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LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


AANSONIAN itr ti-™ 


301919% 
* JULI 41994 * 


4, 
VOLUME II 


OCTOBER 1915-1923 AUGUST 


fional Muse’ 


NUMBERS 1-18 INCLUSIVE 


ao fobs He D BY fH E 5 0 Gaia Ts 
foe ZOOLOGICAL” PARK, NEW YORK 


[or 4 


New York Zoological Society 


GENERAL OFFICE: 101 Park Avenue, New York 


Officers 
Honorary President, HENRY FAIRFIELD OSBORN; 
Vice-Presidents, MADISON GRANT and FRANK K. Sturcis; 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 
Treasurer, CORNELIUS R. AGNEW 


Board of Managers 


Glass nf 1925 
Percy R. Pynr, GrorGE Birp GRINNELL, CLEVELAND H. Dongs, C. Lepyarp Buarr, 
AntHony R. Kuser, Mortimer UL. Scuirr, FREDERIC OC. WALcort, 
BEEKMAN WINTHROP, GEORGE C. CLARK, JR., W. REDMOND Cross, 
HENRY FAIRFIELD OSBORN, JR., ARTHUR A. FOWLER. 


Glass nf 1926 


HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. DIETERICH, GEORGE 
F. BAKER, WM. PizrRSON HAMILTON, Ropert S. BREwstTeR, Epwarp S. 
HARKNESS, WILLIAM B. Oscoop FIELD, EDWIN THORNE, PERCY 
A. ROCKEFELLER, JOHN E. BERWIND, IRvING K. TAYLor. 


Glass nf 1927 


MADISON GRANT, WILLIAM WHITE NILES, FRANK K. SturGIs, OGDEN MILLS, 
Lewis RurHERFURD Morris, ARCHER M. HUNTINGTON, GEORGE D. PRATT, 
T. CoLEMAN puUPoNT, HENRY D. WHITON, CoRNELIUS R. 
AGNEW, Harrison WILLIAMS, MARSHALL FIELD. 


Scientific Stalf 
WituiAm T. Hornapay, Director of the Zoological Park ; 
W. Rem Buatr, Assistant to the Director ; 

CHARLES H. TOWNSEND, Director of the Aquariwm; 
Raymonp L. DitmMars, Curator of Reptiles; 
WituiAM Bresr, Honorary Curator of Birds and Director of 
Department of Tropical Research; 

Lee S. CRANDALL, Curator of Birds; 

GrorGE 8. Huntineton, Prosector ; 

ELWwIn R. SANBORN, Photographer and Editor. 


Editorial Committee 
MADISON GRANT, Chairman; WiLu1AM T. HorNADAY, CHARLES H. TOWNSEND, 
WILLIAM BrresE, ELwin R. SANBORN, Secretary. 


TITLES OF PAPERS 


1—Two Series of Amphibians.......5...--.-- eee eee c esse e noes Deckert 
2—A Tetrapteryx Stage in the Ancestry of Birds................. Beebe 
Silvas Git: eis Aa All ae eeanino cing clgdind odo ome Ue oer oAideor Beebe 
A= Wena Ot .iunmle Webris. (sj. ase ee oe ce see Beebe 
Ga IMs Cane @ iSEil sk po pmasg co sou loeb moe NOduao cme omenmcm” oonoe Gudger 
(INSTI GLE UNIS WRRIIMeN: 53 Ske clo oehert had Glove 6 bicete OOo O cual Or Srreee Le Souef 
7-8-9—Higher Vertebrates, British Guiana.................+-004- Beebe 
10—Habits of the Sage Grouse........... teen eee ete e eee Horsfall 
11—KEclipse Plumage in Domestic Fowl...........-----.+-+0005- Crandall 
2 ter huastory Of the Putters. v.56 cess sec e een Welsh and Breder 
13—Hlermaphroditism of a Croaker.............-.-.«0% Ss OREM: ONG Breder 
i——hood o: Certam Minnowss.0.-..-.--.-20.6+600%% Breder and Crawford 
Sip HPI SINGS One AN iy AO Ole DAV 0c) (3601s ood 's oreiit wis Gre @ cielo ae nis wiseliel © eels Breder 
Mp NVcavinon Ot tHe mVIGAMEI wisTEC foc Go wstecke oe 2.8 acaeies as Friedman 
if NG e WVNTGCHSHES: «of. cisj- cael od oiseca «6, ars ensyerene oa Mellen and Van Oosten 


Pa nerATiCeansomy (UTEG ME TOO... «,<2s\6 veins, «0 oiocieielei e's s/s e's eysteleie) tie a) Noble 


PAGE 


ioe) 


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LIST OF ILLUSTRATIONS 


TWO SERIES OF AMPHIBIANS 
Figures 1 to 6 inclusive 


PAGE 
Il, Isleyelleeitiin  I8MOSS cogdoocopgcbuaboabdondcqdnoondnEducUd coo ADCS 3 
2, ILEBEeyeGl INOS 5 oGog ooo bos oo bb OoOdOnoEgaOoUO GON Con OUODOGQOONr 3 
em VL AIMe NO ACE ete otal chet sic clone ts cr eprieleyeiietersts: che lels) sie. el's. 3/0016 eketneretehs eletenel a 
dh ANISIRAINEN AMOS ooocotononcosdpacnp0d0oo0cagIUUnO UDO DUsOOnG 25 
emer OO A CROCE OOM a oraucpahincoiet otal ies cians win el oe Cieiae) es ovele sida @ sora «i> 28 
Rim UNE: chew MOL e ciara. cx cccoe ore arene os care noni ols /o-allelaon lo shal eYeie( svozc8aiuts as. ose 28 
A TETRAPTERYX STAGE IN THE ANCESTRY OF BIRDS 
Figures 7 to 11 inclusive 
ie ketrapteryx stage im the Ancestry Of Birds) ....2 0. twscae es coms 39 
Gh IDiomisinte IPyReorl SWE) oocooopeoeosoodaee aoqco de male oe sooogaobgc 42 
See UAE Of Vy Ge VV ITCC MOVE <6: asstesate o) ois %aheve's w ale aluis\@ue @ Sees Sie) oowiere 42 
LOS “Detail of Pelvic: Wing of White-Winged Dove...............00.0000% 42 
11, The Evolution of Birds from the Tetrapteryx Stage through 
PRP CHACO PEON RII KG! LARC! iso. /6 Spisisie «ov eins s vier 02 ¢ lela vide as) ole 48 
NOTES ON THE BIRDS OF PARA, BRAZIL 
Figures 12 to 19 inclusive 
ieee neme MIMamon, “ree Or the Birds: <2 ..hec os vadiec Geen tsisae ssiae's 55 
Pepe U IME A UNS Le: SHONEREOUMG 5.5100 oie low aiecnls cole «diese wia.s sielblsis ersieicis 58 
ae ACECO Le AMES SOM hte: NEATH y 0c 55:2 «0 seis xin se ein plermieidlele ow Sain dew care 58 
POM OPANL SIGMA SIGMA tase ais ais is sins s'8 ete Se ES vob elev wikis e's VS eels ele vale’ 60 
HG INES! be) RS 1 cia 01 Sa ae, ee 60 
mame te El AV EPL INAUMMerce os sihase (alae cn cor eae mv. Gis hau oA Oale Moan belle Basitas 76 
PSpeminemeanikcable: Lnseeh HOLM s, ..s\-iscivesied-m 4 se oe s-s Myierale toe i dewis ets 113 
UCIT DLE EMISECE MOLUAS,. sp 546 «6 0.0,n 0100s wisi eile efdisie'w old's abetd clevopn oles 115 
THE GAFF-TOPSAIL 
Figures 20 to 31 inclusive 
Aram ext Op Setll PC AUEMESIN gh tesss enn pile. &:ajee. Hops eteion a aveshie) f'n aisve Wa) @ialelS sideievesalngsis 125 
Sean Ol CXO-CArrVvINe “TALE... 5s, cis: fyreie-s, chelera ss ale/ora le, whe wha a Slaler eles 133 
Beem RE SN NV TE TN LATING scat taici a ts Alot wes Aa apeerer Stonoie ala ig x7 eye arate lattes casa & b= 134 
Smee Winn Ave Se Gratl-LOPSAUL 5 5-3 sai dte =/eelarss sia) sioist slaPUh asesor ny dealeyate loca’ si a'e> os 6 139 
Zeeeeiocn:s Hisurev et the Gaff-topsalll:. 23.3. cm ec. < cae aleve sins > 0 « 139 
Bie Ceitenecastvoe mouth Of males... .0< ace icc mercimeaiye mileleeae we «5 6 147 
HiPeechichimecast OF MOUth.Of Male, . .. .cncnsgwe ainda ve aadltslere arslels's «6 147 


ILLUSTRATIONS 


PAGE 
Fig. 28. Skull of the Gaff-topsail.......... cece cece eee e eee e nee e teens 148 
Fig. 29. Ventral surface of skull............. see e cece eee e eee e eee e ences 150 
Fig. 30. Gaff-topsail that has just burst the shell...........+.ssee+eeeee 152 
Fig. 31. Larvae of Gaff-topsail 1... 0... c cece e eee e eee eee e ener e ees 152 

MAMMALS OF AUSTRALIA IN THE ZOOLOGICAL PARK 
Figures 32 to 68 inclusive 
Fig. 32. Typical forest in the Province of Victoria, Australia........... 167 
Fig. 33. Australia: Great Britain’s Southeastern Empire..............-.- 169 
Fig. 34. Young Gray Kangaroo, M. gigantus..............e.eeeseeeeees 176 
Fig. 35. Wallaroo or Kuro Kangaroo...........sssseeeseceneereeceees 176-177 
Fig. 36. Woodward Kangaroo ..............seeeee eee ceneecsersccease 176-177 
Fig. 37. Black-faced Kangaroo ..4.....-..0.2sececcec cee seeeeccceeees 176-177 
Big. 88. Parry Kangaroo. oi oni. cic eiveeinteteielae ims ice m  o/oelr xe ossta ne iinet 176-177 
Fig. 39. Giant Red Kangaro0..... 2.2. .2e.ee.e ener ses anvenecese ee ema 176-177 
Fig: 40. Kangaroo Island Kangaroo ...........5s.ecessesssecsecseeees 176-177 
Fig. 41. “Bennett. Tree’ Kangaroo. on «cis am - wis- 10 aieieteiaie\e)> ol onele ston oe 176-177 
Fig. 42. Bennett Tree Kangaroo .... 2.0.0.0 cee cet ce nnses ccnsss enene 176-177 
Fig. 43; Captive Tree Kangaroo |... 2 cis. wees ew cine cee ones ce wems wales 176-177 
Pig445* Rufous-necked| “Wallaby %2/.ccgie casts oles sims «else ctemiels ciel hig imei 176-177 
migse45;, (Brush-tavleds WPebascogale ce acta. citaomie, os mister «lorie canis eae 176-177 
Fig. 46: Bennett ree Kangaroo’... «cits. We pjae ote Rleeyo mis «o> sie ie lage 176-177 
ier, ye Vallllovoetoyy Jol PMIeeN tN (od cnicoog bg anaobb oon So UoonabieGeSooooas L76-1717 
Big. 48. Albino, Red-bellacds Wallaby, eciects cecleveiors ele eeaieo'=)siete ail ia 176-177 
Bigs 49, Ring-tailed “Walllaliys: sf cin = loticse crs 2) oop arias (= etalon Menara ete ee 176-177 
LO Os lanaisncienieGh Wellness: erga soenddcacesvdocnccsécanc00s50anccor 176-177 
Big. 51. , "Black? Swamp-W allaby: (3). s:oetsicteus Gv oor ate bie aise reas nee 176-177 
hisio2. Male sofathe Swarnp Wiel alloys creme teicioun nore et chelRete the iictehster aise 176-177 
Bie 53.) HApile Wall al yig coseyciass: 0-1 aye 1e crore cvoncisterefatastcree tot hehe miei cle ele takekt tae eee 177 
Biovo4.. Theyhomevot the wires Wallaloyersadte pa icleciteniete tenet eer eae ut) 
Big. 55.) Short—tailed Walla ys cis c:s) occ qopsier ot enor cnsetie breve ree) ote eiaeereretsene e 192 
Big. 756.7, FRAT IANS aArOO esis vaca al cieisis ote sine owiatese ctckeyeuae geneity mea Neen 192-193 
Hip.ov., Australian iGrayPhalanger ys... aaer tee cence tener 192-198 
Bigs 58. /Moalaor; mative, Beart... nevaeriitemr ee ites aceyahersnen tierra 192-193 
Big. bos Koala in’ its stree-top) haut ermine sii cle ior yer yer tee iets 192-193 
Bigs 60), i at-tailed OpossumpNouses seeemcacmocic cic arielemieaie etek eile 192-193 
Hig Gl, shlyane=Phalan per stcctist vette rer man etl el orci rae ieereiere 192-193 
hips 62Z.e Lalbbit-earedsiamnclicootiet iki eee ee ee ie 192-193 
Bigs 63. Wombat and ayiouns! 22.7.5... .che neers ene eee ete 192-198 
BieG4. Tasmanian: Wiolitges is ein. o% sates ¢ suctors ei diegscis tral te eR nae Ree 193 
Bip. 60.' A family ‘of “Tasmanian Devils .¢ +t... och eae eee eee ee 197 
Hie <665 (Winder surface lof ihe H chicas acis-tyes ascii eee eee 198 
Hig..6%,° Echidna in’-normal attitude... 03...2 06. .5s » Meomiee nee eee ers 198 
Bug. 68; The Platypus: or; Duck pill son set ls ss ee eee eae 201 


ILLUSTRATIONS 


REMARKABLE HABITS OF THE SAGE GROUSE 
Figures 69 to 76 inclusive 


PAGE 
Fig. 69. Sage Grouse displaying ............cceceeeecee eee e enc eeceecens 241 
Big Ol Piling. POU WILD AIP 2... nema rece etnies res eda ewes eee eees 246 
Hig. 71. Stifilesged rum after filling....... 2... cc ee wee cece e ere eeees 246 
Big. 72. Lifting pouch with the wings........5..s.seerevsceseseeenseeees 247 
Mie. jo. oidenview Of Nfl (of the pouch. 2. 2.5. ibe ete tess eels ee 5 aleles 24:7 
Bie ae extreme Of throw Of the pouch. ...5. 602 sms ees cele sedis sce ees 248 
Pe eere hl apy OOM GOL «POUCH: OMNCRESE ai./0- nia xtelaye aa ate siete eats Mle ole aime nin’ os 248 
Be ee rae ei TOMESE ON a6 oe) rater capa ay ai 2) sXe) the aisle lave aPaiwvai'e ope wi RRh a eysiapare isles wis 249 

ECLIPSE PLUMAGE IN DOMESTIC FOWL 
Figures 77 to 79 inclusive 

i oramer Cocke snowing Nackle IM -EClPSEs .. 26.5 sa. see 'aee oe os ves ae e's 253 
iiags “ifeky lateral “Asin Ke Mito} dle boo c a CeO OOO Emo cOdeL CAD Id 0 COMO mn mOnntar 253 
Peenoeebeathers on Games COCK. tiie cisici6 scchciee 06s le we wise cniineye seta desis 256 


A CONTRIBUTION TO THE LIFE HISTORY OF THE PUFFER 
Figures 80 to 96 inclusive 


OT, tel; IRETAVE) Gibns Cebus Cliiveiy Ines eoas sooo dbo deoedocnan oon cboaues OG 261 
Reeser eae WE cee TAL AULD OE MORE 5 aia. 6 ahePase an. 6p). exe, «0 ew ble Osi pe yare +, ohaaiaislel yas are 262 
Ee aN eer Van tb (ISLE Paes, oh wt see gh soe s's.04 8 0 pecie sce Gnd HarmEne wel we 6 266 
ie cp eanvaleishvOnerdayeakter MAGNUS 67... ci... sree see dela oe eels sis cies 266 
iio wore warty) fish fverdays, after hatching. . 0... 0... cs. c cea oe cree 266 
ices eleannalacsie SevenucayS cal ter WnatCMimer. cs. .\. sete cls ovals ols meie a) see 267 
bos MUD CLE CSO ES Soe INP Rb” Ae Ss eee i eee ere ae ae 267 
Bie Sie MOM Sis DALY MAMATE fs: a. 5oic 5yst0avsiaig de» sales) o siiein es 2 wa 6 efecto a 267 
nie s.e0 erat or head Of newly hatehed larval<... 2.0600. .006 cen eunese aes 268 
Sie Soe Metall of vent ot two day dld larva... 6... .i. ecawnee tenn sce ees 269 
eee EO ten OM MMMED ELL OLEAN Fy pies 50 <sne 5 cxe:s Foye ii slelinie, © wSisnee x oe 6 a tie was 270 
Biel mip lOrnbathoar Sixeday: O10 Larva. co osc ¢ siacaet ew slsje, cle sale oiarers ew o's 270 
His 92).. opines from ‘ventral surface of ten day larva......:2....+s00.5. 270 
eee Se ONDINE: OL ASP CCIM stacy areislaheia soere-es'e «aie o\6 spn’ als, 0's eleye wre Sielole. oie 0: 271 
PRO ae) OFS AlekVAGW Ae SANMEH STOECUMNEINeo cacl unl clei a r\eile ery eave. eee ope estates clin ie 271 
eS Demme LOZ OMe Un exe — CATE: v2 ccejenec statale. sere wise crajeier bested cress ci eieidlaveieslersiars 274 
Bie96.  Etorizontall index—ehart ....-...-...:.- ith con UOOG COMA OO OOO E 275 
HERMAPHRODITISM OF A CROAKER 
Figure 97 
Hie Ol. mOLORKer——AMALOMNCAL CHAT E. scie a x ct e'kja\o.6 sfc ein/s se lndisioa aie tediviek oo 44 3 282 
THE FOOD OF CERTAIN MINNOWS 
Figures 98 to 128 inclusive 
Bi eo Smee Vico mOmICOMECHIMG ‘SIC: cs). dee ia erat +) ay ote che eluven Teaver e RTO! bate res aid od 290 
ie ew POG RAN ST) SLPOAUL si. e's <5ie' Vs, 4/e ws Rely MALS RMI Ooldeds «ale oc 292 


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Fig. 129. 


ILLUSTRATIONS 


PAGE 

100. View looking Gown Stream eemn silos cee tee ensccenensins 292 
10 Contents, stomach) and initestuie ecOrn ays iretet ee) e) el) elale) = eielel ele eons 293 
102. ‘Contents, stomach and Untestimesmconnweers ei. © sre elelcle <= itera 293 
HOS=eSemotalus: (Grullaris’ Se erckine-cetae oie eretapebieletenere estore seiete/ssleuesehae 304 
OA GCUUCIS CUS OANA OUSRULUSereeetdcne eater treNens Retort iota) tet sierenetaorehelets 304 
LOB LE wers Cus GCANAOU UWS.) AMATI cay aialel sheleeheletetetetersiislelatelelersiale}s)s/sle)a)einielele 304 
106. Notropis procne.......-..-.- ele felekehels tat neieterereheveneheteh terete lore erakeKorons 304 
107. Notropis COTNULUS  . oe meen eens scenes meserecmecesscserareene 305 
108-< Notropis ‘cornusus, WON. 4 <y-oG - clctel ote telale eiehietoe teeta sete 305 
109; hinichthys QUpOn@sus Aver sstoierelolaielelsferatalatelatt ele mieisiape nici tale i winleat 305 
110. Hwoglosswm maxillangua 2... 5.6. e ieee wee ee tenses sneeseoe 305 
le ISCO (OUI Sada po omoma oro aOba DO bon OU doooD do ODDO BONGO 314 
WR EGO ONS COC OOSMOID sagcasos5b0uses ocopdn UGC adDSoenaoaoa0OaS 314 
IB), AOUFOPDIS (UROGMIO > so dacocnascone dosob dodo sooo MoDGUGOSoOdHUeCOURE 314 
TUS AONE CORMEUUE sondoooe cen ndonouSoooecauasacdzccgacoNsaons 315 
TIS; JRO LUIS. GROWENUS SoocooacocaobosdncbcsouoDOsD OB ODDBSSOE EC 315 
WG, JER OO MOSSE TCOUNGLI ao00coc00cKaedanne coansDeosanesbouEDOS 315 
S225) ee hyman Peale beetin serct-nocueueloiei herd teverseeieke stele rtetalctertne relearn 316 
TOQ=ZSiy Mnibesbimalle wall Saeetecstseeucven eueistcvetetetoker tenekers eVoleucl cvevatensy cccken ere renstete 317 

THE FISHES OF SANDY HOOK BAY 
Figure 129 
Map of Sandy Hook Bay........s.sesesescanesccereececereeee 330 


THE WEAVING OF THE RED-BILLED WEAVER IN CAPTIVITY 


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130. 
131. 


132. 
133. 
134. 
135. 


136. 
137. 
138. 
139, 
140. 
141. 
142. 


Figures 130 to 135 inclusive 


Weaving of Weaver Birds. ..... 0.00 .0.0000sesces es seunseecns 357 
Outline drawings of two nests and part of a play ground built 

TDL. CAP ULVALY oie iais) cance aim «)elersls\eiefepuiievs ell nits ey ehevelnenps sane eees 358 
Types of stitches used by Q. qweled......sseeesceeeeeccseeecees 361 
Shotenesy Ot \yVGener Isidtsls cooadanopapoooadconooagoesanxoanduoS: 363 


Highly specialized correlated action between beak and feet.... 365 
Graph showing color preferences of Q. qweled.....+.sseeeeeeees 370 


THE WHITEFISHES 


Figures 136 to 144 inclusive 


Whitefishes reared in the New York Aquarium................ 373 
Typical scale of Lake Huron Whitefish..............seeeeeees 388 
Scale of New York Aquarium Whitefish, December 1, 1920... .388-389 
Seale of New York Aquarium Whitefish, April 28, 1921...... 388-389 
Scale of New York Aquarium Whitefish, July 13, 1921........ 388-389 
Scale of New York Aquarium Whitefish, August 3, 1921...... 388-389 
Seale of New York Aquarium Whitefish, January 3, 1922...... 388-389 


Fig. 


Fig. 


Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 
Fig. 


143. 


144, 


145, 
146. 
147, 
148, 


149. 


150. 
151. 
152. 
153. 


ILLUSTRATIONS 


PAGE 

Chart of rate of growth of the scales of Open-gill and Double- 
GOONS Igoe cop 200 OUg boon Pak eb nO Snide DOCU oan Can pee eee 401 

Chart of growth curves of Lake Huron and New York Aquarium 
VWs chagGesdsod ocn.cock Gado ow ono OOD monn pric enaon 409 

THE ANDERSON TREE FROG 
Figures 145 to 153 inclusive .... .... 

tine (Chill erlang cpa oe bp ame Hone coc oOe a Oud One Ono OnE Oe core AL5 
Distribmtion| Ole alavanGEenso meiner opatec\e asl -1 ciel cletela(verere evercioele 6 416 
The Pine-Barrens at Lakehurst, New Jersey................. 2 420 

Flashlight photographs illustrating the diversity of the calling 
SIDUONS dosed odd ddsadousospeocoogeouoarodeoaenun Goose. 430-431 

Flashlight photographs of H. andersonii illustrating stages in the 
Fane RNASE I ROCES ae pala Oe hee ars) aro cae! aca i/ator ores ake Sen mores < siie.8 434-435 
Mherovipostionsoh Hija and ersonih).1. 1 sj) 41) ele) elais 2 ed > a oa 438 
Pigmentation of the egg of H. andersonii....... Seater norte 440 
Development of the adhesive organs. ... 4.220062 00) s's anaooar 443 
Madpoles Of Eyl) Wud ersOnib. <.o5. ag sieca + cise tarts @ stale vie oye alas’ elasals 446 


7 eo p : 
‘hey ie 


LOOLOGICA 
SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 1. 


REVIEW 
OF 


“TWO SERIES OF AMPHIBIANS 


By 
RICHARD DECKERT 


er UB LIS H E:D ey, THE. SOO Tt Bory 
foe ZOOLOGICAL PARK, NEW: ¥-0R-K 


a sf OCTOBER, 1915 


OFFICERS 
OF THE. | a 


Nem York Zunlogiral Suoriety 


President: . a 

HENRY FAIRFIELD OSBORN. : a 
Vice-President and Treasurer : a 
Secretary : Prrcy R. PYNE, < 
MADISON GRANT, 30 Pine Street. oe 


11 Wall Street. 


Executive Committee 
MADISON GRANT, Chairman. 


PERCY R. PYNE, FRANK K. STURGIS, 

*SAMUEL THORNE, LISPENARD STEWART, 

WILLIAM WHITE NILES, WATSON B. DICKERMAN, 

Wo. PIERSON HAMILTON, HENRY FAIRFIELD OSBORN, 
ex-officio. 


Auditing Committer 
WILLIAM WHITE NILES, Chairman. - 
H. CASIMIR DE RHAM, LISPENARD STEWART. 


General Officers : . 
Director of the Zoological Park: WILLIAM T. HoRNADAY. 
Director of the Aquarium: CHARLES H. TOWNSEND. 
Prosector: DR. GEORGE S. HUNTINGTON. 
Architect: C. GRANT LA FARGE. 
Consulting Engineer: H. DE B. PARSONS. 
Assistant Secretary: H. J. SHORTER, 


Assistant to the Treasurer: R. L. CERERO. 


*Deceased. . : 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
haw YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 1. 


REVIEW 
OF 


TWO SERIES OF AMPHIBIANS 


BY 
RICHARD DECKERT 


PU biel S HE D SY. Rees Mo SS OsGel hy & 
Stee OoOrkOGiCAL PARK, NEW YORK 


OCTOBER, 1915 


FIG. 1. HARLEQUIN FROGS, DENDROBATES 


Blue-legged Frog, Dendrobates typographus, Keferst. 
Ornate Frog, D. tinctorius, Schneid. Searlet Frog, D. t. ignitus, Cope 


reset 


Richard Deckert, pinx 
FIG. 2. LARGE-EYED FROG, RANA CHRYSOPRASINA, COPE 


GPoanlnanrvra Val TI Nn 1 BRwnmtremoro 


REVIEW 
OF 


TWO SERIES OF AMPHIBIANS* 


By RICHARD DECKERT. 


(Color plates from drawings by the author.) 


INTRODUCTION. 


The species described in these articles belong to the Class 
Amphibia and the living members can be roughly defined as 
Vertebrates which undergo an external metamorphosis, hatching 
from eggs (spawn) as tadpoles or larvae, and gradually assum- 
ing the adult form. These larvae have gills at some stage of 
their development and several species complete the metamorpho- 
sis inside the egg and emerge as gillless and tailless frogs. 


Most amphibians lose these larval gills, except some of the 
salamanders, like Necturus and Proteus, after attaining the adult 
form. Several of the American land salamanders of the genus 
Amblystoma retain the gills throughout life, if the conditions for 
developing into the land form are unfavorable, and have been 
known to breed in this semi-larval state. 


The members of the Amphibia are divided into three Orders, 
as follows: 


The Apoda or limbless amphibians, have a vertebral column 
with rudimentary ribs and amphicoelous vertebrae sometimes to 
the number of 300. Each vertebra is cupped before and behind 
and articulated with the adjoining member by means of a car- 
tilaginous plate. Only one lung is present. The shape of these 
animals is cylindrical, the head is not distinct from the body, 
there is no tail and the anus is placed at the posterior end of 
the body. The body shows no internal rudiments of limbs, is 
naked and ringed by furrows running around it similar to the 
segments of an earthworm. A few species have calcareous de- 


*The specimens described in these reviews have been preserved and do- 
nated to the American Museum of Natural History. Marked with all possible 
data, they are now available for study. 


4 Zoologica: N. Y. Zoological Society. CEE sal: 


posits in the form of bony scales under the skin. The eye is 
small, the mouth usually wide, and the teeth large. These ani- 
mals lead a subterranean life, burrowing in the soil of tropical 
and subtropical countries. The larvae live in the water until 
the absorption of the gills. The eggs are round or oval, and are 
joined together by a gelatinous string. This Order embraces 
fifty species. 


The next order is the Caudata or tailed amphibians. These 
creatures have a spinal column formed of from thirty-seven to 
ninety-eight vertebrae, which are amphicoelous or opisthocoe- 
lous, that is cupped in front and behind or only behind. The 
skin is naked, the head broad, flat and distinct from the body. 
All Caudata have limbs, although some species only rudimentary 
ones (Amphiwma). Sternal apparatus as well as pelvis always 
present, although the latter is sometimes rudimentary (Siren). 
All members of this order have a tail throughout life. Lungs 
are usually present, although the Plethodontinae have no lungs; 
breathing solely through their slimy skin. All salamanders or 
tailed amphibians hatch from eggs and undergo a metamorpho- 
sis from larval to adult form. All of the known species, which 
number about one hundred and fifty, breathe through gills at 
some period of their existence. 


The last order, Salientia, or tailless amphibians, is the one 
to which all species treated in these articles belong. They are 
characterized by their form and the presence of four well de- 
veloped limbs. With all the tail is absent in the adult form. 


The skeleton is simple, with comparatively a large and broad 
skull and a short spinal column, consisting of from five to nine 
vertebrae and which terminates posteriorly in an elongate pelvis. 
This peculiar pelvic arrangement is necessary for the attach- 
ment of certain muscles that are used in leaping or acting as 
springs in giving impetus to the enormous leaps, which are pos- 
sible by most members of this order. Short ribs are present only 
in one family, (Discoglossidae). The limbs are always four in 
number with four digits on the hand and five on the foot, but 
in a few species some of the digits are rudimentary or absent 
(Stumpffia). Some tree-toads have a rudiment of a fifth finger 
on the hand. The skin is either smooth or dry, and more or less 


1915] Deckert: Two Series of Amphibians. 5) 


granular or warty, but always naked. In a few species (Man- 
tophryne, Ceratophrys) there are calcareous deposits in the skin 
in the shape of bony plates or granules. The eye is usually 
large, bright and so very mobile that it can be lowered into the 
skull until even with the top of the head. 


The life habits of frogs and toads present considerable vari- 
ation. Some species are terrestrial, some arboreal, some aquatic 
and others subterranean. The food consists of living insects 
chiefly, but some of the larger species are cannibalistic, and a 
few of the largest will eat small mammals, birds and snakes. 


The order consists of nine families, divided into eleven sub- 
families numbering about 1,200 species. 


In the descriptions to follow it should be particularly noted 
that the color patterns are described from living subjects 
throughout. The greater number of the past descriptions of the 
rarer amphibians are from preserved specimens, and some con- 
fusion has resulted owing to the rapid fading of these animals, 
with a consequent marked change, not only in hue but in pattern. 
The greater number of the colored figures are for the first time 
sketched from life. 


FROGS AND TOADS FROM COSTA RICA. 


On May 20, 1914, Mr. Lee S. Crandall, Assistant Curator of 
Birds at the Zoological Park, and Donald Carter, a student at 
the Park, returned from a six weeks’ collecting trip in Costa 
Rica. The trip resulted in the capture of many interesting birds 
and other animals, among the latter being ten species of frogs, 
toads and tree toads; most of them never before exhibited in 
this country. Following is a list of the species: 


Family BUFONIDAE—Toads. 
Marine Toad, Bufo marinus, Seba. 


Family HyLIpAE—Tree Toads. 
Baudin’s Tree Toad, Smilisca baudini, Dum. & Bibr. 


6 Zoologica: N. Y. Zoological Society. BeCea 


Family CYSTIGNATHIDAE—Arch-Jawed Toads. 
Underwood's Toad, Hylodes underwoodi, Blgr. 
Barred Piping Toad, Hylodes polyptychus, Cope. 
Brown Piping Toad, Hylodes rhodopis, Cope. 


Family RANIDAE—Frogs. 


Subfamily RANINAE—True Frogs. 
Large-eyed Frog, Rana chrysoprasina, Cope. 
Godman’s Frog, Rana godmani, Gthr. 


Subfamily DENDROBATINAE—Harlequin Frogs. 
Blue-legged Frog, Dendrobates typographus, Keferst. 
Searlet Frog, Dendrobates typographus ignitus, Cope. 
Ornate Frog, Dendrobates tinctorius, Schneider. 


Family BUFONIDAE—Toads. 


The toads of this family have no teeth in either jaw, the 
shoulder girdle is arciferous or dilatable, and the vertebrae are 
procoelous, or cupped in front and without ribs. The tips of the 
fingers and toes are either obtuse, (genera Notaden, Pseudo- 
phryne, Nectes, Bufo), pointed, (Myobatrachus, Rhinophrynus, 
Cophophryne) or triangular, and carrying medium-sized or large 
adhesive disks or pads, (Hngystomops and Nectophryne). The 
Bufonidae are distributed over all parts of the globe except, of 
course, the Arctic and Antarctic regions, which have no amphibi- 
ans. Central America and northern South America have the 
most genera, as well as the greatest number of species. The 
species number about one hundred and fifty, grouped in nine 
genera. Of these genera, Notaden (one species) and Myoba- 
trachus (one species) are Australian, Pseudophryne (four spe- 
cies) is Australian and African, Nectophryne (seven species) is 
African and Indian, Nectes (four species) Javan and Sumatran, 
Cophophryne (one species) Indian, Engystomops (three species) 
Central—and South American, Rhinophrynus (one species) is 
Mexican and Bufo (about one hundred and thirty species), cos- 
mopolitan with the exception of Madagascar, New Guinea and 
the Australian region. 


1915] Deckert: Two Series of Amphibians. i 


The habits of this family are terrestrial except the genus 
Nectes, which is aquatic and has enormous webs on the hind 
feet. Nectophryne is more or less arboreal as indicated by the 
enormously dilated and padded fingers and toes. 


Most of the Bufonidae are excellent burrowers, hiding by 
day and coming forth at dusk to hunt insect prey. Few species, 
however, are strictly nocturnal, some of the genus Bufo (Bufo 
fowleri B. calamita B. quercicus) having been observed hunting 
in the brightest sunshine. The genera Rhinophrynus of Mexico 
and Notaden and Myobatrachus of Australia, are almost exclu- 
sively termite and ant-eaters, herein approaching most of the 
species of the family Engystomatidae (the narrow-mouth frogs) 
whom they also resemble in external appearance, small head, 
enormously fat body, short arms and legs and comparatively 
smooth skin. 


In their movements the Bufonidae are not as agile as the 
true frogs (Ranidae), usually proceeding by short hops, walk- 
ing, crawling or in rare cases running. They are excellent climb- 
ers, taking advantage of every unevenness to obtain a hold, and 
maintaining their balance in most trying situations. With this 
they combine great persistence, especially when trying to escape 
from some pit, well or terrarium. They are but indifferent swim- 
mers and divers with the exception of the genus Nectes, and will 
only enter the water to soak their skin, and during the breeding 
season for the purpose of depositing their spawn. 


All species that have come under observation are great feed- 
_ers, eating untold numbers of insects, most of which are injuri- 
ous to plant life. In this they take the place of the insectivorous 
birds on the ground at night and therefore merit our protection, 
which unfortunately has been withheld to a great extent until 
lately. This was probably due to ignorance of their habits, and 
also to the often unprepossessing appearance of these harmless 
creatures. The flesh of the larger species is said to be quite as 
edible as that of some of the true frogs, but is eaten only by a 
few aboriginal tribes of tropical countries. 


In appearance most of the Bufonidae are squat, fat and 
warty. There are, however, some smooth skinned, long legged 
tropical species that remind one of a true frog. 


8 Zoologica: N. Y. Zoological Society. [nies Se 


Marine Toad, Bufo marinus, Seba. (Bufo agua, Latr., 
Bufo ictericus, Spix., Bufo horridus, Daud.) (Boulen- 
ger, Cat.-Batr- Sal, Pa Sib). 


Color: Brownish or greenish-olive, yellowish or reddish- 
brown or plain dark brown above with or without large, black, 
insuliform spots, these usually edged with pale yellow. Some- 
times a light median line from behind the head to above the 
vent. There may be a few scattered whitish spots on the back 
and sides. Arms and legs sometimes distinctly banded with 
dark brown. The cranial crests, parotoids and larger warts are 
usually light, reddish brown. Below this toad is dirty white 
or yellow with or without brown spots. 


Structure: The head is broad and crowned with very 
prominent bony crests diverging from above the nostrils, edg- 
ing the canthus rostralis, curving around the orbit above, and 
sending out and down a branch before and behind the eye, 
several more or less distinct branches towards the median line 
above, and one connecting with the enormous parotoid glands. 
These glands curve down over the arms, and in a toad six and 
a half inches from snout to vent, attain a length of one and 
three-quarter inches and a width of one and three-eighth inches. 
They are studded with large pores. There is a distinct bony 
ridge along the upper jaw on the edge of the mouth. The tym- 
panum is about one-half the diameter of the eye, and very dis- 
tinct. The body is covered with large round warts. The skin 
is loose and much wrinkled and creased. Arms and legs are 
long and powerful, hands and feet large. The ends of fingers 
and toes are tipped with dark brown. The toes are moderately 
webbed. Metatarsal tubercle moderately developed. 


Size: Adults range from five inches to eight and one-half 
inches in length from snout to vent. 


Range: Southern Mexico through Central and South Amer- 
ica to Southern Argentina. Many of the West Indian Islands. 


Thirty-one specimens of the giant marine toad, of all sizes 
from two to seven inches in length were collected. These large 
toads are very common throughout their range and no doubt 
are beneficial in helping to keep the insect life of those regions 


MARINE TOAD, BUFO MARINUS, SEBA 


FIG. 3. 


Face page 9. 


Zoologica Vol. II, No. 1. 


1915] Deckert: Two Series of Amphibians. 9 


within reasonable bounds. This is the largest species of the 
true toads, only exceeded in size by Rana goliath and Rana ad- 
spersa, both African frogs, and possibly Ceratophrys dorsata 
of Brazil. The male can be distinguished from the female by 
the horny pads on the inner fingers, and by a blackish area on 
the throat indicating the presence of a large gular pouch. This 
dark area may be sprinkled with lemon-yellow, especially in 
young males. The writer has often observed the males sitting 
with this pouch partly distended in the pan of water provided 
for their soaking bath in the spacious cage in the lobby of the 
Reptile House, especially after they had fed well, but so far has 
not heard their call. The warts on the upper surface of the 
males are covered with small horny spines, making these toads 
exceedingly rough to the touch, in contrast to the females, whose 
warts are round and smooth. When picked up, the male of this 
species gives voice to a series of harsh squawks similar to those 
produced by some toy animals. The female is silent, and when 
handled will shake the whole body as if in a rage, then puff it 
up enormously but deflating it after a little while with a loud 
hiss. From the enormous parotoid glands, Dr. J. J. Abel, of 
Johns-Hopkins University in Baltimore, has recently extracted 
two distinct and powerful poisons, called respectively Epine- 
phryn and Bufagin. The latter poison, Dr. Abel has found by 
experimenting, has many times the strength of Digitalin, the 
“‘fox-glove”’ poison, and like this, it affects the heart action. This 
species has proven one of the hardiest in captivity, seemingly 
being immune to the festering and bleeding ulcers with which 
captive toads so often are infested, and to which the majority 
of them succumb. In the Reptile House these toads are fed on 
all kinds of insects, and also large earth worms and cockroaches. 
A few of the largest toads sometimes get an extra tid-bit, suen 
as very young mice or rats. 


Although the giant among toads and able to swallow the 
largest of insects and worms as well as small mammals, this 
species is not cannibalistic. The writer had repeatedly tried to 
induce them to eat very young common toads and frogs, which 
they will snap up but reject immediately. In spite of their enor- 
mous size and bulky appearance, the “marine toads” are re- 
markably agile and quick on their feet. When insects are thrown 


10 Zoologica: N. Y. Zoological Society. [List 


into their cage they instantly become alert and with a series 
of rapid hops, almost cat-like in their stealth, come forth from 
the darkest corner of their cage where they usually spend the 
day piled on top of one another and sleeping. Some specimens 
eventually become so tame that they will take insects and worms 
from one’s hand. The tongue of this toad is very flexible, seem- 
ingly more so than in other species. The writer has seen some 
of the larger specimens snap up a grasshopper or mealworm 
fully four inches from the toad’s head. The species breeds 
during the rainy season, the eggs being quite small and laid 
in two strings very similar to those of our own common toads, 
in puddles, ditches, ponds and canals. The matamorphosis is 
comparatively rapid, and the baby toads are tiny, measuring 
scarcely three-eighths of an inch from snout to vent. The call 
of the male is said to resemble the barking of a dog. 


Family HyLipAE—Tree Toads. 


This family is divided into two subfamilies, the Amphigne- 
thodontinae and the Hylinae. It is the latter subfamily only, 
that we have to deal with in this article. The Hylinae or Tree 
Toads are characterized by the dilatable shoulder girdle, the 
presence of teeth in the upper jaw, vertebrae cupped in front, 
absence of ribs, and dilated transverse processes of the sacral 
vertebra. The end-phalanges of the fingers and toes are claw- 
shaped and support more or less prominent, adhesive disks. 
These disks secrete a sticky fluid which, aided by the moist and 
granular surface of the belly, enable the tree-toads to climb 
trees, vines and even the glass sides of a terrarium with perfect 
ease. The skin of the Hylinac is always moist and slimy, thus 
enabling them by means of increased evaporation to withstand 
greater heat than other amphibians. Tree-toads often can be 
seen sitting for hours in the most glaring sunlight. 


Their skin may be quite smooth or covered with warts of 
various sizes. In some species of the genera Hyla, Nototrema, 
Nictymantis and all those of Pternohyla, Corythomantis, Trip- 
rion and Tetraprion, the skin of the head adheres to the skull. 
The Hylinae includes some of the brightest colored and most 


1915] Deckert: Two Series of Amphibians. 1} 


attractive of all the frogs and toads. All of the species have 
great powers of color-change. Some of them are really mar- 
vellous in this respect. This applies especially to the genus 
Hyla. Most of the Hylinae live on the trees, vines, shrubs and 
other plants, but a few, however, have such tiny adhesive disks 
that they are of little use, compelling the creatures to live on 
the ground. All species are insectivorous, although many of 
the larger ones incline to cannibalism. The family Hylidae con- 
tains sixteen genera; one of these, Amphignathodon, belongs to 
a separate subfamily. With the latter there are teeth in both 
jaws. It includes but one species, A. guentheri, of Ecuador, and 
is exceedingly rare. 


The other fifteen genera are included in the subfamily Hy- 
linae. There are about two hundred and forty known species 
of which the genus Hyla alone has about one hundred and 
eighty. The pupil of the eye of the toads of this genus is hori- 
zontal elliptic. The toes are webbed. Adhesive disks distinct, 
sometimes very large. The distribution of the Hylidae is as 
follows: The genus Hyla is almost cosmopolitan with over thirty 
species in Australia and Australasia, about one hundred and 
thirty-five species in Mexico, Central and South America, seven 
species in the West Indies, ten species in North America, and 
one species in Europe and Asia. This latter, Hyla arborea, has 
two subspecies in China and Japan. Of the other genera, Acris, 
(one species, two subspecies) and Chorophilus (five species) are 
North American, Smilisca (one species) ranges from Texas 
through Mexico into Northern South America, and the genera 
Nototrema (eight species), Hylella (seven species), Thoropa 
(one species), Phyllomedusa (fourteen species), Agalychnis 
(four species), Nictymantis (two species), Triprion (two spe- 
cies), Tetraprion (one species), Diaglena (one species), Cory- 
thomantis (one species), and Pternohyla (one species) are Cen- 
tral and South American. 


This distribution seems to show that the original home of 
the Hylidae was South America. One species of the otherwise 
North American genus Chorophilus occurs in the mountains of 
Peru (Ch. cuzcanus). Quite a number of species of this inter- 
esting family are remarkable in their breeding habits. They 


12 Zoologica: N. Y. Zoological Society. pages 


do not lay their eggs in ponds, ditches, lakes or swamps as is 
the habit of most frogs and toads, but use the axillae of large 
broad-leaved parasitic or other plants for this purpose. One 
species, Hyla resinifictriz, of Brazil, uses knot holes which it 
lines with the resinous sap of an aromatic tree (Protium hep- 
taphyllum) and which soon become filled with rain water. The 
gigantic Hyla faber, also of Brazil, constructs nests or nurseries 
of mud, forming circular inclosures about twelve inches in diam- 
eter in shallow parts of ponds. The eggs are deposited and the 
tadpoles reared in these nurseries. Other species, (Hyla goeldt) 
and species of the genus Nototrema carry the spawn in a pouch 
on the back until the tadpoles hatch. This pouch is formed by 
the introverted skin of the back and is possessed by the female 
only. Species of the genus Hyla have the strongest voices cf 
any of the Salientia. The call may be a shrill pipe, whistle, a 
very loud rattle, croak or bark, or a bell-like note (Hyla gratiosa, 
of Florida, H. faber, of Brazil) that can be heard in some cases 
for more than a mile. Each species has its distinctive call and 
the din produced by these and other toads and frogs in tropical 
forests during the breeding seasons is said to be ear-splitting. 


Baudin’s Tree Toad, Smilisca baudini, Hyla baudint 
Dum. & Bibr. (Boulenger Cat. Batr. Sal. P. 371). 


Color: The body color is green of varying shades from 
dark olive through bright pea-green to pale, golden green. A 
dark band from the eye to the shoulder, covering the tympanum 
and sometimes extending to the groin, a light spot beneath the 
eye, and a dark band curving over the upper arm at its inser- 
tion. The groin of both sexes is bright yellow, also the throat 
of the male. Undersides white. These marks are always pres- 
ent. Markings which sometimes disappear with the assumption 
of pale shades are, a broad band on the middle of the back with 
two branches extending on the eyelids, two or three cross bars 
on arms and legs and a few smaller dark spots on the back. 


Structure: The head is broad and flat, canthus rostralis 
acute, eyes large, reddish golden in color and very far’ apart. 
The skin is smooth or very finely granular. The legs long, toes 


1915] Deckert: Two Series of Amphibians. 13 


two-thirds webbed, fingers slightly webbed, with adhesive disks 
smaller than the tympanum. The latter two-thirds the diameter 
of the eye. The vomerine teeth are situated slightly behind the 
internal nares, and arranged in a straight series which is inter- 
rupted in the middle, thus separating the genus Smilisca from 
Hyla. The male has two large gular pouches, one on each side 
of the throat. 


Size: This species is large for a tree toad, reaching a 
length of three and a half inches from snout to vent. The male 
is smaller than the female; two and a half inches being the 
average size. 


Range: From southwestern Texas through Mexico to Pan- 
ama. Four specimens of this fine tree toad were captured near 
Guapiles, Costa Rica, by Mr. Crandall and his assistant. They 
were heard calling at night from a piece of waste ground, and 
their cry was traced to several old tin cans partially filled with 
water in which the tree toads were sitting. In their cage in 
the Reptile House they seem to prefer dark corners, where they 
sleep during the day, coming forth at night and climbing all 
over the glass sides of their vivarium. They have not been seen 
feeding since their arrival, although tempted with all kinds of 
small live insects. 


Family CysTIGNATHIDAE—Arch-Jawed Toads. 


This is a most difficult family to define as it approaches the 
Bufonidae, Hylidae, Pelobatidae, and Ranidae in internal as 
well as in external identification characteristics. Its distribu- 
tion is South American and Australian almost exclusively. One 
species (Liopelma hochstetteri, the only amphibian there), being 
found in New Zealand, where it is rare, and four species enter- 
ing North America. 


The family has the following internal structural character- 
istics: A dilatable shoulder girdle, teeth in the upper jaw only, 
(subfamily Cystignathinae), in both jaws (Hemiphractinae), 
or no teeth at all (Dendrophryniscinae). The terminal phalanges 
or finger and toe ends are never claw-shaped, although some- 


14 Zoologica: N. Y. Zoological Society. Lb Gest 


times carrying adhesive disks as in the Hylidae. The family is 
composed of three subfamilies, the Hemiphractinae, helmet- 
heads, so called from the shape of their enormous heads which 
carry large bony protuberances reminding one of the casques 
or helmets of ancient knights, the Cystignathinae, arch-jawed 
toads, including the greatest number of species, and so called 
from the enlargement of the lower jaw of some species, and 
finally the Dendrophryniscinae, toads without teeth. 


The genera are so numerous and so poorly defined that 
almost every author gives a different number of the same (Cope, 
thirty-seven genera, Gadow thirty-two genera, Werner thirty 
genera, etc.). There are about two hundred and fifty species, of 
which Australia has about thirty and the remainder are from 
South America. 


Their habits are very diversified, some being burrowers, 
some strictly aquatic and a great number of species are arboreal, 
living like tree toads (Hylidae) and resembling them in appear- 
ance. 


Most of the Australian species have a vertical pupil, indi- 
cating nocturnal habits. 


In size the Cystignathidae range from the enormous Cev'a- 
tophrys dorsata, measuring nine inches from snout to vent, and 
Leptodactylus pentadactyles, six to eight inches from snout to 
vent, to the small Pseudis minuta, which measures only three- 
quarters of an inch from snout to vent. Both extremes in size 
are found in tropical America. In South America, Central Amer- 
ica, Mexico and the West Indies the tree-living species pre- 
dominate, whereas in Australia the members of this family 
without exception are burrowers. The largest genus is Hylodes, 
having more than eighty species, Leptodactylus has about thirty- 
five, Paludicola has thirty-two and Ceratophrys has seventeen 
species. The other genera have from one to twelve species each. 


The breeding habits of this family vary considerably. Some 
species like Hylodes martinicensis carry the tadpoles on their 
backs while others like some of the Hylidae lay the eggs in foamy 
masses in the axillae of large-leaved plants; but the breeding 
operations of the majority of the species are unknown The 


1915] Deckert: Two Series of Amphibians. 15 


tadpoles of two species grow to an enormous size; that of Calyp- 
tocephalus gayi, a giant water frog of Chile, reaches six inches, 
the adult frog is six to seven inches from snout to vent, while 
the tadpole of Pseudis paradoxa, also a water frog of the Guy- 
anas, is larger still, one specimen being ten and one-third inches 
long, three and one-third inches of this total is taken up by the 
body and head, and the tail, which is thick and muscular, meas- 
ures almost four inches in width by six and two-third inches 
in length. The size of the larva is all the more remarkable since 
the adult frog measures only two to two and a half inches from 
snout to vent. 


Underwood’s Toad, Hylodes underwoodi, Boulenger 
(Guenther, Biologica Centr. Am.). 


Color: The general color is sepia-brown with a W-shaped 
mark on the shoulders. This mark may be much darker than 
the ground color or very pale, yellowish brown. The rest of the 
back is marbled with dark brown, and the arms and legs of some 
specimens are banded with dark brown.  Undersides bluish 
white, specked with brown. 


Structure: The head is long and the snout pointed. The 
eyes are large with the interorbital space smaller than the eye- 
lid and the skin is rough with large elongated warts, giving this 
frog some resemblance to our cricket frog. The fingers and toes 
are free, subarticular tubercles prominent, and the adhesive disks 
minute, scarcely produced. 


Size: From snout to vent the length is one to one and one- 
quarter inches. The specimens collected were immature and 
from one-quarter inch to three-quarter inches in length from 
snout to vent. 


Range: Known only from Costa Rica. 
Barred Piping Toad, Hylodes polyptychus, Cope (Guen- 
ther, Biolog. Centr. Am.). 


Color: The general color is a dark, brownish olive with a 
white band between the eyes. ‘The arms and legs are indistinctly 


16 Zoologica: N. Y. Zoological Society. fakes 2 


barred, with a rich pink spot on the groin which is hidden when 
the frog is at rest. 


Structure: The skin is finely granulated. The head is 
broad with the interorbital space equal to or larger than the 
diameter of the eye. Tympanum small and distinct. Subartic- 
ular tubercles distinct. Disks on fingers and toes small, but 
distinct. Toes not webbed. 


Size: The single specimen examined was one and a quarter 
inches from snout to vent. 


Range: Costa Rica. 
Brown Piping Toad, Hylodes rhodopis, Cope, (Cope 
Proc. Acad. Phil. 1866). 


Color: Brown prevails above, while below it is bluish white 
with a few scattered brown dots. There is a pale area in front 
of the eyes on top of the head and the canthus rostralis is mar- 
gined with dark brown. 


Structure: The head is long and pointed and the canthus 
rostralis acute. The nostrils are close to the top of the snout, 
the interorbital space is wider than the eye and the tympanum 
distinct and smaller than the eye. The fingers and toes are 
equipped with small disks. Subarticular tubercles distinct. The 
back has several longitudinal rows of warts arranged in the 
shape of a lyre. 


Size: One specimen one and a quarter inches from snout 
to vent was examined. 


Range: Mexico to Costa Rica. 


All these small frogs were shy and delicate and did not 
live long. 


Family RANIDAE—Frogs. 


The Ranidae belong to the second group of tailless amphib- 
ians, the Firmisternia, so-called because the halves of the shoul- 
der girdle are united below, forming a firm median bar or 
metasternum, instead of overlapping as in the Arcifera, to which 


1915] Deckert: Two Series of Amphibians. Ly 


all previously described frogs and toads belong. The vertebrae 
are cupped in front. 


The Ranidae are divided into the following subfamilies ac- 
cording to the arrangements or absence of teeth: 


Subfamily Ceratobatrachinae, having teeth in both jaws 
and consisting of only one genus and species, Ceratobatrachus 
guentheri, of the Solomon Islands. This is a large, huge-headed 
land frog with horn-like appendages on the eyelids, snout, sides 
of body and limbs. 


Subfamily Raninae or true frogs with teeth in the upper 
jaw only. This is the most numerous branch of the family, 
comprising about forty genera with some three hundred and 
seventy species. These are so diverse in identification charac- 
teristics and habits that it would be impossible to describe all 
the genera in this paper, and but a few examples will be men- 
tioned here. 


Genus Polypedatus (Rhacophorus), frogs resembling tree 
toads in having the tips of the fingers and toes with adhesive 
disks, but the end phalanges not claw-shaped as in the Hylidae. 
Some species of this large genus have enormous webs between 
the fingers as well as the toes. They have been called flying 
frogs but do not actually fly, only jumping from great heights 
occasionally and using the large expanse of web as a parachute. 
Fifty-four species are known from southern and eastern Asia, 
and sixteen from Madagascar. Many species of this genus lay 
their eggs between leaves glued together by the female to form 
a sort of funnel which they suspend over a ditch, pond or brook, 
so that when the tadpoles have hatched they will drop into the 
water below their nest. This queer mode of depositing eggs is 
also practiced by the African genus Chiromantis, which resem- 
bles Polypedates, except that it has no web between the fingers 
and that the two inner fingers are opposed to the outer ones, 
enabling these frogs to grasp twigs and stems in climbing. Their 
movements are slow and mechanical, like those of the African 
and Madagascan cameleons. ‘Cameleon frogs” would therefore 
be an appropriate popular name for these queer creatures. 


Hylambates with about twenty species, all African, is also 
a tree frog in the true sense. Our own so-called tree frogs are 


18 Zoologica: N. Y. Zoological Society. felt est 


really tree toads, being grouped with the toads in the super- 
family Arcifera. 


The genus Hylambates has some highly colored frogs with 
odd and picturesque color patterns. The female of one species 
(H. brevirostris) has been found by Boulenger to have a sin- 
gular habit of nursing, carrying the eggs about in her mouth. 
The African and Madagascan genus Rappia is also very numer- 
ous, having about thirty known species. They are mostly small 
tree-living frogs with rather short, stout limbs, all beautifully 
colored and have great powers of color-change. 


The female of one species from Madagascar has the singu- 
lar habit of winding the eggs, which resemble a string of beads, 
around her forelegs. 


Trichobatrachus, only one species of which is known so far, 
is peculiar in the possession of hair-like papillae forming a thick 
fringe on each side of the flank, also on the upper side of the 
thighs. This frog inhabits Central Africa. 


Phyllobates, having five species, all small frogs, is a South 
American genus. The tadpoles of Ph. trinitatis of Trinidad, 
British West Indies, adhere to the back of the male by means 
of their suckers, and are thus carried from evaporating pools 
to more permanent ones. A7throleptis comprises twenty spe- 
cies, mostly African. One species, A. seychellensis, of the Sey- 
chelle Islands, was found on some tree ferns carrying its tad- 
poles in the same manner as the genus described before, with 
the exception that the larvae adhered to the back of the adult 
by means of a sticky secretion. 


Rana, the type genus of the whole family, is also the largest, 
having about one hundred and fifty species, of which fifteen in- 
habit the United States. 


The Indian region including most of the islands of the In- 
dian and Pacific Oceans, has the greater number of species, 
Africa has but a slightly smaller number. 


The structural description of the genus is as follows: The 
pupil of the eye is horizontal and the tongue deeply notched and 
free behind. Teeth on the upper jaw and on the vomers, (small 


1915] Deckert: Two Series of Amphibians. ee 


protuberances in the upper jaw), between or slightly behind the 
internal nares or nostrils. The fingers are free and the toes 
are more or less webbed. The fourth and fifth metatarsal bones 
of the central part of the foot diverge, but are united by the 
web. The terminal phalanges may be simple and pointed or 
T-shaped, sometimes carrying disks. The external ear plate 
(tympanum) is usually distinct. The males of most species have 
vocal sacs, which may be internal (Rana catesbiana, R. sylvatica, 
R. temporaria), or external, protruding through slits under the 
angle of the lower jaw or over the arm insertion when they are 
distended in calling (R. esculenta, R. aesopus, R. tigerina). 


Nuptial excrescences in the shape of horny or spiny pads, 
spikes or granules may be found on the forelimbs and hands of 
the males of many species, reaching their greatest development 
in Rana liebigii of the Himalayan region, India. 


The males of this genus are further distinguished by their 
heavy forelimbs or arms which in the aforementioned species 
are enormously developed. Gadow says in his ‘“‘Amphibia and 
Reptiles” (Cambr. Nat. Hist.): “All species of Rana spawn in 
the water, except those of the Solomon Islands, where the only 
permanent bodies of water are roaring mountain torrents un- 
suitable for the metamorphosis of amphibian larvae.” 


One species from this group of Islands, Rana opisthodon, 
lays its eggs in moist crevices in rocks near the water. The 
larvae undergo the whole metamorphosis from tadpole to frog 
inside the eggs and emerge as perfect frogs, absolutely tailless. 
The tip of the snout of the young frog is armed with a short, 
horny protuberance which is used to perforate the egg and is 
absorbed soon after the animal has emerged. The largest spe- 
cies of all frog-like amphibians is Rana goliath of the Cameroons, 
attaining a length of twelve inches from snout to vent. . Next 
in size are Rana adspersa (nine and one-quarter inches), of 
South and Central Africa, Rana macrodon (nine inches) of In- 
dia and Malaysia, Rana catesbiana (six to eight inches) of 
North America, Rana tigerina (six to seven inches) of India 
and Malaysia, and Rana guppyi (six to seven inches) of the Sol- 
cmon Islands. All these large species are cannibalistic and large 
examples of our own bull frog have been known to swallow half- 


20 Zoologica: N. Y. Zoological Society. eG ae: 


grown rats, small chicks, ducklings, sparrows, toads and young 
snakes. 


Insects, of course, make up the greater percentage of the 
food. 


Some species of this genus are quite terrestrial, only entering 
the water during the breeding season, while others are typical 
water frogs never wandering far from their native stream, pool 
or swamp. 


The genus Gampsosteonyx resembles an ordinary frog, but 
has vertical pupils. The terminal points of the fingers end in 
sharp, bony claws which perforate the skin of the finger tips. 
One species is known: G. batesi from the French Congo. 


Subfamily DENDROBATINAE—Harlequin Frogs. 


These small frogs are separated from the others of the fam- 
ily by the absence of teeth from both jaws and comprise three 
genera: Dendrobates, of Tropical America, Mantella, of Mada- 
gascar and Cardioglossa with one species C. gracilis of the French 
Congo. The frogs of the genera Mantella and Dendrobates are 
very much alike in shape, size and in possession of a striking 
color pattern. Deep black, bright blues, brilliant reds, greens and 
yellows in many contrasting combinations are the colors which 
often form fantastic patterns. 


The tiniest insects are the food of these little harlequins of 
the frogs’ world and they are usually found near fallen decaying 
tree trunks, where they feed on small termites, or in banana plan- 
tations, where they can be seen in numbers about the fallen and 
decaying fruit which attracts myriads of small fruit flies. 


Dendrobates has seventeen species, Mantella nine species 
and Cardioglossa has one. 


Subfamily RANINAE—True Frogs. 
Large Eyed Frog, Rana chrysoprasina, Cope (Boulen- 
ger, Cat. Batr. Sal. P. 49). 


Color: The head is green and the back, sides and limbs a 
yellowish olive with a few brown specks. A dark line extends 


1915] Deckert: Two Series of Amphibians. 21 


from the tip of the snout through the nostril and eye over the 
tympanum and below the lateral glandular fold to the groin. 
The edge of the upper jaw has a few small brown spots, the 
eye is brassy yellow and the sides and belly an immaculate gold- 
en yellow. 


Structure: The head is broad, flat, snout acuminate, pro- 
jecting beyond the mouth and the tympanum two-thirds the 
diameter of the eye, which is very large. The tips of the fingers 
and toes are slightly dilated, fingers very long and slender, and 
the toes webbed four-fifths of their length. The skin is very 
finely pustulated above and smooth underneath. There is a lat- 
eral fold on each side of the body and narrow longitudinal gland- 
ular ridges on the calf of the leg. 


Size: The specimen examined was three and one-quarter 
inches from snout to vent. 


Range: Costa Rica. 


One adult, one young frog and several tadpoles of this beau- 
tiful species were collected near Guapiles. This frog is very 
shy, as most large-eyed frogs usually are, and seeks cover with 
great rapidity when disturbed. Like most water frogs it is a 
good feeder, and so far has proven a very satisfactory captive. 
When taken up it will sit quietly in the open hand and will not 
jump unless frightened by a quick movement. Several of the 
tadpoles have metamorphosed and are living now as young frogs 
in a vivarium with small tree toads on the main floor of the 
Reptile House. In the daytime they usually sit concealed under 
some moss, but come forth with the darkness and occupy the 
pan of water provided for them. 


Godman’s Frog, Rana godmani, Guenther (Biologia 
Centr. Am.). 


Color: The color is greenish olive above with indistinct 
darker spots and whitish below. 


Structure: The structure is like that of Rana clamitans, 
but with much shorter legs. 


oe Zoologica: N. Y. Zoological Society. [ibs 


Size: One young specimen metamorphosed from a tadpole 
is in the Reptile House. This frog is just as shy as the preced- 
ing species, constantly hiding under a large, flat stone in its 
terrarium. The size of the adult is from two and one-half to 
three and one-half inches. 


Range: Costa Rica. 


Subfamily DENDROBATINAE—Harlequin Frogs. 
Blue Legged Frog, Dendrobates typographus, Kefer- 
stein (Roulenger, Cat. Batr. Sal. P. 148). 


Color: This frog is a brilliant red above and below with 
or without tiny black dots. The legs and forearms are brilliant 
dark blue or blue-black, and on the upper side of the thighs 
there is a row of small red dots. The blue of the inner arm 
extends across the breast. 


Structure: The snout is obtuse and the canthus rostralis 
rounded. The tympanum is distinct but small, measuring about 
one-half of the diameter of the eye, and the interorbital space 
twice the width of the eye. The arms and legs are slender and 
moderately long with disks on the fingers and toes equal to or 
exceeding the tympanum in size. The skin is smooth and shiny. 


Size: Adult frogs are one inch or less from snout to vent. 


Range: Costa Rica. Eight of these queer little frogs were 
collected by Mr. Crandall. They were found prowling along the 
decaying timbers of a fallen fence, probably hunting for the 
small white termites that usually infest such places. 


Scarlet Frog, Dendrobates typographus, subspecies ig- 
nitus, Cope (Proc. Acad. Phil. 1874). 


Color: As the name implies, it is brilliant red all over ex- 
cept for a small star-shaped area on the breast and a larger 
one at the junction of the hind legs and the belly, beneath which 
is dark blue. 


1915] Deckert: Two Series of Amphibians. 23 


. Structure: Exactly like D. typographus var. typica. 


Range: Costa Rica. Rarer than the typical form. One 
specimen from Limon. 


Ornate Frog, Dendrobates tinctorius, Schneider, (Bou- 
lenger Cat. Batr. Sal. P. 142). 


Color: The color is very variable, the single specimen col- 
lected by Mr. Crandall was bright emerald green and black; 
the green predominating above and the black below. 


Structure: The snout is truncate and the canthus rostralis 
rounded. The interorbital space is wider than the diameter of 
the eye and the tympanum one-half the.diameter of the eye. The 
arms and legs are slender, with the disks of the fingers and toes 
distinct. The skin is smooth and shiny. The male has a sub- 
gular vocal sac. 


Size: From snout to vent it is one to one and one-half 
inches in length. 


Range: Tropical America. This specimen has proven the 
hardiest of the smaller frogs brought from Costa Rica and lives 
on tiny fruit flies that are enticed into its terrarium with slices 
of banana, apple or pieces of wet bread. It can see a fly at quite 
some distance, and with short hops follows every turn of its 
flight until it alights within reach, when it is greedily snapped 
up. The tongue, which is not notched behind like that of the 
frogs of the genus Rana, can be thrust out for quite some dis- 
tance. This curious little creature does not always hop, but will 
often elevate its body on its long slender legs and stalk around 
as though walking on stilts. The adhesive disks, although tiny, 
are large enough to enable this little frog to climb up the glass 
sides of its terrarium. Owing to their intensely bright colora- 
tion, Mr. Ditmars has suggested the very appropriate name of 
Harlequin Frogs for these odd creatures. 


The frogs of the genus Dendrobates are known for the in- 
tensely virile poison contained in their skin secretion. This 
poison, especially that of D. tinctoriuvs, has been put to several 


24 Zoologica: N. Y. Zoological Society. TT sa: 


uses by the aborigines, one being that of an arrow poison, and 
another a bleaching agent, which turns the green of parrots’ 
feathers to yellow. The poison, like that of the toads, has no 
power to injure by touch, acting only when injected into the 
circulation or rubbed into a deep wound. The life habits of 
these queer little creatures are also worthy of note, especially 
the nursing or carrying about of the tadpoles. These habits are 
shared by the frogs of the Ranoid genus Prostherapis and Ar- 
throleptis, and by species of Hylodes of the Cystignathidae. As 
has been observed by the naturalists J. Natterer, H. S. Smith 
and A. Kappler, frogs of this genus will take their tadpoles upon 
their backs and carry them to another pool in times of drought. 
A. Kappler saw D. tinctorius and D. trivittatus in Surinam go 
into evaporating pools, sit still awhile and then emerge with 
tadpoles, some frogs carrying from twelve to eighteen, which 
adhered to their backs by means of a sticky secretion. Whether 
this secretion is exuded by the frog or the tadpole is not known 
as yet, nor has the sex of the nurse been determined up to the 
present writing. 


Since the arrival of this collection additional material has 
been promised us by several gentlemen who have been to the 
canal zone and who, upon being shown the specimens, said that 
they were fairly abundant in those regions. Other interesting 
frogs, toads and tree toads have also been promised us from 
that region, and it is hoped that the writer will be enabled to 
make further observations on these interesting and little-known 
creatures. Much work is yet to be done in this line of investiga- 
tion, and a large field is open for the student having the oppor- 
tunity to observe these creatures in their natural environments. 


AK 


Richard Deckert, pinx 


FIG. 4. AUSTRALIAN “FROGS” 


Sand “Frog’’, Limnodynastes dorsalis, Gray 
Perron’s Tree Toad, Hyla perronii, Bibron 


(Two color phases) 


FROGS AND TOADS FROM NEW SOUTH WALES.* 


The Australian frogs and toads that are described in this 
paper will be of especial interest, since it is the first time that 
these important species have been exhibited in the Reptile House 
(alive) or the United States. Among them are included the 
following species: 


Family BUFONIDAE—Toads. 
Australian Toad, Pseudophryne australis, Gray. 


Family HYLIDAE—Tree Toads. 
Perron’s Tree Toad, Hyla perronii, Bibron. 
Golden Tree Toad, Hyla awrea, Lesson. 
White’s Tree Toad, Hyla coerulea, White. 


Family CYSTIGNATHIDAE—Arch-Jawed Toads. 
Sand “Frog,” Limnodynastes dorsalis, Gray. 
Silver “Frog,” Heleioporus pictus, Peters. 


Family BUFONIDAE—Toads. 


Australian Toad, Pseudophryne australis, Gray. (Bou- 
lenger Cat. Batr. Sal. P. 277). 


Color: Above, the body is blackish brown with a yellow or 
reddish narrow streak on the posterior back. The posterior 
sides of the arms are bright orange-yellow and the rear sides of 
the thighs usually have a few yellow spots. The throat, abdomen 


and under surface of the arms and legs are marbled black and 
white. 


Structure: The head is rounded; canthus rostralis not pro- 
duced. The interorbital space is as wide as the eyelid. The 
fingers and toes are short without web or dilatations. The skin 
is smooth or with a few indistinct flat warts. With the male 
there is an internal gular vocal sac, and an oval flat gland on the 
hinder side of each thigh. The pupil of the eye is horizontal. 


*A resumé of the Families embraced in this article will be found in the 
preceding pages. 


26 Zoologica: N. Y. Zoological Society. [iso 


Size: One and one-quarter inches, snout to vent. 


Range: Australia. The two specimens examined were from 
near Sydney, N.S. W. This little toad is said to be quite com- 
mon all over Australia in localities favorable for amphibian life, 
and is interesting on account of its breeding habits. The large 
eggs are laid in damp places in numbers up to ninety under 
stones, stumps and other hiding places, and have been found in 
November, January and May near Sydney. Oviposition takes 
place after heavy rains and the next rain is depended upon to 
set the larvae free. 


This may occur within two or three weeks, or three or four 
months. The embryo is very tenacious of life and, as noted 
above, will accommodate itself to remaining in the jelly-like mass 
of the egg for a long time. The actual limit for this has not yet 
been determined; four months being the longest time recorded. 
The two specimens now in our collection seem to be hardy, and 
live in company with Hyla pickeringvi and Dendrobates tinctorius 
in a small terrarium, the bottom of which is covered with very 
damp wood-pulp. 


They feed greedily on all kinds of small insects; the method 
of hunting differing from that of any frog or toad observed by 
the writer. Instead of hopping they slowly and deliberately 
creep up to their intended prey, moving each arm and leg sep- 
arately, first an arm, then the leg of the opposite side, then the 
other arm and lastly the other leg, giving the whole movement 
a singular mechanical appearance. When about a half-inch from 
the insect, the tongue shoots forth with lightning-like speed and 
the insect vanishes. The toad retains its seemingly strained po- 
sition until another victim is sighted, when the whole manceuvre 
is repeated. When disturbed, however, these toads hop in ordi- 
nary fashion. 


Family HyLIpDAE—Tree Toads. 
Perron’s Tree Toad, Hyla perronw, Bibron (Boulenger 
Cat. Batr. Sal:-P. 390). 


Color: Brown above, but subject to great variation and 
eolor-change. There may be a distinct pattern of dark marblings 


1915] Deckert: Two Series of Amphibians. ZA 


or dots, but when the animal is at rest, it is usually dark brown, 
without dark spots, though with bright yellow dots, which are 
lined with black and intermingled with emerald green spots 
slightly larger than the head of a pin. The abdomen is white, 
throat (male) marbled with brown, and the arm-insertion (be- 
hind), the groin and the concealed surface of thigh, calf and 
foot are bright orange, marbled with black. These colors are 
not seen when the frog is at rest. The eye is silver with the 
pupil contracted to a tiny square with four black lines radiating 
from it and dividing the eye into quarters. When active the 
color fades until the whole frog is pale reddish or yellowish 
brown and the arms and legs barred with darker brown. The 
color of the eye also changes, becoming bright yellow as the pupil 
expands into a regular, horizontal oval. 


Structure: The head is broader than lony, snout rounded 
and canthus rostralis rounded. Loreal region slightly concave. 
Interorbital space equal in width of the diameter of the eye. 
Tympanum distinct and two-thirds the diameter of the eye. The 
fingers are half webbed and the toes about three-fourths webbed, 
with large adhesive disks about half the diameter of the eye. The 
upper surface is sometimes smooth and occasionally covered with 
small roundish warts. A fold of skin extends from the eye over 
the tympanum to the shoulder, and another fold across the 
breast. The male has a large subgular vocal sac. The entire 
lower surfaces are granulated. 


Size: It attains a length of two and a half inches from 
snout to vent. 


Range: Northern and Eastern Australia and Tasmania. 

One specimen from’ near Sydney, New South Wales, is in 
the collection of the Reptile House. It usually sits in a corner 
and near the top of its vivarium, with the pupils contracted, ap- 
parently sound asleep. At dusk it becomes active, climbing slowly 
over the glass sides of the case until it spies an insect, when it is 
capable of making enormous leaps. It does not seem particularly 
shy, and will allow itself to be handled, clinging to one’s fingers 
with its sticky toes. When the plants in the Reptile House are 
being syringed in the morning and evening, the sound of the 
splashing water stimulates this tree-toad to giving voice to its 


28 Zoologica: N. Y. Zoological Society. wee 


loud call. This call resembles the noise of the pneumatic drill 
used by structural iron workers, and might be described as a 
loud, metallic rattling. The throat pouch is expanded into a large 
globe, larger than the tree-toad’s head, while the entire body 
vibrates with the force of the exertion used in producing the call. 


Golden Toad, Golden “Frog,” Hyla awrea, Lesson (Bou- 
lenger Cat. Batr. Sal. P. 410). 


Color: The general body color is a bright metallic green, 
sometimes bluish, sometimes yellow in tone. From the tip of 
the snout through the nostrils, over the eyes and tympanum to 
the groin, extends a wide brown band, and on the back there are 
usually a series of spots of varying size and shape, but of the 
same color, which sometimes fuse into longitudinal bands. A 
brown band, which becomes yellow or silver, passes from the 
tip of the snout along the upper margin of the mouth and ends 
at the shoulder. From the nostril through the eye, interrupted 
by the tympanum, is a black streak which ends behind the shoul- 
der. The arms and legs are brown, the former spotted, and the 
latter longitudinally banded with green. All of the brown spots 
and bars may become beautifully golden or coppery bronze, and 
the glandular, lateral fold a pale, golden color. The color at the 
groin is deep blue-black. The sides are green, sometimes with a 
bronze shading, the tympanum bronze and the under sides pure, 
silvery white. The entire toad is subject to strong color changes, 
sometimes becoming plain blue-black with metallic reflections. 
The eye is large, brilliant and of a beautiful, reddish-gold color. 


Structure: In general form it is like a Rana. The head is 
a little longer than broad, the interorbital space narrower than 
the eyelid, tympanum about half the diameter of the eye, canthus 
rostralis distinct and the loreal region concave. The fingers are 
free and the toes almost entirely webbed. The disks of the fin- 
gers and toes are small. The skin may be entirely smooth or 
warty above. A longitudinal fold extends from the eye to the 
groin. The male has two internal vocal sacs. 


Size: It attains a size of two and a half to three and a half 
inches from snout to vent. 


FIG. 5. GOLDEN TREE TOAD, HYLA AUREA, LESS. 


FIG. 6. WHITE’S TREE TOAD, HYLA COERULEA, WHITE 


Zoologica Vol. II, No. 1. Face page 29. 


1915] Deckert: Two Series of Amphibians. 29 


“Range: The Golden Tree Toad is found throughout Aus- 
tralia and some of the Australian Islands. 


This is one of the commonest of Australian toads and is 
called “bell frog” in its native country. In shape, habits and 
actions it is absolutely different from any other species of the 
genus Hyla, so much so that other naturalists have made it the 
type of a distinct genus, Ranoidea, Tschudi. But the internal 
structure is that of a typical Hyla and Boulenger in his admir- 
able “Catalogue of Batrachia Salientia,’” has included it among 
the Hyla, where it rightly belongs. It is a large, gorgeously col- 
ored species and reminds one of a water frog in appearance as 
well as in habits. In its terrarium in the Reptile House it will 
sit for hours on the ground or in the pan of water provided, 
never attempting to climb up the sides of its cage like its cage- 
mates. It is also like the water frogs, distinctly cannibalistic, 
seemingly preferring small frogs to any other food. It is a greedy 
feeder and will eat all kinds of insects. After sundown its call 
can be heard in the Reptile House. This call is a long, drawn- 
out and loud croak, very coarse but sometimes ending in a bell- 
like note, “bong”; also very loud. The species is quite hardy and 
can endure considerable cold. At a low temperature it loses its 
beautiful tints and assumes a plain blackish-olive or blue-black 
color. It is most beautiful and active at a temperature of about 
70 degrees F. When taken up it will struggle violently, and 
upon being released, hop away in a series of rapidly executed, 
enormously long leaps, reminding one of a water frog. Its eggs 
are laid in a white, frothy mass in pools, canals, wells or other 
permanent bodies of water, differing herein from most Austra- 
lian frogs. Oviposition takes place during August and Septem- 
ber. If these months should be dry, however, the species waits 
until the following spring. In its native country this species 
frequents permanent bodies of quiet water, never climbing on 
the bushes and trees surrounding such places, but always remain- 
ing at the water’s margin, plunging into the protecting element 
at the slightest alarm, like a true water frog. The “golden frog” 
is hunted at night by the Australian natives (Bushmen), with 
the aid of lanterns and torches. Numbers of them are spitted 
upon a sharp stick as they are caught, roasted over an open fire, 


30 Zoologica: N. Y. Zoological Society. is 


and eaten with great relish by the natives, without taking the 
trouble to remove the viscera. 


White’s Tree Toad, Hyla coerulea, White (Boulenger 
Cat. Sal. P. 383). 


Color: The general color of the body is a bright, leaf-green 
to dark olive above; undersides pinkish-white, and the concealed 
surfaces of the thighs and arms, fleshy-pink. A few round or 
elongate white spots are sometimes present on the sides, limbs 
and back. 


Structure: The head is large, broad and flat, with rounded 
and truncate snout, canthus rostralis distinct, loreal region con- 
cave and the interorbital space much wider than the diameter of 
the eyelid. A strong fold extends over the tympanum, which is 
from two-thirds to four-fifths the diameter of the eye, to above 
the arm insertion. The hands are large, fingers are webbed one- 
third and the toes almost entirely webbed. The adhesive disks 
are very large, those of the hands equal in diameter to the tym- 
panum, those of the feet smaller. There is a tarsal fold, and the 
subarticular tubercle is prominent. The skin is smooth and 
shiny, much thickened on the head and scapular region, studded 
with large pores, and the under surfaces are granulate. The 
male has a large gular vocal sac. 


Size: This species is one of the largest of the Hylidae, 
attaining a length of five inches; our largest specimen measuring 
four inches from snout to vent. 


Range: Australia, Tasmania, Australasia, Malaysia. The 
nine specimens exhibited in the Reptile House are from New 
South Wales. This large and fine species is a typical Hyla in 
appearance as well as in habits. In its coloration, it is one of 
the most constant of all Hylas, resembling in this respect our 
own Hyla cinerea, s. carolinensis. The upper sides are always 
some shade of green. The iris is a beautiful, golden bronze when 
the frog is awake, and when asleep may be pale, silvery-blue. In 
the Reptile House it shares its terrarium with Hyla aurea. Un- 
like the latter, it is tame and confiding, and when picked up will 


1915] Deckert: Two Series of Amphibians. 31 


cling to the hand, climb around it looking for a comfortable spot 
to sit, and upon finding one will settle down, tucking its hands 
and feet well under. A German naturalist named Riedel relates 
the following amusing incident, illustrating well the phlegmatic 
disposition of this tree-toad: He had taken a specimen to show 
a friend, placing it on the cover of a large beer-stein, such as 
are in general use in Germany. Whenever the owner lifted the 
stein and tipped the cover back to take a drink, he turned the 
tree-toad upside down in doing so. The latter did not jump, but 
clung to the cover with its enormous hands, and when the cover 
was placed in its natural position it immediately settled down in 
its former position, tucking its hands and feet under its body, 
always a sign of contentment. It remained thus on the cover of 
the stein for over half an hour, being frequently disturbed by 
the drinking operations of the owner. 


This toad has the sense of locality very much developed, re- 
turning again and again to the same spot for basking or sleep- 
ing. In this it resembles our Hyla versicolor. On cold nights 
when all the other tree-toads in the collections creep under cover, 
these big fellows will be seen sitting motionless in their accus- 
tomed places. It will also sit for hours in the brightest sunlight, 
apparently as indifferent to the burning heat as it is to the cold. 
The voracity of this species is in keeping with its size. Any- 
thing is welcome from mice and small frogs to all kinds of insects, 
worms and small crustaceans. It will eat until abnormally gorged. 
In cleaning the vivarium, I had occasion to put these large tree- 
toads temporarily in another vivarium containing Hyla arborea, 
Bufo calamita and Rana temporaria. The big fellows were 
hardly placed in this cage when they attempted to capture and 
eat the smaller amphibians, and if they had been left to their 
own devices, would soon have swallowed all the rightful inhabit- 
ants. The call of the male can be heard in the Reptile House 
usually late in the afternoon, and consists of the syllables “kra- 
kra-kra-kra,’ repeated about twenty times. At first it is not 
loud, but gains in volume as the cry proceeds, until near the end 
it resembles the barking of a large dog. The enormous finger 
disks give this frog unusual clinging powers, and when one is 
lifted from its place, it will usually grasp any object near and 
hold on with such strength and tenacity that great care must be 


>) 


32 Zoologica: N. Y. Zoological Society. [ Est 


exercised not to injure the animal in trying to loosen its hold. 
Hyla coerulea breeds in permanent bodies of water such as cis- 
terns, wells, reservoirs and canals. The spawn, like that of 
H. aurea, is laid in the water in large clumps and enveloped in 
a foamy mass that floats on the surface. In its native land, this 
species is found in numbers under the roofs of outhouses, veran- 
das, and other places of concealment; sometimes coming into the 
dwelling houses at night, attracted by the light of lamps and by 
the presence of insects. When there is rain, the frogs around 
a house will set up a deafening din, almost driving the occupants 
to distraction. The species should be useful as an insect-destroyer 
on account of its size and proportionate voracity. The geograph- 
ical distribution of this tree-toad is very extensive, ranging 
throughout most of the islands of the Pacific and even some of 
the Indian Ocean, this being probably due to the ease with which 
the frog can be carried from place to place in cargoes of lumber 
or fruit. The phlegmatic habits of this species, no doubt, also 
favor such distribution. 


Family CYSTIGNATHIDAE—Arch-Jawed Toads. 
Sand “Frog,” Limnodynastes dorsalis, Gray (Boulen- 
ger Cat. Batr. Sal. P. 261). 


Color: The color is variable, usually an olive brown, with 
or without spots. Our specimen is blue-black above with a few 
inky black spots on the sides and posterior back. <A broad, yel- 
low streak extends from in front of the eye to the commissure 
of the mouth, and a series of broad yellow spots, which in some 
specimens is blended into a band, from behind the tympanum to 
the groin. A second series of yellow blotches occurs below this, 
and a number of greenish-yellow spots on the upper eyelids and 
back and a narrow yellow streak on the coccygeal region. The 
outer edge of the tarsus and outer toe are yellow, and the rest 
of the foot bluish white. There are some yellow spots on the 
arms, belly white, and the throat yellow. The eye of both sexes 
is bright, golden-yellow. 


Structure: The head is large and the snout rounded. The 
mouth is very wide, eyes large, very prominent and brilliant in 


1915] Deckert: Two Series of Amphibians. 33 


coloring, pupil vertical and the interorbital space equal to the 
width of the upper eyelid. The tympanum is distinct and two- 
thirds of the size of the diameter of the eye; skin smooth above 
and below; arms and legs short, and the fingers and toes slender. 
Subarticular tubercles prominent. A cuneiform shovel on the 
tarsus. A parotid-like swelling on the calf (male only). The 
male has a subgular vocal sac. There is an elongate, yellow 
gland extending from beneath the eye to above the arm-insertion 
(see color description). 


Size: Our specimen measures two inches from snout to 
vent. 


Range: Australia. 


This odd creature has a startling resemblance to our spade- 
foot toads, Scaphiopus, of the family Pelobatidae, in habits, color- 
ing and structure. In Australia it occurs in sandy regions which 
are dry the greater part of the year—in fact, without permanent 
bodies of water, and is therefore compelled to await the occa- 
sional heavy rains for the consummation of its breeding opera- 
tions. This is likely to happen almost any time of the year. The 
spawn is laid in frothy clumps in the temporary pools formed 
by these rains, and the metamorphosis is correspondingly rapid, 
exactly as in our Scaphiopus. In Scaphiopus, however, the eggs 
are laid in strings. The species is fairly common on the south 
coast and is called ‘‘sand frog” by the farmers. It is sometimes 
dug up in the gardens. Its habits are nocturnal as indicated by 
the vertical pupil, and it feeds chiefly on the large spiders that 
abound on the ground at night. In the Reptile House this frog 
is kept in a small terrarium in company with the Silver Frog 
and seems fairly hardy. It is a good feeder with cannibalistic 
tendencies. By day it lies buried in the fine gravel, but at night 
comes forth, sitting partly buried, alert for any insect that may 
come its way. It progresses by short hops of startling rapidity. 


Silver “Frog,” Heleioporus pictus, Peters (Boulenger 
Gat2Batr. sal. P. 272). 


Color: The body is yellowish or grayish olive with large 
black, brown or green spots. There is sometimes a pale yellowish 


34 Zoologica: N. Y. Zoological Society. [sad 


vertebral line. A long, white gland extends from beneath the 
eye to the shoulder, and a black streak from the nostril through 
the eye to above the shoulder. The belly is white, and the throat 
(male) yellow. The arms and legs barred with the same color as 
the large spots on the back, and the eye is a pale, brassy yellow. 


Structure: The head is moderately large and the snout 
rounded. Eyes close together ; fingers and toes short; subarticu- 
lar tubercles well developed and metatarsal tubercle large. Arms 
and legs short; tympanum indistinct; skin smooth and the toes 
very slightly webbed. 


Size: Adult males one and a half to one and three-quarter 
inches from snout to vent. 


Range: Australia (Victoria and New South Wales). 


This species is not nearly so nocturnal as the preceding, and 
is always ready to feed on insects placed in its terrarium. In 
New South Wales it is called ‘‘silver frog’’ and seems fairly com- 
mon around Sydney. Its eggs are laid in pools formed by the 
heavy rains and the metamorphosis is rapid, keeping pace with 
the evaporation of the water in the temporary pools. This spe- 
cies does not seem to be possessed of great powers of color-change, 
the specimens living in the Reptile House keeping the same dress 
under various conditions, which would induce other frogs, at 
least, to change from light to dark or vice versa. The call is a 
rattling croak which is not very loud, the throat pouch being 
inflated to the size of a small marble during the call. The “‘silver 
frog’ is rather shy, squatting low in its vivarium upon being 
approached, and when further disturbed will dart about in great 
haste, seeking concealment. It is a ready feeder, however, and 
we hope to be able to keep it alive for some time to come. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 2. 


A TETRAPTERYX STAGE IN THE 
ANCESTRY OF BIRDS 


By 


C. WILLIAM BEEBE, 
CURATOR OF BIRDS 


Pees sheD. BY THE SOCIETY 
THE ZOOLOGICAL PARK, NEW YORK 


NOVEMBER, 1915 


OFFICERS 
OF THE 


New York Zonlogical Society 


President: 
HENRY FAIRFIELD OSBORN. 


Vice-President and Treasurer : 
Secretary : Percy R. PYNE, 
MADISON GRANT, 30 Pine Street. 


11 Wall Street. 


Exerutive Committee 
MADISON GRANT, Chairman. 


PERcY R. PYNE, FRANK K. STURGIS, 
*SAMUEL: THORNE, LISPENARD STEWART, 
WILLIAM WHITE NILEs, WATSON B. DICKERMAN, 
WM. PIERSON HAMILTON, HENRY FAIRFIELD OSBORN, 
ex-of ficio. 


Anditing Cammitice ‘ 
WILLIAM WHITE NILES, Chairman. 
H. CASIMIR DE RHAM, LISPENARD STEWART. 


Geueral Officers 
Director of the Zoological Park: WitLIAM T. HoRNADAY. 
Director of the Aquarium: CHARLES H. TOWNSEND. 
Prosector: Dr. GEORGE S. HUNTINGTON. 
Architect: C. GRANT LA FARGE. 
Consulting Engineer: H. DE B. PARSONS. 
Assistant Secretary: H. J. SHORTER, 
Assistant to the Treasurer: R. L. CERERO. 


*Deceased. 


ZOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 2. 


A TETRAPTERYX STAGE IN THE 
ANCESTRY OF BIRDS 


By 


C. WILLIAM BEEBE, 
CURATOR OF BIRDS 


EUS LDS HED Boy THE Sim © Dl Cora 12 ed a 
Tite AOoObLO GICAL PARK, NEW YOR K 


NOVEMBER, 1915 


79 
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avardsyjuotgy “SON ‘[] “JOA PaL60]007 


Spiiq SULAl| Jo Sunod 9yy ul pue *xAtoydoaeyoty ul yueseid sieyoeIeYyO uO peseq SI Suimevap a 
PaIq att Y P Y iq \ P PUL 
sqduid AO AULSHONV AHL NI AOVLIS XAYUALIVULEAL “1 Dla 


A TETRAPTERYX STAGE IN THE 
ANCESTRY OF BIRDS 


By C. WILLIAM BEEBE, 
Curator of Birds. 


I. Introduction. 


II. Pelvic Wing: 
White-winged Dove. 
Domestic Pigeons. 
Jacana. 

Great Horned Owl. 
Archaeopteryx. 


III. Argument. 
PART I.— INTRODUCTION. 


Our knowledge of the genealogy of birds is in inverse ratio 
to the abundance of these organisms on the earth today. We 
know of quite primitive forms of fish—both fossil and living—of 
reptiles and of mammals. But of living birds, those which show 
encouragingly primitive characters exhibit also an equal number 
of extremely specialized ones. 


Some six or seven million years ago in the Cretaceous Period 
we know that there lived creatures which undeniably deserve the 
name of birds. Ichthyornis was a strong-flying, tern-like type 
with numerous, recurved teeth, and Hesperornis was also toothed, 
but practically wingless, essentially a diving bird, and on shore 
probably more helpless than a seal. 


In the Jurassic, let us say four million years earlier, two 
more or less complete fossil skeletons have been discovered, and 
an odd feather or two of the famous Archaeopteryx, the sine qua 
non of avian genealogy. Teeth again we find in a very lizard- 
like head; delicate, weak, wing bones supporting a good-sized but 
rounded wing, and the fore limb terminating in three well devel- 


40 Zoologica: N. Y. Zoological Society (Iis2 


oped, clawed fingers; a long, vertebrated tail, with a pair of 
excellent feathers sprouting at each joint and a pair of feet ad- 
mirably adapted for perching. The unmistakably reptilian traces 
give weight to Huxley’s superclass Sauropsida. 


Slight though this evidence is compared to the imposing 
array of fossil reptiles and mammals, it nevertheless provides an 
unmistakable lead in the direction of small, arboreal, lizard-like 
creatures along a still earlier line of ancestry. 


I do not wish in this paper to discuss, except in one respect, 
the various characters of Archaeopteryx. I am working out a 
life-sized restoration of a flock of seven of these winged creatures 
of ancient times and not until this is completed shall I feel confi- 
dent of expressing any new views on the general character of 
this much discussed creature. 


My present thesis, while in a way independent of Archaeop- 
teryx, yet is given force in dynamic presentation by consideration 
of this strange creature. 


Perhaps the most astounding thing about this being is the 
perfection of its wing and tail feathers. Without going into rea- 
sons, I am convinced that Archaeopteryx was a bird of very 
limited powers of flight. I am not certain that it could flap at 
all and if it could, its aerial feats hardly equalled those of a 
modern tinamou or domestic fowl. It certainly had very excellent 
powers of scaling, and in this direction probably exceeded any 
modern flying squirrel or lemur. 


Whether this be conceded or not is aside from my point, 
which is concerning the origin of this wing. Our knowledge of 
the workings of evolution often enable us to visualize the growth 
and later development of an organ, its subsequent specialization 
and perhaps ultimate degeneration, while we utterly fail to ex- 
plain its origin or early development. It is well within the limits 
of gradual cumulative variation to admit the change from an 
Archaeopteryx to a modern wing. The wing bones increase in 
size and those of the hand coalesce, the fingers become mittened 
in flesh and tendon; the primaries increase in number creeping 
out upon the phalanges, and the muscles wax stronger, become 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds AL 


larger and find adequate place for attachment upon a greatly 
enlarged sternum. But how could the wing have reached its 
Archaeopteryx stage of development? 


In Mexican or Neotropical jungles bordering rivers and 
streams it is a common sight to see great iguanas resting high 
among the upper branches and foliage. When suddenly alarmed 
or toward sunset, these great reptiles do not bother to climb 
slowly down along their back trails which they so laboriously 
mounted earlier in the day. They recklessly launch out into mid- 
air and with legs widely extended, body flattened, toes clutching 
at the empty air, they hurtle downward, landing with a crash 
into the underbrush or with a splash in the water. Perhaps their 
flattening may help somewhat to break their fall, but I doubt if 
this would save their life were they to land upon hard ground. 
Twice, in fact, I have seen iguanas after a had take off, half turn 
in the air, so that they landed in the water on their side or in 
one case actually upside down, when the reptile seemed stunned 
for a minute before it turned over and swam from sight. Here, 
it seems to me, we have a very probable anlage of scaling flight, 
as ultimately perfected in Archaeopteryx. 


But if we arm our imaginations with a prejurassic, para- 
chuting lizard on the one hand, and Archaeopteryx on the other, 
we still have a hiatus which no logical combining of proportional 
characters will bridge. Suppose if you will that the scales along 
the posterior edge of the fore leg and those along the tail begin 
to lengthen. Carry these along to a fair development and then 
start the hopeful organism out into mid-air and it will prove an 
utter failure. The scaly primaries may be sufficient to support 
the front part of the body, but the tail scales would certainly 
not suffice both to balance and to bear up the remainder of the 
lizard. The result would be a woeful sagging which must bring 
instant disaster,—a herpetological Darius Green which could not 
hope to leave offspring to work out their fossil destiny. The 
development of scaling flight with nothing to correspond to the 
great lateral and caudal membranes of flying mammals is incon- 
ceivable. Something is needed to bridge over the very beginnings 
of the parachuting wing function. Even a flying fish has two 
nodes of aerial support during its brief essay into a thinner 


o 


42 Zoologica: N. Y. Zoological Society [II;2 


medium. It spreads capable little planes aft as well as forward. 
Cut off the pelvic fins and I imagine it would plop hindmost into 
the water almost as soon as it emerged.” Some such accessory 
has always seemed to me necessary if we are to complete our 
lizard-to-Archaeopteryx line of ascent. 


Recently, while examining the fresh body of a four-days’-old 
White-winged Dove in the New York Zoological Park, I observed 
on its almost naked body a remarkable development of sprouting 
quills across the upper part of the hind-leg, and extending toward 
the tail across the patagium just behind the femur. A second 
glance showed that this was no irregular or abnormally preco- 
cious development of part of the femoral pterylum, but a line of 
primary-like sheaths, many of which had a very definitely placed 
covert. The iguana-Archaeopteryx puzzle flashed through my 
mind and I at once followed up the clue thus given. For the two 
sketches illustrating my idea of the Tetrapteryx Stage of alar 
evolution I am indebted to Mr. Dwight Franklin. 


PART I].— PELVIC WING. 


The detection of this interesting character occurred in Au- 
gust, too late for observations on many forms of nestling birds. 
I embody in the following notes all that I have been able to gather 
together on the existence of this curious pelvic or femoral wing. 


WHITE-WINGED DovVE. 
Melopelia asiatica (Linn.) 


Several of these birds had been reared during the present 
season in the special breeding cages of the New York Zoological 
Park. On August 19 a four-days’-old squab, the only nestling 
of a second brood, was found dead in its nest and brought to me. 


Its leaden grey body appeared almost bare, being covered 
sparsely with the characteristic short, greyish white, filamentous 
down. Three areas showed precocious development of contour 
feathers, the wing proper, the pelvic wing and the tail. In the 
former, twenty-two flight feathers were developed, of which ten 
were primaries averaging 10 mm. in length, nine were secondar- 


FIG.8. DOMESTIC PIGEON SQUAB FIG. 9. SQUAB OF WHITE-WINGED DOVE 


Showing great development of the pelvic wing. Four days old, with wing and leg extended, 
The leg is flexed, hence this atavistic showing the wide spread pelvie wing 
wing is folded 


FIG. 10. DETAIL OF PELVIC WING OF WHITE-WINGED DOVE, SHOWN IN FIG. 9 


The wing consists of twelve flights and six coverts 


Zoologica Vol. II, No. 2. Face page 42 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds 43 


ies, grading inward from 8 to 5 mm., and three were tiny terti- 
aries. The primaries had only a single row of strongly developed, 
greater coverts. Four rows of secondary coverts were sprouting, 
the central ones pure white, indicating the future color pattern 
of the wing. 


Next in development to the wing proper, were the feathers 
of what, for lack of a better name, I call the pelvic wing. This 
seems inexplicable on any other hypothesis than the vestigial 
secondary plane, which must have been of the utmost importance 
in the ancestral scaling flight. 


This area begins on the anterior outer edge of the crus or 
leg proper, about one-third of the distance down from the knee. 
From this place it extends backward across the tibia almost at 
right angles to the backbone of the body, and, posterior to the 
femur, following the patagium, which lies between the leg and the 
body. It ends on the side of the body at an equal distance from 
the outer tail feather and from the pelvis between the acetabula. 
The areas are similar on both sides. There are twelve fain or 
flight feathers. Feathers 1 to 6, extending from the body out- 
ward along the femoral patagium, all have a well-developed cov- 
ert. The next six flights, numbers 7 to 12, lie close together on 
the flesh of the leg itself and show no signs of coverts. Counting 
from within outward these feathers measure as follows: 


IIL Sean eee cee My Gyr en 0 6 6 6 O25) <5 4 3 1 mm. 
Covers 26 eee ee 3.5 A 4. 5 5.5 5.5 mm. 


The tail is much less advanced than the pectoral and pelvic 
wings, the rectrices and a single row of upper and another of 
under coverts being all equally advanced, measuring uniformly 
3 mm. in length. 


The pelvic wing tract is not apparent in the adult pterylosis 
of Columba livia as given by Nitzsch.’ Its course is approximate- 
ly along the upper margin of the crural tract, and continuing 
toward the tail well into the posterior part of the femoral or 
lumbar pterylum. In fact, the remaining pterylae of the body 
are very indistinctly demarcated in the down of this young squab. 


*Pterylography, Nitzsch, Ed. by Sclater, 1867, plate VIT, fig. 2. 


44 Zoologica: N. Y. Zoological Society flies 


DOMESTIC PIGEONS. 
Columba livia Bonn. (var.) 


The pelvic alar tract is less regular in domestic pigeons than 
in wild birds, but is remarkably well developed. I give the re- 
sults of my examination of four squabs taken at random from 
a large number. 


A. The first was about a week old and the pelvic alar tract 
shows seven flights and four coverts. Always counting postero- 
anteriorly, the measurements in millimetres are as follows: 


Number of feathev........ peo et rel eee I) Qo 830 Ae 526 40-mmnl 
Wen gt ot stlifoints ee ee, GS tf th iy sep 2 
Ieenloth mote COVeTtS er ena eee ere ee A 1655s 3 


B. A squab two weeks old shows ten flights. The posterior 
four are uniform; they have well developed upper coverts, which 
are small and lie close above the main feather. From the 5th 
onward the coverts give place to a row of under coverts. As 
we go forward, the flights and their coverts become less closely 
associated, until only the slight difference in elevation of the 
two most anterior pairs reveals their true relationship. The 
most anterior flight is isolated and covertless. 


Number sOletedtner wc... ees I Que Sie A 5D 16) ieee Ot 
Weng thy of quppenyCOvertsi en. DOO mS. 

NGeRN ste Ok ee ELN ONES. tees nee en ee ee ace 6 7% 8:5 19) 7 16S NGtoosom 
Mength of under coverts pa... 222. 4 3.5 Ab 4.5" 4 


The precocious development of the feathers of this tract may 
be realized when compared with those of the true wing and tail 
in this same bird. The primaries and secondaries are all under 
five millimetres and the rectrices not more than two in length, 
while, as we have seen, seven out of ten of the pelvic flights are 
six millimetres or more, the general average over seven, and the 
maximum length nine millimetres. 


C. A squab about three weeks old shows twelve pelvic flights. 
The arrangement of coverts is as follows: 


Flight No. 1—small upper covert. 
~2—small upper covert. 

3—down covert. 

4—down covert. 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds 45 


5—small under covert. 
6—down under covert. 
7—large under covert. 
8—large under covert. 
9—large under covert. 
10—large under covert. 
11—no covert. 

12—no covert. 


D. A squab of flve weeks shows that no additions occur at 
the posterior end of the pelvic alar tract. The next contour 
feathers to appear at this point form an ascending series of three, 
parallel to the backbone and at right angles to the pelvic alar 
tract. The first four flights with their upper coverts are well 
grown, far ahead of the rest of the body plumage. The coverts 
indeed are quite full grown, downy and white-shafted. As in 
squab C the flights from the 5th onward have under coverts. Al- 
together there are nine flights with coverts, and three anterior 
covertless ones. 


While considering this newly observed character of pigeons, 
I thought of the feather-footed breeds and sent for a pair which 
I carefully dissected. I found no connection between these feath- 
er-footed and feather-legged domestic breeds and any unusual 
development of the pelvic alar tract. The feathers, which have 
been bred to great length, sprout from the scaly covering of the 
tarsus and phalanges and not from the leg proper or the femoral 
patagium, which is the seat of the character under consideration. 


JACANA. 
Jacana jacana (Linn.) 


In a half developed embryo the rectrices and pelvic alar 
feather papillae are well ahead of all others, even of the wing 
proper, and are the only ones which show any trace of pigment. 
In the pelvic alar tract there are four flights and three upper 
coverts, the anterior flight lacking a covert. In a second embryo, 
a day or two older, five flights and four coverts are visible in 
this tract. 


46 Zoologica: N. Y. Zoological Society pues 


GREAT HORNED OWL. 
Bubo virginianus (Gmelin) 


A brief examination of a living bird showed that the great 
development of soft plumage on the leg of this species arises 
from the pelvic alar tract. I was led to expect this from the 
pterylosis of Stria bubo, as given by Nitzsch.* 


In his figure of Columba livia’ there is, as I have said, no 
hint of the great development of the pelvic alar tract in the young 
bird, nor its remarkable disagreement with the lines of demar- 
cation of the pterylae of the adult. 


Judging merely from the pterylosis of the adult, many spe- 
cies of Coraciiformes, Scansores and Piciformes should show 
most interesting developments of this tract in the young birds. 


ARCHAEOPTERYX. 


The foregoing observations on various species of living birds 
were inaugurated and completed before I took up the question 
in regard to Archaeopteryx. I realized that any trace of this 
pelvic alar tract which might be present in this ancient bird 
would be of superlative interest and significance, but until I care- 
fully examined a full-sized photograph of the Berlin specimen 
I was not aware of the existence of feathers other than those on 
the wing and tail. I succeeded in finding distinct traces of 
strongly marked feathers on both sides of the tibia and of still 
larger feathers, lying between the pelvis and the bent back head. 
It seemed to me that such very evident traces could not have 
escaped the observation of other students of this wonderful fossil 
and I began a search of the literature. I was delighted to find 
that the tibial feathers had already aroused considerable dis- 
cussion and I present this in abstract to show how variously the 
scientific mind has reacted to evidence of this character, unsup- 
ported by any other more modern proof. The London Archaeop- 
teryx shows no trace of these feathers, so the whole evidence lies 
with the single fossil in the Berlin Museum. 


1Pterylography, 1867, plate II, fig. 9. 
“Nok, jollenie. VN aang 2 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds AG 


The bibliography of this discussion is a short one: 


Voer: 
“T?Archaeopteryx macrura.—Un intermédiaire entre les oiseaux et les rep- 
tiles.” Revue Scientifique, 1879, (2) LX, p. 245. 
Evans: 
“On portions of a Cranium and a Jaw of the Archaeopteryx, 
reprint, 1881, pp. 4-6. 


” 


Preface to 
Dames: 

“Ueber Archaeopteryx,” Paleontologischer Abhandlungen, 1884, II. pp. 39-41, 
ABEL: 

Grundziige der Palaeobiologie der Wirbeltiere, 1912, p. 343. 
HEILMANN: 

“Vor Nuvaerende Viden om Fuglenes Afstamning,” p. 14. Saertryk of Dansk 

Ornithologisk Forenings Tidsskrift. 

Vogt is the first to mention the feathers whose impressions 
are visible on the leg of Archaeopteryx. “Le tibia était couvert 
de plumes dans toute sa longueur. L’Archaeopteryx portait 
done des culottes, comme nos faucons, avec les jambes desquels 


sa jambe a le plus de resemblance, suivant M. Owen.” * 


J. Evans devotes several pages to these feathers and their 
significance, and a few years later Dames takes issue with him. 
The following sentences present Evans’ view: “ . along the 
outer margin of the right tibia, there is a series of eight or pos- 
sibly nine feathers of much the same character as those along 
the tail, and nearly, though not quite, of the same length.... 
Prof. Marsh has, indeed, already suggested that the power of 
flight probably originated among the small arboreal forms of 
reptilian birds, and has instanced the flight of Galeopithecus, the 
flying squirrels (Pteromys), the flying lizard (Draco) and the 
flying tree-frog (Rhacophorus) as indicative of how this may 
have commenced. Should it eventually prove to be the case that 
there were what may be termed supplementary wing-feathers on 
the hinder extremities of such early forms of birds as the Archae- 
opteryx, his views as to the origin of the powers of flight will 
be satisfactorily confirmed.” 


Under the heading “Das Federkleid,” Dames devotes con- 
siderable space to these tibial feathers. He admits them as 


*Quite unconnected with the present thesis, but interesting as a sentiment ex- 
pressed thirty-six years ago, the following paragraph by Vogt is worthy of note: 
“M. Volger se bercait dans l’espérance que S. M. ’empereur Guillaume achéterait 
la piece (Archaeopteryx) pour la conserver & l’Allemagne. Sa Majesté n’entra 
pas dans ces vues. Ah! si au lieu d’un oiseau, il s’etait agi d’un canon ou d’un 
fusil pétrifié!” 


48 Zoologica: N. Y. Zoological Society pee 


culottes or what we would call a booted feathering, but denies 
their function as assisting in flight. Without following his in- 
volved arguments, he says in part: 


“Zunachst ist es nicht richtig, dass diese Federn dieselbe 
Consistenz gehabt haben, wie die des Fliigels und des Schwanzes, 
denn sie sind weit undeutlicher erhalten, auch betrachtlich kiirzer 
(Lange durchschnittlich 32 mm. lang) als die Schwanzfedern, 
welche im mittleren Theil des Schwanzes ungefaéhr 65 mm. lang 
sind; .... sondern halb so lang .... Wenn man aber trotz alle 
dem doch noch an der Moglich keit, dass Archaeopteryx auch mit 
den Hinterbeinen geflogen sei, festhalten wollte, so ware dem 
noch entgegenzuhalten, dass diese EHigenschaft nothwendig auch 
irgend welchen Ausdruck im Bau der Knochen der Hinterex- 
tremititen erlangt haben miisste.”’ 


The two most recent commenters on this subject differ as 
completely as do Evans and Dames. Abel in his interesting sub- 
heading of “Die mangelhafte Ausbildung des Flugvermogens von 
Archaeopteryx,” writes: ‘Die zweizeilige Befiederung der Unter- 
schenkel spricht dafiir, dass diese Federn den Flug der Archaeop- 
teryx als Fallschirmapparate unterstiitzt haben.”’ 


Heilmann, writing in Danish, gives his opinion in an equally 
pithy phrase; “it is improbable that the feather coating on the 
tibia (as assumed by some authors) was of any importance in 
flight, as it appears too weak.’ ”’ 


PART III.-— ARGUMENT. 


The pelvic alar tract as I have found it in modern birds 
is remarkably uniform in position, originating on the anterior 
outer side of the tibia below the knee, and extending back, along 
the femoral patagium, to the body and toward the tail. The 
feathered patagium between the extended leg and the body must 
have been of the greatest importance, for the feathers sprouting 
in this area in the young bird are of very large size and invari- 


‘Altsaa rimeligvis paa Overgangen mellem Faldskaermssvaeven (som vi f. 
Kiks. traeffer den hos Flyveegern) og ubehjaelpsom Flagreflugt. 


A 
ce ye LS aS 
Ge ae 
((( ath » 


(( a r » 


yA r WN 
YAN 
YAN 


<Cecgy e re en ps 
Ku yy? ee ie coh SS 
ag) Oe poe 
Cea oe (Uys? 
|! { 
VN A 
3 ( i DFranklin # 
Wis 


FIG 11. THE EVOLUTION OF BIRDS FROM THE TETRAPTERYX STAGE (No. 1), 
THROUGH ARCHAEOPTERYX-LIKE STAGE (No. 2), TO THE MODERN BIRD (No. 4). 


The principal changes were the feathering and mittening of the fingers; the great 
strengthening and centralizing of the pectoral wing; the correlated reduction 
of the femoral or pelvic wing; the shortening of the tail and 


the concentration of the tail-feathers 
a 


Zoologica Vol. II, No. 2. Face page 48 


carn 


a 
ty Vise : 
— 
. at 
my vy 
4 ¥) 
a 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds 49 


ably provided with coverts. This is the pterylum which we hope 
to find paralleled or directly represented in Archaeopteryx. 


_ The most cursory examination of the fossil reveals the beau- 
tifully preserved wing and tail-feathers. Very faint and not at 
all certain traces have been thought by several observers to rep- 
resent a ruff of soft feathers at the base of the neck. We have 
already seen the diverse opinions which the two rows of tibial 
feathers have aroused. Besides these feathers there have been 
noted traces of small, soft, covert-like feathers covering the bases 
of the wing and tail feathers. The remainder of the body has, 
wholly without reason, been adjudged as scaly. On circumstan- 
tial evidence, but equally improbably, others have considered it 
as quite naked. The most reasonable hypothesis is that the body 
was fully clothed in soft, rather downy plumage. When the bird 
died, it fell upon the mud of some river or shore and there, like 
the remains of gulls which we may find today, it was slowly dis- 
integrated by the elements to a point where all the soft, body 
plumage was detached and washed or blown away. At the time 
of being imbedded in the fine silt it retained only the strongly 
socketed wing and tail feathers and those clinging to the hinder 
extremities. 


The most perfectly preserved part of the London specimen 
of Archaeopteryx is the tail. From base to tip it is almost with- 
out a flaw, and the relative length and position of the feathers 
are as distinctly seen as in the living bird. The outline of the 
tail as a whole is like that of a broad, truncated feather, tapering 
gradually to the base. I mention these details in order to com- 
pare them with those of the tail in the Berlin specimen of Arch- 
aeopteryx. Here the tip of the tail is lost, but the base is quite 
distinct. We can observe the same gradual narrowing, due to 
the increasing shortness of the feathers toward the pelvis. Be- 
tween the bent-back head and the pelvis, however, we see im- 
pressions of feathers which are longer than any at the base of 
the tail. Their origin is indefinite, somewhere near the pelvis or 
femur, and they arch up and backward as distinctly as many 
of the tail-feathers themselves. It seems reasonable to me that 
this group of feathers, which somewhat resembles a diminutive 


50 Zoologica: N. Y. Zoological Society [il 32 


wing, may represent the pelvic alar tract which is so remarkably 
developed in modern squabs. 


This character is plainly visible in any good photograph of 
the Berlin Archaeopteryx. Lankester’ shows it very distinctly 
in his reproduction of the fossil. As to the much-discussed tibial 
feathers, I agree with Evans and Abel that they seem too pro- 
nounced in outline to be classed with the downy feathers such 
as we see on our bocted falcons. I think they are the distal 
elements of the pelvic wing, of far less importance as a fallschir- 
mapparate than the larger feathers near the pelvis, which prob- 
ably arose from the femoral patagium. Most students of this 
bird have ignored these tibial feathers and in restorations they 
are usually omitted. Miss Woodward in her artistic plate’ shows 
them as soft and fluffy. 


Heilmann has approached the general subject of the origin 
of birds in a most delightful manner. His illustrations show real 
imagination, using that much abused word in its most admirable 
sense. Unfortunately his Danish text limits the possibility of 
wide appreciation. While, as I have shown, he does not believe 
that the tibial feathers were of volant function, yet, curiously 
enough, in his very original and dramatic restoration of Archae- 
opteryx,’ he has indicated a line of large feathers near the pelvis, 
which in position correspond to the inner feathers of my pelvic 
wing. 


The argument of Dames against the possibility of the hind 
leg functioning in aerial activity is at fault. It is naturally im- 
possible to conceive of a skilful flier, flapping with both arms 
and legs, and with ability for sustained and directive flight, to 
have evolved such a complicated dermal apparatus without cor- 
responding changes in muscles and bones. But in Archaeopteryx 
or in our prejurassic Tetrapteryx, the function of the pelvic 
wings would have been merely passive parachutes. In this early 
stage, as probably also in Archaeopteryx, flight was merely glid- 
ing or scaling. The fingers were too free, the arm bones too deli- 
cate, the sternum small or absent, and these facts considered in 


*Extinect Animals, 1905, p. 238. 
“Evolution in the Past, Knipe, 1912, p. 96. (II). 
“Vor Nuvaerende Viden om Fuglenes Abstamning, fig. 11. 


1915] Beebe: Tetrapteryx Stage in Ancestry of Birds 51 


connection with the small, weak pelvis, makes it impossible to 
picture the bird as flying skilfully about. 


In earlier, lizard-like, aboreal forms, the scale-anlagen of the 
wing feathers were correlated with corresponding developments 
along the hind leg, the two increasing equally in size and evolving 
feather lightness with change in structure. 


Even in Archaeopteryx, with its broad, excellent wing, the 
hand shows little or no correlated adaptation. The absence of 
two digits has probably no avian, or certainly no volant signifi- 
cance, for we find identical conditions in the manus of carniv- 
orous, bipedal but terrestrial dinosaurs, such as Ceratosaurus, 
Ornithormimus and Ornitholestes. 


If we admit Archaeopteryx to the direct line of lizard—Tet- 
rapteryx—hbird geneology, we must conceive of it as having 
reached a stage where the pectoral wing was becoming dominant, 
and beginning to afford support to the creature in general. The 
elongated flight feathers were now extending backward and 
superceding the passive function of the pelvic wing. With this 
concentration of motive and supporting power was soon to be 
correlated a shortening and reduction of the long unwieldy tail. 


In sueceeding generations the pelvic wings would become 
more and more reduced. Having arisen from among the sur- 
rounding scales, they had, for a time, volplaned through the air 
of early ages, a character passive and, as future centuries would 
show, of merely transitory function. Yet they were of tremen- 
dous importance in allowing the pectoral scales to develop, to 
become feathers, and then to assume an importance which was 
to make the class of birds supreme in the air. Yet the function 
of the pelvic wings had been so passive and negative that no 
special muscling had been necessary, no increase nor coalescence 
of bony tissue. Little by little the line of feathers and their 
coverts sank again into insignificance and became lost among the 
body plumage. It affords an excellent example of what Profes- 
sor Henry F. Osborn would call the phylogenetic accelleration of 
a character, followed by its gradual reduction. 


Millions of years after they were of use, the feathers of the 
pelvic wing are still reproduced in embryo and nestling. And 


52 Zoologica: N. Y. Zoological Socvety [II;2 


for some unknown reason, Nature makes each squab pass through 
this Tetrapteryx stage. The line of feathers along the leg of 
the young bird reproduces on this diminutive, useless scale the 
glory that once was theirs. No fossil bird of the ages prior to 
Archaeopteryx may come to light, but the memory of Tetrap- 
teryx lingers in every dove-cote. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


ou” 


VOLUME II, NUMBERS 3&4 


No. 3. NOTES ON THE BIRDS OF PARA, BRAZIL 


No. 4. FAUNA OF FOUR SQUARE FEET 
OF JUNGLE DEBRIS 


By 


C. WILLIAM BEEBE, 
CURATOR OF BIRDS 


SPU BLES H-E D BY THE SoG CT Bch 
THE ZOOLOGICAL PARK, NEW YORK 


Frepsruary, 1916 


New York Zoological Soriety 


General Office, 11 Wall St., New York City 
OFFICERS OF THE SOCIETY 


President 
HENRY FAIRFIELD OSBORN. - 


Hirst Vice-President Second Vice-President 
MADISON GRANT. FRANK K. STURGIS. 
Oreasurer Asst. Treasurer 
Percy R. PyNE, FARMERS’ LOAN & TRUST Co. 


Executive Committee 
MADISON GRANT, Chairman. 


Percy R. PYNE, LISPENARD STEWART, 
WILLIAM WHITE NILEs, WATSON B. DICKERMAN, 
WM. PIERSON HAMILTON, ANTHONY R. KUSER, 
FRANK K. STURGIS, ae HENRY FAIRFIELD OSBORN, 

; . “ex-of ficio. 


Enitorial Committee 
HENRY FAIRFIELD OSBORN, Chairman. 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


Auditing Committee 
? WILLIAM WHITE NILES, Chairman. 
H. CASIMIR DE RHAM, LISPENARD STEWART. 


General Officers 
Director of the Zoological Park: WILLIAM T. HORNADAY. 
Director of the Aquarium: CHARLES H. TOWNSEND. 
Prosector: Dr. GEORGE S. HUNTINGTON: 
Architect: C. GRANT LA FARGE. 
Consulting Engineer: H. DE B. PARSONS. 
Assistant Secretary: H. J. SHORTER. 
Bursar: R. L. CERERO. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBERS 3 & 4 


No. 3. NOTES ON THE BIRDS OF PARA, BRAZIL 


No. 4. FAUNA OF FOUR SQUARE FEET 
OF JUNGLE DEBRIS 


By 


C. WILLIAM BEEBE, 
CURATOR OF BIRDS 


Usb Ges A ED Bry, THE »/O°C Livery 
Pattee O Ono GLCAL PARK NEW- YORK 


FresBrRuARY, 1916 


= 


ne 


fies 


FIG. 12. THE CINNAMON TREE OF THE BIRDS 
The Utinga Jungle 


Zoologica Vol. II, No. 3. Face page 55 


Volume 2. Number 3. 


NOTES ON THE BIRDS OF PARA, BRAZIL 


By C. WILLIAM BEEBE, 
Curator of Birds. 


PART I.—INTRODUCTION. 


Belem or Para is a city of about two hundred thousand in- 
habitants. It is a hundred miles from the sea, on the south 
bank of the Amazon delta, and only one hundred and sixty kilo- 
metres south of the equator, built on low swampy land. The 
birds in the vicinity have been collected assiduously and offer 
little chance of novelty to the transient ornithologist. 


During the first part of May of the present year I had the 
opportunity of spending a little time in the jungle in the im- 
mediate suburbs of Para. Through the courtesy of the Gover- 
nor and of Dr. Snethlage I was given the use of a house at the 
water-works, in a large restricted area of jungle known as 
Utinga, and here every facility was afforded for collecting and 
study. Unexpectedly meeting Mr. George K. Cherrie, who had 
just come down the Amazon, I prevailed upon him to share our 
opportunities and with my companion Mr. G. Inness Hartley, 
spent a few days together. I found the region to be of much 
zreater interest than I had expected and by resorting to a rather 
novel method of observation I obtained a new angle upon life 
in these tropical lowlands, and one which profoundly impressed 
me as to future possibilities in this direction. 


PART II].—GENERAL ECOLOGY. 


The Utinga water-works consisted of a pumping station 
from which radiated long open cement water-ways and closed 
pipes leading straight through the jungle. The light jungle be- 
gan at the very edge of the small clearing which was within a 
few minutes’ walk of the tram line leading directly back to the 
heart of the city. 


5d 


56 Zoologica: N. Y. Zoological Society [liga 


It was without question quite the hottest, most humid 
tropical place I have ever encountered. I qualify with humid as 
I have known the dry heat of India to be much greater, as 110° 
F. at eleven P. M. at Agra. But this moist heat was in excess 
of any corresponding temperature I have known in Malaysia, 
Borneo, Mexico or elsewhere. It was the rainy season and the 
first day of our stay bore out the reputation of Para for precipi- 
tation, the rain pouring down much of the day. During all of 
the remainder of our stay, the weather was ideal, clear until 
about 2:30 P. M., when dark clouds and wind came up, the rain 
continuing until 4 P. M. On only one day it rained for twenty 
minutes in the morning, with the afternoon shower as usual. 
The nights were, of course, cool. 


Birds were most abundant from 8 to 10 A. M. and 2 to 5 
P. M. while at mid-day, all songs and chirps ceased and only 
the occasional note of an insect broke the stillness. 


Most of the birds had just passed the breeding season, and 
a goodly proportion of those secured were full-grown young. 
Both young and adults were molting or just completing the molt. 
In general they corresponded to our northern birds in August 
and September. A few, however, were preparing to nest and 
several were building. Blue tanagers had a nest a few yards 
from our house with two young which flew on May 8th. Yellow- 
backed caciques had several small colonies in isolated trees near 
native houses and were breeding. 


Much of the land between the small streams or igarapés 
was marshy and covered with an almost impenetrable cover of 
undergrowth. Occasionally a slight rise resulted in dry ground 
and here the growth became higher, more open and assumed 
the general character of almost primitive tropical jungle. A 
narrow trail opened into jungle of this character only a few 
yards from our house in the pumping-station clearing. It led 
straight northwards for about two hundred yards when it ended 
in open, overgrown fields. Along this trail the undergrowth was 
fairly dense, with here and there a giant buttressed tree, sur- 
rounded by lesser trees of many species. 


1916] Beebe: Birds of Parad, Brazil 57 


On the first tramp I took in the jungle I noticed a number 
of small birds in the upper branches of a tree which grew along- 
side of this trail. Not until I had passed that way several times 
did I come to realize that this particular tree had some powerful 
attraction for birds of many species. Knowing the shortness of 
time at my disposal I determined to concentrate my efforts on 
this tree which was a species of wild cinnamon. The presen‘ 
paper has to do chiefly with the facts thus obtained. 


Once having our attention called to this bird tree, Mr. 
Hartley and I kept on the watch for others. Several hundred 
yards away along a pipe line we discovered another. It was a 
real giant, towering high above all the surrounding growth and 
we named it the toucan tree as it appeared to be especially at- 
tractive to these birds. It was covered with an abundance of 
good-sized scarlet fruit, the size of which accounted for the 
presence of medium and large birds such as toucans, caciques, 
trogons and kiskadees, instead of smaller callistes and fly- 
catchers. A third berry-laden tree half a mile to the eastward 
straight through the jungle, bore oblong, yellow-skinned fruit 
appealing especially to woodpeckers and flycatchers, and from 
the brief glimpses we had as we passed, the constant abundance 
of birds would have furnished as interesting a list here as at the 
tree near our house. 


I began my study of’ bird-life in the wild cinnamon tree 
by stealthy approaches, working my way through the jungle 
until I was close underneath. I soon found that this was quite 
unnecessary, as the birds among the upper branches paid no at- 
tention either to me or the sound of my gun. Three hours of 
constant observation beneath the tree resulted in many hours 
of pain from strained neck muscles. On the third day I brought 
out a canvas steamer chair and placing it in the trail at a con- 
venient spot, found it to be ideal for observation. I could re- 
cline so that looking straight upward was no effort. With gun 
on my knees, glasses around my neck, note-book and dead birds 
on a stump within reach, I had discovered a truly de luwe method 
of tropical bird study. The biting flies, gnats and mosquitos 
made it impossible to sit absolutely quiet for more than a minute, 
and the ants soon found the legs of the chair gave easy access 


58 Zoologica: N. Y. Zoological Society [ike3 


to one’s person. On the whole, however, I was too much ab- 
sorbed in the novelty of the method of work and its unexpected 
results to give any thought to these annoyances. 


The principal jungle bloom was the heliconias, whose 
scarlet, jagged spikes glowed brightly against the dark foliage. 
Variegated leaves were abundant and when the slanting sun 
struck through the jungle, it often appeared vivid with color. 
The jungle about my seat was, of course, more or less impover- 
ished by the nearness of the city and the presence of the water- 
works. Black capuchin monkeys of more than one species were 
hereabouts and I saw as many as nine in a band. Three-toed 
sloths were common as were agoutis and small squirrels. But 
during my periods of watching no mammal came near the tree. 


The more frequent sounds were the common ones of light 
jungle, Tinamou called and answered one another, gold-birds 
lifted their wonderful voices far away in the forest, toucans 
yelped, caciques squeaked and gurgled overhead, cicadas 
shrilled and buzzed and great bees and hummingbirds whirred 
past. The commonest cicada had a note like a person calling a 
cat puss-puss-puss kept up interminably in a high soprano. An- 
other had a shrill, strident note which, when it gained full 
strength, quavered and broke into two alternating tones, which 
finally ran together into a true trill. After the daily rain, the 
tiniest of frogs would each strike up a single, shrill note, un- 
ceasingly reiterated. The most memorable sounds were the deep, 
gutteral voices of great frogs hidden in the igarapés, who re- 
iterated the never answered syllable, wh y? wh y? 


My business was chiefly with the birds which I could ob- 
serve from my canvas seat. I spent from two to six hours each 
day for a period of one week in the immediate vicinity of the 
tree and during that time identified ninety-seven species of birds, 
none of which were more than a few yards from the trail. A 
further division of these is as follows: 


Aerial species flying overhead W000... 7 
Birds; of the: surrounding jjuncle 14 
Birds observed in: then trees = ae eee 76 


a 
~. 


FIG. 13. 


FIG. 14. 


4 


Sea 


f. 


UTINGA JUNGLE FOREGROUND 


PARASOL ANTS ON THE MARCH 


Zoologica Vol. IT, No. 2. 


Face page 58 


baa 
ry 


SN a), 
€ - “ 
i. i 
: a hh 
rh, 


1916] Beebe: Birds of Parad, Brazil 59 


_ I shall reserve the details of the various species until later 
pages, and here give only a résumé of the more general points 
of interest. 


Of the seven aerial species, one was a vulture, one a night- 
hawk, one a swift and four were swallows. These all came 
into view at one time or another across the patch of sky visible 
beyond the upper branches of the tree. Now and then birds of 
prey appeared, but at such great elevations that I was unable 
to identify them. 


The fourteen birds of the surrounding jungle may be 
divided thus: one tinamou, dove, woodpecker, kingfisher, trogon, 
ani and woodhewer, two antbirds, two flycatchers and three 
finches. In one or two instances these were birds of adjoining 
fields which had strayed a little way into the undergrowth. The 
majority, however, were typical of the lower jungle strata, either 
terrestrial or living in the low undergrowth. 


This series of strata of bird life visible to me as | sat 
quietly hour after hour was very striking, a phenomenon which 
would never come to one while moving about through the jungle. 
Bound to the ground were the tinamou, and almost as terrestrial 
were the rustling ground doves. In the lower underbrush finches, 
Synallaxis and antbirds moved restlessly; a little higher mana- 
kins whirred about and woodhewers hitched up the trunks. Then 
came the birds of the upper branches, callistes, tanagers, fly- 
catchers, toucans and parrakeets. Then the low flyers—the 
swallows, martins, swifts and nighthawks, and finally the vul- 
tures, hanging like the faintest of motes in the sunlight high 
above the earth. 


PART II].—CANELLA Do MATTO AND ITs BIRD LIFE. 


The tree which I have already mentioned grew only about 
one hundred yards from our house at the pumping station and 
within five minutes’ walk of the Para tram line. It was at the 
side of a jungle trail, which, while seldom traversed by natives, 
was kept clear of vegetation by the workmen at the pumping 
station. It was smooth-barked, richly decorated with lichens 
and while only about fifteen inches in diameter at a man’s height 


60 Zoologica: N. Y. Zoological Society ites 


above the ground, it was very tall in proportion. The first 
branches were small, mostly dead and about fifty or sixty feet 
up. From this point the trunk split into lesser divisions and 
lifted its topmost foliage into the full tropical light and heat a 
hundred and ten feet above the ground. The berries were small, 
round and three-parted and, like the leaves, slightly acrid, with 
a spicy, aromatic flavor. 


A few minutes after dawn I have counted eight birds in the 
tree and a half dozen would sometimes linger until dusk. Asa 
rule, however, there were few in sight until 7:30 or 8:00 A. M., 
after which there would be a continual coming and going until 
the heat of mid-day drove all to shelter. The larger number of 
afternoon visitors came after the rain was over. Sunshine had 
much to do with the presence of the birds, and a cloudy half- 
hour meant but scanty notes as I sat beneath. With the reap- 
pearing of the sun, the birds would again begin to flock from 
the surrounding jungle. 


Abundance of species and relative fewness of individuals 
is a pronounced characteristic of any tropical fauna. This was 
beautifully shown by my first two days’ collection from the tree, 
collecting too, which was quite indiscriminate in character, very 
different from the more careful picking and choosing with which 
I shot on succeeding days. The first day I secured sixteen birds, 
all of different species. The second morning I got fourteen, all 
different, and only one of which was represented in the lot of 
the previous day. Thus in five hours’ time I secured thirty 
specimens of twenty-nine species. From the entire district of 
Para, three hundred and seventy-nine birds have been recorded. 
In this single tree within a week’s time and during a period of 
intermittent observation I found seventy-six species. 


The bird visitors to the tree arrived in one of two character- 
istic ways. Many came direct and swiftly, singly or in pairs, 
flying straight and with decision as if from a distance. A hun- 
dred yards away in any direction this convergence could fre- 
quently be observed, small birds flying over the summit of the 
jungle revealing a general flight direction treeward. Another 
method of arrival was wholly casual, loose flocks drifting slowly 
from the neighboring jungle, sifting into the tree and feeding 


FIG. 15. GIANT LAND SNAIL 


FIG. 16. NEST OF SAUBA ANTS 


Zoologica Vol. II, No. 3. Face page 60 


' ai Pa : onr'e ‘ “ ; : 
Fil 7 jhe ures ’ r : ) is A oe ii i 
ae aL ¥ M 7 iy 


ut 
ay 
we 


De % 
1a! e af 


oh 


1916] Beebe: Birds of Parad, Brazil 61 


for a time before passing on. When these left it was rather 
hastily and in answer to the chirps and calls of the members 
of their flock who had not been beguiled by the berries of this 
tree and hence had forged steadily ahead. 


These more or less well-defined flocks are very typical of 
all tropical jungles. Little assemblages of flycatchers, callis- 
tes, tanagers, antbirds, manakins, woodhewers and _ wood- 
peckers are drawn together by some intangible but very social 
instinet, and unite day after day in these fragile fraternities 
which drift along, gleaning from leaves, flowers, branches, 
trunks or ground, each bird according to its structure and way 
of life. They are so held together by an invisible gregarious in- 
stinct that day after day the same heterogeneous flock may be 
observed, identifiable by peculiarities of one or several of its 
members. The only recognizable bond is vocal—a constant low 
calling; half unconscious, absent-minded little signals which 
keep the members in touch with one another, spurring on the 
laggards, retarding the over-swift. 


At first I found it almost impossible to identify birds unless 
they were on the lower branches or silhouetted against patches 
of foliage. When in the upper branches and seen against the 
sky, birds with under-parts of black, blue or green all looked 
black. White under plumage appeared grey and buff seemed 
orange. Even when the tree was filled with the most brilliant 
callistes, not a bird was visible as long as they were motionless, 
but when the smallest, most drab of flycatchers moved head 
or tail I could at once detect it, and distinguish it from the mov- 
ing leaves about it. Gradually I came to know all the more 
common species, beginning with the tail-flirting silver-beak tan- 
agers, and before the end of my week’s vigil, I seldom made the 
mistake of shooting a species with which I was already familiar. 


While I watched, there came to my tree one species of 
pigeon, two hawks and two parrots, four hummingbirds and an 
equal number of toucans and woodpeckers. Fifty-nine were 
passerine birds of which there were eight each of the families 
of flyeatchers, manakins and cotingas, and eleven tanagers. 


62 Zoologica: N. Y. Zoological Society Bs} 


Besides the seventy-six species which I positively identified 
by shooting or observation, I saw at least thirty-three more 
which eluded me, and of which a hasty glance told no more than 
that they were of new, and to me, unknown species. 


The following is a list of the birds observed actually in the 
Canella do Matto tree. 


COLUMBIFORMES 1 


Splendid Pigeon, Columba speciosa Gmel. 


ACCIPITRIFORMES 2 


Brazilian Black Eagle, Urubitinga wrubitinga (Gmel.). 
Plumbeous Kite, Ictinia plumbea (Gmel). 


PSITTACIFORMES 2 


Tuipara Parrakeet, Brotogeris tuipara (Gmel.). 
Dusky Parrot, Pionus fuscus (Miill.). 


CORACIIFORMES 


TROCHILIDAE 4 

Red-vented Hermit, Phaethornis ruber ruber (Linn.). 

Great Jacobin Hummingbird, Florisuga mellivora mel- 
livora (Linn.). 

Amazonian Wood-Nymph, Thalurania furcata furca- 
toides Gould. 

Green-breasted Fairy, Heliothrix auriculata phaino- 
laema Gould. 


SCANSORES 


RAMPHASTIDAE 4 
Red-billed Toucan, Ramphastos monilis Mill. 
Double-collared Aracari, Pteroglossus bitorquatus bi- 
torquatus Vig. 
Lettered Aracari, Pteroglossus inscriptus inscriptus 
Swains. 


1916] Beebe: Birds of Para, Brazil 63 


Gould’s Toucanet, Selenidera maculirostris gould 
(Natt.). 


PICIFORMES 


PICIDAE 4 

Spix’s Amazonian Woodpecker, Celeus jwmana jumana 
(Spix). 

Waved Woodpecker, Celeus undatus multifasciatus 
(Mahl.). 

Malherbe’s Black Woodpecker, Campephilus trache- 
lopyrus (Malh.). 

Amazonian Gold-fronted Piculet, Picwmnus aurifrons 
Belz. 


PASSERIFORMES 59 


FORMICARIIDAE 1 


Sclater’s Amazonian Bush-Shrike, Thamnophilus ama- 
zonicus Sel. 


DENDROCOLAPTIDAE 7 

Whiskered Recurved-Bill, Xenops genibarbis genibar- 
bis Ill. 

Wedge-billed Woodhewer, Glyphorhynchus cuneatus 
cuneatus (Licht.). 

Eyton’s Fulvous-throated Woodhewer, Xiphorhynchus 
guttatus eytoni (Scl.). 

Chestnut-rumped Woodhewer, Xiphorhynchus parda- 
lotus (Vieill.). 

Picine Woodhewer, Dendroplex picus picus (Gmel.). 

Layard’s Woodhewer, Picolaptes layardi Scl. 

Buffon’s Barred Woodhewer, Dendrocolaptes certhia 
certhia (Bodd.). 


TYRANNIDAE 8 
Sulphury Flatbill, Rhynchocyclus sulphurescens 
(Spix). 


Zoologica: N. Y. Zoological Society piss: 


Sclater’s Flatbill, Rhynchocyclus poliocephalus sclateri 


Hellm. 

Oily Flycatcher, Mionectes oleagineus oleagineus 
(Hich#t:):. 

Sharp-billed Flycatcher, Tyranniscus acer (Scl. and 
God.). 


Yellow-vented Crested Flycatcher, Hlaenia flavogaster 
flavogaster (Thunb.). 


Gaimard’s Crested Flycatcher, Hlaenia gaimardii 
guianensis Berl. 


D’Orbigny’s Black-headed Flycatcher, Myiarchus tu- 
berculifer (Lafr. and D’Orb.). 


Azara’s Flycatcher, Empidonomus varius (Vieill.). 


PIPRIDAE 8 


Banded-tailed Manakin, Pipra fascticauda Hellm. 


Red-headed Manakin, Pipra erythrocephala rubro- 
capilla Temm. 


Slate-breasted Black Manakin, Pipra leucocilla bahiae 
Ridgw. 

Orange-bellied Manakin, Pipra suavissima Sol. and God. 

Para Opal-crowned Manakin, Pipra opalizans Pelz. 

Schomburgk’s Manakin, Piprites chlorion (Cab.). 

Blue-backed Manakin, Chiroxiphia pareola pareola 
(Linn.). 


Eastern White-breasted Manakin, Chiromachaeris 
manacus purus Bangs. 


COTINGIDAE 8 
Cayenne Tityra, Tityra cayana (Linn.). 
Red-cheeked Tityra, Tityra inquisitor erythrogenys 
(Selby). 
Little Psaris, Platypsaris minor (Less.). 
Cinereous Thickbill, Pachyrhamphus rufus (Bodd.). 


1916] Beebe: Birds of Parad, Brazil 65 


Lichtenstein’s Thickbill, Pachyrhamphus marginatus 
(Licht.). 


Gold Bird, Lathria cinerea (Vieill.). 
Schomburgk’s Attila, Attila brasiliensis (Less.). 
Cayenne Chatterer, Cotinga cayana (Linn.). 


TROGLODYTIDAE 2 
Swainson’s Moustached Wren, Thryothorus genibarbis 
genibarbis Swains. 


Venezuelan House Wren, Troglodytes musculus clarus 
Berl. and Hart. 


TURDIDAE 1 


Cabanis’s White-throated Thrush, Planesticus phaeopy- 
gus phaeopygus (Cab.). 


VIREONIDAE 3 
Chivi Vireo, Vireo chivi (Vieill.). 
Grey-naped Wood Vireo, Pachysylvia thoracica semici- 
nerea (Sel. and Sal.). 


Guiana Vireo-Shrike, Cyclarhis gujanensis gujanensis 
(Gmel.). 


FRINGILLIDAE 2 
Rothschild’s Blue Grosbeak, Cyanocompsa rothschildt 
(Bartl.). 
Great Saltator, Saltator maximus (Miill.). 


COEREBIDAE 5 
Brazilian Flowerpecker, Coereba chloropyga chloropyga 
(Cab.). 
Turquoise Honey-Creeper, Dacnis cayana cayana 
(Linn.). 
Black-backed Honey-Creeper, Dacnis angelica angelica 
Bonap. 


66 Zoologica: N. Y. Zoological Society [ies 


Blue Honey-Creeper, Cyanerpes cyaneus cyaneus 
(Linn.). 

Green MHoney-Creeper, Chlorophanes spiza spiza 
(Linn.). 


TANAGRIDAE 11 


Blue-backed Green Tanager, Chlorophonia chlorocapilla 
(Shaw). 


Northern Violet Euphonia, Tanagra violacea lichten- 
steinu (Cab.). 


Cayenne Euphonia, Tanagra cayennensis (Gmel.). 


Para Blue-bellied Tanager, Tanagrella velia signata 
Hellm. 


Spotted Tanager, Tangara punctata punctata (Linn.). 


White-shouldered Blue Tanager, Thrauwpis episcopus 
episcopus (Linn.). 


Palm Tanager, Thraupis palmarum palmarum (Wied.). 


Silver-beaked Tanager, Ramphocelus carbo carbo 
(Pall.). 


Scarlet-crested Tanager, Tachyphonus cristatus brun- 
neus (Spix). 


Para Crested Tanager, Tachyphonus surinamus insig- 
nis Hellm. 


Guira Tanager, Hemithraupis guira guira (Linn.). 


ICTERIDAE 3 
Great Green Cacipue, Ostinops viridis (Miill.). 
Yellow-backed Cacique, Cacicus cela (Linn.). 


Brazilian Red-rumped Cacique, Cacicus haemorrhous 
haemorrhous (Linn.). 


The great abundance of birds in this particular tree was 
due, of course, to the multitude of ripe berries among its foliage. 
These were the primary cause of attraction. Lacking these, the 


1916] Beebe: Birds of Parad, Brazil 67 


birds would have had no special reason for visiting it more than 
the surrounding jungle. And it was surprising to discover how 
many of the birds which would usually be considered as fly- 
catching or insect eaters, had in this case turned frugivorous. 
It seems worth while to reclassify this arboreal avifauna by the 
raison d’étre of their presence. 


Feeding on Tanagers, 11 Casual Visitors 

tree berries Caciques, 3 ee 
ee ; : Snail-eater Parrot, 1 
-arrakeet, : ; : , 
Toucanets. 3 Hawk, 1 umnninebizds, 4 
Woodpecker, 1 Insect-eaters Toucan, 1 
Flycatchers, 8 of the trunk Woodpecker, 1 
Manakins, 6 Woodpeckers, 2 Bush-Shrike, 1 
ae 7 Woodhewers, 7 Manakins, 2 
Vireos, 3 Insect-eater of Cotinga, 1 
Finch, 1 the branches Wren, 1 
Honey-Creepers, 5 Wren, 1 Finch, 1 


The greedy, noisy parrakeets were restless jungle birds, 
shifting from one feeding place to another, always gorging them- 
selves, tearing off bunches of berries and wasting much more 
than they ate. Of the members of the Ramphastidae, the visitors 
to this tree were almost wholly toucanets, the smaller, more agile 
species which found less trouble perching on the rather slender 
branches. The toucan tree a few hundred yards away, hung 
its larger fruit on stouter branches and attracted the toucans of 
larger size. 


Without exception all the flycatchers which I observed in 
the tree—eight species—were feeding on the berries, in spite of 
their wide gapes and insect-guiding bristles. This was not so 
surprising in the case of the six manakins and seven cotingas, 
but the three vireos and five honey-creepers must have been 
birds of originality to turn thus wholly frugivorous. The tana- 
gers led all in numbers, eleven of them, and were feeding exclu- 
sively on the berries, and the same was true of the three caciques. 


68 Zoologica: N. Y. Zoological Society [iks3 


On the casual visitors it is unnecessary to remark. A wren 
hunted insects among the upper branches one day, and on an- 
other a hawk found a giant snail crawling up the trunk and 
preceeded to devour it. 


The insect-eaters of the trunk were nine in number and 
showed no interest in the berry harvest. Two were woodpeck- 
ers and there were seven species of that interesting tropical 
family of woodhewers. These birds were abundant at Utinga. 
Their labor was confined to a careful search for insects on the 
trunk and larger branches. The smaller woodhewers such as 
Xenops and Glyphorhynchus usually drifted to the tree as mem- 
bers of the loose jungle flocks. The larger woodhewers were 
more independent, and usually seen singly or in pairs. The low, 
plaintive notes of the little wedge-billed woodhewer were typi- 
cally like those of the loose flocks, keeping the members in touch 
with one another. 


Woodhewers are the very essence of protective coloring, and 
their habits of life make of them wandering bits of loose bark, 
yet because of their constant motion, they are very easy to see 
even in the dim light of the under jungle. The moment they 
are quiet they vanish, and the keenest eye in the world could not 
recognize them. This similarity of dress is a remarkable feature 
of this whole family; big and little, short and long-tailed, with 
beaks blunt, sharp, straight, curved, thick or needle-thin. In 
these characters they differ, by these points they must know 
one another. But their pattern shows little variation. Their 
olives or browns almost invariably warm into rich foxy rufous 
on wings and tail, while over head and shoulders a shower of 
light streaks has fallen, bits of sunlight fixed in down. 


And so came to a close my rambling observations on the 
bird life of this single Canella do Matto. Within the space of 
a week I had spent not more than twenty hours of neck-racked, 
vertical observation, shooting whenever necessary, holding up 
my glasses until my arms collapsed with fatigue. In return I 
had been able definitely to identify seventy-six species and to 
record the presence actually in the tree of at least one hundred. 


1916] Beebe: Birds of Para, Brazil 69 


In point of actual numbers I kept no sustained record, but dur- 
ing one vigil of two hours’ length I counted four hundred and 
sixteen birds in the tree. 


When I began I had no conception of such success and as 
I look back and realize the necessary desultory character of my 
observations, the list seems even more remarkable. Relay ob- 
servation on the part of two or three watchers for a correspond- 
ingly greater length of time, or closer observation from a blind 
fixed in a nearby tree, would yield notes of incomparably greater 
thoroughness and value. 


PART IV.—NOTES ON SOME INVERTEBRATES NEAR THE 
CANELLA Do MATTO. 


I made no effort, during the short time at my disposal, to 
carry on any lines of observation, other than upon the avifauna 
of the one tree. Yet as I walked back and forth along the trail, 
or sat quietly during the rather rare periods when no birds were 
in sight, or rambled about in the surrounding jungle and along 
the overgrown igarapés, I made a few desultory notes on certain 
invertebrates of especial interest to me as forming the food of 
jungle birds. 


The great land snail, Strophocheilus oblongus Miller, we 
saw now and then, partly hidden in crevices of bark, and early 
one morning I saw a plumbeous kite in the canella tree, hold- 
ing the shell of one of these mollusks in his talons and devouring 
the inmate. The shells were strongly grained, and of a rich 
brown with salmon-colored mouth. An ordinary sized shell was 
about four or five inches in length, and when the mollusk was 
fully extended the whole organism reached seven and a half 
inches. On a tree-trunk leaning over an igarapé I counted four- 
teen of these mollusks crowded into one very shallow cavity. 


I observed that spiders entered largely into the diet of the 
birds I examined and I was interested in watching the method 
of escape of several common species, whose webs were hung along 
the trail. 


70 Zoologica: N. Y. Zoological Society i Bea 


Acrosoma spinosa Linn., an exceedingly spiny, gaudy spider 
hung in the center of its web. Its scarlet, yellow and black color- 
ing seemed to indicate an unsavory mouthful, and it was corre- 
spondingly slow to take alarm. Its large, round web was swung 
obliquely within a foot or two of the ground. At the center was 
a heart-shaped open space in which the spider hung by six legs, 
the other two being drawn back ready for action. The web 
slanted backward and the spider hung upside down, the brilliant 
colors of the upper side of the body being thus completely hidden. 
When the creature was alarmed, it dropped to the ground along 
a cable which it attached to the point of the heart-shaped space 
and paid out as it fell. The moment it touched land, it slipped 
under a leaf. If no further disturbance ensued it regained its 
courage in about three minutes, and climbed swiftly, winding 
in its cable and apparently swallowing it, as it went. When 
caught in the hand, it turned at once upon its back and feigned 
death. 


A mottled, rectangular, rather flattened and much more 
toothsome appearing spider was Hpeira audax Blk. Its lure 
was usually hung under a stump or a fallen sapling. When dis- 
turbed it invariably ran upward from the center of the web to 
the trunk, where it drew in its legs and squatted. In four in- 
stances its resting place was a bit of mossy or lichened bark, and 
although in full view, it merged perfectly with its surroundings. 
So perfectly, indeed, that the eye had to search carefully to redis- 
cover it each time it sprinted to safety. 


Epieira truncata Keys, was a smallish black spider, with yel- 
lowish-white markings on its back. It had still a third place of 
concealment. Wherever its web was hung, there was always 
some convenient leaf which the spider had half rolled up, tied 
fast with web and lined with silk. At the first sign of danger or 
when heavy rain fell, the architect rushed from the center of 
the web to the prepared sanctuary. 


The commonest spider at Utinga, fat, round, black and be- 
loved by birds was Hriophora purpurascens. Unlike all the others 
its point of vantage was not at the center of its web but in a 


1916] Beebe: Birds of Parad, Brazil (sk 


specially prepared den. The web was invariably hung between 
the leaves of some shrub. At one side, usually above the web 
and in full view, three leaves were drawn loosely together and 
fastened. Between these the spider waited for tell-tale web vibra- 
tions, and in such places inquisitive antbirds and jungle wrens 
found and devoured it. 


One day a short distance from the tree I watched an indeci- 
sive bout between one of these spiders and a small but courage- 
ous wasp. The contest must have been going on for some time 
as about half the web was already destroyed. The spider had 
left its den and was clinging to the center of the slack structure. 
The wasp was exerting every effort to destroy the remaining 
two or three chief supporting cables. She would alight and chew 
them with all her might. After a few futile attempts, buzzing 
with rage, she would fly at the strand, seize it in her mandibles, 
and darting backward in midair, endeavor to snap it. Then she 
alighted on a nearby leaf and carefully cleaned feet, wings and 
head. 


After such a rest she would turn her attention to the spider 
itself, buzzing around as closely as she dared, and making sud- 
den rear attacks. 


Eriophora was never off guard for a moment and raising his 
grasping feet he offered an invincible front. As the wasp was 
only a fourth of the size of the spider she dared do nothing more 
in the line of direct attack. It appeared that all her efforts were 
directed to cutting the spider down to the ground when she 
could probably have mastered him. He evidently did not dare 
to attempt to reach his leaf shelter, and remained quiet, guard- 
ing against attacks, swaying in his half demolished web. Before 
any dramatic crisis could develop, a heavy downpour of rain 
came on and drove both creatures to shelter. 


Caterpillars were abundant at this season and remains of 
them were found in the stomachs of about one bird in every 
three. The most noticeable, however, were too well-armed to 
fear sudden death at the beaks of birds. One appeared on the 


be Zoologica: N. Y. Zoological Society pees 


smooth bark of Miconia, like a great felted mass of long reddish 
hairs, each of which was a veritable barbed and stinging nettle. 
This larva has never been reared to maturity, but it is supposed 
to belong to the Limacodidae. These caterpillars climbed slowly 
up the trunks, making about ten feet an hour. 


Another bizarre larva spent the day hidden on the under 
side of a banana frond, close to the midrib. It looked like a short, 
thick, arrow, notched posteriorly, with a rounded, blunt head 
fringed behind with a row of great spikes. The imago is the 
moth Opsiphanes invires. 


A sphingine snake-head caterpillar of the genus Macloryx 
was seen once. It is unquestionably one of the most startling 
dénouements in nature to see this large, smooth, innocuous look- 
ing larva suddenly bend its head forward and down, and trans- 
form into a vivid representation of a serpent’s head, even to the 
rapidly playing forked tongue. 


The omnipresent saiiba ants (Atta sp.) forced themselves 
on the attention of the most casual observer. All day long their 
interminable lines flowed back and forth from tree-tops to nest, 
conveying myriads of green leaf burdens. The single point which 
impressed itself upon me was the large number of ants getting 
free transportation. Every other leaf had from one to six ants 
of small size clinging to the swaying frond. Where the leaf was 
pliable and of large size they had all they could do to maintain 
their position as it was jerked along. These were doubtless some 
small form of the saiiba citizenry but why the free transporta- 
tion and what their function was I could not determine. 


One of the most remarkable invertebrates which I observed 
was an aquatic hairy caterpillar. This was found in abundance 
in shallow pools and creeks. The first one which I saw seemed 
to be wriggling about in the throes of drowning, having, as I 
supposed, fallen from the overhanging foliage. I charitably 
scooped it out and set it to dry on a bit of palm leaf. It attempted 
to walk away, but in spite of the fact that much of its hairy coat 
dried at once, it staggered about, toppling over at each step and 


1916] Beebe: Birds of Parad, Brazil 73 


appearing more at ease squirming about on its side. Some dis- 
tance further on I saw a dozen more in an open pool and then, 
realizing my mistaken kindness, went back and restored the cater- 
pillar to its strange element. It seems that this is the larva of a 
small moth appropriately named Palustra, which has assumed 
an aquatic life. It swims by vigorous wriggles and uncoilings, and 
occasionally, like a mosquito larva, comes to the surface. It is 
not known, however, whether it breathes directly from the sur- 
face, or from the air entangled in its hairy coat. 


PART V.—NOTES ON THE MOLT OF SOME PARA BIRDS. 


My recent study of the molt, and especially of the tail molt, 
of pheasants has seemed to yield something of value in dividing 
these birds into subfamilies.‘ While disclaiming any preconceived 
belief in the use of this character in other groups, I have never- 
theless lost no opportunity to record whatever data I could find 
in regard to this phenomenon. I intend as rapidly as possible 
to examine molting birds of all orders and to place the results 
on record. With this in view I present the facts derived from 
sixteen species which I examined at Utinga, near Para, in Brazil 
in the early part of May. Fragmentary as they are, they show 
nevertheless that differences exist. Whether these, in some cases, 
are of only specific distinction, or whether of generic or family 
value, only future, more extensive investigations can prove. 


As regards wing molt, I found only two exceptions to the 
regular formula of the primaries molting regularly and succes- 
sively from within out, and the secondaries molting from the 
outermost inward. In the cotinga, Platypsaris minor, the pri- 
mary molt appeared to be 1-2-3-4-10-5-6-7-8-9. The secondaries 
nad two modes of molt. From the outer to the 4th pair; then 
from the 5th pair inward and the 12th pair outward, meeting 
about the 8th pair. 


A specimen of Dacnis cayana cayana showed a similar break 
in the secondary molt, molting in both wings inward from the 
outer pair, and outward from the 9th pair, meeting at the 5th 
or 6th pairs. 


"Zoologica. Vol. I, No. 15, p. 265. 


74 Zoologica: N. Y. Zoological Society Livers 


In attempting to work out tail molt from dried skins in the 
Museu Goeldi I was impressed with the difficulty of accurate 
observation. It is almost impossible to examine thoroughly the 
entire individual rectrices without damaging the appearance of 
the skin, and the dried sheaths which are so often the sole clue 
to recent growth, crumble at the first touch of the pliers. 


To summarize at once my data taken from fresh, unskinned 
birds, I record the following types of tail molt: 


Centripetal, from the outside in, 


Ramphastidae (3) 
Picidae (1) 


Centrifugal, from the center out, 
Dendrocolaptidae (2) 


Vireonidae (1) 
Tanagridae (3) 


Other types of tail molt, 
2 
Pe ear . . 
3 1 4-5-6 Pipridae (2) 
3— 1-2-4-5-6 Cotingidae (1) 


i 24 


<gs Coerebidae (2) 


Ramphastos osculans Ramphastidae. 


Two individuals collected from the same flock, May 9, were in 
almost the same stage of tail molt. 


Ten rectrices. Moit from outside in. 

Specimen A. Central, 2nd and 3rd pairs, old, unshed. 
Ath pair, blood sheath of 28 mm. 
5th pair, growing 98 mm. 


Specimen B. Central, 2nd and 3rd pairs, old, unshed. 
Ath pair, growing 59 mm. 
5th pair, growing 106 mm. 


1916] Beebe: Birds of Parad, Brazil (ss 


In both birds the mode of molt of the primaries traveling 
outward had reached the 5th pair. That of the secondaries mov- 
ing inward, had caused the renewal of eight feathers. 


Pteroglossus imscriptus Ramphastidae. 


Bird collected May 6th. 
Ten rectrices. Molt from the outside in. 
Central pair, just shed. 
2nd pair, one-half grown, 44 mm. 
3rd pair, growing, 84 mm. 
4th and 5th pairs, new, full-grown. 


Selenidera gouldit Ramphastidae. 


Birds shot May Ist. 
Ten rectrices. Molt from the outside in. 
Central and 2nd pairs, old, unshed. 
3rd rectrice (left), just shed. 
3rd _ rectrice (right), blood sheath, 
4 mm. 
4th and 5th pairs, new, full-grown. 


Celeus undatus Picidae. 


Bird shot May 6th. 
Twelve rectrices, ten functional, and an outer vestigial pair 20 
mm. in length. Molt from the outside in. 
Central and 2nd pairs, old, unshed. 
3rd pair, blood sheath just appearing. 
4th pair, growing, 36 mm. 
5th pair, almost full-grown. 
6th pair, full-grown. 


Dendrocolaptes certhia Dendrocolaptidae. 


Bird collected May 8th. 
Twelve rectrices. Molt from the center out. 
Central pair, new, full-grown. 
2nd pair, nearly grown 96 mm. 
3rd pair, blood sheath, 16 mm. 
Ath, 5th and 6th pairs, old, unshed. 


76 Zoologica: N. Y. Zoological Society [Tiss 


Picolaptes layardt Dendrocolaptidae. 


Bird collected May 6th. 


Twelve rectrices. Molt from the center out. 
Central, 2nd and 3rd pairs, new, full-grown. 
4th pair, one-half grown. 
5th pair, blood sheath, 19 mm. 
6th pair, old, unsheath. 


Pipra leucocilla Pipridae. 


Bird collected May 3rd. 


Twelve rectrices. Molt nearly complete; probably like that of 
the following species. 
Central, 2nd and 3rd pairs, new, full-grown 
4th pair, nearly full-grown. 
5th pair, one-half grown, 18 mm. 
6th pair, one-third grown, 8mm. 


Pipra opalizans Pipridae. 

Twe individuals collected on May 8th and 9th. 

Twelve rectrices. Molt about the same stage in both. The 
second and third pairs are shed first and simultaneously ; 
next the central, and then in succession the 4th, 5th and 
6th pairs. This unexpected type of molt received confirma- 
tion from the fact of its occurrence in two individuals shot 
on successive days, in different parts of the Utinga jungle. 


Specimen A. Juvenile male (Fig. 17). 


FIG, 17. TAIL OF MANAKIN 


All twelve rectrices were blood sheaths, only the central, 2nd 
and 3rd pairs having broken through. 


1916] Beebe: Birds of Parad, Brazil 77 


Central pair, 9mm. 
2nd pair, 11 mm. 
3rd pair, 11 mm. 
Ath pair, 6mm. 
5th pair, 4mm. 
6th pair, 2mm. 


Both wings were exactly alike. 


Primaries molting outward; 1st to 5th pairs new. 
6th pair nearly grown. 
7th pair three-quarters grown. 
8th pair blood sheath, 7 mm. 
9th and 10th pairs, old, unshed. 


Secondaries molting inward; outer pair nearly grown. 
2nd pair, blood sheath, 11 mm. 
3rd pair, etc., old, unshed. 


Specimen B. Adult male. 


All twelve rectrices were blood sheaths, just breaking through. 
The sheaths all averaged 6 mm. in length. 


Total Length 

Right Left 
11 mm.—1l1st —11 mm. 
14 mm.—2nd—15 mm. 
13 mm.—srd —14 mm. 
11 mm.—4th —12 mm. 

7 mm.—5th— 6.5 mm. 

7 mm.—6bth — 8 mm. 


In these specimens the very specialized, opalescent crest 
feathers were in full molt, almost all of them ensheathed. These 
sheaths were slender, conical, pointed and lightly fluted. The 
general appearance of the ensheathed crown feathers was of a 
mass of obliquely lying, parasitic cocoons on a caterpillar. 


Platypsaris minor Cotingidae. 


Bird collected May 8th. 
‘Twelve rectrices, all old, unshed. 


78 Zoologica: N. Y. Zoological Society [ihe 


Wings in full molt, both wings the same. Old feathers rufous 
buff; new ones black, with white basal spots. 


Primary molt 1-2-3-4-10-5-6-7-8-9. 
Secendary molt, outer to 4th pair. 


5th pair inward and 12th pair outward, 
meeting about the 8th. 


Primaries, Inner, 2nd, 3rd and 4th pairs, new, full-grown. 

5th pair, nearly full-grown 
54mm. 

6th pair, just breaking sheath, 
20 mm. 

7th, 8th and 9th pairs, old rufous 
feathers. 

10th pair, new, full-grown. 


Secondaries, outer pair, nearly grown, 52 mm. 
2nd pair, short sheath, 4mm. 
3rd and 4th pairs, old, rufous feathers. 
5th, 6th and 7th pairs, new, full-grown. 
8th pairs, short sheath, 8 mm. 
9th pair, etc., new, full-grown. 


Attila brasiliensis Cotingidae. 


Bird collected May Ist, juvenile. 
Twelve rectrices, molt apparently 3-1-2-4-5-6. Old feathers, 
worn, brown almost rufescent; new ones brownish black. 


3rd pair, growing, 40 mm. 

Ist pair, growing, 21 mm. 

2nd pair, blood sheath, 8 mm. 

Ath, 5th and 6th pairs, old, unshed. 


Primaries molting outward, six pairs renewed. 

Secondaries show no molt. 

Cyclarhis gujaranensis Vireonidae. 
Specimen A. Bird collected May 1st, female. 


Twelve rectrices. Molt from the center out. 


1916] Beebe: Birds of Parad, Brazil 19 


Outer four pairs, old, unshed. 


Central pair, new full-grown. 
2nd pair, growing. 


No wing molt. 


Specimen B. Bird collected May 1st, juvenile, female. 


Twelve rectrices. All old except central pair which are nearly 
grown. 


Dacnis cayana cayana Coerebidae. 
Specimen A. Bird collected May 3rd, juvenile, female. 

: 2-4 
Ten rectrices. Molt 1 G = 


Central pair, new, full-grown, 36 mm. 
2nd pair \ 
3rd pair ( 
Ath pair ) 
5th pair f 


growing, 27 mm. 


blood sheath, 11 mm. 


Specimen B. Bird collected May 5th, adult, male. 


Tai! has almost completed molt, outer pairs being nearly full- 
grown. 


Wing molt three-quarters complete, showing an interesting and 
unusual type of secondary molt. Old feathers edged with 
green; new ones with blue. 


Left Wing Right Wing 
987654321 1234 5 6789, ete. 987 6 543821 123456789 
<—— & BB <= »-> <a »— 
primaries secondaries secondaries primaries 


Secondaries, right wing, outer, 2d, 3d and 4th, new, full-grown. 
5th, blood sheath. 
6th, old, unshed. 
7th, still growing. 
8th and 9th, new, full-grown. 


80 Zoologica: N. Y. Zoological Society PUES; 


left wing, outer, 2nd, 3rd and 4th, new, full- 
grown. 
5th, old, unshed. 
6th, blood sheath, 7 mm. 
7th, blood sheath, 17 mm. 
8th and 9th, new, full-grown. 


Chlorophanes spiza Coerebidae. 
Bird collected May 5th. 


Ten rectrices. Molt 1 aces oe 


Central pair new, full-grown. 
2nd pair 
ard pair yl grown 


Ath pair | 


Bth pair f° unshed. 


Thraupis episcopus episcopus Tanagridae. 


Specimen A. Bird collected May 9th. Fledgling, male, first day 
after leaving nest. 


Twelve rectrices well grown, and apparently of equal length. 
Measurements 


Fledgling Adult 


19 mm.—Central— 64 mm. 
23mm.— 2nd —65 mm. 
25 mm.— 8rd —65 mm. 
25mm.— 4th —66 mm. 
23 mm.— 5th —66 mm. 
21 mm.— 6th —65 mm. 


Specimen B. Bird collected May 1st, male. 


Twelve rectrices. Molt from the center out. The whole web of 
the new feather is blue, stronger on the outer web. Old 
feathers are black on the inner web, greenish on the outer. 


1916] Beebe: Birds of Parad, Brazil 81 


Central pair, new, full-grown. 


2nd pair, still growing, 6mm. shorter than 1st. 
3rd pair, unbroken blood sheaths. 

Ath (right), sheath just appearing. 

Ath (left), not yet shed. 

5th and 6th pairs, old, unshed. 


Ramphocelus carbo carbo Tanagridae. 
Specimen A. Bird collected May Ist, male. 


Twelve rectrices. Molt from the center out. 
Central and 2nd pairs, full-grown. 
3rd and 4th pairs, just drying up. 
5th and 6th pairs, not quite full-grown. 


Specimen B. Bird collected May 5th, adult male. 


Tail in full molt, from the center out. 
Central and 2nd pairs, new, full-grown. 
3rd, 4th, 5th and 6th pairs, all with 18 mm. 
sheaths, but total length steeply grad- 
uated. 
Primaries molting outward, two outer pairs still growing. 


Secondaries molting inward, three outer pairs full-grown, next 
three in active growth. 


Tachyphonus surinamus insignis Tanagridae. 
Specimen A. Bird collected May 2nd. 


Twelve rectrices in full molt from the center out, with unusually 
long time hiatus between the central and 2nd pairs. 
Central pair, new, full-grown, 72 mm. 

Outer five pairs all with 11 mm. sheaths. 
Total lengths 2nd pair, 69 mm. 
3rd pair, 54mm. 
4th pair, 40 mm. 
5th pair, 31 mm. 
6th pair, 25 mm. 


82 Zoologica: N. Y. Zoological Society BGs 


Wing molt nearly complete; primaries outward, secondaries 
inward. 


Specimen B. Bird collected May 5th. 
Tail completing molt from center out. 


PART VI.—ANNOTATED LIST OF BIRDS OBSERVED. 
A. BIRDS OF THE WILD CINNAMON TREE. 
Columba speciosa (Gmel.) SPLENDID PIGEON. 


Three were observed on May 2nd, one in the tree feeding 
on the berries, the others on adjoining branches. They flew at 
once when I walked past beneath. 


Urubitinga urubitinga (Gmel.). BRAZILIAN BLACK EAGLE. 


Twice in the same day this bird visited the trail near the 
tree, once perching rather low in the jungle and remaining mo- 
tionless. An hour later it returned and alighted on one of the 
Jower branches of the tree itself, preening its feathers and pay- 
ing no attention to the small birds scolding from the shelter 
of the thick foliage to which they had fled. A specimen secured 
had a large green, blue, red and yellow mantis with a hundred 
or more of its eggs in his crop. 


Ictinia plumbea (Gmel.). PLUMBEOUS KITE. 


Early on May 11th at a time when there were only three 
or four small tanagers in the tree, this bird suddenly appeared. 
IT had stopped watching for a few minutes to rest my fatigued 
muscles, and on looking up I saw this hawk perched in the tree 
on a branch, so slender that it was still swaying from the im- 
pact of his alighting. He seemed to be picking at something on 
the branch beside him, but flew at once when I fired, apparently 
quite uninjured by the small shot which I had to use. I then 
found that he had been devouring a snail of large size in its 
shell (Strophocheilus oblongus). 


1916] Beebe: Birds of Pard, Brazil 83 


Brotogeris tuipara (Gmel.). TUIPARA PARRAKEET. 


Quite common in families or small flocks. Twice observed 
mn the tree feeding on the berries, and one which I secured had 
twenty-three in its crop. The noisiest birds hereabouts. While 
sitting at the foot of the tree, half an hour would seldom pass 
without a pair or more of these parrakeets dashing past high 
overhead, screeching loudly. Other trees seemed to offer more 
permanent attraction than this one. They showed little fear 
and members of their flocks could be shot one after the other 
without frightening the remainder. In the evening they col- 
lected in flocks of thirty or forty and circled about high in the 
air before setting off steadily south-westward toward some dis- 
tant roost. 


Pionus fuscus (Miill.). DUSKY PARROT. 


A pair alighted in the tree on May 4th and remained for 
five minutes before flying off in the direction of the toucan tree. 
I heard them now and then in other parts of the jungle but did 
not again catch sight of one. 


Phaethornis ruber ruber (Linn.). RED-VENTED HERMIT. 


The most abundant hummingbird. Two females spent 
much of their time searching surrounding heliconia blossoms 
for tiny insects and resting from time to time on a lower 
branch of the tree. 


Florisuga mellivora mellivora (Linn.). GREAT JACOBIN 
HUMMINGBIRD. 


Thalurania furcata furcatoides Gould. AMAZONIAN 
Woop-NyYMPH. 


Heliothria auriculata phainolaema Gould. GREEN BREASTED 
FAIRY. 


These three species of hummingbirds were observed perch- 
ing in the tree on several occasions. Two others were not se- 
cured and could not be identified by the glass. 


84 Zoologica: N. Y. Zoological Society [iis 


Ramphastos monilis Mill. RED-BILLED TOUCAN. 


In the cinnamon tree the visit of this large red-billed 
toucan was very evidently accidental as the berry-bearing 
branches were too slight to support his weight. I saw one on 
May 3rd, resting only for a moment before he flew on in the 
direction of the toucan tree. When the afternoon’s rain was 


over, the yelping cries of these birds were the most conspicuous 
sound of the jungle. 


Pteroglossus bitorquatus bitorquatus Vig. DOUBLE-COLLARED 
ARACARI. 


Twice observed in the cinnamon tree, and still oftener in 
the toucan tree. From a flock of eight secured two. Brilliant 
as these birds are, it is remarkable how easily they escape ob- 
servation when in the tree-tops. Even when one of a flock is 
discovered, the closest scrutiny with powerful glasses fails to 
reveal the remainder, until one by one they move and betray 
their whereabouts. When motionless they resemble an irregu- 
lar knot or bunch of leaves. When the broken stub of a branch 
contains water, they all visit it in turn, drinking after eating a 
half dozen or more berries. 


Colors: iris pale yellow, with a antero-posterior extention 
of dark brown pigment, giving the pupil an elongated appear- 
ance. Bare skin around eye blue, lower lid orange yellow; 
facial skin same red color as feathers of nape; upper mandible 
lemon yellow, whitish near base and at tip, black along cutting 
edge; lower mandible black on terminal two-thirds, greenish- 
white near base and along ventral line; legs and feet yellow 
green like the flank feathers. 


Pteroglossus inscriptus inscriptus Swains. LETTERED ARACARI. 


An occasional visitor to the tree, and when a flock of them 
came, they made such a commotion that callistes and other small 
birds could hardly get a foothold. Four out of a flock of five 
were shot about fifty yards from the tree and the following day 
the survivor remained near, through most of the hours of day- 


1916] Beebe: Birds of Parad, Brazil 85 


light, calling, and now and then feeding on the berries. The 
first bird shot was a young one and the rest actually followed 
it to within ten feet of where four of us were standing. Even 
after the third shot, the fourth bird came as boldly as ever in 
answer to the yells of the youngster. Bates and other writers 
speak of being mobbed by toucans in much the same manner. 


Of the four birds, two were males, two females. The young 
molting male had the iris scarlet ; crown above eye pale caerulean 
blue; eyelid, lores, beneath eye and around ear dark livid blue; 
broad line between eye and ear vermillion; skin back of nostrils 
bright blue; bill bright orange yellow and black; legs and feet 
sage green similar to the under tail web. The crop was filled 
with round, black seeds, which stained everything an indelible 
dark blue. 


Selenidera maculirostris gouldii (Natt.). GOULD’s TOUCANET. 


The commonest toucan in the tree, observed on four 
separate occasions in pairs or trios, but remaining only for a 
short time and very wary. The iris is lemon yellow above and 
below, shading off in front and behind into green, which changes 
to black next the pupil, giving it an extremely flattened, elongate 
appearance; bill black and white, with the terminal parts of 
both mandibles pale green; facial skin yellowish and bluish 
green; legs and feet bluish-green. 


Celeus gumana jumana (Spix). SPIX’S AMAZONIAN 
WOODPECKER. 


On May 3rd a single bird hammered at a soft place in the 
bark of the tree for five minutes, then caught sight of me be- 
neath and fled silently. 


Celeus undatus multifasciatus (Malh.). WAVED WOODPECKER. 


Observed by Cherrie in the tree on May 5th. Had been eat- 
ing berries. 


86 Zoologica: N. Y. Zoological Society Pipes 


Campephilus trachelopyrus (Malh.). MALHERBE’S BLACK 
WOODPECKER. 


Late in the morning of May 6th a pair alighted on the trunk 
ten feet from the ground and worked their way upward to the 
small branches before flying off through the jungle. A female 
collected some distance away had the iris pale orange, bill green- 
ish horn, darker along the culmen; legs and feet deep olive green. 
Crop filled with large yellow seeds. 


Picumnus aurifrons Pelz. AMAZONIAN GOLD-FRONTED PICULET. 


While watching a flock of Dacnis in the tree early in May, I 
noticed three small birds which at first glance reminded me of 
nuthatches. I secured two and found they were curious soft- 
tailed woodpeckers or piculets. Whether they came for berries 
or in hope of insect food I cannot say and I did not again have 
opportunity to observe them. The third bird remained motion- 
less in a neighboring tree for some time. Para is a new locality 
for this group, but these individuals seem to be quite typical. 


Thamnophilus amazonicus Scl. SCLATER’S AMAZONIAN 
BUSH-SHRIKE. 


While having no real right in an arboreal fauna I must in- 
clude this species, as a male bird flew up from the underbrush 
when I shot at it and missed, and alighted for a moment on one 
of the lower branches. With several other species it was not 
ancommon in the surrounding jungle. 


A few yards from the tree a little earlier in the day, I had 
stalked the same individual in thick underbrush, where it seemed 
to be at odds with a white-breasted manakin. After the latter 
flew off, the Bush-Shrike kept constantly in one place, close to 
the ground, singing every thirty of forty seconds. It was a 
simple refrain whut! whee-whee! whee-whee! whee-whee! 
When startled it uttered the whut! alone. It was difficult stalk- 
ing ground but only a loud crackle of leaves made the bush- 
shrike shift its perch. The female appeared for a moment and 
the male repeated his song twice very rapidly, and turning close 


1916] Beebe: Birds of Parad, Brazil 87 


to her ruffled all his feathers, making himself into a perfect ball, 
blatantly displaying the usually concealed white patch, and with 
the spotted shoulders protruding conspicuously from the round, 
slate-colored mass. Keeping thus inflated he hopped around 
and around on his perch, completing a half turn at each hop, 
stopping for a second or two between hops and twisting so as 
to face her. At this time his song came irregularly. Twice he 
began it while on the hop, but did not end it. The moment the 
female slipped away, all his excitement ceased and he went hard 
at work on his never ending ditty. Once the shadow of a pass- 
ing vulture fell upon him and cut short the refrain, but only 
fora moment. Great metallic bees buzzed close about the singer 
but were not noticed. I later found his crop crammed with small 
black ants. 


Xenops genibarbis genibarbis Ill. WHISKERED RECURVED-BILL. 


One seen on the tree, and once shot in the depths of the 
Utinga jungle. 


Glyphorhynchus cuneatus cuneatus (Licht.). WEDGE-BILLED 
WOODHEWER. 


The commonest woodhewer hereabouts, and observed al- 
most every day on the tree, moving creeper-like up and around 
the trunk. The slightly upward curve of the beak gives to the 
bird a decidedly nuthatch profile. This species seemed about” 
to nest and two females would have deposited eggs within a very 
few days. Its low, plaintive note often revealed its presence 
befcre it was seen. 


Xiphorhynchus guttatus eytoni (Scl.). EYTON’S FULVOUS- 
THROATED WOODHEWER. 


A pair of these large woodhewers were courting, a process 
which seemed to consist in the constant pursuit of one by the 
other. This took place along the trail on which the tree grew, 
and the birds alighted again and again in the tree but not to feed. 
After resting a moment, panting, they continued their endless 


88 Zoologica: N. Y. Zoological Society. [lls3 


chase. They were silent and only when the pursuer almost 
caught up did the other utter a sharp, querulous note. So fast 
did they fly that the two brown bodies would appear like streaks 
shooting in and out of the tree-trunks. As they were seen in 
the trail every day their nesting site was doubtless not far off. 


Xiphorhynchus pardalotus (Vieill.). CHESTNUT-RUMPED 
WOODHEWER. 


Seen only once and secured from one of the higher branches 
of the tree. 


Dendroplex picus picus (Gmel.). PICINE WOODHEWER. 


Next to Glyphorhynchus the commonest woodhewer seen 
near the tree. Once only did one alight on it, but others were 
seen constantly on the adjoining trunks. Owing to the large 
amount of white it was the most conspicuous of these birds. 
Several times I saw one alight crossways on a branch, the first 
time I have ever seen a woodhewer assume this passerine 
position. 


Picolaptes layardi Scl. LAYARD’S WOODHEWER. 


Dendrocolaptes certhia certhia (Bodd.). BUFFON’S BARRED 
WOODHEWER. 


I saw neither of these species but I examined specimens in 
the flesh shot from the tree by Mr. Cherrie in my absence. 


Rhynchocyclus sulphurescens (Spix). SULPHURY FLATBILL. 


Abundant in tree. A dozen could have been shot at each 
period of observation, had I wished them. An adult and a young 
male which were secured were both feeding on the tree berries. 
The latter was in very much worn juvenile plumage and about 
to moult. 


Rhynchocyclus poliocephalus sclateri Hellm. SCLATER’S 
FLATBILL. 


A male collected in the tree on May 10th had both tree 
berries and small Diptera in its crop. 


1916] Beebe: Birds of Para, Brazil 89 
Mionectes oleagineus oleagineus (Licht.). OILY FLYCATCHER. 


This was the commonest flycatcher which frequented the 
tree. I secured six and could have shot twenty on any of the 
days when I was on watch. Its bright buff breast rendered it 
one of the easiest birds to recognize, and after a day’s observa- 
tion I shot none by accident. Their food consisted both of tree 
berries and small insects. 


Ty’anniscus acer (Scl. and God.). SHARP-BILLED FLYCATCHER. 


These little flycatchers were rather rare and usually early 
comers. I secured none after seven-thirty in the morning, and 
even then they had been feeding for some time. Those collected 
in the tree had fed altogether on the tree berries. They were 
breeding at this season. Even with my powerful field glasses, 
and with knowledge of the points of difference it was absolutely 
impossible to distinguish this species from either of the pre- 
ceding forms of Rhynchocyclus. When eighty feet or more up, 
I do not think identification with glasses of these lesser fly- 
catchers can be accomplished. 


Elaenia flavogaster flavogaster (Thunb.). YELLOW-VENTED 
CRESTED FLYCATCHER. 


Observed several times in the tree feeding on the berries. 
It kept lower down than the other smaller species and was 
recognizable by its clean-cut, white markings. 


Elaenia gaimardti guianensis Berl. GAIMARD’S CRESTED 
FLYCATCHER. 


Only among the top-most branches with other small Fly- 
catchers. On two occasions when seen against a mass of dense 
foliage I detected the half-concealed, white crown, but usually 
the species merged wholly with the Rhynchocyclus and Tyran- 
niscus feeding with it. It was feeding wholly on the tree berries. 


90 Zoologica: N. Y. Zoological Society LIh;3 


Myiarchus tuberculifer (Lafr. and D’Orb.). D’ORBIGNY’s 
BLACK-HEADED FLYCATCHER. 


This was the only species of flycatcher which ever got in 
the least excited over my presence at the foot of the tree. As 
I was getting into position for a prolonged period of observa- 
tion, one or a pair of these birds would occasionally drop down 
from the upper branches and with crest raised, excitedly flutter 
from one branch to another uttering a continual sharp tsip! 
tsip! While the berries were eaten by all I examined, yet in- 
sects were never wholly absent, and more than once I saw birds 
of this species launch out high above the tree after passing in- 
sects. When seen against green foliage, even at a great height, 
the distinct areas of grey and yellow on the lower plumage were 
quite distinct. 


Empidonomus varius (Vieill.). AZARA’S FLYCATCHER. 


A specimen in worn plumage shot from the tree and three 
others near by. All must have been in the tree during the morn- 
ing as all had tree berries in their crops. 


Pipra fasciicauda Hellm. BANDED-TAILED MANAKIN. 


Several times I had watched orange and black manakins in 
the lower branches of the tree and supposed they were the com- 
mon red-headed species (Pipra rubricapilla). It is very prob- 
able that most of them belonged to that species, as all which 
Cherrie and I secured in the neighborhood of Utinga were 
rubricapilla. The single bird which I secured from the tree was 
the banded-tailed manakin. In its crop were two small beetles 
and seven tree berries. 


Pipra erythrocephala rubrocapilla Temm. RED-HEADED 
MANAKIN. 


The commonest manakin at Utinga. Early every morning 
a male would be perched on the same branch of the tree and 


1916] Beebe: Birds of Parad, Brazil sl 


twice I saw him driven away by other manakins. He never fed 
while I watched him, but sat sometimes for fifteen minutes with- 
out moving, paying no attention even to the sound of the gun 
or of the shot as it returned and swished through the leaves after 
I had fired a shot straight upward. 


Pipra leucocilla bahiae Ridgw. SLATE-BREASTED BLACK 
MANAKIN. 


Next to the red-crowned this manakin was most frequently 
seen. It was a female of this species which, with a male opal- 
crowned manakin, I secured from the tree with one shot. They 
had united to chase away the red-crowned bird from his perch 
and at once had flown upward beyond the usual height at which 
these birds are found. In the upper branches they joined a 
small flock which had come out of the jungle, and which soon 
left the tree and went on toward the north. 


Pipra suavissima Sol. and God. ORANGE-BELLIED MANAKIN. 


After a flock of roving jungle birds had left the tree I 
secured this specimen from their number. It had two tree 
berries and a great mass of insect larvae in its crop. I did not 
observe it again during my stay. 


Pipra opalizans Pelz. PARA OPAL-CROWNED MANAKIN. 


The female which I secured was in an adjoining tree, but 
only about twenty feet from the cinnamon tree, and within half 
an hour the small flocks of manakins appeared from which I 
got the male bird. There were five berries in the crop, which 
otherwise was empty. A day or two before seven or eight of 
these beautiful birds had been secured for Mr. Cherrie two 
miles away, by a native collector. Aside from these examples 
we saw nothing of the species. 


Piprites chlorion (Cab.). SCHOMBURGK’S MANAKIN. 


Shooting at what I took to be a flycatcher of some new 
species I secured a female of this species from one of the lower 


92 Zoologica: N. Y. Zoological Society Lites 


branches. It had been hopping about for some time in the neigh- 
boring jungle and its crop contained only small insects. It was 
quite alone and I saw nothing, nothing of its mate or of other 
individuals. 


Chiroxiphia pareola pareola (Linn.). BLUE-BACKED MANAKIN. 


Twice seen and one male secured. On May 5th a male had 
been flying back and forth for some time before I gave it care- 
ful attention. Although well above the ground, it showed its 
crown and back so distinctly that I knew it at once, and watched 
it through the glasses snatching berries and chasing some species 
of Dacnis through the branches. 


Chiromachaeris manacus purus Bangs. EASTERN WHITE- 
BREASTED MANAKIN. 


Although a day seldom passed when I did not see this 
Species near the tree, it was only on the last day of observation 
that I saw it actually in the tree itself. Two manakins of un- 
known species were having a most excited time in the lower 
branches and making all the noise of which they were capable. 
The uproar drew two male white-breasted manakins from the 
jungle undergrowth and they flew up without hesitation to see 
what the matter was. When they reached the branch the row 
soon ended and all concerned sought privacy again. A pair was 
always to be found about one hundred yards from the tree on 
the edge of the jungle where an old cultivated field had grown 
up to dense briery undergrowth. A second pair must have had 
a nest within ten or fifteen yards of the trail, although most 
careful search failed to locate it. While sitting quietly near the 
tree the female often came close and peered at me, hopping from 
twig to twig, and at each flight producing the characteristic 
deep, low whirrrrrrrrr! the wing song by which these little 
jungle people give vent to their emotions—courtship, suspicion, 
fear. 


Tityra cayana (Linn.). CAYENNE TITYRA. 


One of these birds perched for some time in a tree close to 
our house on the first day of our stay at Utinga. I saw no more 


1916] Beebe: Birds of Pard, Brazil 93 


of the species until I found that late in the afternoon just after 
the rain or even while it was still falling, three of these tityras 
came to the tree regularly in company with one or two Cotingas. 
I saw them under these conditions on three separate occasions 
and watched them feeding on the berries at leisure. 


Tityra inquisitor erythrogenys (Selby). RED-CHEEKED TITYRA. 


One early in the morning of May 6th. Not seen again. 


Platypsaris minor (Less.). LITTLE PSARIS. 


At seven A. M. on May 6th the cinnamon tree seemed 
almost deserted. I arranged my canvas chair and lying back, 
searched the upper branches carefully with my glasses for signs 
of life. Suddenly I saw motion in the tip of what I had thought 
was a broken branch stub. Several minutes passed and as I 
could make nothing of it, I secured it and found it to be a female 
psaris. It had evidently been feeding elsewhere as well, as the 
stomach contained a large yellow seed and a green grasshopper, 
while in the crop were three tree berries. 


Pachyrhamphus rufus (Bodd.). CINEREOUS THICKBILL. 


Twice I observed the unmistakable female of this species 
feeding in the tree, but was unable to secure it. On the following 
day we shot a specimen some distance away. Its only food was 
hairy caterpillars. It seems a silent, quiet bird, slow in move- 
ment and stupid in taking alarm at the warning cries or flight 
of other birds. 


Pachyrhamphus marginatus (Licht). LICHTENSTEIN’S 
THICKBILL. 


Quite ignorant of what I was shooting at, I secured a female 
of this thickbill from the very top of the tree where it was feed- 
ing in company with callistes and flycatchers. It had break- 
fasted on a spider and several tree berries. 


94 Zoologica: N. Y. Zoological Society [I1;3 


Lathria cinerea (Vieill.). GOLD BIRD. 


The gold or greenheart birds as they are known in Guiana, 
were found in the Utinga jungle, isolated as usual, vague call- 
ing Voices, penetrating and ventriloqual. A great fig ten yards 
from my cinnamon tree was a favorite perch of one of these 
birds and twice or more each morning it came to the berry tree 
to snatch a mouthful of the fruit and dash back again. It would 
utter its call the moment it alighted, but I never heard it given 
elsewhere than from this perch in the dense heart of the great 
fig tree. 


Attila brasiliensis (Less.). SCHOMBURGK’S ATTILA. 


While observing this species and after I had secured a 
specimen I supposed I was dealing with some unknown form 
of flycatcher, although I had never known any member of the 
Tyrannidae with such a marvellous vocabulary as had these 
birds. Two individuals, one adult and a young male, were in 
the tree early on the morning of May 1st and ultimately I se- 
cured the latter and identified the species. They were exceed- 
ingly active and playful. The full-grown young bird would ap- 
roach its parent, fluttering its wings and begging for food, then 
being chased swiftly through the jungle and back again, or 
swinging around, would pursue the other in turn. The song, 
which was uttered every ten or fifteen seconds was exactly alike 
in the two birds. It was a high, liquid four note phrase, wheedle- 
wheedle-wheedle-wheedle! Four rapid repetitions was the rule, 
more rarely increased to five or six. But this was constantly va- 
rid from the more usual timbre. When uttered while in pursuit of 
one another it became higher and shriller, or when given as the 
overgrown youngster was swallowing a berry it was fairly 
gargled. Again only a single whee! would be uttered, standing 
for some unknown emotion. At least a score of variations or 
shades of utterance were heard in fifteen minutes. The note of 
suspicion or alarm, given when I made too loud a noise or when 
another bird or a squirrel alarmed them, was very different, a 
loud, sharp, woodpecker-like cackle. After this was uttered 
once or twice, during which time the birds were motionless, the 
wheedle call or song commenced, the Attilas becoming at once 
active. 


1916] Beebe: Birds of Para, Brazil 95 


’ They kept to the.tree-tops and only by a quick, long-distance 
shot was I able to secure the young bird. The iris was pale 
hazel-brown; upper mandible horny black; lower also, with a 
large, fleshy-white patch mid-way along the rim on each side. 
The inner gape showed the loose yellow skin so characteristic 
of young birds; legs and feet slaty-blue; soles yellowish-flesh ; 
claws dark brown. 


The most unexpected fact was in connection with its food. 
The crop was full of berries and there were two which had not 
yet been swallowed, but in the gizzard were the recognizable 
remains of a small fish. The only way I can account for this 
unusual item of diet is that the birds must have been drinking 
at a jungle pool near by in which were many small minnow-like 
Tetragonopterus, and the young bird in some way had managed 
to seize and swallow one. 


Cotinga cayana (Linn.). CAYENNE CHATTERER. 


Once or twice these brilliant birds were seen in the mango 
trees near our house, but like the tityras I did not see them 
elsewhere than in the cinnamon tree in late afternoons. There 
were usually two, one in full color and the other a female or 
young bird. Their brilliance absolutely disappeared when seen 
against the bright sky, but in contrast with the green leaves or 
a cloud, lighted by the slanting rays of the sun, they flashed like 
great gems. 


Thryothorus genibarbis genibarbis Swains. SWAINSON’S 
MOUSTACHED WREN. 


For two days in succession a pair of these birds remained 
in the neighborhood of the tree, occasionally visiting the lower 
branches, but only momentarily and, as far as I could judge not 
touching the berries, but intent only on insect prey. One made 
occasional attempts at song, but the season was evidently past 
or had not yet arrived. 


96 Zoologica: N. Y. Zoological Society flies 


Troglodytes musculus clarus Berl. and Hart. VENEZUELAN 
HOUSE WREN. 


This is, of course, not a bird of the jungle and its presence 
in the tree was accidental, and as far as my observation went 
occurred only once. The bird seen was doubtless one of a pair. 
which lived in and about the clearing about our house, and made 
deeper foraging inroads now and then into the jungle. It was 
probably the number and commotion of the small callistes and 
other birds feeding in the tree which drew the inquisitive wren 
thither early in the morning on May 6th. 


Planesticus phaeopygus phaeopygus (Cab.). CABANIS’'S WHITE- 
THROATED THRUSH. 


Not uncommon in isolated pairs through the jungle, and an 
occasional visitor to the tree especially in late afternoons. They 
went about feeding in a business-like manner, apparently filling 
their crops in a short time. The nesting season for them had 
just begun. 


Vireo chivi (Vieill.). CHIVI VIREO. 


At 8 A. M. on May 5th I secured this bird from the upper 
branches of the tree, not knowing at what I was shooting except 
that it had a different carriage from the flycatchers and dacnis 
which thronged the upper foliage. Four tree berries were in 
its crop and a fifth still unswallowed in the mouth. 


Pachysylvia thoracica semicinerea (Scl. and Sal.). GREY-NAPED 
Woop VIREO. 


Cyclarhis gujanensis gujanensis (Gmel.). GUIANA VIREO- 
SHRIKE. 


I secured these birds within ten minutes of one another on 
May 2nd. Both were feeding on the berries of the tree. 


1916] Beebe: Birds of Para, Brazil o% 


'The previous day we had shot elsewhere an adult male 
vireo-shrike and a young male of the year in very worn plu- 
mage. Comparison of these two showed the following differences: 


Adult Juvenile 


SOG inp eee ee ag ey ec a 147 143 
OUI ITCS aE S Ne Oe te Rice See 16 15 
Cimlimens tron NOStEIL oe 10 9 
NIT O2S SR aS ie ea i ee nee eh 70 65 
NSTI eo a A PRD 55 55 
a MENTS UICT ska oie Te Re 20 ZN 
Middletoe.and claws...0 eS. 16 1g 
Bill: adult reddish horn; juvenile slaty grey; tips in both 


whitish. 
Legs and feet: adult brownish blue; juvenile clear slaty blue. 
Iris: adult reddish orange; juvenile hazel, paling outwardly. 
Bare facial area: adult warm flesh; juvenile olive green. 
Forehead: adult rich chestnut; juvenile grey like head. 
Superciliary: adult rich chestnut; juvenile warm buff. 


Cyanocompsa rothschildu (Bartl.). ROTHSCHILD’S BLUE 
GROSBEAK. 


I saw this bird on two occasions feeding on the berries of 
the tree, although it was probably the same individual. The 
second time it descended to one of the lower branches and re- 
mained motionless for many minutes. 


Saltator maximus (Miill.). GREAT SALTATOR. 


Saw but one of this species in the tree and that quite an 
accidental visitor as it perched only for a few seconds on a lower 
limb and then flew straight off through the jungle. Two days 
later we secured a specimen a mile away, but saw no others 
during our stay. 


Coereba chloropyga chloropyga (Cab.). BRAZILIAN 
F'LOWERPECKER. 


A few of these little birds were seen almost every day in 
the tree usually well up near the top, but unlike most of their 


98 Zoologica: N. Y. Zoological Society [11;3 


companions feeding apparently altogether on small insects. The 
first one which I saw in the tree was on a lower branch by itself, 
singing with all its might. Its song was sweet, rather short 
end of a wheezy character with a quaint little lilt. This in 
spite of the fact that it was in very worn, shabby breeding 
plumage. 


Dacnis cayana cayana (Linn.). TURQUOISE HONEY-CREEPER. 


These exquisite little birds were one of the most abundant 
species which frequented the tree. I saw at least fifty during 
each period of two or three hours of observation. All which I 
secured were feeding on the berries. They usually kept to the 
upper branches, flying swiftly from the surrounding jungle 
summits, and moving actively about, now and then catching an 
insect but preferring the tree berries. This was the only species 
of the Family which ever came down to lower branches. When 
well up it was impossible to differentiate between this and the 
next species. The color of the turquoise honey-creeper is re- 
markable. When the bird is held between the observer and the 
light, no matter how oriented, whether sideways, head or tail 
on, it is a deep cobalt blue; when looked at with the light behind 
the observer, it is as intense a clear, shining turquoise. There 
is no position of feather or bird which will alter these colors. 


Dacnis angelica angelica Bonap. BLACK-BACKED 
HONEY-CREEPER. 


Still more active than the turquoise, this bird equalled it 
in numbers, and sometimes twenty were in the top of the tree 
at one time. 


Cyanerpes cyaneus cyaneus (Linn.). BLUE HONEY-CREEPER. 


The blue honey-creeper, perhaps the most beautiful of all 
this group, was much more common at the tree in the afteroon 
than in the morning. I was able to identify the males of these 
birds at any height and found them in the proportion of two in 
the morning to seven in the afternoon. I have counted eighteen 
individuals at one time. They seldom descended to the lower 
branches. In every specimen I examined there were a few in- 
sects in the crop in addition to the tree berries. 


1916] Beebe: Birds of Pard, Brazil 99 
Chlorophanes spiza spiza (Linn.). GREEN HONEY-CREEPER. 


The fourth member of this group, glowing with its green 
iridescence in the sunlight. Instead of insects these birds were 
plucking tree berries with their long curved beaks. They seemed 
equally abundant, whether at daybreak or after the daily rains 
in late afternoon. Eight males and two females were grouped 
together on one of the central branches for fully five minutes 
one morning, excited about something which the most careful 
scrutiny with my glasses failed to reveal. 


Chlorophonia chlorocapilla (Shaw). BLUE-BACKED GREEN 
TANAGER. 


This bird which appears to be new to this part of Brazil 
was shot accidentally. I aimed at a blue-bellied tanager in the 
tree, missed it, and this small, wonderfully-colored species, which 
I had quite failed to observe, dropped from an upper branch. 
it had two tree berries in the crop. 


Tanagra violacea lichtensteinii (Cab.). NORTHERN VIOLET 
KUPHONIA. 


Never present in large numbers but several pairs were sure 
to turn up in the tree during the day. They did not remain long, 
perhaps, because a berry or two must have made a cropful for 
such diminutive chaps. No matter how busy hopping about, 
they always found time every few minutes to stop and burst 
into their jubilant little song. 


Tanagra cayennensis (Gmel.). CAYENNE EUPHONIA. 


Decidedly rare in the tree. Saw four and secured one. Easy 
to identify when not silhouetted against the sky, the two lateral 
patches of orange feathers standing out in strong contrast with 
the blue black of the remainder of the plumage. The specimen 
which I shot had small green seeds in its crop, not those of the 
tree. 


100 Zoologica: N. Y. Zoological Society Pies 


Tanagrella velia signata Hellm. PARA BLUE-BELLIED TANAGER. 


On May 5th Cherrie shot a female of this beautiful bird 
from the tree and within five minutes I secured its mate. On 
three later occasions I observed this tanager, always in pairs 
and in the early morning. It could not be recognized with cer- 
tainity in the upper branches as the yellow of the black was 
usually concealed. They fed greedily on the tree berries. 


Tangara punctata punctata (Linn.). SPOTTED TANAGER. 


Early visitors to the tree, coming singly or in pairs straight 
across the top of the jungle as if from a distance. They knew 
the tree well and began to feed as soon as they arrived. After 
they had eaten several berries they would appear satiated and 
either sit in the sun and preen their feathers or chase one an- 
other about, always returning from the surrounding jungle for 
another period of feeding before they left. 


Thraupis episcopus episcopus (Linn.). WHITE-SHOULDERED 
BLUE TANAGER. 


Blue tanagers were rare at the tree although common else- 
where, and wher they appeared came singly or in pairs. I saw 
them there only three times. This may have been because they 
were nesting at this season, a pair of birds having a nest in a 
mango tree a few yards from our house. There were two young 
birds and these flew on May 8th. 


Thraupis palmarum palmarum (Wied.). PALM TANAGER. 


One bird shot in the tree in company with a flock of silver- 
beaks on May 8th. Its mate fed for some time afterwards on 
the tree berries. Although fairly common elsewhere on the 
borders of the jungle no more were observed in the tree. 


Ramphocelus carbo carbo (Pall.). SILVER-BEAKED TANAGER. 


The commonest bird at Utinga and almost constantly pres- 
ent in the tree. When large numbers of eallistes and fly- 
catchers were gathered together there would sometimes be only 


1916] Beebe: Birds of Para, Brazil 101 


one silver-beak. Then with a rustle of wings a whole flock 
would fly up from the surrounding jungle, twenty or thirty in 
all, and without actual aggression but by sheer numbers would 
disturb most of the smaller birds. They would chase other 
birds half playfully or in turn be pursued by some flycatcher, 
but on the whole the tree-top assemblage of birds was a peace- 
ful one. The quickest glance served to identify these tanagers, 
for though their white beak might be invisible, and their plum- 
age appear jet black viewed against the bright sky, the char- 
acteristic sideways flirting of the tail never failed. Their sharp 
metallic chip! was another positive factor of identification. 
They were restless, never remaining very long in the tree but 
flying off one after the other to work their way slowly through 
the jungle. 


Tachyphonus cristatus brunneus (Spix). SCARLET-CRESTED 
TANAGER. 


One specimen with a number of honey-creepers was secured 
in the tree early on May 2nd. Did not note another during my 
stay. 


Tachyphonus surinamus insignis Hellm. PARA CRESTED 
TANAGER. 


Three or four times I observed this bird at the tree feeding 
on the berries and secured two specimens. Its peculiar mark- 
ings enabled me to identify it at almost any height. On May 
5th a male suddenly swooped down from the upper branches 
and showed great agitation upon finding me in my observation 
chair. I soon discovered that the cause was a female and single 
young in the undergrowth near by, who were attracted rather 
than frightened by the emotion of the male. They soon took 
themselves off, and in a few minutes the male crested tanager 
was again back feeding in the tree. 


Hemithraupis guira guira (Linn.). GUIRA TANAGER. 

Early on May 5th a pair of these birds high up in the tree. 

One of these I secured. The other continued feeding and flying 

about the tree with honey-creepers and flower-peckers for some 
time afterwards. 


102 Zoologica: N. Y. Zoological Society Cites 


Ostinops viridis (Miill.). GREAT GREEN CACIQUE. 


A small colony of these splendid birds was established near 
the toucan tree, to which tree they paid frequent visits. Only 
once did I see one in the cinnamon tree and then only for a 
minute. He snatched two berries, looked carefully about him, 
down at me and flew off through the jungle in the direction of 
the colony, a few hundred yards away. 


Cacicus cela (Linn.). YELLOW-BACKED CACIQUE. 


Five times I saw these birds in the tree feeding greedily on 
the berries. The slenderness of the branches seemed to bother 
them, however, and they never remained long. They constantly 
haunted the toucan tree several hundred yards away, which 
had larger berries and stouter branches. There were three 
separate colonies within the radius of a half mile, the nearest 
enly a hundred yards from the tree and from our house in the 
yard of a native. In certain zones of the jungle the squeaks and 
gurgles of these birds were the dominant sounds throughout 
the day. 


In spite of this indescribable squeaking and yelping, the 
yellow-backed caciques appear to have a consistent call or song. 
It may be written, yank! yank! yank-keou-ke-wonk! 


Cacicus haemorrhous haemorrhous (Linn.). BRAZILIAN RED- 
RUMPED CACIQUE. 


One individual was in the tree on May 6 with four yellow- 
backed birds. I could not secure it but watched it for more 
chan five minutes. 


B.—AERIAL BIRDS. 


Catharista atratus brasiliensis (Bonap.). BRAZILIAN BLACK 
VULTURE. 


Five minutes seldom passed, hour after hour, when one or 
more of these birds did not soar across the bit of sky visible 
above the cinnamon tree. Usually they were very high up, 


1916] Beebe: Birds of Parad, Brazil 103 


soaring, but occasionally just sweeping the tree tops with their 
pelican-like habit of alternate flapping and gliding. At sunset 
scores flew past southward, just clearing the jungle, or else col- 
lected on some dead tree until twenty or thirty had assembled, 
when all flew off in the same direction to some distant roost. 


Chordeiles acutipennis acutipennis (Bodd.). SouTH AMERICAN 
NIGHTHAWK. 


Chaetura spinicauda spinicauda (Temm.). SPINE- 
TAILED SWIFT. 


Tachycineta albinenter (Bodd.). WHITE-VENTED TREE 
SWALLOW. 


Progne chalybea chalybea (Gmel.). GREY-BREASTED MARTIN. 
Atticora fasciata (Gmel.). WHITE-BANDED SWALLOW. 


Stelgidopteryx ruficollis ruficollis (Vieill.). BRAZILIAN ROUGH- 
WINGED SWALLOW. 


The above six species were observed, the first in late after- 
noon and the others throughout the day, hawking about in the 
sky over the tree. None were very rare, the last named, per- 
haps, the most abundant. In clear weather they flew high, but 
as the clouds gathered they settled lower, following the shifting 
strata of volant insect life. 


C.—BIRDS OF THE SURROUNDING JUNGLE. 


Crypturus variegatus (Gmel.). VARIEGATED TINAMOU. 


These tinamou were twice seen and heard daily within a 
few yards of the cinnamon tree. Their plaintive, sustained 
note was one of the commonest sounds of the jungle. They 
would reply to an imitation of their notes and even approach, 
but never close enough for a shot, and no especial effort was 
made to stalk them. The only specimen examined was one in 
the last stages of decomposition which had met its death a few 
feet from the cinnamon tree trail. It was being skeletonized 
by ants and there was left barely sufficient plumage for identi- 
fication. 


104 Zoologica: N. Y. Zoological Society [1133 


Geotrygon montana (Linn.). RED GROUND DOVE. 


Not uncommon on the jungle floor, flushing with a loud 
noise of wings, and at first being confused with small tinamou. 
One which I secured showed no evidence of recent breeding. 


Ceophloeus lineatus (Linn.). GREAT LINEATED WOODPECKER. 


In late morning on May 9th as I sat watching under the 
cinnamon tree I was bothered for ten or fifteen minutes by 
what I thought were two men building a house. The hammer- 
ing was loud and incessant, and I could tell when first one then 
the other began work as their boards gave forth varying tones. 
Often they would go at it together. Finally I heard a resound- 
ing rattle, more rapid and staccato than any hammering car- 
penter could produce, and my suspicion aroused, I walked to the 
end of the trail at the edge of the jungle. Out in a cleared field 
stood a headless, weatherbeaten royal palm and to this were 
clinging a pair of great lineated woodpeckers hammering in- 
termittently and audible half a mile away. 


Ceryle torquata torquata (Linn.). GREAT GREY KINGFISHER. 


On May 8th one of these splendid kingfishers passed over 
the tree on its way from one igarapé to another. 


Trogon melanurus melanurus Swains. BLACK-TAILED TROGON. 


The commonest Trogon hereabouts, more often heard than 
seen. 


Crotophaga ani Linn. COMMON ANI. 


Abundant in the surrounding brushy fields, but seldom ven- 
turing far into even the more open jungle. 


Thamnophilus aethiopes incertus Pelz. PELZELN’S 
BUSH-SHRIKE. 


Not uncommonly seen and heard, and easy to recognize at 
sight after a specimen has been examined. 


1916] Beebe: Birds of Para, Brazil 105 


Cercomacra tyrannina Scl. TYRANT ANT-WREN. 


A common species in the undergrowth of the jungle, fre- 
quently seen close to the base of the tree, hopping about or 
scratching among the dead leaves. 


Synallaxis rutilans omissa Hart. Para SPINETAIL. 


A pair of these birds were nesting very close to the tree, 
and were never quite reconciled to my continued presence. 
Strangely enough, the occasional sound of the gun did not 
seem to alarm them. They kept rather low down, but five min- 
utes seldom passed without one or the other coming to have a 
look at me, and voicing’ its dissatisfaction in a low chut. Speci- 
mens secured elsewhere showed that this species was both pre- 
paring to lay and brooding in early May. 


Pipra aureola (Linn.). ORANGE-HEADED MANAKIN. 


Not uncommon in the surrounding jungle, and I am almost 
certain that some of those in the tree itself were of this species. 


Pitangus sulphuratus sulphuratus (Linn.). KISKADEE 
FLYCATCHER. 


Conspicuous in appearance and vocally. An inhabitant of 
the open places but occasionally flying over or alighting on 
jungle near the tree. 


Muscivora tyrannus (Linn.). FORK-TAILED FLYCATCHER. 


This unmistakable species was seen several times flying over 
the tree. 


Volatinia jacarini splendens (Vieill.). GLOSSY GRASSQUIT. 


Common in the overgrown fields fifty yards beyond the tree 
at the edge of the jungle. Once a pair flew past down the trail 
headed for the pumping station clearing. 


106 Zoologica: N. Y. Zoological Society Boke: 


Brachyspiza capensis capensis (Miill.). CHINGOLO SONG 
SPARROW. 


Heard singing in the nearest open glade and twice seen at 
the base of the tree. 


Arremon silens (Bodd.). PECTORAL SPARROW. 


These beautiful sparrows were not rare in the undergrowth 
at the base of the tree and as I was seated on watch, one or two 
would now and then flit across the trail with sharp chirps, com- 
ing back as closely as they dared to stare at me, hopping about 
nervously. 


Volume 2. Number 4. 


FAUNA OF FOUR SQUARE FEET OF 
JUNGLE DEBRIS 


L 


For a week I had been studying the bird-life of a single tree, 
a Canella do Matto, as I have described in detail in the preced- 
ing number of ZOOLOGICA. On the last day as I was about to go, 
I concentrated my attention on the tree and the surrounding 
jungle, endeavoring to fix it indelibly in my mind. I realized 
that in a few minutes I would leave this place with which I had 
become so intimate, and should very probably never return. I| 
had demonstrated a remarkable concentration of bird-life when 
attracted by the ripened fruit of a single jungle tree. It was 
the unparalleled insurgence of such a variety of organisms as 
can occur only in the tropics. 


Now that there remained only a brief space of time I tried 
to conceive of some last thing I could do to re-emphasize this 
important phase of tropical life. 


As I walked slowly up the trail toward the tree I heard a 
rustling among the leaves at one side, and in deep shadow 
beyond a dense clump of scarlet heliconias, I made out a tyrant 
antwren (Cercomacra tyrannina) scratching with all its might. 
To the kicking power of its small legs it occasionally added sud- 
den flicks with the bill, given with such nice judgment and 
power, that it flung leaves larger than itself into the air and 
backward quite over its body. I had often wondered of what 
the food of these birds really consisted. Anyone could glance 
at the contents of a crop and gizzard and label it “small insects.” 
But the actual details of this varied bill of fare, except in the 
case of very recently swallowed objects, was usually merged and 
lost in the comminuted mass of legs, elytra and antennae. 


107 


108 Zoologica: N. Y. Zoological Society [ier 


Acting on this hint I brought from my camping stores an 
empty war bag, and carefully scraped together a few handfuls 
of leaves, sticks, moss, earth and mold of all sorts. From 
directly under the Canella do Matto, I gathered four square feet 
of jungle debris, filled my bag and shouldered it. Then I said 
adieu to my trail and my tree, a sorrowful leave taking as is 
always my misfortune. For the bonds which bind me to a place 
cr a person are not easily broken. 


In this case, however, the bond ‘was not altogether severed, 
and a week later when the sky line was unbroken by land, when 
a long ground swell waved but did not break the deep blue of 
the open sea, I unlaced my bag of jungle mold. Armed with 
forceps, lens and vials, I began my search. For days I had gazed 
upward; now my scrutiny was directed downward. With 
binoculars I had scanned without ceasing the myriad leaves of 
a great tree. Now with lens or naked eye I sought for signs 
cf life on an infinitely smaller scale; the metropolis of a fallen 
leaf, the inhabitants of a dead twig. When I studied the tree- 
top life in the lofty jungle I was in a land of Brobdingnag; now 
I was verily a Gulliver in Lilliput. The cosmos in my war bag 
teemed with mystery as deep and as inviting as any in the 
jungle itself. 


When I began work I knew little of what I should find. My 
vague thoughts visualized ants and worms, and especially I 
anticipated unearthing myriads of the unpleasant macuins, 
or béte rouge, whose hosts had done all in their power to make 
life in the jungle unhappy. 


For ten days or more on the steamer trip north Mr. Hartley 
and I labored over the jungle debris. After two hours steady 
concentration our eyes rebelled and we had to desist. It seemed 
at times as if the four square feet had increased to forty, but the 
last handful was finally sifted and teased to shreds. Our method 
of work was to place a small pile on a newspaper spread on a 
table under the skylights of the smoking room, and with forceps 
and dissecting needle to search carefully every surface of leaf 
and frond and to split every twig and stem. 


It was found that the safest way to capture the minute 
ereatures which crawled or hopped about was to wet a small 


1916] Beebe: Fauna of Jungle Debris 109 


brush in alcohol, touch them with the tip and float them off in 
the liquid in a very small vial. Thus they were uninjured and 
we could pick them from a mass of earth or fungus without 
including any of the debris itself. Usually we worked with our 
naked eyes, but occasionally hunted over a particularly rich field 
with low power dissecting lenses. 


Day by day our vials increased. Scores of creatures evaded 
our search. Many others, of which I had captured a generous 
number, I allowed to escape. My lilliputian census was far from 
the mere aggregation of ants and worms which I had antici- 
pated, and a review of the whole showed that hardly any great 
group of living creatures was unrepresented. 


Two objects indicated the presence of wild mammals. First 
a bunch of rufous hairs which in size, color and minute struc- 
ture were identical with those of the common agouti, which was 
very common at Utinga. I also found sign of this rodent. Man 
himself was represented by two wads which had dropped from 
my gun-shots sometime during the week. One had already be- 
gun to disintegrate, wet, half decayed and inhabited by half a 
dozen tiny organisms. 


Five feathers were the marks of birds, also doubtless the 
result of my study during the week. A body feather, and two 
primaries from a sparrow-like bird were indeterminate, but two 
brilliant, green plumes came without question from the body of 
a calliste. Of reptiles there was a broken skull of some lizard, 
half disintegrated with a few of the teeth still left. There was 
besides the small egg-shell of a lizard which had hatched and 
gone forth to live its life elsewhere in the jungle. A third rep- 
tilian trace may have been his nemesis—a good-sized shred of 
snake-skin. The group of amphibians was present even in 
this small area of four square feet—a very tiny, dried, black 
and wholly unrecognizable little frog. Fishes were absent, 
although from my knees as I scraped up the debris, I could 
almost see a little igarapé in which dwelt scores of minnows. 


As I delved deeper and examined the mold more carefully 
for the diminutive inhabitants, I found that this thin veneer 
from the floor of the jungle appeared to have several layers each 
with its particular fauna. The upper layer was composed of 


110 Zoologica: N. Y. Zoological Society [Il34 


recently fallen leaves, nuts, seeds and twigs, dry and quite fresh. 
As yet these showed but little change, and only the damage 
wrought by insects and other agencies while they were still on 
the trees. In this layer were small colonies of ants in hollow 
twigs and occasional huge solitary ones. Here lived in hiding 
small moths, beetles and bugs awaiting dusk to fly forth through 
the jungle. The lowest layer was one chiefly of matted, thready 
roots holding together compact masses of earthy soil, mixed 
with a large proportion of tiny bits of quartz. The animal life 
of this stratum was very meagre, occasional mites—especially 
red ones—and a few earth and round worms, the latter in much 
fewer numbers than in the middle layers. 


Between the upper and the middle layers were sprouting 
nuts and seeds, with their blanched roots threaded downward 
into the rich dark mold, and the greening cotyledons curling 
upward toward light and warmth. Thus had the great Canella 
do Matto itself begun life. In my war bag were a score of poten- 
tial forest giants doomed to a death in the salt ocean. 


The middle layer, finally, was the all-important stratum. 
In it lived four-fifths of the small folk. This was composed of 
debris in full course of disintegration; leaves, sometimes partly 
green, usually brown or black, nuts half decayed, twigs half 
rotten. All still preserved their form, although some were ready 
to fall apart at a touch. All were soaked through, or at least 
damp and soggy. Often four or five leaves would be stuck to- 
gether, stitched with the threads of fungi. In such a haven was 
always a host of living organisms. 


Some of the half decayed leaves were very beautiful. Vistas 
of pale, bleached fungus lace trailed over the rich mahogany 
colored tissues, studded here and there with bits of glistening, 
transparent quartz. Here I had many hints of a world of life 
beyond the power of the unaided eye. And here too the grosser 
fauna scrambled, hopped or wriggled. Everywhere were tiny 
chrysalids and cocoons, many empty. Now and then a plaque 
of eggs, almost microscopic, showed veriest pin-pricks where 
still more minute parasites had made their escape. Contracting 
the field of vision to this world where leaves were fields and 
fungi loomed as forests, competition, the tragedies, the mystery 


1916 | Beebe: Fauna of Jungle Debris imal 


lessen not at all. Minute seeds mimicked small beetles in shape 
and in exquisite tracery of patterns; small beetles curled up and 
to the eye became minute seeds of beautiful design. Bits of 
bark simulated insects, a patch of fungus seemed a worm, and 
in their turn insects and worms became transmuted optically 
into immobile vegetation. Scores of little creatures were wholly 
invisible until they moved. Here and there I discovered a life- 
less boulder of emerald or turquoise—the metallic cuirass of 
some long dead beetle. 


Some of the scenes which appeared as I picked over the 
mold, unfolded suddenly after an upheaval of debris, were start- 
ling. When we had worked with the lens for many minutes, all 
relative comparisons with the surrounding world were lost. In- 
stead of looking down from on high, a being apart, with titanic 
brush of bristles ready to capture the fiercest of these jungle 
creatures, I, like Alice in Wonderland, felt myself growing 
smaller, becoming an onlooker, perhaps hiding behind a tiny leaf 
or twig. This feeling became more and more real as we labored 
day after day, and it added greatly to the interest and excite- 
ment. Close by would appear, under the lens, piles of great logs 
and branches protruding from a heaped up bank of precious 
stones. Mauve, yellow, orange’ and cerulean hues played over 
the scene. Over a steep hill came a horned, ungainly creature 
with huge proboscis and eight legs, and shining, liver-colored 
body, all paunch, spotted with a sickly hue of yellow. It was 
studded with short, stiff bristles, and was apparently as large 
as a wart hog and much more ugly. It was a mite, one of the 
biting mites of the tropics, but under the lens a terrible monster. 
We put one of these on our arm to see if its bite corresponded 
to that of the legions of macuins which tortured us daily 
in the jungle. Under the lens I saw the hideous creature stop 
in its awkward progress and as it prepared to sink its proboscis 
we involuntarily flinched, so fearful a thing seemed about to 
happen. 


In the middle layer, that of most active change, and sur- 
charged with life, ants were abundant, together with small 
colonies of termites. These were the only social insects, the 
twigfuls consisting of from five to fifteen members. All the 


112 Zoologica: N. Y. Zoological Society pue! 


other organisms were isolated, scattered here and there. Life 
in these lowly places, so far beneath the sunlight, is an individual 
thing. Flocks and swarms are unknown, and the mob has no 
place here. Each organism must live its life and fulfil its destiny 
single-handed. Even when two individuals were found together 
it was apparently more through accident of environment than 
from any gregarious instinct. In fact the same tropical law 
which holds good in regard to plants and the larger creatures 
of the sunlighted world overhead applies here. I found numbers 
of different species, but very few collections of individuals of 
the same kind. 


Flatworms were rather rare, but small, white ones were 
found now and then flowing slowly along in their characteristic 
manner over the surface of damp, half decayed leaves, as flat- 
worms do the world over. Roundworms, small, white and 
threadlike were present in equally small numbers. Earthworms 
of small size, one or two inches in length, were common. They 
moved slowly along in orthodox angleworm fashion until some- 
thing alarmed them when they instantly became a maze of twist- 
ing, snapping curves, dancing all about in a most unwormlike 
fashion. The head and especially the collar were brightly 
colored, from reddish to an intense scarlet. 


Centipedes and millipedes were common, all small, in keep- 
ing with the diminutive size of the other inhabitants of this 
little world. The largest centipede was less than an inch in 
length and scurried along on eighty-four legs. Very few were 
dark colored. Almost all were dead white, with yellowish brown 
heads and jaws. The larger millipedes were slow moving in 
spite of their abundance of feet, but small ones of various species 
were very agile, and slipped in and out of fungi forests in a 
most disconcerting way. ‘They were about evenly divided be- 
tween the groups of Polydesmoidea, Julioidea and Gerphiloidea. 


Scorpions were decidedly rare, and two small and one 
medium sized specimen were all we could discover. Pseudo- 
scorpions, however, were abundant and conspicuous. I secured 
fifty, and could have taken three or four times as many. They 
would rush out excitedly when disturbed, and unlike all the 
other creatures of the underworld did not seek to hide. Instead, 


* 


rr ‘ys 
nb 


FIG. 18. REMARKABLE INSECT FORMS, CHIEFLY NEW 
Found in the surface of a Tropical Yard of Jungle. 


a, An unknown form, beetle, roach or cricket. 6, The worker of a new genus and species of 
ant, Blepharidatta brasiliensis Wheeler, an extraordinary form, with small-eyed workers fitted 
for a subterranean life. The general structure is very simple and primitive. d. Pseudoscorpion, 
or false scorpion, a member of a compact, widely distributed family of Arachnida, with a pro- 
nounced superficial resemblance to true scorpions. /, Unknown, even as to order. g, A mite, 
one of the vast host of bete rouge, or maquins, the most troublesome pests of tropical jungles. 


Zoologica Vol. II, No. 4. Face page 113 


1916] Beebe: Fauna of Jungle Debris 113 


they bravely sought open spaces, walking slowly and feeling 
ahead with their great pincer-tipped arms, which they bran- 
dished with the greatest ease, although these weapons were as 
long as their entire bodies. When really alarmed, they scurried 
backward, holding up their chelae in readiness. Their bodies 
were whitish, but their arms and pincers deep reddish brown. 
While there were several species, these superficially fell into two 
distinct types. The most abundant kind was pot-bellied, with 
heavy chelae, and was slow in movement. The other had a nar- 
row, lighter body and very delicate slender chelae, and ran with 
great speed when alarmed. These, however, always ran for- 
ward, not backward like the others. 


Harvest men were represented by a single daddy-long-legs 
which looked decidedly out of place among this dense debris. I 
rather fancy he was strolling on the surface when my onslaught 
bagged him and his surroundings. 


Very small and very pale colored spiders lived in the middle 
layer in fair numbers. We saw about two score altogether. They 
were usually slow or moderately gaited, like their more abundant 
relatives, the mites. Only twice did we see a spider dash off 
with any of the speed which characterized those which lived in 
the jungle above ground. 


Next to the ants the mites and ticks were the most abun- 
dant organisms. Hardly a leaf or bit of mold was free from 
them. We could have gathered hundreds. They were of many 
species and all colors, red, brown, purple, black and flesh. Some 
were naked and shining, others clothed in bristly hairs to their 
very feet. All were repulsive, slow, and so awkward that it 
was inexplicable how creatures with such lack of correlation 
could ever manage to find food, much less a mate. They were 
always crawling slowly along, tumbling over every obstacle in 
their path. Ticks were much rarer than mites. 


Numbers of very simple insects were common. Silverfish 
or Thysanura of several species ran like active little ghosts out 
of their hiding places and scurried swiftly to another which they 
fancied safer.. Their nimble movements made them exceedingly 
difficult to capture. Collembolas, almost equally primitive, were 
usually white, but now and then a purple one appeared. Not 


114 Zoologica: N. Y. Zoological Society US 


only were they capable of active running, but when the brush 
wet with alcohol was about to touch them, they leaped to a dis- 
tance of twenty to thirty times their own length. Again and 
again this enabled them to escape. When they landed they re- 
mained motionless for some time and were most difficult to 
discover. Among the specimens collected were Campoclea, and 
many individuals of Collembola, belonging at least to three dif- 
ferent genera Jsotoma, Lepidocyrtus and Schottella. 


Termites, or “white ants,” lived in small colonies of six to 
thirteen individuals in small twigs, in the upper layer of debris. 
Sometimes they seemed to be living in close association with 
real ants with no signs of hostility on either side. 


A very few immature wood roaches represented the order 
Orthoptera, while the Hemiptera or true bugs had only a slightly 
better showing. Earlier stages of these insects lived in the 
middle layer, while those in the upper were quite adult and were 
ready to fly. 


Beetles of small size were abundant and of numerous 
species. Of about fifty which I gathered, about sixty per cent 
were rove beetles. All the others were slow travellers, or on 
discovery pretended to be dead, but the rove beetles were very 
agile, and never lost any opportunity of trying to escape capture. 
There were members of Rhynchophora of the Tribe Tylodini; 
of the Families Thorictidae, Phalacridae, Pselaphidae and 
Tenebrionidae. Also of Clivina, Secyclonaenus, Oxytelus, and 
Platystethus; Staphylinidae were, as I have indicated, by far 
the most numerous. 


Some tiny flies had apparently just emerged from their 
pupae in the upper layer, these being the only representatives 
of their order, while of the Lepidoptera there were only two 
small moths among the dry leaves of the top stratum. 


Ants were the most abundant form of life, both in numbers 
and species. They lived in the upper layers and with the excep- 
tion of the great, black, solitary fellows who apparently had 
been walking about on the top of the leaf stratum, all were of 
small size. Their colonies were apparently complete but very 
small, a very small twig being packed full of individuals from 
six to fourteen in number with a half dozen pupae. 


FIG. 19. REMARKABLE INSECT FORMS, CHIEFLY NEW 
Found in the surface of a Tropical Yard of Jungle 


«, Unknown form. e, An unknown form, possibly the remarkable larva of some Myrmeleonid 
species, related to the Ant-lions. h, The worker of a new genus and species of ant, Glamyro- 
myrmex beebei Wheeler. This is also a subterranean form, living in small colonies in tiny twigs. 
In the colony from which the species was described, there were only three workers, three females, 
and two males. 


Zoologica Vol. II, No. 4. Face page 115 


1916] Beebe: Fauna of Jungle Debris 115 


* Finally mollusks were found in small numbers, all very 
small, some with flat shells, others with steeply turreted ones. 
These were young specimens of two species of Leptinaria and 
several very young Polita, or Vitrea as it is more generally 
called. 


In addition to all these was a host of unknown forms, im- 
mature or in some unrecognizable early stage of development. 
Some had huge jaws and the body encircled with a dense 
chevaux-de-frise of horny, frond-like spikes. Others were so 
simple that their relationships could only be guessed at. 


One thing was evident early in my exploration. I was hay- 
ing to do with a world of small people. No insects of large 
size were in any layer of the debris. The largest would be very 
small in comparison with a May beetle. Another fact which 
impressed me was the durability of chitin. The remains of 
beetles, considering the rareness of living ones, was remarkable. 
The hard wing cases, the thorax armor, the segments of wasps, 
eyeless head masks, all these still remained perfect in shape and 
vivid in color. Even in the deepest layers where all else had 
disintegrated and returned to the elements these shards of death 
were as new. 


Day after day as I worked with my face close to the mold, 
I was constantly aware of the keen, strong, pungent odor. It 
hinted of the age-old dissolution, century after century, which 
had been going on. Leaves had fallen, not in a sudden autumnal 
down-pour, but in a never ending drift, day after day, month 
after month. With a daily rain for moisture, with a tempera- 
ture of three figures for the quicker increase of bacteria, and 
an excess of humidity to foster quick decay, the jungle floor was 
indeed a laboratory of vital work—where only analytic chem- 
istry was allowed full sway, and the mystery of synthetic life 
was ever handicapped and ever a mystery. 


Before the vessel docked we had completed our task and 
had secured over five hundred creatures from this lesser cosmos. 
At least twice as many remained, but in making calculations I 
estimated that the mold had sheltered a thousand organisms that 
were plainly visible to the eve. 


116 Zoologica: N. Y. Zoological Society aie! 


When I had corked my last vial and the steward had re- 
moved the final pile of shredded debris, I leaned back and 
thought of the thousand little creatures in my scant four square 
feet of mold. Then there came to mind a square mile of jungle 
floor with its thin layer of fallen leaves sheltering many more 
than six billion of these creatures. Then I recalled the three 
thousand straight miles of continuous jungle which had lain 
westward up the course of the Amazon, and of the hundreds of 
miles of wonderful unbroken forest north and south. My mind 
faltered before the vision of the unnamable numerals of this 
uncharted census, of the insurgence of life which this thought 
embraced. It seemed quite clear that no Tyrant Antwren need 
ever go hungry, as long as he had strength to turn a leaf. 


Leaving out the hints of vertebrates which in numbers were 
almost negligible, the lower types of creatures may roughly be 
grouped as follows: 


Flatworms (Platyhelminthes) uw... 2% 
Roundworms (Nemathelminthes) 0... 2% 
rie VVOLrmc CV erImMes)® 2) 2 sees ee a eee So 8% 
Myriapods iGMyniapoda)....2-6 225) a 6% 
Scorpions ((Scorpionidd))) t2.. 9 ka 1% 
Pseudoscorpions (Pseudoscorpionida) — 8% 
Harvest. men (Phalangida) 42222 Ae eho 
Spiders: (Araneida) a... ee 3% 
Mites.and’ Picks <(Acarida)) {238 2-2 ae 14% 
Silvertish. Glhysanura)) oso oo ee ee 2% 
Collembola (Collembola) 2.22... 3% 
Vermites:-Clsepveray) 5 ae oe ee 10% 
Roaches (Orthoptera) — Bed iin AL Ni 1% 
Buss Giiemipteray San. aCe te 2% 
beetles: (Coleoptera 7 2... ee eee 10% 
WhiesCDipterd:):> Soc cis a ian ht) ene 1% 
Noths; «(bepidopteray A... ee Ne eee 1% 


Ants CHYyYMenOpveray”. 4 see ec ee eee 30% 


1916] Beebe: Fauna of Jungle Debris 117 


- As shown by this list, ants were the dominant form of life, 
so I have chosen to mention these in detail as representative of 
the interest of this method of investigation. They have been 
thoroughly worked out by Prof. Wheeler,* and the unexpected 
result of this mode of intensive study is well illustrated by a 
paragraph from one of Prof. Wheeler’s letters. Referring to 
the nineteen vials of ants which I had sent him he says: “I have 
just found time to mount them up and to my surprise discover 
among them representatives of two new and remarkable genera! 
That you should have found these is indeed remarkable, because 
Professor Goeldi, formerly the director of the Para Museum, 
collected ants very assiduously in that region and sent them to 
Forel for description. Moreover, one of my students, Mr. Wil- 
liam M. Mann, who has been with me several years, collected 
very extensively in Brazil and recently enumerated all the known 
Brazilian forms with a description of the new species he had 
taken, and neither of these men came across the two very pecu- 
liar little ants which you found. I take it that they did not work 
in the leaf mould as you did and that probably when other col- 
lectors adopt your method an extensive ant fauna will be un- 
earthed even in Brazil, which has been pretty well worked for 
ants within recent years. ...I have named the two new genera 
and species Blepharidatta brasiliensis and Glamyromyrmex 
beebe.” 


The seventeen species of ants which I discovered in this 
cour square feet of jungle mould are as follows: 
1. Pachycondyla harpax Fabr. (workers). 


2. Huponera (Trachymesopus) stigma Fabr. (work- 
ers). 


Ponera opaciceps Mayr. (workers). 
Anochetus mayri Emery (dedlated female). 


oe 


Solenopsis subtilis Emery (workers, males, dea 
lated female). 

§. Crematogaster victima F. Smith. var. (dedlated 

female). 


*Two new Genera of Myrmicine Ants from Brazil, Bull. Mus. Comp. 
Zool. Harvard, LIX, No. 7. 


118 Zoologica: N. Y. Zoological Society [II;4 


7. Pheidole flavens Roger subsp. eaxigua Emery 
(soldiers, workers, males, dedlated female). 

8. Pheidole subarmata Mayr. (workers, dedlated 
temale). 

9. Trachymyrmex sp. (dedlated headless female). 

10. Cyphomyrmex rimosus Spin. (dedlated female). 

11. Rhopalothrix (Octostruma) balzani Emery (work- 
ers, dedlated female). 

12. Strumigenys subedentata Mayr. (dedlated female). 

13. Prenolepis steinheili Forel (workers, males). 

14. Rhizomyrma goeldii Forel (workers). 

15. Camponotus (Myrmothrix) abdominalis Fabr. var. 
(dedlated female). 

16. Blepharidatta brasiliensis Wheeler. 

17. Glamyromyrmex beebet Wheeler. 


The solitary deflated females of the species of numbers 4, 
6,9, 10, and 15 were evidently establishing colonies. At least 
eight of the species, those of the genera 2, 3, 5, 11, 12, 14, and 
the two new genera 16 and 17 are hypogaeic or subterranean 
ants, with small-eyed workers. With the exception of numbers 
1 and 15, all of the species are small or very small. 


Taking ants alone, we thus find that in numbers they formed 
about thirty per cent of the visible fauna of the jungle mould. 
With the exception of the two species all were adapted by their 
small size to life in the leaf mould, and fifty per cent were struc- 
turally fitted for subterranean existence. 


Rie 


I have made a single interesting comparison between 
this fauna of four square feet of tropical jungle debris and that 
of a corresponding area in a temperate and an Arctic latitude. 
In the tropical material, as I have stated, we found, at the very 
lowest estimate, one thousand visible organisms. In four square 
feet of leaves and moss from an uncleared area in the woods 
of the New York Zoological Park were two hundred and sixty 
creatures. From a slightly larger area, approximately a square 
yard, of tundra moss from Labrador, twenty-seven living organ- 
isms were unearthed. This last material consisted chiefly of 


1916] Beebe: Fauna of Jungle Debris og 


white reindeer moss, near a grove of fir trees from the North 
West River on Lake Melville, ninety miles directly west of Rigo- 
let up the Hamilton Inlet. For this I am indebted to A. Sheard, 
Esq., of the Grenfell Association, who was kind enough to gather 
it personally for me. 


The value of this comparison is, of course, relatively super- 
ficial, but nevertheless it is not without interest and should stim- 
ulate effort in this comparatively unworked ecological field.* 


Fauna of Four Square Feet (New York) 

Temperate 
ieee Worms JC \CLMeS): ce ke Moneta ah AE 
Myriopoads (Myriopoda) «20s ee. Brees eet 10% 
Pseudoscorpions (Pseudoscorpionida) 1% 
Pfarvest vena Phalangigda) sos. 0 Soka ee 4G 
Sere@ers (ewranel@ain st tee Se ee ee Lae 
WiitessanGd. Hicks. (AeaATICay 6.0 oe) et 1% 
‘SBE es IS avt CAB) RESET a ehh Weare ete ee cet aS ame fone on Oe . 8% 
Bugs (Hemiptera) Te ane Neha SC ewte. 6 8% 
beetless (@aleanueray Ga: so eA at ewe % 
iIMoths*(Lepidoptera). 28 oe DA ee cee mt 5 
PSO Ayr en OPGERA)) keene hb cee a a 40% 


The lists speak for themselves, the interesting facts being 
the marked diminution in number of general groups, as well as 
species and individuals from the tropics northward. The dom- 
inance of ants in both temperate and tropical cases is worthy of 
notice, and the remarkable number of true worms in the north 
and of mites and ticks in the south. In none of the lists are 
eggs or cocoons included. 


Attempts to identify the tropical organisms have shown 
how little knowledge we have of the life histories of these in- 
vertebrates. It was indeed fortunate when even a genus or sub- 
family could be told. The lack of a great central museum, 
library and collection of types in our country is keenly felt, as 
well as the handicap of the general habit of publishing new 
species in all sorts of magazines and periodicals, wholly unre- 
lated except by the widest of zoological bonds. 


*Consult W. L. McAtee, Science, N. S., Vol. XXVI, pp 447-449, and N 
Banks, id. p 637. 


P= Z00L0GICA 
_ SCIENTIFIC CONTRIBUTIONS OF THE 
- NEW YORK ZOOLOGICAL SOCIETY 


ian insti, 
_ i 
im) r . 


VOLUME II, NUMBER 5. 


THE GAFF-TOPSAIL 
(Felichihys felis) 
A SEA CATFISH THAT CARRIES ITS EGGS 
IN ITS MOUTH 
BY 
E. W. GUDGER 


STATE NORMAL COLLEGE 
GREENSBORO, N. C. 


Me PUBLISHED BY THE. SOCIETY = 
(fae ZOOLOGICAL: PARK, NEW YORK = 


iad 


ee ees : AucusT, 1916 


» Neu Hork Foviogical Bovey 


General Office, 111 Broadway, New York City 
_OFFICERS OF THE SOCIETY 


President 
_ Henry FAIRFIELD OSBORN. 


Hirst Vice-President Second Vice-President 

MADISON GRANT. . FRANK K. SrurGIis. 
Greasurer . Asst. Greasurer 2 

_PrErRcy R. PYNE, FARMERS’ LOAN & TRUST Co. 


e Executive Committee 
es MADISON GRANT, Chairman. 
Percy R. PYNE, | LISPENARD ‘STEWART, 


WILLIAM WHITE NILEs, WATSON B. DICKERMAN, - 
WM. PIERSON HAMILTON, ANTHONY R. KUSER, 3 Bide 
FRANK K. STURGIS, ' HENRY FAIRFIELD OSBORN,  —— 

ce ex-of ficio. 


Enditorial Committee 
HENRY FAIRFIELD OSBORN, Chairman. 
WILLIAM T. HORNADAY, ~~ CHARLES H. TOWNSEND. 


Anditing Committee Sa Aa 
WILLIAM WHITE NILES, Chairman. es 
H. CASIMIR DE RHAM, LISPENARD STEWART. 


General Officers 
Director of the Zoological Park: WILLIAM T. HORNADAY. 
Director of the Aquartum: CHARLES H. TOWNSEND.- 
Prosector: DR. GEORGE S. HUNTINGTON. 
Architect: C. GRANT LA FARGE. 
Consulting Engineer: H. DE B. PARSONS. 
Bursar: R. L. CERERO. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 5. 


THE GAFF-TOPSAIL 
(Felichthys felis) 
A SEA CATFISH THAT CARRIES ITS EGGS 
IN ITS MOUTH 


BY 


E. W. GUDGER 
STATE NORMAL COLLEGE 
GREENSBORO, N. C. 


PWR Ss Bee D BY THE po 0 et Co Ui cae ss 
tite Z, © Or O-GlC AL PARK, NEW -YOR-K 


AUGUST, 1916 


CONTENTE 


INTRODUCTION 


THE SEARCH FOR THE GAFF-TOPSAIL 

Historical Account, 126 
Data Obtained in 1906, 129 

Structure of Spent Ovaries, 129 
The Search Continued, June-July, 1907, 130 

The Structure of Ripe Ovaries, 130 

The Finding of the Fish with Eggs in the Mouth, 131 
Further Search, 1908-12, 135 

Difficulties Due to Heavy Rains, 136 

Effects of Cold Weather, 137 

Difficulties Due to Inexplicable Causes, 138 


THE NATURAL HISTORY OF FELICHTHYS FELIS 
Description of the Gaff-topsail Catfish, 138 
History of the Fish, 139 
Habitat, 140 
Swimming Habits, 142 
Behavior When Caught in a Net, 142 
Defensive Habits, 143 
Food and Feeding, 144 
Parasites, 145 
Use as Food, 145 
Sounds Made by the Fish, 146 
Size of Breeding Females, 147 
Size of Incubating Males, 147 
How the Eggs are Carried, 147 
Size of Mouth Cavity, 148 
Size and Structure of Skull, 148 
Size of Eggs and Number Carried, 150 
Attempts to Hatch Eggs Artificially and the Difficulties 

Met With, 150 
How the Embryological Series Has Been Obtained, 155 


CONCLUSION 


Fig. 


Fig. 


Fig. 


ig, 


Fig. 


Fig. 
Fig. 


Fig. 
Fig. 
Fig. 


Fig. 


Fig. 


20. 


21. 


22. 


23. 


24. 


25. 
26. 


Bilis 
28. 
29. 


30. 


dl. 


FIGURES 


Lateral view of a female Gaff-topsail Catfish, 


1914 inches long, drawn from life Frontispiece 


Photograph of the head of an egg-carrying male 
Gaff-topsail, showing the depressed floor of the 
mouth and the distended gill-covers .............. facing 


Pen sketch from above of a Gaff-topsail, 1744 
inches long. The large head and prominent gill- 
covers give the fish a marked tadpole shape ......... 


Instantaneous photograph of eggs with larvae, 


Marcgrave’s Gaff-topsail. The earliest known 
freTesot tars shishs (Cl G48) 2a ee ee eee facing 


Bloch’s figure of the Gaff-topsail (1794) facing 


Dorsal aspect of the cement cast of the mouth 
of the male fish carrying 55 eggs ow... facing 


Lateral view. of the samevcast, . 0 te facing 
Dorsal view of the skull of Felichthys felis facing 


Photograph of the ventral surface of the same 
skull showing the “Crucifix in the Catfish.” 
CNatural sive) i. ste ee 6 ee facing 


(A. & B.). An instantaneous photograph of the 
dorsal and ventral surfaces of a little catfish 
which has just burst its shell and thrust out its 
HROOIG Pe Ae Ben Led Ee, soe at tA ee ee facing 


An instantaneous photograph of larvae ‘‘skat- 
ing” on their yolk-sacs on the bottom of an aqua- 
1D 00 ipl, aieiine MOI eee ee Boat, Ec ts in ee facing 


133 


133 


134 


139 
139 


147 
147 
148 


150 


152 


152 


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THE GAFF-TOPSAIL 
(Felichthys felis) 


A SEA CATFISH THAT CARRIES ITS EGGS 
IN ITS MOUTH’ 


By E. W. GUDGER, 
STATE NORMAL COLLEGE, GREENSBORO, N. C. 


INTRODUCTION. 


At the 1908 meeting of the North Carolina Academy of 
Science, I gave a short paper on the oral gestation of this fish, 
but, not being ready to publish, the title only appeared in the 
proceedings published in Science (vol. XXVII, p. 991) and in 
the Journal Elisha Mitchell Scientific Society (vol. XXIV, p. 50). 


For the Washington meeting of the American Association for 
the Advancement of Science in 1911, I prepared, but was unable 
to give in Section F., a paper bearing practically the same title. 
However, there was published in Science (1912, vol. XXXV, p. 
192) an abstract, the data of which forms the closing paragraph 
of this paper. 


Since the more technical data obtained in this research will 
be presented in a series of papers which will require several ' 
years for working up and for publication, it has seemed best 
to give without further delay an account of the search for incu- 
bating males of the Gaff-topsail and of the difficulties met with 
in obtaining and hatching the eggs, and to present the general 
data concerning this very interesting phenomenon together with 
the natural history of the fish. 


1Contribution from United States Fisheries Biological Station, Beaufort, N. C. 
Published by permission of the Commissioner of Fisheries. 


*] have pleasure in recording here the hearty thanks due my friends among 
the fishermen of Beaufort, N. C., for testimony as to the carrying of eggs by 
the fish, for advice as to where and how to take the fish, for specimens, and 
for a vast deal of arduous manual labor involved in the very extensive seinings 
carried on for six years. My especial thanks are due to Jack and Southey Mades 
and Henry Congleton; to W. E., C. F., and J. W. Wheatley; and to Wilbur 
Whitehurst, Arthur Newkirk, Pete and Billy Garner, Ed Simpson and Walter 
Longest. 


\ 


126 Zoologica: N. Y. Zoological Society [lis 
THE SEARCH FOR THE GAFF-TOPSAIL. 


HISTORICAL ACCOUNT. 


On July 26, 1906, while at Cape Hatteras, N. C., in company 
with Mr. N. F. Jennett, a pound-net fisherman of Pamlico Sound 
at that place, I was informed that, on the preceding day while 
looking over the fishes brought in from his nets, Mr. Jennett 
had found in his hands some catfish eggs having young on them. 
By their flat barbels, he had readily identified these little fish as 
Gaff-topsails in contradistinction to the other sea catfish. The 
eggs, he thought, were about the size of peas or slightly larger, 
but whether they came out of the mouth or the vent he could 
not say. 


On returning to the laboratory of the Bureau of Fisheries at 
Beaufort, N. C. to which I was at that time temporarily attached 
as investigator engaged in research work in fish embryology, I 
began to look up the literature with a special reference to the 
egg-carrying habits of the catfishes and of other fishes. 


This search has been kept up ever since, but the literature has 
been found to be so voluminous that only the data gathered in 
1906 will at this time be reviewed. This is given in brief form 
that it may afford the reader something of an historical setting 
for the data to be presented herein. However, it is my purpose to 
publish at some future time a paper now in MS. reviewing the 
literature of the world on oral gestation in teleostean fishes. 


Evermann and Bean, in 1898, in their ‘“‘“Report on the Indian 
River and its Fishes,” say of Galeichthys milberti, a near rela- 
tive of the Gaff-topsail, that “Mr. Stypman of Stuart, Florida, 
assures us that eggs are never found in this catfish, but that the 
young are brought forth alive. He says during March the adult 
females are found filled with well-developed young, each rolled 
up in a ball, and the various balls connected in a long string. He 
thinks they hatch out very much like the sawfish. Others give the 
same information and it sems certain that this species is ovo- 
viviparous.” 


Evermann and Goldsborough in 1902 in their ‘‘Report on 
Fishes Collected in Mexico and Central America” describe an- 
other allied but heretofore unknown form, Conorynchos nelsoni, 


1916] Gudger: The Gaff-topsail i WaT 


which carries its young in its mouth. This was a freshwater 
catfish taken in southeastern Mexico, from the Rio Usumacinto, 
125 miles from its mouth. Two fish were collected, both males; 
of these one, 13.5 inches long, had one egg in its mouth, the 
other, 16 inches long, had thirty-nine eggs in the oral cavity at 
the time of its capture. Eight or ten other fish were captured 
at the same time but none carried eggs. All the eggs of the 
second fish save four were lost. These four after being in 
alcohol for two years averaged 10/16 of an inch in diameter. 


Jeffries Wyman, while United States Consul at Paramaribo, 
Surinam, South America, had his attention called in 1857 to cer- 
tain Siluroid fishes belonging either to the genus Bagrus, or one 
closely allied, which were said to carry their eggs in their 
mouths. These reports he verified by visits to the markets 
where these fish were offered for sale for food. He found either 
eggs or larvae in the mouths of various specimens of jarra- 
bakka and njinge-njinge, and was assured by the negro fisher- 
man that koepra and makrede together with one or two forms 
had the same habit. The egg-carriers in all the fishes examined 
by him were males. The eggs of jarra-bakka ranged up to three- 
fourths of an inch in diameter. Of njinge-njinge, eight speci- 
mens nine inches long were examined. The eggs were in differ- 
ent stages of development, and the number contained in the 
mouth varied between twenty and thirty. 


Giinther in 1864 noted this curious habit in specimens of 
Arius fissus from Cayenne in the same region of South America. 
In fishes six to seven inches long, all males, he found some 
twenty eggs about the size of a pea, having on them advanced 
embryos. This habit, he observed, is not uncommon among 
American Siluroids. 


In 1866, Boake described oral gestation in two species of 
Ceylon catfishes of the genus Arius. Their ova were “large” (he 
seems to have made no measurements of either ova or fish) and 
immediately after deposition are “taken up either by the fish 
that has laid them or by another of the same species, and, not 
swallowed but kept in the mouth until they are hatched and able 
to take care of themselves, a period of some weeks.’ Later he 
ascertained (presumably by dissection) that the egg-carrying 
fish was the male. 


128 Zoologica: N. Y. Zoological Society [1135 


Boake sent to England some specimens of these fishes (two 
males and one female) which, coming into the hands of William 
Turner, were described by him in 1867. He confirmed Boake in 
all respects, and noted that one of his male specimens had ten 
eggs in its mouth, whereas one of Boake’s specimens had 
thirteen. The other male, like the female, had the oral cav- 
ity empty. The “large” ova were about the size of grapes or 
small cherries and possessed embryos well along in development, 
measuring 1/2 to 7/10 of an inch in length. Some of Boake’s 
specimens reached Giinther also and he in 1866 described and 
named them. He commented on the habit of the male in carry- 
ing the large eggs in the spacious cavity of the mouth, and com- 
pared the fish with Arius fissus from South America, previously 
(1864) described by him. 


In 1889, Day described the oral gestation of the males of two 
genera of Indian catfishes, Aviws and Osteogeniosus. The former 
had eggs averaging .5 to .6 of an inch in diameter, and each 
carried as many as fifteen to twenty eggs in the mouth. The 
eggs were in stages of development varying from very early 
embryos to larvae nearly ready to take care of themselves. 


Giinther in his “The Study of Fishes” (1880), p. 160, has 
the following brief reference: “The Siluroid genus Avius, the 
males of which take care of their progeny, produces ova 
5-10 mm. in diameter.” On the same page, he gives a figure of 
the ovum of the Avizs boakei marked ‘‘natural size” but meas- 
ing 14 mm. in diameter. Finally Jordan in his “Guide to the 
Study of Fishes,” (1905), vol. 1, p. 128, writes: ‘In certain sea 
catfishes (Galeichthys, Conorynchos) the male carries the eggs 
in his mouth, thus protecting them from attacks of other fishes.” 
Again, in vol. II, p. 179, he says: “In most or all of the ‘sea 
catfish, the eggs as large as small peas are taken in the mouth 
of the male and there cared for until hatched.” 


Having exhausted the literature in the library of the labora- 
tory, I turned to the fishermen of Beaufort and began the tak- 
ing of testimony, and was surprised to find how many of them 
had observed in a general way and now gave such unanimous 
testimony that the eggs are carried in the mouth. One man 
thought that the eggs came out in strings, another was not sure 
on this point, but his best recollection was that this was true, 


1916] Gudger: The Gaff-topsail 129 


while another believed that they were hung in bunches in the 
roof of the mouth. All united in declaring that the eggs came 
out of the mouth and not out of the vent and that in size they 
were about equal to peas. One man phrased it that ‘‘The sea 
cat spits its young out of its mouth”; and all affirmed that when 
the fish are struck on the head or thrown into the boats, the 
eggs would fly out of their mouths. Boake credits the Ceylon 
fisherman with reporting a similar phenomenon when the egg- 
carrying Arius of that island is captured, the eggs being voided 
in such numbers that they are gathered from the bottoms of the 
boats and used for food. 


Several other fishermen testified that the eggs are always 
carried in the mouth, sometimes as many as two handfuls, but 
not necessarily of the same age, and that they may be found in 
May and June. One man declared that the eggs were sometimes 
as large as the yolk of a small hen’s egg and that they were “of 
a pinkish color between white and blood-red.” 


DATA OBTAINED IN 1906. 


Structure of Spent Ovaries. 


About this time, August 3-6, 1906, there were found in the lab- 
oratory pound net considerable numbers of Gaff-topsail catfish, 
all of which were brought in and examined. The ovaries pre- 
sented some very interesting structures, but no extended des- 
cription of them will be gone into here. They were of the 
ordinary teleostean type, bifurcated in front, but united behind 
to form the short oviduct which opens out in the urinogenital 
pore behind the anus. In the ovaries, only the anterior region 
bore eggs of any size, some of them being as large as small 
peas. Each of these eggs was enclosed in an investing follicle 
richly vascularized and was carried on a short pedicel. The 
middle section had only pedicels from which the eggs had 
broken away—the follicles having disappeared probably by re- 
sorption—together with small, almost microscopic ova in be- 
tween. The posterior or oviducal part was very curiously 
plicated or folded longitudinally like the oesophagus of the frog 
and so abundantly vascularized that while alive it was of a rich 
beef-steak-red color. 


130 Zoologica: N. Y. Zoological Society [Ils5 


Until August 13, our pound net daily gave us several of these 
catfish, the females predominating. These fish were all dissected 
and from the reduced condition of the reproductive organs— 
the ovaries running 2 to 214 inches in length—it was clear that 
the breeding season was long over and that no further work 
could be done during this summer. Fig. 20, frontispiece, shows 
one of these spent females, but it might well represent a non- 
breeding male since there is nothing to distinguish the two 
sexes at any time other than the breeding season. 


The structure of the ovary of Felichthys, in conjunction with 
the reported habits of Galeichthys, seemed to indicate that pos- 
sibly the fish was viviparous, that the eggs might descend into 
the oviduct and there remain until hatched, nourished by trans- 
fusion of oxygen and food materials from the richly vascular- 
ized walls of the oviduct by which they might become partly 
enveloped. On the other hand there were the positive state- 
ments of a number of fishermen, men who presumably knew what 
they were talking about, that the fish incubated the eggs in its 
mouth. In this state of uncertainty, the question, owing to the 
lack of material, had to go over until the next summer. 


THE SEARCH CONTINUED—JUNE-JULY, 1907. 
Structure of Ripe Ovaries. 


As soon as it became known that I had reached the Beaufort 
laboratory in June, 1907, the fishermen began to look out for 
catfish material for me. On June 4 they brought in a 21-inch 
female Gaff-topsail catfish which they had split open from snout 
to anus without finding any eggs. They had even cut into the 
ovary, following the idea, which had been advanced to them the 
previous summer that the eggs were possibly carried there, but 
had found in this only eggs still bound up in their stalked fol- 
licles. This ovary was about 41% inches in length and was 
crowded with eggs in size from 10 mm. down. Having never 
before seen such enormous eggs in a teleost and finding ruptured 
follicles from which eggs had been evaginated not many days 
before, I came to the conclusion that these eggs must be ripe 
and that 10 mm. was the normal size for such. 


1916] Gudger: The Gaff-topsail awl 


Two days later the same men brought in two Gaff-topsails 
which had been caught but a few hours before and which were 
unmutilated. One fish, 1634 inches long, proved on dissection to 
be a female with ovaries 31% inches in length from tip to opening 
of oviduct. These organs were much distended with eggs 15 
mm. in diameter, and occupied 50 to 60 per cent. of the body 
cavity. The second fish, also a female, was 21 inches long and 
had the most enormous and remarkable ovaries I had ever seen 
in any fish. They were 51% inches long, tightly distended with 
eggs from 18 to 20 mm. in diameter and occupied from 75 to 
85 per cent. of the body cavity. The other organs were very 
much reduced in size and crowded out of their normal position. 


On June 13 there was brought in from our fyke net a 22-inch 
female with an ovary 5 inches in length, which by its flabby 
condition showed plainly that the season’s eggs had been lately 
discharged. From all this mass of evidence it was clear that 
the breeding season of Felichthys was at hand, but, being in- 
tently occupied with the completion of another research in fish 
embryology, I was unable at the time to devote myself to this 
problem. 


A week later, my other investigation having ended, I was 
ready to take up this research; but our fyke net having ceased 
to yield specimens and the much-expected new pound net hav- 
ing failed to arrive, the fishermen were called on to help. 


The Finding of the Fish with Eggs in the Mouth. 


On June 22, we went to the Narrows of Newport River 
some seven miles northwest from the laboratory. Here where 
the river proper enters the estuary of the same name, there are 
oyster reefs at the edges of extensive mud flats. As the tides 
swirl around these reefs, they dig out deep holes and in these 
holes the catfish congregate at low tide when their mud flat feed- 
ing grounds are nearly laid bare. At the uppermost of these 
reefs, after several unsuccessful hauls elsewhere, a big catch 
of Gaff-topsails was made. The number of these was unfor- 
tunately not noted, though the lengths of thirty-two egg-carriers 
were recorded. Probably there were from sixty to seventy-five 
of them in all. 


132 Zoologica: N. Y. Zoological Society flr¢s5 


From the mouths of these catfish there were obtained between 
200 and 300 eggs. This is a minimum estimate, for, in the 
confusion and excitement due to such good fortune, no record 
was kept and afterward at the laboratory other eggs (to be 
described later), as they died, were put in the same bottles in 
which this day’s catch was preserved. The fact that all these 
save thirteen were put into killing fluids was due to this same 
confusion and excitement which was enhanced by the threshing 
about of sharks and rays and the splashing of a large number 
of small fishes in the bottom of our boat, together with my being 
called on every minute to receive a new lot of eggs. Fortunately 
at the very last of the haul it occurred to me to try to carry 
in to the laboratory one of the ovigerous males, and to make 
sure that the thirteen eggs contained in the mouth were not lost, 
this was loosely sewed up with a bit of small cord. Although 
the fish was then put into a bucket of water which was renewed 
from time to time, it died, but the eggs reached the laboratory in 
good condition and when put into running salt water soon re- 
covered and seemed perfectly normal. 


In this connection Wyman may be quoted that in the bagre 
of Surinam “In many instances the foetuses were still alive 
through the parent had been dead for many hours.” The con- 
text not indicating that the parent had been kept in water, it is 
probable that these larvae had lived because the moisture in the 
mouth of the parent had kept the egg-shells damp and hence 
permeable to oxygen. 


Omitting small numbers, one catfish gave up eleven eggs, an- 
other thirteen, another fifteen, and others sixteen, twenty, twen- 
ty-one, twenty-six, the total amounting as stated above, to be- 
tween 200 and 300 eggs. These eggs showed considerable 
variation in size, the extremes being from about 17 to 25 mm. 
in diameter, the average being from 18 to 20 mm. Their large 
size and great weight together with the extreme fluidity of their 
yolks, made them very difficult to handle for fear of hurting 
the embryos lying on the dorsal side. These embryos, as exam- 
ination later showed, were in different stages, from that wherein 
the future fish was in the form of an axial rod with forming 
eye vesicles, to the young in the black-eyed free-tailed stage 
about 17 mm. long. 


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FIG. 21. HEAD OF AN EGG-CARRYING MALE GAFF-TOPSAIL CATFISH 


The photograph shows the depressed floor of the mouth and the distended gill-covers 


Zoologica Vol. II. No. 5. Face page 133 


1916] Gudger: The Gaff-topsail 133 


FIG. 22. GAFF-TOPSAIL CATFISH FROM ABOVE 


The large head and prominent gill-covers give the fish a marked tadpole appearance. 
Pen sketch from a specimen 172 inches long. 


The method of procedure in obtaining the eggs was very sim- 
ple. The fishermen, standing in water and mud up to their 
waists “fished” in the net, keeping the lead line on the bottom 
to prevent the escape of any fish. As the net came in the fisher- 
men would grasp the fish just back of the head and in front 
of the dorsal spine, and keeping their mouths shut to prevent 
the escape of the eggs, would turn to the boat; then holding 
the fish with its tail upward, they would allow the eggs to fall 
or run out into a vessel of water. After being looked over for 
a minute, or counted in case there was a considerable number 
of them, they would be transferred to the killing fluid to make 
room for others. 


On this trip only some half dozen egg-carriers were dissected, 
but in every case the fish was ascertained to be a male. In all 
the testis was small, stringy, often almost insignificant in size, 
indicating that the breeding season was past. With the proof 
by dissection that the male is the carrier of the eggs, the Gaff- 
topsail catfish falls in line with all other egg-carrying catfishes. 


134 Zoologica: N. Y. Zoological Society files 


The eggs were loosely held in the mouth, some being pushed 
into the spaces between the branchial arches, but nowhere was 
there any evidence of arrangement. It was very noticeable that 
the mouth cavity, ordinarily so large as to be called enormous, 
was increased by a distension of the branchial region, but even 
more noticeably by a marked outpushing or rounding out of the 
whole hyoid and branchiostegal region. Instead of being flat or 
insunken as in most fishes and as in all the Gaff-topsails save 
ovigerous males, this region showed a rounded contour corre- 
sponding to the enlargement of the oral cavity, giving the fish a 
“double chin” appearance. 


Figure 21 is a photograph of the head of a living male Gaff- 
topsail carrying eggs in his mouth. Note the depressed hyoid 
region and the out-spread gill covers. The mouth is held some- 
what closed to prevent the escape of the eggs. Figure 22 is a 
pen drawing of a living Gaff-topsail, 1714 inches long seen from 
above. Attention is called to the great size of the head and 
especially to the width in the region of the gill covers. 


Several other collecting trips were made during the summer 
of 1907 and considerable numbers of eggs were obtained. In 
one batch of eggs the embryos averaged 20 to 25 mm. in length 
and fully 3 mm. wide from inside of eyes, while their tails were 
about half as long as the bodies. Black pigment was noticeable 
along the dorsal region, the caudal fin rays were visible, and 
the yolk blood-vascular system was well developed, giving the 
eggs a beautiful pink or reddish color. (Here recall the fisher- 
man’s description.) The heads of the little fish were deeply 
sunken in the yolk and even the tails occupied little grooves. 
In this connection an interesting correlation was noted. The 
heads of the little fish were all twisted, right or left, only one 
eye being visible, and likewise the tails were twisted right and 
left. If the right eye was sunk, then the tail was twisted to 
the right, and vice-versa. This may be seen by careful inspec- 
tion of figure 23 made from a photograph of the live eggs. How- 
ever, it seemed to be a matter of indifference to which side the 
body was bent, for of the 114 eggs on hand, fifty-four had the 
embryos bent to the right and sixty to the left. 

The last trip for 1907 was taken up Newport River on July 
18. Again former experiences were repeated, for no catfish 


FIG. 23. EGGS WITH LARVAE 


Showing the insunken heads, curled tails and prominent yolk circulation. 


From an instantaneous photograph. 


Zoologica Vol. II, No. 5. Face page 134 


1916] Gudger: The Gaff-topsail 135 


were taken until the mud bottom of the upper oyster reef was 
hauled. Here was secured one male, 18 inches long and from his 
mouth were taken 21 actively wriggling larvae, one of which 
was lost. One of these, of average size, died on its way to the 
laboratory (probably having been injured in being taken from 
its father’s mouth). After being hardened for 24 hours in 10 
per cent. formalin, it measured as follows: Length all over from 
point of snout to tip of upper lobe of caudal 57 mm.; width, 
between eyes outside to outside, 11.25 mm.; length of dorsal 
spine, 14.65 mm. The yolk was 18 mm. in diameter measured in 
the long axis of the fish, and 16.5 mm. in diameter at right angles 
to the above—the vertical measurement or depth of the fish was 
unfortunately not noted. On these fish the black stippling was 
quite thick on the head and along the dorsal region of the body. 
It was thickest at the roots of the dorsals and on the body it 
was arranged in distinct lines following the myomeres. Although 
the eyes were well along in development, the faint choroid slit 
could still be seen on the ventral side. The anal papilla showed 
as a projection in the center of a triangle formed by the pelvic 
and anal fins. In the nasal region, between the two orifices, a 
flap-like papilla-shaped organ was noticeable. 

Figure 31 shows these little fish in the stage of development 
referred to. It is from an instantaneous photograph made in 
direct sunlight. The larvae are as yet unable to lift their heavy 
yolks. Their mode of progression is by ‘“‘skating’”’ on their yolk 
sacs over the smooth bottom of the aquarium. 


FURTHER SEARCH, 1908-12. 


My summers during the years 1908-1912 were spent in the 
Fisheries Laboratory at Beaufort in strenuous endeavors to fill 
out my embryological series of the eggs of the Gaff-topsail and 
to find out if possible how the eggs are transferred. In this 
work the few successes were interspersed amid many failures. 
Had artificial fertilization been possible the first task would have 
been much lightened, and had the fish been small enough to 
keep in even large aquaria, the latter might have been possible. 
But as it was I was never able to get breeding males and females 
at the same time save once and then artificial fertilization failed ; 
while for the second point there was no tank in the laboratory 


136 Zoologica: N. Y. Zoological Society [11;5 


large enough to hold these considerable-sized fish. In addition 
there were other hindering causes which at times defeated all 
efforts.to collect the ovigerous males. A brief recital of these 
will enable the reader to form an idea of the great difficulties 
under which research in the habits of fishes is pursued. Later 
the difficulties met with in the effort to hatch the eggs will be 
discussed. 


Difficulties Due to Heavy Rains. 


In 1908 I reached Beaufort on the afternoon of May 28. On 
the following day 4.02 inches of rain fell. On May 22 it had 
rained 3 inches, and the total rainfall from May 22 to 29 was 
7.79 inches. The total rainfall for the month was 11.07 inches, 
being 8.05 inches above normal. Again, on July 9-10, 4.48 inches 
of rain fell in 24 hours, while in a similar period toward the 
close of the month the precipitation amounted to 5 inches. 

The result of all this abnormal downpour was that the water 
at the head of the estuary of Newport River was so freshened 
that the catfish, especially males with eggs, were driven out of 
the deep holes along the mud flats at the Narrows and dispersed 
in the lower and broader reaches of the estuary where it was 
exceedingly difficult to find them. Thus it came about that the 
first lot of eggs was not obtained until June 11, the second lot 
on June 15, and the third and last on July 7. Consequently, the 
batch of eggs gotten on the first of these dates was far too old 
to furnish the early stages so earnestly desired, since, by reason 
of a grant’ from the Carnegie Institution of Washington, an 
artist was at Beaufort to draw figures to illustrate the embry- 
ology of the fish. 

Again in 1912, the search for the Gaff-topsail was greatly 
hampered by heavy downpours. On May 22 (the day of my 
arrival at the laboratory) the rainfall was 1.31 inches, and the 
precipitation from May 6-22, inclusive, was 7.51 inches. This 
so freshened the estuary of Newport River that the catfish were 
driven into the lower harbor, and possibly into the ocean. At 
least none were taken by any drag-net fishermen visited in New- 
port River, while the menhaden fishermen reported the taking 
“outside” of considerable numbers—more than usual. 


1Figures 20, 22 and 28, reproduced in this paper, were drawn under this grant 
by Mr. E. A. Morrison. The photographs were all made by the author. 


1916] Gudger: The Gaff-topsail 137 


Effects of Cold Weather. 


In 1911, more strenuous efforts than ever were made to obtain 
the early eggs. Having heretofore always reached Beaufort 
after the beginning of the breeding season, I made two trips this 
year. It should be noted, however, that the spring of 1911 was 
a late and cold one, extending well into May. There was a light 
frost in Beaufort on May 8, while toward the 20th it turned 
suddenly quite warm. The cold weather greatly delayed the 
breeding season, while it was greatly accelerated beyond the 
normal when the warm weather came. 


On the first trip the laboratory was reached on May 13. On 
that day, and on the 15th, trips were made to our favorite fishing 
grounds. Here great numbers of catfish were taken, mainly 
large Gaff-topsails with enormous ovaries distended with many 
large eggs. While their bellies were tremendously swollen, their 
genital orifices were but little reddened, and no eggs could be 
obtained though vigorous efforts were made to spawn them. 


The males, though smaller than the females, were adult, but 
from none could milt be obtained. None of the males of the 
first day’s catch had ‘“‘double chins” indicative of a readiness to 
receive eggs. However, those of the second day’s collecting did 
have the depressed hyoid region, the throat enlargement, but 
none carried eggs nor could milt be obtained from any. 


On the trip of May 18, not a single Felichthys was taken. 
Evidently it was too early, the ripening of eggs and sperms hav- 
ing been greatly retarded by the cold weather. It was necessary 
for me now to return to my college duties, but on May 20 Mr. 
Henry D. Aller, Director of the Laboratory, seined for me, but 
without getting a single cat. 


On May 25, I returned to Beaufort and personally supervised 
another seining, from which were obtained the youngest lot of 
eggs but one ever gotten during the whole of this research. 
These eggs had on them blastoderms with forming embryos, but 
neither these nor any others ever taken showed the coveted seg- 
mentation stages. Further seinings brought in only older eggs, 
and failure and disappointment were the result of this expendi- 
ture of time and money. 


138 Zoologica: N. Y. Zoological Society pel Ress 


Difficulties Due to Inexplicable Causes. 


The most disheartening failures of all during this collecting 
work were those for which no cause could be assigned, for nei- 
ther rain nor cold weather interfered. At various times during 
all these six years’ work, but especially during the latter half of 
the season of 1910, trip after trip was made to all our hauling 
grounds, where in times past boatloads of catfish had been taken, 
but all were “water hauls,” few fish and no eggs being taken. 
At one period some six or eight trips, covering two weeks, did 
not bring in a single egg. To make these trips, it was necessary 
to leave the laboratory from 3 a. m. to 7 a. m., in order to reach 
the seining ground at or before low water. 


In the meantime many fishermen were visited. Some of these 
used seines 1,200 feet long and drifted over a mile of river. 
They caught few small catfish or none at all, and none carried 
eggs. A few undersized females had in their stomachs small 
gray holothurians, which are to be found “outside” only, and 
hence, it seems to be a possible conclusion that for some un- 
known reason the catfish had left the brackish river for the 
saltier ocean. 


THE NATURAL HISTORY OF FELICHTHYS FELIS. 
DESCRIPTION OF THE GAFF-TOPSAIL. 


Felichthys felis, (felis, cat; ichthys, fish), whose portrait 
forms the frontispiece of this paper, is one of the two kinds of 
sea catfish found at Beaufort, the other being the much smaller 
Galeichthys milberti previously referred to. The body is elon- 
gated, but, as figure 22 shows, very large in the head region, the 
greatest depth being at the anterior edge of the first dorsal fin. 
The nostril is double. The eye, which has a vertical pupil, sur- 
rounded by a red iris, is placed low on the side of the head and 
just above the insertion of the maxillary barbel. This latter is 
flat and very long, reaching almost to the anterior base of the 
pelvic fin. The pectoral and dorsal spines are continued in long 
filaments, and these, together with the long, flat maxillary bar- 
bels, are such marked features as to make it impossible to con- 
fuse the Gaff-topsail with any other catfish found in the salt or 


FIG. 24. MARCGRAVE’S GAFF-TOPSAIL 
The earliest known figure of this fish (1648). 


FIG. 25. BLOCH’S FIGURE OF THE GAFF-TOPSAIL (1794) 


Zoologica Vol. II. No. 5. Face page 139 


1916] Gudger: The Gaff-topsail 139 


fresh waters of the United States. The caudal fin is large and 
deeply forked, the upper lobe being slightly the larger. The 
color of the fish is a beautiful silvery blue, darker above and 
lighter below, best seen in lateral view. The fins, especially 
those on the ventral part of the body, show a reddish tinge as 
first noted by Mitchill (1815). 


HISTORY OF THE FISH. 


The discoverer of our fish seems to have been George Marc- 
grave, in whose Natural History of Brazil (1648) there is fig- 
ured and described a catfish with flat barbels and long filaments 
to dorsal and pectorals which is apparently the fish under con- 
sideration. Figure 24 is a photographic copy of Marcgrave’s 
figure. This figure, as I have shown elsewhere (Gudger, 1912), 
was probably painted by Marcgrave himself while in Brazil 
sometime between 1638 and 1644. It nas suffered many things 
at the hand of the engraver, who seems to have been one De Bray. 


His description may be translated: ‘This Bagre, though of 
another kind, is in size and shape like the preceding; but it has 
a beard made of four ray-like barbels, two of which are eight 
digits long and wide like straps, and two are short ones. To 
the dorsal fin there is [attached] a similar strap nine digits 
long, and behind the gills barbels of the same kind. The other 
[fins] are similar to those of the preceding fish.” 

Attention is called to the four barbels, the two maxillary ones 
being long and flat or strap-shaped; and to the strap-shaped 
(ligula) dorsal and pectoral filaments. Another interesting 
point is to be found in the rays of the dorsal fin. While Marc- 
grave in 1644 knew nothing of the use of fin rays in distinguish- 
ing the genera and species of fishes, he has portrayed his fish 
with one spinous and seven soft rays in its dorsal fin, the correct 
number. 

Marcgrave’s figure and description have been copied by a 
large number of the old writers. Without going into details 
there may be named: Piso (1658), Willughby (1686), Ruysch 
(1718), Johnston (1758), and Bonnaterre (1788). However, 
that greatest ichthyologist of them all, Marcus Elieser Bloch, 
in 1794 figured and described a sea catfish from Surinam which 


140 Zoologica: N. Y. Zoological Society pees 


he says is identical with Maregrave’s. Figure 25 here is a photo- 
graph of Bloch’s fish. He notes that the nostrils are double, 
that the oblong eyes with black pupils and red irises are near 
the angle of the mouth. His figure shows four barbels, the two 
maxillaries being long and flat; and also the long filaments to 
dorsal and pectoral fins. 


Doubt has been expressed as to the correctness of Bloch’s 
identification, but the Eigenmanns in their great monograph on 
South American catfishes (1890) have identified Bloch’s sea 
catfish with the Gaff-topsail. 


Bloch’s description gives the fin rays as follows: dorsal, I-8; 
pectoral, I-12; pelvic, 8; anal, 24. His figure has 26 rays in 
the anal, but he notes that Gronow counted 23 in the anal of an- 
other specimen; it also has I-7 in the dorsal which is correct, 
though his description says I-8; the figure likewise has 8 rays 
in the pelvic whereas the true number is 6. However, Bloch in 
1794 may be forgiven for a miscount of the fin rays in his figure 
when Jordan and Evermann (1900) in their figure 52, plate 
XXIII, have the dorsal fin I-6, and anal 22. 


The earliest American describer of the Gaff-topsail was Mit- 
chill in 1815, who took it in the waters of New York. Indeed 
he definitely gave this fish a place in zoological literature by his 
splendid description which, however, need not be repeated here. 
The name Felichthys felis, by which the Gaff-topsail is known 
today, was assigned by Jordan and Evermann in 1900. 


HABITAT. 


This fish is a sub-tropical form ranging as far north as Cape 
Cod, but is especially common along the South Atlantic and Gulf 
Coasts where it is abundant in brackish waters, for which it 
seems to have a predilection. Bloch as early as 1794 noted that, 
“This fish (the saltwater katfish) is found not only in Brasil 
but also in the great rivers of North America.” By this he prob- 
ably meant in the estuary mouths of these rivers which are 
brackish. The EKigenmanns note (1890) that it is found along 
the Atlantic coast of America from Cape Cod to Rio de Janeiro. 


The earliest account given of the occurrence of catfish in 
North Carolina coastal waters is found in Thomas Ash’s ‘‘Caro- 


1916] Gudger: The Gaff-topsail 141 


lina,” published at London in 1682. The reference to seamen 
indicates that the fish in question was a marine form, and, since 
(as will be shown later) the Gaff-topsail is the more abundant 
of the two marine Siluroids on our coast, we may conclude it to 
be Ash’s fish. “ Cat-fish, whose head and glaring eyes re- 
semble a Cat; it’s esteem’d a very good fish, it hath a sharp 
thorny Bone on its Back, which strikes at such as endeavor to 
take it; which by seamen is held venemous.”’ 


Again, John Lawson says (1714), “Catfish are round, black- 
ish fish with a Great Flat Head, a wide mouth, and no scales. 
They sometimes resemble Eels in taste. Both this sort and an- 
other that frequents the salt water, are very plentiful.” The 
“another sort that frequents salt water’? was in all probability 
the Gaff-topsail. Brickell (1737), whose data seems largely 
to have been taken from Lawson, does little else than repeat the 
statements above given. 

The first definite record of the occurrence of the Siluroid fish 
known as the Gaff-topsail in North Carolina waters was made 
by Yarrow in 1877. Since his day the fish has been well known 
and often recorded. 


At Beaufort Felichthys is taken everywhere in the “rivers,” 
which are really brackish estuaries. My best catches have been 
made at the very head of Newport estuary, within one mile of 
the limit of tide water, where at dead low water the density 
was 1.007. 


It is very abundant on both coasts of Florida; in the Indian 
River so much so as to be a great nuisance to the fishermen. It 
is also abundant in all the sound-like lagoons and the estuarine 
river mouths opening into the Gulf of Mexico. 

H. M. Smith (1907) says that the smaller relative of the Gaff- 
topsail, Galeichthys (weasel-fish) milberti, is the most abundant 
of the sea catfish at Beaufort, but I have not found it so in my 
many years of seining there. The Gaff-topsail is found in large 
schools, and I have often taken 100 or more at a haul, while my 
fishermen on one occasion caught a wagon-load, estimated at 
over 500, ranging from 20 to 25 inches in length. On the other 
hand I have never taken more than a half dozen at a time of the 
small-mouthed catfish. It seems to be a shy and possibly a rather 
solitary fish. 


142 Zoologica: N. Y. Zoological Society [I1;5 


SWIMMING HABITS. 


The Gaff-topsail is a bottom liver, and generally not a very 
rapid swimmer. The strong tail and deeply forked caudal fin 
might lead one to think to the contrary, but, if the fish is viewed 
from above (Fig. 22), it is readily seen that the enormous head 
parts would render it impracticable if not impossible for the 
fish to get up much speed. 


Although a bottom swimmer, nevertheless as the net comes 
in, the Gaff-topsail has the interesting habit of swimming near 
the surface of the water with the dorsal fin, or at any rate the 
filament, projecting above the water. This habit of carrying 
the dorsal fin and filament in an elevated position gives it its 
name, Gaff-topsail. DeKay as early as 1842 made note of this 
peculiar swimming habit. In Newport River at dead low water, 
when the fish, driven off the mud flats as the water lowers, col- 
lect in deep holes, this same habit may be noticed. 


Furthermore, for two weeks in the summer of 1910 I kept a 
12-inch Felichthys in a 4 by 6 foot wooden tank in the laboratory 
at Beaufort, and during this time it persisted in swimming at 
the surface of the water with its dorsal filament carried high 
out of water. At the same time two Galeichthys in the same 
tank as persistently swam at the bottom 6-8 inches below. Gaff- 
topsail larvae also show a marked tendency to swim at or near 
the surface of the water of their aquaria. Even more marked is 
their habit of ‘‘hanging’’ motionless at the surface, much as a 
frog does. 


BEHAVIOR WHEN CAUGHT IN A NET. 


When caught in a seine, the Gaff-topsail has the very annoying 
habit of rolling itself up and very effectively entangling its dor- 
sal and pectoral spines in the meshes. So firmly imbedded does 
it sometimes become that it is necessary to break its spines or 
to cut the net to get rid of it. 


It is also a great annoyance to the fishermen in another way. 
All fish give off a slimy mucus which is very destructive to nets, 
causing them to rot rapidly, but of all fish known to me the 
Gaff-topsail, when caught, gives off not only the most slime, but 


1916] Gudger: The Gaff-topsail 143 


the most tenacious. Only repeated washings and rubbings will 
take it off the hands, and it is almost impossible to get it off 
the nets. Fishing for and handling this catfish is a very nasty 
matter. 


DEFENSIVE HABITS. 


So far as I have been able to ascertain the Gaff-topsail has 
no Offensive habits. Twelve and fifteen-inch specimens kept in 
tanks with various other and smaller fishes showed no tendency 
to molest these latter. It is true that Felichthys is sometimes 
found with fish in its stomach, but it is not impossible that these 
were dead or at any rate disabled ones which were not able 
to escape the relatively slow moving catfish. Certain it is, as 
will be shown in the next section, that the food of this species 
is mainly crustacean. 


However, weapons of defense are present in the shape of 
dorsal and pectoral spines and are capably used. If the fish is 
caught and held by the tail it will swing violently and convul- 
sively to the right and left almost through an are of 180°, en- 
deavoring to strike with its pectoral spines. If picked up in- 
cautiously it will almost surely wound one. The only safe way 
to grasp it.is across the back of the head in front of the dorsal 
spine, the thumb on one side and the fingers on the other behind 
the pectoral fins. Held firmly thus (see Fig. 21), it is almost 
incapable of inflicting a wound. Such wounds, while quite pain- 
ful, are not especially dangerous, though bacteria carried in with 
the slime may set up an inflammation and the slime itself may 
possibly be toxic. 


These points were covered by Thomas Ash, two and one- 
third centuries ago. Writing in 1682, he says of a marine ecat- 
fish on the coast of North Carolina (for reasons given before, 
presumably the Gaff-topsail): “. . it hath a sharp thorny 
Bone on its Back, which strikes at such as endeavor to take it; 
which by Seamen is held venemous; yet, I saw one of our Sea- 
men, the back of whose Hand was pierced with it, yet no poy- 
sonous Symptoms of Inflammation or Rancor appeared on the 
Wound, which quickly healed, that I concluded it was either 
false, or that of this Fish there were more kinds than one.” 


144 Zoologica: N. Y. Zoological Society [5 


FOOD AND FEEDING. 


The Gaff-topsail is an omnivorous feeder, almost anything 
being meat that comes to its mouth, whether fish or crab or 
worm. It seems to affect mud flats and after them submerged 
sand flats as feeding grounds. The water in the Beaufort re- 
gion, where the sea cat-fish is found, is too muddy for any obser- 
vations to be made on the feeding habits, but if one may judge 
of these by analogy after observing the habits of the young 
(4 to 6 inches long) in a large aquarium, it probably feeds by 
swimming a few inches above the bottom with its long barbels, 
tactile organs, just touching the surface of the mud. Whenever 
these touch anything edible, there is a quick turn, a sudden 
opening of the cavernous mouth and it is gone. I have often 
experimented with the young, and have found their barbels 
exceedingly sensitive te bits of oyster dropped into the aquarium. 
I have seen the little fish thus arrested, stop in full flight and 
even turn a somersault in its eagerness to get at the oyster. 
The fish, of course, would readily perceive moving objects and 
:f these were edible, would snap them up. The feeding described 
above is more that of a scavenger. 


The food of the adult is—anything edible. I have on dissec- 
tion found the stomach filled with fish, worms, crabs. The latter, 
however, is its staple food, and I have taken from the stomach 
blue crabs so large that it was difficult to see how they could 
have been taken into the mouth and down the oesophagus. 
Autopsy has revealed the presence of ascidians, and, during one 
summer, certain small gray holothurians as noted above (page 
138). H. M. Smith, (1907), notes similar feeding habits and 
food for the smaller ocean catfish, Galeichthys milberti, at 
Beaufort. 


Before leaving this subject it may be noticed that in Florida 
this and the other marine catfish are accused of feeding on hu- 
man feces. I have had a very detailed account of this from a 
man who is absolutely reliable, and his account has been cor- 
roborated by a scientific friend who has personally seen the fish 
thus engaged. 


1916] Gudger: The Gaff-topsail 145 


PARASITES. 


As might be expected from its omnivorous feeding habits, the 
Gaff-topsail harbors a considerable number of worm parasites. 
However, as the greater number of my autopsies have been 
performed at the fishing ground with other points in view and 
while greatly pressed for time, my notes merely record the find- 
ing of worms in the stomach. Further, however, it is interest- 
ing to note that another investigator, working at Beaufort on 
internal protozoan parasites, has found in the intestine of our 
fish considerable numbers of a large potato-shaped ameba having 
remarkably clear protoplasm and a rapid rolling motion. This 
and other results have not yet been published. 


USE AS Foon. 


The value of the Gaff-topsail as a food fish is, irrespective 
of other points, considerably diminished by the large size of its 
head and by the bony cuirass extending back to the origin of 
the first dorsal. Nevertheless, it has been, and is used as food. 
Bloch, (1794), says that it is eaten, but that its flesh is not 
especially palatable. Ash, (1682), remarks of the marine cat- 
fish that: ‘“. ... it’s estem’d a very good Fish.” The older 
American ichthyologists thought highly of it as a food fish. 
Thus Mitchill, (1815), says, “It is an exquisite fish for eating.” 
While De Kay, (1842), writes: “Its flesh has been represented 
to me by those who have eaten it as having an exquisite flavor.” 
But Jordan, (1884), while remarking that its flesh is palatable, 
says that it rarely is saved for food, for the most part being 
thrown away. 


Various authors, Jordan and Gilbert (1883), Henshall (1891, 
1895), Evermann (1899), Gregg (1902), and others, writing 
of this fish in our southern waters, say that by reason of the 
abundance of other and far better fish it is rarely eaten, save 
by negroes. I never knew the fish to be eaten at Beaufort, nor 
was there any demand for it for export save in one season. There 
was a considerable shortage of fish in 1908 and a New Bern fish 
dealer, who had a “buy boat”? anchored in Newport River, bought 
Gaff-topsails along with other common (non-choice) fish to sell 


146 Zoologica: N. Y. Zoological Society ess 


to the negroes of that town. I have eaten its flesh, in order to 
be able to report on it, and have found it not unpalatable, but 
not particularly appetizing. Perhaps, however, it was not well 
prepared. 


SOUNDS MADE BY THE FISH. 


Felichthys felis makes two distinct sounds, one a croaking 
and the other a rasping sound. The first is the more common 
and is produced by the swim bladder. If the fish be grasped back 
of the pectorals, distinct pulsations may be felt with every croak. 
These are very apparent in a fresh and vigorous fish, especially 
if it shows signs of anger. Larvae also croak and by holding 
them in the fingers it will be noted that, as in the adults, pulsa- 
tions may be felt in the body wall. | 

The rasping sound made by the Gaff-topsail was first thought 
to be due to the fish rubbing its superior and inferior pharyn- 
geals together. However, it was soon noticed that the grating 
or rasping sound was accompanied by a spamodic jerk of the 
pectoral spines, and that if these were held immovable no rasp- 
ing sound could be perceived although the croaking continued, 
the gritting noise beginning again when the spines were re- 
leased. So it seems that these sounds are made by the spines 
as they rotate in their sockets. 

On one occasion, after some resistance on her part, I took a 
large active female cat from the water and laid her down in the 
dip net on a small pile of oyster shells, whereupon she made a 
spitting noise for all the world like an angry tabby cat. I am 
not sure how it was done, but it was possibly a combination of 
the two sounds previously described, and the pile of oyster shells 
may have acted as a resonator aiding in combining the two 
sounds. This was the only occasion on which this peculiar sound 
was noticed. 


SIZE OF BREEDING FEMALES. 


It will be of interest briefly to consider the size of breeding 
fish, and first of the females. In fish generally these run larger 
than the males, and in our catfish this is especially true, due 
largely to the enormous ovaries filled with huge eggs ranging 
up to 25 mm. in diameter. 


— 


~ 
= 


-“ 


uy 
a 
: 


x. 
Dy 


: a 


neleht 
a mL 3 


FIG. 26. CEMENT CAST OF THE MOUTH OF THE MALE FISH CARRYING 55 EGGS 


Dorsal view. 


FIG. 27. CEMENT CAST OF MOUTH 


Lateral view of Fig. 25. 


Zoologica Vol. II. No. 5. Face page 147 


1916] Gudger: The Gaff-topsail 147 


The fish taken on May 138 and 15, 1911, are quite typical of 
breeding females. These had not spawned their eggs and hence 
had enormous bellies. The largest, taken May 15, measured: 
191% inches, 1; 20 inches, 2; 2214 inches (15 inches in girth), 
2; 23 inches, 1; 2814 inches (14 inches in girth), 1; 2414 inches, 
1; 25 inches, 1. This last was the most enormous catfish I ever 
have seen. She measured 19 inches in circumference just back 
of the dorsal fin, the filament only of which (whence the name 
Gaff-topsail as noted above) was 334 inches long. She had, 
however, not reached her maximum size for her eggs were not 
ripe—they could not be spawned. 


SIZE OF THE INCUBATING MALES. 


This can best be set forth by giving the sizes of 32 egg- 
carriers measured on the initial trip taken in this research, June 
22, 1907. It will be noted that their sizes run very uniform, 
but that they are markedly smaller than the females. On this 
day there were measured: 1, 1534 inches over all; 4, 1714 inches; 
Sas menesr a cleo ig, loloe 2s 18387 so WO. 1199842 7, 20: 1, 
2014; 2, 21: 32 in all. Of these 32, 23 range from 171% to 18% 
inches; and generally speaking later observations confirm these 
figures as being the average. 


How THE Eaccs ARE CARRIED. 


These breeding males, as previously noted, carry the eggs 
loosely in the mouth, the gill-covers being widened outwardly 
and the hyoid distended downward to make the “double chin” 
previously referred to. In this way the cavity of the mouth 
is enlarged and its capacity increased. As may be seen from 
figure 22, which is a pen and ink sketch of a 171% inch breeding 
male, the Gaff-topsail, like most siluroid fishes, is largely head, 
and the head is mainly mouth. Figure 21 shows the mouth 
distended in the hyoid region to accommodate the eggs. 


148 Zoologica: N. Y. Zoological Society [II;5 


SIZE OF MOUTH CAVITY. 


A number of casts were made of the mouths of fish carrying 
large numbers of eggs, but of them only the largest will be 
considered here. This was of a 22-inch male burdened with 
fifty-five eggs, the largest number ever obtained from any Gaff- 
topsail in the course of this work. This fish was carried to the 
laboratory, seven miles away, that the capacity of its enormous 
““Keim-hohle’”’ might be made. However, there was but a small 
quantity of plaster of Paris in the laboratory and none in Beau- 
fort. In this predicament, Director Aller came to the rescue 
with the suggestion that cement be used, there being a barrel 
at hand. So a tolerably thick grout was made and the mouth 
filled with it, a towel being wrapped around the gills to prevent 
the escape of the cement before it had hardened. The head was 
then cut off, put out in a cedar thicket, where covered with a 
box it was left until the ants had eaten off the flesh. Later it 
was cleaned, shellacked, and photographed. Figures 26 and 27 
are dorsal and lateral, views of this huge cast. The volume of 
this cast, up to the insinking in the oesophageal region, is 580 ec. 


SIZE AND STRUCTURE “OF THE SKULL. 


In intimate connection with the size of the mouth is the 
matter of the magnitude of the skull. Reference to figures 20 
and 22 shows that the head makes up a large part of the body, 
about one-quarter of the length and possibly an equal part of 
the bulk. The buccal cavity, as has been shown in the preceding 
section, is enormous. In order that the reader may get a clearer 
idea of what gives it this great size, two views of the skull are 
given. Figure 28 is Mr. Morrison’s drawing of.the dorsal sur- 
face, while figure 29 is a photograph of the ventral surface of 
the same skull. The buccal cavity extends the whole length 
of the under surface of the skull, the hinder part, the beginning 
of the oesophagus, being formed under the coalesced vertebrae. 


Since such would be apart from the purpose of this paper, 
no attempt will here be made to work out the osteology of this 
very interesting skull. However, attention may be called to its 
armor-clad dorsal surface. This will explain why so much 


SKULL OF THE GAFF-TOPSAIL 


Dorsal view showing the fontanelle. 


FIG. 28. 


Face page 148 


Zoologica Vol. IT, No. 5. 


‘on 


1916] Gudger: The Gaff-topsail 149 


clubbing on the head is required to subdue an active catfish. 
There is, however, one easy method of quickly and compara- 
tively easily killing a catfish. This is by inserting a knife 
blade through the slit in the anterior median line in the bony 
armor. Through this the brain is easily reached. This open 
space in the roof of the brain is called a fontanelle. In the 
higher bony fishes it is closed, and its presence here is an 
evidence of the lowly position of the catfish in the class Pisces. 
It is an inheritance from its shark ancestor, which had a very 
marked fontanelle in the corresponding region of its skull. 


The only person, who, so far as has been found, seems ever 
to have noticed this structure in the skull of the catfish and 
consequently this method of killing it, was John Luccock. Luc- 
cock made a journey in 1808 to Rio de Janeiro and the River 
Plate, and twelve years later published a very interesting ac- 
count of his travels which contains many valuable natural his- 
tory notes. In speaking of the Bagre caught in the La Plata 
off Buenos Aires, he says: “It lives long out of water and is 
with difficulty killed by blows. I observed in the plate of the 
skull, between the eyes, a small aperture, covered with a thin 
whitish membrane, and imagined that through this, it might 
be killed by touching the brain. We accordingly introduced 
a filament taken from one of the bass cables, which produced 
an immediate paralysis and the fish died without further suf- 
fering.” 


Turning to the ventral surface of the skull of F'elichthys we 
find some equally interesting structures. In the posterior region 
are the large round paired bullae containing the ear stones. 
Ventral and posterior to these we have a curious bony formation 
very like a crucifix, the two little semi-circular bones behind 
(above) it looking somewhat like a halo. I have the indefinite 
recollection of having somewhere read of the feeling of semi- 
veneration paid to the catfish skull showing these structures 
by the superstitious inhabitants of the Guianas and the neigh- 
boring islands but I have been able to lay hands on but one 
reference. 

The Beebes in their charming book, “Our Search for a Wilder- 
ness”, (1910) say that, while their vessel was anchored in one 
of the mouths of the Orinoco: “‘At the bottom, our hooks would 


150 Zoologica: N. Y. Zoolegical Society BD oes 


be taken by great fierce-whiskered cats, bedecked with long 
streamers, which gave no end of trouble before they were quieted. 
They were pale yellow, and the head and back were encased in 
bone; Maestro the cook called them the Crucifix fish, and later 
showed us why. On the under surface of the bony armor is 
a large cross with a halo about it just above the arms. The 
crew never caught one of these fish without making the sign 
of the cross in their right palm”’’. 


The Beebes give a photographic reproduction of “the cruci- 
fix in the catfish’, but their figure seems to have been made 
from a skull that had suffered considerable erosion. It does 
not have the sharpness and clearness of detail found in figure 29. 


SIZE OF EGGS AND NUMBER CARRIED. 


The ripe eggs vary in size as is to be expected, running from 
15-25 mm., but the average diameter is 19-20 mm.. The small- 
est number of eggs found in the mouth of any gestating male 
was two. Two fish were found, 13 and 1514 inches long over 
all, each with two eggs. Once, it is true, a large male was 
found carrying only one egg, but from the great size of his 
buccal cavity there is good reason to believe that other eggs 
had been thrown out in the process of capture. This I have 
known the Gaff-topsail to do. Large numbers of eggs are by 
no means unusual; a dozen fish have been taken with eggs in 
the thirty’s; forty-five eggs have been taken twice; fifty were 
obtained from a 22-inch male; and greatest of all fifty-five from 
another fish of the same size. 


ATTEMPTS TO HATCH THE EGGS ARTIFICIALLY AND THE DIFFI- 
CULTIES MET WITH. 


Karly in the course of this investigation it was seen that it 
would be necessary to carry these eggs by artificial means to 
the point of hatching and beyond, if an embryological series 
was to be obtained. However, it seemed doubtful if eggs accus- 
tomed to such a highly specialized brooding chamber could pos- 
sibly be carried on to hatching in open jars of running sea-water. 


FIG. 29. VENTRAL SURFACE OF SKULL 


Showing the “Crucifix in the Catfish.” 
Photograph of same skull as Fig. 28 


Zoologica Vol. II. No. 5. Face page 150 


1916] Gudger: The Gaff-topsail 151 


Unfortunately it was never found possible to bring incubating 
males to the laboratory, for the fish either died or at best became 
sick and spat out their eggs. The eggs however could readily 
be brought in in pails of water renewed at intervals. 


But for all this great difficulty was experienced in keeping 
these eggs alive. At first they were kept in shallow glass aquaria 
under running salt water, but the fine sediment from the water 
so thickly covered their shells that the supply of oxygen was 
to no inconsiderable degree cut off. That this was not the only 
cause of their “going bad” was, however, afterwards ascertained. 


To remedy this, some of the eggs were put in filtered sea- 
water with Ulva and placed near a window but not in direct 
sunlight. Though some died, others did fairly well for a while. 
The greater number, however, were put in baskets of a coarse- 
meshed galvanized wire netting and hung in aquaria 9 inches 
deep under running salt-water—the idea of course being that 
the greater part of the sediment would fall through to the 
bottom. These baskets were hung with copper wire covered 
with thread and erroneously supposed to be paraffined. Pos- 
sibly this combination made a weak electrolytic apparatus. At 
any rate, on the day following their suspension thirty-nine 
dead eggs were taken from the baskets. The baskets were then 
suspended by zinc-coated wires, but the mesh being rather large, 
the heavy eggs settled down in it in such a way as to cause 
congestion in the yolk circulation and finally death. 


After the death of all the eggs, as noted in the preceding 
paragraph, another trip was made and another lot of eggs in 
fine condition was brought in. Each had an embryo on the top 
nearly ready to burst the shell. Each little fish rested in a 
groove in the yolk, the head lying flat with both eyes above the 
groove. The tail of each was bent, the caudal fin covering one 
eye and reaching to the edge of the other. Here again see 
figure 23. 


The eggs continuing to die daily, in seeking to remove all 
possible causes, it was thought that the density or saltness of 
the water might be too great. This at the laboratory averaged 
1.021, while at the Narrows of Newport River, where the fish 
were caught, it was at the surface 1.007. To obviate this pos- 
sible cause a number of eggs were put into filtered sea-water 


a 


152 Zoologica: N. Y. Zoological Society (ils 


diluted with filtered rain-water to a density of 1.016 and were 
placed under running water of the same kind carried over by a 
siphon. These eggs all died, possibly because the flow of the 
siphon was insufficient to aerate the water in which they were 
placed. Presently but one egg remained. The larva on this 
had burst its tough shell on the dorsal side, and through the 
slit had thrust its head and the anterior part of the body. 
With its projecting eyes, black with a golden ring, and its 
head parts faintly stippled in black, it presented a striking and 
beautiful appearance. Fig. 30, A and B, from photographs made 
on this day, give some idea of the dorsal and ventral surfaces 
of this egg; only colored drawings could do it justice. Probably 
as a result of the handling incidental to the photographing of 
this egg, is was found dead the next morning. 


All other methods having failed to bring about the hatching 
of these eggs, it was determined to try the hatching jar devised 
by former Commissioner of Fisheries MacDonald and named 
after him. In this apparatus water is admitted to the bowl- 
shaped bottom of a tall glass jar, whence it rises up through 
the eggs and escapes by means of a pipe at the top. The pur- 
pose is to keep the eggs continually in motion and to carry off 
all sediment, bacteria, and mold spores by the escape pipe at 
the top. With the catfish eggs it was hoped that the current of 
water would be sufficiently strong to keep the eggs agitated, to 
lift them up enough to prevent the congestion of the ventral 
yolk-sac circulation consequent upon the considerable weight of 
yolk plus embryo. However, the outcome was only partially 
successful, for even in the hatching jars the mortality was very 
great. 


On July 7, 1908, a trip to the Narrows was made in the hope 
that hatching eggs might be gotten. In this we were successful 
for two cats were taken. One, 1714 inches long, carried thirty- 
six eggs. The little cat on one of these eggs had burst its prison 
and had thrust its head out of the rent in its shell as shown in 
figure 30 and the others were about ready to do so. The other 
male carried in his mouth six larvae 53-55 mm. long over all, 
sitting on yolk sacks so heavy that they could not yet lift them. 
Here again see figure 31. This would seem to fix the first week 
in July as the approximate hatching time. 


A B 
FIG. 30—A « B. A LITTLE GAFF-TOPSAIL THAT HAS JUST BURST THE SHELL 


A—Dorsal view. B —Ventral view. 
From an instantaneous photograph. 


FIG. 31. LARVAE OF THE GAFF-TOPSAIL CATFISH 


From an instantaneous photograph of the little catfish skating on their yolk-sacks at the 
bottom of an aquarium. 


Zoologica Vol. IT. No. 5. Face page 152 


1916] Gudger: The Gaff-topsail 153 


Great difficulty was experienced in hatching these eggs. Some 
died with congested yolk circulations as described above for the 
previous year, others gradually grew pale and finally died with- 
out any definite cause being found. And now there was devel- 
oped a new trouble which threatened to carry off all the remain- 
der at hatching time. Their shells seemed to grow rotten so that 
the fishlets by vigorous twistings could burst them at one place 
or another. When this took place in what may be called the an- 
terior region, so that the head could be thrust forth, all was well. 
For this see figure 30. But when as more commonly happened, 
the shell burst elsewhere and the compressed yolk pushed out the 
investing wall with its plexus of blood vessels forming a hernia, 
death shortly ensued from strangulation of the circulation unless 
the trouble was relieved at once. At this stage in the history of 
the little cats, life was conserved only by constant vigilance. I 
kept watch by day and until 11 o’clock at night and the night 
fireman thereafter; and, whenever a “herniated” egg was dis- 
covered, the shell was torn partly or completely off the egg and 
the hernia pressed back into place with the smooth handle of a 
scalpel. This was a rather rest-disturbing matter since I was 
frequently awakened three or four times in one night. But the 
operation was for the most part successful since some 75 per 
cent. of the young so treated recovered. The majority of deaths 
in these “hulled” eggs resulted from the congestion of the ventral 
yolk circulation brought about by the weight of the fish and yolk 
or by the continued wriggling of the fishlets. Those from which 
the shells had been removed suffered especially, since their yolks 
flattened down greatly, while those whose shells were merely 
torn open, but not removed, suffered far less since their yolks 
were partially supported. All were put on beds of cotton wool 
at the bottoms of aquaria under jets of sea-water. 

During the season of 1909 in endeavoring to hatch the eggs, 
the experience of the past seasons was repeated. The eggs 
went forward very well till they neared the time when they might 
be expected to hatch, then they died by fives and tens and twen- 
ties. An interesting phenomenon may here be noted which may 
offer a possible explanation for some of these deaths. On July 
1 there was noticed inside the shell of an egg a small mass of 
greenish-yellow matter looking much like the fecal matter given 


154 Zoologica: N. Y. Zoological Society [11;5 


off by a young baby and noted as such at the time. Further an 
embryo at the time of hatching or a few minutes thereafter had 
hanging from its vent a string of fecal matter. A third egg 
on the same day exhibited a similar state of affairs. All this 
leads one to question whether it may not be that some of these 
eggs which die just at the time of hatching are poisoned by 
fecal stuff given off by the embryos and confined within the egg 
shell. 


The embryos from the paternal mouth, which were just ready 
to hatch, had far less difficulty in ridding themselves of their 
shells than the ones brought up in MacDonald hatching jars. 
And while it is probable that there is some mortality among 
the eggs incubated in the mouth of the father, there is no doubt 
that it is nothing like so great as that among eggs hatched ar- 
tificially. 


Just as the percentage of fish hatched in the paternal mouth 
is greater than that of those brought up in a hatching jar, so 
is it probable that the young incubated therein mature earlier 
than in the jars. This seems to be confirmed by this fact. On 
July 20, my fishermen, men whom I know well and in whom I 
have great confidence, brought in a little cat about four inches 
long taken from the mouth of a 20-inch male. This little fish 
was grown in the sense that its body walls had completely coal- 
esced over the yolk. The men reported that they saw several 
little ones in their net. These they tried to catch, but they es- 
caped through the meshes of the seine, the large fish, however, 
retained one in his mouth. This young one gave much trouble by 
jumping out of the bucket of water into the bottom of the boat, 
and finally escaped by jumping overboard as it was being handed 
to me at the wharf. 


In contradistinction to the early hatching noted above in the 
mouth of the father, of the larvae in captivity, taken on July 7, 
1908, the older ones did not close over their yolk sacs until Au- 
gust 5-8, while the corresponding dates for the younger captives 
was August 15-17. Making all allowance for difference in time 
of beginning incubation, the two or three weeks’ interval separ- 


1916] Gudger: The Gaff-topsail 155 


ating the periods of yolk disappearance in the two sets of larvae 
is too long to be accounted for satisfactorily save on the ground 
that the young in the mouth of the paterfamilias develop more 
rapidly. This must be due to the fact that they feed while 
therein. 


How THE EMBRYOLOGICAL SERIES WAS OBTAINED. 


For six years (1907-1912) the search for the fish and its eggs 
went on, and with few successes and many defeats the series 
of eggs was pushed both backwards and forwards. By fitting 
in one egg here and another there and a third elsewhere, the 
series is now complete from invagination to the free swimming 
young in which the walls of the belly have closed over the di- 
minished yolk sac and have coalesced into a raphe on the median 
line. The inability to obtain the segmentation stages is a great 
disappointment, for they only are lacking for the complete em- 
bryology, but, since the most strenuous efforts continued for six 
years have failed to obtain them, I have come to the reluctant 
conclusion that chance in the case of this fish will play more 
part in the collecting of eggs and embryos than any amount of 
hard and long continued effort. 


CONCLUSION. 


How the eggs are extruded, fertilized, and transferred is not 
known, but when these processes are effected the male incubates 
the eggs in his mouth not only until they are hatched by the burst- 
ing of the shell, but until the yolk has been absorbed and the 
young are able to take care of themselves. The largest num- 
ber taken from the mouth of one male was fifty-five. A cement 
cast of his mouth had a volume of 580 cc. The volume of an 
average sized egg is 3.75 cc., of the fifty-five eggs 206.3, add 
25 per cent for interstices; total space occupied by the fifty-five 
eggs equals 258 cc. This fish was 22 inches long and of aver- 
age size. The eggs average 19-20 mm. in diameter, and the 
young fish at the end of the period of incubation are 85-100 mm. 
long. The length of this period can not be stated definitely, 
since it has been found to be impossible artificially to carry the 


156 Zoologica: N. Y. Zoological Society Bee 5° 


eggs and embryos to the stage of the free-swimming young. 
However, it is probably about seventy days. During all this time 
the paternal nurse does not seem to feed. The large eggs would, 
if spawned on sandy or shelly bottoms, be quickly destroyed by 
crabs or by other fish; if laid on a mud bottom (where the breed- 
ing fish are caught) their considerable weight would cause them 
to sink into and be smothered by the mud. This habit is com- 
mon in estuarine catfish in all tropical and warm temperate 
regions. These data are based on six summers’ work at the 
Beaufort laboratory of the United States Bureau of Fisheries, 
in which time scores of male fish carrying eggs and larvae have 
been captured and autopsied. 


BIBLIOGRAPHY 


Asu, THoMmas. 
1682 Carolina; or a Description of the Present State of that Country, and 
the Natural Excellencies thereof, etc., p. 26. London. 


Brrese, Mary B. ann C. Wi1tam. 
1910 Our Search for a Wilderness, pp. 13-14, fig. 5. New York. 


Biocn, Marcus Enieser. 
1794 Naturgeschichte der Ausliindische Fische, Achter Theil, p. 26. Berlin. 


Boake, Bancror’. 
1866 On the Air-Breathing Fish of Ceylon. Journal Ceylon Branch Royal 
Asiatic Society for 1865-66, pp. 128-142. 
BoNNATERRE, PIERRE JOSEPH. 
1788 Ichthyologie: Tableau Encyclopedique et Methodique des 'Trois Regnes 
de la Nature, p. 154. Paris. 


BRICKELL, JOHN. 
1737 The Natural History of North Carolina. Dublin. 


Day, Francis. 
1889 Fauna of British India: Fishes, vol. I, pp. 169-170. London. 


De Kay, JAMES. 
1842 Zoology of New York, or the New York Fauna, Part IV, Fishes in 
Natural History of New York, p. 179. 
E1cENMANN, C. H. ann R. S. 
1890 A Review of the South American Nematognathi or Catfishes. Occa- 
sional Papers California Academy of Science, vol. I, p. 36. 


EverMANN, B. W. 
1899 Report on Investigations by the United States Fish Commission in 
Mississippi, Louisiana and Texas in 1897. Report United States 
Fish Commission for 1898, pp. 285-310. 


EverMANN, B. W., anv Bean, B. A. 
1898 The Indian River and Its Fishes in The Fisheries of Indian River, 
Florida. Report United States Fish Commission for 1896, p. 240. 


EvermMann, B. W., ann GoxpsBoroucH, E. L. 
1902 Notes on Some Mexican Fishes. Science, n. s., vol. XV, pp. 746-749. 
1902 Report on Fishes Collected in Mexico and Central America. Bulletin 
United States Fish Commission, vol. XXI for 1901, pp. 140-146. 


Gupcer, Kk. W. 
1912 Oral Gestation in the Gaff-topsail Catfish, Felichthys felis. (Abstract). 
Science, n. s., vol. XX XV, p. 192. 
1912 George Marcgrave, the First Student of American Natural History. 
Popular Science Monthly, pp. 250-274. 
Guntuer, A. C. L. 
1864 Siluroids in Catalogue of Fishes in the British Museum, vol. V, p. 173. 
1866 Description of a New Siluroid- Fish from Ceylon. Annals and Maga- 
zine of Natural History, ser. 3, vol. XVII, pp. 473-474. 
1880 An Introduction to the Study of Fishes, p. 160. Edinburgh, 
Grece, W. H. 
1902 Where, When, and How to Catch Fish on the East Coast of Florida, 
New York and Buffalo. 


HensHat., J. A. 

1891 Report upon a Collection of Fishes Made in Southern Florida during 
1889. Bulletin United States Fish Commission, vol. IX for 1889, 
jos, wilh 

1895 Notes cn Fishes Collected in Florida in 1892. Bulletin United States 
Fish Commission, vol. XIV for 1894, p. 211. 


157 


BIBLIOGRAPHY. 


JONSTON, JOANNES. 
1758 Historia Naturalis Piscibus et Cetis, Libri V, tab. 38, fig. 3. 
Jorpan, Davin STarr. 
1884 The Catfish Family in Natural History of Useful Aquatic Animals, 
Sec. I of The Fisheries and Fishing Industries of the United 
States, G. Browne Goode, Editor. 
1905 A Guide to the Study of Fishes, vol. I, p. 128; vol. II, p. 179. New 
York. 
Jorpan, Davin Starr, AND Evermann, B. W. 
1900 The Fishes of North and Middle America, vol. IV, fig. 52, plate 
XXIII, and p. 3196. 
1883 Notes on Fishes Observed about Pensacola Harbor, and Galveston, 
Texas, etc. Proceedings United States National Museum, vol. 
V for 1882, p. 245. 


Jorpan, Davin Svarr, anp» Gitpert, C. H. 
1883 Notes on a Collection of Fishes from Charlestown, South Carolina, 
etc. Proceedings United States National Museum, vol. V_ for 
1882, p. 584. 


(LAWSON, JOHN. 
1714 The History of Carolina: Containing the Exact Description and Nat- 
ural History of that Country. Reprinted Charlotte, N. C., 1903, 
p- 94. 


Luccock, JoHN. 
1820 Notes on Rio de Janeiro and the Southern Parts of Brazil, ete., p. 
148. London. 


MarccGraveE, GEORGE. 
1648 Historiae Rerum Naturalium Brasiliae (De Piscibus, Liber Quartus), 
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George Marcgrave, Amstelodami. 


Mrrcuitt, Samvuet L. 
1815 The Fishes of New York Described and Arranged. Transactions Lit- 
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Piso, WiILLiamM. 
1658 De-‘India Utriusque Re Naturali et Medica, p. 64, Amstelodami. 


Ruyscu, Henry. 
1718 Historia Naturalis Piscium in Theatrum Universale Animalium, etc., 
p. 143. 


Smiru, Hucu M. 
1907 The Fishes of North Carolina. Vol. If of N. C. Geological and Ec- 
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Turner, WILLIAM. 


1867 On a Remarkable Mode of Gestation in an Undescribed Species of 
Arius. Journal of Anatomy and Physiology, vol. I, pp. 78-82. 


WitiucHsy, FRANCIs. S 
1686 Ichthyographia. De Historia Piscium, Libri Quatuor, p. 140, Oxonii. 


WYMAN, JEFFRIES. 
1857 Exhibition of and remarks on Some Fishes of Surinam and on the Con- 
ditions under which their Eggs are Developed. Proceedings Bos- 
ton Society of Natural History, Sept., 1857, pp. 268-269. 
Yarrow, H. C. 
1877 Notes on the Natural History of Fort Macon, North Carolina, and 
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158 


e ZOOLOGICA? 
SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 6. 


MAMMALS OF AUSTRALIA IN THE 
ZOOLOGICAL PARK 


Illustrated with photographs by the Author and Elwin R. Sanborn 


By W. H. D. LE Souer, 
Director Zoological Gardens, Melbourne. 
Author ‘‘The Animals of Australia,” ‘Wild Life in Australia.” 


Pou Bol tS H- ED Bere oe SS OOGNE Bp ey 
THE ZOOLOGICAL PARK, NEW YORK. 


JANUARY, 1919 


New York Zoological Suriety 


General Office, 111 Broadway, New York City 


Officers 
President, HENRY FAIRFIELD OSBORN; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; © 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 
Treasurer, PERCY R. PYNE. 


Board of Managers 
Glass nf 1919 
PERcY R. PYNE, GEORGE BIRD GRINNELL, GEORGE C. CLARK. 
CLEVELAND H. DopcE, C. LEDYARD BLAIR, FREDERICK 
‘GILBERT BOURNE, EMERSON -McMILLIN, ANTHONY 
R. KUSER, WATSON B. DICKERMAN, MORTI- 
MER L. SCHIFF, FREDERIC C. WALCOTT, 
BEEKMAN WINTHROP. 
Glass of 1920 
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, Gaines F. 
DIETERICH, GEORGE F’. BAKER, WM. PIERSON HAMILTON, 
ROBERT S. BREWSTER, Epwarp S. HARKNESS, 
WILLIAM B. Oscoop FIELD, A. BARTON 
HEPBURN, WILLIAM WOODWARD, 
EDWIN THORNE, PERCY A. 
ROCKEFELLER. 
Glass of 1921 
Levi P. MorTON, ANDREW CARNEGIE, MADISON GRANT, WILLIAM 
WHITE NILES, HENRY A. C. TAYLOR, FRANK K. STURGIS, 
GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFORD 
Morris, ARCHER M. HUNTINGTON, HENRY ~ 
M. TILFORD, E. C. CONVERSE. 


Srivutific Staff 


WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 

WILLIAM BEEBE, Honorary Curator of Birds and Director of the 

Tropical Research Station; 
LEE S. CRANDALL, Curator of Birds; 
GEORGE S. HUNTINGTON, Prosector; 
GEORGE A. MACCALLUM, Pathologist; 
W. ReEID BLAIR, Veterinarian; 
ELWIN R. SANBORN, Photographer and Editor. 


Editorial Committee 


HENRY FAIRFIELD OSBORN, Chairxman; MADISON GRANT, 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


’ 


—— — ee, 
Aysonian Instip~ 
r Me Oe, » 


VOLUME IJ, NUMBER 6. 


MAMMALS OF AUSTRALIA IN THE 
ZOOLOGICAL PARK 


Illustrated with photographs by the Author and Elwin R. Sanborn 


By W. H. D. LE SoueEr, 
Director Zoological Gardens, Melbourne. 


Author ‘‘The Animals of Australia,” ‘‘Wild Life in Australia.”’ 


Pel sbi. k Sohn D Boy Tell SOC Ey 
tebe ei GolC Ad PYAR K, NEW YORK 


JANUARY, 1919 


PREFACE 


DIRECTOR LE SOUEF AND THE AUSTRALIAN FAUNA. 


In length and breadth of departure from the recognized 
standards of mammalian anatomy and physiology, the mammals 
of Australia are, per capita, the most odd and remarkable of any 
continental group. With the exception of the dingo, a few rodents 
and bats, all those species that do not lay eggs are marsupials, 
and carry in the abdominal pouch the astoundingly minute newly- 
born young until it grows to a size fit to take a small place in 
the outer world. A newly-born kangaroo cannot possibly be ap- 
preciated by a stranger until it is seen. 


The Australian marsupials display a remarkable line of ra- 
diating development that is quite inexplicable to zoologists. This 
relates to the production of forms within an order, that strik- 
ingly parallel in external appearance the characteristic forms of 
members of various orders of mammals. It would appear as if 
the scheme of evolution among the Australasian marsupials 
tended to produce an aggregation of pouched mammals that in 
form and habits would cover the strange absence of other orders. 
The Tasmanian “wolf” may be cited as an example and the ant- 
eating echidna, with its porcupine-like quills, as another. There 
are carnivorous, fox-like phalangers, marsupial “mice,” the wom- 
bat—in form and habits like a gigantic woodchuck, and the flying 
phalanger, which latter animal is precisely like a flying squirrel 
in form and actions. Yet more remarkable is a marsupial mole. 


The New York Zoological Park always has been rather 
strong in Australian mammals. They are so universally inter- 
esting as to be irresistible. Our Australian collection is now 
very rich. As a contribution to public interest in these strange 
creatures from the continent wherein Nature has done every- 
thing differently, the distinguished Director of the Melbourne 
Zoological Gardens has been prevailed upon to write a series of 
short, popular sketches of the Australian species now or recently 
exhibited here, and illustrate many of them with photographs 
taken by him in Australia. 


Mr. Le Souef is a man of charming personality and success- 
ful habit. He visited and lectured in America about eight years 
ago, and thereby greatly strengthened the bonds of interest be- 
tween the zoologists of his country and ours. He is the author 
of books on the wild life of Australia that are at once deeply 
interesting and thoroughly reliable. The titles of those best 


164 Preface 


known are “Wild Life in Australia” (London, 1907), and ‘The 
Animals of Australia,” by A. H. S. Lucas and W. H. Dudley Le 


Souef (London, 1909). 
W. T. HORNADAY. 


CONTENTS 


PREFACE . sn he ee Se, Te} 
INTRODUCTION OMe be: errr | lol 
MEAVENMEAL CON NUSTRALIA >.> ss 0 sue. +. wD 
DiCOMMET ..2- so Sd eee Pere Tes: a ek eee 
Picea s .)  NSNiiaots nce Meme te ee ae a ea 
RATS Pence Pn en eee en ng es el 2 ae 
EAURETS) Go Se RS ee MEM etectis 20 es 9 i a ae dea a a!) | 
PEEK AMET A TG ery ey nen tt Mourne Sb se 2) A pr cg ee ell them 
RETARD EO AT Stetie ot Arie eta se eed eS ae 
Saar ees, naires i et cael AM al oS oc ee 
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FIGURES 


33—Australia: (Map) . 
34—Young Gray Kangaroo 
35—Wallaro 
36—Woodward Kangaroo 
37—Black-Faced Kangaroo 
38—Parry Kangaroo 
39—Giant Red Kangaroo 
40—Kangaroo Island, Kangaroos 
41—Bennett Tree Kangaroo 
42_Bennett Tree Kangaroo 
43—Captive Tree Kangaroo 
44-—Rufous-Necked Wallaby 
45—Brush-Tailed Phascogale 
46—Tree Wallaby 
47—Albino Red Kangaroo 
48—Albino Red-Bellied Wallaby 
49—Ring-Tailed Wallaby 
50—Brush-Tailed Wallabies 
51—Black Swamp-Wallaby 
52—Male of the Swamp-Wallaby 
53—Agile Wallaby J 
54—_The Home of the Tree Wallaby 
55—Short-Tailed Wallaby 
56—Rat Kangaroo 
57—Australian Gray Phalanger 
58—Koala or Native Bear 
59—Koala in its Tree-top Haunts 
60—Fat-Tailed Opossum Mouse 
61—F lying-Phalanger 
62—Rabbit-Eared Bandicoot 
63—Wombat and Young 
64—Tasmanian Wolf J 
65—A family of Tasmanian Devils . 
66—Under surface of the Echidna . 
67—Echidna . ; 
68—The Platypus, or Dick Bill 


Fig. 32—A typical forest in Victoria, Australia . Frontispiece 
Fig. : J 

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Facing 169 


Between pages 
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MAMMALS OF AUSTRALIA IN THE 
ZOOLOGICAL PARK 


By W. H. D. LE SOuEF, 
Director Zoological Gardens, Melbourne. 
Author “The Animals of Australia,’ ‘““Wild Life in Australia.” 


ILLUSTRATED WITH PHOTOGRAPHS BY THE AUTHOR AND 
ELWIN R. SANBORN. 


INTRODUCTION. 


Australia is a large country, approximately 2,000 miles 
square, and is very sparsely populated, therefore, although good 
laws exist for the protection of native game, it is very difficult 
to see that they are enforced in the thinly populated districts. 
For example, Queensland has an area of 670,000 square miles, 
but its population is only about 190,000 whites, and approxi- 
mately 9,000 aborigines. New South Wales is better, having an 
area of 309,460 square miles and a population of 1,847,214. 
Victoria has an area of only 87,884 square miles, but has a popu- 
lation of 1,397,977, so is considerably more dense than the other 
States. South Australia consists of 380,070 square miles and 
has 433,616 people, but Western Australia had the large area 
of 975,920 square miles and a population of only 308,806. 


The Northern Territory, also is a large district, consisting 
of 523,620 square miles, and inhabited by only 4,767 people, ex- 
cluding natives. The island of Tasmania has 26,215 square 
miles, with a population of 199,925. 


In glancing over these figures one can easily realize the dif- 
ficulty in fully enforcing game laws. The only way that native 
animals surely can be preserved for those that come after us 
is, to form Reserves in various types of country. This is being 
done in many of the States, but only to a limited degree at pres- 
ent, because the subject is a difficult one. Introduced foxes and 
domestic cats that have gone wild, to say nothing of rabbits, 
cannot well be kept out of these Reserves. The foxes and cats 
prey on the protected game, and the rabbits destroy the native 
grass and shrubs that it is sought to preserve. Of course, these 
animals are not al! over Australia yet, but they certainly will 


168 Zoologica: N. Y. Zoological Society Te G 


be in course of time, despite fences, and we cannot possibly 
estimate the havoc they will play with the ground game and wa- 
ter fowl. It is quite possible that some species will become ex- 
tinct before we realize it. 


Then again, parts of Australia are subject to severe droughts, 
and thousands of small animals, as well as birds and kangaroos, 
perish, and emus cannot migrate as they used to do, on account 
of fences and settlements. The sheep and cattle help to denude 
the country and drain the waterholes. Therefore, in some dis- 
tricts where certain forms of life formerly were in evidence, 
none are seen now. Take as an example about fifty miles inland 
from Rockhampton in Queensland: there the beautiful parra- 
keet, (Psephotus pulcherrimus) was fairly plentiful, but since 
the drought in 1896 not a bird has been seen in the whole dis- 
trict. The pig-footed bandicoot was comparatively common in 
the southern districts of Australia, but now one is rarely, if 
ever, found. | 


Gilbert’s rat-kangaroo, (Potorous gilberti), of southwest 
Australia, apparently is extinct. The so-called native cat, (Dasy- 
urus) was exceedingly plentiful in Victoria, but now they are 
just as scarce as they once were plentiful. It is difficult to say 
why these various animals have almost disappeared. Of course 
the settlements and what they bring with them might account 
for a good deal, but certainly not for all. We really know little 
as to the unaccountable disappearance of small mammals in dis- 
tricts where they were numerous, and when we wake up to the 
fact that they have gone, it is usually too late to take measures 
of protection. Probably the same thing occurs in America, and 
elsewhere. 


The introduction of foxes into Australia by private persons 
is bound to cause the destruction, and possibly, extinction of 
certain ground game. Inasmuch as much of the country has 
been cleared of scrub, the game does not have the same cover 
that it had formerly. The animals that live in burrows prob- 
ably will hold their own longer than those that make their nests 
on the surface. Tasmania being of comparatively small area, 
is sure to lose the marsupial wolf or thylacine before long, as 
the dense bush is cleared and the country becomes more thickly 


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1919} Le Souef: Mammals of Australia 169 


settled. Even now it is scarce, and the settlers snare and destroy 
it whenever they get the opportunity. The Government has 
lately established a large Reserve for it near Hobart. 


In Queensland there are ninety-two Honorary Rangers, and 
that State is trying to protect its animal life, but having so much 
sparsely populated country it is difficult. In the near future, 
they probably will convert more Crown Land into Reserves. 
So far only four have been made for animals and fifty-two for 
birds. It is now under consideration to take up the subject of 
the exportation of the skins of Australian native wild animals, 
and to place this important matter on a proper basis. It prob- 
ably will be under the control of the Commonwealth Govern- 
ment, and further efforts then will be made to preserve our fast 
disappearing fauna. In every country there are what are popu- 
larly called ‘“‘game hogs,” heedless men as well as thoughtless 
boys, who seek to destroy the fauna of the country for their own 
individual benefit, and with no thought for posterity, or whether 
they are exterminating the fauna or not. Simply for what they 
eall “sport,” they carelessly destroy all they can, making little 
use of what they do kill. Persons of this class always are with 
us, more or less. 


THE MAMMALS OF AUSTRALIA. 
DINGO. 


The Dingo, (Canis dingo) is found over the whole of Aus- 
tralia, but curiously enough not in Tasmania or the adjacent 
islands to the north. It probably is one of the most ancient of 
wild dogs, and its anatomy shows it to be an intermediate form 
between the wild dogs of South America and those of the old 
world. It is a true wild dog. 


These animals are usually met with singly or in pairs and 
only occasionally in small packs, and then they probably would 
be parents and young. They never attack a human being. They 
usually hunt at night, but their scent is so keen that they fre- 
quently capture ground birds by stalking them and then catch- 
ing them as they rise on the wing from the ground. They are 
the size of a sheep dog, and the ears are short and erect. The 


170 Zoologica: N. Y. Zoological Society [TL 3/6 


fur on the under part of the body is gray, the longer hairs, 
which give the body coloring, are reddish-yellow to much darker 
Shades and in some specimens the saddle is almost black. The 
Western Australian dogs are, as a rule, darker than those in 
Victoria. The feet and tip of the tail are usually white. 


Albinos sometimes occur and as these often breed true, a 
white race could easily be established. The females always seem 
to predominate. They are fleet and strong and can overturn a 
sheep or calf with ease, should the mother of the latter be absent. 
However tame they may appear in captivity, they cannot resist 
the temptation of killing a fowl, should they get an opportunity. 
They bite with a snap like a wolf, and animals bitten by them . 
seldom recover. 


In the open country these dogs can be ridden down by a 
good horse and despatched with a stirrup iron or waddy, or even 
caught if necessary. They are readily poisoned with strychnine. 
When the aboriginals of Australia found Dingo pups they used 
to rear them with care, tame them partially, share their bed 
and food with them, and the dogs then would only follow their 
owners. They were never struck by the natives, but these par- 
tially tamed dingoes often joined their wild comrades in the 
bush and did not return. The natives used them for hunting, 
but they apparently only followed their master. These animals 
breed freely with European dogs and consequently it is now 
difficult to obtain a pure bred Dingo. They never bark, but can 
howl dismally. 


The fossil bones of these animals have been found in several 
parts of Australia in a formation that is considered to be Plic- 
cene, so that apparently they were in existence in Australia 
long before human beings. 


WATER RAT. 


The Water Rat, (Hydromys chrysogaster). This is a large, 
handsome rodent fully twenty inches long from nose to tip of 
the tail. They are dark buff above, a rich orange-brown below, 
and the tail is white toward the end. These animals are still 
fairly plentiful, but being nocturnal are seen rarely. They are 
purely aquatic and frequent inlets of the sea as well as rivers 


1919} Le Souef: Mammals of Australia nara 


and lakes, where they feed on the shell-fish, crustaceans and 
vegetation. They are found in all parts of Australia and Tas- 
mania. The western species is slightly darker than those found 
in the eastern side of Australia and the amount of white on the 
tail also varies. A small rat, dark grey on the back, called 
Thomas’ Rat, (Xeromys myoides) is found in Queensland. Its 
body is only four and one-half inches long and its tail three and 
one-half inches. 


RATS. 


Due to their remarkable fecundity, rats are very plentiful 
in Australia, as in other parts of the world, both in species and 
individuals. Both rats and mice occasionally increase during an 
unusually good season, when food and cover are plentiful, to 
almost incredible numbers. During the year of 1917, there was an 
abundance of rain in southern Australia throughout the summer 
which produced a great growth of grass and other vegetation. 
Therefore, as food was plentiful and the vegetation so dense, 
the rodents were securely hidden from their enemies. They in- 
creased so enormously that food became scarce, especially where 
the grass began to get dry, and they had to migrate in their 
many millions. They were then preyed on by snakes, carnivorous 
animals and birds, but despite this, the destruction caused by 
these little rodents was very great. At the wheat stacks alone 
at eight railway stations in Victoria, about thirteen tons of mice, 
representing approximately 892,000 animals, were caught in 
three days. The same migrations occur in other countries, espe- 
cially among the lemingss in Norway. 


MICE. 


The members of the genus Mus are represented in Aus- 
tralia by twenty-eight species so far known. But this number 
is certain to be increased later on. As our knowledge of these 
animals is very incomplete at present, an authentic list cannot 
well be given. Of our twenty-eight species, only five can be 
termed mice, although it is difficult to draw a dividing line be- 
tween the two species. They are met with practically every- 


py Zoologica: N. Y. Zoological Society [II;6 


where, increase rapidly, and all burrow more or less. One of 
the commonest species is the Dusky-Footed Rat, (Mus fuscipes), 
is found in South Australia, the islands of Bass Straits and Tas- 
mania, usually near water or on swampy land. Its body length 
is about six inches and the tail is about four inches. The fur 
is conspicuously long. 


A closely allied form, (M. assimilis) or the Allied Rat hav- 
ing very soft fur, which is light brown above, is found from 
N. E. Queensland to S. W. Australia, usually in thickly timbered 
country. They live largely on fruits and seeds. On the Dar- 
ling Downs in Queensland, the M. sordidus is found. It is black- 
ish-brown above and measures about six and one-half inches, 
with a tail five inches long. Generally in open country, and at 
the extreme N. E. of Australia, the White-Footed Rat, (M. terrae- 
reginae) is plentiful. Its back is dark brown, with longer black 
hair and tail with light colored rings. It measures eight and 
one-quarter inches and the tail seven and one-quarter inches.* 


M. gouldi is reddish-yellow in color, with numerous long 
black hairs above. It measures four and one-half inches and 
tail three and one-half inches. It is found in south, eastern and 
central Australia. And M. greyi inhabits central and north- 
eastern Australia and is reddish brown in color with longer dark 
hairs. It is six inches long and its tail four and three-quarter 
inches. One small species in Western Australia, M. nanus, is 
only four inches long and its tail three and one-half inches. It 
is brown in color and has a white patch under the tail. The 
Pigmy Mouse, (M. delicatulus) from Northern Australia is only 
two and one-half inches and the tail about the same length. It 
is yellowish-brown in color and the upper parts and the sides of 
the body are yellow. 


The Greyish-White Mouse, (M. albocinereus) is found near 
the seashore in S. W. Australia. Its body is covered with long, 
soft, light grey hair and the tail and feet are white. The Brown 
Rat, (M. decumanus) and Domestic Mouse, (M. musculus) have 
spread over most of Australia and have become numerous in 
many districts in the country, as well as in the towns. The 


*In a short article like this full descriptions of the various species cannot 
well be given. (Author.) 


1919] Le Souef: Mammals of Australia Ms 


former have from two to three litters a year each containing 
from nine to fourteen young. The European Black Rat is also 
in evidence, not only in the cities, but chiefly in the country, as 
they have the habit of building their nests in hollows in trees 
and are therefore largely arboreal. 


JERBOA RATS. 


The interesting family of Jerboa Rats of the genus Coni- 
lurus (Hapalotis) is confined to Australia where they are also 
plentiful in many districts, their long ears and tails making them 
conspicuous. The fossil specimens that have been found are 
of great interest as illustrating in the rodents a mode of pro- 
gression similar to that of the kangaroos. These little animals 
advance by leaps and bounds like the Jerboas of Africa and Asia 
and the jumping mice of North America. Fourteen species are 
known so far, mostly confined to the interior, although some 
varieties live in the coastal districts of north Queensland. 


They have from three to four young, but have no pouch; 
the young being attached firmly to the nipple and also grasping 
their parent with their claws. They are strictly nocturnal, rest- 
ing during the day in nests of dried leaves and grass in hollow 
fallen branches. The largest varieties, C. boweri and C. hirsutus 
are nearly two feet in length and are found in north Queensland. 


The White-Footed, (C. albipes) is found in the southeastern 
districts. This animal is greyish-brown, black around the eyes, 
and has a body length of ten inches and a tail nine and one-half 
inches. The nest-building Jerboa Rat, (C. canditor) from the 
interior of eastern Australia is only six inches long, and its tail 
five inches. They combine together and make large nests of 
grass, sticks and bits of bark sometimes over three and one-half 
feet high, usually around a small bush, the branches of which 
help to strengthen the structure. One family or more may oc- 
cupy a nest and each family has its own compartment, which is 
connected with the others by passages that put one very much 
in mind of a beaver’s lodge. This animal is greyish-brown and 
is darker on the center of the back and head. 


174 Zoologica: N. Y. Zoological Society [II;6 


The Long-Tailed Jerboa, (C. longicaudatus) inhabits West- 
ern Australia and is seven inches long, with a tail nine inches. 
It is pale buff on the back and the end of the tail is white. It is 
usually found in scrub-covered country. The Fawn-Colored 
Jerboa, (C. cervinus) is four and one-half inches long and the 
tail five and one-half inches long. It is found in the central 
districts of South Australia. The large ears of this delicate 
looking little animal are much lighter in color than its back and 
are very conspicuous. It is all white below. A rat named the 
Dusky Broad-Toothed (Mastacomys fuscus) is found in Tas- 
mania. It is only five and one-half inches long and is dark grey- 
ish-brown in color. 


GIANT RATS. 


The largest rat in Australia, the Giant Rat, (Uromys ma- 
cropus) is found from northeastern Australia to the Arnheim 
Islands. It is fourteen inches long with a tail about the same 
length, and rarely is seen in captivity. It is reddish-grey above 
and white below and is probably destructive to birds’ eggs and 
young during the nesting season. The Buff-Footed Rat, (U. 
cervinipes) found in Western Australia only, is but six inches 
long with a tail a little over five inches. It is light brown above, 
with buff-colored feet. The scales on the tails of these animals 
do not overlap but are set edge to edge. 


MUSK RATS. 


We now come to the animals that are strictly marsupial, 
and in Australia they are naturally numerous. The Australian 
Musk Rat, (Hypsiprymnodon moschatus) usually found in the 
scrub-covered country of the coastal districts of Queensland, is 
a graceful little animal, with soft and orange-grey colored fur, 
diurnal in habit and living on insects, snails, fruit and seeds. 
It has two young at a time in its pouch, and its length is about 
ten inches and tail six and one-half inches. It is rarely seen and 
the perfume of musk easily identifies it. 


1919] Le Souef: Mammals of Australia 175 


KANGAROOS. 


All kangaroos have more or less the same habits and are 
usually found in small companies in country where they are 
not disturbed. They are protected for the whole year in Vic- 
toria and soon increase if undisturbed. Partial protection is 
given to them in New South Wales, but not in Queensland, except 
in certain districts. The number of skins annually sent to other 
countries from Australia, especially from Queensland, runs into 
many thousands. Of this number, the United States receives a 
large share; sometimes over 80,000 in one year. 


Many men make their living entirely by shooting kangaroos 
with a rifle; one man I know having shot over 400 last year 
(1917). This means that in the course of comparatively a few 
years, these interesting animals will become very scarce, as the 
skins of all species, including wallaby, are used for leather. The 
introduction of the fox into Australia will not help matters as 
they are sure to kill some of the young ones. These animals 
fortunately live and breed freely in confinement, having but 
one young at birth, although twins have been known to occur 
occasionally. 


They are hunted on horseback with the aid of a large dog 
of the grey-hound type, known as a kangaroo dog, and if the 
country should be sufficiently open, they usually are caught and 
killed. When hard pressed, they often will take refuge in a 
river or in swamps standing waist deep in the water and await- 
ing their enemies. Should a dog swim out to them, they will 
hold it under water with their fore arms and eventually drown 
the venturesome animal. When attacked on land, the old males 
that are not as speedy as the females, often stand with their 
back to a tree ready to fight with the dogs; and they are usu- 
ally quite a match for any single dog. Young kangaroos are 
often caught and reared by hand, when their mother has been 
shot or otherwise killed. Their backs are easily damaged if 
roughly handled. When leaning forward to feed on short grass, 
they often rest on the upper part of their paws, as well as on 
the under part in the ordinary way. When in this position, the 
young that may be in the pouch, and old enough, can nibble on 
the grass at the same time. 


176 Zoologica: N. Y. Zoological Society [II;6 


The Grey Kangaroo, (Macropus giganteus) is found across 
the entire southern part of Australia as well as in Tasmania. 
The species from that island (M. fuliginosus) is now very scarce. 
It has long, dark fur and the under parts are white. The female 
is much lighter in color than the male. Those on the western 
side of the mainland usually are darker, but generally melanism 
is more pronounced among the animals in the western portion 
of Australia than in the eastern. These animals are only a 
little inferior in size to the red kangaroo, and the fur is longer 
and coarser. The males are a dark grey and the females and 
young much lighter. They are found in open forest country 
and frequently are called locally the Forester Kangaroo. The 
variety from Tasmania and Kangaroo Island (M. fuliginosus) 
is now very scarce. It has long, dark fur, the under parts being 
white. The female is paler than the male. 


The Wallaroo of Euro Kangaroo, (M. robustus) have long 
and coarse fur; the color of the male being dark reddish- 
grey and that of the females more bluish-grey. Farther north 
in Queensland, the color is often dark greyish-brown in the males. 
The exact tint varies considerably. This variety is found in the 
central districts of Australia, as well as towards the coast. They 
live only on the rocky ranges and are thickset and strong and 
adepts at bounding over the often rough country where they 
are found, and where frequently it is difficult for a dog to follow 
them. 


Several sub-species of this animal, (M. woodwardi) from 
northwest Australia, have been described. The color of the short, 
close hair of the male is bright red and that of the female, fawn. 
The head and body measures four feet and the tail three feet. 
The fur of M. alligatoris from north Australia is also short 
and the color more or less rufous, with the neck, arms and fore- 
back, fawn. Another sub-species from southwestern Australia, 
M. cervinus, is lighter in color, and lastly M. isabellinus from 
Barrow Island off west Australia, has a dark rufous back with 
the front of the neck white. In the southern districts of Aus- 
tralia, in the drier and frequently sandy country where the 
mallee eucalyptus grows, is found a darker and more slender 
variety of kangaroo, the Black-Faced, (M. melanops). However, 


Photograph by W. H. D. LeSouef 
FIG. 34. YOUNG GRAY KANGAROO, M. gigantus 


Immature specimen just born and placed in the pouch. 
Beside it is the the nipple to which it would have 
been fastened about life size. 


FIG. 35. WALLAROO OR EURO KANGAROO 
New York Zoological Park. 


Zoologica, Vol. II, No. 6. Face page 176 


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KANGAROO ISLAND KANGAROOS 
New York Zoological Park. 


FIG. 40. 


Photograph by E. R. Sanborn 


FIG. 41. BENNETT TREE KANGAROO 


The tail is not prehensile, but is used in balancing. The forefeet are adept in grasping. These 
animals sometimes leap to the ground from a height of fifty feet. 


New York Zoological Park. 


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FIG. 47. ALBINO RED KANGAROOS 


Melbourne Zoological Gardens 


Photograph by W. H. D. LeSouef 
FIG. 48. ALBINO RED-BELLIED WALLABY 


Tasmania 


FIG. 49. RING-TAILED WALLABY 


New York Zoological Park. 


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1919} Le Souef: Mammals of Australia i 


as this country is being cleared rapidly for wheat-growing, this 
animal will become scarce, as it is destroyed by the farmers. 


Kangaroos are diligently sought for their skins, and although 
they are well protected in Victoria, and to a certain extent in 
other parts of Australia, their numbers are diminishing. A 
small, slender species, Parry’s Kangaroo, (M. parryii) is found 
in the hilly coastal districts of Queensland and northern parts 
of New South Wales. The short, soft and light bluish-grey fur 
marked with a white line on each side of its face as well as on 
the neck, and the long, thin tail, have suggested its local names, 
the Pretty-Face or Whip-Tail Kangaroo. It usually frequents 
scrubby country and often may be seen in the Darling Downs 
district from railway carriages when one is travelling from Bris- 
bane to Sydney, or vice-versa. It is three feet in height and its 
tail is two and one-half feet in length. The Red Kangaroo, (Ma- 
cropus rufus) is probably the largest of the kangaroos. The 
short, woolly fur is red in color in the male and bluish grey in 
the female. When standing upright, practically on its hind toes, 
and resting the weight of its body on the end portion of the tail, 
it measures about six and one-half feet; otherwise four to five 
and a half feet. 


Old males get very pugnacious and frequently fight one an- 
other. They do so by scratching, if possible, with their fore 
paws, and also by leaning back and resting the weight of their 
body on the extreme end of their tail, only about six inches, and 
striking forward with the hind feet. The claws are sharp and 
although they do not often do much damage to each other, they 
can easily rip up an unwary dog should one tackle them. These 
animals live on the plain country of New South Wales and south- 
ern Queensland, generally remaining during the heat of the day 
under the shade of the trees that fringe the plains. They can 
easily travel at the rate of twenty miles an hour when pursued, 
and exceed that speed when pressed. They cover about twelve 
feet at a jump and can clear a fence eight to ten feet high. Oc- 
casionally they are pursued on the plains with motor cars, al- 
though I hardly think that is a fair way to get them, as they 
have no chance, unless. they get into a belt of timbered or rough 
country. However, the sport is not destined to be very popular 


178 Zoologica: N. Y. Zoological Society [Li 


as motoring over the plains at over twenty miles an hour is 
usually a very bumpy experience. A female kangaroo when hard 
pressed in flight if she should be carrying a heavy young one, 
or joey in her pouch, will take the young one out and conceal it 
under a bush, coming back when all danger is over, should she 
have a chance. 


The only safe way to hold a kangaroo is by the tail, and it 
takes a strong man to hold one. The young are born in the ordi- 
nary way, but in a very immature state. They are about an 
inch long, the fore feet are twice the size of the hind feet and 
the tail very small. It is placed on the nipple in the marsupium 
by the mother and the pressure of the milk forms a small bulb 
at the end of the nipple at the back of the mouth. This swelling 
being larger than the entrance to the mouth of the young one, 
holds it on. If the young kangaroo is pulled off at an early stage 
it cannot be replaced. 


The Antilopine Kangaroo, (M. antilopinus) is found in the 
Coburg Peninsula in north Australia, and very little is known 
of this animal. It is of a heavy build with short fur, rufous in 
color with underparts white. The female is smaller and of a 
fawn color. The head and body are four and one-half feet and 
the tail two feet long. 


THE TREE WALLABY. 


These interesting animals are found in the mountain ranges 
near the coast of northern Queensland as well as in New Guinea. 
Two varieties inhabit Australia, namely Lumholtz’s, (Dendro- 
lagus lumholtzi) and Bennett’s, (D. bennettianus). The former 
which has long yellowish-brown fur with a black chin and white 
chest is found in the more southern districts near Cardwell, and 

-the latter which has long dark brown fur, further north near 
Cooktown. Like most of the other grazing Australian animals, 
they rest during the day and feed chiefly at night. They live 
almost exclusively in trees or on the tops of granite boulders 
that are covered with vegetation. I once saw one of these ani- 
mals that I disturbed when in the latter situation, jump to a rock 


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1919] Le Souef: Mammals of Australia 79 


below, a distance of about forty-five feet. It is wonderful the 
way they are able to jump from bough to bough and rarely make 
any miscalculation. Their long tail is not prehensile and is used 
for balancing only, and the soles of their hind feet are serrated 
and that prevents them from slipping. 


The natives obtain them by going to the upper portions of 
the scrub-covered ranges in the early morning with their dogs, 
and the latter are frequently able to pick up the scent of a 
kangaroo that has gone from one tree to another or to track 
it to the tree in which it is feeding. Some of the natives then 
climb any tree in proximity to the one that shelters the wallaby. 
to prevent it escaping into it, while another of their number 
climbs the tree that harbors the animal, and either catches it 
by the tail or forces it to jump to the ground in its endeavors 
to escape. The other natives with the dogs are there on the 
lookout for it and generally secure it. 


The wallabies frequently get from one bough to another by 
going along towards the end of a branch, and clinging to it with 
their fore paws, as it bends they are enabled to get a foothold 
on one at a lower level. They can also jump a considerable dis- 
tance from one bough to another. As a rule, when they are on 
the ground they lean well forward and keep their tail clear of 
the soil. In ascending a tree, they do so by clinging with their 
fore paws round a creeper and moving both hind feet up at the 
same time; and they go up very quickly. They cannot ascend 
an ordinary trunk of a tree, but as the forests in the districts 
where they are found abound with creepers, practically every 
tree having one or more, they have no difficulty in climbing. 
They also can climb an ordinary two or three-inch rope with ease, 
or even a gas or water pipe; their serrated pads enabling them 
to get a secure hold. Should one escape on board a vessel, it 
quickly climbs the ropes and easily gets to the mast-head. These 
animals in their native state are more or less infected with two 
or three kinds of parasites and often have sore places caused by 
them. As their food consists of leaves of various shrubs, they 
live well in captivity. 


180 Zoologica: N. Y. Zoological Society [Tie 


WALLABIES. 


There is little difference between kangaroos and wallabies. 
The members of the genus Macropus whose head and body are 
over four feet in length are called kangaroos, and those three 
feet and under, usually wallabies. One of the largest of the lat- 
ter is the Black-Tailed, (Macropus nalabatus) measuring just 
under three feet and the tail two feet. This animal, which is of 
rather a heavy build, and not nearly so active as many of the 
smaller kinds, is always found in scrubby country. Its color is 
very dark brown or reddish-grey and light rufous underneath. 
They formerly existed in countless numbers in the densely tim- 
bered portions of the coastal districts of New South Wales and 
Victoria, and hundreds of thousands of their skins have been 
exported. But trappers and settlements have so reduced their 
numbers that they are now protected in Victoria. During the 
day they usually remain well hidden, coming out in the evening 
to feed. A very closely allied variety, (M. apicalis) is found in 
the same class of country in the coastal districts of Queensland. 
It has shorter fur and the rufous color is more intense. Prob- 
ably the largest of these animals is the Red-Necked Wallaby. 
It is of more slender build than the preceding species, is greyish- 
fawn in color, with a reddish neck and rump, and measures three 
and one-half feet and its tail two and one-half feet. It is found 
in the eastern parts of Australia from southern Queensland to 
Victoria where it usually inhabits the open forest country. 


The Tasmanian form of this wallaby, (M. bennettii) has 
thicker and longer fur and is slightly darker in shade. Its neck 
and rump are dull brown instead of red. It also is found in 
southern Victoria and on the islands in Bass Strait. Formerly 
it was very plentiful, especially in the Islands, but now the 
hunters with their dogs have completely exterminated them also, 
except in Tasmania, where they still hold their own in the rough 
parts. In captivity they easily become tame and do not knock 
themselves about in the way other species often do. In Victoria 
and South Australia, Grey’s Wallaby, (M. greyi) is found. It 
is a Slender animal and can travel very fast. The color is grey- 
ish-fawn with a rufous tinge on the neck. It measures about 
two and one-half feet in length and its tail slightly under. An- 


1919] Le Souef: Mammals of Australia 181 


other fine wallaby, the Black Striped, (M. dorsalis) is found in 
the inland districts of New South Wales and southern Queens- 
land. The general color is grey with a reddish tinge on the fore- 
quarters, and it is readily distinguished by a narrow black line 
down the center of its back. It measures slightly over two and 
one-half feet and its tail two feet. 


The Black Gloved Wallaby, (M. irma) from southwestern 
Australia is a well-marked animal, with soft fur of a bluish- 
grey above, white below the chin, cheeks stripes also of the same 
color, and another white stripe on the neck, edged with darker 
color. They measure about three feet and tail two and one-half 
feet, thrive well in captivity and make very docile pets. The 
Agile Wallaby, (M. agilis) is a heavier animal with short, coarse 
dark sandy-colored fur, very short ears and a long tail that 
easily distinguishes it. The habitat of this species is southeast 
New Guinea, as well as in the northeastern portion of Australia. 
The Cape York Wallaby, (M. coxeni), another species from 
northeast Australia, is also a dark sandy color, darker on the 
back with white underparts and a white hip-stripe. These ani- 
mals are small, being only twenty-eight inches long and their 
tail fourteen inches. 


The Branded Wallaby, (M. stigmaticus) is found also in 
northeast Queensland, but usually further south than the before- 
mentioned species. It is of slender build, the fur is short, of a 
bright reddish-grey color, with less red on the neck and fore- 
quarters. There is also a pale cheek-stripe, the hip-stripe is 
yellowish and prominent, and the underparts white. It meas- 
ures twenty-nine inches in length and its tail fourteen inches. 
The Red-Legged Wallaby, (M. wilcoxi) is very similar to the 
preceding one, but the color is duller, and the hip-stripe hardly 
visible. It is found in the southern districts of Queensland and 
the northern parts of New South Wales. 


The Pademelon Wallaby, (M. thetidis), found in eastern 
Australia from southern Queensland to Victoria, is a light, grace- 
ful little animal, grey in color, reddish on the neck and white 
below. Its ears are long, and the hip-stripe is very faint. It 
measures twenty-six inches and its tail sixteen inches. The 
Dama Wallaby, (M. eugenii) from West Australia, the islands 


182 Zoologica: N. Y. Zoological Society [II;6 


off that coast and South Australia, has thick, dark grey fur with 
reddish shoulders and a pale cheek-stripe. This little animal is 
about the same size as the preceding one. 


The Parma Wallaby, (M. parma), very closely allied to the 
former species, has an even reddish-grey color with a distinct 
cheek-stripe and a white front. It is found in eastern New 
South Wales. The Rufous-Bellied Wallaby, (M. billardierz) 
used to be exceedingly numerous in Victoria and especially on 
the islands in Bass Straits, as well as in Tasmania, but those on 
the islands have been nearly cleared out. The hunters with 
packs of kangaroo dogs, used to burn the thick patches of scrub 
in which they knew the wallabies had taken refuge during the 
day and their dogs caught the unfortunate animals as they ran 
out. Their habitat is in the dense scrub and although their runs 
are very numerous in such places, they are fairly safe under 
ordinary circumstances. Many hundreds of thousands of their 
skins have been exported. These animals are of stout build, 
have thick, soft fur of a dark greyish-brown color, face and head 
olive-grey and no face markings. The body measures twenty- 
seven inches and the tail which is very short, only fourteen 
inches. 


The Short-Tailed Wallaby, (M. brachyurus) from Western 
Australia is the smallest of the wallabies. Its body measures 
twenty-three inches and its tail ten inches. Its fur is long and 
coarse and is a uniform greyish-brown. The ears are small and 
rounded. Its habits are identical with those of the rufous-bellied | 
wallaby. 


The Rock Wallabies, (Petrogale) are found all over Aus- 
tralia, but not in Tasmania. As their name implies they live 
only in rough rocky country, whereas the members of the family 
Macropus are usually found in the more level districts. The 
Rock Wallabies lean well forward, using their long, bushy tails 
only for balancing and not as a third support, as do the Ma- 
cropus family, especially the larger forms. The underside 
of the toes are covered thickly with small tubercles that prevent 
the animals from slipping on the rocks, especially when they are 
wet. They usually take refuge during the day in caves or under 
rocks, coming out to feed in the evening and at night. The 


1919] Le Souef: Mammals of Australia 183 


wonderful way that they can bound freely and without hesita- 
tion from rock to rock, sometimes onto excrescences that can 
hardly be seen, is extraordinary. A dog naturally and fortu- 
nately has little chance of catching them. In the many runs 
among the rocks that have been used by countless numbers of 
these animals for many years past, the rocks are perfectly pol- 
ished and shiny. No ordinary fence will stop this active animal, 
and, should they escape from captivity they seem to enjoy hop- 
ping about the roofs of buildings, apparently quite at home and 
where they cannot well be followed. 


The largest of the group is the Yellow-Footed, (P. xan- 
thopus). The body measures thirty-two inches and the tail 
twenty-four inches. The fur is long, soft and grey in color. It 
has a prominent white cheek-stripe, an orange spot above each 
eye, and long ears. A black line extends from the head to the 
middle of the back, there is a brown patch behind the elbow and 
a white lateral line that runs to the hip. On top of the knee 
there is another brown patch, and alongside of it a patch of 
white, and the arms, bands, legs and feet are yellow. The tail 
is marked above with alternate bands of dark brown and pale 
yellow. This handsome animal lives in South Australia. 


Another variety, the Brush-Tailed, (P. penicillata) found 
in the eastern coastal districts of Australia, is a thick set animal 
with long, coarse brown fur, a light cheek-stripe and short ears. 
It measures thirty inches long and the tail, usually tipped with 
yellow, is twenty-four inches. 

The Western Australian form, (P. lateralis) is smaller in 
size. The body being two feet long and its tail one and one-half 
feet. The fur which is light grey, is long and soft and its cheek- 
stripe yellowish. It also has a dark line on the center of the 
upper part of the back, a dark brown patch just behind the el- 
bow, and a prominent white stripe running down to the hip. 
The latter half of its tail is black. 


Another form from the northwest coastal districts of Aus- 
tralia is the Short-Eared, (P. brachyotis). This little animal is 
also slender, with short greyish-brown fur, has practically no 
face markings, the body markings only just visible and is grey- 


184 Zoologica: N. Y. Zoological Society [II;6 


ish-white below. The body measures twenty-two inches and the 
tail sixteen inches. 


There are three wallabies belonging to the genus Onychogale 
(Nail-Tailed). They are well marked animals having fairly 
long tails, crested at the ends and provided with spurs. These 
are the only marsupials that have such an excrescence. Among 
mammals, the lion is the only one that has a similar spur. The 
Nailed-Tailed Wallaby, (O. unguifera) from northwest and 
north-central Australia is a slender and graceful fawn-colored 
animal, with a darker medium band, and white hip-stripes and 
under parts. The body measures twenty-six and the tail twenty- 
eight inches. The tail is long and white on the upper side with 
a few, faint brown rings showing towards the end, which is 
black. The spur is flattened laterally and hidden in the long 
hairs. 

Another variety, the Bridled, (O. frenata) is found in the 
eastern districts of Australia from South Queensland to Vic- 
toria. These slender little animals having a body length of but 
twenty-three inches and tail of eighteen inches, make charming 
pets and when hopping along have the habit of holding their 
fore paws straight out in front of them. Their grey fur is soft 
and thick. The cheek-stripe is indistinct, the center of the back 
of the neck is black, and there is a white shoulder-stripe and a 
very indistinct hip-stripe. 

The West and South Australian form, Cresent, (O. lunata) 
is very similar to the preceding species. The fur is a soft dark 
grey, and the neck is rufous. The white shoulder-stripe is promi- 
nent and there is a faint stripe above the hip. It measures twen- 
ty-one inches and the tail fifteen inches. 


The Hare-Wallabies, (Lagorchestes) of which three varie- 
ties are known, form another interesting group of these animals. 
The name was given because of the likeness in form, size and 
habits to the hare. They are found usually in plains country, 
can travel at great speed and are adepts at dodging any dog that 
may be chasing them. The Spectacled Wallaby, (L. conspicil- 
latus) is found on the islands off the west coast of Australia. 
On the mainland, the closely allied variety of this species has 
been named L. leichhardti. Their color is yellowish-grey with 


1919] Le Souef: Mammals of Australia 185 


a-reddish band round the eye and two light lateral bands: They 
measure about twenty-one inches and tail eighteen inches. The 
colors of the mainland form are brighter and the bands and 
under parts are white. 


The common Hare-Wallaby (L. leporoides) is found in the 
interior districts of New South Wales and South Australia. It 
is light in structure and its general color is like that of the 
common hare, with the exception of a black patch on the elbow 
and also the reddish band round the eye. It measures about 
twenty inches and the tail about thirteen inches. The last va- 
riety is the Rufous, (L. hirsutus). It is found in the more south- 
ern districts of West Australia and is greyish in color, with a 
reddish tinge behind, and no black patch on the elbow. It meas- 
ures eighteen inches and the tail fifteen inches. 


In the same part of Australia, the graceful Banded Wallaby, 
(Lagostophus fasciatus) is found. It usually lives in thick, scrub- 
by country and is still plentiful in places. It is greyish-brown 
in color and has three sets of fur. First, grey under-fur, then 
grey, coarser hair and then long well separated piles projecting 
well beyond the others. It has no marks on its face or flanks. 
The lower part of the back has dark and light transverse bands 
that identify this little animal readily. 


KANGAROO RATS. 


We now come to the Rat-Kangaroos, or as they are called 
in Australia, Kangaroo Rats. They formerly were exceedingly 
plentiful, but dogs and foxes have taken a heavy toll of them 
and in the settled districts they have almost disappeared. They 
are about eighteen inches in length with a tail fourteen inches, 
and are of a sandy-grey color. They usually sleep coiled up in 
their nests during the day, coming out to feed in the evening and 
at night. The largest of them, readily distinguished by its reddish- 
grey color, the Rufous, (Aepyplymnus rufescens) is found only 
in New South Wales, and is twenty-one inches long with a tail 
fifteen inches. It has an indistinct stripe in front of the hips. 


There is another family of these active little animals, namely 
the Bettongs, (Bettongia), characterized by the fact that they 


186 Zoologica: N. Y. Zoological Society (Tr 


are the only ground animals having prehensile tails, which they 
use for carrying bundles of grass for the construction of their 
nests. A hollow is first scraped out in the ground and in it a 
dome-shaped nest of grass is built; the top being about level 
with the surface of the ground. The animal upon entering the 
nest draws a bunch of grass after him, and closes the entrance 
so perfectly that the nest cannot be observed, easily. A fox or 
dog, however, can readily detect the presence of the owner; if 
he happens to be at home. If he should be, there is little hope, 
for the intruder simply pounces down on the nest and usually 
secures the owner. The varieties of these animals are closely 
allied externally and are difficult to identify without knowing 
the locality from which they came. The Tasmanian variety, (B. 
cuniculus) is slightly the largest.and has white feet instead of 
brownish, as in the others. The underparts are white and, gen- 
erally, there is a white tip on its tail. The New South Wales 
variety, (B. gaimardi) has hair of a more woolly texture than 
the others and white hind paws; also a few white hairs at the 
tip of the tail. The Brush-Tailed Rat-Kangaroo, (B. penicillata) 
is found all over southern and central Australia and has a body 
length of fourteen inches. The tail which is twelve inches long 
has a black crest along the upper surface of the lower portion, 
but not white hairs at the tip. The phinarium is bare of fur 
as in the other species. This is the variety that is usually seen 
in captivity. 


The West and South Australian variety, (B. lesuewri), also 
found on some of the islands off the coast of West Australia, has 
a tail twelve inches long. Some specimens have a light stripe 
across the hip, and a white tip on the tail. The Plains Rat- 
Kangaroo of which there is but one species, (Caloprymnus cam- 
pestris) found in the plains country of South Australia, is a 
slender animal with a broad face, and sandy-colored, thick, soft 
fur, which is darker on the back. The feet are white, and the 
middle of the chest usually has a bare patch. It is very quick 
in its movements when chased, and dodges about so cleverly that 
it is difficult to catch. 


Probably, the rat-kangaroo that has the widest range is the 
Common, (Potorous tridactylus). It is found all over south- 


1919] Le Souef: Mammals of Australia 187 


eastern Australia and Tasmania. There are only three varieties 
of this family. The hind feet are short, and the tail short and 
tapering. When travelling fast, they use their fore feet as well 
as their hind ones. They never kick with their hind feet as the 
other rat-kangaroos do. The head of the Common is narrow 
and long, and is larger than the other two varieties. It meas- 
ures seventeen inches and its tail nine and one-half inches, but 
the size varies; those found in Tasmania probably being the 
largest. The fur is long, coarse and greyish-brown in color, and 
the ears short and rounded, and the tail usually is tipped with 
white. 


Gilbert’s Rat-Kangaroo, (P. gilberti) is found in the south- 
ern districts of West Australia and is smaller than the afore- 
mentioned; otherwise externally identical. The other form, (P. 
platyops) is also from southwestern Australia and is still smaller, 
measuring only eleven and one-quarter inches, and tail seven and 
one-half inches. It is called the Broad-Faced; which character- 
istic is very noticeable. It is otherwise externally the same as 
the others. An interesting little animal Hypsiprymnodon mo- 
schatus has been described from the tropical scrubs in northern 
Queensland. In size and form like a large rat, it is reddish-grey 
in color, and the large ears are bare as are also the fingers. It 
is seen but rarely in captivity, as the animal is hard to detect 
and capture. It hops like a bandicoot and lives on insect life, 
as well as fruit and vegetation. It has two young. 


PHALANGER OR AUSTRALIAN OPOSSUM. 


The Australian Phalangers, locally mis-called opossums, are 
not carnivorous like the American opossum, but feed entirely on 
vegetation; eucalyptus leaves forming the principal part. They 
live entirely in trees, and are nocturnal in their habits, sleeping 
during the day in some hollow or in their domed nest. They once 
existed in very large numbers, despite having formed the princi- 
pal article of food of the aboriginals in days gone by, but as their 
fur is of value for rugs they have been shot and trapped unmer- 
cifully and practically cleared out of many districts; over a mil- 
lion skins sometimes being exported annually from Australia. 
Foxes are now taking their toll of them in southeast Australia; 


188 Zoologica: N. Y. Zoological Society (II; 6 


catching them as they pass on the ground from one tree to an- 
other. They climb the trees by jumping quickly upwards with 
all feet at once; the sharp claws being extended to their fullest 
extent and thus securing a firm hold in the bark. They choose 
the upper side of a tree, should it be reclining in any way, and a 
defined track will be made on the bark of the one that is much 
used, which serves to guide the natives in finding the hollow in 
which the opossums are coiled up asleep. They usually are 
caught by placing a long, thick branch of stick against the tree, 
and the animal will always ascend by this in preference to going 
up the straight trunk. In descending the branch, the animal ad- 
vances head first, thrusts its head through a wire noose that has 
been placed on the stick, and thereby meets its fate. Many are 
shot; a moonlight night being chosen for the purpose, as the 
animals then can be distinguished against the face of the moon. 
The skins from the animals that have been shot are not as valu- 
able as those that have been snared. 


The smaller race of phalangers, called the Ring-Tailed, 
(Pseudochirus) is found in Tasmania, Australia and New 
Guinea. They also construct bulky, domed nests of sticks and 
leaves near the top of some thickly growing shrub, on which their 
tracks are not easily seen. They have from two to three young 
at birth which, on leaving their mother’s pouch, hang to her back 
for some weeks, by clinging with their claws to her fur, and are 
carried about until they are able to look after themselves. As 
their tail is prehensile and frequently used for clinging, the un- 
derpart of the end of it is rough and bare. Sometimes when 
shot and badly wounded they will hang on by their tails before 
life leaves them, and remain in that position after death for a 
considerable time; frequently a day. The end of the tail is usu- 
ally white. In the Herbert River district in Queensland, a small 
lemur-like variety, (P. lemuroides) is found. The soft, woolly 
brownish-grey fur is darker on the shoulders and lighter on the 
hips, and the head is brown and the tail black. It measures 
fifteen inches and tail twelve inches. 


Another closely allied form, P. herbertensis, is found in the 
Same district. It is dark brown in color, without markings, and 
is about the same size, but has a longer tail. Some specimens 


1919] Le Souef: Mammals of Australia 189 


have white rings round their limbs. The common Ring-Tailed 
Phalanger, (P. peregrinus) has a wide range, being found from 
eastern Queensland to the southern parts of South Australia. 
They still are very plentiful in places, even near the large cities, 
where they find refuge in private gardens, and frequently are 
seen in captivity where they soon become very tame. I often 
have seen them walking along the thin telephone wires in the 
grounds of the Melbourne Zoo. They use their tails as a balance, 
moving them quickly from side to side as necessary, and if 
knocked over they will continue their progress by hanging on 
by their claws. These animals do not live on eucalyptus leaves 
like the large kind, but eat almost any vegetation and fruit and 
are fond of rose vines and other leaves in flower gardens. Their 
fur is short and its color varies; being varying shades of dark 
rufous-grey with a patch of white on the edge of the ear. They 
measure about fourteen inches and their tail about twelve inches. 


The form in Western Australia is closely allied but has no 
rufous shade and the underparts are whiter, and usually there 
is more white on the end of the tail. There is also little differ- 
ence in the Tasmanian form, (P. cooki). Its thick fur is more 
woolly in texture and dark rufous-brown in color with more 
white on the ears. The Yellow Ring-Tailed is found in the coastal 
districts of central Queensland. Its color of yellowish-green and 
white below renders it, therefore, easily distinguished. Four 
forms are found in New Guinea, namely, P. albertisi and P. 
schlegeli from the Anfak Mountains, P. canescens from Pamoi, 
and P. forbesi from Sogere. The large Phalanger, or as it is 
universally called in Australia, the opossum, belongs to the genus 
Trichosurus. The species feed chiefly on the leaves of the euca- 
lyptus, resting during the day in hollows in these trees. It is 
the skins of this phalanger that have been exported in such quan- 
tities and are so largely used for fur rugs, etc. The variety that 
is by far the most plentiful, 7. vulpecula, is found over the whole 
of Australia, except on the extreme northeast. Although form- 
erly so abundant, in many districts they have been almost com- 
pletely exterminated; but when afforded protection, which they 
now have in many places, and especially in Victoria, they soon 
increase. They are easily tamed and would be kept as pets more 
often if they did not sleep coiled up all day long and become 


190 Zoologica: N. Y. Zoological Society [II;6 


lively only at night. Their thick, woolly fur is grey in color, the 
under parts whitish, the end of the bushy tail is black and bare 
on the under side, and there always is a reddish patch on the 
chest. In making rugs, the bare part of the tail is usually cut 
off and the backs only used. The ears are long and the body 
measures eighteen inches and the tail eleven inches. 


The Tasmanian variety, T. fuliginosus, is larger and has 
longer fur. Many specimens are dark grey, tinged with rufous, 
but others are a dark rufous-brown with a black tail. The skins 
of this latter handsome variety are of considerable value, and 
the animal has to be rigorously protected to prevent its exter- 
mination. In the heavily timbered uplands of Victoria and New 
South Wales is fonud the Short-Eared Phalanger, (T. caninus). 
Its beautiful fur is very thick and of a dark grey color, although 
in some specimens it is dark reddish-brown. The short, rounded 
ears, about one-half the length of the other varieties, render it 
easily distinguished. This animal is closely allied to the Tas- 
manian form. 


A black form of caninus, (T. c. nigrans) is found in the 
coastal scrubland of New South Wales and Queensland. The 
interesting point is, that the underparts are just as dark as the 
upper surface. During the summer, however, there frequently 
is a rutous shade. 


KOALA OR NATIVE BEAR. 


The Koala, or Native Bear as it is always called, of which 
there is but one species, (Phascolarctus cinereus) is found in all 
the eastern districts of Australia. They are strictly arboreal, 
living in the eucalyptus trees, and sitting during the day in a 
coiled-up position in a fork, where they are more secure. On 
account of this habit they are therefore easily shot, or killed by 
the heavy bush-fires. Like phalangers, the young cling to the 
back of the parent when they become too large for the pouch. 
The querelous, high-pitched note of the little ones is exactly like 
the crying of a child, but the old animals utter a prolonged, deep 
bass note. These animals are grey with white feet, and have 
thick, woolly fur. They do not possess a tail. An interesting 
little animal, Tarsipes rostratus, that is not often seen in cap- 


1919] Le Souef: Mammals of Australia 191 


tivity is found in Western Australia. It is only thirty-one inch- 
es in length, and its tail three and one-half inches. The color is 
grey, striped dorsally with dark brown. Its principal food is 
insects and it also sucks honey from the flowers. 


FLYING-PHALANGER. 


Australia possesses several forms of Flying-Phalangers, or 
as they are popularly called, Flying-Squirrels. When the Pha- 
langers stretch the feet well out, the loose skin that acts as a 
parachute holds the air sufficiently to allow the animal to glide 
from the higher branches of one tree to the lower trunk of an- 
other; the long, furry tail acting as a rudder. As they alight, 
a quick upward movement is made, the sharp claws enabling 
them to hold on to the bark, when they quickly can ascend the 
tree again and repeat the performance. All the species have 
beautiful long, soft fur. 


The Pigmy Flying-Phalanger, (Acrobates pygmaeus), well 
distributed over the eastern parts of Australia, is a delicate look- 
ing little creature, three inches long and with a tail three and 
one-half inches. The soft, silky fur is greyish-brown, the under- 
surface is white and the edge of the parachute is tipped with the 
same color. They usually have four young. On the timbered 
ranges of the coastal districts of Victoria and New South Wales, 
a large form of flying-phalanger is found, called the Yellow- 
Bellied, (Petaurus australis). Their color is greyish-brown, but 
varies in shade. The claws are strong and much curved, to en- 
able them to get a good hold of the tree trunk when alighting. 
The body length of the species measures eleven and one-half inch- 
es and the very long and bushy tail is sixteen and one-half inches. 
The Squirrel Flying-Phalanger, (P. sciureus), a much smaller 
form measuring only ten inches and tail ten and one-half inches, 
is light grey with a dark line on the crown. They are easily 
tamed and make interesting pets. They are found in eastern 
Australia. 


A still smaller form, found in the same districts, namely the 
Lesser, (P. breviceps) is only seven inches in length and its tail 
slightly longer. It also is light grey and can be distinguished 
from P. sciwreus only by its smaller size. The Papuan form, 


192 Zoologica: N. Y. Zoological Society [I1;6 


(P. papuanus) is also closely allied, even in markings, but has 
shorter fur and usually yellowish underneath and is smaller in 
its measurements. It is found in the Papuan sub-region of New 
Guinea, as well as on the adjacent islands. 


The Striped Phalanger, (M. dactylopsila trivirgata) , a strik- 
ing animal with a white body marked by dark, longitudinal 
stripes and a long bushy tail with a black line on its upper sur- 
face, is found from northern Queensland to New Guinea and on 
the Aru Islands. In the eastern districts of Australia, from 
southern Queensland to Victoria, is found the Taguan Flying- 
Phalanger, (Petauroides volans). It measures seventeen and 
one-half inches and the tail slightly longer. The ears are also 
very large. The fur, generally white below, is dark ashy grey, 
but it varies much in shade, some being lighter and others again 
nearly black. The Queensland form is usually smaller and has 
been named P. minor. Probably one form gradually runs into 
the other. 


A small, mouse-like animal, the Dormouse-Phalanger, (Do- 
micia) is closely allied to the flying-phalangers, but has no flying- 
membrane. Two forms are found in Tasmania. One, D. lepida 
measuring only three inches with a tail of the same length, is a 
graceful little animal, light fawn in color with fine, soft fur like 
all the others, numerous long whiskers and large ears. The 
other form, D. nana, fawn in color, but: with the legs usually 
grey, also found in Victoria and New South Wales, is slightly 
larger, measuring four inches and its tail slightly longer. They 
have four young at atime. These little animals often have fatty 
accumulations on the body. The Long-Tailed, (D. caudata), 
which is the larger of the genus, comes from northwest New 
Guinea. It measures four inches in length, tail five and three- 
quarter inches and the general color is rufous, with two dark 
lines on each side of the face. 


The smallest form, the Lesser, (D. concinna), found in 
South and West Australia, only measures three and one-half 
inches, with the tail slightly longer. Their color is fawn, and 
the underparts white. These little animals live well in captivity. 
Another genus, Gymnobelideus leadbeateri has been described 
from the mountainous districts of southeast Australia (Gipps- 


t61 and aony “9 “ON ‘JT JOA ‘n2160])007 


“yaed [Bo1so[oo7 y4IoX MON 
AGVTIVM GHTIVL-LYOHS “S$ “Old 


Photograph by E. R. Sanborn 


RAT- KANGAROO 


New York Zoological Park. 


FIG. 56. 


“yared [Bo1s0[007 YIOX MAN 
UAONVIVHd AVUD NVITVULSNV “LS “Old 


ULOQUDS “YW fq ydvsbo,0YT 


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LNOVH dOL-GHUL SLI NI VIVOM ‘69 “Old 
SenosaT “THM fq ydvsbojoyq 


SNAAIUVI SNIA DIOISDYT 


dvdd AAILVN YO VIVO ‘8S ‘DI 


Photograph by E. R. Sanborn 
FIG. 60. FAT-TAILED OPOSSUM MOUSE 


FIG. 61. FLYING-PHALANGER 
New York Zoological Park. 


8 


Photograph by E. R. Sanborn 


RABBIT-EARED BANDICOOT 


FIG. 62. 


New York Zoological Park. 


"yale [B01S0[007 yaoK MON 
ONNOA GNV LVEANOM °89 “Old 


ULOQUDS “YAY fig yavAbozoyT 


&61 abnd aD “9 “ONT ‘JT “JOA ‘n91d01007 


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uULogung “Yq fig ydv..b0,0y,7 


ie RST Le Souef: Mammals of Australia 193 


land), and is very rarely found. It is five and three-quarter 
inches long, tail a little longer, and the color of the body is grey 
with a dark line on the top of its head. 


CUSCUS. 


The Cuscus (Phalanger family), widely distributed over the 
islands in the Southern Pacific, are large and arboreal. They 
subsist on leaves and other vegetation. The Spotted Cuscus, (P. 
maculatus), the only one that is found in Australia, and that, only 
in the extreme north near Cape York, is also found in the Moluc- 
cas, Wai, Amboina, Ceram, Aru, and New Guinea as well as 
on some of the smaller islands. They live fairly well in captivity. 


The fur of these animals is thick and woolly and the general 
color mottled white, black and red in various combinations. The 
females are generally dark grey and black without any white, 
but the markings vary, some color predominating more than 
others in different individuals. The Grey Cuscus, (P. orientalis) 
is found in Bourn, Amboina, Ceram, Waigin, Aru Islands and 
New Guinea. The males of these species are sometimes quite 
white, but the color of individuals varies much in shade. The 
Short-Eared variety, (P. breviceps), slightly smaller and darker 
than the Grey Cuscus, comes from Duke of York, Solomons, San 
Christoval and other small adjacent islands, and the Ornate 
Cuscus, (P. ornatus), which is smaller and lighter than the Grey 
Cuscus, is found in Morty, Ternate and Batchian Islands. In 
the Celebes and adjacent islands of Sanghir is found another 
form of Grey Cuscus, (P. celebensis). 


THE BANDICOOTS. 


The Rabbit Bandicoot, (Peragale lagotis), from southwest 
Australia, is about the size of a rabbit. The fawn-grey fur is 
long and silky, head and ears are long, an indistinct dark line 
runs vertically from the back on the sides of the rump, the 
underparts are white and the tail towards the end is also white 
and crested. Bandicoots are destroyed chiefly by dogs and cats, 
and were far more plentiful in the days gone by than they now 
are. They are nocturnal; resting during the day in burrows, and 
feeding at night upon insects, grubs, earthworms, fungus and 


194 Zoologica: N. Y. Zoological Society [I1;6 


roots. They are marsupial and the pouch opens backwards. 
Their general color is olive-grey, with bars across the lower part 
of the back. The smaller Bandicoots belong to another species, 
namely Perameles. The Short-Nosed, (P. obesula) is the most 
widely distributed, being found all over southern Australia and 
Tasmania. It is fourteen inches long and the short, coarse fur 
is grizzled-yellow and black, the underparts are white and the 
ears Short and broad. They are usually found in swampy locali- 
ties where the vegetation is very dense. The North Australia 
Bandicoot, (P. macrura) is closely allied to the former, but it 
is larger and darker in color and its tail is longer. The Long- 
Nosed Bandicoot (P. nasuta), found in the central districts of 
eastern Australia, also is larger than the Short-Nosed, has very 
coarse hair and is brown in color without any markings. The 
ears are long and pointed, the underparts are white, and it meas- 
ures fifteen and one-half inches and its tail five inches. Gunn’s 
Bandicoot, (P. gunni) is about the same size as the last men- 
tioned, but has soft fur, yellowish-brown in color and with bands 
across the rump. 


The form in West Australia is the Striped Bandicoot, (P. 
bongainville:) is of small size being only nine inches long and 
has coarse olive-brown fur. Another form, (P. fasciata) in 
southeast Australia is closely allied to the former, but the stripes 
are very conspicuous and the tails are white. The Pig-Footed 
Bandicoot, (Cheropus castanotis), now becoming rare in places, 
is a small species measuring ten inches, and tail four inches. 
The name indicates its identity. 


THE WOMBATS. 


Wombats, (Phascolomys) are still fairly plentiful in south- 
eastern Australia and Tasmania. They dig deep burrows and 
are safe there from foxes and dogs, as they are quite able to 
defend themselves. Their length averages about forty-four 
inches. They prefer scrubby, mountainous country and their 
food is entirely vegetable. They are nocturnal, resting during 
the day in their burrows. They have no tail. 


The common variety, (P. mitchelli) is found in Victoria and 
New South Wales. They vary in color from dark yellowish-grey 


1919] Le Souef: Mammals of Australia 195 


to black. The Tasmanian, (P. tasmaniensis) is smaller and usu- 
ally of a dark greyish-brown color, and the Flinders Island form, 
(P. uwrsinus), the form originally but incorrectly described as 
from Tasmania, is yet smaller, being thirty-six inches in length. 
The Hairy-Nosed (P. latifrons), grey in color with the end of 
its muzzle white, is found only in South Australia. They are 
not as uniformly colored as the other varieties. These animals 
are very strong and burrow with great rapidity with their pow- 
erful claws; a habit that makes them very troublesome to settlers, 
as they dig under and damage wire-netting fencing. In walking, 
they shuffle along in a clumsy manner. They live well in cap- 
tivity, but are very subject to skin disorders. 


TASMANIAN WOLF. 


The Marsupial Wolf, (Thylacinus cynocephalus). These 
rare animals probably will become extinct before very long, as 
the settlers are prejudiced against them on account of their de- 
struction of sheep and other stock. The dark marks across the 
back are so very striking and distinctive, that the animal is usu- 
ally called locally the Tasmanian Tiger. They utter a peculiar 
coughing bark, rapidly repeated and something like that of the 
kangaroo. They have a fair sized pouch which opens backwards 
and usually bear from one to two young at a time. 


These animals resemble in form some of the short-legged 
wolves, but have short, close hair. During the day they gener- 
ally sleep in hollow logs, holes, under rocks, and pursue their 
prey in the evening and at night. They are not very fleet of 
foot but have a keen scent and usually spring on their prey, 
which consists, besides the stock of settlers, of wallabies, rat- 
kangaroo’s and other ground game. They swim well and readily 
cross rivers in pursuit of their prey, one having been recently 
observed swimming a river after a wallaby; quickly overtaking 
it. They are now found only in Tasmania, but their bones have 
been found in Australia. Why they disappeared from the main- 
land, it is difficult to say. 


196 Zoologica: N. Y. Zoological Society [I1;6 
TASMANIAN DEVIL. 


The Tasmanian Devil, (Sarcophilus ursinus) is strictly ter- 
restrial and is now found only in Tasmania, although formerly 
it was plentiful in the southern districts of Victoria, judging by 
the remains found. But these animals were apparently extinct 
before the arrival of Europeans. They are strong but sluggish 
beasts and, having powerful canine teeth, are a match for any 
ordinary dog. They are carnivorous and can bite severely, while 
the molar teeth enable them to crush bones with ease. As the 
Tasmanian Devil is comparatively slow of movement, it usually 
catches its prey by a sudden spring, afterward devouring it 
greedily bones and all. It utters a disagreeable kind of snort. 
The three to four young, when too large to remain in their moth- 
er’s pouch, generally cling to her back. They lie up in hollow 
logs or burrows during the day, coming out at night to catch their 
prey. They are destructive to poultry and lambs, consequently 
have few friends and are being killed out of all settled districts. 
Their color is jet black with a white horse-shoe mark on the 
chest, but they often have patches of white on other parts of 
the body. So far I have not seen an albino specimen such as 
one finds among the kangaroos and wallabies. 


SPOTTED DASYURE. 


The Spotted Dasyure or, as it is popularly called in Aus- 
tralia, the Native Cat, formerly used to exist in thousands in 
South Australia, but from some unknown cause these pretty little 
animals have now disappeared from many districts. There are 
three varieties, namely, the North Australian, (Dasyurus hal- 
lucatus), which is small, only measuring eleven inches and its 
tail eight inches; the Black-Tailed, (D. geoffroyi) from all Aus- 
tralia except the extreme north and the coastal districts of the 
southeast, and the Common, (D. viverrinus) from eastern New 
South Wales, Victoria, South Australia and Tasmania. These 
animals are marsupial, having about six young ones at birth. 
They are usually of a reddish-grey color, but also often black and 
are well marked with white spots, but not on the tail which is 
usually white at the tip. They measure about seventeen inches 
and their tail ten inches. A larger variety, the Spotted-Tail, (D. 


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1919 | Le Souef: Mammals of Australia 197 


maculatus) found from eastern Queensland to Tasmania, is more 
than twice the size of the other species. These animals can climb 
well although they are mostly terrestrial, taking refuge during 
the day in hollow logs and among rocks, etc. If several are kept 
in the same compartment in captivity, they are liable to turn 
cannibals; the stronger eating the weaker. They are carnivorous, 
taking birds and their eggs, mice, rats, bandicoots, and other 
game, and are very destructive to poultry. Consequently, they 
are not spared by the settlers. They are plentiful in Tasmania; 
more so than on the mainland. 


MARSUPIAL MICE. 


A form of Pouched Mouse with habits the same as the oth- 
ers, is the genus Phascogale. Their pouch is hardly visible. They 
bear from six to ten young ones at birth, living principally in 
holes in trees and lining their nests with grass and leaves. Nine 
species have been described, namely: the Crest-Tailed, (P. cris- 
ticardata), which measures about five and one-half inches with 
a tail three and one-half inches much thicker at the base, is found 
in central and southern Australia; P. macdonnellensi from cen- 
tral Australia has an abnormally thickened tail at the base. The 
Lesser Brush-Tailed Pouch Mouse, (P. calura), also found in 
southern and central Australia, is five inches long and has a tail 
six inches. P. penicillata or Greater Brush-Tailed Mouse is ten 
inches long and tail nine inches, is the largest of this genus. They 
are found all over Australia except at the extreme north, and 
live almost entirely in trees, making their nests in the hollows of 
the branches. They have thick tails and the end is covered with 
long, black hair, forming a brush. There is a very small variety 
named P. minutissima, found in southern Queensland and New 
South Wales, that is only three inches long. The Yellow-Footed, 
(P. flavipes) with yellow legs and feet, as its name indicates, 
ranges from eastern Australia to New Guinea. In Tasmania and 
the adjacent islands, a small variety is found, P. minima, or 
Little Pouched Mouse, with a body length of five inches, and 
tail three and one-half inches, and in Queensland a still smaller 
form is P. minutissima or Pigmy Pouched Mouse, only three inch- 
es long and tail two and one-half inches. Tasmania and southern 


198 Zoologica: N. Y. Zoological Society [Ti6 


Victoria has still another variety, P. swainsoni which has long 
soft fur. In West Australia is found the Freckled, (P. apicalis), 
which is freckled, reddish-grey above. Australia is well off for 
Marsupial Mice, and other varieties certainly will be found, as 
these little animals are easily overlooked. Insects form a'large 
part of their food. 


Marsupial or Pouched Mice, (Sminthopis) are slender and 
active little animals, from three to four inches long, with large 
ears and a well developed pouch in which they carry the three to 
four young they have at birth. They are terrestrial, insectivo- 
rous and do not often burrow. Their grey fur is soft and fine. 
Six species have been described, S. lavapinta, from Central Aus- 
tralia near the Finke River and the Fat-Tailed, (S. crassicau- 
data), which is found all over Australia except the north. Both 
of these little animals have the basal portion of their tail much 
thickened. The Common, (S. murina) is found over the south- 
ern and central portions of the continent. S. psammophila lives 
near Lake Amadeus in Central Australia, among the sand hills 
covered with tussocks of porcupine grass. The White-Footed, 
(S. leucopus) extends over eastern Australia from Cape York 
to Tasmania and is plentiful in certain localities. S. virginiae is 
known only in eastern Queensland. It is five inches long. 


An interesting animal found in southwestern and southern 
Australia, of which but one species is known, is the Marsupial 
Anteater, (Myrmecobius fasciatus). It frequents both the 
ground and hollows in the trees and its food consists of insects, 
generally. The fur is short and strong, of a general rufous color 
which darkens to black toward the tail, with prominent bands 
of white. It therefore is distinguished readily. The underparts 
are light yellowish. The females have no pouch, the young ad- 
hering to the nipples. It leaps along the ground like a squirrel 
with the tail slightly raised. They make charming pets and 
never attempt to bite. It measures ten inches long and its bushy 
tail seven inches. 


THE ECHIDNA. 


Of these most interesting animals, the Echidnas, or as they 
usually are called in Australia the Porcupine Anteater, there are 


Photograph by E. R. Sanborn 
FIG. 66. UNDER SURFACE OF THE ECHIDNA 
New York Zoological Park. 


2 we _ ates me 
Photograph by EB. R. Sanborn 


FIG. 67. ECHIDNA IN NORMAL ATTITUDE 
New York Zoological Park. 


<li 


Zoologica, Vol. II, No. 6. Face page 198 


1919] Le Souef: Mammals of Australia 199 


three varieties, namely, that found in New Guinea, (Echidna 
lawesii) , about fourteen inches in length, which has short spines; 
the Brown Echidna, (E. setosa) from Tasmania, with long hair 
almost concealing the spines, and the Australian form, (H. acu- 
leata) which is slightly larger than the New Guinea variety, 
being about seventeen inches long and without as much hair 
as the Tasmanian species. These animals live entirely on the 
ground and their food consists of small insects, and, especially 
termites and ants, which they easily can obtain by digging with 
their powerful claws into the ant’s nests or termites mounds. 
The tongue which is covered with sticky saliva is then protruded 
and when covered with ants is drawn back into the mouth. In 
captivity they are fed on finely chopped raw meat, eggs and milk. 
They have no true teeth, but have small spines at the back of 
the tongue. Their tail is rudimentary and the feet short and 
strong. When in danger, the animal rolls itself up something 
like a hedgehog. The sharp spines not only are presented to 
its enemy, but also are stuck into the ground, making it harder 
to lift up. The only way it can be carried conveniently is by 
grasping it by its hind feet, so that its head hangs down. It 
can dig in any hard soil by the aid of the spines as well as by 
the strong claws, and it is remarkable how quickly it seems to 
sink into the ground. It also can hold so tightly to the soil that 
it is only with difficulty that it can be raised, even by the aid 
of a spade or strong stick. It also is very difficult to dislodge 
from the corner of a room, and can climb over almost any wire 
fence and also out of any ordinary box at the corners, and unless 
the lid is very firmly nailed on, will push it off, and get through 
a very small aperture. The strength of the animal is astonish- 
ing and even if tightly fastened by a cord around one of its hind 
legs, is almost certain to get it off. These animals generally 
hibernate during the winter; usually under the surface of the 
ground, and frequently by a rock or rising ground. It is at this 
time that the egg is laid and the young hatched. The shell of 
the egg is soft and not calcified, and measures about half an inch. 
The mother by rolling herself up helps to protect the young 
which are in her pouch, and as the female has no nipples the 
young one when hatched has to lick the milk from the folds in 
the pouch. The young Echidna leaves the pouch just as the 


200 Zoologica: N. Y. Zoological Society [II1;6 


spines begin to appear and when it is a little over three inches 
long. The pouch then gradually disappears until the next breed- 
ing season. Like the kangaroo, it is very rarely that two young 
are born. These animals are more or less nocturnal, as are 
nearly all the Australian animals. 


PLATYPUS. 


The Duck-Billed Platypus, (Ornithorhynchus anatinus) is 
of great interest. Like the Echidna it belongs to the genus Mono- 
tremata, but passes its time in water and not on land, except 
when coiled up in its burrow with its tail tucked underneath, 
which usually is most of the day. It seeks its food generally in 
the evening or sometimes during the day in some very sheltered 
spot, feeding on earthworms, shell-fish, crustaceans and water 
insects, generally; a certain amount of which it can store in its 
cheek pouches. Although the young have rudimentary teeth, 
they have none when they reach an adult stage; horny plates 
developing in the place of them which enables the animals to 
masticate their food, which they usually do when lying on the 
surface of the water. The fur, which looks very much like that 
of a seal when the longer hairs are removed, is of two kinds; 
the longer being shiny and crisp and the under fur soft and short. 
The bill is soft and leathery, but shrinks considerably when dry, 
as in museum specimens. The underparts are lighter in color, 
usually greyish white. The tail is broad and flat and of a dark 
color above. The underpart is usually devoid of hair, especially 
in the older animals. Their eyes are very small, but as their 
bill is usually sensitive, they generally can find their insect prey 
by a sense of touch. 


Being unable to raise the body high from the ground like 
ordinary animals they can only shuffle along in an awkward 
manner. It is a burrowing animal and makes a long upward 
tunnel in the river banks, sometimes thirty feet in length, usu- 
ally starting at the roots of a tree that grows to the water, with 
the entrance generally under the surface of the water. At the 
end is a small chamber lined with leaves and grass, generally 
not so far from the surface of the ground, so that the natives 
frequently can tell where the nest is by striking the surface of 


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“APAdES UL UMOYS PUB YIOX MON 07 YSnoIq aq ued sndAze[d SurAll B 324} edoy A0F punoas 91931] 
SToroyy, “BI[B1ISNY FO episzNo dAl[e USES TOASU SI I ‘[BULLUE a[qeyABWAL JSOUI S14} JO S7IGBY POOF OY} 07 SUIMO 7eY} ABS 0} Yoadar aM 


TIIG-MONd YO ‘SNdALWTd AHL °89 “Dla 


1919] Le Souef: Mammals of Australia 201 


the ground above and listening for the echo. When swimming, 
the claws and web are stretched out to their full extent, but on 
land the extended web is always doubled up underneath; the end 
of the claws then coming in contact with the ground. In nearly 
all specimens in museums, the web is expanded beyond the claws, 
although the animal is represented as being in the ground. But 
that is incorrect.. These animals are very timid, and though 
they possess no external ears, they are very quick at hearing, 
and any suspicious sound makes them dive out of sight. The 
male measures about eighteen inches, tail six inches. The spur 
on its heel is larger than that on the female. The latter animal 
is about fourteen inches in length: The shell of the egg is of a 
tough, leathery texture, and from two to four eggs are laid at 
a time. As the mother has no pouch, she practically makes one 
by rolling herself up in her nesting chamber. She has no nip- 
ples, but the mammery glands which are in two groups, are un- 
derlying the skin on the underside, and the milk is pressed out 
by a contraction of the muscles and the young takes its food by 
applying its flat face and tongue to the lacteal surface. These 
animals are found in the rivers of Tasmania and eastern Aus- 
tralia, except the extreme north. 


MOLE. 


Australia possesses a marsupial mole, (Notoryctes typhlops) 
found in central and western Australia, but, naturally it never is 
seen in captivity. As a matter of fact they will not live in cap- 
tivity. It is about six inches long, with a curious ringed tail 
about an inch in length, and much thickened at the base. The 
nose has a hard shield. The fur is soft with an irredescent effect, 
and varies in color from a yellowish tint to chestnut-brown. They 
have two young at birth, live underground entirely, are without 
eyes and subsist on insect food. 


ZOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 


1590.53 


NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBERS 7,8 AND 9 


THE HIGHER VERTEBRATES OF BRITISH 


GUIANA 


WITH SPECIAL REFERENCE TO THE FAUNA OF 
BARTICA DISTRICT 


No.7. LIST OF AMPHIBIA, REPTILIA AND MAMMALIA 
No. 8. BIRDS OF BARTICA DISTRICT 


< 


_ No.9. LIZARDS OF THE GENUS AMEIVA 


By WILLIAM BEEBE 


Honorary Curator of Birds and Director of the Tropical 
Research Station 


Pot ene ER D BY THE SO) Cor ¥ 


tee OOLOGICAL PARK, NEW. YORK 
May, 1919 


Now York Zoological Suciety 


General Office, 111 Broadway, New York City 5 


Offirers 
President, HENRY FAIRFIELD OSBORN; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 
Treasurer, PERCY R. PYNE. 


Board of Managers 
Glass of 1919 , 
PERCY R. PYNE, GEORGE BIRD GRINNELL, GEORGE C. CLARK 
CLEVELAND H. DopGr, C. LEDYARD BLAIR, FREDERICK 
GILBERT BOURNE, EMERSON MCMILLIN, ANTHONY 
R. KUSER, WATSON B. DICKERMAN, MoRTI- 
MER L. SCHIFF, FREDERIC C. WALCOTT, 
BEEKMAN WINTHROP. 
Olass of 1920 ; 

HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. 
DIETERICH, GEORGE F’. BAKER, WM. PIERSON HAMILTON, 
ROBERT S. BREWSTER, EDWARD S. HARKNESS, 
WILLIAM B. Oscoop FIELD, A. BARTON 
HEPBURN, WILLIAM WOODWARD, 

EDWIN THORNE, PERCY A. 

ROCKEFELLER. 

Glass uf 1921 
Levi P. MorTON, ANDREW CARNEGIE, MADISON GRANT, WILLIAM 
WHITE NILES, HENRY A. C. TAYLOR, FRANK K. STURGIS, ~ 
GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFORD 

Morris, ARCHER M. HUNTINGTON, HENRY 
M. TILFoRD, E. C. CONVERSE. 


Scientific Stat 
WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 
WILLIAM BEEBE, Honorary Curator of Birds and Director of the 
Tropical Research Station; 

LEE S. CRANDALL, Curator of Birds; 

GEORGE S. HUNTINGTON, Prosector; 

GEORGE A. MACCALLUM, Pathologist; 

W. REID BLAIR, Veterinarian; 

ELWIN R. SANBORN, Photographer and Editor. 


Editorial Committee 
HENRY FAIRFIELD OSBORN, Chairman; MADISON GRANT, 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


: 
Lae +, , 

eS eS eae | 

ae See Se 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBERS 7, 8 AND 9 


THE HIGHER VERTEBRATES OF BRITISH 


GUIANA 


WITH SPECIAL REFERENCE TO THE FAUNA OF 
BARTICA DISTRICT 


No. 7. LIST OF AMPHIBIA, REPTILIA AND MAMMALIA 
Nos: =BIRDS OF BARTICA DISTRICT 
No.9. LIZARDS OF THE GENUS AMEIVA 


By WILLIAM BEEBE 


Honorary Curator of Birds and Director of the Tropical 
Research Station 


Plies El So ED Buys THE SOy Cal aay. 
eer AO) OG Tl CA ie PARK NE Wo -Y ORK 


May, 1919 


Volume II, Number 7 


HIGHER VERTEBRATES OF 
BRITISH GUIANA 


WITH SPECIAL REFERENCE TO THE FAUNA OF 
BARTICA DISTRICT 


By WituiaAmM BEEBE 
Director, Tropical Research Station 


In the course of recent studies of neotropical vertebrates, with 
reference to future ecological work in the field at the Tropical Re- 
search Station of the New York Zoological Society, I have been in 
constant need of lists of the Vertebrata of British Guiana.’ 

Finding no résumé available of the Amphibia, Reptilia and 
Mammalia of this colony, I have gone through the literature at 
hand and made my own lists. These I offer as a preliminary 
enumeration of the species thus far recorded in literature, or in my 
own collections, from this British Colony. They form a tangible 
basis for future increments—the many new species and the radical 
extension of present known distributions which intensive study of 
these phyla in British Guiana is certain to achieve. Check-lists of 
mere names such as these are wholly foreign to the future zoological 
work of the Tropical Station, but they are absolutely necessary as 
a basis for identification and investigation, and it is in this spirit 
that this preliminary work has been undertaken. 

I have made no attempt at a thorough search of literature for 
priority or for confirmation of names or other similar phases of tax- 
onomy, deeming this the special province of the literary systematist. 
I have merely sought to utilize the most recently accepted names of 
herpetologists and mammalogists. In general I have followed the 
classification of Gadow in the Cambridge Natural History, Volume 

‘A preliminary survey of the fishes has been made by Prof. C. H. Eigenmann 
in his ‘‘ Freshwater Fishes of British Guiana,’’ and lists of the birds are avail- 
able in Brabourne and Chubb’s ‘‘ The Birds of South America,’’ in Chubb’s 


unfinished ‘‘ Birds of British Guiana,’’ and in my ‘‘ Tropical Wild Life,’’ published 
by the Zoological Society in 1917. 


205 


206 Zoologica: N. Y. Zoological Society LED SAg 


VIII, for Amphibia and Reptilia, and that of Osborn in “‘The Age of 
Mammals” for Mammalia. Dr. J. A. Allen has kindly criticized 
the mammalian names, and to John Tee-Van, a member of my 
Tropical Station staff, I am indebted for untiring search through 
many scores of volumes for British Guiana records. 


The lists of names are unannotated, and in each case I have given 
one authentic record for British Guiana, not necessarily the first in 
published priority. An asterisk before a name indicates that the 
species has been collected within a radius of ten miles of the Zoological 
Society’s Research Station in Bartica District. This locality is 
indicated on the adjoining map. 


The total number of species is as follows: 


British Bartica 
Guiana District 


I. Class AMPHIBIA (Coecilians, Frogs and Toads) 52 35 


Me OClasseREP TMA AC o.oo. So bos ie oa ae 2 54 
CHELONIA (Turtles and Tortoises) eae egal 5 
CROCODIMIA(Crocodiles)= 79 30> ae re 4 1 
MACK ie EMMA iChiZards) nd ee ee eee 34 ral | 
OPRHTDEANSHakes)i. . yt (oe en 2 eee as 63 27 

Pie Class: MEAMIMIALTA = 22.22) We Sa Be oye eee 119 48 
MARSUPRTIALTIA(Opessums)io. 22 10 5 
CHIROP TGR An(Bats\e Wo ates once eee 36 3 
CARNIVORA (Jackals, Raccoons, Otters and 

Cats} as olenres) IMEI e ed iaare CI Mes AnN, 16 a 
RODENTIA (Agoutis, Porcupines, Rats and 

NICS: Cagle, eee Nl a re ont he Oa rat 8 
EDENTATA (Sloths, Anteaters and Arma- 

UNOS) ah rsa tetas 08 Se annie ee nh 8 cee Delp ae 8 8 
PRIMATES @ilonkeys))) "ae O22 See ee § 5 
ARTIODACTYLA (Peccaries and Deer) 018 4 
PEWS oODACTVALAY Gaping). "So .cce eee 1 1 
BLRIEIN TA (Sea-cows)i) (5.45) Sh ee 1 il 
ODONTOCETI (Dolphins and Whales) .... 3 if 


1919) Beebe: Amphibia, British Guiana 207 


CLASS: AMPHIBIA 
Order APODA; Limbless, Burrowing Coecilians 


Family Coecilidae 


Coecilia gracilis Shaw 
Boul. Cat. Batr. Brit. Mus. 1882. 


Siphonops annulatus (Mikan.) 
Beebe Coll. Bartica Dist. 1916. 


Order ANURA; Frogs and Toads 
Family Pipidae 


*Pipa americana Laur 


Boul. Cat. Batr. Brit. Mus. 1882, p. 459 


Family Bufonidae 
Bufo guttatus Schneid. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 291 - 
*Bufo molitor Tschudi 
Beebe Coll. Bartica Dist. 1916 


*Bufo marinus (Linné) 


Boul. Cat. Batr. Brit. Mus. 1882, p. 316 


*Bufo typhonius (Linné) 
Beebe Coll. Bartica Dist. 1916 


*Bufo sternosignatus Keferst. 
Beebe Coll. Bartica Dist. 1916 


Family Hylidae 

*Hyla maxima (Laur.) 

Boul. Cat. Batr. Brit. Mus. 1882, p. 349 
*Hyla crepitans Wied 

Boul. Cat. Batr. Brit. Mus. 1882, p. 352 
*Hyla indris (Cope) 

Boul. Cat. Batr. Brit. Mus. 1882, p. 353 
*Hyla pardalis Spix 

Beebe Coll. Bartica Dist. 1916 

Hyla albomarginata Spix 
Boul. Trans. Linn. Soc. London, (2) VILL, 1900, p. 56 


*Hyla punctata (Schneid.) 
Beebe Coll. Bartica Dist. 1916 


208 Zoologica: N. Y. Zoological Society 


Hyla granosa Boul. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 358 


Hyla fasciata (Cope) 
Schomb. Reise Brit. Guiana, III, 1848, p. 660 


*Hyla boans Daud. 
Beebe Coll. Bartica Dist. 1909 


Hyla leprieurii Dum. & Bibr. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 362 


Hyla taurina (Steindachn.) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 363 


Hyla venulosa (Laur.) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 365 


Hyla aurantiaca Daud. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 388 


Hyla acuminata Cope 
Boul. Cat. Batr. Brit. Mus. 1882, p. 403 


*Hyla rubra Daud. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 404 


*Hyla lineomaculata Werner 
Beebe Coll. Bartica Dist. 1909 


Hyla evansi Boul. 
Proc. Zool. Soc. London, 1904,-II, p. 106 


Phyllomedusa bicolor (Boddaert) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 427 


Phyllomedusa hypochondrialis Daud. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 430 


Family Cystignathidae 
*Pseudis paradoxa (Linné) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 186 


*Ceratophrys cornuta (Linné) 
Beebe Coll. Bartica Dist. 1909 


Eleutherodactylus marmoratus (Boul.) 


Boul. Trans. Linn. Soe. London, (2) VII, 1900, p. 56 


*Leptodactylus lineatus (Schneid.) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 207 


Leptodactylus hylaedactylus (Cope) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 240 


(7. 


1919] Beebe: Amphibia, British Guiana 


*Leptodactylus longirostris Boul. 
Beebe Coll. Bartica Dist. 1916 


*Leptodactylus pentadactylus (Laur.) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 242 


*Leptodactylus poecilochilus (Cope) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 244 


*Leptodactylus typhonius (Daud.) 
Beebe Coll. Bartica Dist. 1916 


*Leptodactylus ocellatus (Linné) 
Beebe Coll. Bartica Dist. 1909 


*Leptodactylus caliginosus Girard 
Beebe Coll. Bartica Dist. 1916 


*Leptodactylus gaudichaudii (Dum. & Bibr.) 
Beebe Coll. Bartica Dist. 1916 


Family Engystomatidae 
Oreophrynella Quelchii Boul. 


Boul. Ann. Mag. Nat. Hist. (6) XV, 1895, p. 522. 


Oreophrynella Macconnelli Boul. 


Boul. Trans. Linn. Soc. London, (2) VIII, 1900, p. 55 


*Otophryne Robusta Boul. 


Boul. Trans. Linn. Soc. London, (2) VIII, 1900, p. 56 


*Atelopus Proboscideus Boul. 
Beebe Coll. Bartica Dist. 1916 


*Atelopus varius (Stannius) \) 
Beebe Coll. Bartica Dist. 1916 


*Atelopus flavescens (Dum. & Bibr.) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 154 


*Atelopus pulcher Boul. 
Beebe Coll. Bartica Dist. 1916 


*Engystoma microps Dum. & Bibr. 
Boul. Cat. Batr. Brit. Mus. 1882, p. 163 


Family Ranidae 


*Prostherapis inguinalis Cope 


Boul. Cat. Batr. Brit. Mus. 1882, p. 189 


*Prostherapis trinitatus (Garman) 
Beebe Coll. Bartica Dist. 1916 


*Dendrobates tinctorius (Schneid.) 
Beebe Coll. Bartica Dist. 1916 


209 


210 Zoologica: N. Y. Zoological Society [Th3% 


*Dendrobates trivittatus (Spix) 
Boul. Cat. Batr. Brit. Mus. 1882, p. 144 


*Rana palmipes Spix 
Beebe Coll. Bartica Dist. 1916 


CLASS: REPTILIA 
Order CHELONIA ; Turtles and Tortoises 


Suborder Crypropira 


Family Cinosternidae 


*Cinosternum scorpioides (Linné) 
Beebe Coll. Bartica Dist. 1916 


Family Testudinidae 
*Nicoria punctularia (Daud.) 
Boul. Cat. Chel. Brit. Mus. 1889, p. 124 
*Testudo tabulata Walb. 
Beebe Coll. Bartica Dist. 1916 


Family Chelonidae 


Chelonia mydas (Linné) 
Leechman; Handbook Brit. Guiana, 1918, p. 134 


Suborder PLeuRopIRA 


Family Pelomedusidae 
Podocnemis unifilis Trosch. 
Schomb. Reise Brit. Guiana, III, 1848, p. 647 


*Podocnemis expansa (Schwigg.) 
Beebe Coll. Bartica Dist. 1916 


Podocnemis tracaxa (Spix 


) 
Schomb. Reise Brit. Guiana, III, 1848, p. 646 


Family Chelydidae 
*Chelys fimbriata (Schneid.) 
Schomb. Reise Brit. Guiana, III, 1848 
Hydraspis tuberosa (Peters) 
Boul. Cat. Chel; Brit.’ Mus. 1889, p. 223 
Hydraspis gibba (Schweigg.) 
Boul. Cat. Chel. Brit. Mus. 1889, p. 224 


Platemys platycephala (Schneid.) 
Boul. Cat. Chel. Brit. Mus. 1889, p. 228 


1919] Beebe: Reptilia, British Guiana ZU) 


Order CROCODILIA ; Crocodiles 
Family Crocodilidae 


Caiman niger Spix 


Schomb. Reise Brit. Guiana, III, 1848, p. 647 


*Caiman sclerops (Schneid.) 


Schomb. Reise Brit. Guiana, III, 1848, p. 648 


Caiman trigonatus (Schneid.) 
Boul. Cat. Croc. Brit. Mus. 1889, p. 296 


Caiman palpebrosus (Cuv.) 
Schomb. Reise Brit. Guiana III, 1848, p. 648 


Order LACERTILIA ; Lizards 


Family Geckonidae 
Gonatodes annularis Boul. 
Bouls PoZe SS. 188 pps Lda 


*Thecadactylus rapicaudus (Houtt.) 
BoulCat. Liz.-Brit.Mus. 1, 1885,i:p.-112 


*Hemidactylus mabouia (de Jonnés) 
Beebe Coll. Bartica Dist. 1916 


Family Iguanidae 
Anolis alligator Dum. & Bibr. 
Boul. Cat. Liz. Brit. Mus. III, 1887, p. 500 


*Anolis sagrae Dum. & Bibr. 
Beebe Coll. Bartica Dist. 1916 


*Anolis fusco-auratus D’Orb. 


Beebe Coll. Bartica Dist. 1916 


*Anolis ortonii Cope 


Beebe Coll. Bartica Dist. 1916 


Anolis punctatus Daud. 
Boul. P.Z. S887 -p. 158 


*Anolis chrysolepis Dum. & Bibr. 
Boul. Cat. Liz. Brit. Mus. II, 1885, p. 90. 


Norops auratus (Daud.) 
Boul Cat. liz? Brit. Mus. Il; 1885; p. 96 


*Polychrus marmoratus (Linné) 


Boul. Cat. Liz. Brit. Mus. II, 1885, p. 99 


212 Zoologica: N. Y. Zoological Society 


Ophryoessa superciliosa (Linné 


) 
Boul. Cat. Liz. Brit. Mus. II, 1885, p. 112 


Tropidurus torquatus (Wied) 
Schomb. Reise Brit. Guiana III, p. 650 


*Plica umbra (Linné) 
Boul. Cat. Liz. Brit. Mus. II, 1885, p. 180 


*Plica plica (Linné) 
Boul. Cat. Liz. Brit. Mus. II, 1885, p. 181 


Urocentron azureum (Linné) 


Boul. Cat. Liz. Brit. Mus. II, 1885, p. 183 


*Iguana iguana (Linné) 


Boule ‘Cat; Liz, Brit. Muse i, 1885,-p. 190 


Family Teiidae 


Tupinambis teguixin (Linné) 


Schomb. Reise Brit. Guiana III. p. 65 


*Tupinambis nigropunctatus Spix 


Boul. Cat. Liz. Brit. Mus. II, 1885, p. 337 


*Centropyx intermedius (Schleg.) 
Boul, Cate iz. Brit? Mus.) 11,7 1885, ps 841 


*Centropyx calcaratus (Spix) 
Boul. Cat! Liz-? Brita. Mus. Il, 1885; pas42 


*Ameiva surinamensis (Laur.) 


Boul. Cat. Liz. Brit. Mus. II, 1885, p. 353 


*Ameiva punctata Gray 
Boul..Catz* lize; Brit; -Musa 11, 1885;p., 360 


*Cnemidophorus murinus (Laur.) 
Beebe Coll. Bartica Dist. 1916 


*Cnemidophorus lemniscatus lemniscatus (Daud.) 


Boul. .Gat.. Yaz., Brit: Mus. Wf, 1885, p.364 


Crocidilurus lacertinus (Daud.) 


Schomb. Reise Brit. Guiana III, p. 650 


*Prionodactylus oshaughnessyi Boul. 
Beebe Coll. Bartica Dist. 1916 


*Cophias flavescens (Bonnat.) 
Boul. "P2Ze5. 168% sp. 153 


Iphisa elegans Gray 
Boul, Cat. * Liza Brit. “Muss ile Ass5sp.425 


Calliscinopus agilis Ruthven 


Ruth. Occ. Papers Univ. Mich. No. 22, 1916, p. 3 


ed ee 


1909) Beebe: Reptilia, British Guiana 


Family Amphisbaenidae 


*Amphisbena fuliginosa Linné 


Boul. Cat. Lig. Brit. Mus. II, 1885, p. 438 


Family Scincidae 


Mabuia punctata (Gray) 
Boul. Cat. lig. Brit: -Nius’ Itt, 1685; p: tol 


*Mabuia aurata (Schneid.) 
Boule Gas big. Brite: Mus. hi, 1885s"p...190 


Mabuia agilis (Raddi) 
howl Cate lige prt. Mus. IPE. 18855" p.- i91 


Order OPHIDIA ; Snakes 


Family Typhlopidae 


*Typhlops reticulatus (Linné) 
Boul. Cat. Snakes Brit. Mus. I, 1893, p. 28 


Typhlops lumbricalis (Linné) 
Boul. Cat. Snakes Brit. Mus. I, 1898, p. 31 


Family Boidae 


*Epicrates cenchris (Linné) 


Boul. Cat. Snakes Brit. Mus. I, 1898, p. 95 


Corallus cookii Gray 


Boul. Cat. Snakes Brit. Mus. I, 1893, p. 100 
*Corallus hortulanus (Linné) 

Boul. Cat. Snakes Brit. Mus. I, 1898, p. 102 
*Corallus caninus (Linné) 

Boul. Cat. Snakes Brit. Mus. I, 1898, p. 103 
*Eunectes murinus (Linné) 

Boul. Cat: Snakes’ Brit. Mus. f,. 1893, p. 115 


*Boa constrictor (Linné) 
Beebe Coll. Bartica Dist. 1916 
Family Ilysiidae 


*Ilysia scytale (Linné) 
Boul. Cat. Snakes Brit. Mus. I, 1898, p. 134 


213 


214 Zoologica: N. Y. Zoological Society (TL; 7, 


Family Colubridae 


*Helicops angulatus (Linné) 
Boul. Cat. Snakes Brit. Mus. I, 1898, p. 279 


Drymobius boddaertii (Sentzen) 
Boul. Cat. Snakes Brit. Mus. IJ, 1894, p. 13 


*Phrynonax sulphureus (Wagl.) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 19 


Phrynonax fasciatus (Peters) 
Boul. Cat. Snakes Brit. Mus. IJ, 1894, p. 22 


Spilotes pullatus (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 23 


*Coluber corais Boie 


Boul. Cat. Snakes Brit. Mus. II, 1894, p. 32 


*Herpetodryas sexcarinatus (Wagl.) 
Beebe Coll. Bartica Dist. 1916 


*Herpetodryas carinatus (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 74 


*Herpetodryas fuscus (Linné) 
Boul. Cat. Snakes Brit. Mus. 11, 1894, p. 75 
*Leptophis liocercus (Wied) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 114 
Liophis typhlus (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 186 


Liophis reginae (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 138 


*Xenedon colubrinus Gunther 


Beebe Coll. Bartica Dist. 1916 


*Xenedon severus (Linné) 


Boul. Cat. Snakes Brit. Mus. II, 1894, p. 149 


*Xenedon merremii (Wagl.) 


Boul. Cat. Snakes Brit. Mus. I2; 1894, p: 150 


Aporophis lineatus (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 158 


Rhadinaea cobella (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 167 


Rhadinaea purpurans (Dum. & Bibr.) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 168 


Urotheca bicincta (Hermann) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 184 


TOLD Beebe: Reptilia, British Guiana 


Dimades plicatilis (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 186 


Hydrops triangularis (Wag].) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 187 


Petalognathus nebulatus (Linné) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 294 


Atractus torquatus (Dum. & Bibr.) 
Boul. Cat. Snakes Brit. Mus. II, 1894, p. 309 


Atractus trilineatus Wag. 


Boul. Cat. Snakes Brit. Mus. II, 1894, p. 312 


*Lycognathus cervinus (Laur.) 
Beebe Coll. Bartica Dist. 1916 


Trypanurgos compressus (Daud.) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 59 


Himantodes cenchoa (Linné) 


Schomb. Reise Brit. Guiana III, 


*Leptodeira albofusca (Lacép.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 96 


Oxyrhopus petolarius (Linné) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 102 


Oxyrhopus trigeminus Dum. & Bibr. 
Boul. Cat. Snakes Brit Mus. III, 1896, p. 104 


Oxyrhopus cloelia (Daud.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 109 


Oxyrhopus coronatus (Schneid.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 112 


Rhinostoma guianense (Trosch) 


Schomb. Reise. Brit. Guiana, III, 1848, p. 653 


Thamnodynastes nattereri (Mikan.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 117 


Thamnodynastes punctatissimus (Wagl.) 


Boul. Cat. Snakes Brit. Mus. II], 1896, p. 117 


Philodryas viridissimus (Linné) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 129 


Oxybelis fulgidus (Daud.) 
Schomb. Reise. Brit. Guiana III, 1848 


*Oxybelis acuminatus (Wied) 
Beebe Coll. Bartica Dist. 1916 


Erythrolamprus aesculapii (Linné) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 202 


215 


216 Zoologica: N. Y. Zoological Society [aS 
*Tantilla melanocephala (Linné) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 216 


Apostolepis quinquelineata Boul. 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 235 


Family Elapidae 
Micrurus surinamensis (Cuv.) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 414 


*Micrurus psyches (Daud.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 427 


Micrurus marcgravii (Wied) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 429 


Micrurus lemniscatus (Linné) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 430 


Family Amblycephalidae 


Leptognathus catesbyi (Sentzen) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 449 


Leptognathus pavonina (Schleg.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 450 


Leptognathus variegata Dum. & Bibr. 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 451 


Leptognathus leucomelas Boul. 
Beebe Coll. Bartica Dist. 1917 


*Dipsas bucephala (Shaw) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 462 


Family Crotalidae 


*Lachesis mutus (Linné) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 534 


*Lachesis lanceolatus (Lacép.) 


Boul. Cat. Snakes Brit. Mus. III, 1896, p. 536 


*Lachesis atrox (Linné) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 539 


*Crotalus terrificus (Laur.) 
Boul. Cat. Snakes Brit. Mus. III, 1896, p. 575 


1919] Beebe: Mammalia, British Guiana 


217 
CLASS: MAMMALIA 


Order MARSUPIALIA ; Opossums 


Family Didelphidae 


*Didelphis marsupialis marsupialis Linné 
Linnaeus Guiana Opossum; Crab-eating Yawarri 
Thomas, Cat. Mar. Brit. Mus. 1888, p. 327 


*Metachirus opossum opossum (Linné) 
White-faced Opossum; Quica 


Thomas, Cat. Mar. Brit. Mus. 1888, p. 332 
Metachirus nudicaudatus nudicaudatus (E. Geoff.) 
Bare-tailed Opossum 

Quelch, Animal Life in Brit. Guiana, 1901, p. 79 


Lutreolina crassicaudata (Desm.) 
Thick-tailed Opossum 


Thomas, Cat. Mar. Brit. Mus. 1888, p. 336 
Caluromys philander (Linné) 
Woolly Opossum 

Thomas, Cat. Mar. Brit. Mus. 1888, p. 338 


*Marmosa murina murina (Linné) 
Linnaeus Mouse Opossum; Little Yawarri 
Thomas, Cat. Mar. Brit. Mus. 1888, p. 346 
Marmosa chloe Thomas 
Chloe’s Opossum 
Thomas, Ann. Mag. Nat. Hist. (7) XX, p. 167 


Marmosa cinerea demerarae Thomas 
Demerara Ashy Opossum 


Thomas, Ann. Mag. Nat. Hist. (7) XVI, p. 313 


*Peramys brevicaudatus (Erxl.) 
Short-tailed Opossum 


Quelch, Animal Life Brit. Guiana, 1901, p. 80 


*Chironectes minimus (Zimm.) 
Water Opossum; Yapock 


Thomas, Cat. Mar. Brit. Mus. 1888, p. 370 


218 Zoologica: N. Y. Zoological Society euiie 


Order CHIROPTERA ; Bats 


Family Emballonuridae 


Rhynchincus naso (Wied) 
Guiana River Bat 
Dobson, Cat. Chir. Brit. Mus. 1878, p. 369 


*Saccopteryx leptura (Schr.) 
Young, Timehri, (2) X, 1896, p. 44 


Saccopteryx bilineata (Temm.) 
‘Thomas,,-Ann. Mag. Nat?Hist. <7) Vili 1901 pe 240 


Peropteryx canina (Wied) 
Thomas, Ann. Mag. Nat. Hist. (7) VIII, 1901, p. 140 


Cyttarops alecto Thomas 
Thomas, Ann. Mag. Nat: Hist. (8) XI, 1913, p. 136 


Family Noctilionidae 


Noctilio leporinus leporinus (Linné) 


Dobson, Cat. Chir. Brit. Mus. 1878, p. 416 


Dirias albiventer (Spix) 
Young, Timehri, (2) X, 1896, p. 45 


Family Phyllostomidae 


Micronycteris megalotis megalotis Gray 


Thomas, Ann. Mag. Nat. Hist. (7) VIII, 1901, p. 142 


Dolichophyllum macrophyllum Wied 
Quelch, Timehri, (2) VI, 1892, p. 97 


Phyllostomus hastatus hastatus (Pallas) 
Dobson, Cat. Chir. Brit. Mus. 1878, p. 486 


Phyllostomus discolor Wagner 
Young, Timehri, (2) X, 1896, p. 45 


Phyllostomus elongatus Geoff. 
Young, Timehri, (2) X, 1896, p. 45 


Phyllostomus latifolius Thomas 
Thomas, Ann. Mag. Nat. Hist. (7) VIII, 1901, p. 142 


Vampyrus spectrum (Linné) 
False Vampire 
Dobson, Cat. Chir. Brit. Mus. 1878, p. 471 


ewe Beebe: Mammalia, British Guiana 219 
Glossophaga soricina (Pallas) 
Quelch, Timehri, (2) VI, 1892, p. 97. 


Hemiderma perspicillatum (Linné) 


Thomas, Ann. Mag. Nat. His. (7) VIII, 1901, p. 143 


Rhinophylla pumilio Peters 
Young, Timehri, (2) X, 1896, p. 46 


Uroderma bilobatum Peters 
Quelch, Timehri, (2) VI, 1892, p. 97 


Artibeus planirostris fallax Peters 
Anderson, Proc. Zool. Soc. 1908, p. 243 


Artibeus cinereus cinereus (Gervais) 


Anderson, Proc. Zool. Soc. 1908, p. 291 


Artibeus quadrivittatus Peters 
Young, Timehri, (2) X, 1896, p. 46 


Family Desmodontidae 


*Desmodus rotundus (Geoff.) 
Common Vampire; Colony-Doctor 
Quelch, Timehri, (2) VI, 1892, p. 97 


Diaemus youngi (Jent.) 
Young’s Vampire 
Young, Timehri, (2) X, 1896, p. 46 
Family Natalidae 
Natalus stramineus Gray 
Young, Timehri, (2) X, 1896, p. 44 
Family Furipteridae 
Furipterus horrens (F.. Cuv.) 
Thomas, Proc. Zool. Soc. 1887, p. 151 
Family Thyroptidae 
Thyroptera tricolor Spix 
Quelch, Timehri, (2) VI, 1892, p. 97 
Family Vespertilionidae 


Myotis nigricans (Wied) 
Young, Timehri, (2) X, 1896, p. 44 


220 Zoologica: N. Y. Zoological Society [Lei 


Eptesicus hillarii (Geoff.) 
Thomas, Proc. Zool. Soc. 1887, p. 151 


Lasiurus borealis borealis (Miill) 


Quelch, Timehri, (2) VI, 1892, p. 97 


Dasypterus intermedius (H. Allen) 
Young, Timehri, (2) X, 1896, p. 44 


Dasypterus ega ega (Gerv.) 
Quelch, Timehri, (2) VI, 1892, p. 97 


Family Molossidae 
*Molossops planirostris Peters 
Dobson, Cat. Chir. Brit. Mus. 1878, p. 409 


Eumops abrasus (Temm.) 
Dobson, Cat. Chir. Brit. Mus. 1878, p. 416 


Eumops maurus (Thomas) 
Thomas, Ann. Mag. Nat. Hist. (7) VIII, 1901, p. 141 


Molossus rufus Geoff. 
Young, Timehri, (2) X, 1896, p. 45 


Molossus obscurus Geoff. 


Quelch, Timehri, (2) VI, 1892, p. 97 


Order CARNIVORA ; Dogs, Raccoons, Otters 
and Cats 


Family Canidae 


*Cerdocyon thous thous (Linné) 
Guiana Jungle Jackal; Rough Fox; Crab-dog 
Quelch, Animal Life Brit. Guiana, 1901, p. 43 


Cerdocyon thous savannarum Thomas 
Savanna Jackal 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 146 


Icticyon venaticus Lund. 
Guiana Hunting Dog; Bush-dog; Warracabra Tiger 
Quelch, Animal Life Brit. Guiana, 1901, p. 42 


Family Procyonidae 


*Procyon cancrivorus (G. Cuvier) 
Crab-eating Raccoon; Crab-dog 
Quelch, Animal Life Brit. Guiana, 1901, p. 45 


1919) Beebe: Mammalia, British Guiana 


*Nasua nasua nasua (Linné) 
Black Coati; Kibihee; Quashi 
Quelch, Animal Life Brit. Guiana, 1901, p. 46 


*Nasua rufa (lIll.) 
Red Coati; Kibihee 
Quelch, Animal Life Brit. Guiana, 1901, p. 45 


*Potos flavus flavus (Schreber) 
Guiana Kinkajou; Potto:; Night Monkey 
Quelch, Animal Life Brit. Guiana, 1901, p. 46 


Family Mustelidae 


*Tayra barbara barbara (Linné) 
Guiana Tayra; Galictis; Hacka 
Quelch, Animal Life Brit. Guiana, 1901, p. 47 


*Grison allamandi (Bell) 
Guiana Grison 
Quelch, Animal Life Brit. Guiana, 1901, p. 47 


*Pteronura brasiliensis (Zimm.) 
Fin-tailed Otter; Water-dog 
Quelch, Animal Life Brit. Guiana, 1901, p. 49 


Family Felidae 


*Panthera onca (Linné) 
Jaguar; Tiger; Black Tiger; Maipurie Tiger 
Quelch, Animal Life Brit. Guiana, 1901, p. 32 


*Felis couguar Kerr 
Puma; Deer Tiger 
Quelch, Animal Life Brit. Guiana, 1901, p. 36 


*Herpailurus jaguarondi unicolor (Traill) 
Guiana Jaguarondi; Hacka Tiger; Eyra 
Quelch, Animal Life Brit. Guiana, 1901, p. 38 


*Leopardus pardalis tumtumari (Allen) 
Guiana Ocelot; Tiger Cat; Labba Tiger 


Allen, Bull. Am. Mus. Nat. Hist. 34, 1915, p. 


Margay tigrina tigrina (Schreber) 
Margay Cat; Wild Cat 
Quelch, Animal Life Brit. Guiana, 1901, p. 41 


Family Viverridae 


Mungos mungo (Gmelin) 
Mongoose 
[Introduced] Beebe Coll. Georgetown, 1916 


632 


221 


PP? Zoologica: N. Y. Zoological Society pags 


Order RODENTIA ; Agoutis, Porcupines, Rats 
and Mice 


Family Hydrochaeridae 


*Hydrochaerus hydrochaerus Linné 
Capybara; Waterhaas; Waterhare 
Quelch, Animal Life Brit. Guiana, 1901, p. 28 


Family Caviidae 


Cavia porcellus guianae Thomas 
Guiana Guinea Pig 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 152 


Family Dasyproctidae 


*Dasyprocta croconota prymnolopha (Wagl.) 
Agouti; Accourie 
Quelch, Animal Life Brit. Guiana, 1901, p. 27 


*Agouti paca paca (Linné) 
Paca; Labba 
Quelch, Animal Life Brit. Guiana, 1901, p. 26 


*Myoprocta acouchy (Erxl.) 
Pigmy Agouti; Adourie 
Quelch, Animal Life Brit. Guiana, 1901, p. 28 


Family Erethizontidae 


*Coendou prehensilis (Linné) 
Common Tree Porcupine 
Quelch, Animal Life Brit. Guiana, 1901, p. 31 


Coendou melanurus (Wagner) 
Black-tailed Tree Porcupine 
Quelch, Animal Life Brit. Guiana, 1901, p. 31 


Family Octodontidae 


*Proechimys cayennensis Desm. 
Cayenne Spiny Rat 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 152 


Loncheres guianae Thomas 
Guiana Spiny Rat 
Quelch, Animal Life Brit. Guiana, 1901, p. 31 


BOLO Beebe: Mammalia, British Guiana 223 


Family Muridae 


Mus musculus musculus Linné 
Common House Mouse 
[Introduced] Beebe Coll. Georgetown, 1916 


Epimys rattus alexandrinus (Geoff.) 
Black Rat 
[Introduced] Beebe Coll. Georgetown, 1916 


Rhipidomys sclateri Thomas 
Thomas, Proc. Zool. Soc. 1887, p. 152 


Rhipidomys macconnelli de Winton 
De Winton, Trans. Linn. Soc. (2), VIII, 1900, p. 52 


Rhipidomys nitela Thomas 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 148 


Rhipidomys milleri Allen 
Allen, Bull. Am. Mus. Nat. Hist. XXXII, 1918, p. 602 


Holochilus guianae Thomas 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 149 


Nectomys squamipes Brandt 
Thomas, Proc. Zool. Soc. 1887, p. 151 


Sigmomys savannarum Thomas 


Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 150 


Oryzomys navus messorius Thomas 


Thomas; Ann. Mag. Nat. Hist. (7), VIII; 1901, p. 151 


Zygodontomys stellae Thomas 
Thomas, Ann. Mag. Nat. Hist: (7), VIII, 1901, p. 152 


Oecomys rex Thomas 
Thomas, Ann. Mag. Nat. Hist. (8), VI, 1910, p. 504 


Oecomys nitedulus Thomas 
Thomas, Ann. Mag. Nat. Hist. (8), VI, 1910, p. 505 


Family Sciuridae 


*Sciurillus pusillus pusillus (Desm.) 
Common Dwarf Squirrel 
AllensaB ull. Amie Wrus. Nat. ddist. SOX TV, 19155 p. 197 


Sciurillus pusillus glaucinus Thomas 
Grayish Dwarf Squirrel 
Thomas, Ann. Mag. Nat. Hist. (8), XIII, 1914, p. 575 


*Guerlinguetus aestuans aestuans (Linné) 
Common Jungle Squirrel 
Allen, Bull. Am. Mus. Nat. Hist. XXXIV, 1915, p. 257 


224 Zoologica: N. Y. Zoological Society (TTS 


Guerlinguetus aestuans macconnelli (Thomas) 
Macconnell’s Jungle Squirrel 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 148 


Guerlinguetus aestuans quelchii (Thomas) 
Quelch’s Jungle Squirrel 
Thomas, Ann. Mag. Nat. Hist. (7), VIII, 1901, p. 147 


Order EDENTATA ; Sloths, Anteaters and 


Armadillos 


Family Bradypodidae 


*Bradypus tridactylus Linné 
Three-toed Sloth; Gray Sloth; Ai 
Quelch, Animal Life Brit. Guiana, 1901, p. 64 


*Choloepus didactylus (Linné) 
Two-toed Sloth; Brown Sloth; Unau 
Quelch, Animal Life Brit. Guiana, 1901, p. 67 


Family Myrmecophagidae 


*Myrmecophaga tridactyla tridactyla Linné 
Guiana Great Anteater; Antbear; Tamanoir 
Quelch, Animal Life Brit. Guiana, 1901, p. 73 


*Tamandua tetradactyla tetradactyla (Linné) 
Guiana Tamandua; Lesser Anteater 
Quelch, Animal Life Brit. Guiana, 1901, p. 74 


*Cyclopes didactylus didactylus (Linné) 
Guiana Silky Anteater; Two-fingered Anteater 
Quelch, Animal Life Brit. Guiana, 1901, p. 75 


Family Dasypodidae 


*Priodontes giganteus (Geofir.) 
Giant Armaaillo 
Quelch, Animal Life Brit. Guiana, 1901, p. 70 


*Dasypus novemcinctus Linné 
Lesser Armadillo; Nine-banded Armadillo; Yesi 
Quelch, Animal Life Brit. Guiana, 1901, p. 72 


Tatoua unicinetus (Linné) 
Small Armadillo; Tatouay 
Quelch, Animal Life Brit. Guiana, 1901, p. 71 


1919] Beebe: Mammalia, British Guiana 225 


Order PRIMATES ; Monkeys 


Family Callitrichidae 


*Cercopithecus midas (Linné) 
Midas Marmoset 
Hho. Mon.vof Prim. 1,-1913, p: 191 


Family Cebidae 


*Alouatta seniculus macconnelli Elliot 
Guiana Howling Monkey; Red Howler; Baboon 
Elliot,- Mon: -of- Prim: 1; 1918; p. 281 


Pithecia pithecia (Linné) 
White-headed Saki; Red-bellied Saki 
Elliot, Mon. of Prim. I, 1918, p. 294 


Pithecia satanas (Hoff.) 
Black Saki 
Mihot-sVon..of Pam. 1, 1913;-p> 29% 


Pithecia chiropotes (Humb.) 
Red-backed Saki; White-faced Beesa 
Elliot; Mon. of Prim: 1, 1918, p: 298 


*Saimiri sciureus (Linné) 
Squirrel Monkey; Sackiwinki 
Elhot: Mon. of Prim: I; 1913; p.. 310 


Aotus trivirgatus (Humb.) 
Night Monkey 
Elbo Mon of Prim: Al, 1913,- pz 16 


*Ateleus paniscus (Linné) 
Red-faced Spider Monkey; Quata 
Hiliot, iow: of Prim. Il, 1913,.p. 28 


*Cebus apella apella (Linné) 
Guiana Ring-tailed Capuchin 
Elliot, Mon. of Prim. II, 1913, p. 80 


Order ARTIODACTYLA ; Peccaries and Deer 


Family Tayassuidae 


Tayassu pecari pecari Fischer 
White-lipped Peccary; Bush-hog; Kairuni 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV. 1915, p. 377 


*Pecari tajacu (Linné) subsp.? 
Collared Peccary; Black Bush-hog; Abouyah 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV, 1915, p. 381 


226 Zoologica: N. Y. Zoological Society [Then 


Family Cervidae 


Odocoileus virginianus gymnotis (Wieg.) 
Orinoco Virginia Deer 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV, 1915, p. 174 


Odocoileus virginianus spinosus (Gay & Gerv.) 
Guiana Virginia Deer 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV, 1915, p. 175 


Mazama americana americana (Erxl.) 
Guiana Red Brocket 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV, 1915, p. 202 


*Mazama americana tumatumari Allen 
Tumatumari Red Brocket 
Allen, Bull. Amer. Mus. XXXIV, 1915, p. 536 
*Mazama simplicicornis simplicicornis (IIl.) 
Guiana Brown Brocket; Wellibicirie 
Lydekker, Cat. Ung. Mam. Brit. Mus. IV, 1915, p. 211 


Blastocerus dichotomus (IIl.) 
Marsh Deer; Guazu 
Quelch, Timehri, (2) We so2e pela 


Order PERISSODACTYLA ; Tapirs 
Family Tapiridae 
*Tapirus terrestris terrestris (Linné) 


Guiana Tapir; Bush Cow; Maipurie 
Lydekker, Cat. Ung. Mam. Brit. Mus. V, 1916, p. 42 


Order SIRENIA ; Sea-cows 
Family Trichechidae 


*Trichechus manatus Linné 
Manatee; Water Cow; Water Mama 
Quelch, Animal Life Brit. Guiana, 1901, p. 58 


Order ODONTOCETI; Dolphins and Whales 
Family Delphinidae 


*Delphinus delphis Linné 
Common Dolphin 
Quelch, Animal Life Brit. Guiana, 1901, p. 62 


LOL Beebe: Mammalia, British Guiana 


Family Platanistidae 


Inia geoffroyensis (Blainville) 
Guiana Fresh-water Dolphin 
Quelch, Animal Life Brit. Guiana, 1901, p. 62 


Family Balaenidae 


Eubalaena australis (Desm.) 
Southern. Right Whale 
Quelch, Animal Life Brit. Guiana, 1901, p. 62 


227 


utease- 
{SG ay 


i 
ke he 


Volume II, Number 8 


BIRDSsOr BARTICA DISTRIC# 
ADDITIONAL LIST 


At the time of publication of Volume I of ‘Tropical Wild Life’’* 
the list of birds observed by ourselves or collected by Whitely in 
Bartica District numbered three hundred and fifty-one forms. 

Since then a considerable addition to this list has accumulated 
from various sources, and these seventy-five new names I now 
present, bringing the total to the really remarkable number of four 
hundred and twenty-six different species of birds, all occurring 
within a radius of ten miles of our Research Station. These lists 
are, of course, to be considered as merely convenient check-lists, 
preliminary to the ultimate elaboration of the life histories of the 
various species. 

The number preceding the name corresponds to that in Brabourne 
and Chubb’s “‘List of the Birds of South America.” 


183 Claravis pretiosa pretiosa (Ferrari-Perez) 
Blue Ground Dove 


189a Leptotila verreauxi macconnelli Chubb 
Guiana Rusty Dove 

228 Aramides axillaris Laur. 
Venezuelan Woodrail 


238 Porzana carolina (Linné) 
Sora Rail 


255 Gallinula galeata galeata (Licht.) 
Florida Gallinule 


258 Ionornis martinicus (Linné) 
Purple Gallinule 


267 Heliornis fulica (Bodd.) 
Finfoot 


323 Sterna hirundo Linné 
Common Tern 


330 Sterna superciliaris Vieill. 
Least Tern 


*New York Zoological Society, New York, 1917. 
229 


230 Zoologica: N. Y. Zoological Society 


375 Pluvialis dominicus dominicus (P. L. S. Miill) 
American Golden Plover 


479 Tigrisoma lineatum (Bodd.) 
Lined Tiger Bittern 


549 Phalacrocorax vigua vigua (Vieill.) 
Guiana Cormorant 


585 Micrastur brachypterus (Temm.) 
Pied Hawk 


586 Micrastur mirandollei (Schl.) 
Mirandolle’s Hawk 


591 Geranospizias caerulescens (Vieill.) 
South American Blue Hawk 


597 Nisus superciliosus (Linné) 
Eye-browed Sparrow Hawk 


732 Ara ararauna (Linné) 
Blue and Yellow Macaw 


743 Orthopsittaca manilata (Bodd.) 
Red-bellied Macaw 


869 Deroptyus accipitrinus accipitrinus (Linné) 


Hawk-headed Parrot 


928 Chordeiles acutipennis acutipennis (Bodd.) 
South American Nighthawk 


946 Hydropsalis schomburgki Sclater 
Schomburgk’s Nighthawk 


960 Antiurus maculicaudatus (Lawr.) 
Spotted-tailed Nighthawk 


987 Reinarda squamata (Cassin) 
Fork-tailed Palm Swift 


988 Panyptila cayanensis (Gmelin) 
Cayenne Swift 


1001 Glaucis hirsuta hirsuta (Gmelin) 
Hairy Hermit 

1055 Campylopterous hyperythrus Cab. 
Rufous-breasted Sabre-wing 


1082 Agyrtrina fimbriata fimbriata (Gmelin) 
Lesson’s Emerald 

1125 Hylocharis cyanus viridiventris (Berlep.) 
Green-bellied Sapphire 


1199 Chrysolampis elatus (Linné) 
Ruby and Topaz Hummingbird 


Pits 


1919} Beebe: Birds, British Guiana 231 


1288 Vestipedes vestitus vestitus (Lesson) 
Glowing Puff-leg 


1449 Discosura longicauda (Gmelin) 
Racket-tail 

1544 Pteroglossus acarari atricollis (P. L.S. Miill.) 
Roraima Aracari 


1586 Galbula ruficauda ruficauda Cuvier 
Rufous-tailed Jacamar 


1631 Malacoptila fusca (Gmelin) 
White-breasted Puffbird 


1703 Melanerpes cruentatus (Bodd.) 
Little Black Woodpecker 


1749 Celeus undatus (Linné) 
Waved Woodpecker 


1784 Picumnus spilogaster Sunde. 
Sundevall’s Picilet 

1799 Picumnus buffoni undulatus Harg. 
Undulated Piculet 


1867 Taraba major major (Vieill.) 
Great Bush-shrike 


1900 Hypolophus canadensis canadensis (Linné) 
Black-crested Bush-shrike 


1992 Myrmotherula behni Berl. & Lever. 
Behn’s Antbird 


2096 Sclateria naevia naevia (Gmelin) 
Surinam Ant-creeper 

2493 Xenops rutilus heterurus (Cab. & Hein.) 
Red-tailed Recurved-bill 

2540 Xiphorhynchus guttatus sororius (Berl. & Hart.) 
Venezuelan Woodhewer 


2678 Mecocerculus leucophrys setaphagoides (Bonap.) 
Bonaparte’s Chat-tyrant 


2715a Colonia leuconota poecilonota (Cab.) 
Northern Long-tailed Tyrant 


2747 Platyrhynchus saturatus Sal. & God. 
Guiana Flatbill 


2767 Rhynchocyclus poliocephalus sclateri Hellm. 
Sclater’s Flatbill 


2829 Perissotriccus ecaudatus (d’Orb. & Lafr.) 
Short-tailed Pygmy Tyrant 


232 Zoologica: N. Y. Zoological Society 


2927 Tyrannulus elatus elatus (Lath.) 
Yellow-crowned Tyrantlet 


2934 Tyranniscus gracilipes Scl. & Sal. 
Slender-footed Flycatcher 


3080 Myiarchus tyrannulus tyrannulus (P. L. 8. Miill.) 
Rusty-tailed Flycatcher 


3095 Myiarchus nigriceps Scl. 
Black-headed Flycatcher 


3104 Tyrannus melancholicus satrapa (Cab. & Hein.) 
Lesser Yellow-breasted Kingbird 


3160 Chiroprion pareola pareola (Linné) 
Blue-backed Manakin 


3191 Scotothorus chrysocephalus (Pelz.) 
Golden-crowned Manakin 


3221 Pachyrhamphus rufus (Bodd.) 
Cinerous Thickbill 


3227 Pachyrhamphus niger cinereiventris (Scl.) 
Grey-bellied Thickbill 


3263 Attila thamnophiloides Spix 
Spix’s Attila 

3390 Thryophilus albipectus albipectus (Cab.) 
Schomburgk’s White-faced Wren 


3498 Donacobius atricapillus atricapillus (Linné) 
Black-capped Mocking-thrush 


3573 Pachysylvia pectoralis (Scl.) 
Cinereous-headed Woodbird 

3617 Compsothlypis pitiayumi pitiayumi (Vieill.) 
Olive-backed Warbler 

3641 Geothlypis aequinoctialis aequinoctialis (Gmelin) 
Equinoctial Warbler 

3645 Granatellus pelzelni pelzelni Scl. 
Pelzeln’s Red-throated Chat 


3648 Setophaga ruticilla (Linné) 
American Redstart 


3659 Myioborus castaneicapillus (Cab.) 
British Guiana Redstart 


3678 Basileuterus roraimae Sharpe 
Roraima Warbler 


3713 Cyanocompsa cyanoides rothschildii (Bart.) 
Rothschild’s Blue Grosbeak 


(Tiss 


1919} Beebe: Birds, British Guiana 233 


3760 Sporophila americana (Gmelin) 
Gmelin’s Seedeater 


4076 Dacnis bicolor (Vieill.) 
Blue-grey Honey-creeper 

4138 Tanagrella velia velia (Linné) 
Blue-bellied Tanager 


4281 Thraupis palmarum melanoptera (Scl.) 
Western Palm Tanager 


4361 Nemosia pileata pileata (Bodd.) 
Hooded Tanager 


4437 Schistochlamys atra atra (Gmelin) 
Black-faced Grey Tanager 


The various sources of these additions are as follows: 


A—Collected by William Beebe near Kalacoon in 1909 during the 
expedition described in the volume “Our Search for a Wilderness;’’ * 
Numbers 479, 549, 591, 732, 1586, 1631, 1703, and 1784. 


B—Collected at Bartica, in Chubb’s account of McConnell’s 
collection, ‘“The Birds of British Guiana,” Vol. I., together with a 
manuscript list of those to be published in Vol. II; Numbers 16, 188, 
189, 228, 238, 255, 258, 267,. 585, 586, 597, 748, 869, 946, 1001, 
1055, 1082, 1125, 1199, 1544, 1992, 2096, 2498, 2540, 2678, 2715, 
Q7TAT, 2767, 2829, 2927, 2934, 3080, 3095, 3104, 3160, 3191, 3221, 
3227, 3263, 3390, 3498, 3573, 3617, 3641, 3645, 3648, 3659, 4678, 
3713, 3760, 4076, 4138, 4281, 4861 and 4437. 


C—Recently identified specimens collected in Bartica District in 
1916; 928, 960, 987, 1449, 1749, 1867, 1900, 2715, 3191, 3617, 3645, 
3659, 3678, and 4861. 


D—Additions made during the short visit to the District in 1917; 
2a SoU ees O295,000,, 980, 1256;,1 (998 292K « 


Volume II, Number 9 


LIZARDS OF THE GENUS AMEIVA IN 
BARTICA DISTRICT 


NOTES ON THEIR COLOR AND PATTERN 
VARIATION 


Preparatory to intensive studies of the ontogenetic and phylo- 
genetic evolution of color and pattern among neotropical birds, 
many notes have been made of the remarkable variations found 
among lizards, especially in such abundant forms as Amewa. While 
reserving the publication of these until another season’s work will 
have made them more complete, I make mention here of the bearing 
which color and pattern variation have on the present classification 
of certain Ameiva. 

““A Revision of the Lizards of the Genus Ameiva’ is the title of 
a very excellent and thorough paper by Thomas Barbour and G. 
Kingsley Noble.* I wish especially to speak of the forms which the 
authors recognize as Ameiva ameiva ameiva, A. a. bilineata, A. a. 
melanocephala, and A. a. petersir. 

The following quotations have to do with these subspecies: 


KEY TO THE SPECIES 

i! Dorsal surface with heavy confluent seer, of black. j! Throat ee with a 
few black spots. . ig ro _ameiva amewa 
j? Throat smoky. k! SBrahehialea in Fhe rows a subequal cee 2 _atrigularis 

ie Brachials in one row of very large scales and three rows of emailer ones 
ameiva melanocephala 
i? Dorsal surface with a few black eg not confluent. j! A broad lateral band of 
brown on each side of the adult. aA ode #. 0... amet ONURE AIG, 
j? Lateral stripe indistinct or wanting. . ee seen (NITCTNOMDCLETS UD 


HABITAT 

Ameiva a. ameiva. Widely distributed over the northeastern part of South 
America from the Demerara River in British Guiana as far south as Bahia, Brazil, 
inland along the Amazon to as far west as the Madeira River. 

Ameiva a. bilineata. Apparently confined to the region between the Demerara 

and Orinoco Rivers. 

Ameiva a. melanocephala. Probably widely distributed throughout Venezuela. 

Ameiva a. petersii. Found along the upper Amazon from the Madeira River 
westward. 


*Bull. Mus. Comp. Zool. Harv. LIX. No. 6, 1915, pp. 417-479. 
Bos 


236 Zoologica: N. Y. Zoological Society paber 


As regards the two subspecies inhabiting British Guiana, the con- 
clusions are based on six specimens, one from Tumatumari and 
five from Dunoon. The detailed descriptions are from individuals 
measuring from one hundred and six to one hundred and sixty-two 
millimeters in length—individuals which I should hardly be inclined 
to call fully adult. The average of a half hundred specimens col- 
lected in one locality I have found to be almost twice this length, 
while extreme individuals reach over five hundred millimeters. 


By a minor geographic error the ranges of the two British Guiana 
forms are made to overlap, as Tumatumari, the locality for Ameiva, 
is a cataract on the Potaro, a western tributary of the Essequibo, 
far west of the Demerara and hence well within the indicated range 
of bilineata, ‘between the Demerara and Orinoco Rivers.’’ Dunoon 
is on the Demerara River. 


The main point I wish to make is that within an area of about 
five hundred yards around the Research Station of Kalacoon, and 
within a period of one week, I have collected several score of per- 
fectly typical specimens of ameiva, bilineatus, melanocephala and 
peters; and others which, judged by still more extreme variation of 
color and pattern, deserve still further subspecific differentiation. 
A certain proportion of this remarkable variation is due to age—to 
ontogenetic pigmental and pattern changes, but, on the other hand, 
all the forms are represented in my collection by fully adult lizards, 
that is, by individuals three hundred millimeters or more in length. 
This extremely local variation would seem to indicate either that all 
four forms were only intermediate variations, or, rather improbably, 
that Bartica District is a meeting place for a quartet or more of 
geographic subspecies. This collection of several score specimens 
from one locality opens up most interesting questions, and the cor- 
relation of observations carried on more widely, should quickly solve 
these rather superficial problems of diagnostic characters of color 
and pattern. 


The ontogenetic phase can be certainly demonstrated. Young 
specimens of Bartica ameiva averaging one hundred and fifty milli- 
meters in length are almost invariably of an extreme bilineatus type, 
exceeding the description of that form as given by Barbour, in con- 
centration of pigment as much as it in turn is said to differ from 
ameiva. The dorsal and ventral surfaces are immaculate, while a 
broad black band begins at the snout and extends back to the thigh, 


Lore | Beebe: Genus Ameiva, British Guiana 237 


narrowly bordered above and below with blue or white. In about 
ten per cent. of these small specimens the black body bands are 
broken up by vertical rows of faint dots. 


A second, larger stage, averaging about two hundred and fifty 
millimetres in length shows black spotting on the throat, and a more 
decided penetration of the black body bands by the rows of dots. 
But a new pattern in this phase is a double dorsal series of large 
black spots, which is found neither in larger nor in smaller forms 
of this lizard. In my notes I have distinguished this as brpwnctata. 


Passing through larger stages we find a typical lizard of four hun- 
dred millimetres in length with much of the upper surface covered 
with large confluent blotches, and the black lateral bands practically 
gone, the bluish-white vertical rows of dots of great size, and furnish- 
ing the dominant color and pattern note. It haschanged from a brown, 
banded lizard to a green, spotted one. It is nearer to the description 
of typical ameiva than anything else, but lacking such relatively 
immature characters as the white flank stripe, and the lateral, black, 
caudal stripe. 

Finally, we find a few big bluish-green giants over five hundred 
millimetres in length, dotted rather than blotched above, and with 
the lateral green spots large, isolated and framed in black—this 
framing being all that is left of the broad solid bands which form 
such a dominant feature in many smaller specimens. 

While much more material is needed and will be secured in the 
near future, yet even in this collection, a hint of still another problem 
is presented. In all the sizes and color patterns we find occasional 
individuals which appear melanistic—either in part or as a whole. 
Thus a specimen which will pass as extreme melanocephala, has the 
entire sides of the head, lower jaw, chin, throat and lower neck to 
between the forelegs, jet black. This anterior concentration of 
pigment seems to have been directly at the expense of the dark 
pigment in other parts of the body. The dorsal and lateral regions 
are quite green, with the isolated pale lateral dots lacking even 
their black frame, so drained are all the posterior parts of the animal 
of their black pigment. It has had a rush of melanism to the head, 
giving the superficial appearance of a very remarkable pigmented 
mutation. 

A second and more abundant melanistic form goes farther, and 
while above presenting a dull ameiva or petersi color and pattern, is 


238 Zoologica: N. Y. Zoological Society pubes) 


quite uniformly smoky black below. For purposes of record I have 
called this melanoventer. 


Studies of the color and pattern variations in living and recently 
killed specimens; recording of sexual characters in hundreds of indi- 
viduals, especially of mating pairs; scrutiny of the variations within 
a single brood of these lizards; uninterrupted observation of onto- 
genetic changes in individuals from the egg to the five hundred milli- 
metre stage; all these will surely contribute to the solution of such 
intensely interesting problems as the following: 


(a) Are ameiva, bilineata, petersii, bipunctata, melanocephala and 
melanoventer recognizable geographic subspecies or variations, which 
meet and live close together within a few yards radius. 


(b) Are they ontogenetic phases of one or more species? 


(c) Is it possible that in different localities they combine in part 
both a and 6; a paedogenetic acceleration or retardation such as we 
find in Axolotl? 


(d) Are these characters environmental or hereditary? 


(e) Are the similar variations which aden has so well demon- 
strated in Mexican Cnemidophorus to be considered as parallel or 
convergent when compared with those of Amewa, in addition 
perhaps to being a striking instance of orthogenesis? 


*A Contribution to the Study of Evolution Based Upon the Mexican Species of 
Cnemidophorus, Proc. Zool. Soc., London, 1906, I, p. 277. 


CRS ie 
‘ 


“LOOLOGICA |. 
SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


oA 
SOR Sen 
mR 
asa gee : 


MAY 4 7099 
Letlone: Muse’ 


VOLUME II, NUMBER 10 


REMARKABLE HABITS OF 
THE SAGE GROUSE 


_ AS OBSERVED IN SOUTHEASTERN OREGON 
IN MAY, 1918 


By R. BRUCE HORSFALL 
With Illustrations Drawn from Life by the Author 


- 


PUBLISHED BY: THE -SOCIETY 
THE ZOOLOGICAL PARK, NEW YORK 


APRIL, 1920 


oo” 


New Vork Zoological Society 
‘ General Office: 111 Broadway, New York City 


Officers — : i‘ 

President, HENRY FAIRFIELD OSBORN ; ; one” 

Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; | ee 

Secretary, Chairman, Exec. Committee, MADISON GRANT; 
Treasurer, PERCY R. PYNE. 


Board of Managers 
5 Class of 1921 
LEVI P. MORTON, MADISON GRANT, WILLIAM WHITE NILES, ~ 
HENRY A. C. TAYLOR, FRANK K. STURGIS, GEORGE J. 
GOULD, OGDEN MILLS, LEWIS RUTHERFORD MORRIS, - 
- ARCHER M. HUNTINGTON, E. C. CONVERSE, 
GEORGE D. PRATT, T. COLEMAN DUPONT. 


Class of 1922 
PERCY R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE, 
C. LEDYARD BLAIR, EMERSON McMILLIN, ANTHONY R. 
KUSER, WATSON B. DICKERMAN, Mortimer L. 
SCHIFF, FREDERIC C. WALCOTT, BEEKMAN 
WINTHROP, GEORGE C. CLARK, 
W. REDMOND CROSS. ‘ 


Class of 1923 
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. 
DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON, 
ROBERT S. BREWSTER, EDWARD S. HARKNESS, 
WILLIAM B. OSGooD FIELD, A. BARTON 
HEPBURN, WILLIAM WOODWARD, 
EDWEN THORNE, PERCY A. 
ROCKEFELLER. 


Scientific Staff 
WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 
MEDEA NM BEEBE, Honorary Curator of Birds and Director of the 
Tropical Research Station; 
LEE 8. CRANDALL, Curator of Birds; 
GEORGE S. HUNTINGTON, Prosector; 
GEORGE A. MACCALLUM, Pathologist; 
W. REID BLAIR, Veterinarian; 
ELWIN R. SANBORN, Photographer and Editor. 


€ditorial Committee 
HENRY FAIRFIELD OSBORN, Chairman; MADISON GRANT, 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


SAGE GROUSE DISPLAYING AND STRUTTING AT SUNRISE 
Painted from Life by R. Bruce Horsfall 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 10 


REMARKABLE HABITS OF 
THE SAGE GROUSE 


AS OBSERVED IN SOUTHEASTERN OREGON 
IN MAY, 1918 


By R. BRUCE HORSFALL 
With Illustrations Drawn from Life by the Author 


Pei tS 0 D B:¥ THE sO GLE TY 
tate 2OOoLOGICAL PARK, NEW YORK 


APRIL, 1920 


FIGURES 


Fig. 69—Sage Grouse Displaying................ Frontispiece 
Figs. 70-75—Studies of Sage Grouse Dance.......... 246-248 
Filling, pouch “with: aifc 00. « 2 ooo oe ee oe 246 


Stiff-legged run after filling the pouch. 


Liftine--pouch with the. wines: «5 sn .1.cc eee ee 247 
Side view of lift of the pouch. 


Extreme of-throw-or. the pouch .......< acces a eee 248 
Slap down of pouch on the chest. 


Fig. 76—Sage Grouse (male) Neck Feathers............ 249 


Volume II, Number 10 


REMARKABLE HABITS OF 
THE SAGE GROUSE 


AS OBSERVED IN SOUTHEASTERN OREGON 
IN MAY, 1913s 


By R. BRUCE HORSFALL 
With Illustrations Drawn from Life by the Author 


Stealthily and carefully we picked our way along the mud- 
flat road to the high gate in the lava-rock ranch fence, and peered 
through the bars. 


“There they are!” burst from our lips in an excited whisper, 
as we caught the glint of white spots a few hundred yards beyond. 


Cautiously we clambered over the irregular loose rocks and, 
like Indians on the warpath, crawled, squirmed and wriggled our 
way to a low outcropping of volcanic rock. From this vantage 
point we had an unobstructed view of the broad and bare flood- 
plain before us. 


Cramped and strained in every muscle and bone, we remained 
hidden among those rocks till supper time, loath to leave the 
wonderful sight. 


In the open at intervals of from twenty-five to fifty feet were 
sixty magnificent sage cocks strutting around with puffed-out 
chests and tails spread like miniature turkey gobblers, making 
noises for all the world like the popping of corks on the pier at 
Atlantic City, a sound which took me back in memory to the 
hotel in Colombo, Ceylon, where a hundred guests were served 
soda water at every meal, and the bottles were always opened 
at the tables. 


Evidently these rocks had been a favorite place for Indians in 
years gone by, for all about us in the sand were obsidian chips 
and charcoal, with bits of arrow.and spear heads. We after- 
wards picked up many perfect specimens on the open flat. 


243 


244 Zoologica: N. Y. Zoological Park [II; 10 


Here the Klamath Indian had lurked at evening to secure, 
with his twanging bow and bits of flying glass, a sage cock for 
the morrow’s dinner, making arrow heads in the middle of the 
day when no birds were about. To the Indian a strutting sage 
cock was but an easy mark and a quick lunch. His stolid nature 
probably did not marvel at the wondrous performance, and no 
question entered his mind as to how and why. 


To us, however, it was a sight which satisfied a great hunger ; 
not the hunger of the body, but of the mind. We had spent days 
and nights in travel to see that phase of nature, to gain that 
‘scrap of knowledge; and we feasted to a great content, though 
many questions remain unsatisfied as to the how and the why. 


The opportunity for these observations had come after two 
years’ delay, and we were accordingly appreciative to the utmost. 


In the summer of 1915, we had been viewing the Klamath 
Lake Pelican Colonies with the game warden, Mr. J. J. Turber, 
and had gone on to Laird’s Landing, at the foot of the lake, to 
remain over night. While we were there Mr. Laird told us of 
sage grouse coming down on the flat at the eastern end of the 
pasture every spring to do their courting, but at that date, May 
31st, they had stopped for the season. 


It was now May, 1917, and we were to have our opportunity 
to observe and picture the birds in action. 

Mr. Turber had brought William L. Furley, Stanley G. Jewett 
and the writer from Klamath Falls to Mr. Laird’s place. It was 
afternoon when we rounded the lower end of the lake, and a few 
grouse were on the open alkali flat. We cached our camera and 
blind, and went on to the house to arrange for beds and board, 
leaving as soon as possible for our first close view of the birds. 
After supper that evening we set up our blind near the grassy 
slope reaching from the sage-covered hill at the eastern side of 
“the wash.” 


The waters that had formed this flood plain came down from 
the forest-capped Van Brimmer mountain away off to the south. 
Westward of this wash lay hillocks and ridges of dark lava rock. 
About eight level acres, near the shore of the lake, were bare of 
vegetation, and it was there that the sage cocks came from miles 
around to dance and strut—and “plop.” 


1920] Horsfall: Habits of Sage Grouse 245 


The strut was made up of four movements. First, the filling 
of the air pouch, accompanied by a grunting sound; second, a 
short stiff-legged run in which neither pouch nor wings touched 
the ground; third, the bird stopped suddenly, spread his tail as 
it raised to the perpendicular, threw back his head and with a 
forward movement of the wings pushed the air-filled pouch well 
up on the chest; fourth, there was a sudden upward throw, fol- 
lowed by a more vigorous and snappy toss, and the tightened 
pouch came down again on the extended chest with a rubbery 
“plop.” This plop was repeated three times, then the bird eased 
down for another rumbling gurgle and another run. 

Mr. Laird assured us that these antics take place from early 
March till the first of June; in fact, through the mating season. 
However, it was evident that this was not a courting action; be- 
cause when in the course of the morning or evening performance, 
two or three hens meandered through the throng no notice what- 
ever was taken of them. A real courting performance of a 
different character may take place in the daytime, far from the 
watering place, on the sage covered hills. 


Each bird appears to have a private spot on which no other 
dares to trespass. In reaching those proprietary spots, collisions 
sometimes occurred, and quiet cock-fights took place much after 
the manner of China pheasants. With rump feathers erect, 
lowered heads and tails, and dragging pouches, the birds sidled 
around and struck with their wings; all the while scolding in 
a trumpeting, gurgling grunt, as the owner actually pushed the 
intruder off his domain. A few feet one way or another, the 
belligerents would separate and go on plopping as before. 


Some birds began to perform well up in the sage brush, and 
plopped all the way out to the dancing spot; others walked quietly 
into their respective claims before beginning to show off. 


An examination of the pouch of the sage grouse discloses a 
peculiar development. In front are two yellowish-green bare 
spots separated and surrounded by short stiff feathers, shortest 
and stiffest immediately surrounding the bare area. Probably 
it is these spots which make the sounding plop, after the man- 
ner of a wet drum-head. In the fall, when new, these feathers 
are soft and exceedingly friable, and by the time the birds are 
ready for strutting, have broken away to stiff, sharp bristles 


246 Zoologica: N. Y. Zoological Park 


Fic. 70. FILLING POUCH WITH AIR 


Fic. 71. STIFF-LEGGED RUN AFTER FILLING THE POUCH 


STUDIES FROM LIFE OF SAGE GROUSE DANCE 
Pen Drawings by R. Bruce Horsfall 


[irs 10 


1920] 


Horsfall: Habits of Sage Grouse 


ma 


\ | | bs is EY 


S44 


4 


Fic. 72. LIFTING POUCH WITH THE WINGS 


Fic. 73. SIDE VIEW OF LIFT OF THE POUCH 


STUDIES FROM LIFE OF SAGE GROUSE DANCE 


Pen Drawings by R. Bruce Horsfall 


247 


248 Zoologica: N. Y. Zoological Park [II; 10 


Fic. 74. EXTREME OF THROW OF THE POUCH 


Fic. 75. SLAP DOWN OF POUCH ON CHEST SHOWING DISTENDED 
BARE SPOTS 


STUDIES FROM LIFE OF SAGE GROUSE DANCE 
Pen Drawings by R. Bruce Horsfall 


1920] Horsfall: Habits of Sage Grouse 249 


Fic. 76. SAGE GROUSE (MALE) NECK FEATHERS 


September feather of loose structure which accounts for the wear on 
the spring feather. Upper—Taken from near bare spots on the 
pouch. Lower—Feather from breast beneath pouch 


Pen Drawings by R. Bruce Horsfall 


250 Zoologica: N. Y. Zoological Park eG) 


which will in no way interfere with the vigor of the snap, as 
soft feathers would do. 


The morning after our arrival, the 11th of May, no birds came 
near enough to the blind for photographing. At nine o’clock it 
began to rain, and we spent the remainder of the morning search- 
ing for nests on the adjacent hillside. We found two from which 
the young had hatched, and one that had been rifled of its con- 
tents, probably by a coyote. In the afternoon the birds came in 
too late for pictures. 


On May 11, no birds came near the blind, so we moved it to 
another spot. The afternoon was windy and stormy, and only 
a few birds came in at four o’clock. 


On the 13th of May, we arose at three A. M. and after a 
hurried breakfast, stumbled and wabbled along the ruts of the 
road. No wind was stirring; which was a very unusual thing 
for this high plateau region. A heavy cloud hung over us, as 
only clouds in an arid region can hang, black as night, but the 
faint yellowish light of dawn was slowly brightening as we crept 
into the blind, at four o’clock. Dark as it was, a few birds were 
already there, and by five-fifteen we were able to make the first 
exposure. As usual, there were about sixty birds in the field. 
Actual counts of birds within our range of vision at various times 
were 51, 52, 51 and 54. By seven o’clock all had left for the 
sage-brush hills; but we had had several birds within thirty feet 
of us most of the time. 

We returned to the blind at 4:15 A. M. A few birds were on 
the shore when we arrived and by seven o’clock, the usual time 
for the birds to scatter, we had secured moving and still pictures 
of every action from birds within twenty-five feet of us. 


As we left, a lone coyote yapped to us a long farewell. 


Publications 
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ZOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 11 


ECLIPSE PLUMAGE IN DOMESTIC FOWL 


a 


AMS 


BY 


LEE §S. CRANDALL 
CURATOR OF BIRDS 


Pte. 1 oat Do BY. -T AR rokind BA Sap) Us (fib Rae 
tote ACOLOGICAL PARK, NEW YORK 


OcToBER 15, 1920 


"s 


New York Zonlogical Soriety 


General Office, 111 Broadway, New York City 


President: 
: HENRY FAIRFIELD OSBORN. 
First Vice-President: Second Vice-President: 
MADISON GRANT. FRANK K. STURGIS. 


Treasurer: PERCY R. PYNE, 20 Exchange Place. 
Assistant Treasurer: THE FARMERS’ LOAN & TRUST COMPANY. 
Secretary: MADISON GRANT, 111 Broadway. 


Executive Commitier 
MADISON GRANT, Chairman. 


Percy R. PYNE, LISPENARD STEWART, 
WILLIAM WHITE NILES, WATSON B. DICKERMAN, 
WM. PIERSON HAMILTON,’ ANTHONY R. KUSER, 
FRANK K. STURGIS, HENRY FAIRFIELD OSBORN, 
Ex-Officio. 
Geurral Officers 


WILLIAM T. HorNADAY, Director, Zoological Park. 
CHARLES H. TOWNSEND, Director, New York Aquarium. 
GEORGE S. HUNTINGTON, Prosector. 

GEORGE A. MACCALLUM, Pathologist. 

C. GRANT LA FARGE, Architect. 

H. DE B. PARSONS, Consulting Engineer. 

R. L. CERERO, Bursar. . 


Annditing Commitiee 
WILLIAM WHITE NILES, Chairman. 
LISPENARD STEWART, ANTHONY R. KUSER. 
Pension Board 
WILLIAM PIERSON HAMILTON, Chairman. 
Percy R. PYNE, Treasurer. 


Dr. Lewis R. MorkrIs, CHARLES H. TOWNSEND, 
GEORGE BIRD GRINNELL, H. R. MITCHELL, 
WILLIAM WOODWARD, R. L. DITMARS. 


HERMANN W. MERKEL, Secretary. 


Editorial Committee 
HENRY FAIRFIELD OSBORN, Chairman. 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


ee te 


FIG. 77. GAME COCK, SHOWING HACKLE IN ECLIPSE 


FIG. 78. Left: RED JUNGLE FOWL (Gallus gallus) SHOWING HACKLE IN ECLIPSE 
Right: RED JUNGLE FOWL (Gallus gallus) SHOWING FULL HACKLE 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 11 


KCLIPSE PLUMAGE IN DOMESTIC FOWL 


BY 


LEE S. CRANDALL 
CURATOR OF BIRDS 


Po iteb i. 1.8) Eb. D By ¥% as (ae 3 ee fee eT 
PEE 7.00 hOGICAL PARK, NEW YORK 


OcTOBER 15, 1920 


ECLIPSE PLUMAGE IN DOMESTIC FOWL 


By LEE 8. CRANDALL 
Curator of Birds. 


From the time of Darwin, ornithologists almost unanimous- 

ly have agreed that the Red Jungle Fowl (Gallus gallus) of 
India, the Malay Peninsula, Indo-China and the neighboring is- 
jands, is the sole ancestor of domestic fowls. Practical poultry- 
men, on the other hand, are equally sure that certain breeds, 
especially of the groups known as Asiatics and Oriental games, 
must have come from some unknown, gigantic ancestor. There 
are some, also, who feel that the case of the Red Jungle Fowl’s 
ancestral relation to breeds other than these Eastern types, is 
not fully established. While it is not the purpose of the present 
paper to enter fully into this discussion, it is felt that the facts 
herein presented have a strong bearing in this seemingly inter- 
minable controversy. 
On October 22, 1916, the New York Zoological Park received, 
through the medium of Ansel W. Robinson, of San Francisco, 
a pair of Red Jungle Fowl. They were young birds but the male 
was in full color, with flowing hackle and saddle feathers. The 
pair had been obtained in the Philippine Islands, where the 
species is believed to have been introduced by man. Both speci- 
mens were typical, which is more than can be said for most of 
the Red Jungle Fowl! seen in captivitv. They were very wild 
when first received but gradually became tamer, though they 
never would allow themselves to be handled or even touched. 

The birds were kept in warm quarters during the winter 
months, and by spring were in very good feather and condition. 
In June, the cock began to shed his long neck hackles and dropped 
the sickle feathers of the tail at the same time. 

Close watch was kept on the bird during this period and it 
was found that he was going through the oft-mentioned but sel- 
dom described eclipse which is typical of the species. As the red, 
black-centered, pointed hackle feathers dropped out, they were 
replaced by short, round-tipped black ones. Those in the upper 
portions of the neck were broadly margined with orange, but as 
they approached the neck, they became wholly black. 

Late in July, the cock caught a sudden cold, and died on 
August 2, 1917. The eclipse was not quite complete, the small 


255 


256 Zoologica: N. Y. Zoological Society [i Tee ate 


feathers on the head, just behind the comb, not having been 
molted. It is quite possible that this is as far as the normal 
eclipse goes. The new tail sickles had not yet begun to grow. 
This individual seems to correspond in general with Ogilvie- 
Grant’s* designation of June-September as the eclipse period. 


This eclipse plumage of the male Red Jungle Fowl, while 
not generally well known, has always proved a stumbling block 
to those who maintain that the species is the sole ancestor of our 
domestic fowl. It has been difficult to explain the absence of the 
eclipse in domestic birds, since it is so characteristic of their 
wild progenitors. 

There seems to be no record of eclipse plumage in domestic 
cocks, though in the light of the present evidence, this seems due 
rather to oversight than to lack of occurrence. 


In the autumn of 1919, Mr. Prescott Van Wyck, of Sum- 
mit, New Jersey, brought to the Zoological Park, a Black- 
breasted Red Pit Game cock, for the writer’s inspection. The 
bird was a yearling, in his first adult molt. Examination showed 
that the sickle feathers of the tail had been cast and that the 
neck was in eclipse plumage, exactly as in the Red Jungle Fowl. 
This is all the more interesting, as, in color, the Black-breasted 
Red game fowl is an almost perfect counterpart of the Red 
Jungle Fowl. The eclipse feathers of the game cock were orange, 
with broad black margins, exactly the opposite of the colors in 
the Jungle Fowl eclipse. To complete the comparison, the short, 
red, pointed feathers of the head, as in the Jungle Fowl, had not 
been replaced. 


Unfortunately, this cock was not the property of Mr. Van 
Wyck, and could not be left for observation. However, a photo- 
graph was made, showing the hackle in eclipse. In the fore- 
ground of the photograph, which is presented herewith, a single 
unmolted hackle feather may be seen. 


A few months later, the change to full plumage had been 
accomplished, and specimens of the normal hackle feathers were 
obtained, which may be compared with those of the eclipse. 


It seems reasonable to suppose that this example of the 
eclipse plumage in domestic fowls, is not unique but that its 
occurrence has been overlooked. Careful examination of molt-.- 
ing cocks, particularly game fowl, no doubt would reveal total or 
partial eclipse plumage as at least fairly common. 


Pee oe ise Sia le W. R. A Hand-book of the Game Birds, 1897, Vol. II, 
Pp. 40-9. 


FIG, 79, Above: FEATHERS FROM ECLIPSE HACKLE OF GAME COCK 
Below: NORMAL HACKLE FEATHERS FROM SAME BIRD 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE - 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 12 


(Papers from the New York Aquarium) 
(Contribution Number 5) 


A CONTRIBUTION TO THE LIFE HISTORY OF 
THE PUFFER, SPHEROIDES MACULATUS 


(SCHNEIDER) 
oneal 

By W. W. WELSH _vsvinta | 

United States Bureau of Fisheries Si 
AND 
C. M. BREDER, JR. (% MAR. 14 1929 
New York Aquarium ine 
! N, - 
Hon 9 | Mi x 


PA ea See De BYY';- THE SOCIETY 
THE ZOOLOGICAL PARK, NEW YORK 


JANUARY, 1922 


New York Zoological Society 


if General Office: 111 Broadway, New York City 


Officers ; 

President, HENRY FAIRFIELD OSBORN ; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 

Treasurer, Corne ios R. Acnew 


Board of Managers 


Glass of 1923 
HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. 
DIETERICH, GEORGE F. BAKER, ‘Wo. PIERSON HAMILTON, 
ROBERT S. BREWSTER, EpWwArp S. HARKNESS, 
WILLIAM B. Oscoop FIELD, A. BARTON 
HEPBURN, WILLIAM WoOopWaARD, 
EDWIN THORNE, PERCY A. 
ROCKEFELLER. 
Glass nf 1924 
MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, 
GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFURD 
Morris, ARCHER M. HUNTINGTON, GEORGE D. 
PRATT, T. COLEMAN DUPONT, HENRY D. WHITON, 
EDWARD HATCH, JR., CORNELIUS R. AGNEW 
Glass of 1925 
Percy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DopGE, 
C. LEDYARD BLAIR, EMERSON McMILLIN, ANTHONY R. 
KUSER, WATSON B. DICKERMAN, MortTIMER L. 
SCHIFF, FREDERIC C. WALCOTT, BEEKMAN 
WINTHROP, GEORGE C. CLARK, 
W. REDMOND CROSS. 


Srivutifc Staff a 
WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 
WILLIAM BEEBE, Honorary Curator of Birds and Director of the 
Tropical Research Station; 
LEE S. CRANDALL, Curator of Birds: 
GEORGE S. HUNTINGTON, Prosector; 
GEORGE A. MACCALLUM, Pathologist; 
W. Rep Buair, Veterinarian; 
ELWIN R. SANBORN, Photographer and Editor. 


Enttorial Committer 
HENRY FAIRFIELD OSBORN, Chairman; 
WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


Corrected to January, 1922. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 12 


(Papers from the New York Aquarium) 
(Contribution Number 5) 


A CONTRIBUTION TO THE LIFE HISTORY OF 
THE PUFFER, SPHEROIDES MACULATUS 
(SCHNEIDER) 


By W. W. WELSH 


United States Bureau of Fisheries 
AND 


C. M. BREDER, JR. 


New York Aquarium 


Poesia D BAY.) fa Sr O°cG.L By TY 
Piro OhOnG ke AL “PARK. NEW - YORK 


JANUARY, 1922 


“WU ¢G"Z ‘YI SuUe] penjoyw 
ONIHOLVH YALAV SAVG AAIA VAUNVT 08 ‘DI 


Volume II, Number 12 


A CONTRIBUTION TO THE LIFE HISTORY OF 
THE PUFFER, SPHEROIDES MACULATUS 
(SCHNEIDER) ' 


By W. W. WELSH 
United States Bureau of Fisheries 


C. M. BREDER, JR. 
New York Aquarium 


INTRODUCTION. 


The present paper embodies the results of studies on 
Spheroides maculatus (Schneider), carried out at Atlantic City, 
New Jersey, chiefly during the month of August, 1920, where 
a temporary field laboratory was established on Young’s Million 
Dollar Pier through the courtesy of Captain E. L. Young. All 
material for study was obtained from the two pound-nets oper- 
ated upon this pier. 


The temperature of the water during this season was ab- 
normally low, being comparable to that normally encountered 
below the five fathom line in this region. Although conditions 
in general were rather unsatisfactory for this type of work, no 
difficulty was encountered in securing adequate material for the 
study of this species. 


The drawings of the eggs and larvae were made from liv- 
ing material by means of the camera lucida. In most cases a 
small quantity of chlorotone was added to the sea water con- 
taining the larvae, to quiet them. All illustrations, excepting 
Fig. 87 have been executed by the junior author. 


*Published by permission of the United States Commissioner of 
Fisheries. 


262 Zoologica: N. Y. Zoological Society | (Ths 02 


ONTOGENY. 


Spawning.—Ripe females of this species were taken from 
July 30 to August 27, which were the first and last dates on 
which specimens were examined. There seemed to be no in- 
crease or decrease in the proportionate number of ripe fish, 
which fact suggests that the peak of their sexual activity ex- 
tends well over the period in this region. Hard and spent in- 
dividuals were constantly taken with the ripe ones. The ova 
passed freely on the application of very slight pressure, issuing 
forth in a stream about 3 mm. in diameter. Ripe males were 
frequently taken, but equal success in fertilization was obtained 
by macerating the testes of fish from which milt would not 
flow. The sexes were present in approximately equal numbers. 


Eggs.—The eggs are transparent, spherical, and invested 
with a smooth adhesive covering which is irregular in outline. 
They are demersal and readily become attached to any sub- 
merged object, or caked in a mass, owing to their adhesive na- 
ture. Where numbers adhere to a side of the container, close 
together and in a single layer, the adhesive envelope assumes 
a somewhat hexagonal appearance. The surfaces of the eggs 
are finely reticulated, rather resembling crepe paper. The eggs 
average about 0.874 mm. in diameter, varying from .85 to .91 
mm., while the enveloping adhesive coat increases the diameter 
to an average of about .954 mm. A large number of colorless 
oil globules of low refractive index are present in a foamy 
cluster, which averages about .34 mm. in diameter, and a very 
faint yellowish olive tinge can be detected in the area in which 
the blastoderm is to develop. Fig. 81, A represents the unfertil- 
ized egg. 


Embryology.—The eggs, of which the development was 
studied, were incubated in small bowls, with a daily change of 
water, at a temperature which averaged about 67°F. At fifteen 
minutes after fertilization no great change was apparent, but 
within the following two hours the first cleavage had been com- 
pleted (Fig. 81,B), and by sixteen hours after fertilization the 
blastodise had commenced to indicate the approaching differen- 
tiation (Fig. 81,C). By twenty-four hours the embryo was quite 


FIG. 81. MAGNIFICATION 36 X. 


A—Unfertilized egg. B—Egg with blastoderm of two cells. Two and 
one-half hours after fertilization. C—Egg showing early stage in the 
differentiation of the embryo. Sixteen and one-half hours after fertili- 
zation. D—Egg showing a moderately advanced stage of differentiation 
of the embryo. Forty hours after fertilization. E—Egg with moderately 
advanced embryo. Seventy hours after fertilization. F—Egg with ad- 
vanced embryo. Ninety hours after fertilization. 


264 Zoologica: N. Y. Zoological Society IPs i 


distinct and in the succeeding twenty it had reached more than 
half way around the yolk (Fig. 81,D) At seventy hours the tail 
was free at its tip, vertebral somites were visible, and con- 
vulsive squirmings had commenced. Scattered black chromato- 
phores had appeared along each side and the eyes were quite 
distinct. The oil globules were chiefly located in the dorsal half 
of the yolk (Fig. 81, E). At ninety hours, in addition to the black 
chromatophores, red and orange ones were scattered along the 
sides. The anterior ventral surface of the yolk was well covered 
with large black chromatophores, rather dendritic, and there was 
a suggestion of the oil globules consolidating into a somewhat 
lesser number of larger spheres. A few small black chromato- 
phores and punctulations were located in the posterior part of 
the iris and on the tip of the snout (Fig. 81, F). At this stage the 
tip of the tail overlapped the head and the embryo exhibited 
much activity. The eggs began hatching at about one-hundred 
and twelve hours after fertilization. At this time they were 
steadily increasing in pigment content. The pigmentation ter- 
minated posteriorly in abrupt fashion about midway between 
the vent and the tip of the tail, and at this point a brilliant 
opaque chrome yellow spot was apparent on the dorsal surface. 
Numbers hatched however before the chrome yellow spot ap- 
peared (Fig. 82). Most of the larvae emerged from the egg tail 
first, although in a few cases the head came first. Both the 
eggs and larvae were decidedly variable during the entire 
period they were under observation, especially in the matter of 
pigmentation. 


The frontispiece (Fig. 80) represents the typical coloration 
of the fry at the age of five days. Various shadings have been 
used in the text figures in an attempt to represent the very strik- 
ing coloration exhibited by this species. Red has been represented 
by large closely placed dots, yellow by small ones, and the vari- 
ous green, orange, and purple markings by short lines, the lo- 
cation of each being explained in the text. By comparing the 
line drawings with the colored plate, a fair idea of the pig- 
mentation at each stage may be gained, since, while the amounts 
of pigment present in the various stages differ, the actual colors 
shown by the chromatophores differ little from those repre- 
sented in the plate. 


1921] Welsh & Breder: Life History of the Puffer 265 


Larval development.—The newly hatched larvae averaged 
about 2.41 mm. in length. The yolk sac was small and still 
contained oil globules. The head was somewhat deflected and 
the eyes, the pupils of which were not yet dark, were directed 
a trifle forward and downward. The pectoral fins were difficult 
to see, owing to the heavy pigment lying beneath them. The 
coloration was brilliant; red, orange, yellow and black chroma- 
tophores being thickly distributed over the body. Deep purplish 
black chromatophores invested the anterior end of the yolk sac 
and some were scattered through the iris, which also contained 
a cluster of heavy black ones in the dorsal and posterior quad- 
rant. Numerous minute tubercles were present over practically 
the entire body. The newly hatched larvae were very active 
and possessed considerable vitality. Fig. 88 shows a detailed 
profile of the head at this age. 


As development advanced the red pigment became rela- 
tively reduced, the orange and yellow coming more into promi- 
nence. About twenty-four hours after hatching the pupil be- 
came black, the nostrils were plainly visible, the pectorals more 
distinct, and what seemed to be beginnings of lateral line or- 
gans had appeared. Seven pairs of these organs were observed 
in a two day old specimen, and probably more were present but 
not distinguishable from the surrounding tissue. Three pairs 
have their origin over the eyes, a small and a large pair are 
posterior to the pectorals and another small pair over the vent, 
as well as a pair somewhat forward of that point (Fig. 83). On 
treating one specimen with chlorotone a secretion appeared to 
exude from these bodies, which indicated their glandular na- 
ture. Fig. 89 shows in detail the vent and location of the two 
posterior glands, and Fig. 90 represents the appearance, in 
profile, of the large one just behind the left pectoral in a speci- 
men seven days old. 


At two days the vent and mouth were open, and in addition 
to the other pigments, various green markings had appeared, 
especially in the iris. The yolk was materially reduced and at 
three days the mouth was functioning. Viewed by reflected 
light the colors were brilliant and metallic. The fry were active 
and decidedly heliotropic. The chrome yellow caudal spot, pre- 


FIG. 82. NEWLY HATCHED FISH. 
Actual length, 2.41 mm. 


FIG. 88. LARVAL FISH ONE DAY AFTER HATCHING. 
Actual length, 2.50 mm. 


FIG. 84. LARVAL FISH FIVE DAYS AFTER HATCHING. 


Actual length, 2.55 mm. 


FIG. 8. LARVAL FISH SEVEN DAYS AFTER HATCHIN 
Actual length, 2.62 mm. 


FIG. 86. POST LARVAL FISH. 
Actual length, 7.35 mm. 


FIG. 87. ADULT FISH, PARTLY INFLATED. 
Actual length, 200 mm. 


268 Zoologica: N. Y. Zoological Society TIT se 


FIG. 88. DETAIL OF HEAD OF NEWLY HATCHED LARVA. 
Actual length of entire fish nearly 2.40 mm. 


viously noted, and a few black ones on the head constituted the 
only dorsal pigments. 


On the fifth day numerous tubercles of considerable size 
were observed on the ventral surface. The eyes were coming 
into a more nearly lateral position and a small spine was de- 
veloping on the operculum. The black chromatophores on the 
abdominal region (the yolk being practically absorbed and the 
oil globules gone) were decidedly dendritic. The pectoral was 
assuming a less spatulate shape, a rather pronounced point ap- 
pearing near the upper margin, and the maxillary was well 
formed and prominent (Fig. 84). The tip of the tail, shown by 
Fig. 91, indicates the tuberculated appearance of the body in a 
six day old specimen. At this age the otoliths were becoming 


1921] Welsh & Breder: Life History of the Puffer 269 


FIG. 89. DETAIL OF VENT OF TWO DAY OLD LARVA. 


Actual length of entire fish nearly 2.52 mm. 


‘more complicated in conformation and the body was becoming 
opaque. At seven days, (Fig. 85), little further change in ap- 
pearance could be observed. The tubercles, especially on the 
ventral surface steadily increased in size, and the maceration of 
some from that location disclosed small barbed hooks as shown 
in Fig. 92. At ten days the larvae were all dead or dying, the 
largest having reached a length of 2.65 mm. 


It is believed that these eggs and larvae would prove excep- 
tionally satisfactory for detailed studies of embryology and larval 
development; the eggs, because of extreme hardihood, trans- 
parency, and possession of an adhesive coat maintaining them 
in a definite position; and the larvae because of the large and 
varied chromatophores that develop so strikingly, and their 
great tenacity to life. 


Fig. 86 was drawn from a specimen 7.35 mm. long taken by 
the U. S. S. Grampus July 31, 1918, station No. 10081, over a 
depth of eleven fathoms off the New Jersey coast near Barne- 
gat, the net haul being from ten fathoms to the surface. At this 
stage most of the diagnostic characters of the adult have been 


FIG. 90. PROFILE OF LATERAL ORGAN, POSTERIOR OF LEFT 
PECTORAL FIN, FROM A SEVEN DAY OLD LARVA. 


Magnification nearly 3860 X. 


FIG. 91. TIP OF TAIL OF SIX DAY OLD LARVA 
SHOWING TUBERCLES. 
Magnification nearly 104 X. 


FIG. 92. SPINES FROM VENTRAL SURFACE 
OF TEN DAY OLD LARVA. 


Magnification nearly 360 X. 


FIG. 98. OUTLINE OF A SPECIMEN 22.5 MM. LONG, SHOWING PROPORTIONS 
ASSUMED ON INFLATION BY THE POST LARVAL FISH. 


From United States National Museum collection. Aug. 11, 1892, Quissett Harbor, Mass. 


FIG. 94. DORSAL VIEW OF SPECIMEN SHOWN IN FIGURE 93. 


De Zoologica: N. Y. Zoological Society | [TE 12 


acquired, although the skin surrounding the body is more dis- 
tensible than in the mature fish. The transparent membranes 
covering the eyes move away from them on inflation, adding 
to the grotesqueness of appearance. The skin is literally turned 
inside out over the caudal, dorsal and anal fins, the fish inflat- 
ing to such an extent that these fins are buried in furrows in 
the distended skin. In preserved specimens the dorsal outline 
over the eye is extended by a fold of this expansive membrane, 
which shows in the deflated specimen illustrated by Fig. 86. 
The projecting pectorals are practically the only prominences 
of note on the otherwise spherical surface of an inflated speci- 
men. As shown in Figs. 938 and 94, by the time the species 
attains a length of 22.5 mm. ‘their inflatibility does not exceed 
that of the adults. The entire skin, however, is still loose, and 
upon inflation, as the drawing indicates, the membrane covering 
the eye becomes pulled down to a certain extent.. 


Latham, 1916, reports having seen specimens 2.5 to 7.5 
em. at Orient, Long Island, during November and the early 
part of December. It may be inferred that the smaller ones 
at least must have been hatched that same year, when it is con- 
sidered that specimens of over 7 mm. were taken in July by the 
Grampus. This, together with the fact that specimens in 
graduated sizes from a few mm. in length to adult size have 
been taken at most diverse seasons, indicates a long spawning 
season. 


Preserved material—Examples of all stages studied were 
preserved in strong formalin. The most notable change in the 
younger eggs was the change of the oil globules to a decidedly 
amber color, and their migration from the interior of the yolk 
to its outside wall, at which surface they mostly adhered. The 
blastodise and the early embryos became white and opaque. All 
chromatophores lost their color and were difficult to differenti- 
ate, excepting the black ones. The prominent chrome yellow 
spot nearly disappeared, and the lateral glands were almost in- 
distinguishable. Staining with alizerin red failed to aid much 
in bringing out these details. 


*Latham, Roy—Migration notes of fishes from Orient, Long Island. 
Copeia, March 24, 1916, No. 41, p. 22. 


1921] Welsh & Breder: Life History of the Puffer 213 


FOoop. 


The stomachs of 102 specimens taken between July 30 and 
August 4 were examined immediately on capture. The range of 
total lengths was from 14 to 24 em. Some individuals were 
spawned out, others still hard, although the majority were ripe. 
No variation in diet could be correlated with size, sex or condi- 
tion. The males were fifty-nine in number and the females 
forty-three. 


The accompanying table indicates the number of stomachs 
containing each constituent of their food. 


Material Number of stomachs. 
SUPT eA NM RUIGE pete ek eee eae Sh 9 OAL Eee ure ts 14 
Wnidentined crustaceans (20.000) 20) 28 Be Zi 
LIPEISSYeViC Ce tad SE IGM Se a Make De any oni cette SEO ea? Oe eee i 
LENDS EU A SS So cee SO ec pt SP SRR RRO Se it 
VG PTrE Sra TN elo IY See ene sie oes Oe BORE EN ecien 82 
JE TTO TAY yee Oke 8 UR See UAT En ensue elles eee ae Soma 16 


The crabs were all of small non-commercial species. The 
unidentified material consisted largely of matter reduced to an 
offensive blue or yellowish paste. The apparent discrepancy in 
numbers is accounted for by the fact that some specimens par- 
took of more than one kind of food. 


Linton, 1905°, found on examining fifteen specimens dur- 
ing July and August at Beaufort, N. C., that their food in- 
cluded fragments of oysters, scallops, mussels, razor-clams, gas- 
teropods, barnacles, crabs, shrimp, sea-urchins, worms, ascid- 
ians, bryozoans, and watermelon seed, which gives the species 
a much more varied diet than found in the present investi- 
gation. 


RELATIVE SIZE OF THE SEXES. 


The females of this species average somewhat larger than 
the males. Inspection of Fig. 95 shows that there is a differ- 
ence of 3 cm. between the modal lengths of the two sexes. The 


“Linton, Edwin.—Parasites of the fishes of Beaufort, N. C. In Bulletin, 
U. S. Bureau of Fisheries, Vol. XXIV, 1904, p. 402. 


274 Zoologica: N. Y. Zoological Society 7 (Thee 


FIG. 95. 


Horizontal index—Total lengths in centimeters. Vertical index—Number of individ- 
uals. Males represented by hatching from upper left to lower right corner. Females 
represented by hatching from upper right to lower left corner. 


difference of the averages is slightly less, being about 2.85 cm. 
The 102 specimens examined for food determination were 
measured for plotting this graph. It is evident that fish more 
than 22 cm. long are almost surely females, while those less 
than 17 cm. are nearly all males. The sole female of 14 cm. 
length was probably a large specimen of a younger group, the 
remainder of which probably passed through the meshes of 
the net or were chiefly located in another area. 


This knowledge proved very useful when dissecting the 
fish for eggs and milt, as by simply selecting the largest and 
smallest examples of a catch, both sexes could be had with al- 
most entire certainty. 


While both gonads were functioning in ripe fish, in all 
cases the left one was considerably larger than the right. 


THE RELATION OF LENGTH TO WEIGHT. 


The total lengths and weights of the 102 specimens of both 
sexes formed the basis of Fig. 96 which is arranged to show 
the weight of a fish of any given length or vice versa. These 
are plotted against each other, the larger circles indicating the 
points obtained by averaging the weights of all specimens in 
their respective length groups, which are in steps of 1 cm. The 
small circles on either side of the large ones indicate the high- 


1921] Welsh & Breder: Life History of the Puffer 21D 


RIG. 96. 


Horizontal index—Weights in grams. Vertical index—Total lengths in centimeters. 

Large circles—Average weight of corresponding length group. Small circles—Extremes 

in weight for corresponding length group. Numbers near large circles—Number of 

specimens making up average. Sex symbols—Average weight of sex indicated for 

corresponding length group. Center heavy curve—Line of average weights, smoothed. 

Outer heavy curves—Lines of extreme variation of weights. Heavy dashed curve— 
Line of formula, w=1°/56. 


est and lowest weights found in the individual groups. The 
central heavy curved line is the theoretical curve obtained by 
smoothing the dotted line connecting the large circles. It 
graphically represents the change in relation of length and 
weight as growth increases. The two outer heavy curved lines 
represent the limit of individual variation as established by the 
plotted points. The sex symbols connected by light solid lines 
shows the average weight for the indicated sex in each length 
group. This shows well the fact that males weigh considerably 
less than females of a given length. The modal weight of the 
males is about eighty-two grams less than that of the females, 
while the difference of the average weights is about eighty- 
seven grams. 


It is evident that having the length or weight of a fish of 
this species, the other may be had by inspection of the curve, 


276 Zoologica: N. Y. Zoological Society BO bee: 


the central lines giving the probable measurement, while the 
outer two limit the known variation in the breeding season. If 
the specimen in question is a male its weight will in all proba- 
bility fall between the central line and the left hand outer one, 
while if it is a female it will probably be located on the other 
side of the central line. 


On the assumption that the weights of fish of this species 
vary as the cube of their length multiplied by some constant, as 
is usual in fishes, the formula w=1*/56 was calculated in 
which w = weight in grams, 1= length in ecm., and 56 =the 
constant. The heavy dashed curve was plotted from this equa- 
tion. While it does not follow the smoothed curve perfectly it 
keeps well within the limitations required and the small dis- 
crepancies can be accounted for by the fact that the specimens 
weighed varied in sex, development of the gonads, and amount 
of material present in the alimentary tract. 


SUM MARY. 


1. Spheroides maculatus has a long spawning season, prob- 
ably lasting all through the warmer months. 

2. Artificial fertilization is readily accomplished and ow- 
ing to the transparency, adhesive nature, and hardihood of the 
ova, as well as the vitality of the larvae, the species is ideally 
suited for studies of early development. 

3. Incubation occupies about three days and ten hours at 
an average temperature of 67°F. 

4. The critical period for the larvae of this species ap- 
pears to be on or about the tenth day after hatching. 

5. Preservation in formulin destroys the finer details but 
fails to make the eggs or larvae unidentifiable. 

6. The food of Spheroides does not include Goes TSI of 
any considerable commercial value. 

7. The females average larger than the males. 

8. In specimens of equal length the females are some- 
what heavier than the males, at least in the breeding season, 
although this is irrespective of the condition of the gonads. 

9. The relation of length to weight in this species is ap- 
proximated by the formula w = 1°/56. 


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Destruction of Our Birds and Mammals (Hornaday).. 15 
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The Origin and Relationship, Large Manna: North 
PATO TIGA CRTILIED Fora ics Legis = sie aya sig adh awed & oP ae als, woe.ohe wees 


The Rocky Mountain Goat (Grant)*.............000045 
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OBJECTS OF THE SOCIETY. 


A PUBLIC ZOOLOGICAL PARK 
A PUBLIC AQUARIUM 
THE PRESERVATION OF OUR NATIVE ANIMALS 
THE PROMOTION OF ZOOLOGY 


ZOOLOGICA 
SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME Il. NUMBER 13 


(Papers from the New York Aquarium) 
(Contribution Number 6) 


HERMAPHRODITISM OF A CROAKER, 
MICROPOGON UNDULATUS (LINNAEUS) 


By C. M. BREDER, JR. 


New York Aquarium 


NV. 
‘4; Laney 
tional Muse’ 


RU ee ins. Ae. Di Boye. TB 5,0; OC: TP Bhetsy 
THE ZOOLOGICAL PARK, NEW YORK 


JANUARY, 1922 


New York Zoulngical Society 


General Office: 111 Broadway, New York City 


Officers . 

President, HENRY FAIRFIELD OSBORN; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 

Treasurer, Cornrtivs R. AcNew 


Board of Managers 
Glass nf 1923 
HENRY Been OSBORN, LISPENARD STEWART, CHARLES F. 
DIETERICH, GEORGE F. BAKER, Wo. PIERSON HAMILTON, 
ROBERT S. BREWSTER, EDWARD §. HARKNESS, 
WILLIAM B. Oscoop FIELD, A. BARTON 
HEPBURN, WILLIAM WOODWARD, 
EDWIN THORNE, Percy A. 
ROCKEFELLER. 
Glass nf 1924 
MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, 
GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFURD 
Morris, ARCHER M. HUNTINGTON, GEORGE OD. 
PRATT, T. COLEMAN DUPONT, HENRY D. WHITON, 
EDWARD HATCH, JR., CORNELIUS R:. AGNEW 
@lass nf 1925 
PreRcY R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE, 
C. LEDYARD BLAIR, EMERSON McMILiIN, ANTHONY R. 
KUSER, WATSON B. DICKERMAN, MORTIMER L. 
SCHIFF, FREDERIC C. WALCOTT, BEEKMAN 
WINTHROP, GEORGE C. CLARK, ; 
W. REDMOND CROSS. 


Sricntifce Staif 
WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 
WILLIAM BEEBE, Honorary Curator of Birds and Director of the 
Tropical Research Station; 
LEE 8S. CRANDALL, Curator of Birds; 
GEORGE S. HUNTINGTON, Prosector: 
GEORGE A. MACCALLUM, Pathologist; 
‘W. REID BLAIR, Veterinarian; 
ELWIN R. SANBORN, Photographer and Editor. 


Enitorial Committee 
HENRY FAIRFIELD OSBORN, Chairman; 
: WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 


Corrected to January, 1922. 


ah Insti Gi, 
\ gona 
4 hs 
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; } ¢ 1929 
fa 5 


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LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW VORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 13 


(Papers from the New York Aquarium) 
(Contribution Number 6) 


HERMAPHRODITISM OF A CROAKER, 
MICROPOGON UNDULATUS (LINNAEUS) 


By C. M. BREDER, JR. 
New York Aquarium 


Betiebe lal Son ED. 5B. Y) ) Tene » OC LEE Y 
DiiteesAQOLOGlLe Ah, “PARK. NEW . YORK 


JANUARY, 1922 


ana Up 


Volume II, Number 13 


HERMAPHRODITISM OF A CROAKER,} 
MICROPOGON UNDULATUS (LINNAEUS) 


By C. M. BREDER, JR. 
New York Aquarium 


During the months of July and August, 1920, a large num- 
ber of croakers, Micropogon undulatus (Linnaeus), were taken 
in the pound nets operated at Young’s Million Dollar Pier, 
Atlantic City, New Jersey. They appeared to be approaching 
the spawning season rapidly and some males were found from 
which milt would flow. No females were taken with ripe eggs 
however, so on August 9, when what appeared to be a female 
turgid with eggs was taken, considerable interest was aroused. 
Its proportions were as follows: 


ALORS SUT nee eee ae OER eRe ed CL ee, Ie 32. (cin: 
SSLeneli emt OM Ob Wiha tee ee Ne 2 em: 
GG amOCi bin. emer nis . - bee OU bares 9.5 cm. 


Externally it appeared to be normal in all respects and 
when it was found that stripping was not possible, curiosity 
prompted dissection. The explanation of its great body depth 
was apparent when it was seen that perfect sets of both ovaries 
and testes were present. The testes lay dorsal of the ovaries, 
but in all other respects the internal anatomy was of the usual 
character. 


In the accompanying semi-diagramatic sketch of the 
ventral aspect of the dissected specimen all fatty tissue and 
mesentary membranes have been omitted. Only the digestive 
tract and sexual organs have been shown in detail. The stom- 
ach appears as a very small appendage, but that is the normal 


*Published by permission of the United States Commissioner of 
Fisheries. 


a ; “16 “OIA 


VOAaVo 
StMHOTAd LYMVSH 


SHBUS2UE 


Rasy 


a 


1921) Breder: Hermaphroditism of a Croaker 283 


condition of the organ when the individual has not been 
feeding. 


The only displacement of the viscera, other than that 
caused by the spreading due to opening the fish, is that of 
moving the anterior ends of the ovaries outward so as to show 
the testes lying dorsally of them. Before this separation, the 
ovaries were nearly parallel to each other and appressed against 
either side of the intestine. The paired fins indicated are the 
ventrals, the pectorals not being shown as the open flaps of 
body wall hide them completely. 


The junction of the ova, sperm and urinary ducts ap- 
peared to be at the genital pore. This is shown in the diagram 
just posterior to the vent. The spermatic ducts can be seen 
passing around the rear end of the distended ovaries, which 
extend backward past the genital pore. The urinary duct 
passes between the posterior ends of the ovaries as it descends 
from the kidneys lying dorsally of the swim bladder, and is 
flexed forward to the external opening. 


Both sets of gonads seemed well developed and in the state 
most frequently found in normal individuals of this species 
taken about the same time as this sportive example. The testes 
were soft and flocculent, and easily ruptured, the milt stream- 
ing out from such injuries, but they were not quite ripe enough 
to strip, and the ovaries were also a little too green for that 
operation. 


Most previous records of hermaphroditism in fishes tell of 
one set being much in advance of the other in regard to devel- 
opment, but this specimen suggests speculation on the possi- 
bility of self-fertilization, which mechanically, at least, appears 
to be entirely possible. Scale examination and size indicate that 
this fish was about five years old and has therefore passed 
through at least one spawning season. The age of the fish to- 
gether with the fact that the gonads were normal in themselves, 
strongly suggests that this is a case of functional hermaphro- 
ditism of which we have three possible methods of function: 
that is, self fertilization, and mating with other fish, either as 


284 Zoologica: N. Y. Zoological Society [ItetS 


the male or female element or alternately, as first one and then 
the other. 


To the best of the writer’s knowledge bi-sexuality in tele- 
osts has been recorded only from the following families and 
orders; Cyprinidae, Clupeidae, Salmonidae, Esocidae, Poecilii- 
dae, Gasterosteidae, Mugilidae, Percidae, Serranidae, Sparidae, 
Scombridae, Labridae, Squamipinnes, Gadidae and Plewronec- 
tidae.? 

It is believed that no additions have been made to the above 
list up to the present time, so this notice stands as record of 
the addition of the Sciaenidae to it. 


C. Stewart, in the Journal of the London Linnean Society 
(Zool.) 24 pp. 70-71, mentions the most nearly similar case, 
which is among the Scombridae. This is apparently the only 
other record in which the abnormality has approached sym- 
metry; but even in the case of the mackerel the organs failed 
to reach the degree of symmetry found in the present specimen. 


This example of abnormal hermaphroditism is now de- 
posited in the U. 8S. National Museum, number 66140. 


2This list, excepting the Poeciliidae and Serranidae was recorded by 
James E. Gemmill in his publication, “The Teratology of Fishes”, 1912, 
James Maclehase & Sons, Glasgow, Pub. The occurrence of this type of 
monstrosity among the Poeciliidae was recorded by H. H. Newman, 1908 
“A Significant case of Hermaphoditism in fish”, Biol. Bull. Woods Hole, 
Mass., pp. 207-214. This reference however was listed in Gemmill’s bibliog- 
raphy. The Serranidae are included in this list on D. S. Jordan’s authority. 
In his “Guide to the Study of Fishes” 1905, p. 124, Holt & Co., N. Y., Pub., 
he states that Serranus is “sometimes truly hermaphroditic.” 


Piublications 


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Four royal quarto volumes, illustrated with reproduc- 
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Knight, Fuertes and Jones, and many photogravures’ * 


- and maps. Sold im sets only. Vols. I and II, ready. 


Paper 
Our Vanishing Wild Life (Hornaday)......:.......... 
Destruction of Our Birds and Mammals (Hornaday). . 15 
Notes on Mountain Sheep of North America (Hor naday) 40 
OE OUE G7 ILO Diora aoe cela oe See A UR .40 
The Origin and Relationship, Large Mammals, North 
PETC AC Ce TILL DS oi in idee vs cen oy AGS. A Vio id ch ADEE aT ola 
Phe: Rocky Mountain Goat (Grant). oar ee ee a 
en and Horns Brochure (Hornaday), Nos. 1 and 2, 
is PETE Rs Ge au racg Hike eae Sle a ee eee: 1.00 
Tropical Wild Life (Beebe; Hartley; Howes)........... 
Zoologica, Vol. I, Nos: 1-20 incl. (In parts)............ 3.85 
Zoologica, Vol. a Nos.) 1-20 melo (bound) 25 325 seakioes 
Zoologica, Vol. IL, POSH (1532; SO, COMY, sins wien ae Ve ieee 25 
Aoolorica Viel AT ING: Gs) CODY coos 3 oe es 2s Sal ee Wage ren 75 
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Zoopathologica, Vol. Te Nos tbo Dy CODY 2a. es Roe a ae 25 
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Official Guide, Zoological Parke Hornadat):c 2s. bse ee .50, 
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- Northern Elephant Seal (Townsend), paper............ AT a 
Porpoise in Captivity (Townsend), paper............... 2b 
Natural History of the Whale Shark (Gudger)......... .25 
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Illustrated Guide to the Aquarium (Townsend), boards. .50 
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x 


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All publications of the Society for sale at the Zoological Park, 185th St. and 
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Corrected to January, 1922 


Y) 


‘THE PRESERVATION oF OUR NATIVE . 


ASey 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 14 


THE FOOD OF CERTAIN MINNOWS 


A STUDY OF THE SEASONAL DIETARY CYCLE OF SIX 
CYPRINOIDS WITH ESPECIAL REFERENCE 
TO FISH CULTURE 


By C. M. BREDER, JR. 
New York Aquarium 


AND 


D. R. CRAWFORD 
United States Bureau of Fisheries 


Bi Seda dose De BOY Tae SF. O.Cl Beegiy 
THE ZOOLOGICAL PARK, NEW YORK 


August 15, 1922 


Nem York Zoological Sorivty 


General Office: 111 Broadway, New York City 


Officers 
President, HENRY FAIRFIELD OSBoRN; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; 
Secretary, Chairman, Exec. Committee, MADISON GRANT; 
Treasurer, CORNELIUS R. AGNEW 


Board of Managers 
Glass of 1923 


HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. 
DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON, 
RoBERT S.' BREWSTER, EDWARD S. HARKNESS, 
WILLIAM B. Oscoop FIELD, WILLIAM 
WOODWARD, EDWIN THORNE, 

PERCY A. ROCKEFELLER, 

JOHN E. BERWIND. 

Glass nf 1924 
MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, 
GEORGE J. GouLp, OGDEN MILLS, LEWIS RUTHERFURD 
Morris, ARCHER M. HUNTINGTON, GEORGE D. 
PRATT, T. COLEMAN DUPONT, HENRY D. WHITON, 
EDWARD HATCH, JR., CORNELIUS R. AGNEW 

! / Glass of 1925 
anew ee PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE, 
a OF LEDYARD BLAIR, ANTHONY R. KUSER, WATSON 
B. DICKERMAN, Mortimer L. SCHIFF, 
FREDERIC C. WALCOTT, BEEKMAN 
WINTHROP, GEORGE C. CLARK, 
W. REDMOND CROSS. 


Srientific Stait 
WILLIAM T. HORNADAY, Director of the Zoological Park; 
CHARLES H. TOWNSEND, Director of the Aquarium; 
RAYMOND L. DITMARS, Curator of Reptiles; 
WILLIAM BEEBE, Honorary Curator of Birds and Director of the 
Tropical Research Station; 
LEE 8. CRANDALL, Curator of Birds; 
GEORGE 8S. HUNTINGTON, Prosector; 
' GEORGE A. MACCALLUM, "Pathologist; 
W. REID BLAIR, Veterinarian: 
, ° ELWIN R. SANBORN, Photographer and Editor. 


Editorial Conunitteer 


HENRY FAIRFIELD OSBORN, Chairman; 
WILLIAM T. HORNADAY, . CHARLES H. TOWNSEND. 


Corrected to July, 1922 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 14 


THE FOOD OF CERTAIN MINNOWS 


A STUDY OF THE SEASONAL DIETARY CYCLE OF SIX 
CYPRINOIDS WITH ESPECIAL REFERENCE 
TO FISH CULTURE 


By C. M. BREDER, JR. 


New York Aquarium 


AND 


D. R. CRAWFORD 
United States Bureau of Fisheries 


Ptrogalio hi D BY THE SO Col Er y 
THE ZOOLOGICAL PARK, NEW YORK 


AucusT 15, 1922 


Volume II, Number 14 


THE FOOD OF CERTAIN MINNOWS 


A STUDY OF THE SEASONAL DIETARY CYCLE OF SIX 
CYPRINOIDS WITH ESPECIAL REFERENCE 
TO FISH CULTURE 


By C. M. BREDER, JR. 
New York Aquarium 
AND 
D. R. CRAWFORD 
United States Bureau of Fisheries 


INTRODUCTION 


The smaller members of the family Cyprinide although often 
neglected, are without doubt the most important of the small 
fishes which inhabit our fresh waters. To the practical fish 
culturist they are of greater importance than is usually recog- 
nized since they represent the food of many game and food 
fishes, while their diversity of species and habits, together with 
their interesting ecological relations, and availability, make them 
of considerable interest to the scientific student. A study of 
their intimate relationships and habits, therefore, is at once im- 
portant from an economic and scientific standpoint. With these 
considerations in mind, this study of the food and feeding habits 
was essayed, since they are undoubtedly among the chief factors 
in the lives of these fishes. 


FISH CULTURAL VALUE 


The fact of primary importance to fish culture is that these 
minnows enter largely into the diets of the larger game fishes. 
At least twenty-three different predatory fishes are known to sub- 
sist largely upon various cyprinoids. They are as follows: 


Lepidosteussossems, (Winneus) = 2. ...-1.<- seine Long-nosed Gar or Billfish 
Hiodon alosoides (Rafinesque)............+++: Golden-eye or Northern Moon-eye 
Ligodormtennasius) Le iSUeui-. safe ci <2 1s 2c eee > Golden-eye or Moon-eye 
Pomolobus chrysochloris Rafinesque..........-- Skipjack 


Salmomsevagom (Girard). oy ccc ae + cis stesso 1s oheneks Sebago Salmon 


288 Zoologica: N. Y. Zoological Society MU! 


Cristovomer namaycush (Walbaum).......... Lake Trout 

Soak? CaaGaemny (Given) .onccoonossoneucuec Little or Bonded Pickerel 
Eoxenenculatusm Wexsueun) memento eee Pickerel or Pike 

sox lucius, Winn euss ces cro ners sree © ee eee Pike 

Esoxnminasquinongy aMutchilll eae sere neers Muskallunge 

A phredoderus sayanus (Gilliams)............. Pirate Perch 

Pomoxis annularis Rafinesque.........,...-.--- Crappie or Croppie 
Pomoxis sparoides, (Wacépede))..5.-+-4--+4 +--+ Black Crappie or Calico Bass 
Ambloplites rupestris (Rafinesque)............ Rock Bass 

Chaenobryttus gulosus (Cuvier and Valenciennes) Warmouth 

Micropterus dolomieu Lacépéde............... Small Mouthed Black Bass 
Micropterus salmoides (Lacépéde)............ Large Mouthed Black Bass 
Stizostedion vitreum (Mitchill)............... Wall Eyed Pike 
Stizostedion canadense (De Kay)............. Gray Pike or Sauger 
Pepea WEeSeGne (MI OwIl)) 5 soccegc00san500000 Yellow Perch 

IROGEHS- CUI HG (IREMWINESC WS) so s0ccc0b0bG000ccc White Bass 
Roacusmiuneartism (Bloch) epee Striped Bass 

Morone americana (Gmelin)................. White Perch 


After listing a number of species (included above), Forbes and 
Richardson (’08), add: ‘‘That th’s list might be considerably en- 
larged by more extensive studies of the food of fishes is beyond 
a doubt, and it is safe to say that no fish-eating fish would, if 
hungry for fish, refuse a minnow of any kind unless it seemed 
too small to be worth the trouble capturing. . . . Moreover, 
by their great numbers, by their various adaptations and corre- 
sponding ecological d'stribution, and by their permanently small 
size, the minnows must distract in great measure the attention 
of carnivorous fishes from the young of the larger species, upon 
which, without them, the adults of these larger species would 
fall with the full force of their voracious appetites. . . . It 
is not too much to say, consequently, that the number of game 
fishes which any waters can maintain is largely conditioned upon 
its permanent stock of minnows.” 


Since the successful stocking of any stream with game fishes 
depends upon the food supply, it is essential to know how this 
important item may be maintained. As Forbes (’83) remarked, 
“Really intelligent fish-culture on any large scale, implies a full 
acquaintance with the food of the native species.” 


Fortunately, most of our streams east of the Rocky Moun- 
tains seem to be plentifully supplied with Cyprinoids, numerous 
species being found frequently in one stream. The availability 
of these fishes as a staple food for larger fishes depends upon 
their abundance, which, in turn, depends upon the abundance of 
their food. It is important, therefore, to know what these min- 


1922] Breder & Crawford: Food of Minnows 289 


nows eat at various seasons of the year. For this reason a con- 
siderable portion of this paper is devoted to the analysis of the 
stomach contents of specimens taken at various seasons. 


SCIENTIFIC VALUE 


Ecologically and taxonomically the Cyprinide form a puz- 
zling group. A study of the affinities of the various species is 
needed, both with reference to anatomical details and environ- 
mental conditions. In this connection, a knowledge of the food 
of the group throughout the seasons is of prime necessity and the 
results embodied in this paper, it is hoped, may be of value to 
other workers. 


MINOR VALUES 


This group of fishes furnishes a large per cent of the bait 
to fishermen, sport for many a small boy, and a few of the larger 
species are used for human consumption. 


Any or all of the different species of suitable size may be used 
as bait, many being hardy as live bait, but Notropis cornutus is 
probably the favorite among most of the fishermen. Semotilus 
bullaris is a wary fish and in some localities, at least, it is regarded 
as a minor sport fish. 


Lastly, many of the species, in fact all of those which were 
collected in connection with this work, are attractive aquarium 
fishes, most of them becoming adapted to balanced aquaria, al- 
though they become adapted more readily to conditions in an 
aquarium supplied with running water. 


FIELD METHODS 


In order to determine the nature of their food, collections 
were made of six of the common species of cyprinoids occuring 
in the District of Columbia. 


A small stream known as Oxon Run was chosen for this 
purpose because of its accessibility and various physical features 
which will be subsequently described. It was decided that collec- 
tions should be made once a month for one entire year. This plan 
was followed, except that no collection was made in August, but 
the collecting dates were so arranged that the greatest gap be- 


290 Zoologica: N. Y. Zoological Society . [Tks 14 


WATER LEVEL 


SHRUBS 


PROFILE Y RUN et A—B 


SHRUBS \\ NV 


QXON RUN 
CONTOUR INTERWAR 
ON STREAM BED - 5° 
CONTOUR INTERVAL 
ON STREAM BANK - 2' 

SCALE 


FIG. 98. MAP OF COLLECTING SITE 


The two curved lines crossing the stream mark the limits of the area in which the 
collections were made. This locality is indicated on the Washington Biological 
Sociely’s Map (MacAtee °18) as S. E. G. 12. 


tween any two was only 49 days, which period spanned that 
month. 


Oxon Run is about 7.5 miles long and flows into the Potomac 
River near the southeast boundary of the District, opposite Alex- 
andria, Va. In every way it is a typical steam such as found in the 
coastal plain region of Virginia and Maryland. It was observed 
that there was a considerable fluctuation in the amount of flow, 
depending on the amount of rainfall, and consequently, the turbid- 
ity varied accordingly. These changes were not detrimental to 
the work, although the swift current increased the difficulty of 
hauling the seine. The location shown on the map (Fig. 98) is 
about 214, miles above the mouth and was chosen because there is 
a rather deep eddy, on one side of which is a small beach provid- 
ing an ideal place for hauling out the seine. The bank opposite this 
beach is rather high with overhanging shrubs, grasses and other 
plants. Upstream, the channel is nearly flat and has a gravelly 


1922] Breder & Crawford: Food of Minnows 291 


TABLE NO. I 
PHYSIOGRAPHICAL CONDITIONS AT OXON RUN DURING 1920 


Temp. (Fahr.) CONDITION OF WATER WEATHER 
DATE AiR WATER ‘TursipiTy, ETc. DEPTH CONDITIONS 
Tens ls Cees meee 48 37 Rather swift, 12 inches Partly 
fairly clear cloudy 
Ble al areca eves a 29 32 2 inch ice over Greatest, Snow flurries. 
area seined 36 inches Cloudy 
March 14 ..... 42.5 38 Slightly Evidence of Fair and 
murky previous high windy 
water 
ATA 27. eos 59 58 Fair 
Waly i235) juiesr-ie's of 63 Partly cloudy; 
warm 
[one TE eae 89 71 Clear Quite low Partly cloudy 
Jilly Wire 2st 80 74 Very roily from Sunny; hot 
children swim- 
ming 
SG@ptteAeetva ave. 83 66 Very clear Low Clear or partly 
cloudy 
OGE Anas arco sys 75 57, Very clear Very low Fair 
INOweei sean tere 61 48 Clear Low Cloudy 
Def oRI ST ahs ater ae 65.5 49 Turbid, swift Highest noted 


bottom that slopes gently, the water being about six inches deep 
at the low water stages. Down stream there are deeper places 
which were found unsuitable for seining. The table of physio- 
graphic conditions (Table No. I.) shows the general conditions 
and seasonal variations under which the collections were made. 


All of our collections were made within the limits shown on 
the map by means of a ten-foot seine of one-quarter inch mesh. 
The bottom was quite free from snags and weeds of any kind 
and the only cover provided for the fishes was such debris as 
dead leaves and other materials which may have collected in the 
eddy. Even when the water was clear, few fishes were to be seen, 
yet the number of specimens collected gives some evidence as to 
their abundance. No definite number of hauls was made, since 
the catch each time was found to vary considerably, collecting 


292 Zoologica: N. Y. Zoological Society [II; 14 


being continued until a sufficient number of specimens had been 
secured, or until the site had been exhausted temporarily of fish. 
The specimens were placed immediately into formal alcohol and 
sorted later in the laboratory. Formal alcohol was found to be an 
efficient killing fluid since it acted quickly thus preventing further 
digestion of the stomach contents and hardened the specimens 
without perceptible shrinkage. 


LABORATORY METHODS 


After the specimens were brought to the laboratory, the solu- 
tion of formal alcohol was poured off and 75 per cent. alcohol sub- 
stituted for permanent preservation. Each species was preserved 
in a separate bottle and the different monthly collections were 
also segregated. Each fish was measured, the standard length 
being recorded because this measurement was used in construct- 
ing curves of growth which are discussed further on. Since this 
is the only measurement referred to throughout the paper it is 
mentioned subsequently simply as length. Each specimen was 
provided with a paper tag numbered serially to provide a ready 
reference to each specimen. In all, there were 1554 specimens 
including six species. 


The entire digestive tract of each specimen was removed 
and the contents pressed out on a glass slide. The material was 
examined with a low power of the compound microscope supple- 
mented by higher powers when necessary. Pierce’s method 
(Pierce ’15)! was used to estimate the various quantities of food 
present. Itis apparent that such a method can yield only a rough 
estimate, but none other was found to be feasible on account of 
the time required to make more accurate volumetric determina- 
tions. It is pointed out that frequently only small quantities of 
food, such as the leg of a beetle or wing of a fly, would be found 
yet such material had to be listed as 100 per cent. coleopterous, or 
dipterous remains, as the case might be, since there was no evi- 
dence of other food having been eaten. However, these errors 
would be naturally compensating rather than cumulative and in 
the tables we present giving the averages of each collection, they 
lose significance. 


1 Briefly, in this method the contents of each example is considered as unity, the various 
items being expressed in terms ci percentage by volume as estimated by inspection. 


In t 


FIG. 99. 


aking these photos the camera was located a little beyond the upper 


VIEW LOOKING UP STREAM 


tions were made. Taken in early October. 


FIG. 100. VIEW LOOKING 


See Map, page 290. 


DOWN STREAM 


and lower limits, respectively, of the area in which the collec- 


Tol. TI, No. 13 
? 


Zoologica J 
ace page 29 


15 


FIG. 101. ENTIRE CONTENTS OF THE STOMACH AND INTESTINE 
OF A SPECIMEN OF NOTROPIS CORNUTUS 7.3 CM. IN LENGTH 
TAKEN IN DECEMBER. 4X 


Laue ene Wevees ee Your 


ye & fo ae 
. y : a 
eet % i 


Ps ae pe t. ach a. 
Yas ee se ei 
af oe cathe a Ace. a 3 


FIG. 102. ENTIRE CONTENTS OF THE STOMACH AND INTESTINE 
OF A SPECIMEN OF NOTROPIS CORNUTUS 5.7 CM IN LENGTH 
TAKEN IN DECEMBER. 4X 

Photomicrographs showing the degree to which the food was usually found 

to be macerated. 


Zoologica Vol. TT, No. 14 
Face page 293 


1922] Breder & Crawford: Food of Minnows: 293 


The food was found to be masticated usually beyond the pos- 
sibility of positive identification, as illustrated by figs. 101 and 102. 
This condition, doubtless, was caused by the action of the rap- 
torial pharyngeal teeth possessed by these species. However, it 
was possible to distinguish insect remains from those of other 
organisms and often the various orders of insects such as 
Coleoptera, Diptera, etc., could be separated. 


Our thanks are due Mr. J. T. Nichols, of the American 
Museum of Natural History at whose suggestion the problem was 
undertaken, to Dr. W. C. Kendall, of the U. S. Bureau of Fisheries, 
for determining the identity of certain specimens, and to Dr. R. 
EK. Coker, for his kindness in allowing the use of the facilities of 
the Division of Inquiry of the latter institution. All illustrations 
have been made by the senior author. 


OBSERVATIONS 


Arrangement of Treatment by Species—Complete state- 
ments and tables of the stomach contents of the six species on 
which this paper is based follow. A few general notes are fol- 
lowed by a discussion of the analysis of the data obtained. Fol- 
lowing this, is the table of foods given in volumetric percentage 
by months. Opposite each date of collection is given the number 
of specimens which contained food and the number which were 
found to be empty. The modal, maximum and minimum lengths 
in millimeters are given next to convey a general idea of the size 
of the specimens examined. In cases in which no modal length is 
given the specimens were so scattered or few in number that no 
distinct mode was discernable from the frequency graph. Of 
course, in other cases the significance of the mode is directly 
proportional to the number of variants measured. In plotting 
these on graphic paper groupings of 4.0 mm. each were used 
throughout, beginning with 0 to 4 as the first group. 


The body of the table in which the various organic substances 
are arranged in systematic order follows next, with the unidenti- 
fied materials placed last. The figures for any one month indicate 
the averages for that collection. They have been reduced in prac- 
tically all cases to whole numbers because figures closer than 0.5 
per cent. have no particular significance, owing to the variation 
of the estimates and other uncontrollable factors. All such mate- 


294 - Zoologica: N. Y. Zoological Society 2) ABS 


rials present in quantities of less than 0.5 per cent. are indicated 
simply by a plus sign indicating a mere trace. The general aver- 
age for the whole year differs slightly from that which would be 
obtained by averaging the monthly numbers since the original 
calculations with all their fractional parts have been used because 
frequently the aggregate number of plus signs was sufficient to 
form a whole number. This grand average then also was 
smoothed in a similar manner to the method used for the monthly 
averages. Here, also, the value of the results is directly propor- 
tional to the number of specimens. 


- The table is followed in each case by a list of annotations 
amplifying and explaining certain features not expressed in the 
table itself or its preceding analysis. 


Semotilus bullaris (Rafinesque) FALLFISH 


Examples of Semotilus bullaris ranging from 1.6 to 12.2 cm. 
in length were taken. Those of the larger size were at once dis- 
tinguished from S. atromaculatus of which a considerable number 
were taken also, by the black spot at the base of the anterior rays 
of the dorsal fin, from which the species takes its name. The 
very small specimens, however, were extremely difficult since 
most of the adult charactertistics had not yet appeared, and it 
was only after considerable study that they were separated to 
our satisfaction. S. bullaris is separated from S. atromaculatus 
chiefly by its larger scales and the crowded conditions of the 
anterior costal scales of the latter which is not apparent in very 
small specimens of S. bullaris. Most of the specimens taken were 
immature. None below 4.0 cm. were mature while most of those 
of greater length were sexually developed. Graphs were con- 
structed plotting the lengths with their frequencies. Those below 
4.0 cm., which formed the majority, composed a well-defined 
group forming a single mode as is indicated in Table No. II. No 
second mode was formed since the larger specimens were too few 
in number to show any tendency in that direction. The small 
number of mature individuals suggests that probably the adults 
run up the stream to spawn. This would be in accordance with 
observations of this species in various lakes where it is known 
that the adults spawn in the streams. Fowler (’08) records this 


1922] Breder & Crawford: Food of Minnows 295 


species as reaching a length of eighteen inches which is several 
times the length of our largest specimen. 


Judging from the modal lengths of each collection these fish 
appeared to be chiefly of two-year classes; that is, most of those 
collected from January to July inclusive were undoubtedly of the 
spawning season of 1919. In September those of the 1920 spring 
began to appear in the collections, being by this time large enough 
to be unable to escape through the meshes of the seine. During 
the months of March, June, September, November, and December, 
a few larger individuals probably of greater age were taken. 


Analysis of Stomach Contents:—Reference to the accom- 
panying table plainly shows that this species is decidedly insec- 
tivorous, since these invertebrates formed 87 per cent. of the en- 
tire food of the 242 fish which were found to have been feeding. 
The plant remains amounting to only 5 per cent. apparently were 
taken incidentally. In September these remains were found to 
be present among the food in their greatest quantity (17 per 
cent.) but this relatively large amount in no way invalidates the 
conclusion that they were accidentally ingested, because none of 
the diatoms or algae were found to be plasmolized which would 
have been the case if any of them had been acted upon by digestive 
fluids; or if they had been macerated, the alcohol would have 
completed that action. This, together with the fact that the 
unidentified debris also amounted to only 5 per cent., suggests 
that these fish are not bottom feeders, and as they do not have a 
superior mouth, (Figs. 103 and 111), the inference is that they 
captured most of their prey as it fell through the water at some 
point below the surface. It was observed that specimens in the 
aquarium usually fed in this manner. Assuming this to be the 
truth, the lack of diatoms and algae would be explained since in 
the locality where the collections were made these plants must 
have been taken from the bottom because of the absence of 
larger plants, brush, etc., to which such growths could adhere 
and flourish, while the swiftness of the current precludes the 
existence of pelagic forms. Some of what they took might well 
have been broken fragments drifting down stream from above 
which were snapped up in passing. Additional evidence that this 
species indulged in bottom feeding to only a slight extent is 
furnished by the fact that not a single grain of sand was found 


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1922] Breder & Crawford: Food of Minnows 29% 


in any stomach, although in several of the other species sand 
was found quite commonly. 


No correlation of feeding habits and size was discerned in 
any case since all sizes fed on essentially the same types of organ- 
isms. The few exceptions noted are attributable simply to me- 
chanical differences due to size. As an example, in March a 
Boleosoma was eaten by a fish 86.5 mm. in length. It is obvious 
that some of the smaller specimens could not have eaten a Boleo- 
soma since many were smaller than the Darter itself. It may be 
noted here that not a single case of cannibalism was observed in 
any of the six species studied. 


The tabulations of the number of fish found to be empty 
plain!y shows that these fish feed considerably less during the cold 
months. On February 1, the coldest day on which a collection 
was made, when ice two inches thick was broken in order to 
operate the seine, twice as many of the stomachs were found to 
be empty as those which contained food and of the latter, two- 
thirds contained very little food. 


This species appeared to be nearly free of intestinal parasites 
and no other kinds were noted among the entire series of 298 
specimens. Only ten contained parasitic worms in the alimentary 
tract. These parasites were distributed as follows:—January, 4; 
February, 1; March, 4; April, 1. None was found in any suc- 
ceeding months. 


All foods other than insect were present in such small quan- 
tities that they cannot be considered important. 


It should be noted here also that such bottom forms as the 
larval Plecoptera and Ephemeroptera were taken only in the 
colder months when fewer terrestrial insects were available, and 
the fish were naturally seeking deeper water on account of low 
temperature at the surface. 


Incidentally fourteen specimens of S. atromaculatus were 
examined and it was found that they had partaken of food prac- 
tically identical to that taken by S. bullaris. 


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S}U9}JU0D YORWO}S JO UONE[NGe], SN]NSLOpuvaA SnIsisnaT III ON 292. 


1922] Breder & Crawford: Food of Minnows 299 


Leuciscus vandoisulus Cuvier and Valenciennes 
ROSY-S{DED DACE 


The specimens of Leuciscus vandoisulus that were collected 
for this study appeared to be of normal size. Fowler (’08) gives 
a length of from 134, to 3 3/16 inches which compares well with 
our maximum and minimum of 6.4 and 2.0 cm., (2.11 and 1.27 
inches). This species has been recorded as having a maximum 
length of 5.0 inches. Our series which of course included very 
young fish, did not include any which reached Fowler’s largest. 
There was no particular difficulty experienced in identifying the 
mature specimens because of the beautiful rose red streak on the 
sides brought out in the preservative both on the males and 
females. The young which did not have this distinctive mark 
were easily identified by their large gape of mouth and a certain 
dark pigment along one of the lateral fascia which showed 
through the skin and scales as a diagonal dark streak from shoul- 
der to tail. (Breder, ’20, b). This well served to separate this 
species from the other twenty-three taken at this locality. Ripe 
fish about to spawn were taken in May. 


At least four of the collections show two distinct modes in 
the frequency graphs which clearly divides the fishes into two 
year-classes (Breder ’20, a). The group of smaller specimens 
represents those which hatched in the spring or summer of 1919, 
while that of the larger fishes were mostly of the 1918 season, a 
few possibly being referable to 1917. In September, examples 
of fish hatched in the spring of 1920, entered the collections but 
so far overlapped those of 1919 as to merely shift the mode a 
trifle and lower the minimum lengths. 


No correlation appeared to exist between size and feeding 
habits of this species, mechanical limitations alone entering. 


Analysis of Stomach Contents:—The food of this species is 
practically identical with that of the preceding differing only in 
very minor details. In this species the insects amounted to 88 
per cent. and the vegetable content to only a trace, reaching only 
4 per cent. in the month of its greatest amount. In these fish the 
diatoms and alge also were found in an unplasmolized condition, 
and the unidentified debris averaged only 2 per cent. 


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$]U9}U0D YORUIO}S FO UOT}E[NQe J, auro4g SigosjON AI ‘ON PIGPL 


1922] Breder & Crawford: Food of Minnows 301 


The strongly oblique and capacious gape, (Figs. 104, 105, and 
112,) suggests that this species is inclined to feed from the sur- 
face. This is well borne out by the table of foods, since in all 
probability the Thysanura were taken while being supported on 
the surface film. In aquaria it was noted that this cyprinoid 
tended to keep nearer to the surface than any of its associates. 


In the specimen which had taken a larval caddis fly was 
found a small amount of detritus which was absent from the 
remaining specimens, 294 having been examined. This suggests 
that bottom feeding is only occasional. As in Semotilus, it is 
quite evident that the only regular bottom feeding was performed 
in the winter months with the exception of the above mentioned 
caddis fly and a dragon fly nymph taken in September. The low 
temperature and lack of food at the surface probably caused the 
descent of this species although some food could, no doubt, have 
been taken at times as it was released from melting blocks of ice 
as they drifted down stream. 


Lighter feeding in winter is not indicated by the number of 
empty stomachs or their distribution in time, although if this 
species had been taken in February and December such a condi- 
tion might have been suggested. Attention is called to the pos- 
sibility of a semi-hibernation or dormancy, because no specimens 
were taken on the two collecting days which were most cold. 


Notropis procne (Cope) DELAWARE MINNOW 


The examples of Notropis procne which were collected ranged 
in length from 1.6 to 5.2 cm. Fowler (’08) gives their lengths 
as reaching up to 2 11/16 inches which is a little in excess of 
what our maximum examples showed. It was easily distinguished 
from any of the other species which we took. However, we care- 
fully scrutinized each specimen in order to exclude N. bifernatus 
which has not been recorded from this region, but has been taken 
in Maryland, just north of the District. 


Five collections show what appears to be a double mode but 
the overlapping of the extremes is so great that it quite obscures 
any attempt to read the age of the various groups from such data 
alone. There is no differentiation of food with the advance in 
size. 


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19227) Breder & Crawford: Food of Minnows 303 


Analysis of the Stomach Contents:—It may be seen at a 
glance that the feeding habits of this species are somewhat differ- 
ent from those of the two preceding forms. Here the vegetable 
remains exceed those of the insect, the respective percentages 
being 47 and 36. Also, the comparatively large amount of sand 
indicates a bottom feeding habit. In aquaria this was not ob- 
served to be especially noticeable, since most of the specimens 
kept well up in the middle water. The rather subterminal mouth 
does not suggest that these fish are bottom feeders to a much 
greater extent than Semotilus, although the food points strongly 
to that conclusion. 


The number of empty stomachs and the months in which 
they were found indicates that this species also feeds less heavily 
in the winter than at other times. 


Notropis cornutus (Mitchill) REDFIN 


Our specimens of this species varied in length from 2.0 to 
9.6 cm. According to Fowler (’08) this species appears to reach 
a length of about twice that of our largest. Many of our larger 
specimens were breeding fish. All above 4.0 cm. were mature 
while all those of less length were juvenile. In the frequency 
graph this point coincides with the gap or point of greatest de- 
pression between two modes when such were distinctly present. 
Collections in which only one mode was evident a few scattering 
specimens were always to be found on the other side of this line 
of demarcation. Obviously there were two year-classes repre- 
sented here, and the group of smaller specimens was evidently 
from the spawning of the previous year, while the mature fish 
had passed through two or more winters. Here we have the same 
length of time required to reach maturity as was found necessary 
for Leuciscus. Ripe fish about to spawn were taken in May. 


No correlation between size and the food taken could be 
found. 

Analysis of Stomach Contents:—This species feeds upon 
about twice as much insect as vegetable matter. Judging from 
specimens in aquaria, they seem to be given to rather promiscuous 
feeding. As observed in captivity, they were noted to rise to the 
surface with both the force and grace of a trout although they 
seemed also to be quite adept at securing food from the bottom. 


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FIG. 103. SEMOTILUS BULLARIS 
Standard length 12.0 cm. 


FIG. 104. LEUCISCUS VANDOISULUS 
Standard length 6.3 cm. 


FIG 105. LEUCISCUS VANDOISULUS, imm. 
Standard length 3.8 cm. 


FIG. 106. NOTROPIS PROCNE 
Standard length 4.6 cm. 


FIGS. 103-106. FISHES FROM OXON RUN 


Zoologica Vol. II, No. 14 
Face page 304 


FIG. 107. NOTROPIS CORNUTUS 
Standard length 9.0 cm. 


FIG. 108. NOTROPIS CORNUTUS, imm. 
Standard length 5.3 cm. 


FIG. 109. RHINICHTHYS ATRONASUS 
Standard length 3.9 cm. 


FIG. 110. EXOGLOSSUM MAXILLINGUA 
Standard length 6.3 cm. 


FIGS. 107-110. FISHES FROM OXON RUN 


Zoologica Vol. II, No. 14 
Face page 305 


1922] Breder & Crawford: Food of Minnows 305 


The percentage of empty stomachs and the months in which 
they occurred, in this case also, indicates a cessation of active 
metabolism during the cold season. 


Rhinichthys atronasus (Mitchill) BLACK-NOSED DACE 


This readily recognized species required close examination 
only to prevent the possibility of confusing it with R. cateractz, 
which, however, was not taken in any of our collections. Our 
specimens varied in length from 1.6 to 5.6 em. Fowler (’08) 
found his maximum to be 314 inches, which is somewhat larger 
than ours. The smallest mature fish had a length of 3.0 cm. 
although a few above that size were immature. The two collec- 
tions showing double modes on the frequency graphs presented 
the point of greatest depression between them at approximately 
the 3.0 cm. point. However, the overlapping of the year-classes 
was so great that very little information could be deduced there- 
from. Nothing except the fact that more than one year-class 
was present could be satisfactorily determined. 


Analysis of Stomach Contents:—This species appears to be 
intermediate in feeding habits between Semotilus and Leuciscus 
on one hand and the two species of Notropis on the other, the 
vegetable and insect remains appearing as 24 and 61 per cent. 
respectively. The unidentified debris amounted to a considerable 
quantity, constituting the remaining 15 per cent. 


The number of fish which were not feeding before the time 
of capture roughly suggests a lighter feeding in the colder months. 


This species was the most heavily infested with intestinal 
parasites, although the fish seemed to show no ill effects from the 
presence of these worms since they were uniformly fat and 
healthy in appearance. In all, 22 examples out of the 257 speci- 
mens that were examined contained one or more parasites. 


The vegetable remains were found to be almost exclusively in 
an unplasmolized state. 


Specimens in the aquarium appeared to feed almost indiffer- 
ently from either the bottom or middle of the tank, and occasion- 
ally rose to the surface. 


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$]Ua}UOD YORUIOJG FO a]qeT, DNbuljpixvi wWNsso,b0x 7 IIA “ON 2927, 


1922] Breder & Crawford: Food of Minnows 307 


Judging from the relative frequency of Chetopods, Mayfly, 
and Stonefly larve, these fish feed more frequently from the bot- 
tom in the colder months, in a manner comparable to the be- 
havior of the other species. 


Exoglossum maxillingua (Le Sueur) CUT-LIPS 


This unique and striking species presented no difficulty to 
identification in any way, the peculiar three-lobed mandible being 
entirely sufficient to at once isolate the species. The lengths of 
the specimens of our series varied from 2.0 to 6.4 cm. Fowler’s 
(08) maximum or 4% inches (12.38 cm.) was about twice that 
of ours, although many of the latter were adult and in breeding 
condition. This cyprinoid has been recorded as reaching a length 
of 6.0 inches (15.24 cm.). The smallest mature fish was 4.3 cm. 
and the largest immature specimen was 5.8 cm. This excessive 
overlapping of the year-classes together with the unfortunate 


paucity of the collection precluded drawing of any conclusions as 
to age. 


Analysis of Stomach Contents:—Something decidedly differ- 
ent in the food of this species might be expected judging from the 
peculiar formation of the lips, (Figs. 110 and 116). However, 
this was not found to be so, as practically all of the food of this 
species was similar to that taken by the others. The large amount 
of debris and vegetable matter suggests that these fish were 
primarily bottom feeders. In the aquarium, this was observed 
to be the case, the specimens for the most part poking around in 
little nooks and crannies among the rocks and negotiating with 
difficulty all but the smallest particles of food. However, they 
were seen to rise occasionally to the surface as small particles of 
food were descending through the water. 


In this case, also, feeding was less heavy during the winter 
season. In those specimens which contained diatoms, the diatoms 
appeared partly digested. 


DISCUSSION 


Comparison of Foods:—Table No. VIII shows the foods of 
each species for the year side by side to facilitate comparisons. 


308 Zoologica: N. Y. Zoological Society [ii 


Table No. VIII 


Showing averages of foods taken by each species for entire collection 
(Comparison by percentage) 


a 
. oe 2a 
2 2 S 2 = = 
= S XS Rg = Ss 
FOOD S 3 < s = i 
§ Sy 5 ee me SR 
yk ae cea ee hee ~ 
Diatomsiee ascites eae ee OS ee ee 02 +- 39 25 22 08 
Rilamentoussallewyer-acnan soc ee ee + + 06 07 02 + 
Unidentitted plants nemains-a-- seen eee 02 — 02 01 + 07 
TOtAISUCG CLOUT ALLER ere er er ree 05 + 47 3)3) 24 ike} 
Unsesmentedawormsesen sae cee ie — — + 4 — -- 
Chetopodialessitc scum csterae toned teoke eeepc 01 + 13 02 04 30 
Decapoda * 5.2 :onerOate cose oh Ce Ee ee 01 + — _ —_ —_— 
Diplo podamssncaceoct occ, te ee ee 01 — —_— _— = = 
Mdult why sanuTa™ ee oat ecer et ee eis ces teooe — + —- — —_ — 
Larval -Ephemeropteras-75 s5.00stee eee 01 06 02 07 + — 
Odonata ny mphns en eyes eae ee cee — a aL ao — — 
Karvalvblecopteral seryaetnccerts coe ieee aces 02 02 01 02 05 — 
Marval eiricioprena: ec ciel ses tana nae ores — -+ — — 02 
Avdinlt Ortho pce nas wrists cic ane iene — _ — — — — 
KanvialiColeoptera saeiaecmerceoe eee ee a 02 +. + 01 — 
Adults C€oleopterwal ey-rcier: arc acter ateterisiehe ie 30 20 04 08 04 01 
larval epidopteraissat. sac ast eer ieee < 01 01 -= 02 —_— 
Crysilid@ltepidopterasse eer en eet + — — — — 
Wainy alle pte nas oar 1a senerd fe eres aeons 01 — — + — — 
AdultaDiptenatcen, str tyes cys ea tens Seite ae 06 07 -—— — + _— 
ANolalte JER RONNIE Songoasqeuconoencaancone 03 08 — 01 + — 
Wnrdentified insect -ee ote te eee eee 43 53 28 39 43 34 
EOLA AASEGL ser RC a Te &7 98 36 57 57 515) 
VAN Cate Cal Mererty Mace: caro trc ples RE oon Gio ae eee — — a al. — — 
Arte hind eresedes.co sen aioe Re Ue eee: — + — — + —_— 
Miollluscatecteacasc 5 2 sks eho a eo eee eee — - — — — — 
Beas Iae sere s roots occte <n aerate ehh a ease eee werner + — — — — 
Bhi SO Mae eed Soc se Ne a TCS oct ope ee —_ — — — — a 
Unidentificdadebris. Aoki een oe eee 05 02 04 08 152220 


The table clearly indicates that all of the species are rather in- 
sectivorous. Hymenoptera and Diptera were represented mostly 
by various minute chalcid flies and midges. Lepidoptera were 
represented by the larve of Geometride. All of the insects with 
few prominent exceptions were terrestrial species which evidently 
had fallen into the water. 


Table No. IX shows in still more epitomized form to what 
degree each species was carnivorous. The species have been ar- 
ranged in this table with the most carnivorous coming first. A 
superficial examination of this table no doubt would suggest to 


1922] Breder & Crawford: Food of Minnows 309 


Table No. IX 


Showing degree to which each species is carnivorous 
(Comparison by percentage) 


ee = = 
ria eee g. 
SPECIES 5 ie) ¢ . E 2 = 

a= bp 2 o > oN 

5 o 2. a 2°5 L 

< >a, = Os > 
ILC CRIGERTO Po 8 Ban Ob peioe heme eee 98 02 98 trace trace 02 
INEMIDOTILIES hese Tete tiem ace citin nave i 90 10 87 03 05 05 
EGOGLOSSUTT Jet cloctays cae Wis 2.0) s aie 65 35 35 30 15 20 
TUTE GIDLIEYS Maree cutter ate otal of 02 Sis adk 61 39 57 04 24 15 
IS, DGS AoA Oc ble See ee 59 41 57 02 33 08 
NE OF ATED RO ES CROCS SCT I 49 51 36 13 47 04 


the reader that N. procne was decidedly vegetarian, the others 
somewhat so, but to a lesser degree, progressing upward toward 
Leuciscus which shows only 2 per cent. of vegetable matter and 
debris combined, for the whole year. 


However, since most of the vegetable matter was found to 
have been undigested and all of these species have a short diges- 
tive tract and decidedly raptorial pharyngeal teeth, it seems un- 
likely that N. Procne or any of the other species are intentionally 
vegetarian. It must be noted, however, that in the case of N. 
procne the pharyngeal teeth present a greater grinding surface 
than in any of the other species, a few of the teeth being simply 
obliquely truncated cylinders. (Fig. 119). 


The lack of vegetable matter in Lewciscus and Semotilus is 
easily explained by assuming that they feed above the bottom, as 
was pointed out in the individual treatments, pages 301 and 295 
respectively. Also, the presence of considerable vegetation in the 
stomachs of Hxoglossum and Rhinichthys may be explained by 
their known habit of nosing around near the bottom. All consid- 
erations point respectively to such habits of these four species: 
the food, the structure of the mouth, and the habits as observed 
in the aquarium. : 


However, in the case of N. cornutus and N. procne it is an- 
other matter. The trout-like grace of N. cornutus in no way sug- 
gested a typical bottom feeding fish, although a relatively large 
amount of plant and vegetable debris was found in this species. 


310 Zoologica: N. Y. Zoological Society “thas 


Table No. X 


Specimens containing no food by months 
(Comparison by percentage) 


H tu Fh 
p> © 2 ee 

SPECIES a) So hee. S 8 aes 

ww 4 ee = vo a =) ° S ro) 

So rg Ge SS 620s * Oo We eee 

re SS = <q SS St = 1 © Zi a) 

SOROUHIG sadlec scopes s 11 66 51 1 00 S00 2000) 00 OCS 
ECUCUSGUSH a. ae 09 — 00 01 02 00 09 00 11 00 — 
OG, oy ROG? Aaa eet alka Berea 22 58 47 13 02 00 00 00 00 06 00 
INES GORIUILTUSI ie ere oe 09 — 27 10 00 O00 OO O00 00 00 800 
Rhinichthys Sei tapenoe sites 32 50 50 46 10 43 14 00 22 03 36 


EXO GLOS SIUM e eee 1005 — 55 25 10 09 0089 05) 00) Ome 


In the case of N. procne, the only suggestions of bottom feeding 
were the very slightly inferior mouth and the rather prominent 
grinding surfaces of the pharyngeal teeth. 


These considerations taken as a whole are not convincing 
that these fish are partly vegetarian. The impression left by this 
study was that most if not all of the vegetable matter was in- 
gested accidentally along with the invertebrates which are usually 
associated with diatoms and filamentous alge. The presence of 
these plants in such quantities can be explained by the fact that 
peristalsis in these fishes depends largely upon the mechanical 
action of forcing the food backwards as more food is eaten and 
consequently digestion is rather a slow process. Owing to this, 
the presence of the silicon and cellulose covered cells, of diatoms 
and alge, for a more or less protracted period would be expected. 
Furthermore even the chitonous parts of insects can be discerned 
at times even in the excrement of these fishes. 


In view of the fact that none of the vegetable matter seemed 
to have been acted upon by digestive juices, with the possible ex- 
ception of diatoms in some specimens of Exoglossum previously 
noted (page 307), it seems to be an unwarranted conclusion from 
the examination of the stomachs of these fishes that they are 
vegetarian, or derive any perceptible amount of nourishment 
from vegetation ingested. 


These fishes appear to be very adaptable to changing condi- 
tions of food and feeding. In an aquarium they take anything 


1922] Breder & Crawford: Food of Minnows all 


and everything that is given to them, living or dead, both the 
food which might possibly have been taken in their native haunts 
and that which it would have been impossible for them to secure 
under ordinary conditions, i.e., unnatural foods, such as boiled 
egg, shredded wheat, cooked meat, boiled potatoes, et cetera. 


It is easily demonstrated by means of simple experiments 
that these minnows will snap at any small particle alighting upon 
the surface of the water, or settling to the bottom. Whether or 
not the particle is ingested seems to depend upon its physical 
properties. To the fish it seemed to be immaterial whether one 
kind of food or another was presented provided it was of proper 
size. 


“Tt seems likely to be a general rule,” Forbes has written, 
(80), “that a fish makes no more than a mechanical selection 
from the particles of food accessible to it, taking almost indiffer- 
ently whatever edible things the water contains which its habitual 
range and its peculiar alimentary apparatus enables it to ap- 
propriate, and eating of these in about the ratio of their relative 
abundance and ease with which they can be appropriated at any 
time and place.” The positive identification, therefore, of the 
different constituents of the food is of small importance when 
carried beyond a certain point. 


The following tabulation seems to include all factors directly 
concerned with the act of taking food. 


EXTRA ORAL 
I. Habitat of food. 


A. Geographical distribution. 
The ranges of the food and feeder must overlap. 
This is necessary for “B”’ to be possible. 


B. Exact, or local position. 
The exact local habitats of the food and feeder must 
overlap and finally the individual feeder, and food 
unit must be in proximity. 


ale Zoologica: N. Y. Zoological Society [Tr s14 


II. Size and condition of food. 


A. Limitations. 
Upper. Too large to be swallowed and unsuitable for 
nibbling into small pieces. 
Lower. Too small, or well concealed to be perceived 
by any of the sense organs. 


INTRA ORAL 
I. Tactile and taste reactions. 


A. Tactile limitations. 
Upper. Hard, sharp, rough, unmasticatable. 
_ Lower. Soft, flocculent, “‘melting in mouth.” Such 
food is not rejected intentionally, but most of it 
usually escapes through the operculum. 


B. Taste seems to be unimportant since strongly medi- 
cated food only is rejected, and then not always. 


Table No. X showing the percentages of empty stomachs for 
the various months, which by comparison with Table No. I 
demonstrates lighter feeding for all species during the three cold- 
est months, January, February, and March. In February, only 
three of the species were represented in our collections which 
suggests hibernation or a secretive habit during the extreme cold 
weather. There appears to be no cessation of feeding in any case 
during the spawning season which in all species was indicated as 
extending from sometime in April to well into the summer. 


Little can be said of the high percentage of empty stomachs 
in the series of Rhinichthys in June since it was composed of only 
seven specimens. 


The general feeding habits of these six species appears to be 
remarkably similar even as much, or more so, than their anatom- 
ical structures. It is rather striking that so many members of 
the same family of fishes should become suitably adapted for 
survival in the same localities and still retain their specific iden- 
tity, although in many cases the young were difficult to differen- 
tiate. It seems that all fill practically the same ecological niche 
and very likely we have here a case of intra-family convergence, 
rather than divergence from a recent common prototype. 


1922] Breder & Crawford: Food of Minnows 313 


COMPARISON OF ALIMENTARY TRACTS 


On the whole, there is no vast difference between the alimen- 
tary structures of the six species under consideration. How- 
ever, the minor details vary considerably as shown by the dissec- 
fions- (Figs. 111-116). 


In making these dissections, the body wall of the left side 
was removed by cutting it through along a median line from the 
vent forward to a point just posterior to the cardiac cavity, from 
whence it was passed diagonally upward to a point dorsal of the 
pharingeals, thence following back just ventrally of the gonads 
to the vent. In this way, only the visceral cavity was laid open, 
the heart, gonads and air bladder and kidneys remaining hidden. 
Two perpendicular cuts were made; one above, and one below 
the oral cavity extending from the snout to the opened visceral 
cavity. Between these two cuts, all substance was removed.down 
to the median plane, thus showing the mouth and pharynx in true 
cross section. In making the semi-diagramatic sketches all un- 
necessary detail was omitted for the sake of clarity. Thus the 
. structures above and below the buccal cavity are indicated simply 
by conventional cross-hatching, as is likewise the pelvic girdle. 


The intestine, in all cases a simple tube containing a single 
sigmoid flexure, was spread out somewhat. That is, the forward 
arm was drawn down and the posterior one upward. By simply 
swinging them up and down respectively to a nearly horizontal 
position and allowing their distal ends to act as fulcra, the 
normal location of the parts can be found. The forward arm 
would thus lie to the right and the posterior two to the left, one 
above the other. The liver with its two lobes drops curtain-like 
to either side of the intestine. Their derangement has been 
trivial, but slightly different in most of the sketches in order to 
show the parts to their best advantage. 


The representation of all mesentary membranes and fat has 
been omitted in order that the details which the drawings were 
designed to show might not be obscured. Lettering of the parts 
also has been omitted since the shape and locations of the organs 
together with the descriptive text following is sufficient explana- 
tion. 


314 Zoologica: N. Y. Zoological Society eS ee 


FIG. 111. SEMOTILUS BULLARIS 
Standard length 12.0 em. 


BIG 2s VSB UIGLSGUS: WAND OLE: 
Standard length 6.3 cm. 


FIG. 113. NOTROPIS PROCNE 
Standard length 4.6 cm. 


FIG. 111-113. VISCERAL DISSECTIONS 


22] Breder & Crawford: Food of Minnows 315 


FIG. 114. NOTROPIS CORNUTUS 
Standard length 9.0 cm. 


FIG. 115. RHINICHTHYS ATRONASUS 
Standard length 3.9 cm. 


FIG.116. EXOGLOSSUM MAXILLINGUA 
Standard length 6.3 cm. 


FIGS. 114-116. VISCERAL DISSECTIONS 


316 Zoologica: N. Y. Zoological Society [TEs 


For those readers who are not familiar with the internal 
anatomy of this group of fishes, it should be mentioned that the 
stomach is simply the widened anterior arm of the tube extending 
from gullet to vent, while the posterior portions are considered 
as intestine. The two-lobed liver (shown by light stippling) is 
attached to the anterior end of the stomach and the spleen, in all 
cases a small body (shown by dark stippling) is attached further 
back. The gall bladder is embedded in the liver and not shown. 
The pancreas in all cases (also shown in light stippling) lies ven- 
trally to the stomach and it is bound to it with mesentary and is 
so embedded in fatty tissue that it is hard to differentiate. In all 
cases, except one, the point of attachment to the stomach wall was 
at its forward end. In Leuciscus, it was apparently attached at 
the first bend of the sigmoid flexure. However, this is a debateable 
point and in reality it may have been attached similarly to the 
others, although the drawing was made as shown only after many 
specimens were opened. 


The structure of the mouth, its inclination and general con- 
formation, are here, as in many other groups of animals, a very 
fair index to the food and feeding habits. The only species in 
which the mouth can be considered properly other than terminal 
are Exoglossum, Rhinichthys and Notropis procne. However, 
in the latter two, the mouths are very nearly terminal. The buccal 
cavity is much alike in all of them, being lined with an epithelium 
of the same general structure throughout. The most pronounced 
variation in this respect is in the odd formation of the mandible 
of Exoglossum which is so modified that it is divided into three 
lobes, the central one alone pressing firmly to the upper jaw. 
(Hig. 116): 


There is some similarity in the pharyngeal teeth in all of 
the species. The comparative drawings show that the greatest 
difference in number of teeth between any two species is three. 
Most of the teeth are decidedly raptorial, with small grinding sur- 
faces. The greatest variation is seen in N. procne in which some 
of the teeth are blunt and flat across the end with a fairly well 
developed grinding surface. 


It should be noted here that the teeth were held in various 
positions to show the shape and number of teeth to the best 


FIG. 117. SEMOTILUS BULLARIS FIG. 118. LEUCISCUS VANDIOSULUS 


Standard length 5.5 cm. Standard length 5.7 cm. 


FIG. 119. NOTROPIS PROCNE FIG. 120. NOTROPIS CORNUTUS 
Standard length 4.9 cm. Standard length 7.6 cm. 


RIG. 121; RHINICHTAYS ATRONASUS FIG. 122. EXOGLOSSUM MAXILLINGI 
Standard length 4.4 cm. Standard length 3.9 cm. 
FIGS. 117-122. PHYRANGEAL TEETH 
Diameter of black disc = 1. cm. 


Zoologica Vol. II, No. 14 
Face page 316 


FIG, 123. SEMOTILUS BULLARIS FIG. 124. LEUCISCUS VANDIOSULUS 
Standard length 4.2 cm. Standard length 4.0 ecm, 


FIG. 125. NOTROPIS PROCNE FIG. 126. NOTROPIS CORNUTUS 
Standard Jength 39 cm. Standard length 4.0 cm. 


FIG. 127. RHINICHTHYS ATRONASUS FIG. 128. EXOGLOSSUM MAXILLINGUA 
Standard length 4.0 ecm. Standard length 4.0 cm. 


FIGS. 123-128. INTESTINAL WALLS. 28X 


Zoologica Vol. II, No. 14 
Face page 317 


1922] Breder & Crawford: Food of Minnows alr 


advantage in each case.* This causes foreshortenings of the sup- 
porting pharyngeal bones in various ways, depending upon the 
angle from which they were viewed. In all cases the pharyngeal 
bones must bear the same relation to each other in order to accom- 
plish their purpose of masticating the food, but their shapes de- 
pend upon the correlated differences of other related anatomical 
parts. 

The stomach, or first arm of the digestive tract, is very simi- 
lar in all cases, being a simple, nearly straight tube capable of 
considerable distention, and normally extending to a point slightly 
past the pelvic girdle. At its anterior end, the liver and pancreas 
are attached, except in Leuciscus in which the latter appears to 
be attached to the second flexure of the tube, as previously noted, 
although when the disarranged viscera are re-assembled it seems 
probable that this organ may be attached to either or both points 
since they are appressed closely in life. In this position, what 
appears to be the free end of the pancreas is brought in apposition 
to a possible point of attachment similar to that of the organ in 
the other species. In Rhinichthys and Exoglossum the pancreas 
appears to be reduced to a small, almost indistinguishable mass. 
The liver on the left side extends forward of the digestive tract in 
life and infolds it. The spleen is practically identical in all of the 
species, except for size and shape, both of which seem to be quite 
variable in individuals. 


The differences in length of the alimentary tracts are too 
small to be used as an index of food habits. However, the two 
species of the single genus Notropis have the longest tracts. In 
the case of N. cornutus the tube has various curves and bends 
while that of N. procne is made up of three rectilinear sections. 
Semotilus and Rhinichthys follow in the order named and Leucis- 
cus and Exoglossum possess the shortest digestive tracts. The 
lengths of all are characteristic of carnivorous fishes. 


Heinrich Rathke (’24) recognized folds on the intestinal 
walls of certain European Cyprinoids similar in a general way to 
those of these species, illustrated by (Figs. 123-128). His list in- 
cludes Cyprinus gobio, C. jesus, C. aspius, C. vimba, and C. caras- 


* The accompanying wash drawings were modified from line drawings made through the 
camera lucida applied to a low power microscope. The teeth were removed carefully and 
cleansed, after which they were placed on a block of black parafin, which held them in any 
desired position and formed a contrastive background. 


ols. Zoologyca: N. Y. Zoological Society Bs 2! 


sius. Our figures of the intestinal walls of Leuciscus, N. cornutus, 
and N. procne appear to represent one type of folds which under 
low magnification give an effect somewhat similar to that of the 
herring-bone weave of cloth. These evidently are the “Zickzack 
falten” of Rathke. Rhinichthys and Exoglossum represent an- 
other type in which the folds anastomose to a great extent. These 
folds can be traced for a considerable distance along the top of 
the ridges without coming to a blind end. This is markedly dif- 
ferent from the short, straight, or curved folds of the former 
three. Semotilus represents a still different formation. Here the - 
ridges for the most part are short and heavy, and in many in- 
stances they are branched but are not anastomosing in any sense 
of the word. While these absorptive folds fall into three general 
groupings, each is distinctive and would serve as a character 
separating at least these six species. 


Each of the figures’ (123-128) represents a small piece of the 
intestinal wall taken from a point just posterior to the second 
angle in the modified sigmoid flexure of the intestinal tract. All of 
these sections were taken from as nearly similar places as pos- 
sible and from specimens of about the same size in order to make 
as close comparisons as desirable. 


Thus it is seen that the similarity of these six species extends 
to their alimentary structures and food. This fact, as previously 
pointed out, furnishes an illustration of the well-known principle 
that organisms living in the same environment and under similar 
conditions tend to develop similar habits and converge in details 
of their anatomical structures. It seems unlikely that this simi- 
larity is due to a divergence from a single ancestral type since 
these fishes are coexistant in the same locality and subject to the 
same conditions. 


FISH CULTURAL SIGNIFICANCE OF THESE CYPRINOIDS 


By study of the accompanying tables, listing the foods of the 
Six species dealt with in this paper, the conclusion is drawn that 
these species, at least while they are young, will not destroy the 


3 The photographs are not all that could be desired, since it was found to be impracticable 
to thoroughly remove the mucus without destroying the ridges. However, the usefulness of 
the illustrations is not impaired. Alcoholic specimens were used. After the material had been 
cleansed as well as possible the pieces were immersed in glycerine to clear the tissue a tritle. 
The photomicrographs were taken with the aid of an Edinger projector, owned by the Bureau 
of Fisheries. 


1922] Breder & Crawford: Food of Minnows 319 


young of important food and game fishes. However, it is pointed 
out that insects and other small animals enter into the diet of all 
small fishes to a certain extent, and in so far as they do, therefore, 
these minnows must compete with all other fishes of a similar 
size and habitat. Nevertheless, it is difficult to overestimate the 
importance of these cyprinoids to fish culture. No doubt con- 
tinued success in the cultivation of basses, crappies, and other 
species in ponds might be greater if suitable food could be pro- 
vided for the growing fish and thus check their well-known can- 
nibalistic tendencies. Similar difficulties in stocking streams with 
the various salmonoids might be overcome to a large extent by 
providing them with minnows for food. 


All of these cyprinoids are not of equal value in practice and 
due regard for their variations in habit must be taken into con- 
sideration. For instance, if a stream is found to be suitable in 
every way for the introduction of trout, except that it is deficient 
in natural food, the deficiency might be overcome by introducing 
some of the smaller species of cyprinoids, such as Exoglossum 
maxilingua, or Rhinichthys atronasus, which are found in clear, 
rather swift streams. Other species are better adapted to a lacus- 
trine habitat, such as Semotilus bullaris, or S. atromaculatus. In 
view of the fact that S. bullaris is predatory and sometimes 
reaches a length of thirteen inches and a weight of a pound or 
more, this species should be introduced with caution. 


Since very little has been recorded concerning the rate of 
growth of these cyprinoids, success in propagating them in large 
quantities for fish food is problematical. It is known that some 
of the species will spawn in aquaria and that the young when 
reared with small tropical fishes will become adapted to the warm 
water conditions necessary for the latter. This suggests a method 
of procedure in stocking a pond with minnows, but the details of 
the experiment must be worked out for each species under local 
conditions. 


There is no reasonable doubt that a pond may be stocked 
successfully with minnows provided that they are suitable. Brood 
stocks should be available in many places since these minnows 
are widely distributed and generally easily captured with a small 
seine. The suitability of a species selected for introduction will 
depend, of course, upon the similarity of its natural habitat to 


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1922] Breder & Crawford: Food of Minnows 321 


that into which it will be introduced. Success may not be obtained 
by stocking ponds with species taken from swift, clear streams, 
unless, as suggested, the young are hatched and reared under 
pond conditions, or otherwise properly acclimatized. 


It seems desirable to know, 


(1) the spawning time, habits and place, in order that compar- 
isons can be made between them and those of the species 
which will subsist on the minnows. It is obviously desirable 
to arrange conditions in such a way that the young fish under 
cultivation will find a supply of young minnows at the time 
when they are most needed; also 


(2) how prolific each species of minnows is and length of 
time required to mature. However, certain factors, such as 
good protection to the eggs, etc., might compensate for small 
numbers; and 


(3) the natural conditions under which the minnows live, so that 
these conditions may be reproduced as nearly as possible and 
thus assure a continuous supply. 


No doubt, many natural ponds which have become over-run 
with chubs or other species of minnows because of their rapid 
growth, or because they are able to subsist better on the existing 
food supply to the exclusion of more desirable species, such as 
brook trout, could be made a valuable asset to the fish culturist 
if properly manipulated. The young of all of these cyprinoids 
are excellent and acceptable food for the young of all piscivorous 
fishes and if economical ways are developed for handling them, 
another natural resource may be developed. In all cases it is pre- 
supposed that all primary factors will be under the control of the 
fish culturist. If chubs are to be introduced into ponds when 
young, they should not be left to reach a large size and their 
introduction therefore, should depend upon the ease with which 
they may be removed, i.e., small ponds may be found more effec- 
tive. Minnows seem to thrive best in streams or ponds where 
the banks are grown with overhanging vegetation which supports 
an abundance of insect life. If they are introduced into similar 
places where they will find their natural food in abundance it is 
unlikely that they will prey upon the eggs or young of other fishes. 


322 Zoologica: N. Y. Zoological Society. [ips 


NOTES ON ASSOCIATED ORGANISMS 


In the course of making these collections, as shown by Table 
No. XI, seventeen other species of fish and fish-like vertebrates 
were accidentally seined. Two of these are new to the region, 
namely Notropis whipplii (Girard), (Crawford, ’21), and Lamp- 
tera wildert (Jordan and Evermann), (Breder and Crawford, 
’22). Up to the present, regional lists of the District of Golumbia 
have shown twenty-three species of cyprinoids. With the addition 
of N. whipplii, the list is now twenty-four which shows that cypri- 
noids comprise over 25 per cent. of the fishes of this region. The 
three specimens which we took were small, being not over 5.4 em. 
in length. 

The only representative of the Petromyzonide hitherto 
known from the District is Petromyzon marinus Linneus, the 
young of which Lampetra resembles. Considerable time and ef- 
fort was consumed in establishing the status of our specimens 
which are now deposited in the American Museum of Natural 
History. 

One of the specimens of Erycimba buccata presented the 
only decided abnormality taken in the entire collection, in that 
it was a fair example of the pugheaded condition which is quite 
common among other fishes. A few other minor abnormalities 
were noted, such as notches in the operculum and small irregular- 
ities in the fins. As a whole, however, the collection consisted of 
typically normal individuals. 


The eels taken were both small, while two others were seen 
which we judged to be about 30.0 and 45.0 cm. in length respec- 
tively. 

Previous to these collections, in September, 1919, several 
small specimens of Schilbeodes insignia were taken at this point 
and in February, 1921, a larger one was netted a short distance 
down stream. These records suggest that the number of species 
occasionally to be found at this collecting site may be considerably 
greater than our list shows, although collections were consistently 
made all through one year as well as scattering ones before, after 
and between the regular dates listed. 


When all other organisms living associated with these fish, 
both vertebrate and invertebrate, are considered the aggregate 


1922] Breder & Crawford: Food of Minnows 320 


number is rather surprising. Small aquatic salamanders were 
found frequently, as well as frogs and their tadpoles. Land 
turtles visited the place at least occasionally. Kingfishers were 
seen frequently about and they no doubt preyed upon the fishes 
of this stream almost entirely. They were, no doubt, summer resi- 
dents nesting in some sand or clay bank nearby of which there 
were plenty of suitable size and location. A drowned mole was 
noted and various bird and mammalian tracks furnished evidence 
of the presence of these warm-blooded creatures of sizes which 
varied from that of the song sparrow to man. 


Among the invertebrates the crayfish and insects were the 
most prominent. A representative list of the latter is to be found 
among the list of foods in Table No. VIII. Many gomphid 
nymphs were present although but few of our cyprinoids were 
large enough to negotiate such formidable and sizeable insects. 
Practically no snails were present although a single specimen of 
Physa was taken from the stomach of a Boleosoma. Aquatic 
vegetation was entirely absent except for the very minute and 
ubiquitous representatives of alge, none of which appeared to be 
particularly plentiful. The nearest approach to the larger 
aquatics was a very sparse growth of the semi-aquatic Ludwegia, 
which appeared to be gaining a foothold in the little cove just up 
stream of the beach on which the seine was hauled out. None was 
noted either above or below our collecting site. Fair growths of 
Fontinalis were noted beyond the limits of the map and seasonal 
fluctuations in the growth of this plant were observed. 


SUMMARY. 


1. .The six species of cyprinoids treated in this paper are chiefly 
insectivorous. 

2. Vegetable matter ranks second in volume of materials swal- 
lowed, but this material appears to have little or no nutritive 
value since the fish do not seem to be able to digest it. 

3. The amount of food taken in the winter months is relatively 
small. 

4. The seasonal abundance of the more prominent items of 
diet is reflected in the food of the six species. 

5. No correlation exists between the size of the fish and the food 
taken except that which is entirely mechanical. 


324 


10. 


aha 


Zoologica: N. Y. Zoological Society . eG fae le! 


There is apparently no cessation of feeding during the breed- 
ing season. 

A seasonal change from pelagic to benthotic feeding is shown 
more or less strongly by all species as the cold weather ap- 
proaches. 

The fish of this locality appear to be exceptionally free of 
parasites and abnormalities. 

The alimentary structures are much alike in the various 
species. 

These fishes are recorded as the food of many important food 
and game fishes. 

It is believed that, if properly propagated, some of the species 
discussed in this paper would be of great value to fish cul- 
turists as food for other fishes. 


BIBLIOGRAPHY 


BEAN, B. A., AND WEED, A. C. 


1911 Recent Additions to the Fish Fauna of the District of Columbia. 
meee, Wash. Bio. Club. June 16, 1911. Vol. XXIV, pp. 
lear 


BREDER, C. M., JR. 


1920 a. Some notes on Leuciscus vandoisulus Cuv. & Val. Copeia 
No. 82. May 20, 1920. New York. pp. 35-38. Fig. 1. 
b. Further notes on Leuciscus vandoisulus Cuv. & Val. Ibid. 
No. 87. Oct. 15, 1920. pp. 87-90. Fig. 1. 


BREDER, C. M., JR., AND CRAWFORD, D. R. 


1922 Lampetra wilderi Jordan & Evermann in the District of Colum- 
bia. Copeia No. 108. Feb. 15, 1922. pp. 11-13. 


CLEMENS, W. A. 


1917 An Ecological Study of the Mayfly Chirontentes. University of 
Toronto Studies. Bio. Series, No. 17. pp. 1-43. 5 plates. 


CoKER, R. E. 


1918 Principles and Problems of Fish Culture in Ponds. The Scientific 
ey Vol. 7, No. 2. August, 1918. pp. 120-129. 2 
plates. 


CRAWFORD, D. R. 


1921 <A Record of Notropis whipplii from the District of Columbia. 
Copeia No. 101. December 20, 1921. New York. pp. 87-89. 
Fig. 1. 


CONTRIBUTIONS TO CANADIAN BioLocy. Fasc. II. 


1915 Freshwater Fish and Lake Biology (various authors). Ses- 
sional Paper No. 396. Ottawa. pp. 222. 21 plates. 


DAVENPORT, C. B. 


i899 Statistical Methods. Special reference to Biological Variation. 
pp. 220. John Wiley & Sons, Publishers, London. 


DycHE, L. L. 


1910-1914 Ponds, Pond Fish and Pond-fish Culture. State Depart- 
ment Fish and Game. Kansas. 3 parts. 


ForsBEs, S. A. 
1878 The Food of Illinois Fishes. Bull. Ill. State Lab. Nat. Hist. 
pp. 81-89. 
1880 a. The Food of Fishes. Ibid. pp. 18-65. 
b. On the Food of Young Fishes. Ibid. pp. 66-79. 
1883 The Food of the Smaller Fresh Water Fishes. Ibid. pp. 65-94. 
1888 a. Studies of the Food of Fresh Water Fishes. Ibid. Vol. II. 
Art. VII. pp. 433-473. 
b. On the Food Relations of Fresh Water Fishes; a Summary and 
Dicussion. Ibid. Art. VIII. pp. 475-538. 


326 Zoologica: N. Y. Zoological Society fits t4 


FORBES, S. A., AND RICHARDSON, R. E. 
1908 The Fishes of Illinois. Illinois State Lab. 


FowL_er, H. W. 
1908 <A Synopsis of the Cyprinide of Pennsylvania. Proc. Acad, Nat. 
Sci. Philadelphia. December, 1908. pp. 517-553. Plate 
XXVII. 
HAGEN, H. A. 


1861 Synopsis of Neuroptera of North America. Smith. Mise. Coll. 
Vol. IV. Art. I. xx plus 347 pp. 


HANKINSON, T. L. 


1920 Occasional Papers of the Museum of Zoology. Univ. Mich., Ann 
Arbor. No. 89. July 15, 1920. 


KENDALL, W. C. 
1913 Fishes and Fishing in Sunapee Lake. B. of F. Doc. 783. pp. 21. 


MARSHALL, W. S., AND GILBERT, N. C. 
1905 Notes on the Food and Parasites of Some Fresh Water Fishes 
from Lakes at Madison, Wisconsin. Rept. U. S. Com. Fish. 
1904. Appen., pp. 513-522. 


McATEE, W. L. 


1918 A Sketch of the Natural History of the District of Columbia. 
Bull. Bio. Soc. Wash. No. 1. 


MooreE, EMMILINE, Ph.D. 
1915 The Potomogetons in Relation to Pond Culture. Bull. U. S 
Bok. Vol. XXX, 4913; “ppy 251-292: 
1920 Some Plants of Importance in Pond-fish Culture. Appen. iv to 
Rep. U. S. Com. Fish, 1919. Doc. 881. 


MorGan, ANNA H. 
1913 A Contribution to the Biology of Mayflies. Annals of the Ento- 
mological Society of America. Vol. 6, pp. 371-413. 13 plates. 
Columbus, Ohio. . 


NEEDHAM, JAMES G. 
1905 The Mayflies and Midges. In Mayflies and Midges of New York. 
Third Report on Aquatic Insects. By J. G. Needham, 
Kenneth Morton and O. A. Johannsen. N. Y. State Mus. 
Bull. No. 86, pp. 17-36. Albany. 


NEEDHAM JAMES G., AND BETTEN, C. 


1901 Bull. 47, N. Y. State Mus. September. Univ. of State of 
New York. 


NEEDHAM, JAMES S., AND LLoyp, J. T. 
1916 The Life of Inland Waters. pp. 412. Comstock Co., Ithaca, 
N. Y., publishers. 


1920 Burrowing Mayflies of Our Larger Lakes and Streams. Bull. 
U, Se Bes a Vol exexvile Palo nanos: 


PEARSE, A. S. 
1915 On the Food of the Small Shore Fishes in the Waters Near 


Madison, Wisconsin. Bull. Wis. Nat. Hist. Soe. Vol. XIII. 
Noni pps f-22: 


1922] Breder & Crawford: Food of Minnows 327 


1918 The Food of the Shore Fishes of Certain Wisconsin Lakes. 
Bull. U.S. B. F. Vol. XXXV for 1915-16. pp. 245-292. 


1919 Habits of the Black Crappie in inland lakes of Wisconsin. Re- 
port U. S. Commissioner Fisheries, 1918, Appendix III. 
(Suggestions on technical methods.) 
PEARSE, A. S., AND ACHTENBERG, HENRIETTA 
1920 Habits of the Yellow Perch in Wisconsin Lakes. Bull. U. S. 
B. F. Vol. XXXVI. 1917-18. (Research methods.) 
RATHKE, HEINRICH 
1824 Uber den Darmkanal und die Zeugungsorgane der Fische, in 
Schriften der Naturfarschenden Gesellschaft in Danzig. 
REIGHARD, JACOB 


1910 Methods of Studying the Habits of Fishes with an Account of 
the Breeding Habits of the Horned Dace. Bull. U.S. B. F. 
Vol. XXVIII. pp. 1112-1136. 


1915 An Ecological Reconnaissance of the Fishes of Douglas Lake, 
Sheboygan County, Michigan, in Mid-summer. Bull. U. S. 
Be EB. Vol. XXOMITT, 19135 pp. 215-249. 
SHELFORD, V. E. 


1911 Ecological Succession III. A Reconnaissance of Its Causes in 
Ponds, With Particular Reference to Fish. Bio. Bull. Mar. 
Bio. Lab. No. 22, pp. 1-38. Also I and II. 


1914 Suggestions as to Indices of the Suitability of Bodies of Water 
for Fishes. Trans. Amer. Fish Soc. Vol. XLIV. No. 1, 
pp. 27-32. 
SmituH, H. M., AND BEAN, B. A. 
1899 List of Fishes Known to Inhabit the Waters of the District of 
Columbia and Vicinity. Bull. U. S. B. F. 1898. pp. 
179-187. 
Warp, H. B., AND WHIPPLE, G. C. 
1918 Fresh Water Biology. p. 1111. J. Whiley & Son, Publishers, 
London. 
WELCH, PAUL 
1912 Insect Life of Ponds and Stream. Part II. Nature Study 
Review. VIII. pp. 181-198. 
WILSON, C. B. 


1920 Dragonflies and Damselflies in Relation to Pond-fish Culture, 
With a List of Those Near Fairport, Iowa. Bull. U. S. 
Be Vol) XXXVI,” 1917-18. 


Besides the works listed, a number of lesser importance were 
consulted. They were omitted from the list because of their age 
and because they have been superceded by more general works 
or because they seemed less pertinent to the discussion of the food 
and feeding habits of the cyprinoids studied. References to other 
phases of such studies as the breeding and spawning habits, etc., 
may be found by consulting the various bibliographies appended 
to the different works cited in the foregoing list. 


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Corrected to July, 1922 


) 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 15 


THE FISHES OF SANDY HOOK BAY 


By 
C. M. BREDER, JR. 


New York Aquarium 


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Volume II, Number 15 


THE FISHES OF SANDY HOOK BAY 


By C. M. BREDER, JR. 


New York Aquarium 


The work of collecting local marine fishes undertaken by the 
New York Aquarium’s wellboat Seahorse for the exhibits of that 
institution has made it possible for the writer to compile an an- 
notated list of the fishes of Sandy Hook Bay during the summer 
of 1921, which is herewith presented. Much of the material was 
gathered by visiting the various pound nets at times when the 
owners were removing their catches. There were thirty-five such 
traps within the limits of the Bay, and as the accompanying map 
shows, they were well scattered over the area, in this way inter- 
cepting at least a few individuals of practically all species enter- 
ing the Bay which might be taken in such fishing gear. Late in 
the season when fykes were set they were likewise visited. They 
were six in number and located along the shore between Port 
Monmouth and Atlantic Highlands as indicated on the map. A 
seine, three hundred feet in length, was frequently used on the 
Beach of the Government Reservation at Sandy Hook, while less 
often smaller ones were dragged in tide pools and back waters. 
The shores along which the seines were operated are so marked 
on the map. Spermaceti Cove was seined only in 1920 as it was 
found to contain less material than numerous other places, which 
were more accessible. The beach at Atlantic Highlands was tried 
but once during 1921 when it was found to be rather unsatisfac- 
tory on account of the many submerged snags there present, 
upon which the net continually caught. 


The first records of 1921 were made on June 1 ae the last 
on October 21, which dates mark the beginning and end of the 


Bye Zoologica: N. Y. Zoological Society [ths 


boat’s activity for the year. Between these dates records were 
made on fifteen trips varying in length from two to five days, 
the writer accompanying the boat on all except three. On these 
Mr. S. A. Callisen, of the New York Aquarium, was kind enough 
to record and report such notes as were of particular interest. 
Credit is also due him for giving various kinds of assistance in 
the compilation of this list. 


Records relative to 1920 were made by Dr. C. H Townsend, 
Director of the Aquarium, who has permitted the use of his log 
book for that year. His notes extend from June 1 to November 
8 and consist of data taken on twenty-four trips of from two to 
four days duration. As the trips were approximately equidistant 
throughout: both seasons the collections and observations so made 
give avery fair idea of the fishes to be encountered during the sum- 
mer in these waters. It is unfortunate that it was impossible to 
make trips before and after the dates mentioned as the seasonal 
aspect of the Bay for an entire year would be particularly valu- 
able in regard to the study of the migrations of certain forms. 
However, it is doubtful if many species not mentioned in the list 
are present in any considerable numbers in winter. 


The kindness of Mr. J. T. Nichols, of the American Museum 
of Natural History, in allowing a perusal of his scrap book of 
records and notes on local fishes leaves little doubt as to the au- 
thenticity of the early and late dates recorded in this list. 


The most recent regional paper on the fishes of New Jersey 
is “A List of the Fishes of New Jersey” by Henry W. Fowler, 
Proceedings of the Biological Society of Washington, Vol. 38, 
pp. 139-170, Dec. 30, 1920. This publication gives, under specific 
captions, the region in which each species is found, by counties. 
It was rather surprising to find that seventeen species included 
in the present list have not been recorded,' according to Fowler, 
from Sandy Hook Bay or indeed from any of the waters of Mon- 


1A short note on thirteen species found in 1920 has been published in 
Copeia, No. 91, Feb. 15, 1921, by C. H. Townsend and J. T. Nichols and a 
non-annotated list of sixty species taken the same year is given in 
“The Twenty-fifth Annual Report” of the New York Zoological Society, 
July, 1921. The substance of the former and all species included in the 
latter have been incorporated in this paper for convenience in reference. 


1322] Breder: Fishes of Sandy Hook Bay 300 


mouth County.* When it is considered that Monmouth County 
has a long coast line on the open sea as well as on small bays, and 
that it is to the entire county that Fowler refers, the actual lack 
of accurate knowledge concerning the distribution of the fishes of 
this region is at once apparent. Notice to Fowler’s omissions is 
given under the respective specific headings. Unfortunately the 
available regional lists of fishes found near New York City con- 
tain little or no information concerning exact geographical loca- 
tions and are therefore practically impossible to compare. 


Fowler, in addition, records about forty other marine species 
from Monmouth County, not in the present tabulation, but as no 
definite section is indicated, it is not known exactly how many 
have been taken in the Bay. This rather formidable looking list 
of names is made up, however, chiefly of pelagic and southern 
fishes of which comparatively few are likely to find their way into 
this body of water. While the writer’s enumeration in no way 
pretends to be complete and is admittedly preliminary in nature, 
it is hoped that it will form an added step to a better knowledge 
of New Jersey fishes in which direction Mr. Fowler already has 
made such splendid advances. 


The vernacular name following the technical one is, in each 
case, the local appellation known to be in actual use by the fisher- 
men engaged in procuring their livelihood from this body of 
water, except in the cases of those few for which no local name 
appeared to be in current usage. In these instances the common 
names applied to the species in question in adjacent territories 
or proposed by other writers are given in brackets. All measure- 
ments given are standard lengths except where otherwise stated 
and in the case of the sharks where the total length is understood. 
The metric system is used throughout with the approximate Eng- 
lish measure following. The annotation following each specific 
heading is separated into two parts according to the year in which 
the data was collected, so that the fauna of the Bay may be com- 
pared for two consecutive years and in this way be of aid to stu- 
dents of problems to whom such data might be of service. It is to 


*Monmouth County has a coast line extending from a point a few 
miles south of the mouth of the Raritan River, on Raritan Bay to the mouth 
of the Manasquan River on the seaward coast. 


334 Zoologica: N. Y. Zoological Society [iis a5 


be understood that the specimens were taken in pound nets if no 
other type of gear is mentioned. 


GALEIDA 
1—Mustelus canis (Mitchill) DOGFISH, DOGGIE 


Common practically all season during 1920. In the warmest 
weather few were taken except in the pounds set in the deepest 
water near the point of Sandy Hook. During 1921 this species 
was few in number until October 10, after which time it was 
abundant, varying in length from about 40 to 102 em. (1614 to 40 
inches) and in weight from about .20 to 3.20 kilograms (14 to 7 
pounds). A single specimen was taken in a seine near the point 
of the Hook. On June 1, several copepods were noted on some 
examples. 


2—Carcharhinus milberti (Miiller and Henle) 
SHARK, SAND SHARK, GRAY SHARK 


Some examples upward of 1 meter (about 3 feet), as well 
as smaller ones were taken in 1920 from July 15 to September 23. 
In 1921 one small example was taken on each of the following 
dates: July 8, 15, 27, September 19, and October 19. This last 
is a late date for the species, being over a month later than any 
previous record from this region. One specimen of larger size 
was taken in mid-July. Fowler gives a single record of this 
species, which he admits as being a questionable identification, 
from Monmouth County as being made at Perth Amboy, which 
locality actually is in Middlesex. 


SPHYRNIDZ 
38—Sphyrna zygena (Linneus) HAM MERHEAD 


Small examples, less than 60 cm. (about 2 feet), were seen 
during both years in mid-summer. Fairly common. None seen 
before July 14 or after August 26, during 1921. 


CARCHARIIDA 
4—Carcharias taurus Rafinesque SHARK, SAND SHARK 


Thirty or more individuals seen between June 13 and October 
4 in 1920, varying in length from about 60 to 214 cm. (2 to 7 


19227] Breder: Fishes of Sandy Hook Bay 335 


feet). One about 121 cm. (4 feet) long was taken on July 28, 
1921, and what was apparently another about 153 cm. (5 feet) 
was seen basking about five miles south of Coney Island on Sep- 
tember 29, which position, however, is really outside the confines 
of the Bay. 


SQUALIDZ 
5—Squalus acanthias Linneus SPINED DoG, THORNED DOGFISH 


Taken late in fall of 1920 up to November 8. Many small 
ones, mostly under 30 cm. (about 1 foot) taken between October 
Wieand=215°1921.. 


RAJIDZA 
6—Raja erinacea Mitchill SKATE 


Taken on June 1 of both years, and on October 13 and later 
in 1920. A single example was taken on October 10, 1921. Ap- 
parently absent from these waters during the summer months. 


7—Raja eglanteria Bosc SKATE 


Fairly common during season of 1920 except in September. 
During 1921 quite common from June 1 to July 29, and from 
October 10 to 21. This species, while generally more common 
than the preceding, tends likewise to absent itself from this local- 
ity in the warmest weather. October 21 is a late date for this 
Skate. 


8—Raja stabuliformis Garman BARNDOOR SKATE 


Taken in November, 1920. One example about three feet in 
length of disc taken in October, 1921. 


DASYATIDZA 
9—Dasyatis centrura (Mitchill) STINGER, STINGAREE 
Two were taken between June 20 and September 23, 1920. 
In 1921 a single example was taken on September 16 and another 


on September 20, both with a length of disc less than 45 cm. 
(about 18 inches). 


336 Zoologica: N. Y. Zoological Society A Be 5) 


MYLIOBATIDA 
10—Rhinoptera bonasus (Mitchill) STINGER, STINGEREE . 


A few taken between June 13 and September 11, 1920. One 
example with a disc of more than 60 cm. (about 2 feet) in length, 
taken on August 4, 1921. 


ACIPENSERIDA 
11—Acipenser sturio Linneus STURGEON 


Fairly common in fall of 1920. One example 45 cm. (about 
18 inches) long was taken on September 15, 1921. Fishermen 
reported having taken a few others about this time. 


12—Acipenser brevirostrum Le Sueur STURGEON 


A few small examples were taken in the fall of 1920. Fowler 
makes no mention of this species being taken in Monmouth 
County. 


ELOPIDA 
13—Elops saurus Linneus SALMON 


One example 20 cm. (about 8 inches) long was taken on 
October 12, 1921. Fishermen spoke early in the season of tak- 
ing “Salmon” but the identity of the fish they had in mind could 
not be determined from their descriptions. Very likely they re- 
ferred to this species. Cape May is the only New Jersey county 
Fowler records it from. 


CLUPEIDZA 
14—Etrumeus teres (De Kay) [ROUND HERRING] 


One example of 37 cm. (about 15 inches) was taken in a fyke 
on September 20, 1921. The fishermen admitted to not having 
seen this species before and had no name for it. According to 
Fowler it has not been recorded from Monmouth County before. 


15—Clupea harengus Linneus HERRING 


A few medium sized examples were taken from June 1 to 28, 
and one each on July 7 and October 17, 1921. This species is 
known to be irregular near here, but these observations might be 
taken to indicate that possibly the Herring is absent from this 


1922) Breder: Fishes of Sandy Hook Bay aot 


Bay in summer. Fowler does not record it as being found in the 
waters of this county. 


16—Pomolobus pseudoharengus (Wilson) ALEWIFE, SHADINE 


One example of medium size taken on June 1, 1921. Prob- 
ably a few of this and the following two species were present 
later, but as it was not always possible to examine each catch in 
the detail necessary to identify such inconspicuous fish in a boat 
load of Menhaden it can not be said with certainty. 


17—Pomolobus xstivalis (Mitchill) SHADINE 


From June 1 to July 7, 1921, a few medium sized examples 
were taken. 


18—Alosa sapidissima (Wilson) SHAD 


Several were taken in 1920 and a few on June 1, 1921. 
19—Opisthonema oglinum (Le Sueur) 
THREAD HERRING, SAW-BELLY 


A few were taken in mid-summer in 1920. Medium sized 
examples were taken from July 7 to 29 and one on October 21, 
1921. Very likely they were absent in the interim. Fowler 
records this species only from Cape May and Atlantic Counties. 


20—Brevoortia tyrannus (Latrobe) 
BUNKER, BANKER, MOSSBUNKER, MENHADEN 


Taken at all times during both seasons. Both young and 
adults abundant, and taken in all gear. As the season of 1921 
came to a close their numbers became noticeably less, especially 
after the middle of September, at which time the Weakfish re- 
placed them to a certain extent. This species forms by far the 
major part of the pound netters’ catch, sometimes to the exclu- 
sion of almost everything else. On July 28 a skiff was taken 
up the ‘‘Creek” that has its mouth at the ‘‘Horseshoe” on Sandy 
Hook, and many young Menhaden 10 to 13 cm. (about 4 to 5 
inches) long, were seen as far up as it was possible to force the 
skiff, which was almost a mile, measuring along the bank. This 
“Creek” is apparently purely salt water, at present, at least, being 


338 Zoologica: N. Y. Zoological Society St ee 


merely an arm of the Bay. On July 27 and 28 countless numbers 
of these fish of a similar size were seen in great schools from 
the pier at Atlantic Highlands. At this point a large sewer 
empties just below the surface and the water is consequently 
heavily charged with various kinds of evil smelling debris. These 
young fish were in the thick of it, stemming the rather forceful 
current which the flow from the sewer pipe caused. It appeared 
that they were feeding, either on the finely divided sewage itself, 
or else on some small organism drawn there by the large amount of 
decaying organic matter, although it hardly seems possible that 
a small pelagic invertebrate, minute enough for these fish to 
engulf could sustain itself in the strong current urging the debris 
from the pipe. 


About the middle of September several decidedly greenish 
adults were seen which the fishermen say always appear at this 
season and which they dub “Irish Bunkers.” From a casual ex- 
amination it was seen that they had fed on some green substance 
‘and that the internal organs were likewise suffused with the same 
color. 


ENGRAULIDIDZ 
21—-Stolephorus brownii (Gmelin) [STRIPED ANCHOVY ] 


One example was taken on June 7, 1921, in company with 
the following species. Not previously recorded from Monmouth 
County according to Fowler. 


22—-Stolephorus mitchilli (Cuvier and Valenciennes) 
[COMMON ANCHovy ] 
Taken in small numbers on June 7, 29, September 22, October 
18 and 21, 1921. Probably present all summer but passing 
through the meshes of most gear. Large schools of anchovy-like 
fishes were seen which were very likely composed of this species 
with a few individuals of the preceding, in proportion to their 
relative abundance. 


ANGUILLIDA 
23—Anguilla rostrata (Le Sueur) EEL 


Taken uniformly on practically all trips and in all gear. 
Both large and small examples were seen. 


1922] Breder: Fishes of Sandy Hook Bay 339 


PQ@CILIIDA: 
24—Fundulus majalis (Walbaum) KILLY 
Taken whenever fished for with small seines in back waters, 
tide pools, and runs. Especially common in the “Horseshoe.” 
Ripe males and females taken June 1 to 23, 1921. In company 
with the following species but not as abundant. 


25—Fundulus heteroclitus macrolepidotus (Walbaum) KILLY 


Abundant, especially in back waters and tide pools, particu- 
larly so in “Dredged Pool.” Taken whenever such places were 
seined. Ripe examples of both sexes were taken June 1 to 23, 
1O2t 


26—Cyprinodon variegatus Lacépede KILLY 


Taken only in the large and deep ‘“‘Dredged Pool” in company 
with the two preceding species. Males with brilliant coloring 
seen from June 1 to 23, 1921. Apparently breeding at this time 
as were the other Peciliide. Specimens apparently spent were 
taken on July 14 and 22, 1921. 


BELONIDA 
27—Tylosurus marinus (Walbaum) BILLFISH 
Taken in the late fall of 1920. One example 45 cm. (about 
18 inches) long was taken in seine on September 27, 1921 and 


another in a pound net on October 14, the latter being 60 cm. 
(about 2 feet) long. 


GASTEROSTEIDA 

' 28—Aneltes quadracus (Mitchill) STICKLEBACK 
One small ripe female taken in back water of the ‘‘Horse- 

shoe” on June 1, 1921, and a few, of which the condition was not 

noted, on July 20 in ‘“‘Dredged Pool.” Fowler gives this species 

as being unrecorded from Monmouth County. 


SYNGNATHIDA 
29—Syngnathus fuscus Storer PIPEFISH 


Taken in mid-summer of 1920, especially abundant in Sper- 
maceti Cove. Taken between June 23 and October 12, 1921. 


340 Zoologica: N. Y. Zoological Society BMS 


Common near the point of Sandy Hook and in the “Horseshoe” 
in seines. Many males with brood pouches full were observed in 
mid-summer of this year. At times while riding at anchor in 
calm weather these fish could be seen swimming or drifting past, 
a few inches below the surface of the water. On June 21 a post 
larval example of 13 mm. (about 14 inch) was taken in a bolting 
cloth tow net at the surface near the mouth of the bay. 


30—Hippocampus hudsonius De Kay SEAHORSE, HORSEFISH 


One specimen taken during 1920. On August 24 and 26, 
1921 each, one fair sized example was picked off the leaders of 
the pound nets by the fishermen. Several others were reported 
about this time. 


ATHERINIDA 
31—Menidia menidia notata (Mitchill) SPEERING, WHITEBAIT 


Seined in mid-summer in 1920. Taken in seines from June 
21 to September 22, 1921. Very young fry and ripe adults were 
seen in early part of season, while larger fry and spent adults 
were seen later. 


MUGILIDA 
32—Mugil cephalus Linnzus MULLET, LEAPING MULLET 


Common in fall of 1920. A few small examples were taken 
on June 29, many large ones from July 20 to October 21 during 
1921, chiefly in seines and fykes. The adults were exceptionally 
well nourished and fat. Fowler records this species only from - 
Atlantic and Cape May Counties. 


SPHYRAINIDA® 
33—Sphyrxna borealis De Kay BARRACUDA 


On June 20, 1920, a few very small examples were taken. 
Three specimens were taken on October 17, 1921 in the seine, two 
of which measured 1614 and 17 cm. (614 and 634, inches) respec- 
tively. Several others of about the same size passed through the 
meshes at this time. 


1922] Breder: Fishes of Sandy Hook Bay 341 


AMMODYTID2 

34—Ammodytes americanus De Kay [SAND LANCE, SAND EEL] 

A single example, which was probably originally about 5 cm. 
(2 inches) in length, was taken from the stomach of a small blue- 
fish caught on June 29, 1921. However, it is possible that it was 
captured at some point outside the Bay. The reason why this 
common species, abundant at other points close by should be so 
scarce in this Bay is not clear, although no doubt later in the 
fall they appear in numbers. 


SERRANIDZ 
35—Roccus lineatus (Bloch) STRIPED BASS 


Fairly common in fall of 1920. One good sized example was 
taken in seine on June 22, 1921, which was blind, another in a 
pound net on September 19, and several small and medium ones 
were taken in fykes between October 17 and 21. 


36—Morone americana (Gmelin) WHITE PERCH 


Common in fall of 1920. Taken in fykes and pounds. A 
few medium sized examples were taken in fykes between October 
MWieand 21 1921: 


37—Centropristes striatus (Linnzus) SEA BASS 


Taken in late fall of 1920, but not common. After their first 
appearance on September 28, 1921, this species increased in 
numbers until the close of the season, at which time they were 
abundant. 


LOBOTIDA 
38—Lobotes surinamensis (Bloch) TRIPLE-TAIL, FLASHER 


One large, very dark example was taken in one of the outer- 
most pound nets on July 13, 1921. 


HAIMULIDA 
39—Orthopristes chrysopterus (Linnzus) PIGFISH 


A single individual was taken during 1920. 


342 Zoologica: N. Y. Zoological Society : Geils 


SPARIDA 
40—Stenotomus chrysops (Linnzus) PorGy, SCUP 


A few adults were taken during 1920. Common from June 
1 to 23, 1921, males with running milt; spent fish later, July 14 
to October 21, decreasing in numbers as the season drew to a 
close. From October 10 to.21 many small ones of about 3 cm. 
(114, inches) in length were taken in seines and fykes. 


41—Lagodon rhomboides (Linneus) SAILOR’S CHOICE 


One example was taken during 1920. 


KYPHOSIDA 
42K yphosus sectatrix (Linneus) BREAM 


One example was taken in 1920. Fowler’s list fails to men- 
tion this fish as being known from New Jersey waters at all. 


SCIZANIDA 


43—Cynoscion regalis (Bloch and Schneider) 
WEAKFISH, BLACKTAIL 


Present at all times during both years, increasing in numbers 
as the menhaden decreased. Sometimes taken in fykes as well 
as pounds. Many silvery below, while others were more or less 
golden. The fishermen believe the latter to be visitors from out- 
side waters, while the former are supposed to be residents of the 
Bay. There was no opportunity to gather accurate data as to the 
actual cause of the two phases appearing simultaneously, how- 
ever. Several were examined on June 29, 1921, and a number 
found to be ripe. Their stomachs contained squid, prawns and 
the remains of small fish. One example contained what appeared 
to be a small Batrdiella chrysura which in turn held a specimen 
of Stolephorus sp. A few examined on September 14 had fed on 
menhaden. These weakfish varied in length from 32.5 to 50.5 cm. 
(13 to 20 inches) and contained fish about 11.5 em. (414 inches) 
long. In most cases three such young menhaden were found in a 
stomach. 


1922] Breder: Fishes of Sandy Hook Bay 343 


44 Bairdiella chrysura (Lacépéde) WHITE PERCH 


One medium sized example was taken on June 28 and an- 
other on September 14, 1921. No distinction is made by the 
fishermen between this and Morone americana. 


45—Scizxnops ocellatus (Linneus) CHANNEL BASS, RED DRUM 


One example 82.5 cm. (2 feet 814 inches) in length was 
taken on September 13, 1921. 


46—Leiostomus xanthurus Lacépéde SPOT, LAFAYETTE 


Common all season in 1920. Present sparingly all summer 
in 1921, most plentiful in September, falling off in numbers on 
either side of that month. Much less common than in the pre- 
ceding year. 


47—Micropogon undulatus (Linneus) CROAKER 


Common during 1920, being taken at times by the boat load 
between June 18 and October 22. One or a few examples were 
taken on each of the following dates in 1921: June 28, July 7, 28, 
August 25, September 14, 28, and October 20. Some were of a 
fair size while some were not over 15 cm. (about 6 inches) in 
length. 


48—Menticirrhus saxatilis (Bloch and Schneider) KINGFISH 


Only small examples were taken both years, averaging about 
15 cm. (6 inches) in length in fykes and seines. Fairly common 
from July 28 to October 21, 1921. 


49—Pogonias cromis (Linneus) DRUMFISH, BLACK DRUM 


Two examples were taken in September, 1920. One was 
about 914 cm. (3 feet) and the other about 1219 cm. (4 feet) in 
length. 


POMATOMIDA® 
50—Pomatomus saltatrix (Linnzus) 
BLUEFISH (Young—SNAPPER) 


Fairly common in 1920. Taken in 1921 from June 21 to 
October 21. The adults became more common as the season wore 


344 Zoologica: N. Y. Zoological Society - LIT: 15 


on, with a drop in numbers, however, near the end. The young 
were common, being taken in seines and fykes as well as the 
pounds. By October 11 most of the fish hatched early in the 
spring had reached a length averaging 15 cm. (about 6 inches), 
which was well established by a comparison of the serial collec- 
tions, showing this species to have an extremely rapid rate of 
growth. 


RACHYCENTRID2& 
51—Rachycentron canadum (Linnezus) CRAB-EATER 


One large example was taken in 1920. 


STROMATEIDA® 
52—Peprilus paru (Linnzus) HARVEST FISH, BUTTERFISH 


Taken in mid-summer in 1920. Adults were fairly common 
from June 28 to September 17, 1921. Fowler fails to mention 
this species at all in his list. Apparently an oversight, as it is 
mentioned in a paper’ of his of 1906, in which he writes “It is 
known only from the record of Dr. Abbott. It is properly a native 
of southern waters and can only be said to be a straggler on our 
shores.” Nichols, writing of the fishes within fifty miles of New 
York City says “The Harvestfish is occasionally common in 
summer - - - .” The experience of the writer has certainly 
confirmed the latter statement and it seems remarkable that as 
late as 1906 only one record of this species had found its way into 
regional literature. 


538—Poronotus triacanthus (Peck) BUTTERFISH 


Very common during both seasons. Abundant in fall of 
1920. Most numerous during the early part of the season of 1921, 
their numbers falling off prominently after September 28. Adults 
and juveniles were seen in approximately equal numbers from 
June 7, which is an early date for the species, to October 21, 1921. 


*1906—The Fishes of New Jersey, by Henry W. Fowler. Report of the 
New Jersey State Museum, 1905. 


“Fishes of the Vicinity of New York City, by John Treadwell Nichole: 
American Museum of Natural History, Handbook Series No. 7, 1918. 


1922] Breder: Fishes of Sandy Hook Bay 345 


CARANGIDA 
54—Seriola zonata (Mitchill) PILOT, PILOTFISH 
A few of fair size were taken in 1920, from July 21 to 
November 8. Only medium sized examples were taken in 1921, 
mostly less than a foot in length. Seen from July 27 to October 
21. Most common in the latter part of August and early Sep- 
tember. 


55—Selar crumenophthalmus (Bloch) [GOGGLE-EYED SCAD] 


Two were taken in fall of 1920. A few examples 25.5 cm. 
(about 10 inches) long were taken on September 14, 1921. Not 
recorded north of Beesley’s Point, Cape May County by Fowler. 


56—Caranx hippos (Linnzus) JACKFISH, RUNNER 


A few were taken in 1920, and several medium sized ex- 
amples between September 13 and 29, 1921. Recorded from 
Atlantic and Cape May Counties only by Fowler. 


57—Caranxz chrysos (Mitchill) 
YELLOW MACKEREL, RUNNER, GOLDFISH 


A few were taken in 1920. Common in 1921, all small and 
most abundant in September. Taken from August 18 to October 
ZAG 1921 


58—Alectis ciliaris (Bloch) THREADFIN, RIBBON FISH 


One example was seen on July 29, 1921, and another on 
August 12, both 10 cm. (about 4 inches) long. Recorded from 
Cape May County only, by Fowler. 


59—Vomer setipinnis (Mitchill) MOONFISH, LOOKDOWN 


A few small ones were taken in 1920. Two fair sized ex- 
amples were taken on July 14, 1921, and another one of about 
4 em. (114 inches) on October 17. Fowler records this from 
Cape May County alone. 


60—Selene vomer (Linnzus) MOONFISH 
Recorded in 1920. 


346 Zoologica: N. Y. Zoological Society [Irsas 


61—Trachinotus carolinus (Linneus) POMPANO 


Several large examples were taken in fall of 1920 as well as 
a few small schools of young fish. Many small examples 7.5 to 
10 cm. (about 3 to 4 inches) were taken in seines and fykes from 
September 27 to October 18, 1921. Scattering small individuals 
were taken from September 27 to October 21, 1921. Much more 
common than the preceding year. 


SCOMBRIDZ 
62—Scomber scombrus Linnzeus MACKEREL 


A few were taken in 1920. A few large and many small ex- 
amples, 15 to 21.5 cm. (6 to 8 inches) were seen June 7 to 
July 8, 1921, while from September 18 to October 21, only small 
ones were observed. Taken in all gear. Small ones 65 to 120 mm. 
(about 214 to 484 inches) taken on July 7 were found to contain 
schizopods, copepods, amphipods, stolephorus sp. and other man- 
gled small fish. 


63—Scomber colias Gmelin MACKEREL 
Recorded in 1920. 


64—Sarda sarda (Bloch) BONITO, HORSE MACKEREL 
A few dozen examples 914 cm. (about 3 feet) long were 

taken between September 13 and 17, 1921. 

65—Scomberomorus maculatus (Mitchill) SPANISH MACKEREL 
One medium sized example was seen on August 25, 1921. 


LABRIDA 
66—Tautogolabrus adspersus (Walbaum) CUNNER, BERGALL 


A few were recorded in 1920. Three small examples were 
seined in ‘Dredged Pool” on July 21, 1921. 


67—Tautoga onitis (Linnzus) BLACKFISH 


Fairly common in the fall of 1920. Large and medium ex- 
amples were taken in pounds in 1921, and by seines in tide pools 
rather sparingly all season, but becoming more common with the 
approach of cold weather. 


1922] Breder: Fishes of Sandy Hook Bay 347 


BALISTIDA 
68—Balistes carolinensis Gmelin TRIGGERFISH 


Several small examples were taken in 1920. Two fair sized 
specimens were taken on October 17, 1921, which is a late date 
for this species. 


MONACANTHIDA 
69—Stephanolepis hispidus (Linnzus) 
FILEFISH, GRANNY WHALE 
Several were taken in 1920. A few small examples were 
taken from September 13 to October 12, 1921. 


70—Alutera schepfii (Walbaum) GRANNY WHALE, HAMBAG 


A few large and several small examples were’ taken in 1920. 
One large specimen was taken on July 28, 1921, and many small 
ones, about 20 cm. (8 inches) long, with juvenile markings from 
then on to October 21. Recorded by Fowler from Atlantic, Cape 
May and Ocean Counties only. 


TETRAODONTIDA 


71—Lagocephalus levigatus (Linnzus) 
RABBITFISH, SMOOTH PUFFER 


One dead specimen was seen in the possession of a fisherman 
on September 15, 1921. 


72—Spheroides maculatus (Bloch and Schneider) 
SWELLFISH, SWELLTOAD, TOADFISH, PUFFER, PUFF-BALL 


Common practically all season in 1920, but less so in fall. 
Common from June 1 to July 15, 1921, becoming scarcer toward’ 
the latter date. Many of these fish were nearly ripe and some 
of the females emitted ova. Later, July 27 to October 21, young 
were taken in seine from 2.5 to 13 cm. (about 1 to 5 inches) and 
longer, while adults were irregularly abundant, but all apparently 
were spent. 


348 Zoologica: N. Y. Zoological Society — [Ins 5 


DIODONTIDZ 

73—Chilomycterus schepfii (Walbaum) 
SPINY PUFFER, BURRFISH, PINCUSHION 
Not rare during the fall of 1920, and a single specimen was 
taken as early as July 27. One large example was reported on 
July 8, 1921, which is an early date, and small ones fairly common 
from September 13 to October 21. These latter averaged 10 cm. 

(about 4 inches) in length. 


MOLIDA® 
74—Mola mola (Linnzus) HEADFISH, SUNFISH 


A few were reported by fishermen in 1920. One small speci- 
men was reported on June 22, 1921, and others were mentioned 
by fishermen about this time. 


COTTIDA 
75—M yoxocephalus octodecimspinosus (Mitchill) 
- DADDY SCULPIN 
A few were taken in the spring and fall of 1920. 


TRIGLIDA 
76—Prionotus carolinus (Linnzus) SEA ROBIN 
A few were seen occasionally all through both seasons but 
not as common as the following species; medium and small ex- 


amples only. In 1920, some as long as 20 cm. (about 8 inches) 
were seen in the autumn. 


77—Prionotus evolans strigatus (Cuvier and Valenciennes) 
SEA ROBIN 


Common both years. Seen from June 21, 1921, to season’s 
end. None above medium size. Small examples, 10 cm. (about 
4 inches) in length abundant in latter part of season. 


ECHENEIDIDA# 


78—E cheneis naucrates Linneeus 
SUCKER, SHARK SUCKER, PILOT, FISH SUCKER, SUCKING FISH 


One was taken on October 11, 1920, which is a late date. Sev- 
eral medium sized examples were taken from July 7 to 15, and 27 


1922] Breder: Fishes of Sandy Hook Bay 349 


to 29, 1921. It is likely significant that their appearance was 
made only when Carcharhinus milberti was also present during 
this year. 


BATRACHOIDIDA® 
79—Opsanus tau (Linneeus) SALLY GROWLER 


Medium and small examples were taken in all inshore nets 
at practically all times both years, but most common in fall. 
Taken in all gear in 1921, from September 13 to October 21. None 
of the fishermen encountered knew this species as Toadfish, the 
appellation generally given it elsewhere. 


MERLUCCIIDZ 

80—Merluccius bilinearis (Mitchill) 
LING, HAKE, SILVER HAKE, SQUIRREL HAKE 
Taken in spring and fall of both years. Seen on June 1, 
1921, and from October 17 to 21, being absent during warm 
weather. In spring the specimens were of good size, while in 
fall they were small, being about 20.5 cm. (8 inches) long. At no 

time was this species very common. 


GADIDZ 
81—Microgadus tomcod (Walbaum) TomMcoD, TOMMYCOD 


Taken in fall of 1920. One example was taken on July 21, 
1921, in “Dredged Pool” and a few in fykes from October 17 to 
21. All small fish. 


82—Gadus callarias Linnzeus Cop, (Small, up to 2 feet—Scrop) 


Taken in lobster pots at mouth of the Bay during November 
and possibly later in 1921. First reported November 1. Fish 
45 cm. (about 18 inches) long. A few taken in the outermost 
pounds. 


883—Phycis regius (Walbaum) HAKE, LING 
A few were taken in the fall of 1920. 


350 Zoologica: N. Y. Zoological Society [11:15 


84—Phycis chuss (Walbaum) HAKE, LING 


Taken in spring and fall of 1920. Taken in spring of 1921. 
This, like the less common preceding species, disappears in warm 
weather. One example examined on June 1 was found to be 
crammed full of large prawns. 


PLEURONECTID2& 
85—Paralichthys dentatus (Linneus) FLUKE 


Fairly common at all times both seasons. The largest in- 
dividual recorded measured 60 cm., 24 inches in standard length, 
67.5 cm., 27 inches in total length, and was a spent female. It 
was seined on June 21, 1921. Small examples 5 to 15 cm. (2 to 
6 inches) frequently were taken in the seine, while most of the 
large ones were met with in the pounds, and occasionally in the 
fykes. 


86—Pseudopleuronectes americanus (Walbaum) FLOUNDER 


All small examples taken both years, varying from 5 to 20 
cm. (about 2 to 8 inches) in length. Taken throughout the season 
of 1921 and chiefly in seines. 


87—Lophopsetta maculata (Mitchill) WINDOWPANE 


Plentiful both seasons, none seen over 18 cm. (7 inches). 
Absent in August and September of 1920, but present at all times 
in 1921. The stomach contents of several examined on June 1, 
1921, consisted of crustacean remains, probably schizopods. On 
August 25, a small example was seined in which the right side 
was almost as well pigmented as the reverse. Only the under 
part of the head and abdominal region was white, the remainder 
duplicating the normal color of the left side. Fowler does not 
record this species from Monmouth County. 


88—EHtropis microstomus (Gill) | [SMALL-MOUTHED FLOUNDER] 


One example, 7.5 cm. (about 3 inches) long was taken in a 
seine on September 28, 1921. This species is not included in 
Fowler’s list as being known from Monmouth County. Deposited 
in the American Museum of Natural History. 


1922] Breder: Fishes of Sandy Hook Bay 351 


SOLEIDZ 
89—Achirus fasciatus Lacépéde HoG CHOKER 


One example of 15.5 cm. (about 6 inches) in length was 
taken on July 6, 1921. 


LOPHIIDA® 
90—Lophius piscatorius Linnzeus 
CARPET BAGGER, ANGLER, TOADFISH 


A few were taken in the fall of 1920. Two examples, each 
76 cm. (about 214 feet) long were taken on October 20, 1921. 


In concluding this list a few remarks on general considera- 
tions may not be amiss. As might be expected, the pound nets 
lying nearest to the shore were found to usually capture the 
smallest specimens as well as the majority of typical shore loving 
fishes, while those further out most often caught the largest 
examples and also contained the highest number of pelagic forms. 
In addition the latter took nearly all the southern representatives, 
which reach here in the latter part of summer and early autumn. 
The combination of a large number of factors is responsible for 
the above mentioned condition, but probably chief among them 
are, salinity, temperature, and water currents. A through study 
of these three elements would almost certainly throw consider- 
able light on the problem of the distribution of the fishes of 
such a small bay as this, besides adding something to the knowl- 
edge of the life histories of a number of species. 


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LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 16 


THE WEAVING OF THE RED-BILLED 
WEAVER BIRD IN CAPTIVITY 4 


By HERBERT FRIEDMANN / > 


‘a 


Poth Behe Ss Heb: D.” BEY} > T:H-E oO. C FH Ey 
Preise Z2OCOLOGTICAL “PARK, ~NEW YORK 


Aucust 23, 1922 


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Corrected to August, 1922. 


LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II, NUMBER 16 


THE WEAVING OF THE RED-BILLED 
WEAVER BIRD IN CAPTIVITY 


By HERBERT FRIEDMANN 


Pee cenncp . Bey. THE = OGLE Ty 
Rin A7ZO0T0GICAL PARK, NEW YORK 


Avucust 23, 1922 


INTRODUCTION 


It is among the weaver birds that we find the art of nest- 
building developed to its greatest perfection. Their nests are 
models of bird architecture and represent the climax of avian 
effort at home making. 


“Instinct and necessity have made these birds wonderful 
architects. Natives of a land where the rays of a vertical sun 
alternate with tropical rains; where monkeys, serpents, and all 
kinds of other enemies abound, the weaver bird has learnt to 
avoid these manifold dangers for his progeny. ... In its details 
the nest of each species of weaver bird varies, but all of them 
are more or less ball-shaped. The roof is always very thick and 
substantial enough to keep off the heaviest downpour, as well 
as to protect the inmates from the tropical sun. The nest is 
invariably suspended from frail branches or reeds just strong 
enough to bear its weight, but never strong enough to tempt 
any predatory animal to climb up. The entrance to the nest is 
invariably from underneath, a sort of ridge dividing the nest 
proper from the entrance, and preventing eggs or young from 
falling out. No bird of prey can therefore possibly see the con- 
tents of a weaver bird’s nest, much less commit any ravages on 
a brood.” ! 


REVIEW OF THE LITERATURE 


For a long time ornithologists have been interested in the 
nests of the Ploceidae and while a great deal has been written 
about them, no one, as far as I have been able to ascertain, has 
ever described the actual details of the methods of nest construc- 
tion in any species of weaver birds, or, for that matter, in any 
species of bird. This may seem to be a hasty statement, but a 
thorough search in the literature of this subject has failed to 
produce a single article of the nature sought for, the nearest and 
best attempts being those conveniently collected by Dr. A. G. 


*B-ackston, W. A., Swaysland, W., and Wiener, A. F. The Book of 
Canaries and Cage Birds, British and Foreign, p. 404. 


355 


356 Introduction 


Butler in his work on “Foreign Finches in Captivity.”2 But 
these deal only with the larger features of nest-building. Plenty 
of good descriptions of these larger features have been published 
for many of the Ploceidae, notably those just referred to, those 
in the works of Shelley,? Chapin,‘ Stark and Sclater,’ Bates,® 
etc., to mention just a few of the more important ones. Bartlett’s - 
unfinished monograph of the weaver birds does not go into as 
great detail in the matter of nidification as a monograph might, 
and, as far as the present paper is concerned, adds nothing to 
those mentioned above. 


As far as the particular species under discussion in this 
paper, Quelea quelea (Quelea sanguinirostris), is concerned, 
good descriptions of its nesting habits have been published by 
Blackston, Swaysland and Wiener,’ by Butler, Shelley, and 
others, but all, as I said before, deal only with the gross aspects 
of the building process. 


In view of the lack of the literature on this point, it may 
not be amiss to present herein a study of the actual weaving of 
one of the commonest species of Ploceidae, the Red-billed 
Weaver. 


* Butler, A. G., 1899, Foreign Finches in Captivity. 

* Shelley, G. E., 1896, Birds of Africa, Vol. IV. 

* Chapin, J. P., 1917, Classification of the Weaver birds. Bull. A.M.N.H. 
XXXVII, -Art. IX, pp. 248-280. 

° Stark and Sclater, Birds of South Africa, Vol. I. 

* Bates, G. E. Ibis., Jan., 1909, p. 44; Ibis, 1911, p. 589. 

™See’ pp. 408-409. 


Vol. 


— 


FIG. 130. WEAVING OF WEAVER BIRDS IN THE PERCHING BIRDS’ 
HOUSE IN THE ZOOLOGICAL PARK 


1. Nest recently started, showing vertical loop. 2. Completed nest. 3. Part of playground. 
Photographed two years after the studies were made. 


Zoologica Vol. Il, No. 16. Face Page 35/7. 


Volume II, Number, 16 


THE WEAVING OF THE RED-BILLED WEAVER 
BIRD, QUELEA QUELEA IN CAPTIVITY 


By HERBERT FRIEDMANN 


1. General Features of Nest-building 


The general features of nest construction in this species 
have been described in more or less detail by several writers 
referred to above. Therefore I shall pass over this phase of the 
subject in a hasty manner and, were it not for the fact that in 
captivity the birds build nests quite unlike those built by the 
same species in nature, I would scarcely have touched upon it. 


Not only did the nests built in the Zoological Park differ 
from those in nature, but also from those built in the aviaries 
of Dr. A. G. Bulter in London where, to judge by his descrip- 
tions, the nests built in captivity agree with those built in a 
state of nature. I cannot, and do not attempt any explanation 
for the discrepancies between the nests in captivity as I found 
them and those in nature or in the London aviaries. 


In nature, the nest of the species is described as a more or 
less globular nest with an entrance hole on one side. In cap- 
tivity the nest was really cup-shaped and had an arched roof 
which was attached to the nest proper at the two ends of the 
major axis of the slightly elliptical margin of the cup, and free 
on the sides. If the roof had been adnate to the bowl around its 
entire edge, the nest would have been truly globular. Mr. 
Chapin, whose wide field acquaintance with the Ploceidae gives 
his statements the stamp of authority, writes me that ‘“ordi- 
narily no weaver would leave a part of the roof open as in your 
sketch. Building is generally begun by weaving an upright 
ring at about the middle of the nest-to-be, and then adding the 
back of the nest, and the front, with entrance. So far as I know 
they never begin by constructing a simple cup-nest like a vireo.” 


In Reichenow’s Vogel Afrikas III, p. 109, von Heuglin is 
quoted to the effect that while the subspecies aethiopica built 


357 


FIG. 131. OUTLINE DRAWINGS OF TWO NESTS AND PART OF A PLAY- 
GROUND BUILT IN CAPTIVITY. 


P1922 Friedmann: Weaving of Weaver Birds 359 


purse-shaped nests in the Gardens at Khartoum, especially in 
Parkinsonias, they seemed never to lay in them. 


The South African species is said to be parasitic. This is 
probably an error. 


The two nests built by this species in captivity and part 
of the “playground” are shown in outline sketches (Fig. 131). 
It will be noticed that the two nests are very dissimilar in the 
degree of curvature of the roof but that they agree in having 
the roof partly free on the sides. The “playground” consists 
of perches, arches and runways, connecting the nests. 


When building, Quelea quelea usually selects a fork of a 
branch and weaves a small mass of fiber or whatever material 
it may have (raffia in this case) right in the crotch of the fork. 
From this as a basis it forms a hoop nearly vertical as a rule. 
“From this hoop it works, starting from the bottom and gradu- 
ally filling in the back, finishing off with the front, in the center 
of which it leaves a small hole to enter by.” * 


2. Details of Weaving 


At the time this study was begun, there were nests already 
built, and also a large mass of what we may collectively call the 
“playground.” This gave the birds two different types of sites 
for weaving—the first being the bare twigs around which they 
might weave, the second being the already existing woven foun- 
dations (nests and playground), to which they might add by 
weaving. 


There seemed to be different types of stitches employed by 
Quelea quelea when weaving on a previously woven foundation 
than when weaving around a branch or twig. In describing 
stitches, I believe that diagrams are clearer and more eloquent 
than words, and consequently this text is largely an explanation 
of the accompanying plates. 


_ When weaving around a twig or branch, Quelea quelea used 
three types of stitches as illustrated (Fig. 1382). The arrows 
indicate the direction of the progress in making the stitch; the 


* Butler, A. G., 1899. Foreign Finches in Captivity, p. 229. 


360 Zoologica: N. Y. Zoological Society Piisate 


dotted lines represent the strand of raffia as being on the far 
side of the twig, i.e., the portion that would be hidden by the 
twig in this view. (1) shows what seems to be a stitch used 
chiefly in conjunction with that shown in (2). Here the bird 
places a strand longitudinally along the branch, pushes one end 
around the twig, catches it on the other side, carries it up and 
over, tucks it under the part of the strand lying lengthwise 
along the branch, pulls it through and then tightens it with 
jerks of its head, seizing the strand nearer the knot with each 
jerk, until the knot is tight. In all weaving, the bill is the weav- 
ing organ, taking the place of a hand, the feet being used merely 
to clamp the straw down to the twig and hold it there. 


In (2) is illustrated the simplest and most commonly used 
of all the stitches. The diagram should be self-explanatory, the 
stitch consisting in merely laying a strand longitudinally along 
a branch for about half its (the strand’s) length and winding 
the remaining half around and around to hold it there. When 
the end of the strand is reached it is tucked in as in (1). 


A variation of the type shown in (1), is illustrated in (3). 
This type is apparently rare as it was seen but once out of hun- 
dreds of stitches observed. It was used to tuck in the end of the 
strand at the completion of a stitch of the type shown in (2). 
All these three types were used when weaving on straight limbs. 


When weaving at a fork the birds did one of two things: 
Generally they wove a solid compact mass of straw on the two 
arms of the fork before stretching a single strand across. How- 
ever, they sometimes, though seldom, stretched a strand across 
before weaving any foundation on either side. They would 
straddle the fork, one foot on each side, just as far apart as pos- 
sible. At times the distance between their feet was more than 
twice the width of the body! Then they would fasten the ends 
of the strand on either end by a stitch as shown in (2) and (1), 
Fig. 132. However, when the birds did act as above, they in- 
variably drew the straw down to the crotch of the fork after 
they were through. Evidently they have no liking for frail sus- 
pension bridges. 


FIG. 132. TYPES OF STITCHES USED BY Q. QUELEA WHEN WEAVING 
AROUND A BRANCH 


(4) after drawing by James P. Chapin. 


361 


362 Zoologica: N. Y. Zoological Society | PIL: 9G 


When the birds were weaving on an old foundation of straw 
previously woven, their stitches were quite different from any 
of the preceding. In Fig. 133 the stitch is shown in detail in 
(1 to 10 inel.). The arrows indicate the direction of movement 
of the straw, dot-and-dash lines represent the straw being pulled 
through the mass of straw foundation, and dotted lines indicate 
the straw being on the other side of the mass. 


The letter A marks the place in each case where the bill was 
applied. Briefly, the stitch is as follows: The bird holds a strand 
near one end in its bill and pushes it through the already existing 
woven mass (represented in the diagram by the space between 
the two parallel horizontal lines) as in (1). Bending over, it 
pulls the strand until one end is through as in (2-4). Then the 
bird takes the strand around the back of the mass (5-6), and 
repeats the process (6-9), the next time winding it in front of 
the mass as in (10). This stitch formed by far the greatest 
part of all the weaving done by Quelea quelea. 


Especially interesting are (6’) and (7’) as indicative of 
the intelligence of these birds. In (6’) the bird made a knot 
by pulling the strand through the loop. Then on pulling at 
(A) (6’), to draw the knot tight, the bird evidently noticed that 
the part of the strand (B) (6’) was being drawn through the 
woven mass more and more with each tug at (A). The bird 
then tucked (B) under the loop (C) (7’) and then went back 
and jerked at (A) without any danger of pulling the strand 
out! 


The weaving done by Quelea quelea is not only intricate 
and beautiful but it is strong and serviceable. I tried to pull 
down some straw the birds had woven on the wire netting of 
the cage. In one case the straw was attached only at one end, 
the other end dangling freely. I pulled with a force that I esti- 
mated to be over ten pounds, and the straw broke but the knot 
did not undo itself! On the contrary it seemed to become tighter. 


The speed with which the birds weave is subject to great 
variation. All the straws used were of approximately equal 
length (one foot) so that in comparing speed, the comparison 
was a fair one. The speed varied from forty-eight seconds to 


FIG. 133. STITCHES OF WEAVER BIRD OVER AN OLD FOUNDATION. 


364 Zoologica: N. Y. Zoological Society ARTIS Rat 


over thirty minutes per straw. Of course in winding around two 
twigs it would naturally take less time to use up a straw on a 
twig of greater diameter than on a smaller twig, there being > 
fewer revolutions necessary in the former case. 


The relation between the bill and the feet in weaving is in- 
teresting. The birds always pick up the strands with their bills, 
but invariably hold them down on the twigs or woven masses 
with their feet. A highly specialized case of correlated action 
is shown in Fig. 134. The bird pushed the strand underneath 
its toes as in (1-8). Then it took one end of the strand around 
the twig as in (4), and tried to push it under between its toes. 
(4A) shows the same position as (4) but from another view. 
Then to facilitate matters, the bird raised its middle toe, thereby 
loosening the straw and allowing the bill to work its way under 
and make the knot. This was observed but once, so that it is 
evidently not a general practice with Quelea quelea, but never- 
theless serves to emphasize the degree of skill and intelligence 
with which these birds are endowed. 


The discrimination shown by these birds in their nest mak- 
ing is little short of amazing. The location for permanent weav- 
ing is chosen only after many trials of various places. They are 
extremely critical of their weaving, often pulling out part of 
their nests and weaving it over again. In one case a bird pulled 
the same straw out eight times before it was satisfied with the 
manner in which it was woven. The general practice was this: 
A bird would weave in a strand, wipe its bill on the twig and 
then view its work from all sides. Then if not satisfied*it would 
try to mend it or pull it out entirely and try again. The weaving 
had to be compact or it was not satisfactory. Often the birds 
would pull and pull at a straw, each time jerking their heads 
back and forth with such force that it was a wonder that they 
could stand the strain. The discrimination of Quelea quelea 
with regard to color is fully described under Color Preferences. 
Suffice it to say here that red was the favorite color and orange 
next, while green, black, blue and violet were not used to any 
great extent. 


The birds also showed considerable discrimination with re- 
gard to the width of the straws used. They preferred thin, fine 


FIG, 134. HIGHLY SPECIALIZED CORRELATED ACTION BETWEEN BEAK 
AND FEET. 


(1-4) Figure of foot after drawings by James P. Chapin. 


365 


366 Zoologica: N. Y. Zoological Society [iS 26 


straws to coarse, heavy ones. In fact, on numerous occasions, 
after stretching a strand across a fork, a bird would peck at the 
middle of the strand until the raffia would split longitudinally. 
Then it would pull on one side, thereby elongating the split until 
the entire length was divided into two thin strands where there 
had been but one wide strand. Quelea quelea is about as active 
and tireless a nest builder as one can want. The birds are al- 
ways building new nests or playgrounds and when they have no 
building material, they busy themselves with repairing or even 
tearing down the old nests only to rebuild them and tear them 
down again. In his activity and “restless anxiety to weave nests, 
anything comes handy to the Red-beaked Weaver-bird, and a 
small finch coming near him would at once find himself minus 
a wing or tail feather, a friendly pecking at the neighbor’s plum- 
age being more convenient to the Red-billed Weaver than a 
search after a bit of fibre.’” 


This restless desire to build seems not to be restricted to 
Quelea quelea for Bates'® writes of Ploceus cucullatus that 
SERN See FS tearing down their nests only makes them buiid the 
more furiously. They have a perfect mania for building, and 
when not building new nests are all the time repairing the old 
ones. They often destroy palm trees by stripping them bare 
of their leaves.” 


There was much keen rivalry between birds for straws. If 
one bird picked up a straw and dropped it, another bird was 
sure to pick it up in preference to any other straw, regardless 
of color or width. It was not uncommon to see two birds, one 
on each end of the same strand pulling in opposite directions. 
The birds sometimes tried to frighten each other away from their 
nests by spreading out their wings, somewhat in the style of the 
intimidation display of the white-breasted nuthatch, as described 
by Allen", lowering the head, opening wide the bill and uttering 
a harsh scolding note. 

* Blackston, W. A., Swaysland, W., and Wiener, A. F. Book of Canaries 
and Cage birds, British and Foreign, p. 404. 

* Bates, G. L.: Tbis; Jan:; 1909; py: 44. 

* Allen, A. A., Bird Lore, Vol. XXI, No. 1. 


1922 Friedmann: Weaving of Weaving Birds 367 


The presence of crowds of people tended to make the birds 
more active, as on Sundays when thousands of people watched 
them during the day. Ordinarily the birds were most active 
from 10:00 A.M. to 11:00 A.M.; least active, or rather inactive 
from 11:00 A.M. to 2:00 P.M.; and active again from 2:00 P.M. 
to 4:00 P.M. If no crowds were present the birds would tend 
to sleep from 11:00 A.M. to 2:00 P.M., a habit reminiscent of 
their lives in tropical Africa. 


COLOR PREFERENCES 


' In studying the stitches used by the birds it was found con- 
venient to use various colors of raffia so that each stitch would be 
easy to follow through. Incidentally it was found that the birds 
seemed to have a definite preference for certain colors, chiefly 
red and orange. 


In testing for color preferences, the method used was as 
follows: Seven colors of raffia were used, the raffia being of ex- 
actly the same texture as the raffia the birds had been using for 
some months previously. The colors used were red, orange, 
yellow “natural,” blue, green, violet and black. There was no 
noticeable difference between these straws in any respect except 
as to color. They were tested for taste and no difference in taste 
was found for any color. They were tested for weight, and 
found alike. Tests were also made for strength and texture, and 
all gave similar results. Therefore it was safe to say that the 
straws were exactly alike except in color. (The term straw as 
used in this paper refers to a piece of raffia. The term is used 
merely for convenience.) 


Thirty-six equal sized pieces of each color raffia were dis- 
tributed over the floor of the cage at the end of the day, care 
being taken to see that the colors were evenly scattered. The 
birds did not venture to touch the strange material until the next 
morning. By watching them all the next day (from 10:00 A.M. 
to 4:00 P.M.), I was able to record just how many pieces of each 
color each species took, used or rejected. Then, at the end of 
this day, I added to the raffia previously put in, the same number 
of pieces of each color as the birds used up during the day. 


This was repeated each day for four days. Then the ex- 


368 Zoologica: N. Y. Zoological Society [ibs 16 


periment was repeated nineteen days later and carried on for 
three days. Thus every day the birds had thirty-six pieces of 
each of the seven colors or two hundred and fifty-two pieces of 
raffia in all.to start with. Therefore, by adding together the 
results of the different days’ tests for each species, I was able to 
find what colors each preferred and what each disliked. The 
possibility of the birds using up the colors of their preference 
and then, through lack of these, having to use other colors was 
eliminated by starting them off in the beginning with more of 
each kind than they could use up in a day and by adding each 
day just what they used up as described above. 


All the straws used in the experiments were of equal thick- 
ness and about a foot long, this length being chosen because it 
satisfied two conditions: it was long enough for the birds to use 
with comfort, and at the same time it was short enough to en- 
able each bird to use quite a few pieces each day. This latter 
condition was essential if any appreciable number of records 
were to be obtained. 


The experiments were extended not only to include Quwelea 
quelea, but also its close relative Quelea russi, the Russ masked 
weaver bird. There were five individuals of the Red-billed 
Weaver and three of the Russ Masked Weaver under observa- 
tion. The following table illustrates graphically the substance 
of the present paragraph: 


RED-BILLED WEAVER, Quelea quelea 
(5 individuals) 


RED ORANGE | YELLOW GREEN BLUE VIOLET BLACK 


Used 59 27 22 8 11 iz 3 
Rejected 0 0 oes 10 3 5 2 

Total 59 27 25 18 14 12 5 
Percent are |) ylOOm 1 200 88 44 78 58 60 


Percentage | 0 12 56 22 42 40 


Rejected 


1922 Friedmann: Weaving of Weaving Birds 369 


Russ MASKED WEAVER, Quelea russi 
(3 individuals) 


| RED ORANGE | YELLOW GREEN | BLUE VIOLET BLACK 
Used | 21 20 fi 3 1 3 8 
Rejected | 0 0 0 3 fa 0 0 
Total | 21 20 7 6 2 3 3 
Percentage 100 100 | 100 50 50 100 100 
eet 20 0 0 50 50 0 0 


It will be seen at a glance that in numbers of straws used 
by Quelea quelea, red is more than double orange which, in turn, 
is greater than yellow, etc. In numbers of straws rejected, that 
is, picked up by the birds and then voluntarily (apparently) re- 
jected, green leads with ten, while red and orange were never 
rejected. In this connection I may say that I counted as rejected 
those straws, the rejection of which appeared to be voluntary 
on the part of the birds. Several times a bird picked up a straw 
and was chased by another bird or frightened by some noise, 
and dropped the straw and flew off to a perch. These cases are 
not counted here, as they evidently have nothing to do with color 
rejection. Several red and orange straws were rejected in this 
way. In view of this it may be that the figures given in the 
table are not wholly correct but the general results are probably 
very nearly true. While the preferences may not be as marked 
as the figures would indicate, we must admit the existence of 
these preferences. The accompanying graph (Fig. 135) repre- 
sents the color preference of Quwelea quelea as interpreted by the 
number of straws of each color used by that species. 


If we add the number of straws used and the number re- 
jected for each color and then find that what not that percent of 
the total number of each color was used we find the preference . 
to be: 


t ii a 
cat | HEH 
7 u B 
44 
‘1 
i 
T 
t t 
re} j T 
t t 
{ 
a uy Buea t t 
a T +t 
of 2 Oe rt { t ttt 
i “ wa aa a co i t 1 t i t to t 
amt wee bee u { T 1 i ro 

seepepes iB ReESRSESESEE HH + rH 

pecacese Het t tH H 
wage 4 2 T L 

eececscet eal SEBGBGGH coeue sunuaGi att 4 4 

ivan ima ol t t r at t u 

Senn eoews eas +t t 

in it SEC Et ae ee rot eect or 
But iaresovend es Es Cosouezscd GEaTeEonsesasodarest enate ares potatoe 
Oo es SS * ° . s . es . HET, tte ¥ 
DMDEONYTONHOMRDEONYNNHOMDLCONGTNNHOMDL ONGTNNAHOMDDLCONYNNAHOMADrONYNANHO 
w wo = a Ce) "9 nN N ca a 


Guasn SMVULS AO WHAIWAN 


VIOLET BLACK 


ORANGE YELLOW GREEN BLUE 


GRAPH SHOWING COLOR PREFERENCES OF @Q. QUELEA. 


RED 


ae S35. 


FI 


370 


1922 Friedmann: Weaving of Weaving Birds ey 


1. Quelea quelea—red 100%, orange 100%, yellow 88%, 
blue 78%, black 60%, violet 58%, green 44%. 


2. Quelea russi—red 100%, orange 100%, yellow 100%, 
violet and black 100% (not significant as there 
were only three straws of each color picked up 
as compared with twenty-one red and twenty 
orange), and green 50%, blue 50%. 


While there seems to be a fairly well defined color prefer- 
ence for each species, yet there is considerable variation among 
the individuals of the same species. Thus, one individual of 
Quelea quelea used in one day one red, one orange, one violet, 
and one blue straw, while another individual of the same species 
used in the same day three red, three orange, one yellow (and 
rejected one yellow) and one green straws. Still another indi- 
vidual of this species used on the same day four red, one orange, 
one yellow, and one blue straws. Yet each individual, of the five 
used at least one red and no more than one of any other color 
except orange on that day. Not only is there variation between 
individuals of the same species, but the same individual may 
vary from day to day. 


SUMMARY AND CONCLUSIONS 


1. In captivity the birds built abnormal nests, but later 
(two years after), they built normal ones, as shown in Fig. 
130. 


2. Normally they begin by weaving a vertical hoop, and, 
beginning at the bottom, fill in the back and then the front, in 
the middle of which they leave an entrance hole. 


3. Different types of stitches were used in weaving on bare 
twigs and on previously woven masses. 


4. In weaving around bare twigs, three types of stitches 
were used, as shown in Fig. 1382. On this plate (2) shows the 
commonest stitch used. 


5. The type of stitch used in weaving on a previously 
woven mass is shown on Fig. 133 and needs no further mention 
here. 


abe Zoologica: N. Y. Zoological Society [1G 


6. All actual weaving is done with the bill; the toes being 
used to pick up and hold in place the straws used. 


7. A considerable degree of correlation seems to exist be- 
tween the bill and toes in weaving. 


8. The birds exhibit a remarkable amount of discrimina- 
tion in respect to the color and width of the straws used and in 
regard to the compactness of the weaving. 


9. The birds preferred red and orange to all the other 
colors used, the colors being taken in the following order: 


Red-orange-yellow-green-blue-violet-black. 


The last three are not significant due to the paucity of 
records. 


ACKNOWLEDGMENTS 


The work on which this paper is based was conducted 
at the New York Zoological Park through the courtesy of Mr. 
William Beebe and Mr. Lee S. Crandall of the Department of 
Birds. I take this opportunity of acknowledging my indebted- 
ness to both Mr. Beebe and Mr. Crandall for their many kind- 
nesses and to Professor A. J. Goldfarb of the College of the 
City of New York for helpful suggestions and criticism. For 
permission to use the figure of a ploceine foot Fig. 134 (1-4) I 
am indebted to Mr. James P. Chapin of the American Museum 
of Natural History, the original of this figure having appeared 
in his paper on the classification of the weaver birds. Mr. 
Chapin very kindly has given me also a drawing of a Red-billed 
Weaver, Fig. 132 (4), for use in this paper. 


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OBJECTS ‘OF THE SOCIETY 


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ZOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 17 


ian Instip~ 
ry it ett 3 ~ 
s1aNB) ly ue 


(Papers from the New York Aqu 


(Contribution Number 9 co ‘G, > 


THE WHITEFISHE®"! ~’ 


(Coregonus clupeaformis) js a 

“ a2 To nal NM W3e™ 

WHITEFISHES REARED IN THE NEW YORK AQUARIUM 
By IpA M. MELLEN 


New York Aquarium 


A STUDY OF THE SCALES OF WHITEFISHES OF 
KNOWN AGES 


By JOHN VAN OOSTEN 
Field Assistant, U. S. Bureau of Fisheries 


EBL 135 ED Bee, ts 2 a me Ct TY 
Bae .2OOLOGICAL PARK, NEW YORK 


June 19, 1923 


590.5 


\ 


wt 
~~ ’ 


New York Zonlogical Society 


General Office: 111 Broadway, New York City 


Officers 
President, HENRY FAIRFIELD OSBORN; 
Vice-Presidents, MADISON GRANT and FRANK K. STURGIS; 
Secretary, Chairman, Exec. Committee, MADISON GRANT: 
Treasurer, CORNELIUS R. AGNEW 


Board of Managers 
Glass of 1924 
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GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFORD 
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PRATT, T. COLEMAN DUPONT, HENRY D. WHITON, 
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Corrected to April, 1923. 


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LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
NEW YORK ZOOLOGICAL SOCIETY 


VOLUME II. NUMBER 17 


(Papers from the New York Aquarium) 
(Contribution Number 9) 


THE WHITEFISHES 


(Coregonus clupeaformis) 


WHITEFISHES REARED IN THE NEW YORK AQUARIUM 


By IpA M. MELLEN 
New York Aquarium 


A STUDY OF THE SCALES OF WHITEFISHES OF 
KNOWN AGES 


By JOHN VAN OOSTEN 
Field Assistant, U. S. Bureau of Fisheries 


Pie Ets. Ei. D BY fa = ia a € LE TY 
THE ZOOLOGICAL PARK, NEW YORK 


June 19;-1923 


THE WHITEFISHES 


(Coregonus clupeaformis) 


WHITEFISHES REARED IN THE NEW YORK AQUARIUM 


By IDA M. MELLEN 
New York Aquarium 


A STUDY OF THE SCALES OF WHITEFISHES OF 
KNOWN AGES 


By JOHN VAN OOSTEN 
Field Assistant, U. S. Bureau of Fisheries 


Volume II, Number 17 


THE WHITEFISHES 


(Coregonus clupeaformis) 


REARED IN THE NEW YORK AQUARIUM 
By IDA M. MELLEN 


New York Aquarium 


The New York Aquarium is showing a unique exhibit of 
whitefishes (Coregonus clupeaformis) hatched in the Aquarium 
in January, 1913; unique in that no similar exhibit ever has been 
seen there or anywhere else. Once or twice whitefishes were 
reared in the Aquarium to the age of a month, and once to the 
age of a year; but those now on exhibition—nine years old at the 
present writing—are the only whitefishes ever reared in cap- 
tivity from egg to maturity. They are the survivors of a few 
hundred specimens retained from the fry that hatched out of a 
consignment of a million eggs received in December, 1912, from 
the United States Bureau of Fisheries. The eggs came from 
western Lake Erie waters, and were shipped from the fisheries 
station at Put-in-Bay, Ohio. The remainder of the fry were dis- 
tributed to state waters of northern New York and New Jersey. 


By checking up our findings with those of other observers, 
we have learned some interesting facts about whitefishes, the 
probable age when they arrive at sexual maturity, their longevity, 
etc. As ours are the only fishes of this species in existence the 
age of which is positively known, certain biologists engaged in 
the study of lepidology (the scales of fishes) have been provided 
with scales from the Aquarium specimens, whitefishes being one 
of the species in which the age is written in concentric rings on 
the scales. Of the various kinds of scales in fishes—placoid, 
ctenoid, cycloid, ganoid or rhomboid, ete.—those which char- 
acterize the whitefish are of the cycloid type, 7. e., thin, rounded 


375 


376 Zoologica: N. Y. Zoological Society [II; 17 


scales, having concentric rings as just described, and with no 
spiny projections. 


A cycloid scale strongly reminds one of a finger print. It 
has its central whorl, called the focus, which starts the series of 
concentric rings, called circuli, and there are lines of radiation 
called radii. But unlike the finger print, groups of the circuli 
are marked off in definite dark and light bands, two such bands 
representing one year of the fish’s life. That is, there is a large 
area of light summer bands and a small area of dark winter 
bands or annuli, growth being more rapid during the months 
when temperature is higher and food more abundant. 


Dr. Jacob Reighard’s assistant in the Department of 
Zoology, University of Michigan, Mr. John Van Oosten, who is 
also in the employ of the United States Bureau of Fisheries, 
made a study of whitefish scales from our Aquarium specimens. 
One dead fish and three scales from a live fish were sent him 
each month for twelve months, in order that he might observe 
progressive changes in the scales of specimens the age of which 
was known, though it was not to be supposed that fishes that 
had spent all their lives in captivity, with limited swimming 
space and little variety in their food, would compare favorably 
in development with wild fishes. The method followed in remov- 
ing the scales was suggested by Mr. Van Oosten and consisted in 
segregating two specimens, removing scales from one fish one 
month and from the other the next, thus giving each fish a rest 
of two months between operations. The scales were removed 
with a small forceps. 


Whitefishes are so fragile that they are sometimes killed 
merely by transference from one tank to another, or by an acci- 
dental stroke of the brush when their tank is being cleaned. It 
was therefore with some trepidation that the writer commenced 
this series of very delicate operations. The first specimen oper- 
ated on died within fifteen minutes; but this tragedy was never 
repeated. All subsequent operations were made on specimens 
held by the head and tail in a shallow pan of water, the gills 
being kept continually moist, and great care being taken to 
remove scales not too near the lateral line (in fact, all scales 
were removed from a spot about half way between the lateral 


1923] Mellen: New York Aquarium Whitefishes 377 


line and the back, the section anterior to the dorsal fin being 
selected) ; also to remove them rapidly and apply a strong solu- 
tion of permanganate of potassium at once to the injured spot, 
returning the fish quickly to the tank. The operation always 
exhausted the fish and it would lie quite still while the perman- 
ganate was poured on. When returned to the tank it floated on 
its back, breathing hard for a few minutes, then gradually 
equilibrated itself and recovered. 


Mr. Van Oosten found that the scales of the Aquarium 
whitefishes revealed the stunted growth of the specimens in the 
growth of their circuli, but by experience he gradually learned 
that by selecting scales of a certain shape, he would obtain scales 
which showed the annuli or year-rings so clearly that any inex- 
perienced person could read them with ease. In fact, he had 
various such people read the scales though they knew nothing 
about the age of the fish from which the scales were taken. He 
found that while the circuli are formed throughout the year, the 
annulus is truly a winter-band, being formed only during the 
winter months, 2. e., after October. All of which is explained 
in detail in his paper here published. 


So many of these notable whitefishes died in 1921 that it 
was imagined they might have lived the natural span of their 
existence; yet they had never been observed to spawn. It was 
hinted that they must have done so and the eggs had been eaten 
so fast that no one ever saw them; but Mr. Robert J. Lanier, of 
the Aquarium staff, to whom belongs the honor of having reared 
these fishes, has kept watch over them all their lives and was 
entirely certain that they had never spawned. Was it possible 
that captivity prevented them from attaining sexual maturity? 
If so, the case was a unique one. 


In January, 1922, when the fishes were exactly nine years 
old, females were observed swollen with eggs which, however, 
lacked vitality to such an extent that they could not be fertilized! 
Were these whitefishes reaching sexual maturity and their 
natural span of life at the same time? It seemed impossible. 


The writer knew of only one way to solve the riddle. Dr. 
Wilbert A. Clemens, of the Department of Biology, University 
of Toronto, who had also received some scales from the Aquarium 


378 Zoologica: N. Y. Zoological Society ee by 


specimens, had studied the scales of wild whitefishes. Perhaps 
he had found some that were older than ours! We wrote him 
about the eggs, their lack of vitality, the supposition that nine 
years might be the natural term of the life of the fish, etc., and 
asked if he had ever found a wild specimen more than nine 
years old. 


His answer, as the following quotation from it shows, was 
fraught with interest: 

“T have indeed taken whitefishes much older than those 
you have at the Aquarium. I spent last summer on Lake 
Nipigon and according to my records the largest whitefish 
we took was twenty-one and a quarter inches in length and 
was at least sixteen years old. I have records of two others 
about twenty inches in length, which I have recorded pro- 
visionally as sixteen and seventeen years old, but possibly 
eighteen or nineteen. 

“We do not know as yet at what age whitefish first 
spawn. In Lake Erie I suspect it occurs at the end of the 
fifth or sixth summer. In Lake Nipigon it probably occurs 
considerably later. On November 11 of last year I received 
from the spawn-takers on Lake Nipigon four whitefish which 
they said were the smallest they had taken spawn from. 
These were scarcely fifteen inches in length and were nine 
and ten years of age. So it may be that the whitefish in the 
Aquarium are just reaching the spawning age, and although 
the eggs are few and weak this year they may be normal or 
nearly so next year. It will certainly be interesting to see 
what happens.””! 


Mr. Van Oosten’s paper describes the condition in which he 
found the sex organs of the Aquarium whitefishes. 

This species of whitefish normally attains a length of two 
feet or more, but none of the specimens in the Aquarium meas- 
ures more than fifteen inches. We have not infrequently 
observed that fishes and even snails are stunted by captivity. It 
is known, however, that fishes do not always stop growing when 
they reach sexual maturity or decline in years, many continuing 
to grow as long as they live; and we believe that these whitefishes 
are still growing, though they probably never will be of normal 
size. 


1In December, 1922, when the fishes were nearly ten years old, a female was 
again observed carrying eggs. She was stripped and the eggs fertilized; but they 
were weak and did not develop beyond the morula stage. 


1923] Mellen: New York Aquarium Whitefishes 379 


The last time these specimens were counted was in 1919, 
when they were transferred from one tank to another and num- 
bered two hundred and sixteen. They have dwindled to about 
eighty-four, and it is hoped that we may some day be able to 
repeat the remarkable feat of rearing some to maturity from 
the eggs—a feat many times attempted both at the Aquarium 
and at Government hatcheries, but only once performed. 


Jordan and Evermann (Fishes of North and Middle 
America) state that the common whitefish (Coregonus clupea- 
formis) “is subject to considerable variations, dependent on food, 
waters, etc.”’ The food of fishes is indeed a factor of so great 
importance in their growth and development that breeders of 
fancy varieties believe the food controls not only the size and 
health of the fish, but the actual shape and beauty of its fins. 
Like other vertebrated animals, they require, for perfect devel- 
opment, foods that are both nourishing and bone-building. 


In a state of nature infant whitefishes, judging from those 
in the Aquarium tanks, remain near the surface for a.time, their 
first food consisting of plankton—live floating matter of both 
vegetable and animal character: protozoa, diatoms, minute crus- 
taceans such as the young of the shrimp (Gammarus) and water 
fleas (Cyclops, Cypris, Daphnia), etc. Later they subsist entirely 
on minute crustaceans of these and other species. In the earliest 
period of their lives, after the absorption of the yolk sac, Dr. S. 
A. Forbes, of Illinois, has observed that the fry are provided with 
four curved teeth in the lower jaw, which are of no possible 
service and are subsequently lost. 


Gradually they descend to the bottom for food, and there, 
according to those who have examined the stomach contents of 
adult wild specimens, they feed on small live invertebrates, prin- 
cipally crustaceans, snails, insect larvae and water beetles. 


The crystalline gray-whiteness of mature whitefishes is 
exceedingly attractive to the eye, and the specimens in the New 
York Aquarium have long furnished one of its most pleasing 
exhibits for the casual visitor, as well as one of its most im- 
portant economic exhibits for the fish culturist. This species of 
whitefish is not only the largest, but the most delicate in flavor 
of all the whitefishes of the Great Lakes. 


The Aquarium specimens have never known the excitements 
of wild life, or what it means to hunt or be hunted. 


New York Aquarium, April 1922. 


Ef. 


III. 


IV. 


INTRODUCTION. 


Description of a Typical Whitefish Scale. 

Scale Method and Its Application to Life History Work. 
Assumptions of the Scale Hypothesis. 

Summary of Literature. 

Statement of Problem. 

Description of Apparatus and Method Employed. 
Acknowledgments. 


mai Se Ove ee hes 


NEW YORK AQUARIUM WHITEFISH SCALES. 


1. Annuli and Number of Winters of Life. 
2. Marginal Growth and Time of Formation of Annuli. 


3. Correlation between Annual Growth in Length of Body 
and Scales. 


LIFE HISTORY OF AQUARIUM WHITEFISH AND FACTORS OF 
ANNULI FORMATION. 


1. Food. 

2. Temperature. 

3. Sexual Maturity and Spawning. 
4. Annual Rate of Body Growth. 


SUMMARY. 


BIBLIOGRAPHY. 


380 


Volume II, Number 17 


THE WHITEFISHES 


(Coregonus clupeaformis) 


A STUDY OF THE SCALES OF WHITEFISHES OF 
KNOWN AGES’ 


By JOHN VAN OOSTEN 
Field Assistant, U. S. Bureau of Fisheries. 


INTRODUCTION. 


During recent years many investigations of scientific and 
economic importance have been conducted on the age and the 
rate of growth of fishes as determined from a study of their 
scales. This has involved the interpretation of certain rings 
found on the exterior surface of scales. 

In order to illustrate clearly the mode of the formation of 
these rings, Fig. 137.is presented. It represents a typical scale of 
a whitefish, 197 mm. in length, captured October 22, 1917, at 
East Tawas, Mich., on Lake Huron. Near the center of the scale 
is a small, clear area, the focus (F), which represents the 
original scale in the young specimen. Around this focus are 
numerous, more or less relieved striations, concentric or nearly 
so with the margin. These are termed circuli (C) and like the 
rings in a tree mark successive stages in the growth of the scale. 
Running from the focus to the periphery of the scale are four 
more or less conspicuous radiating ridges (AR, PR), which 
divide the surface of the scale into four roughly triangular areas 
or fields. When the scale is in position in the fish the area to the 
right in the figure is directed towards the tail and is therefore 
designated as the caudal or posterior area (Caudal). The area 
opposite the caudal is the anterior (Anterior), while the two 
areas which separate the caudal from the anterior are the lateral 
or the dorsal (Dorsal) and the ventral (Ventral). The borders 
of these four areas which form the periphery of the scale are 
accordingly termed the caudal, anterior, dorsal and ventral 


2Contribution from the Zoological Laboratory, University of Michigan, 
published with the permission of the U. S. Commissioner of Fisheries. 


381 


382 Zoologica: N. Y. Zoological Society ib Bes Ly 


borders. The radiating ridges are either antero-lateral (AR) 
or postero-lateral (PR). The greatest antero-posterior diameter 
which bisects the caudal area of the scale is its length (L-L). 


By careful examination two distinct zones may be seen in 
this scale, an inner characterized in general by more closely 
spaced lines and an outer in which the lines are further apart. 
The two zones are more readily seen when the figure is viewed 
from such a distance as to somewhat obscure the details. The 
inner zone represents, according to current theory, the entire 
growth of the first year, while the outer zone represents the 
growth of the second summer. If the lines of growth in the 
lateral field be followed from the center outward and downward 
along the antero-lateral ridge, it may be seen that the first twenty 
are complete and uniformly spaced. With occasional breaks and 
irregularities they may be traced entirely around the scale. The 
next six are incomplete and the outermost of them ends (or 
begins) near the antero-lateral ridge. Following this last in- 
complete line to the anterior field, a region is encountered within 
which the individual circuli can no longer be traced with cer- 
tainty, for they are less distinct, much broken, anastomosed and 
closer together. This zone of faint, approximated and much 
broken circuli, when contrasted with the preceding and succeed- 
ing areas of strong, complete and widely spaced circuli stands 
out as a rather sharply defined band. This band may be traced 
around the whole scale and is perhaps better defined in the 
posterior field where it appears as a lighter zone with very little 
detail. 


To account for these structures, it is contended that the 
completed and comparatively widely separated circuli are 
formed during periods of rapid growth, the incomplete lines 
during periods of decreased growth, and the short, weak, much 
broken lines during periods when growth has nearly ceased. 
As the cessation or retardation of growth is thought to occur in 
the winter, the much broken area is accordingly designated as a 
winter-band or annulus (A). 


When the scale resumes its growth in the spring a complete 
circulus is again formed which in the process of uniting, as it 
were, the incomplete lines bends sharply at the antero-lateral 
ridge. This circulus is considered the limit of the annulus it 
encloses and is so employed in the measurements of scales. 


The twenty-five circuli of the second summer, in this scale, 
are much more widely separated than those of the first, which 
indicates a much more rapid growth during the former season. 


1923] Van Oosten: Scales of Whitefishes 383 


Of these twenty-five circuli the last five or six at the margin are 
incomplete, which indicates the occurrence of a retardation in 
growth. No approximation of the circuli is yet visible, nor is 
there apparent the area of weak and broken lines. A complete 
cessation of growth has not yet taken place. This conclusion 
appears to be reasonable as the specimen was caught in October 
preceding the period of low temperatures when growth is greatly 
retarded or ceases altogether. 


The foregoing account of the mode of formation of annuli 
is accepted by the majority of those investigators who make use 
of scales in determining the lengths and rate of growth of fishes. 


The application of the above hypothesis to the study of the 
life-histories of fishes is a simple matter. By enumerating the 
annuli on the scales, the age of the individual is determined in 
years. Thus the specimen whose scale (Fig. 137) illustrated the 
method of growth is found to be at the end of its second year. 
The length of an individual at the end of each successive year 
may also be ascertained from its scales. Given the total length 
of the fish and of one of its scales and the length of that part of 
this scale included in an annulus formed during a given year, 
the total length of the fish at the end of this given year may be 
computed by the following formula in which the third term is 
the unknown: 

length of scale formed at end of year X _ 
total length of scale 


length of fish at end of year X 
length of fish at the time of capture. 

Repeating this formula for each year of the fish’s life, the 
lengths attained at the end of the several years are calculated 
and by a simple subtraction the increments of growth for each 
year are determined. 


The soundness of the scale method of determining the length 
of a fish at successive years of its life and its annual growth 
increments depends on the validity of the following propositions : 

1. That the scales remain constant in number and identity 

throughout the life of the fish. 

2. That the annual increment in the length (or some other 

dimension which must then be used) of the scale main- 
tains throughout the life of the fish a constant ratio 
with the annual increment in body length. 


3. That the annuli are formed yearly and at the same time 
each year. 


384 Zoologica: N. Y. Zoological Society i Gees a 


Incidentally the following questions are raised, but the 
validity of the scale method of computation is not affected 
by them: 

4. Whether the annuli represent periods of retarded or 

arrested growth of the scale. 

5. Whether the growth of the fish in length is retarded or 

arrested at the time of formation of the annuli. 
6. What factors are responsible for the arrest or retarda- 
tion of growth in fish and scales. 


Considering now the first three propositions listed above, it 
is believed that the first two are fairly well established and that 
the last one forms the crux of the whole problem. If the age of 
a fish can be determined with certainty, the establishment of the 
validity of the third proposition becomes a comparatively easy 
matter in a group of fishes whose scales show growth rings. 
Indisputable evidence of a correlation between the number of 
annuli on the scales and that of the years of life of their bearer 
can only be obtained by observation on fish of known age in the 
field and in the laboratory. And the value of the results rises 
with the number of years for which this correlation is found 
to exist. 


An extended review of the literature is reserved for a later 
paper. Here I indicate briefly the chief differences of viewpoint. 


Both Hoffbauer (1898, 1899, 1901) and Walter (1901) 
believed that the age-hypothesis does not hold for carp older 
than four years. . Likewise Brown (1904) and Tims (1906) 
contradicting Thomson (1904) held the scale method entirely 
unreliable as applied to the Gadidae. Even Thomson concludes | 
from his experiment that a well-fed whiting may pass the winter 
without forming an annulus on its scales. Arwidsson (1910) 
concludes from his study of a series of salmon, 4 to 36 months 
old, that the completion of the first annulus does not occur at a 
definite time of the year nor at a definite age, but only at a 
definite length of the fish, viz., at 60 mm. Masterman (19138) 
asserts that it is a well known fact that the otoliths.or “ear- 
stones” often used for age-determinations cease growing in the 
plaice after 6 or 7 years, and that scales also are unreliable after 
the first 4 or 5 years, though the latter statement is questioned 
by Hutton (1914). Likewise Scott (1906) expresses the opinion 
that otoliths do not show the exact age of their possessors. Many 
other authors may be quoted as opposing the age-hypothesis, but 
an overwhelming majority assume the validity of the theory and 


apply it. 


1923] Van Oosten: Scales of Whitefishes 385 


Much diversity of opinion exists as to the relation between 
the formation of annuli and the growth of scales and body. The 
majority of students believe that the annuli are due to seasonal 
variation in body growth, that they correspond to retarded 
growth; but Cunningham’s (1905) observation and Cutler’s 
(1918) experiments contradict this view in part, while Taylor 
(1916) denies such a correlation entirely. 


Much controversy also obtains relative to the factors gov- 
erning the formation of annuli. According to Hoffbauer (1898, 
1899), Thomson (1904), Fraser (1917) and others food is the 
primary factor and not temperature. Taylor (1916) and Cutler 
(1918) conclude from their experiments that food is not the 
factor involved. Fraser (1917) holds that neither salinity nor 
density nor temperature has any factorial significance, while 
Cutler (1918) believes that temperature alone is causative. 
Rich (1920) refers to the factor as “a changed environment,” 
Jacot (1920) calls it “migration,” while in the case of some trout 
and salmon the later annuli correspond to a spawning and con- 
sequently are transformed into “spawning-marks.” 


Masterman (1913) wrote, “Experience shows that each 
species of fish must be investigated separately by the method 
best suited to it,” implying that the establishment of the validity 
of the hypothesis for one species does not necessarily make it 
applicable to other species of fish. 


The scales of the whitefish (C. clupeaformis) have never 
been critically studied. During the course of an extended investi- 
gation of the scales of the Coregonine fishes of the Great Lakes, 
the writer was fortunate in obtaining scales of this species of 
known age—nine years. This material forms the basis of an 
attempt to test the underlying assumptions of the scale method 
of computation as applied to this species. It also is believed to 
throw light on the relation between annuli and rate of growth; 
while the accompanying data permit a discussion of the environ- 
mental factors involved in annulus formation. 


Here I wish to present a brief description of the apparatus 
used for the measurement of scales, as my method differs from 
those ordinarily employed. The instrument is constructed on 
the principle of a photomicrographic apparatus in which the 
image is projected on the ground glass. The apparatus consists 
of a rectangular wooden frame, 14 inches square and 34 inches 
long. Into one end of the frame is fitted, flush with the exterior 
surface of the frame, a piece of ground glass, 12 inches square. 
A tapering bellows made of ordinary chart cloth painted black is 


386 Zoologica: N. Y. Zoological Society [lisa 


attached to the ground glass end of the frame. The bellows 
when fully stretched extends about three-fourths the length of 
the wooden frame. The tapered end of the bellows is attached 
to a small square wooden frame into which is tightly fitted a 
wooden block in the center of which a hole large enough for the 
insertion of the microscope tube is bored. When the apparatus 
is used in the vertical position the microscope is simply placed 
beneath and extended into the bellows. It is much easier to use 
this instrument in the horizontal position. In this case the micro- 
scope stand is attached to a board at the base of the wooden 
frame (the end opposite that into which the ground glass is 
fitted) and the microscope tube drawn into the horizontal posi- 
tion. The open base of the frame is then covered with a sheet of 
black paper into which a hole is cut so as to allow the light to 
enter the condenser of the microscope. A special Bausch and 
Lomb lamp with a 108-Watt bulb furnishes the illumination and 
is placed about two feet from the base of the frame. A special 
aspherical condenser accompanying the special lamp is used in 
the place of the ordinary condenser. The light concentrated 
upon the hole in the black paper passes through the condenser, 
microscope tube and bellows, and projects the scale upon the 
ground glass. 


A mechanical stage is always used. To each adjustment 
button of the mechanical stage is attached, by means of a uni- 
versal joint cut from a piece of tin, slender wooden rods which 
extend a little beyond the ground glass end of the frame. In 
a similar way another rod is attached to the coarse adjustment 
screw of the microscope. By means of these rods the scale can 
be moved into place and properly focused from the ground glass 
end of the frame. The projected scale is measured with an 
accurate wooden or transparent millimeter rule which is held in 
place against the ground glass by two strips of steel, four of 
which are screwed on the wooden frame, one at each corner. To 
facilitate the counting of the circuli of each scale an ordinary 
reading glass is used. The whole apparatus is placed upon a 
long table and may be covered with a black cloth. No dark room 
is required as the lamp is strong enough to project a clear image 
on the ground glass in a room illuminated by electric lights; 
during the day the curtains of the room must be drawn. 


The advantages of this method of scale reading over those 
which use the camera lucida, ocular micrometer or micrometer 
eyepiece are many. In the first method the scales can be highly 
magnified without any part being lost to view as is the case in 
the microscope tube; the circuli and the distances between the 


1923] Van Oosten: Scales of Whitefishes 387 


annuli of such highly magnified scales can be more accurately, 
more quickly, and more easily enumerated and measured; and, 
if the illumination is properly adjusted, scale work can be done 
with much less straining of the eyes. 


When the apparatus is used in the horizontal position it is 
necessary that the scales be mounted in a stiff medium. Each 
scale is therefore cleaned in water with a small bristle brush and 
mounted in a medium of glycerine to which has been added 
filtered gelatine and a little carbolic acid. The glycerine and 
gelatine are mixed in such proportion that the solution will 
stiffen immediately upon cooling. When in this medium, the 
scales can be stored as permanent mounts and can also be photo- 
graphed. The photomicrographs (Figs. 137-142) are of scales 
mounted in a gelatine-glycerine solution. 


I wish to express my appreciation to Dr. Charles H. Town- 
send, the Director, and to Miss Ida M. Mellen, the secretary and 
scientific assistant of the New York Aquarium, through whose 
kindness and efficient cooperation I have been able to obtain the 
whitefishes and scales for this work. These whitefishes, the only 
ones known to have been reared in captivity, form a valuable 
exhibition at the Aquarium so that it has been no small sacrifice 
to part with even a few of them. I am also indebted to Dr. 
Walter Koelz of the U. S. Bureau of Fisheries who has kindly 
given me access to his field data and manuscript on the Core- 
gonine fishes of Lake Huron. I would further express my obli- 
gations to Prof. Jacob Reighard, who read the manuscript and 
generously gave assistance in the course of the work. To Mrs. 
Alvina M. Woodford, of the University of Michigan Library, I 
am indebted for many valuable suggestions relative to the pho- 
tographing of the scales. 


NEW YORK AQUARIUM WHITEFISH SCALES 
ANNULI AND NUMBER OF WINTERS OF LIFE 


Twenty-seven preserved specimens of the Aquarium white- 
fish, hatched January, 1913, were received. These had died (or 
had been killed) at intervals between August 13, 1920, and 
January 3, 1922, as shown in Table I—a period of sixteen months. 
The fish received had died (been killed) during every month of 
the year except November. The lengths of each specimen at 
the time of death is shown in column K of Table I and is fol- 
lowed by the formula which indicates the sex and the condition 
of the sex organs (see p. 403). The remaining entries in Table I 
are calculated values and will be referred to in another place. 


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388 


(i it} Nid I \\ Ae 
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~ = 

ANI 


Ze ae 
yy 


SS 


AR PR 


‘Ventral - 


Fic. 1387. Typical scale of Lake Huron Whitefish Geese clupea- 
formis Mitchill) from East Tawas, Michigan. Length of fish, 197 mm., 
captured October 22, 1917. L-L, length of scale; F, focus; C, ‘circuli: A, 
annulus of first winter; 


AR, antero-lateral ridges; ARs postero- lateral 
ridges; Dorsal, Ventral, Anterior, Caudal border and area. X-2 


Pee Vol. if, No. 17 
—Face Page 388 


“hae 4 ie i HY, Hof) 
JHE MEY Wi a 4 
it Li Wy Y WY Yj 
i i i i yp yy Ue Ui 74 

1 5 iy H Uf Z 
2 UT 
3 TM | \ 
4 SU | 


Fic. 138. Scale of New York Aquarium Whitefish (C. clupeaformis) 
hatched January, 1913; killed December 1, 1920. U. of M. Museum No. 
54507; Male, 265 mm. long, 7 years, 10 months old. Scale shows 7 com- 
pleted annuli and a marginal growth. X-22. 


Zoologica Vel. II, No. 17 
Fig. 138—Pages 388-389 


v 
(ie 
[ey 

ty 


Uf fy Vy 
(dG: 


SS 
=< 
Sa 
J aN i 
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Fic. 139. Scale of New York Aquarium Whitefish (C. clupeaformis) 
hatched January, 1913; killed April 28, 1921. U. of M. Museum No. 54513; 
Male, 278 mm. long, 8 years, 3 months old. Scale shows 8 completed annuli, 
the eighth at the margin. X-18. 


Zoologica Vol. II, No. 17 
Fig. 139—Pages 388-389 


A 
HK 


Ri ifs Hy 
y i | Woe A 
=D 


3 Kaa 
5 Nad ‘ i 
TA AML 


Fic. 140. Scale of New York Aquarium Whitefish (C. clupeaformis) 
hatched January, 1913; died July 13, 1921. U. of M. Museum, No. 54516; 
Female, 282 mm. long, 8 years, 5% months old. Scale shows 8 completed 
annuli and a marginal growth. X-18. 


Zooiogica Vel. II, No. 17 
Fig. 140—Pages 388-389 


SSW TERIA NMOL 
CE LN 
Be RS ALARA CN TA ii 
2 ee AN \\y INU 

LCE RY 
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SAAN 
8 


ANNE 
an wi 


SE: 
Za 
z Lo 
ree = 
= ao 
LIL 


Fic. 141. Scale of New York Aquarium Whitefish (C. clupeaformis) 
hatched January, 1913; died August 3, 1921. U. of M. Museum No. 54523; 
Male, 339 mm. long, 8 years, 6 months old. Scale shows 8 completed annuli 
and a marginal growth. X-16. 


Zoologica Vol. II, No. 17 
Fig. 141—Pages 388-389 


} 
| 


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Ws) 
i I Ai 
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i 
Uh 


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ah 


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rUbhay NU \ 
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Fic. 142. Scale of New York Aquarium Whitefish (C. clupeaformis) 


hatched January, 1913; died January 38, 1922. U. of M. Museum No. 54531; 
Female, 347 mm. long, 9 


years old. Scale shows 8 completed annuli and 
a marginal growth. X-19. 


Zoologica Vel. II, No. 17 
Fig. 142—Face Page 389 


1923] Van Oosten: Scales of Whitefishes 389 


Figs. 138-142 are photomicrographs of scales taken from 
Aquarium fish killed at different ages. On each photograph refer- 
ence lines have been drawn which indicate the positions of the 
annuli of the different years. On comparing the scales of these 
aquarium whitefishes with those of the wild whitefish, it may 
be seen that the former are much stunted in growth; their circuli 
are more irregular and crowded together, while their annuli, in 
some instances, nearly come into contact in the posterior or 
exposed area of the scale. In some scales the annuli are difficult 
to observe, but by the use of various magnifications and by the 
manipulation of the light source they can be determined. In most 
of the fish several scales are necessary for an age-determination. 
I gradually discovered, however, that those aquarium scales 
whose caudal area is longer than its anterior area and whose 
form approaches the elliptical possess annuli that can be more 
readily determined than those of scales without these characters. 
Such scales were usually obtained from the area between the 
pectoral fin and the lateral line. Those photographed and em- 
ployed in this study were taken from the left side of the body. 


The first specimen died August 13, 1920, at the age of seven 
years and seven months. Its scales possessed seven annuli with 
a small amount of marginal growth. The number of annuli, 
thus, corresponded with the number of years of life of the indi- 
vidual. The next seven specimens received (54506 to 54512, 
Table I), which ranged from seven years and nine months to 
eight years and two months in age, also possessed scales with 
seven completed annuli and various amounts of marginal 
growth. Fig. 138 represents a photograph of a scale from the 
specimen killed December 1, 1920, and shows seven annuli with 
the eighth year increment at the margin. The eighth annulus is 
completed in April. Fig. 139, a photograph of a scale from the 
April specimen, shows the eighth annulus situated at the margin. 
In the remaining eighteen specimens (54514 to 54531, Table I), 
which ranged from eight years and four months to nine years 
in age, the eighth annulus is entirely removed from the margin 
and surrounded with various amounts of the ninth year incre- 
ment. Figures 140, 141 and 142 are photographs of scales from 
specimens that died July 13, 1921, August 3, 1921, and January 
3, 1922, respectively. Each figure shows eight annuli and, 
presumably, a completed ninth year’s growth. 


It thus appears (1) that the specimens received from 
August, 1920, to March, 1921, which represent ages from seven 
years and seven months to eight years and two months possessed 


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390 


1923] Van Oosten: Scales of Whitefishes a9L 


scales with seven annuli and various amounts of marginal growth 
and therefore belong to the same growth year, the eighth, even 
though the three fish received from February to March actually 
were in their ninth year of life; (2) that those received from 
April, 1921, to January, 1922, which vary from eight years and 
four months to nine years in age possessed scales with eight 
annuli and different amounts of marginal growth. 

The annuli are thus actually proved to be of the same num- 
ber as that of the winters of the fish’s life, if we exclude the first 
one in which the fish was hatched. 


MARGINAL GROWTH AND TIME OF FORMATION OF ANNULI 


Two specimens of the aquarium fish were segregated in the 
New York Aquarium and kept living with the purpose of taking 
scales from them at monthly intervals in order to follow the sea- 
sonal changes in the scales from November until June. Owing 
to deaths a total of six fish was employed during this interim, 
three of which lived for periods of two, six and seven months. 
In Table II these fish are designated by names indicating slight 
.physical peculiarities and by their museum numbers. The 
names should not be taken as indicating that the fish were notably 
deformed. They were the smaller, poorer fish, less desirable for 
exhibition purposes. Table II shows the average total length 
in mm. of the magnified scales taken from these fish in different 
months, the average total length in mm. of the marginal growth, 
the percentage of its length in the antero-posterior diameter of 
that part of the scale included in the last annulus, the difference 
in the percentages of successive months and the number of scales 
upon which each average is based. The percentages in this 
table are, however, only approximately correct as they vary with 
the areas on the body of the fish as well as with the scales taken 
from the same area. However, as all the scales, except those 
removed from the dead fish, have been taken from the same area, 
the variability of the scale values has been reduced to a mini- 
mum so that they may be used with confidence in drawing cer- 
tain conclusions. 

The discrepancy in the fall or winter percentages (which 
seem to show scale absorption) of Crooked-back and Open-gill 
are presumably due, then, to the variability of their scales. The 
small difference (+0.4) between the two winter percentages 
(8.6 and 9.0) of Double-crook which represent a period of two 
months, likewise may be looked upon as due to this same varia- 
bility and thus can have no significance. Presumably, then, the 
percentages in column v8/V7 or v9/V8 remain constant for 


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392 


1923] Van Oosten: Scales of Whitefishes 398 


each fish during the fall and winter, 7. e., the marginal growth 
of the scales is arrested during the period from October to 
March. Marginal growth is resumed sometime in April (or 
March?). On April 26, 1921, the new marginal growth of 
Double-crook showed a percentage of 2.2, which value was in- 
creased to 7.4% on July 13, two and one-half months later. 
Similarly, Open-gill showed a new marginal growth of 7 .0% on 
May 26, 1921, which percentage was increased to about 9.0 on 
June 25, one month later. The percentages of Double-crook and 
Open-gill are thus entirely consistent with and comparable to 
each other from April on and show that rapid scale growth is 
resumed sometime in April (or March?) and is continued at 
least until July. 


In order to show approximately by comparison how much of 
the new year’s growth was completed by Double-crook and Open- 
gill at the time of death, I computed percentages, similar to those 
of Table II for the preserved specimens listed in Table I. Table 
III shows in column v7/V6 the percentage of the scale increment 
of the seventh year in the total length of the scale included in 
the sixth annulus, in column v8/V7 the percentage of the scale 
increment of the eighth year in the total scale length included 
in the seventh annulus, and in column v8/V7 or v9/V8 the per- 
centage of the incomplete marginal growth of the eighth or ninth 
year in the total scale length included in the last completed 
annulus (7th. or 8th.). The average for each series is given at 
the bottom of each column. 


From this table it may be seen that the average percentage 
of the completed seventh year scale increment in the total scale 
length of the sixth year for 27 specimens is 8.9 and somewhat 
less, as is to be expected, for a similar percentage for the next 
year (7.6) for 19 fish. When these averages (8.9 and 7.6), 
which represent the completed seventh and eighth growth years 
are compared with the percentages of Double-crook and Open- 
gill (7.4 and 9.0) and when it is remembered that the percentage 
for the ninth year may reasonably be expected to be somewhat 
less than that for the eighth, it may safely be assumed that the 
scales of the segregated fish have just about completed their 
ninth year’s growth and certainly would have done so by August 
or September. 


If we now assume that August closes the period of scale 
growth and compute the average of the percentages of the incom- 
plete ninth year from August to January and compare this 
average with those of the two preceding completed years and 


394 Zoologica: N. Y. Zoological Society [Tiong 


with the percentages of the two segregated fish we may obtain 
a criterion which ascertains roughly the probability of the cor- 
rectness of our assumption relative to the time of the cessation 
of scale growth. Table III shows that such a ninth year average 
is 6.5%, which compares favorably with those of the two pre- 
ceding completed years (8.9 and 7.6), and may therefore be 
considered as representing the average of a completed ninth 
year. The ninth year average (6.5) thus suggests the comple- 
tion of scale growth by August not only in the fish of Table II 
(with 9th year percentages of 7.4 and 9.0), but also in those of 
Table I. Again, when the average of the percentages of the 
incomplete eighth year from August to March is compared with 
that of the completed eighth year, it is found that the former 
(5.8) compares fairly well with the latter (7.6) when it is 
remembered that the fish sacrificed first (included in the 5.8 
average) were the poorer and less valuable specimens. Thus 
again scale growth may presumably be considered complete by 
August. Also, Tables II and III show that the percentages of 
the incomplete eighth and ninth years show no consistent increase 
from August and October to April, while those of the ninth year 
from April to June or July do. 


In the light of the preceding discussion it now appears rea- 
sonable to accept the interpretation presented on page 391 rela- 
tive to the constancy of the fall and winter percentages given in 
Table II. 


Table III, column v8/V7 or v9/V8, further corroborates 
Table II and shows that marginal growth is resumed in April. 
All the fish killed after April, 1921, completed the new annulus 
and showed various amounts of new marginal growth on their 
scales. 


The percentages of Table II show, then, (1) that there was 
no marginal growth present in November and December of 1920, 
and in January, February and March(?) of 1921, and, (2) that 
a new annulus was recognizable in April, 1921, and was cor- 
related with a resumption in scale growth. The percentages 
of Table III corroborate the conclusions based on Table II, and 
in addition show in conjunction with Table II that marginal 
growth was presumably arrested by August in 1920 and 1921, 
and certainly by September. The data of both tables (II and 
III) therefore prove that the annulus is a winter-mark due to a 
retardation or cessation of scale growth and is completed upon 
the resumption of rapid scale growth in the spring of the year. 


19235] Van Oosten: Scales of Whitefishes 395 


Table IV—Showing for 76 Alpena whitefishes collected September, 
1917, the relation of the average length of the diameter (v), anterior (ac) 
and posterior (pc) radius of the scale to the average body length (K), all 
lengths expressed in mm., for fish in years III to VII inclusive.* 


Year III IV V VII 

1 ES ve ge 269 (47) 315 (14) 3852 (9) 456 (6) 
\/ ee ea ee ae as Srhsy (Zh), 6.75 (14) 7.49 (9) 9.58 (6) 
ONC. 2 sitar ft Lea etapa ae 3.19 (47) 3.86 (14) 4.34(9) 5.80 (6) 
IC eeee a Eee 2.56 (47) 2.89 (14) 38.15 (9) 3.78 (6) 
Reva eto Po Oe tas 46.78 46.67 46.99 47.60 
eae Ree 84.33 81.61 81.11 78.62 
IGA OY Glee Sea ne Ue See 105.08 108.99 111.75 120.63 


“Numbers in parentheses following averages indicate the number of 
specimens employed; the sixth age-group contains only one specimen, there- 


fore, omitted. 
CORRELATION BETWEEN ANNUAL GROWTH IN LENGTH 
OF BODY AND SCALES 


It now remains to examine for the whitefish the correlation 
between the annual increment in length (or other dimension) of 
scales and length of body. Had it been possible to measure the 
body lengths of the living aquarium whitefish accurately at the 
time of the removal of their scales, this correlation could have 
been established by direct observation. Obviously, Table I does 
not afford material for this purpose as the number of specimens 
received each month is too small to warrant valid averages of 
monthly growth increments in body and scales. The’ available 
aquarium whitefish therefore cannot show the proportionate 
growth of body and scale. 


Wild whitefishes may be used to show this correlation. In 
this case it is necessary that a large amount of strictly homo- 
geneous material be used, 7. e., the fish of the several age-groups 
must all belong to the same race and have similar rates of annual 
growth increments, and only scales from corresponding body 
areas must be employed. These requisites necessitate the acqui- 
sition of a large collection of fish taken at the same time and 
at the same locality. At present no such whitefish material is 
available. There are, however, at hand, series of body and scale 
length measurements of a small collection (76 fish) of Lake 
Huron whitefishes taken September, 1917, at Alpena, Mich. A 
summary of their data is given in Table IV. 


In row K, Table IV, is shown the average length in mm. of 
the fish of each age-group, the age-group referring to the year 
of life in which the fish were captured. The number of specimens 
in each age-group is shown in parenthesis. In row (v) is given 
the average length in mm. of the scale diameters of the fish of 
each age-group. In rows (ac) and (pc) the same averages are 


396 Zoologica: N. Y. Zoological Society Be yi 


Table V—Showing for each year for Alpena whitefish (Table IV) in 
the seventh year the average length in mm. calculated from the diameter 
(v), anterior (ac) and posterior (pc) radius of scales (Table IV), and the 
difference between the calculated averages and those of the age-groups 
obtained from actual measurements (K). 


III IV V Vil 
Kegan in inate ik HR Pon CUS 269 315 352 456 
Calculated (K) 
TOW CVA) ees et ec ee 274 321 357 
Calculated (K) 
nih e(O) cabo (eNO) pens Cate Pees <a Ie bie oe 251 3038 341 
Caleulated (K) 
from" (pe) ba RC Uma 8 hie 3809 349 380 
Difference 
(CKO) ice (KGehT Orava) eae ae een eee +5 +6 apd 
Difference 
(QQ) 15 GER Sanctoy ana EXO) oc —18 ——12 —] fi 
Difference 
(KS) Ree CORSET Om ap) eos eee +40 +34 +28 


given for the anterior and posterior radius of the scale respec- 
tively. In the last three rows are shown the body-scale ratios for 
each age-group based on the diameter (v), anterior (ac) and 
posterior (pc) radius respectively. 


The number of fish in each age-group is not as large as one 
could wish, but may be sufficiently large to show roughly 
the relation of the length of the various scale dimensions to body 
length. It may then be seen that the K/v ratio is about the same 
for the third and fourth age-group and rises slightly in the fifth 
and again in the seventh. This means that the diameter increases 
in length in a simple proportion to the increase in the length of 
the body during the fourth year and increases at a slightly 
slower rate relative to the body in the fifth and seventh years 
and presumably also in the sixth for which no values can be 
given. The K/ac ratio is found to decrease with age, while the 
K/pe ratio increases with age. This means that the anterior 
radius of the scale grows faster relatively than the body with 
age, whereas the posterior radius grows more slowly with age. 
None of the measured scale dimensions therefore grow strictly 
proportionate to the body. 


In order to show roughly in a practical way which dimen- - 
sion most nearly acquires this proportionate growth and fur- 
nishes the most accurate estimated length values, I calculated the 
average length for each year of the fish in the seventh year, using 
the average scale dimensions of Table IV. The estimated lengths 
are shown in Table V. From this table it may be seen that there 
is a high degree of correspondence between the lengths eal- 
culated from the diameter and the actual lengths, and that the 


1923] Van Oosten: Scales of Whitefishes B97 


former are somewhat higher than the latter. The lengths cal- 
culated from the anterior radius are lower, while those calculated 
from the posterior radius are much higher than the actual 
lengths. It is, however, realized that these calculated and actual 
values may not be strictly comparable as the two series of values 
represent different year-classes which may have varied con- 
siderably in their rate of growth. Strictly, series of fish of the 
same year-class collected in the same season of different years 
and at the same locality are required for an absolutely valid 
check on calculated values. 


Table V also shows that the calculated values from diameters 
are consistently found between those based on the radii. This 
was also found to be true for the individual fish. Experience. 
has shown that in practically every species of fish whose scales 
were studied the calculated values based on radii (no other 
dimension is ever used) and checked with the actual values were 
always found to be too low for the early years of life. The 
Coregonines prove to be no exception. In the light of past ex- 
perience therefore, and in view of the fact that diameter meas- 
urements of whitefish scales raise the calculated values, espe- 
cially those for the earlier years, above those based on the an- 
terior radius, it may be deemed advisable to test the various 
dimensions of the scales, where possible, before undertaking any 
extensive scale work for a species. 


Tables IV and V show for the whitefish (1) that the 
diameter, anterior and posterior radius of scales increase in 
length at very different rates with respect to the rate of increase 
in the length of the body and that consequently the calculated 
length values based on the different scale dimensions vary sig- 
nificantly, (2) that none of the three scale dimensions consid- 
ered increase in length at a rate strictly proportionate to the 
rate of increase in length of the fish, (3) that, presumably, the 
diameter of the scale increases in length at a rate more nearly 
proportionate to the rate of increase in body length than either 
the anterior or posterior radius and therefore should be used 
for the calculations of length values, and (4) that the calculated 
values based on the diameter always lie between those based on 
the anterior and posterior radii. 


LIFE HISTORY OF THE AQUARIUM WHITEFISH AND THE 
FACTORS OF ANNULI FORMATION 
In view of the great diversity of opinion among investiga- 
tors, and in view of the disagreement and contradiction of the 
conclusions reached by various authors from laboratory experi- 


398 Zoologica: N. Y. Zoological Society [rEny 


ments and field observations relative to the factors responsible 
for the formation of annuli, it was deemed advisable to obtain 
as complete a life history as possible of the New York Aquarium 
whitefishes and thus, perhaps, determine to a certain degree the 
relative significance of each environmental factor in the forma- 
tion of annuli. It is realized that these life history data are not 
equivalent to those of carefully planned and executed series of 
experiments in which only one factor of the environment is 
altered at one time and the results checked with those of a con- 
trol. Yet, it will be seen that experimental requirements are 
partly fulfilled for the fish were regularly supplied with food in 
amounts controlled by their appetites, while the temperatures 
varied from those of summer to those of winter and vice versa. 
And as the period of rapid scale growth and the time of the 
formation of an annulus are known the effect upon the scales 
of a change in a factor may be approximately ascertained. I 
am indebted to Miss Mellen and to Mr. Robert J. Lanier, who 
reared the whitefishes, for most of the life history data. I alone 
am responsible for the conclusions derived from them. 


It has now been proved (p. 394) that the formation and com- 
pletion of an annulus is dependent upon the retardation or cessa- 
tion and resumption of scale growth, and that scale and body 
growth are closely correlated; therefore, any factor that can 
affect the growth rate of the body may have primary signifi- 
cance in the formation of an annulus. But to hold such a factor 
responsible it must be established that a change in this factor 
occurred previous to or synchronously with the change in the 
rate of growth and that no resumption of rapid scale or body 
growth can occur until the change in the factor is reversed or 
its effectiveness lost. The primary factors may be different from 
year to year or even from season to season in the same year. 


(a) Foop 


The Aquarium whitefish were fed in about the same way 
throughout the year. Miss Mellen writes: “I have consulted 
our superintendent (Mr. Robert J. Lanier), who reared the 
whitefishes born in 1913, as to their food since hatching, and 
find that in early infancy they had the advantage of some live 
food, receiving first herring roe, next a few mosquito larvae, 
third, the fry of pike perch, which happened to be hatching at 
just the right time, although all would not take this food, as, 
unlike most fishes, they do not normally eat their own kind, and 
lastly, minced beef-heart. Feedings were very frequent during 
the first couple of vears, after which they were fed daily on beef- 
heart. From 1915 to 1918 inclusive they were fed exclusively on 
beef-heart three times a week; and it is only since 1919 that they 


1923 | Van Oosten: Scales of Whitefishes 399 


have been fed with beef-heart exclusively three times a week in 
summer and beef-heart once, clam twice a week in winter.” 

The amount of food, however, actually consumed by the fish 
varied somewhat with the seasons. In the earlier years when 
the fish were fed daily, Mr. Lanier gave them less food during 
January, February and March in spite of the fact that they were 
always ready to eat. During their third year an apparent change 
in their appetites occurred, so that thereafter they were fed 
only three times a week. Since this year (1915) it was also 
noticed that the fish ‘“‘did not eat quite as much during January, 
February and March,” though the number of feedings remained 
constant throughout the year. 


It is obvious that since the character of the food and the 
number of feedings remained constant summer and winter for 
several consecutive years these factors could not have altered 
the rate of scale or body growth. The amount of food, however, 
offered or taken by the fish, did vary with the season. How 
much the amounts varied is not known. In the fall of 1920 
this change in the amount of food could not have had any sig- 
nificance inasmuch as scale growth was arrested before Janu- 
ary, 1921, when this factor was first altered. In April, 1921, 
the rations were increased for the same reasons that they were 
decreased in January, viz.—a change in the appetites of the 
fish. It was previously shown that scale growth was resumed 
in March (?) or April (p. 393). It thus appears that food and 
growth are correlated in April, 1921, but in this case the former 
is presumably only of secondary importance. As nothing defi- 
nite is known about the distribution of growth in the other years 
of life of these fish, the relative significance of food as a factor 
in these years can only be conjectured and therefore requires no 
discussion here. 


(b) TEMPERATURE 


Table VI shows the monthly and yearly mean temperatures 
in degrees, Fahrenheit, of the fresh water entering the New 
York Aquarium, from 1913 to 1920, inclusive. As soon as the 
water reached a temperature of about 60°F (15.6°C) in the sum- 
mer, it was refrigerated and maintained at a temperature which 
varied from 54° to 57°F and averaged about 55°F (12.8°C), 
except during the summer of 1921, when the range was changed 
to 50° and 54°F, and the average reduced to about 52°F 
(11.1°C). The months in which the refrigerating plant was 
started and stopped are indicated by the letters r and s respect- 
ively in Table VI. In Fig. 143 are plotted a curve (T) based on 
the average monthly temperatures of Table VI and growth 
curves of the scales of Double-crook (D) and Open-gill (O) 


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400 


Van Oosten: Scales of Whitefishes 401 


1923] 


AN; FEB, MA 


O-curve 


which represent the total scale growth 
f Double- 


emperatures 


D-curve based on the per- 


hown in Table VI. 


LOZ 


? 


n between the rate of growth of the 
crook (D) and the temperature (T) of 


to J une 
which represent the total scale growth o 


’ 


refrigerated water s 


based on the percentages of Table II 
1921 


curve based on the average monthly t 


’ 


g the relatio 


) and Double 


T 
ch 


, 


Showin 
-gill (O 
gill from Mar 


1S RY 


scales of Open 
of the refrigerated and non 


centages of Table II 


the aquarium water. 
crook from March 


of Open 


1921. 


to July, 


, 


1921 
based on the percentages of Table II 


renewed growth of the 


’ 


o 
He 
SiS) 

(o) 
=e 
g b 

o 
Ae 
fe) 
= 

rs 
om 
Sr 
cae 

= 
a 
mo 
= 
mo 

oO 

— 

oy 

(9) 

fe) 

— 

(5) 

{S) 

M 


curves are only approximately correct as the scales of each fish 


were taken in alternate months. 


No growth occurred during 

February and March, 

he time of 

the greatest rise in temperature in April and May. The maxi- 


Fig. 148 shows that the curves of scale growth adhere very 
however, continued, at least in Open-gill, throughout June. Scale 


refrigerated and maintained throughout the summer at an 
average constant temperature of about 52°F. Scale growth, 


and the greatest increase in growth took place at t 
mum temperatures were reached in May when the water was 


the period of low temperatures in January, 


closely to the curve of temperature. 


402 Zoologica: N. Y. Zoological Society [Itsy 


growth therefore was not arrested by the temperatures of the 
refrigerated water. As these temperatures do not represent 
those of the water in the whitefish tanks, they may be a little 
too low, especially those of the non-refrigerated water. This, 
however, would not alter the general appearance of the tempera- 
ture curve nor the conclusions based on it.. 


It is thus seen that scale growth and the temperature of the 
water are correlated in the spring of 1921, the latter presum- 
ably having primary significance. That a low temperature is 
here a primary factor in growth seems corroborated by the fact 
that a decrease in the amount of food consumed occurred syn- 
chronously with a decrease in temperature and an increase in 
food consumption with an increase in temperature. The effect 
of low temperatures upon the metabolism of the body is well 
stated by Dr. Fulton (1904), who writes (p. 170): ‘‘Tempera- 
ture is active in modifying the rate of growth by acting directly 
upon the metabolism of the fish and also by affecting the rapidity 
of digestion. In very cold water the fishes give up feeding alto- 
gether, because the ferments upon which digestion depends do 
not act, or act very slowly, at low temperatures, and in fishes, 
as in other animals, appetite waits on digestion, and this is, on 
the other hand, correlated with the metabolism in the tissues. 
It has been shown by Krukenberg that the pepsine or analogous 
body in the stomach of fish acts as well at 20°C as at 40°C, at 
which, among mammals, digestion is most active, and that the 
rapidity of its action is closely related to temperature; and 
Knauthe and Zuntz have shown that the same thing applies to 
the metabolism in fish, the vital activities being more active in 
the higher temperature, as shown by the excretion of carbonic- 
acid gas and other products of metabolism.” 


But, as both food and temperature remained practically 
constant during the summer until November, what factor must 
then be held responsible for the arrest in scale growth in August 
or September? As the aquarium water is well aerated its gaseous 
content cannot be held responsible; nor could any probable 
changes in its mineral or salt content account for the decrease 
in growth. The only remaining variable is sexual maturity. 


(c) SEXUAL MATURITY AND SPAWNING 

Before attempting to describe the conditions of the ovaries 
and testes of the various New York Aquarium whitefishes at the 
time of death, it seems advisable to describe first the normal 
natural conditions of the sex organs as found in wild whitefishes 
of various ages. The following description is based on many 
specimens of Lake Huron whitefishes. 

In order to determine the sex of immature wild whitefishes 
the sex organs must be closely scrutinized often with the aid of 


1923] Van Oosten: Scales of Whitefishes 403 


a magnifying lens or microscope. This difficulty of determina- 
tion is due to the fact that the ovaries and testes are quite 
Similar in appearance in the very young whitefishes. Both con- 
sist of narrow, thin, flat strands of soft, whitish material and 
extend from the anterior to the posterior end of the body cavity 
along its dorsal wall. In the larger immature fish the two kinds 
of sex organs can be distinguished by their structure. When an 
ovary is picked up and stretched transverse folds or layers 
become evident, while the testis under like treatment appears 
to be a compact homogeneous structure. In still older fish color 
may also be critical; the ovary becomes yellowish, while the 
testis retains its whitish color. 


In a maturing female the ovary gradually increases in size, 
the enlargement beginning at the anterior end of the body cavity. 
Minute round yellowish eggs appear in the ovary folds. At the 
time of spawning the ovaries have usually enlarged to such 
extent as to distend and fill the entire body cavity. The con- 
dition of the ovary and the size of the eggs thus indicate the 
stage of sexual maturity in the female. A ripe egg is about 
3 mm. in diameter. The maturing testis also increases in size, 
the enlargement commencing at the anterior end of the body 
cavity. As the testes grow they extend further into the abdom- 
inal cavity and increase in width and thickness. At the time of 
spawning they nearly fill the body cavity, but usually do not 
distend it, as do the ovaries. The size of the testes is then a 
rough index to sexual maturity in the male. When the eggs 
and sperm are ripe they are easily pressed out of the body. The 
sex organs of a spent fish are soft and flaccid. This condition 
is more easily determined for females than for males, and even 
among the former doubtful cases arise, the magnitude of the 
doubt depending partly, I presume, upon the length of the 
ee between the spawning and the date of the capture of 
the fish. 


In describing the various conditions of the sex organs of the 
New York Aquarium whitefishes I devised a number of phases, 
each phase indicating rather definite conditions. As the indi- 
viduals represent nearly every month of the year the conditions 
of their sex organs intergrade more or less imperceptibly, and 
it is therefore impossible always to refer a specimen to one 
phase; parts of different phases may be found in one individual. 
The ovaries alone permit of a rather definite classification and 
they only must be considered reliable in a discussion. 

The various stages of development® are considered under 
five principal phases designated by the letters A, B, C, D and E 
as follows: 


3These descriptions are based on the sex organs of the Aquarium fish 
preserved in 5% formalin and later transferred to 70% ethyl alcohol. 


1923] 


Zoologica: N. Y. Zoological Society 404 


:@—anterior 14 of ovary enlarged; eggs microscopic or 


very small in size. : 
é—anterior 1/4, of testis enlarged into a flat, white gland; 
remainder transparent. 


:@—anterior 14 of ovary enlarged; ovary flat, rigid, 34 


in. at its widest; eggs of year are round, whitish, 

easily visible to naked eye, 1 mm. or less in diameter. 
g—anterior 14 of testis enlarged; compact gland thicker 

and wider than in (E), about 14 in. at its widest. 


: 9? —entire ovary enlarged; eggs 1-2 mm. in diameter. 


é—anterior 34, and posterior end of testis enlarged, 4 
in. at its widest. 


:9—compact ovaries fill body cavity; yellowish eggs 


nearly ripe, 2-3 mm. in diameter; fish about ready to 
spawn; body may be pearled. 

gé—hard testes fill body cavity; fish about ready to 
spawn; body may be pearled. 


:?—matured eggs retained and in process of absorption. 


Coincident with this condition is that of (EF) or (D), 

i. e., eggs of the year are evident. Ovaries may be 

soft, flaccid; retained eggs found either in ovary, in 

body cavity or in both; eggs average 3 mm. in 
diameter when round and smooth; eggs are found in 
various conditions, several of which may be found 
in one individual; these conditions are designated by 

number under (A): 

1. Eggs hard, round, smooth; each egg partly turned 
a brownish yellow. 

2. Eggs wrinkled, indented, of brownish yellow 
color; float in water. 

3. Eggs wrinkled, indented, peripheral portion of 
each egg of a dull or dirty transparency; float in 
water. 

4. Eggs flattened, crushed, sometimes a little brown 
color left; contents absorbed. 

5. Eggs with dark reddish color, and a solid, glassy 
appearance; filled with minute oil globules; eggs 
about 1 mm. in diameter and scattered among 
the eggs of the year in a soft ovary; glassy eggs 
show no evidence of absorption. 

6. Ovary soft, flaccid with many eggs of the year, 
but no retained eggs evident. 

¢ —testes smaller than in (B) ; flabby, 7. e., fish spent, or 
testes compact, of reddish color with sex products 
seemingly retained and undergoing absorption; body 
may be pearled. 


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405 


406 Zoologica: N. Y. Zoological Society BO Se 


The distribution of these phases in time among the aqua- 
rium whitefishes is indicated in Table VII. The number in 
parenthesis following each phase refers to the day of the month 
on which the specimen died. Each such number represents one 
individual. The minus or plus sign following a letter indicates 
an early or a late phase respectively. Two or more phases in 
one individual are designated by a combination of the proper 
letters. The phase of each specimen is also shown in Table I. 


Table VII shows (1) that no immature female whitefish 
were received, (2) that two females were ready to spawn in 
December and January, (8) that retained eggs were present 
from December to August (no females were received in Feb- 
ruary, April, October and November), but that after January all 
the retained eggs, the glassy ones excepted, were undergoing 
absorption or disintegration, (4) that the eggs of the year, 
whether found with the retained eggs or not, were in the same 
phase of development from January to September, those of fish 
received July and later being a little more advanced than those 
of females received before July, and (5) that the ovaries of the 
specimens that died July 13 and September 18 were soft, but 
did not contain wrinkled eggs. 


From these facts we conclude, (1) that the Aquarium 
whitefish were sexually mature in both their eighth and ninth 
years, (2) that in most fish spawning conditions were present 
in December and January and thus that the sex products of the 
Aquarium whitefish ripen later in the season than those of the 
wild whitefishes which usually spawn sometime in November 
and (or) early December, and (3) that the majority of the 
Aquarium whitefish do not spawn, but retain their eggs in the 
Ovaries or in the body cavity where they undergo a process of 
absorption or disintegration. From the condition of the eggs 
of the year and of those undergoing disintegration in different 
months, it appears highly probable that the phase of rapid 
growth is initiated in the eggs sometime after September and 
completed sometime before March or February (?). The con- 
clusion that the aquarium whitefish do not spawn agrees with 
the statements of Mr. Lanier to the effect that no spawning was 
ever observed among these fish. It may also be observed here 
that sexual maturity is not correlated with the size of the white- 
fish, but rather with its age. Thus, for instance, in Table I the 
eight and nine-year-old females of January 28 and May 28, which 
measure 219 mm. and 210 mm. respectively, are no larger than 
a two-year-old whitefish from Lake Huron; yet, no two-year-old 
whitefish has ever been known to spawn. 


That sexual maturity can be a factor in the formation of 
annuli becomes evident when it is recalled that many species of 


1923] Van Oosten: Scales of Whitefishes 407 


fish, especially the salmon to which the whitefish are related, 
practically cease growing when developing a spawning condi- 
tion, even though the amount of food ingested remains prac- 
tically constant. It is also a fairly well established fact for 
many species that the annual increments in body growth of the 
young fishes are very noticeably reduced in that year when 
sexual maturity first occurs. Since it was found that the sex 
products of the Aquarium whitefish began their development 
after June, entered the phase of rapid growth sometime after 
September and completed this phase sometime before March or 
February(?) sexual maturity could account for the cessation in 
body and scale growth in August or September. Sexual ma- 
turity would then be a primary factor in the formation of annuli 
in the adults. 


It has thus far been shown that the scales of the Aquarium 
whitefish ceased growing sometime in August or September and 
resumed growth in April or March(?), that sexual maturity was 
reached sometime between September and March or Febru- 
ary(?), that the lowest temperatures of the aquarium water 
occurred in January to March inclusive, and that the amount of 
food required by the fish was less for the months of January to 
March inclusive than for the other months of the year. It was 
suggested that food could only have had secondary significance 
in the formation of annuli since the reduction of food was caused 
by some other factor which affected the appetite of the fish. It 
was further suggested that since reduction and increase in food 
consumption occurred synchronously with the decrease and in- 
crease in temperature respectively, and since the scales resumed 
their growth at the time of a rise in temperature in April, when 
sexual maturity could have had no influence on growth, tem- 
perature must be considered a primary factor in the formation 
of annuli. Lastly, since the sex products began their develop- 
ment at approximately the same time when a retardation or ces- 
sation of scale growth occurred in late summer, when the environ- 
mental factors of food and temperature were known to have been 
constant, it appears reasonable to assert that sexual maturity 
is also a primary factor in the formation of annuli in scales. If 
sexual maturity is not such a factor, then it must be conceded 
that the retardation or cessation of scale or body growth, and 
consequently the formation of annuli, is caused by some unknown 
physiological factor or factors of annual recurrence. 


The year of life in which sexual maturity first occurred can 
only be conjectured. The break in the growth curve (a) of 
Fig. 144, which represents the average annual increments of the 
Aquarium whitefish, suggests that most of these fish attained 
sexual maturity in the third year of life. This suggestion agrees 


408 Zoologica: N. Y. Zoological Society Baa be 


with the statements of Evermann and Smith (1896, p. 300) and 
the U.S. Fish Manual (19038, p. 110) to the effect that the white- 
fish reach sexual maturity in the third or fourth year; and with 
the statement on p. 120 of the Fish Manual where it is asserted 
that three-year-old whitefish artificially reared in the hatchery 
at Northville, Mich., yielded a large number of eggs, a fair per- 
centage of which were fertilized. My data on the wild white- 
fishes indicate that sexual maturity may be attained in the fifth 
and sixth years by whitefishes from Lake Huron proper and in 
the fourth year by those of the North Channel and Georgian 
Bay; but, an overwhelming majority of the available individuals 
of these years and localities appear to be immature. The wild 
whitefishes perhaps attain sexual maturity for the first time at 
an older age than has previously been assumed, or the first year 
of spawning may vary with the locality. 


The attainment of sexual maturity in the third year by the 
Aquarium fish would also account for the apparent change in 
their appetite in the third winter of life. With sexual maturity 
eliminated, food and temperature must have been the only en- 
vironmental factors of growth in the first two years of life. 
Temperature did not seem to impede the metabolism of the body 
very seriously as the young fish were always ready to eat. 
Restriction in food must then have been the principal factor in 
the formation of annuli in the first two years of life. 


(d) ANNUAL RATE OF GROWTH 


The lengths in millimeters attained by each Aquarium fish 
at the end of each winter of life (K,, K., etc.), and the annual 
growth increments in mm. of each year (k., k,, etc.) are shown 
in Table I. These values were obtained by the method described 
on pp. 385 and 386 and the formula given on p. 383 from the 
diameter of scales. The average length and increment for each 
year is given at the bottom of the respective column. Below the 
calculated averages are Shown corresponding averages of actual 
measurements of 238 Lake Huron whitefish taken at different 
localities in the fall of 1917 and 1919. In Fig. 144 are shown four 
growth curves based on the averages of Table I. Curve (a) 
represents the annual growth increments of the New York 
Aquarium whitefish, while curve (b) represents those of the 
Lake Huron whitefish. Curve (c) shows the total length 
reached by the aquarium fish at the end of each winter of life, 
while curve (d) shows the same thing for the Lake Huron fish. 

From curves (c) and (d) it is at once evident that the 
Aquarium whitefish have been greatly retarded in growth. 
Knowing the limitations of their food and swimming space re- 
tarded growth is to be expected. Curves (a) and (b) show that 
in no year did the Aquarium fish attain the rate of growth of 


Fic. 144. Showing growth curves which represent the average total 
length in mm. attained at the end of each winter of life by the Lake Huron 
Whitefish (d) and the New York Aquarium Whitefish (c), and the average 
annual growth increments in mm. of each year of life of the Lake Huron 
Whitefish (b) and the New York Aquarium Whitefish (a). A!l curves are 
based on the averages of Table I. 


the wild fish and that the greatest retardation in the growth of 
the Aquarium fish occurred in the first year of life. After the 
second year the annual growth increments were much reduced, 
but remained nearly constant to the eighth year in both series 
of fish. 

The validity of the scale method is sometimes questioned 
because of the great range in the extreme lengths of the speci- 
mens placed by it in the same age group. This criticism is 
wholly refuted in column K, Table I. Specimens 54509 and 
54510 of the same age and with the year’s growth completed 
measure 219 and 334 mm. respectively, showing a difference in 
length of 115 mm., while specimen 54514, several months older 
than the two just mentioned, measures 9 mm. less than the first. 


SUMMARY. 


1. Scales were examined from whitefish hatched January, 
1913, from eggs from the Put-in-Bay hatchery and reared 
in the New York Aquarium. The scales studied were re- 
ee from fish that were in their eighth and ninth year 
of life. 


410 


10. 


Le, 


12. 


Zoologica: N. Y. Zoological Society [II; 17 


The annuli in the scales of the Aquarium whitefish are of 
the same number as that of the winters of the fish’s life, 
the first one in which the fish were hatched excluded. The 
age of whitefish may therefore be determined from their 
scales. 

The annuli in the scales of the Aquarium whitefish are 
winter-marks formed by the retardation or cessation of 
scale growth in late summer and winter and completed by 
the resumption of scale growth in the spring of the year. 
The different dimensions (diameter, anterior and posterior 
radius) of whitefish scales grow at very different rates 
with respect to the rate of growth of the body and conse- 
quently the lengths and growth increments calculated from 
these dimensions vary significantly. 

The anterior radius of the whitefish scale grows relatively 
faster than the body and consequently the lengths calculated 
from it are too low. 

The posterior radius of the whitefish scale grows relatively 
much more slowly than the body and consequently the 
lengths calculated from it are much too high. 

The diameter of the whitefish scale, on the whole, grows 
relatively only a little more slowly than the body and con- 
sequently the lengths calculated from it are only a little 
too high. 

The lengths calculated from diameters are always higher 
than those calculated from anterior radii and lower than. 
those calculated from posterior radii. 

The diameter of scales seems to be a better basis for the 
length calculations of whitefish than the anterior radius 
which has nearly always been used in the past for many 
species of fish. 

A rather complete life history of the New York Aquarium 
whitefish is given, which includes a statement as to the 
character of the food consumed, the number of feedings, 
the changes in the amount of food consumed, the tempera- 
ture of the water in different months of different years, 
sexual maturity, spawning and the rate of body growth. 
The rate of body growth of the Aquarium fish is compared 
with that of Lake Huron whitefish. The significance of 
each of the above life history facts, rate of growth ex- 
cepted, as a factor in the formation of annuli, is discussed. 
Food is only a secondary factor in the formation of annuli 
in the adults studied, but may have been a primary factor 
in the immature fish. 

Temperature appears to be a primary factor in the forma- 
tion of annuli in the adults, but only a secondary one in the 
immature fish. 


1923] Van Oosten: Scales of Whitefishes | 411 


18. Sexual maturity appears to be a primary factor in the for- 
mation of annuli in the adults. 


14. The method employed in the study of the scales of the 
Aquarium whitefish differs somewhat from those described 
and used by other investigators. The method and the ap- 
paratus used are therefore briefly described. 


BIBLIOGRAPHY 


ARWIDSSON, IVAR 


1910 Zur Kenntnis der Lebensgeschichte der jungen Lachse in den 
Flissen von der Hinabwanderung ins Meer. Conseil Per- 
manent International pour l’Exploration de la Mer. Pub- 
lications de Circonstance, No. 54, 86 pp., Copenhagen. 


Brown, A. WALLACE 


1904 Some Observations on the Young Scales of the Cod, Haddock and 
Whiting before Shedding. Proc. Roy. Soc. of Edinburgh. 
Vol. XXIV, for 1901-1903. pp. 437-438. Edinburgh. 


CUNNINGHAM, J. T. 


1905 Zones of Growth in the Skeletal Structures of Gadidae and 
Pleuronectidae. 23d Ann. Rep. Fish. Board Scotland, pt. 
III. App. V. pp. 125-140. Glasgow. 


CUTLER, D. WARD 


1918 A Preliminary Account of the Production of Annual Rings in 
the Seales of Plaice and Flounders. Journ. Marine Biol. 
Assoc., n. s. Vol. XI. No. 4. pp. 470-496. Plymouth. 


EVERMANN, B. W., AND SMITH, H. M. 


1896 The Whitefishes of North America. App. 4, Report U. S. Com- 
missioner of Fish and Fisheries for 1894. Washington. 


FRASER, C. MCLEAN 


1917 On the Scales of the Spring Salmon (Oncorhynchus tschawyt- 
scha.) Contr. Can. Biol., 1915-1916. Ottawa. 


FULTON, T. W. 


1904 The Rate of Growth of Fishes. 22nd Ann. Rep. Fishery Board, 
Scotland. pt. III. pp. 141-241. Glasgow. 


HOFFBAUER, C. 

1898 Die Altersbestimmung des Karpfen an seiner Schuppe. Allge- 
meine Fischerei-Zeitung, nr. 19, Oct. 1, 1898. Jahrg. XXIII. 
Art. III. pp. 341-343. Miinchen. 

1899 ——————. Jahres Bericht des Schlesischen. Fischerei-Vereins. 

1901 Weitere Beitrage zur Bestimmung des Alters und Wachstums- 
verlaufes an der Struktur der Fischschuppe. Jahresber. 
der Teichwirthschaftl. Versuch-Station zu Trachenberg. 


HuTTON, J. ARTHUR 


1914 Recent Investigations upon the Life History of the Salmon by 
the Board of Agriculture and Fisheries. Salmon and Trout 
Magazine. No. 7. 28 pp. London. 


412 Zoologica: N. Y. Zoological Society (itsag 


JACOT, ARTHUR PAUL 


1920 Age, Growth and Scale Characters of the Mullets, Mugil cephalus 
and Mugil curema. Trans. Amer. Micros. Soc., July. Vol. 
XXXIX. No. 3. pp. 199-229. 


MASTERMAN, A. T. 
1913 Report on Investigations upon the Salmon with Special Refer- 
ence to Age Determinations by Study of Scales. Board of 
Agric. and Fish. Fishery Investigations. sr. I. Salmon 
and Fresh-water Fish. Vol. I. pp. 1-80. London. 


RicH, WILLIS H. 


1920 Early History and Seaward Migration of Chinook Salmon in 
the Columbia and Sacramento Rivers. Bull. U. S. B. F. 
Vol. XXXVI, 1919-20. Doc. No. 887. Washington. 


Scott, THOMAS 


1906 Observations on the Otoliths of Some Teleostean Fishes. 24th 
Ann. Rep. Fish. Board, Scotland. pt. III. App. II. pp. 
48-52. Glasgow. 


TAYLOR, HARDEN F. 
1916 The Structure and Growth of the Scales of the Squeteaque and 
the Pigfish as Indicative of Life History. Bull. U.S. B. F., 
Vol. XXXIV, 1914, Doc. No. 823, Sept., pp. 289-330. Wash- 
ington. ‘ 


THOMSON, J. S. 


1904 The Periodic Growth of Scales in Gadidae as an Index of Age. 
Journ. Marine Biol. Assoc., N. S. VII, pp. 1-109. Plymouth. 


Tims, H. W. MaretTtT 


1906 The Development, Structure and Morphology of the Scales in 
Some Teleostean Fish. Quarterly Journ. of Microscopical 
Science, Vol. XLIX, No. 193, n. s. Oct., 1905, pp. 39-68. 
London. 


U. S. FisH MANUAL 
1903 Artificial Propagation of the Lake Trout, Grayling and White- 
fish. Washington. R 


WALTER, E. 


1901 Altersbestimmung der Karpfen nach der Schuppe. In: Knauthe, 
Die Karpfenzucht, pp. 88-122. Neudamm. 


New York Zonlogiral Society 


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SCIENTIFIC CONTRIBUTIONS OF THE 
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VOLUME ll. NUMBER 18 al 


THE ANDERSON TREE FROG. 


(Hyla anderson Baird) 


OBSERVATIONS ON ITS HABITS AND LIFE HISTORY 
By G. KINGSLEY NOBLE AND RUTH C. NOBLE 


The American Museum of Natural History 


PB, bse i, D Be ¥ THE Spe Je Gisele thd Wy 
Men Z2ZO0DOGLCAL PARK, NEW YORK 


ce 


August 20, 1923 


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LOOLOGICA 


SCIENTIFIC CONTRIBUTIONS OF THE 
new YORK ZOOLOGICAL SOCIETY 


VOLUME ll. NUMBER 18 


THE ANDERSON TREE FROG 


( Hyla andersonii Baird) 


OBSERVATIONS ON ITS HABITS AND LIFE HISTORY 
By G. KINGSLEY NOBLE AND RUTH C. NOBLE 


The American Museum of Natural History 


D Bay T.H.E SOF Ce Lata 
GAC Avice) (PeAGR VK’, (NCE WwW: YO OURS 


August 20, 1923 


FIG. 145. THE CALL 


Male H. andersonii calling from a pitch pine on the edge of a pine-barren bog, Lakehurst, 
New Jersey. Flashlight Photograph. 


Zoologica Vol. II, No. 18. 


CONTENTS 


LTE OSS) CONGO INS 02 -s Uh on a A17 
[SUISUOTET GAG 2025 5) Sle meee a a eed a EY GURL” cen Sn Pt 418 
‘DCO ATTSIUHSOIN| ee eS ae 2 Re ie A a a a A19 
LASETE TNR os 6 ul S06 Nea SG etd I Bat roe cc a Ne 422 
LS TEBIBIDILS{CETS IAS ON Ase oi na eas lel oe OS aa ae A22 
 CETGIB fee gue ere a lo Ac ee 426 
Pein COG NEREON sya. 0. tr, Sc PAs ahaa wnt. hs Yoel oA vd ale 426 
SHE TCEUAT) IDEA C07S2. 2) EE S17 1S aa sh age Mee ae 2 sees 432 
TR REMOUROn GWTPOSMPION 6.) ue sels sete sda. Dadian gh 433 
EE MeCeANG MTS CAPSULES 6. 2030. oot Egan eae Poe 439 
DEVELOPMENT OF REE INGG 325202 <0 ae! Loa tft oie tee Seen Vas A41 
THE ADHESIVE ORGANS AND THEIR DEVELOPMENT.......... 442 
DEVELOPMENT OF THE TADPOLE. Y.:. yes oes cicd al. oa vk ea 443 
DESCRIPTION OF THE MATURE TADPOLE. ....0.5..0. 66... 005. 446 
PSUS TENT Sso(Q DUIS TBS DN 01210 16) DY aA Le ees eg Cae NU Se Ba eg 448 
LPUSGID, IG UATSUTS\ ONS So Sie ates er ie er hr Pana a Se a a 448 
LEELA INISTSUIES ODN Sees At SOE 2 Span ae 6 nan Peres Ae 449 
I CMUSIONS ar es etl See Tarek. eats A52 
[SI TSIIO GLE sha DRY TURN i oft eens ee een Oe ee 453 


First form on press August 20, 1923. 


FIG. 146. DISTRIBUTION OF HYLA ANDERSONII 
Spots indicate locality records. 


416 


-Volume II, Number 18. 


THE ANDERSON TREE FROG 


(Hyla andersonii Baird) 
OBSERVATIONS ON ITS HABITS AND LIFE HISTORY 


By G. KINGSLEY NOBLE AND RuTH C. NOBLE. 


The American Museum of Natural History 


INTRODUCTION. 


The Anderson Tree Frog has often been considered the most 
attractive of North American hylas, and yet no attempt has hitherto 
been made to study its life history in detail. Former observations 
were made as opportunity permitted and none were continued at 
frequent intervals throughout a season. 

The following observations represent only a single season’s work, 
but they were made with a definite plan in view. The question 
which we have had before us throughout the work was: what are 
the relationships of H. andersoni so far as these may be deduced 
from its habits and life history? Wright (1914), by comparing the 
life histories of certain American batrachians with those of European 
species, has brought forth some clear-cut evidence as to the re- 
lationships of the species he considered. In the present paper we 
have made no attempt to discuss morphological or embryological 
data which do not have a direct bearing on our main problem. 

The following observations were made by the writers at inter- 
vals during May, June and July, 1922. To these observations there 
have been added others made by one of us on various occasions 
during the three preceding years. Field observations have been 
supplemented by studies in the laboratory. Our knowledge of the 
life history of the Anderson Tree Frog is stil] far from complete. In 
the hope that someone more favorably situated than ourselves will 
continue this work, we have made some attempt to give a complete 
picture of what is known concerning the life history of this delightful 
tree frog. Net only have most of the published accounts been 


417 


418 Zoologica: N. Y. Zoological Society ) (Tis 


available to us, but we have been fortunate in having near at hand 
Messrs. W. T. Davis, J. P. Chapin, W. De W. Miller, G. S. Myers, 
K. P. Schmidt, C. L. Camp and others who are very familiar with 
the Anderson Tree Frog in the field. Many of the problems which 
arose we have discussed with these gentlemen. Wherever their 
observations have supplemented our own we have included them 
below with full acknowledgment. It was obvious from the first 
that no complete study of the life history of H. andersonii could be 
made in a few months time. Emphasis was therefore laid upon 
those features which might be expected to shed light upon the re- 
lationships of the species. These features we have discussed under 
separate headings. 


HISTORICAL. 


Hyla andersoni has until comparatively recent years been 
considered one of the rarest of American batrachians. When Cope 
wrote his standard work on “The Batrachia of North America,’’ 
only two specimens had ever been taken: the type specimen, de- 
scribed by Baird (1854) captured at Anderson, South Carolina, and 
a second specimen collected by Professor J. Leidy “in a cedar swamp 
near the town of Jackson in New Jersey, sixteen miles east of 
Philadelphia.” Cope (1862) gave a description of this second speci- 
men from life. This description, with a few emendations, was later 
repeated by him (Cope 1889) and again by Fowler (1907). A 
third specimen of Hyla andersonii was recorded the year Cope’s 
general work appeared (Peters 1889). It was captured the previous 
year “‘on the border of a pine barren at May’s Landing, N. J.” 
Abbott (1890) who had this specimen in captivity, criticises Peters 
for his description of the call. A few years later Moore (1894) 
published a short but informing account of observations he made at 
Pleasant Mills, N. J. This was followed by the capture of an 
additional specimen at Clementon, N. J., and Stone (1901), in a 
note concerning the specimen, states that “the species would no 
doubt prove more abundant if specially sought for, the comparative 
remoteness of the New Jersey barrens, where most of the specimens 
were found, and the retiring habits of the animal both tending to 
make its detection difficult.” 

This prophecy was soon substantiated. Davis (1904, 1905 and 
1907) made a series of observations at Lakehurst and Farmingdale, 
N. J., tending to show that the species was not rare in the New Jersey 


1923] Noble: The Anderson Tree Frog 419 


pine-barrens. We owe to Davis cur first account of the tad- 
pole, and its metamorphosis. Fowler (1909) has given an excellent 
color description based on additional specimens. Of equal value is 
the splendid color plate given by Miss Dickerson (1906). She kept 
under observation one of the specimens secured by Mr. Davis and 
has given a good account of the species in captivity. Recently 
Barbour (1916) captured (July 8 at Lakehurst) “well up in a 
pine tree”’ two pairs while in embrace. These laid the same evening 
under abnormal conditions and no record was made of the egg- 
masses or eggs. More recently Miller (1916) has extended the 
known range of the species by finding it in several localities, “‘all 
in the sandy pine-barren ‘island’ north of the Pine-Barrens proper.” 
Since 1916 the species has been studied at Lakehurst on one or more 
occasions by Messrs. C. L. Camp, K. P. Schmidt, R. Deckert, G. S. 
Myers, the writers and perhaps others. Lastly it may be pointed 
out that mention has been made of H. andersonii by various authors 
not listed above. Among these are Boulenger (1882), Sherwood 
(1898), Ditmars (1905), Stone (1906) and Deckert (1918). Further, 
Davis (1922) has very recently reported the occurrence of the 
species in North Carolina. 


DISTRIBUTION. 


Hyla andersoni is a tree frog of the pine-barrens. The type 
specimen is credited with coming from Anderson, South Carolina 
and the species was named by Baird after that town. But as 
Anderson lies neither in the pine-barrens (Livingston & Shreve 1921) 
nor even in the coastal plain, we suspect that the type specimen 
actually came from some other locality. As shown in the accom- 
panying map, all the other locality records for H. andersonii lie 
within the pine-barrens or their outlying “‘islands.’”? Although 
H. andersoni is abundant throughout most of its range, this range 
does not extend the entire length of the pine-barrens. The species 
has not been recorded south of South Carolina nor in the pine- 
barrens of Long Island. It is, however, widely spread throughout 
the New Jersey pine barrens occuring even in the pine-barren 
“island” just south of the Raritan River. 

lt should be noted, however, that the pine-barrens of New 
Jersey have a flora somewhat distinct from those of North Carolina. 
Mr. W. T. Davis who has studied both regions assures us that the 
facies of the country about Southern Pines, N. C., where he and 


abs 


THE PINE-BARRENS AT LAKEHURST, NEW JERSEY 


A. Portion of the square mile of bog that was intensively studied. Four pairs of 


H. andersonii were taken while in embrace in a small pool on the left side of the road, directly 
B. Cedar 


Not one was found to breed in the deep water to the right. 
It is from these cedars that most males call. 


FIG. 147. 


behind the figure. 
swamps bounded our area on two sides. 


420 


Zoologica Vol. Il, No. 18. 


1923] Noble: The Anderson Tree. Frog | 421 


Dr. Chapin collected H. andersonii is very different from that of 
the New Jersey pine-barrens. The soil of both regions, however, 
is sandy, and much of the vegetation is the same. Until more 
extensive investigations can be made in the Carolinas, it is perhaps 
most conservative to say that H. andersonii is confined to the 
“‘pine-barrens.”’ 

It will be noticed that the distribution of H. andersonii cannot 
be expressed by a single term such as ‘‘Carolinian Life Zone’’ or 
“Coastal Plain.”’ To be sure, it is to be found in these regions, but 
its range does not agree at all closely with either area. Many have 
attempted to reduce the distribution of life in North America to 
a few terms,—to a few zones—or to a few physiographic areas. 
But always objections have been brought forth in opposition to these 
attempts. 

Confronted by such conflicting views, one is at first inclined to 
deny that the distribution of any one form can be explained in terms 
of another. The physiology of no two related animals is the same, 
—why should we expect that related animals would react the same 
to temperature? If we should pick two animals at random that 
have the same reaction to temperature, we might find that they 
reacted differently to ten additional stimuli. Further, the dis- 
tribution of one species might be due to one factor, and of another 
species to another. There is no reason why one should expect 
to discuss the distribution of frogs in terms of the distribution of 
birds. But one may, I believe, make a distinct advance by speaking 
of the distribution of one frog in terms of another’s range. During 
the breeding season batrachians congregate in distinct habitats. 
At other times of the year they do not wander far from these 
habitats. There is some reason to suppose that closely related 
species will have somewhat similar breeding habits. It was on the 
basis of this that Wright made comparisons between the affinities 
of certain European and American batrachians. 

Hyla andersoni is a “coastal plain form.”’ Many other species 
seem to be typically coastal plain forms in our area. These may 
be listed with their breeding habitats: 

(1) Scaphiopus holbrookii—Temporary pools in sandy regions. 

(2) Acris gryllus—Weedy, especially water-lily, ponds. 

(8) Hyla andersonii—Small pools in sphagnaceous bogs of pine- 

barrens. 

(4) Rana virgatipes—Larger pools of pine-barrens. 


422 Zoologica: N. Y. Zoological Society [II; 18 


(5) Rana piprens—Shallow ponds. 

(6) Bufo fowleri—Shallow ponds. 

Whether or not it is because the last two species are not par- 
ticular as to the exact nature of their breeding cite, the fact remains 
that they have a much greater distribution in our local region than 
the other four species. Further, these two species often extend their 
range beyond the coastal plain, and probably because of this toler- 
ance. Scaphiopus holbrookii has been taken in various parts of 
New England where the soil was sandy. H. andersoni and R. 
virgatipes are ecnfined to the pine-barrens. Why R. virgatipes has 
a more extensive range to the south than H. andersonii it is im- 
possible to say at this time. Acris gryllus is not found in the pine- 
barren bogs, and probably because of its preference for less acid 
water. When the life histeries of all our American Amphibia are 
known, the species arranged according to their breeding cite pref- 
erences, many of the present anomalies of distribution among our 
Batrachia will be explained. 


HABITAT. 


The pine-barrens of New Jersey have been well described by 
various botanists (Stone 1911, Harshberger 1916, etc.). H. ander- 
sonii is not limited to any association within the pine-barrens. 
Specimens have been captured in many different kinds of bushes 
and trees. When pursued, H. andersoni invariably leaps to the 
ground and, with a series of short jumps, disappears among the 
grass and sphagnum of the bog. Specimens captured by us on the 
ground have been taken only in sphagnaceous areas. 

Specimens of H. andersonii are taken most easily by running 
down the calling males at night, with an electric hand Jamp. Such 
specimens have been captured on the ground, on low bushes, on 
the top of bushes six to eight feet in height and rarely in trees or 
tall bushes more than eight feet from the ground. We have c¢ap- 
tured only two specimens in the latter position, but Mr. Myers 
has observed a third. 


BREEDING SEASON. 


Previous to our work during the spring of 1922, the eggs of 
H. andersonii had only been observed twice and both under ab- 
normal conditions. Barbour (1916) reports the capture of pairs in 
embrace at Lakehurst, New Jersey, on July 8. Eggs were laid in 


1923] Noble: The Anderson Tree Frog 423 


a crowded vessel during the night. Messrs. Davis and Chapin 
captured a pair at Southern Pines, North Carolina, which later 
embraced and laid eggs the night of June 13, 1922, and also on the 
following night under abnormal conditions. No detailed obser- 
vations were made on the egg laying process and no record made of 
the egg form. Previous to this, Davis had found tadpoles of 
H. andersonii in “all stages of development” at Lakehurst on 
July 21, 1907. From these observations it seemed probable that 
the breeding season of the Anderson Tree Frog at Lakehurst ex- 
tended through June and part of July and may have begun earlier. 

At Lakehurst, New Jersey, during 1922, the breeding season 
had apparently not begun by May 14. A few males were calling, 
but these from concealment on the ground (with a single exception, 
Camp), and mostly from among the sphagnum. On May 20 and 21 
at exactly the same place in the bog, the breeding season was well 
under way. ‘Two females were cbserved before embrace and taken 
during oviposition; a third female was taken just before oviposition. 
The chorus of the males was loud, approximately twenty-five to 
fifty calling within an area of one square mile. On June 4 and 5, 
approximately one hundred males were calling in this same area. 
One female was captured just before the embrace, and four pairs 
were taken in embrace. Although eggs had been found on May 20 
and 21, no tadpoles could be found June 4 and 5. On June 18-19, 
the chorus of H. andersoniz within the square mile of territory we 
were studying reached its maximum. At least two hundred and 
fifty males were calling within this area. Although no mated pairs 
were found, tadpoles of a wide range of sizes (up to appearance of 
hind limbs) were captured. On July 22-28, the chorus had dimin- 
ished. Only about forty males were calling. Tadpoles of all sizes 
from shortly after the formation of the operculum up to meta- 
morphosis were captured in this same region. 

These observations make it clear that the breeding season of 
H. andersonii is not an explosive one, but is more or less protracted. 
Rains occurred at Lakehurst on the days preceding our observations 
of May 20-21, June 4-5 and June 17-18. The days, therefore, 
were comparable so far as humidity was concerned. (Unfortu- 
nately, no thermometers were used to determine this exactly.) 
Comparing this data with random observations made on previous 
years at Lakehurst, the breeding season of H. andersoni seems to 
be more or less dependent on the rains and may extend from the 
middle of May to the middle of July. 


424 Zoologica: N. Y. Zoological Society | (Tiss 


It was perhaps not surprising to find that in correlation with 
this protracted breeding season, the species breeds in more or less 
isolated pairs. As this method is not the general rule among our 
local Salientia, it may be of interest to make some comparisons 
between the breeding season of H. andersonii and that of other 
local forms. So little is known concerning the factors controlling 
these differences that only the most obvious will be mentioned. 


I. Internal factors 

1. Maturing of gonads, may be either 

a. Uniform for all individuals of aspecies within an area, 
or 

b. Irregular, small colonies breeding at different times. 

2. Physiological cycle (correlated with the development of 

gonads) may be 

2. Genetic (closely related species living under similar 
conditions may have different breeding season), or 

b. Acquired. It is well known that Amphibia in cap- 
tivity gradually modify their breeding season, and 
may eventually lay at very irregular times. The 
life cycle characteristic of the species may have been 
induced by the environment, just as it has been 
modified by the new environment of captivity. 

II. External factors 

1. Temperature 

a. Air temperature,—affects those batrachians hiber- 
nating on land. These are, fer the most part, early 
breeders (Wright, 1914). 

b. Water temperature,—affects the species hibernating 
in the bottoms of ponds. These are chiefly late 
breeders. 

2. Rains (often correlated with a change in temperature)— 
are the chief controlling factors in the breeding of Batrachia 
within the tropics. This has been shown experimentally 
by Bles (1906) but field observation has further demon- 
strated the fact. In temperate regions, the rains play a 
vital part in the life cycle of some forms (as Scaphiopus), 
but much less in the case of others. 


As a result of the interaction of these external and internal 
factors, the various species of Batrachia within our local area are 
found to breed at different times. We may group them according 
to the time and period of their breeding season into several cate- 
gories with the understanding that these categories are subject to 
fluctuations according to variability of both external and internal 
factors: 


1923] Noble: The Anderson Tree Frog 425 


I. Explosive Breeders 

1. Temperature controlled; mostly early breeders which have 
hibernated on land. Bufo americanus, Rana pipiens and 
Rana sylvatica. 

2. Rain controlled; Scaphiopus holbrookii, often selecting a 
poor breeding cite. 

II. Protracted Breeders 

Temperature controlled; including a few early but mostly 
late breeders, forms which have either hibernated in the 
water, Rana palustris, Rana clamitans, Rana catesbeiana 
and Rana wrgatipes, or others on land,—Pseudacris tri- 
seriata (?), Hyla crucifer, Hyla versicolor, and Acris gryllus. 

2. Rain controlled; Bufo fowler:, Hyla andersonit may choose 
a poor breeding cite but more often one favorable to the 
welfare of the tadpoles. 


The distributions made above are only approximate, and, 
moreover, apply chiefly to our local area. At Ithaca, New York, 
Wright (1914) found that temperature was the chief factor controll- 
ing the breeding season of the Salientia. Wright was able to arrange 
the species he considered in a series according to their first appearance 
and time of breeding. South of Wright’s locality, even in the New 
York region, the rains begin to play an important part in the breeding 
of the Salientia. In the tropics, as we have recently seen in Santo 
Domingo, it is the rains—and apparently only the rains (correlated 
of course, with slight changes of temperature)—which initiate the 
breeding season. 

In the case of H. andersoni our observations have not been of 
sufficient frequency to determine exactly the part played by the 
rains and the part by temperature in determining the breeding 
season. By comparing several seasons’ observations, it would 
seem that both rain and temperature are effective, but as the season 
is a long one—possibly due to an irregular maturing of the gonads 
in the colony—it would seem that the rains may be the more im- 
portant factor, although, of course, not as effective as in the case of 
explosive breeders such as the Spadefoot toad (Scaphiopus). 

We may add, in passing, that not sufficient consideration has 
been given to the rains in initiating the laying process of Amphibia 
in general. The breeding season of not only some northern Salientia, 
as Scaphiopus, but even of some northern Caudata, may be con- 
trolled primarily by the rains. This has been very thoroughly 
demonstrated in the case of Hynobius nebulosus by Kunitomo 
(1910). It is to be hoped that more exact data of this sort will be 


426 Zoologica: N. Y. Zoological Society [11; 18 


forthcoming for other northern species. It would be interesting 
to know whether the breeding season of a single species may be 
primarily rain or temperature controlled, according to the northern- 
ness of its breeding site. 

VOICE. 


One of the most characteristic features of H. andersonii is its 
distinctive voice. As in the case of all known hylids, only the male 
is thus provided. The call has been variously described as a 
“‘keck-keck”’ (Abbott), a “peep-peep”’ (Peters), a ‘“‘quack-ack”’ 
(Moore), a “whang” (Deckert) or a “aquack-aquack-aquack”’ 
(Davis). One would gather from the literature that the Anderson 
Tree Frog had a variety of calls. Our common tree toad (Hyla 
versicolor) has two distinctive calls. Although Wright (1914, p. 46) 
states that ‘‘an individual tree-toad may give one or two voice- 
forms totally unlike the normal and better-known call,’ undoubtedly 
these “‘abnormal calls” will be found referable to what Overton 
(1914, p. 33) has called the “turkey root.’’ Since Overton’s paper 
appeared, we have watched on several occasions Hyla versicolor 
give its “turkey root.’’ On none of these occasions was there any 
bobbing of the head such as Overton describes. The call, variable 
as to number of syllables, is given with a half inflated pouch. When 
H. versicolor calls with fully inflated pouch it gives its characteristic 
trill; when it calls with only half inflated pouch, a series of mournful 
notes arise. : 

H. andersoni has only one call and that is given with fully 
inflated pouch (Fig. 148). It is a series of ten to twenty, or even 
more, resonant nasal notes, usually increasing in volume. Each 
note is a sonorous, high nasal quank. If one tries to shout the 
word quank while holding the nostrils closed, a sound is produced > 
not unlike the note made by this frog. The call sounds somewhat 
different from a distance, especially when several frogs are calling 
at once. Then the notes tend to run together, each note having 
two syllables, a-quank, a-quank. 


SEX RECOGNITION. 


Although the call of H. andersonzi is loud, often carrying nearly 
a mile on quiet evenings, nevertheless, little or no significance has 
been attributed te it, or to the voices of any frogs and toads, in the 
breeding season. Voice is stated not to control the direction of: 
migration towards the breeding grcunds, or the movements of in- 


1923] Noble: The Anderson Tree Frog 427 


dividuals on the grounds (Boulenger, 1912, p. 22, Cummins, 1920, 
p. 325). It is generally believed that “courtship does not take 
place in any of the tailless batrachians. The female is seized by 
the first comer, ”” (Boulenger, 1898, p. 68). Some years ago 
it was pointed out by Courtis (1907, p. 678) and later by Miller 
(1909, p. 650) and by Wellman (1917, p. 107) that the breeding 
female toad (Bufo) may respond positively to the trill of the male. 
But Cummins (1920, p. 248, italics his) has recently shown that in 
the case of frog material the “‘voice does not direct the movement 
of the frogs into the pond” and “that ‘sex recognition’ . . . results 
from the differential behavior of the two sexes when clasped,. . .” 

The following observations made during 1922, on Hyla ander- 
sonii would tend to show that in that species, and by inference in 
tree frogs in general, the voice plays a considerable réle in bringing 
the two sexes in contact. 

At Lakehurst on May 13-14, H. andersoni was calling only from 
concealment (with one exception) on the ground, chiefly among the 
sphagnum of the bog. On May 20-21 some were calling from the 
ground, but mostly from trees and bushes a few feet from the 
ground. On all other occasions during June and July, H. andersonii 
called chiefly from some point of vantage above the ground; only 
very few were seen calling from the ground. 

As pointed out above, the breeding does not occur simultan- 
eously throughout the Jimited region under observation. By means 
of the flashlamp, individual frogs were studied for several consecutive 
hours on the days indicated above. The following observations 
were made the night of May 20-21. 

Case I. 11:30 P.M. A male was observed calling from edge 
of small and weed-grown ditch, about one and one-half to two and 
one-half feet across, one to ten inches deep; bottom covered with 
sphagnum and water weed. 

Female was first observed three feet away from male, hopping 
in his direction. Without hesitation the female leaped on back 
of the male; but the latter gave a slight wriggle, which threw her 
off his back and he centinued calling. 

Female turned and leaped again on the back of the calling male. 
Again he threw her off with a wriggle, but this time, as she moved 
again toward him, he caught sight of her and quickly turned about 
and embraced her with the normal supra-axillary amplexus. The 
pair maintained their position about eight inches from the water 


428 Zoologica: N. Y. Zoological Society (II; 18 


(Fig. 149c) until 12:30 A.M., when the female leaped with her mate 
into the stream, and a moment later ovulation began. 

Case II. A male was observed calling at 3:00 A.M. (May 21) 
from the edge of the same stream about one hundred yards away 
from place where the above observations were made. The character 
of the stream was identical to that of Case I. The male was sitting 
on a soaking mass of sphagnum close to the water’s edge. The 
female was first observed two feet away, making short leaps toward 
the male. The female approached to within six inches of the male 
and while he continued calling, she hopped rapidly about him twice. 
In making these two circuits she had to splash through about one- 
half inch of water. In completing the last circuit, the female ap- 
proached so close to the male that the side of her body touched his 
side and she nudged him with her limbs as if to draw as close to him 
as possible. On completing his call, he turned and embraced her. 

The pair in embrace hopped for about six yards along the edge 
of the stream and began to ovulate in the water. They were finally 
placed in a vessel where ovulation was completed. 

The following observations were made during the night of 
June 4-5: 

Case III. A male that was calling from a bush, and photo- 
graphed twice, (Fig. 149a) seemed particularly nervous because he 
frequently changed his position a few inches. Suddenly, at 12:30 
A. M., he left the bush without warning. He was followed with the 
flash-light to which he paid no attention, and although we changed 
the position of the light frequently, the frog hopped straight across 
the bog over ruts and small depressions to a small stream flowing 
in a sphagnum-grown ditch. There he took up a position (Fig. 
149b), three inches from the water and approximately thirty feet 
from his first calling station, and began to eall. 

Half an hour later a female came hopping across the bog. She 
was first observed about fifteen feet from the male. She came 
straight toward the male which continued calling. When about 
four inches from him she turned slightly and hopped past him 
but he paid no attention to her. She then continued her journey 
toward a thicket where over thirty males were calling. The female 
began to cross through a weed-grown bog, and as it was apparent 
that we could not follow her, she was collected. Later we found 
only a few yards ahead a flooded but very shallow bog in which 
there were four mated pairs. It was over one hundred and fifty 


1923] Noble: The Anderson Tree Frog 429 


yards from the point where the female was first observed to the 
place where she was taken. In making the journey, the female 
had passed no less than four calling males, but she came close only 
to the first. 

From these data the following tentative conclusions may be 
reached: 

1. H. anderson begins calling in early May from con- 
cealment on the ground. 

2. In the middle of May and throughout June and July 
the males cal] chiefly from bushes or from trees. 

3. At various intervals throughout May and June (and 
some years, in July) when the rains have flooded the bogs and 
changed the ditches into small sphagnum-choked streams, the 
males leave their calling stations and make their way to the 
nearest of these small streams. This migration occurs about 
midnight. The males begin calling again from .their new 
positions near the sphagnaceous streams. 

4. The females are attracted toward the male by his call. 
This attraction is so great that it causes the female to leap upon 
the calling male. 

5. After a more or less persistent courtship on the part of 
the female, during which she strikes the male one or more 
times, the male recognizes the female and embraces her. 

6. It is possible that the female may, under certain cir- 
cumstances, climb a tree after the calling male. At least, this 
seems to be the most feasible explanation for the observation 
made by Barbour (1916) of a pair in embrace while high in a 
pee: 

7. The female may exercise some choice in the selecting of 
a mate. The call is not the only factor involved in bringing 
the sexes together. 

Although no one has observed this method of female courtship 
in any species of Rana and only part of the phenomenon has been 
witnessed in Bufo material, we believe it will be shown to be the 
rule in the case of Hyla. On June 7 at 11:00 P.M. near Patchogue, 
Long Island, we observed a female H. versicolor swiftly approach a 
calling male from behind and leap directly on his back. The male 
broke off his call at once, turned, and embraced the female. 

When the movements of individuals of other species of frogs 
have been followed throughout the night, we believe it will be clearly 


FIG. 148. FLASHLIGHT PHOTOGRAPHS ILLUSTRATING THE DIVERSITY OF 
THE CALLING STATIONS. 
a. Beginning the call from top of a pine-barren maple; note the inflation of the body, 
and the rain drops on the leaves. 0. The height of the call. c. Calling from the wheel 
of an old abandoned cart on the edge of the pine-barrens. d. The height of the call. 


430 
Zoologica Vol. II, No. 18. 


FIG. 148. FLASHLIGHT PHOTOGRAPHS ILLUSTRATING THE DIVERSITY OF 
THE CALLING STATIONS. 


e. Beginning the call from the top of a bush about three feet high. f. The height of 
the call. g. The rest between calls from a blue-berry-bush. h. The call; note the position 
of the hands and feet. 

431 


Zoologica Vol. II, No. 18. 


432 Zoologica: N. Y. Zoological Society (iieas 


demonstrated that voice plays a considerable réle in bringing the 
two sexes together. The problem of sex retention is a more dif- 
ficult one, and can be determined only by careful experimental work. 

It will very probably be shown that voice plays a considerable 
réle, not only in bringing the two sexes in contact, but also in attract- 
ing individuals together to form breeding colonies. We have found 
that males of Scaphiopus holbrooki, Bufo americanus and H. ander- 
soni. often came toward us when we imitated their cal]. In the 
case of one male H. andersonii, the hand-light was directed from a 
variety of angles and yet when we called, the frog always came 
toward us. We have never noticed a male H. andersonii jump ona 
calling male of the same species. It therefore seems likely that the 
attraction of the call is not as great in the case of the male as in that 
of the female. It, nevertheless, may play a considerable roéle in 
the ecology of a species, and to our tentative conclusions above we 
may now add two others. 


1. The gregariousness of a species during the breeding 
season is a function of the attracting power of the call upon 
males of the same species. For example, Scaphiopus holbrookii 
is more gregarious than Rana palustris because it is readily 
attracted by the calling of the colony. 


2. When several species are breeding in one marsh, the 
species are usually separated into colonies because of the 
specific attraction of the different calls. 


SEXUAL DIMORPHISM. 


With the several excellent descriptions of coloration of H. 
andersoni available, especially the color plate of Miss Dickerson 
(1906, color plate VII), it would be superfluous to give a new 
description of this form. Nevertheless, it has not hitherto been 
pointed out, although recognized by some (Davis, Myers), that 
there is a distinct sexual dimorphism in this hylid. I find the fol- 
lowing constant differences between the two sexes. These differences 
are sufficiently marked to permit one to distinguish between the 
two sexes in the field. 

1. Throat of breeding male, purplish gray; throat of 
breeding female pale gray or white, rarely as dark as the throat 
of palest male. 

2. Green patch below angle of jaw broadly edged with 


1923] Noble: The Anderson Tree Frog 433 


white in all females, without a white border in the breeding 

male, or with a very narrow and indistinct one. 

3. Breeding females distinctly larger than males. Average 
head and body length (snout to vent) in the ten females taken 
in embrace is 40.9 mm. (extremes 44 and 38 mm.). Average 
head and body length of fifteen breeding males is 36.7 mm. 
(extremes 35 and 38 mm.). 

In addition to these three characters of which the best field 
mark is the second, there are the two sexual differences to be ex- 
pected. First, the vocal pouch of the breeding male is always more 
or less indicated even in quiet individuals. Secondly, the male 
bears on the inner and upper side of the thumb (prepollex region) 
a patch of minute pigmentless asperities hardly recognizable without 
a lens. The female bears in this same position glandular skin as 
smooth as the adjoining regions. 

As pointed out by Dickerson (1906), the color pattern of H. an- 
dersoni is very constant, the change of coloration being limited to 
a darkening or lightening up of the tones. These changes of color 
are correlated with at least three factors,—(1) excitement, (2) light, 
and (3) humidity. 

A pair in embrace are usually of a different color. If the female 
is ovulating and moving only short distances about the pool, the 
male is much the darker probably because of the sexual excitement 
accompanying fertilization; but if the female is moving rapidly 
along the edges of a bog, the male merely retaining his position on 
her back, the female is the darker. These facts were clearly shown 
in the three cases discussed above. 

The effect of light and moisture on the color of these hylids 
may be readily demonstrated by keeping them in terraria of various 
degrees of humidity. Individuals in cold, wet terraria are very 
dark. 


METHOD OF OVIPOSITION. 


Amplexus in H. andersonii is supra-axillary. No other type or 
no abnormal amplexus was observed. The partly closed hand of 
the male is dug into the sides of the female just behind the head of 
the scapula, and just below the diapophyses of the anterior vertebrae. 
These diapophyses prevent the hand of the male from slipping 
dorsally, the scapula prevents it from slipping anteriorly, while the 
viscera of the female prevents it from sliding posteriorly. 


FIG. 149. FLASHLIGHT PHOTOGRAPHS OF H. ANDERSONII ILLUSTRATING 
STAGES IN THE BREEDING PROCESS. 
a. Beginning the call from ferns and bushes on a bank about two and one-half feet 


above the level of the bog (see text, Case III). b. The new calling station. about 30 feet 
from the first, close to a smal] sphagnaceous stream. a. and b. same individual. 


434 
Zoologica Vol. II, No. 18. 


FIG. 149. FLASHLIGHT PHOTOGRAPHS OF H. ANDERSONII ILLUSTRATING 
STAGES IN THE BREEDING PROCESS. 


c. The embrace. After the female had flung herself twice upon the calling male, he 
finally turned and embraced her (see text, Case 1). A male Bufo fowleri is calling close at 
hand (left of picture). Oviposition did not take place until an hour later. d. The beginning 
of oviposition; note the position of the male and the sphagnum projecting above the water. 


435 
Zoologica Vol. II, No. 18. 


436 Zoologica: N. Y. Zoological Society | ea be, 


Oviposition takes place always in the water and only in small 
basins, or slow-moving streams on the pine-barrens, never, however, 
in stagnant water. All eggs, tadpoles and pairs in embrace were 
found in puddles and streams lined with sphagnum. The water of 
the pine-barrens is always more or less coppery, even after hard 
rains. The color is due to tannin derived from the roots of the bog 
plants. This tannin makes the water slightly aseptic. Eggs will 
develop in other kinds of water. We have found that several lots 
developed normally in tap water. We were not, however, successful 
in raising the tadpoles in tap water. Dilutions of bog and tap water 
proved more satisfactory. From our observations it would seem 
that the bog water is essential to the normal development of the 
tadpole. 

The details of oviposition were studied in four pairs, two in the 
field and two in the laboratory. Mirrors were used to determine the 
exact course of the egg. The process of egg-laying was found to 
be the same in both captive and wild specimens. 

In Cases | and II, discussed above, oviposition began almost 
immediately after the female leaped from land into the deeper water 
of the streamlet. The characteristic attitude in the water is shown 
in the photograph of another pair (Fig. 149d). 

The egg-laying process seems to be initiated by the female. 
She bows her back suddenly, at the same time protruding the 
cloaca. A bunch of eggs appears in the orifice of the cloaca and 
as the female bows her back, either these eggs or the cloaca of the 
female touches the male between the legs. Immediately the male 
wriggles. It is assumed that the male emits the spermatozoa at 
this moment, but these are invisible. In the fully bent position 
(Fig. 150) the cloaca of the female is anterior to that of the male. 
As soon as this position is reached, the female straightens her body, 
suddenly ejecting the eggs. The eggs may have received a wash 
of sperm as they were protruding from the cloaca. It is more 
likely, however, that they strike the spermatozoa as they are shot 
from the cloaca. It will be observed that this method of fertiliza- 
tion is unlike that of the spring peeper, H. crucifer. Here the egg 
(rarely 2 eggs) is held in the orifice of the cloaca not only during 
the upturning of the cloaca, which, as in H. andersoni, seems to 
function in stimulating the male, but also during the return move- 
ment and subsequent forward movement of the whole posterior 


1923] Noble: The Anderson Tree Frog 437 


region of the female. Hence, the egg of H. crucifer is apparently 
fertilized while held in the cloacal orifice of the female, while it 
may or may not be fertilized at this time in the case of H. andersonii. 
The ovipositions of H. andersonii and H. crucifer may be compared 
as follows: 


H. crucifer H. andersonii 
(1) Cloaca upturned, and egg (1) Back bowed greatly, and 
or its capsule (rarely two cloaca upturned; part of a 
eggs) appears in orifice of bunch of eggs (7 to 14) ap- 
cloaca. pear in orifice of cloaca. 


(2) Cloaca of female usually (2) Same. 
touches posterior ventral 
surface of male’s body in 
upward movement. 

(3) Emission of spermatozoa (3) Same. 
apparently takes place as 
cloaca touches or passes 
near ventral surface of 


male. 

(4) Back straightened and (4) Back straightened and 
cloaca of female brought eggs are shot out against 
forward beneath body body of male to which they 
where the egg (or eggs) is do not adhere, but glance 
shot out against some ob- off:to the bottom of the 
ject to which it adheres. pond. 


The remainder of the egg-laying process of H. andersoni can- 
not be compared in detail with that of H. crucifer. At the moment 
the eggs are extruded, the hind limbs of the female are convul- 
sively straightened, forcing the pair forward in the water. The 
female makes one or two nervous half strokes which continue the 
headway of the first stroke. In one to three seconds they have 
again come to rest, the female again bows her back and the process 
is repeated. After two to ten (possibly more) layings, the pair come 
to rest and oviposition may not continue until half an hour later. 
The exact Jength of these “rests”? between sexual periods were not 
determined in the field. In the laboratory this “rest’’ was éx- 
tremely variable. 

The most remarkable feature of this egg-laying is the course 
taken by the eggs (Fig. 150). The female bows her back very 
much as in the ease of the ovipositing H. versicolor, and even makes 


438 Zoologica: N. Y. Zoological Society : iGers 


FIG. 150. THE OVIPOSITION OF HYLA ANDERSONII 


This bombardment of the male with eggs seems to he a specialization derived from the 
method of oviposition found in H. versicolor, where the female lifts her cloaca above the water. 
In this diagram, only one egg has been represented (instead of nine), and the legs of the male 
have been abnormally extended in order to show better the course of the eggs. 


some attempt to raise the cloaca above the water, as in that species, 
but the eggs never (or very rarely) reach the surface of the water. 
On being shot from the cloaca they strike the male on his ventral 
surface immediately below his cloaca and are carromed off to the 
bottcm of the pool. Of the many times we watched this bombard- 
ment of eggs both in the two pairs studied in the field, and the labor- 
atory specimens, only twice—and then in laboratory specimens— 
did we see the eggs miss the posterior part of the male’s ventral 
surface. In these cases the eggs missed the male entirely and fell 
considerably to the rear of the pair. 

This phenomenon of egg bombardment is of special interest 
from a phylogenetic point of view. The eggs of most species of 
Hyla float. In the case of H. versicolor, it would seem that the air 
bubbles entangled in the jelly when the female raised her cloaca 
above water, caused the eggs to float. In other forms it would seem 
more likely that it was some phenomenon of surface tension (Harrison 
1922) holding the eggs near the surface where they were laid. Now 
in H. andersoni, the eggs cannot reach the surface for a very def- 
inite mechanical reason,—namely, the male is in the way. Never- 
theless, the female goes through all the movements as if intending to 


1923] Noble: The Anderson Tree Frog 439 


lay the eggs on the surface of the water. It would thus seem that 
in H. anderson, its habit cf laying bottom eggs has been derived 
from the more characteristic surface egg habit. 

The eggs of H. andersonti, although shot from the cloaca in 
bunches of from seven to fourteen (average, nine), do not adhere to 
one another. They fall to the bottom of the pool where they usually 
adhere to sphagnum or debris. Here they swell rapidly and fre- 
quently lose their attachment to the sphagnum. At Lakehurst, 
many eggs (in late cleavages) were found lying free on the bottom 
of the sphagnaceous streams. 

In nature, H. andersonii was estimated to lay eight hundred to 
one thousand eggs. None of our laboratory animals laid more than 
eight hundred eggs. 


THE EGG AND ITS CAPSULES. 


The eggs of H. andersonii may be readily distinguished from all 
other eggs found in the pine-barrens by the following characters: 
The eggs are— 

1. Single, not adhering to one another, usually scattered 
among the water weed. 

2. Attached to sphagnum (rarely debris), or free and rest 
on bottom. 

3. Found on bottom of small, non-stagnant pools, or in 
slow-moving streams of the pine-barrens. 

4. With dark cap of the animal] pole extending only over 
one-third of the surface of the egg. (Early cleavage stage.) 
Before cleavage the cap on the animal pole is usually dark brown, 

the other two-thirds of the egg, creamy-white. As the cleavage 
continues, new pigment is formed. At the end of cleavage, before 
any gastrulation has begun, about two-thirds the surface of the egg 
is pigmented (Fig. 151). Harrison (1922) has noticed a similar 
phenomenon of pigment increase during segmentation in some of 
the Australian hylas he studied. In making a comparison of the 
eggs of H. andersonii with the eggs of other frogs, care should be 
taken to use only eggs very recently laid, not those which have gone 
beyond the 382 cell stage. 

As gastrulation continues, the egg becomes much lighter in 
color. The late gastrula is pale brown, often with streaks of a 
darker tone. : 

The egg is surrounded by the vitelline membrane and by the 


Zoologica: N. Y. 


a b 
FIG. 151. PIGMENTATION OF THE EGG OF H ANDERSONII 


a. Hight cell stage viewed from side of the gray crescent. 06. Late cleavage stage, 
to show the increase of pigmentation. 


two gelatinous membranes of the usual type. The gelatinous cap- 
sules vary enormously according to their age, and treatment. The 
following measurements are taken from a series preserved in formalin. 
They agree well in size with some living specimens. 
Diameter of Ovum—1.2-1.4 mm. 
‘““ Inner Capsule—1.9-2.0 mm. 
“ Outer Capsule—3.5-4.0 mm. 

The vitelline membrane may be best demonstrated just after 
maturation when the animal pole is slightly flattened leaving a space 
between membrane and ovum. 

In passing, a word may be said in regard to egg membranes in 
general. The eggs of most batrachians possess two gelatinous 
capsules in addition to the vitelline membrane. European pelo- 
batids are stated by Boulenger (1898) to possess only the inner 
capsule, while Wright (1914, p. 16) allows us to infer that some 
American Salientia may lack the same. We should like to emphasize 
that the outer egg capsule of all batrachians is subject to great 
modification, even within aspecies. Thus, our pelobatid Scaphiopus 
holbrookii has two layers of “‘jelly’’ about the eggs. At oviposition, 
the outer layer is extremely adhesive. As development continues, 
this outer capsule swells rapidly, losing its adhesive quality, and 
changing its appearance.! We do not believe that the presence or 


1 The eggs of S. holbrookii are not twisted around the grasses in the spiral 
manner indicated by Deckert (in Overton 1914); on the contrary, they are laid 
on the upper side of grass stems which have been flattened down by the female. 
As the outer gelatinous membrane swells, the egg-masses take on the appearance 
of being arranged in a spiral, having much the same form as the egg-masses of 
European pelobatids. 


ce 


1923] Noble: The Anderson Tree Frog 441 


absence of the outer capsule in all batrachian eggs can be determined 
without investigating the structure of these membranes at the 
moment of oviposition. 

We have had no difficulty in distinguishing two gelatinous cap- 
sules in the living eggs of H. crucifer, although Wright (1914) figures 
only one capsule. 


DEVELOPMENT OF THE EGG. 


The egg of H. andersonii, in spite of its reduced pigmentation, 
shows some indication of a “gray crescent.’’ At least one side of 
the fertilized egg is paler than the other. The pale region is in no 
sense a crescent, but is an area probably homologous to the gray 
crescent of Rana. The first cleavage plane tends to cut the mid- 
point of this “crescent” at right angles. However, some irregular- 
ities occur. The second cleavage plane is as usual meridional, but 
in most eggs it cuts the first cleavage plane not at its mid-point, 
but nearer the “crescent” side. As a result, the two cells con- 
taining the crescent material are usually smaller than the opposite 
pair. The third cleavage is latitudinal. It cuts the egg at right 
angles to its axis and at such a point that on the “‘crescent”’ side the 
third cleavage furrow sharply demarcates the pigmented from un- 
pigmented region. Later cleavages are usually irregular. This is 
probably due to the fact that the second cleavage plane does not 
cut the egg systematically, but leaves less material on the “‘crescent”’ 
side than on the other. It would be interesting to know the con- 
ditions in other species of Hyla. 

With the little comparative material available to us it does 
not seem advisable to discuss the later stages in any detail. The 
changes of pigmentation which accompany the development within 
the egg have been mentioned above. The late gastrula shows some 
dorsal flexure as in Bufo and Rana. No marked differences be- 
tween the gastrulation in these groups were noted. 

Eggs laid in the laboratory hatched in four days. This is 
_ probably a much shorter period than would occur in nature. Al- 
though the period of development within the egg may be greatly 
modified by temperature, not all eggs placed under identical condi- 
tions develop in the same time. Wright (1914, p. 19) found that 
the eggs of the species he considered all developed in about the 
same time under laboratory conditions,—namely, in four or five 
days. But Boulenger (1898) has found a marked difference in the 


442 Zoologica: N. Y. Zoological Society . [Teas 


developmental period of two such allied genera as Alytes and Dis- 
coglossus. We have found that the eggs of Scaphiopus hatched with- 
in thirty-six hours, while the eggs of most species of Rana require 
five days under the same conditions. 

Lastly, a word may be said in regard to laboratory conditions. 
If a breeding pair is placed in a very small container, the chances 
that all the eggs will be fertilized and develop are very much better 
than if they are in a large jar. We experienced nene of the dif- 
ficulties with our material that Wright (1914) mentions. 


THE ADHESIVE ORGANS AND THEIR DEVELOPMENT. 


Very few have studied the adhesive organs of batrachian larvae. 
Although the form of these organs differs in the various species and 
may be utilized as a character diagnostic of the species, these organs 
have been described in only a few hylids. It is, therefore, perhaps 
not surprising that we should find that the adhesive organs of 
H. andersonui bridge, during their ontogeny, the gap supposed to 
exist between the bufonid and hylid types. 

Thiele (1888, pl. 10) found that the adhesive organs of H. arborea 
arose as two swellings, one on either side of the midline in a way 
very similar to the ontogeny of these organs in Rana agilis. Thiele 
pointed out that the more primitive method of development was 
that of Pelobates and Bufo, where the organs arise by modification 
from a crescentic furrow. It is, therefore, of considerable interest 
that we should find some indication of this crescentic type of de- 
velopment in H. anderson. 

In the early embryo of H. andersoni there appears a crescentic 
swelling on the ventral surface of the head (Fig. 152). This becomes 
slightly more pigmented than the surrounding region, but never 
invaginates to form a furrow as in European species of Bufo. As 
development continues, the two horns of the crescent increase in 
size and gradually differentiate into the definitive adhesive organs, 
while the posterior part of the crescent (Fig. 152) becomes less and 
less distinct. 

By the time the tadpole is ready to hatch, the adhesive organs 
have assumed a position lateral to the mouth. It will be noticed 
from fig. 152 that these organs, when fully formed, are not as far 
anterior as the adhesive organs of H. arborea. In H. crucifer we 
find that the adhesive organs have a similar position lateral and 
posterior to the mouth. 


¢23] Noble: The Anderson Tree Frog 443 


FIG. 152. DEVELOPMENT OF THE ADHESIVE ORGANS. 


All but the last stage occur within the egg capsules. The frontal organ (extreme anterior 
end) is conspicuous in all but the last stage. 


It will be noticed from fig. 152. that the frontal organ in H. 
andersonii is very distinct. This would indicate that it must have 
considerable functional significance. 


DEVELOPMENT OF THE TADPOLE. 


Eggs raised in the laboratory in shallow watch glasses hatched 
four days after oviposition. The recently hatched tadpoles varied 
somewhat in size, an average specimen measuring 4.5 mm. in total 
length. The color of these tadpoles was pale yellow finely stippled 
or suffused with brown. As the tadpoles grew older the pigment 


444 Zoologica: N. Y. Zoological Society [11; 18 


became darker. Approximately five days after hatching some in- 
dication of the distinctive pattern of the mature tadpole appeared. 
The pigment of the head first increased on the inner wall of the lymph 
space just anterior to theeye. This gave the tadpole a “pathological 
appearance’’ as though it carried two blisters, one on either side of 
the snout (Fig. 1530). Pigment developed slowly in the outer wall 
of this lymph space. Jt was not until just before the appearance of 
the posterior Jimb buds that the tadpole lost these “blisters.” 

The color pattern became well established in tadpoles of 11 mm. 
in length. Living specimens were uniform dull, chocolate brown 
above, golden or bronzy below. A dark stripe early made its ap- 
pearance on the upper half of the fleshy part of the tail (Fig. 153c). 
An irregular series of blotches of the same dark brown developed 
above the stripe on the upper tail fin and a few smaller ones on the 
lower fin. (Fig. 158c.) 

Only two external gills ever develop in the tadpole of H. ander- 
soni. These are pigmented like the body. Each gill consists of a 
single stalk with four branches. Three of the branches of each of 
the anterior gills become well developed while the fourth remains a 
mere bud. Only two of the branches of the posterior pair of gills 
elongate, the other two branches of each gill remaining as short 
stumps. The longest gill measures only .7 mm. (three days after 
hatching). It is about as long as the diameter of the eye (which, 
although hidden beneath the skin, is visible in both living and 
preserved specimens). 

In laboratory specimens the operculum grew over the external 
gills six days after hatching. The tadpoles at this age averaged 
8 mm. in total length. The pale coloration of the early tadpole 
had darkened, and some indication of the dark tail streak had 
appeared. Although the mandibles were well formed and pigmented, 
no horny teeth had yet developed. The vent at this stage had just 
begun its asymmetrical growth which soon resulted in its character- 
istic dextral twist. 

The horny teeth began to develop immediately. As the tadpole 
matured, the teeth increased in number. This may be seen in our 
series of preserved specimens. Ina tadpole of 11.5 mm. total length, 
the number of teeth in each row, reading the rows from above down, 
is as follows: 38/10 + 10//31/24/5. In one of 17 mm., the formula 
is: 52/21 + 21//47/45/21. In a mature tadpole of 32.5 mm. total 
length and having the limb buds well developed, the formula is as 


1923] Noble: The Anderson Tree Frog 445 


follows: 96/36 + 35//67/82/40. The number of teeth in each row 
is closely correlated with the relative extent of the row. Hence, 
little may be said about the diagnostic value of the teeth rows of 
H. andersoni unless they be compared with those of a tadpole of 
the same age. The mouth parts figured above (Fig. 153a) are those 
of the tadpole figured (Fig. 153c). 

Perhaps the most distinctive feature of the tadpole of H. 
andersonit is its short and narrow tail jfin (Fig. 153c). This gives 
the tadpole a Rana-like appearance, or at least permits one to readily 
distinguish it from the tadpole of H. versicolor, or that of H. arborea. 
The question may be raised: is this reduced fin an adaptive feature? 
It may possibly be such, but the evidence at this time is by no means 
clear. The tadpoles of H. versicolor live for the most part in quiet, 
weedy ponds and these broad fins aid them to make quick turns 
very much in the same way that flattened or deep-finned fishes are 
able to dodge suddenly when avoiding an enemy. But let us look 
further. Most of the vertebrate inhabitants of the pond are deep- 
bodied or deep-finned. The ambystomid larvae have a back fin 
which undoubtedly serves them in their jerky dashes. A glance at 
a neighboring brook and we have a different picture. Here most of 
the forms have reduced the fin and have adopted better “stream 
lines.’ This is especially noticeable in the larvae of Desmognathus 
and Hurycea. The rule holds true for practically all mountain 
brook salamanders as Rhyacotriton in this country and many exotic 
genera. In the puddles of the slower streams we sometimes find 
the narrow-finned tadpoles of Rana pipiens or Rana clamitans. 
The comparison, however, between brook salamanders and brook 
tadpoles may not be drawn too closely. Frequently narrow-finned 
tadpoles occur in the ponds. It is interesting, however, that the 
tadpole of H. andersonii dwells primarily in the slow streams of the 
pine-barrens, and as if in adaptation to the current, it has given up 
its broad “pond life fin’ for the sake of a more efficient one. We 
say “given up,” for it seems probable from what has appeared above, 
that the species was evolved from a type having much the habits of 
H. versicolor. Whether or not we have pushed too far this compar- 
ison of brook salamanders and brook tadpoles, the fact remains that 
H. andersonii with its short fin has much more the habits of a Rana 
or a Bufo tadpole than it has the characteristic Hyla mannerisms 


(see below). 
Metamorphosis first occurred at Lakehurst in the area under 


446 Zoologica: N. Y. Zoological Society [15-28 


Sele Peas cas tae 


5 SED 
Drei! maa 


FIG. 153. TADPOLES OF HYLA ANDERSONII. 


a. The mouth parts of mature tadpole. 06. Early tadpole, showing the conspicuous lymph 
sacs on either side of the snout. c. Mature tadpole, showing the characteristic pattern. 


observation on July 23. None of the laboratory animals reached 
metamorphosis, probably because of our limited supply of bog water. 
Metamorphosis usually followed the day after the right fore-limb 
appeared. The left fore-limb appeared usually a day before the 
right limb. The spiracle became greatly widened to permit the 
passage of the left limb through it. The usual phenomena of 
metamorphosis occurred. The head widened, the mouth changed 
its shape and the body decreased in size. The head and body 
length (excluding the tail) of five tadpoles, having only the posterior 
limbs present, averages 13.1 mm., the head and body length of 
five others, having all four limbs present and the head already 
changed in form, averages 12.1 mm. Metamorphosing tadpoles 
became slightly greener in coloration but did not assume the full 
color of the adult while in the water. 


DESCRIPTION OF A MATURE TADPOLE. 


A tadpole is said to be mature at the time of the appearance of 
the hind limbs. In some forms this does not correspond to the full 


1923] Noble: The Anderson Tree Frog 447 


development of larval characteristics. In H. andersonii the larval 
color pattern may or may not be complete at this stage. For this 
reason, we have utilized slightly older specimens in drawing up the 
following description. As is customary in the describing of adult 
batrachians, the detailed description is based upon a single typical 
specimen, the diagnosis upon several specimens. 

Diagnostic Characters Spiracle sinistral], anus dextral, eyes visible 
from the vental surface, upper fin crest not extending beyond the 
vertical of the spiracle, distance from spiracle to base of hind limb 
contained about 1.3 times in its distance from the snout; labial 
teeth 2/38. Uniform brown above, yellowish on the tail, a con- 
spicuous irregular stripe of dark brown extending the length of the 
tail. Greatest length of tadpole, 35 mm. 

Detailed Description Length of body contained 2.6 times in the 
tail length; width of body 1.7 times in its own length; nostril nearer 
the eye than the tip (midpoint) of snout; eye dorso-lateral, visible 
in part from the ventral surface, nearer the snout than the spiracle; 
distance between nostrils contained 1.66 times in the intererbital 
width, exactly equal to the width of the mouth; spiracle sinistral, its 
distance from the base of the hind legs 1.29 times in its distance 
from the snout; anus dextral; depth of the muscular portion of the 
tail at its base contained 2.5 times in the greatest depth of the tail. 

Upper labium with two series of teeth, a boundary row of teeth 
and an inner or lateral row on each side (Fig. 153a); the median 
space between these two lateral rows only a third the length of one 
of the lateral rows; three continuous rows of teeth on the lower 
labium, the second or median longest, the outer or boundary row 
slightly more than half as long as the median one; a complete circlet 
of papillae around the mouth, broken only for a short space along 
the upper median margin; a clump of papillae at either corner of the 
mouth, medial to the boundary papillae. 

General color (formalin preservation) of the body, chocolate 
brown above, translucent below; tail yellowish; an irregular streak 
of dark brown running the length of the fleshy part of the tail just 
dorsal to the median line; lower border of the fleshy part of the tail 
irregularly spotted with a slightly paler brown; tail fin both above 
and below streaked or finely spotted with brown; the streaks some- 
times forming irregular stellate figures but never a network. 

In life the color pattern was the same, the brown and yellow 
tones of about the same intensity. The belly was very different. 


448 Zoologica: N. Y. Zoological Society [Tbs tse 


It was golden, irridescent or whitish, according to the direction of 
the light. The throat was yellowish. The iris was golden, ver- 
miculated with black; the black pupil was large and round. Some 
indication of internal structure visible, especially the nasal passage 
and two of the posterior cranial nerves. Lateral line system feebly 
indicated. 


Measurements 
PotalLength::.6 ae set: eee eee 31.0 mm. 
Greatest length of head and body ...... Zhe 
Greatest denethiof tail. sats cecal aalBip 
Greatest depth of tail................ Lae 


HABITS OF THE TADPOLE. 


Larvae raised in the aquarium and those studied in the field 
had similar habits. The larvae of H. andersonii are not active 
swimmers. At Lakehurst, these tadpoles seek out the shallows 
whether or not these be weed-grown. In such favored places, great 
numbers of tadpoles were found resting motionless just below the 
surface with dorsal crest touching the surface film. When ap- 
proached they dived quickly into the nearest masses of sphagnum. 
Tadpoles of H. anderson exhibited similar resting and. diving 
behavior in the laboratory. No other tadpoles with which we are 
familiar make such erratic plunges into concealment. 

Laboratory specimens ate some fish food (dried shrimp) and 
some of the water weed in their aquaria. They invariably skeleton- 
ized within a day any of their companions that died. 


Foop HABITs. 


The food habits of H. andersonai are in no way specialized. 
This is to be expected since it has been shown elsewhere (Noble, in 
press) that the food habits of most tree frogs do not radically differ 
from those of frogs living near the water’s edge. Frogs and toads 
seize anything of small size moving in their vicinity. 

The stomachs of ten males which were captured during June 
while calling from bushes or low trees contained the following food: 
5 grasshoppers (two species); 2 beetles, 3 ants (2 species), 1 dipterous 
insect, 2 dipterous pupae (tabaniid?), and some unidentifiable 
insect remains. None of the specimens taken in embrace contained 


1923] Noble: The Anderson Tree Frog 449 


food in their stemachs, but only a few pairs were killed immediately 
after oviposition. 


RELATIONSHIPS. 


H. andersoni has been generally considered a close relative of 
the European Tree Frog. Long ago Cope (1889) said of H. ander- 
soni, “in proportions and general appearance similar to Hyla 
arborea of Europe.” Since then everyone who has had the occasion 
to consider the relationships of H. andersonii has agreed that the 
resemblance was very close. More recently, Barbour (1914, p. 239) 
has expressed the opinion that Hyla pulchrilineata of Santo Domingo 
was allied to Hyla arborea. We have recently had the occasion to 
study H. pulchrilineata in the field, and could find very little re- 
semblance between these two species in either color, structural 
characters, voice, vocal-pouch, breeding habits, or in any other than 
generic characters. In a paper now in preparation, we have con- 
cluded that the two species are only distantly related. 

As pointed out above, the object of the present paper is to 
describe those features of the habits and life history of H. andersoni 
which might shed light on its relationships. As no one has pre- 
viously attempted to ally H. andersonii to any other species than 
H. arborea, it is important that we should first consider the re- 
semblances and then the differences between the two species. Our 
information in regard to H. arborea is taken chiefly from Boulenger 
(1898). 


Resemblances between H. andersonii and H. arborea. 


1. General color and proportions. 

2. Small size. 

3. Many structural features,—as smooth skin, position of 
vomerine teeth, form of nuptial asperities, form of vocal pouch, 
etc. 


Differences between H. andersonii and H. arborea. 


1. Color pattern differs in many details of which the most 
noteworthy are as follows: the lumbar and the dorsal spots 
frequently found in H. arborea are never present in H. ander- 
soni; the ground tone of H. arborea is subject to variation of 
color, of H. andersonii, to only a change of intensity; the details 
of coloration of thighs, throat and often the appendages differ 
remarkably in the two species. 


450 Zoologica: N. Y. Zoological Society [II; 18 


2. Contracted pupil of H. arborea diamond-shaped; not 
so in H. andersonit. 

3. Fingers slightly webbed in H. arborea; free in H. ander- 
sonit. 

4. A strong odor of “raw peas’”’ from H. andersoni after 
handling; no such odor from H. arborea. 

5. A marked sexual dimorphism in H. andersoni; not so 
in the other species. In H. andersonii, as pointed cut above, 
there is a difference between the sexes in the size, in the color 
on the sides of the throat, and in the ground tone of the throat. 
These differences do not appear in the several specimens of 
H. arborea before us. <A breeding pair taken at Blois, France, 
measures 42 mm., total length in both sexes. A female of 
H. arborea from Germany measures 41 mm., while four non- 
breeding males from Germany measure 40.5, 40.5, 36.5 and 35 
mm. respectively. Thus, there might be a slight difference 
in size between the sexes of H. arborea, but this difference is 
not constant. There seems to be no sexual dimorphism in 
H. arborea other than some indication of breeding asperities 
in the male, and sometimes a difference in size between the sexes. 

6. The call of the two species is radically different. The 
following notes have been kindly given us by Dr. J. P. Chapin. 

“In company with Dr. R. E. B. McKenny, at Blois, on the River 
Loire, France, during April, 1918, I found eight or ten individuals of 
Hyla arborea assembled just after dusk in a temporary pond in an open 
grassy field. Their notes, by which we were attracted, bore no resemblance 
to the voice of Hyla andersonii, with which I was very familiar; on the 
contrary, they produced a confused, hoarse, croaking chorus, which re- 
minded me far more of the voices of common European toads. There was 
nothing of the curious nasal resonance of the “‘quank’’ of andersonii. 
The behavior of the individual frogs, too, was very different. They were 
all in the water while calling, not perched in bushes, as is usual with 
andersonii; and from the number of them in one or two small pools, I 
might describe them as far more sociable. In view of the striking external 
resemblance between andersonii and arborea, I was greatly impressed by 
the dissimilarity of their voices and actions.” 

7. The habitat of the two species differs greatly,—H. ander- 
sonit being confined to the pine-barrens, while H. arborea has 
a wide distribution in many types of country throughout 
Europe. 

8. The breeding cites of H. anderson are always shallow 
sphagnaceous streams or puddles on the pine-barrens, while 


1923) Noble: The Anderson Tree Frog A451 


H. arborea selects ‘‘deep pools or ponds of clear water, more 
or less richly endowed with vegetation,’ (Boulenger, 1898, 
p. 258). Thus, H. arborea agrees with H. versicolor in the 
selecting of a breeding cite and differs remarkably from H. 
andersonii. Deep ponds are available to H. andersoni but it 
selects only the small pools. 


9. Eggs of H. arborea are deposited “in several lumps, 

. attached to weeds below the surface of the water”’ 
(Boulenger, 1898, p. 259) Many more eggs are laid at one 
time by H. arborea than by H. andersonzi, and these are adherent 
in the former species, not in the latter. From the form of the 
egg-masses, their attachment to weeds, and the number of eggs, 
it is apparent that the methed of oviposition in H. arborea 
must be very unlike that of H. andersoniv. 


10. The external gills of H. arborea are “unbranched or 
bifid’; in H. andersonii there are only two pairs of gills, the 
posterior pair having two well developed branches, the anterior 
pair, three such branches. 


11. The adhesive organs arise separately in H. arborea; in 
H. andersonii they develop from a crescent somewhat as in 
Bufo. The final position of the adhesive organs is more 
anterior in the former than in the latter species. 


12. The mature tadpole of H. andersoni differs radically 
from that of H. arborea in the extent of its fin crest, and in its 
coloration both above and below. It also differs in having 
its eyes more dorsal and in having somewhat different pro- 
portions. 


13. The mature tadpole of H. andersonii differs from that 
of H. arborea in its habits. It is a slow-moving form, accus- 
temed to bask in the sun with dorsal fin in contact with the 
surface film. The description of the tadpoles of H. arborea given 
by Boulenger reminds us very much of Scaphiopus tadpoles for 
they are found “swimming about like fish in every direction.” 

14. Activity of the adults of the two species differ. H. 
andersonii does not stick well to smooth surfaces; when it 
climbs it frequently grips the branch with opposed fingers 
(Fig. 1482) and the feet wrap around the support. The 
movements of H. arborea agree well with the majority of hylas. 


452 Zoologica: N.Y. Zoological Society (11; 18 


The above differences are far too numerous to be disregarded, 
for some of these differences are of considerable consequence. The 
resemblances between H. arborea and H. andersonii do not outweigh 
the differences. We have examined specimens of H. regilla which 
seem as nearly like H. andersonii as do some specimens of H. arborea. 
The resemblance in color pattern may be due to convergence, for 
Boulenger looks upon the spotted pattern as the more primitive 
and ancestral to the unspotted type, at least in the H. arborea 
group of forms. Further, Boulenger (1898, p. 252) considers that 
Hyla immaculata, described by Boettger, from China as a race of 
arborea “‘cannot be united with H. arborea ... as it lacks the 
web between the fingers.” H. andersonii, too, lacks the web 
between the fingers, and the question is immediately raised whether 
it might not be closely allied to H. ammaculata. 

It may be further pointed out that both botanically and her- 
petologically there areas good a prior? grounds for seeking the ancestral 
stock of a form, at present restricted to eastern United States, not 
in western Europe, but in eastern Asia. Cryptobranchus and 
Leiolopisma are two striking examples of American forms having 
close allies in China. To this list we may now add H. andersonit. 


CONCLUSIONS. 


1. H. andersoni is not closely related to H. arborea nor to H. pul- 
chrilineata. 

2. H. andersonii has been derived from a group of hylas which laid 
surface eggs, its method of oviposition being a modification 
of their method. 

3. H. andersoni exhibits a primitive method of adhesive organ 
formation. 

4. Voice plays an important réle in the mating of H. andersoni and 
probably in other American tree frogs. 

5. H. andersonii, by its coloration (including sexual dimorphism), 
method of oviposition, distinctive tadpole, and restricted 
habitat, occupies an isolated position among American species 
of Hyla. 

6. The relationships of Hyla andersonii are to be sought in Chinese 
forms and probably in H. immaculata (Boettger). 


BIBLIOGRAPHY 
ABBOTT, CHARLES C. 
1868 Catalogue of Vertebrate Animals of New Jersey, in Cook. Geology 
of New Jersey, Appendix E. 
1890 Voice of Hyla andersonii. Amer. Nat., Vol. XXIV, p. 189. 
BAIRD, SPENCER F. 
1854 Descriptions of New Genera and Species of North American 
Frogs. Proc. Acad. Nat. Sci. Phila., Vol. VII, pp. 59-62. 
BARBOUR, T. 
1914 <A Contribution to the Zoogeography of the West Indies, with 
Especial Reference to Amphibians and Reptiles. Mem. Mus. 
Comp. Zool., Vol. XLIV, pp. 204-359, 1 pl. 
1916 A Note on Two Interesting New Jersey Amphibians. Copeia, 
No. 26, pp. 5-7. 
BLES, E. J. 
1906 The Life-History of Xenopus laevis, Daud. Trans. Roy. Soe. 
Edinburgh, Vol. XLI, pp. 789-821, pls. 1 to 4. 
BOULENGER, G. A. 
1882 Catalogue of the Batrachia Salienta . . . of the British Museum. 
London. 
1898 The Tailless Batrachians of Europe, Parts I and II. London. 
1912 Some Remarks on the Habits of British Frogs and Toads,..Proce. 
Zool. Soc. London, pp. 19-22. 
Cope, E. D. 
1862 On some new and little-known American Anura. Proc. Acad. 
Nat. Sci. Phila., Vol. XIV, pp. 151-159. 
Courts, S. A. ; 
1907 Response of Toads to Sound Stimuli. Amer. Nat., Vol. XLI, 
pp. 677-682. 
CUMMINS, HAROLD 
1920 The Rédle of Voice and Coloration in Spring Migration and Sex 
Recognition in Frogs. Journ. Exp. Zool., Vol. XXX, pp. 325- 
343. 
DAVIS, WILLIAM T. 
1904 Note on Hyla Andersoni, Baird. Proc. Nat. Sci. Assoc. Staten 
Island, Vol. IX, p. 26. 
1904a Hyla andersonii and Rana virgatipes at Lakehurst, New Jersey. 
Amer. Nat., Vol. XXXVIII, p. 893. 
1905 Further Note on Hyla andersonii and Rana virgatipes in New 
Jersey. Amer. Nat., Vol. XX XIX, pp. 795-796. 
1907 Additional Observations on Hyla andersonii and Rana virgatipes 
in New Jersey. Amer. Nat., Vol. XLI, pp. 49-51. 
1922 Insects from North Carolina. Journ. N. Y. Entom. Soc., Vol. 
XXX, pp. 74-75. 


453 


454 Zoologica: N. Y. Zoological Society (11; 18 


DECKERT, R. 
1918 Two Interesting Tree Toads. The Aquarium Bulletin, Brooklyn, 
N.-Y..,. pp: 125-126. 
DICKERSON, MAry C. 
1906 The Frog Book, New York. 
DITMARS, RAYMOND L. 
1905 The Batrachians of the Vicinity of New York City. Amer. 
Mus. Journ., Vol. V, pp. 161-206. 
FOWLER, HENRY W. 
1907 The Amphibians and Reptiles of New Jersey,in Ann. Rept. N. J. 
State Museum for 1906, pp. 238-250. 
1909 Notes on New Jersey Fishes, Amphibians and Reptiles, in Ann. 
Rept. N. J. State Museum for 1908, pp. 349-408. 
HARRISON, LAUNCELOT. 
1922 On the Breeding Habits of Some Australian Frogs. The Austra- 
lian Zoologist, Vol. III, pp. 17-34. 
HARSHBERGER, J. W. - 
1916. The Vegetation of the New Jersey Pine Barrens, An Ecological 
Investigation. Philadelphia. 
KUNITOMO, K. 
1910 Uber die Entwicklungsgeschichte des Hynobius nebulosus. Anat. 
Hefte, Vol. XL, pp. 193-283, pls. 13-16. 
LIVINGSTON, B. E. and SHREVE, F. 
1921 The Distribution of Vegetation in the United States, as Related 
to Climatic Conditions. Publ. Carnegie Inst., Washington, 
No. 284. 
MILLER, NEWTON 
1909 The American Toad (Bufo lentigenosus americanus, Le Conte). 
A Study in Dynamic Biology. Amer. Nat., Vol. XLIII, pp. 641- 
668. 
MILLER, W. DE W. 
1916 Notes on New Jersey Batrachians and Reptiles. Copeia, No. 34, 
pp. 67-68. 
Moorg, J. PERCY ; 
1894 Note on the Occurrence of Hyla andersonii in New Jersey, Amer. 
Nat., Vol. XXVIII, pp. 1045-1046. 
NOBLE, G. K. 
1922 The Phylogeny of the Salientia I. The Osteology and the Thigh 
Musculature; Their Bearing on Classification and Phylogeny. 
Bull. Amer. Mus. Nat. Hist., Vol. XLVI, pp. 1-87. 
1923 A Contribution to the Herpetology of the Belgian Congo III, The 
Amphibia. (In press.) 
OVERTON, FRANK 
1914 Long Island Fauna and Flora, Vol. III, pp. 21-40, pls. 2-13. 
PETERS, JOHN E. 
1889 Another Specimen of Hyla andersonii. Amer. Nat., Vol. X XIII, 
pp. 58-59. 


1923) Noble: The Anderson Tree Frog 455 


SHERWOOD, WILLIAM L. 
1898 The Frogs and Toads Found in the Vicinity of New York City. 
Proc. Linnaean Soc. N. Y., pp. 9-24. 
STONE, WITMER. 
1901 Occurrence of Hyla andersonii at Clementon, N. J. Proc. Acad. 
Nat. Sci. Phila., Vol. LIII, p. 342. 
1906 Notes on Reptiles and Batrachians of Pennsylvania, New Jersey 
and Delaware. Amer. Nat., Vol. XL, pp. 159-170. 
1911 The Plants of Southern New Jersey, with Especial Reference to 
the Flora of the Pine-Barrens and the Geographic Distribution of 
the Species. Ann. Rept. N. J. State Museum for 1910, pp. 25- 
288, 120 pls. 
THIELE, J. 
1888 Der Haftapparat der Batrachierlarven. Zeitsch. f. Wiss. Zool., 
Vol. XLVI, pp. 67-79. 
WELLMAN, G. B. 
1917 Notes on the Breeding of the American Toad. Copeia, No. 51, 
pp. 107-108. 
WriGcut, A. H. 
1914 North American Anura, Life Histories of the Anura of Ithaca, 
New York. Publ. Carnegie Inst. Washington, No. 197. 


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