
deductions; and ask most interesting and suggestive questions which

of any of these books will give the student about the same facts, but
Klng^le^, and LiiniUe and Kellx, compel the student so far as a book
can do so, to observe and think, which is the most desirable service
to the memory-laden vouth. It reallv makes little difference how

method of stuch^is of primarv importance.

Soim ot PioU.MH lie link's <hagiam> are not A\ho]h correct For

n ,n Munnh . hn . , I, In ah \I <„ phnh,q„ d x<r 1(,S), i. labelled

a^ a iunah h i. Hu spcnuliKt, an<l not the oMdu( t as m Herrick'
hguie, ^\ iiu h opens at tlu base of the last thorat u limb Tlie foll()^^-
ing statement from page 33 of the text book is an examjjle ot defective
fact and theorv,— " It may seem strange that the oldest animals (Pro-
tozoa) are the simplest, but it is true. It is probablv due to tlic fact

probably not \er> difh h nt t.uln tioni \\\\\x it a\ is u't- i^o Hence

tll<^ ha^e uinaimd mm h tlu s.une ' It is lundlis. u) sax that'Pro-

n, i,u h.Jilx d. \dopMl animals haxe been exoKed in Axater

In, niou a.Uan.ed stu<lents Dr (nlman A Drexx, Professor of

594

THE AMERICAN NATURAIJST

[Vol. XLI

596

THE AMERICAN NATURALIST [Vol. XLI

tion of his butterfly-house, a green-house devoted to rearing plants and
insects together, he writes as follows;— "In my younger days I myself
amassed an extensive collection of butterflies and moths. . . .1 have
the collection still, and never look upon it without pride. Friends love
to gaze upon the Scarlet Tigers, Clouded Yellows and mammoth
Death's Heads ; white bearded fossils come down from afar and beam
upon it — but when all's said and done what else is it but a collection
of corpses? Beautiful though they may be to look upon, arranged
systematically with pinions outstretched upon the clean white paper —
how much more beautiful to gaze upon the living form flashing its
gorgeous wings in sunlight, throbbing with the exuberance of lifeV
This is a frank statement from a collector, but one which is character-
istic of the times ; interest is being transferred from collections to nature
itself.

F. T. L.

Birds of Labrador and of the Chicago Area.— The Birds of Labrador
are well presented by Dr. Charles W. Townsend and Glover Allen
(Proc. of the Boston Soc. of Xat. Hist., vol. 33, pp. 277-428). In the
introduction the authors describe first their visit to T.al)ra.l()r in the sum-
mer of 1900; then the topograjihy, the fauiuil an-as. paths of migra-
tion, and ornithological history of the region; and finally the hird and
egg destruction which in 1833 fllled Audulnm with "liorror and (hs-

and the authors hope that "the wonderful nursery tor \\ai> r l)ii(is in
Labrador will not be entirely depopulated but that snllic ieiit jjrotcction
tor the breeding birds will be givt>n and that sj)ee<lily, befon; it is too

The introduction is followed l)y an annotated list of all Labrador
birds, and the book concludes with a table showing the approximate
number of each species observed by the authors, a l)ibliography, and
a map. This publication (which is sold separately) will be of interest
to the large number of students of local birds in eastern United States,
for Labrador is the destination of many familiar migrants. It is

Thr Hints nj fhr Chiraqn Ami arc similarlv ircatcl l.v FranL M.
Woodnitf ilJnII. (1 <.r Nnt. I /is/. S„ m C /nrrn,n AmJ . nj Sri., 221
pp.). ^ The ininH|,i< ii,,n runK-iin^ not«-^ ..n ilir taxoral.K' ln. ;,lii i... f.,r

catalogue of spcci.-s 1. rings fogrtlh'r a larg.- iwidy c.f fa«'ts covering a
long period of observation. Sometimes, however, the list of synonyms

No. 489]

NOTES AND LITERATURE

597

occupies more lines than the account of the bird, as with the savanna
sparrow and purple martin. An amusing feature of the extensive
bibHography is the transhition of the titles of newspaper articles into
intelligible form, for example, - Li»qn- in Winffrs- Lap. (An acc.niit
of birds which delayr.i tlirir niiirralio.i.) - Uird.s a/ M i,.frr>i nt l.a/.r

or bird haunts include a photograph of a colony of great blue herons,

^ F. T. L.

The Excess of Male Births. - In the Mav Natiori//\-f (vol. -11, p.
303) A. H. Pike discu.vsed the .ignifi( ..ncc <>t th.- cv cf male births

born for every 100 females. In the -lune issue ot the I'ror. of the
Cambridge Phil Soc. (vol. 14, p. 122) ^Valter Ileape presents the
best available statistics for dogs. Among 3(),Sr)7 pups of regi.stered
stock there arc 117 males for every 100 females. Of some twenty
breeds considered, all showed an excess of males except two, the
figures for which were based on limited returns. The excess of males
is apparently greater in large breeds of dogs than in small ones. Mr.
Heape believes that the latest moment at which sex of oH'sjirinu- can
be determined is the time of fertilization. However, .inc.- nnfritiun
of the i)arents .nay aher the sex-.let(M-minino- tael..r> in tlieir -penna-

trolled, at le:ist to some exlenl .

V. 'V. L.

Recent Publications Concerning the Structure of Insects. - Tlir

No. 489] NOTES AND LITERATURE

600

THE AMERICAN NATURALIST

[Vol. XLI

The Suboesopkageal Body of Insect Embryos. — The question as to-
the origin and the morphological significance of the suboesophageal
body which has been found in certain insect embryos is an open one.
Hirschler* has studied this structure in embryos of Donacia and has
added much to our knowledge of its nature. He finds that it is ento-
dermal in origin and that from an unpaired rudiment at the end of
the stomadeal invagination there arise four rounded, paired masses
which finally communicate directly with the lumen of the mid-intestine.
These persist until at least the third day of larval life — their further
fate has not been studied.

Hirschler's results apparently confirm the theory of Nusbaum and
Fulinski, '06, that the suboesophageal body is to be homologized with
the hepatopancreas, or glandular diverticula of the mid-intestine, of
the Crustacea.

Stridulation Rhythm of Crickets.— According to A. F. Shull (Can.
Ent., vol. 39, p. 213), in the chirping of the snowy cricket "exact
synchronism is comparatively rare" and exists only between two or
three neighboring individuals. Thus two crickets five feet apart were
observed to time their chirps in unison as if they heard each other.
The rate of stridulation is independent of wing length; in general it
increases with rise in temperature, but Dolbear's and Bessey's formulae
to express this relation are only approximately correct. Under the
same conditions the rate in different individuals varied from 93 to 110
chirps per minute. Except on cool nights, from 600 to 800 chirps
are usually performed continuously; one cricket was found to chirp
2,640 times without interruption.

Notes.— Bull. 110 of the X. Y. State IVIuseum, preparatory to a
monograph of the Cecidomyiidae, presents descriptions of 203 new
species belonging to this group. The Cecidomyiidae, or gall gnats,
are dipterous insects from 0.5 to 3.0 mm. in length which produce
various leafy galls including tlie "willow cones." Bull. 109 of the
\. Y. State Mu^rum is devoted to the tussock moth and rim leaf beetle.

W. A. Riley.

' Hirsehler, J.
Entuickhmf,' l>ei

im., 1907, 31, pp. 766-770.

und ihre embryonale

No. 489] NOTES AND LITERATURE

601

BOTANY

Plant Geography.— The Scamlimivian fiom. Several naturalistic
have considered tlie origin of the biota of the Scandinavian peninsula.
During the glacial period most of the higher forms of life must have
disappeared, leaving the peninsula to be repopulated by immigrants
from other regions as the ice receded. This immigration was early
thought to have had two sources: the central European lowlands and
the Russo-Siberian region. The biota of the former is supposed to
have come in by wav ot one or more Baltic land connections, and that
of the latter is thought to have gained access by way of Finland and
northwestern Russia. But besides these a third element, called bv
Blytt "the Atlantic group" ot plants, was discerned, as the flora, especi-
ally of the western part, became better known.

This so-called "Altantic" element is discussed bv Stejneger (Swith.
Misc Coll (,nnt ■! 4-.S--)l ^ ]')()7) hoin both th( /oologi. il an<l

mnUun in Noiu u .vpt .l..n^ tlx c .i Imiumii .n i .n.l
Krlstlcln.sundor^^lur( tlu ^ m u Ik slumnfluxt iKuidtinMl l,..ni
this secondary center of distribution. 1 Ins associatiim shows a sironii
resemblance to the biota of Scotland and noriliwcstcni Ireland, and
Stejneger thinks that the similaritv is not dur to parallel drvelopiiiciii
but that It ukIk n.s . dii((ti^(n<li( (onn.Mion I.uumm tli. luo 11,.

602

THE AMERICAN NATURALIST [Vol. XLI

south and the niicrvciuno' ivuion is drv, and ju(l<,nn(r troni tjie contigura-

nionsoon from September to December. The siuiimit of the moun-
tahi, however, is l)athed in mist and consequently affords an isolated
"moist region" vejretation, practically confined to a few acres within
100 vertical feet of the stiimnit which it must have reached by leaping
at one bound o\er the inierxcning 40 miles of dry lowland tliat separate
it from the :\ratalc hills to liie south or over the 280 miles that separate
it from the hills of southern India.

A flora of 144 Howering plants and ferns is found at or near the
summit. Of these, 41 belong to the dry re-ion an<l cons(-(,uently have

103 remaining sj)ecies, 24 have in all i)ro!)al)ility liecn iiurodnced by
birds; and 49, of which 24 are ferns or Ivcopods. have evidentlv l)een
brought by the ^^ind. Thu- only 3(1 remain ^^ hosc method of intro-
duction is doubtful; tliese Dr. Willis <liscuss(>s in detail.

Xo. 489] NOTES A XI) 1. 1 TKU. 1 T I h'l-: 603

Held ^vlll(•ll r::u 1h" inv..k.'.l l,. (-xj.hiin tlic tads.-

J. AHTfUK IIahkis

Lock on Progress in the Study of Variation, Heredity and Evolution.*

— This attnunive little voltnnc c.mtai.is iwc intv..(iu.l„rv cliai-ters on
the genera] concei.tioiis of cvolniiou. .mr on tl..' lUvnvy of natural

604

THE AMERICAN NATURALIST \Yo\. XLI

The work can be regarded only as an introduction to the modern!
experimental and biometric study of evolution, since much space is-
devoted to elementary principles, but it is a commendable effort to bring
the newer work before a wide circle of readers.

J. A. H.

Notes. — The origin of the cow-fea has been investigated by Wight
(r. Dept. of Acjric, Bur. of PI. IruL, Bull. 102. 1907.) who con-
cludes that this legume is a native of India and the region nortliwest-
ward to the trans-Caspian district. Its cultivation in that region is-
very ancient and it also extended to ('hina at an early period. As
early as the beginning of the Christian era it was known in Arabia and '
Asia :Minor and was cultivated in at least one of the countries of
soutlicrn Europe at about the same time. Its introduction into cen-
tral Europe occurred much later and independently. It seems to-
have been introduced into the West Indies in the latter half of the
seventeenth century and probably reached the mainland during the
first half of the eighteenth century.

No. 489] NOTES AND LITERATURE

605

extensively used as food in Japan and now being introduced into the
United States. Funatsu (/. c, 469) gives the composition of a chrysan-
themum flower used as food. Albahary (Compt. Rend., 145:131-133.
1907) publishes analyses of the fruit of the tomato. Jaffa (Yearb.
U. S. Dep. Ag. 1906; 295-312. 1907) considers the value of nuts
as food.

Ybarra (Smith. Misc. Coll., quart, iss. 3:428-457. 1907) has done
a service to those interested in the natural history of America by pub-
lishing an annotated translation of a letter by Dr. Diego Alvarez
Chanca, physician to the fleet of Columbus, dated 1494, relating to
the second voyage of Columbus. The letter embraces observations
made between November 4, 1493, and the last week in January 1494.
Of course but little space could be devoted to botanical matters but
several of the references to economic plants are of considerable interest.

J. A. H.

The difficulties of botanists in capitalizing specific names are illus-
trated in " The Flora of the Gulf Biologic Station," recently issued
by the Louisiana Board of Agriculture. It refers to Vrrhcsinn Vir-
ginka, Commelina rirtj!nir(i. Ijirium Vii/f/arr, K/mcIinris M iifafa, etc

Ipomoea pes-caprae may Ix- coiiiiKircd with I'aui'-iiiii ('nis-(/(ilii. I'lii-
form decapitalization would prcvcm such contusion. This Mora,"
which records some twcuty-si\ phanerogams not ])irviously reported
in Louisiana — astate which " is to-day ahnost unknown hoia nieally "
— is to be sup})lemente.I l.v further pul.Hcations. Its author, R." S.
Cocks, refrains from naming prematurely several new forms.

PUBTJC.VriOXS l{K(^i:iVKl)

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BALTIMORE, MD., U. S. A.

The first of a series of Colored
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