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Deere ww TS oy AS I Nd a) \wad ed ag el ead I AGS IGYY| y WIG AS LIGIISYISY So SIU we we ed ~yewewwywy Yew YS» Welle ivy nad et Oe Oe UMW yall YY ev revere e tenet es CUCUUR IE 29 NINN) 1S dag ade de vv AIG Sw : ' Vi WN wl Vw wv wd ed ed y Wit, MISS gery yu AG a4 SY OC Nat lal ele eee ruedereeseaee eos ove SIGS aly MWS oer Ne ee Scien" ube A Oy VOVONUNYYNTTON Elta devvvt of bdlit IOS J She i ~ / ~~) \. SO JS “iw WOseee SS | | wee viv SSAC 4 . ee Uveveereeeveueceuusuy . SI TRAS Ned Si \ peeeoe SS auch WWW SSS PISS SUEY sl Ww Oe Bag MONS CY Vell aly AAU ls YSOLES AAS A, WEIS UU SSSSGIS EU UCN vosvotonwaeiwoeipeo ooo coiee ewe OV SO nik AA Oi) Ae iS i NA Ww iS } ' , 4 ed we ~~ | Vis } WISE ES JJISEUSSSSY MSI FL AA AAAS z te DIIGYS : II Ne iS ANS ADA) Mer MOMACUMAALECEE en re we INNS US SSMU oul! the SII Mavele wk error voveueeNth ue yuvgee ele \ ed | Wey wy SIS YS SS ca ay Sede www veecooasnneeee | Mehe) Yh rfid iy) vey we aye wwe wulvu eu SACS Wi vue wl ISS DAA GOO ce I_w Mosse sive gy SAN SHAFUSONY| jones, Toaatht VIN CH | CHS AGRAGAYS 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) Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 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 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 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 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 a oo - 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 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 aondsyuotg °¢ ‘ON “TT ‘J0A 091601007 ‘eFl] WO Surmerp e@ worl “SUO] SOU! Je}JeNb-suo pue usejaulU ‘a[BUIOF B JO MOIA [e107 e"T - HSIALVO 'TIVSdOLAAVS 02 ‘Dla 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. ns a : Pr # ’ — a ar . | ’ AL? ; ! erties tebe ye Gee) sath a SD - . ' ‘ mae yy Le ie, a" A 4 ‘in ie i rey! he Dey A wh te ee : as f r ‘ ys F ‘ é i 4 t 4 i 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), 174. In Historia Naturalis Brasiliae by William Piso and George Marcgrave, Amstelodami. Mrrcuitt, Samvuet L. 1815 The Fishes of New York Described and Arranged. Transactions Lit- erary and Philosophical Society of New York, vol. I, pp. 433-434. 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- onomic Survey, pp. 62-63, Raleigh. 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 Vicinity. .No. 3—Fishes. Proceedings Academy Natural Sci- ences of Philadelphia, vol. X XIX, p. 216. 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 ORI SIGE TEGO) SRE = plas LD UC aie ca a er U9 Me RE MNVATIUAR NG sul cis Mime 80 iss Sn tae 2 an, Se PPI Ie ot Wiehe etek eegare sabato oo iy ee yO LANE NROO ut ATS one) eee, cite Ries ais een ce I LOO ESEWAIGAINCER arene ate miicay UIA. crigtitiast as sede Mba hor ke og BE AG tr Nate e eo et am ea ea a Fah oe ee es ata ee gag ce LOG UDG ETAT ANCHE oes Gelet ec Soren eke, cha a) 1 et BOE PUSOUSIM lata nro Ugete ee ore tn RS OS BANDICOOTS Eo a eS t=, Se arte, Ober imi i Saar eae Fs 72" WOMBATS SE REE mie elas meat eer s.) ee oe SRP MEAN VW OUN Et ot: hacer en ek ae eS aa LOD TSOP “DEVEbA@ ratte ta ee enl se dent -e > LOO SHOE DM OAGTURE 5) ot pepe te te ae fy ew el-eee-) LOG MARSUPIAL MICE Me Gera ee Pees at as tae ens Wegner Nd ECHIDNA Mes

ee ae oO MARSUPIAL MOLE 2 ; : : : : : : ee OH 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 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Facing 169 Between pages 176 and 177 Facing 179 Between pages 192 and 193 Facing 197 Facing 198 Facing 198 Facing 201 L9T abnd aony "9 ON ‘TT “0A “parbo 007 . ABp Joy B UO JISIA 0} SadR[d [NJIYSI[ep cae SABMI[E SOl[[NS usog,, “WAOMYIIEY JURIDH pu JOYSYSULY WUBI ‘SUIqoY poys¥eIg-esoYy PUB pesvetg-MOT[O X ‘AQBII@M P2leL-YoV[_ CuI ‘palgq a1AT o1eI Mou oy} JO JUNVY 9}IMOART 9Y} OSTe SI ULag oeL], [NJIynveq ey} Jo ouloY SIY4T, VIIVULSNV ‘VINOLOIA AO AONIAOUd AHL NI LSHYON TVOIMAL V “Ge “Old fonogeT “CHM fq ydv.ubozoyg 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 691 abpd voDqy "9 “ON ‘J[ “WA ‘*VvI1b0j007 “OFT P[[M Jo UONwAdoSei1d 94} AOJ SalienjouRS Jo Salles s[qeyIeuWiet e@ puB ‘Suapies [BO1dO[00Z Jo ULBYD Joy DULUIEUIEU A[SNOIOSIA SI BI[BAISNW ‘1eM JY} LOJ AQUOUI PuB Sd1ys ‘SUOI}IUNUI ‘usu JO SulInod}no snopusuiet} Jay SUIPUeISYy}IMION {eljeusny usle{saM PUB YI419g JULISIP-IBJ YIM BII0JOIA pue Asuplg ‘eueqsiig Syul] MOU ABATIVI [BJUaUIUODSURI} JeaIs B BY. MOUY SUBOIIOUIW AUBUI MOFT ‘UOIZeUIZBUIL 9y} SUNAINO eeISnY JO JUsUIdO[eAep au, FuIdWa NYALSVALNOS S.NIVLIYE LVAD :-VITVULSNV “8 ‘Dla dusiod “s0NWH Gg Or 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 ‘yale [Bo1So[ooZ yAoK MAN OOUVONVY GUVMAGOOM ‘96 “DIA rE . « " - 5 a _ . i age: ULOQUDS “Yay fig ydvsboj0Yd oy s oe o ao ee Jee kg ae Photograph by E. R. Sanborn BLACK-FACED KANGAROO FIG. 37. New York Zoological Park gage 253427! we 835 tdetaer: eee” eaeqne ast @ todedi ease 33 " a SedEt Qe CHW? proche PARRY KANGAROO FIG. 38. New York Zoological Park. *yadeq [Bolsoj00Z yIOK MON OOUVONVHA Gay LNVIO ‘68 Old ew | SERS88 GO Oe ee o4-g SER BA ER fence Eee A oe 8 ekg OR eae ig lene eee ta) Sea es. a reese ee ee: teseccaseaaS sarees S22": 7° o eRee ae Peres bebbee verse cet : oa : Seon co ee ree SESE EERE 8 wee 8 Fees eet pode bode peek Bac es RESTA ME ARES Teese = ee | ees se oO a Se ees oe PEOMON RRS Cee ae eo ooo. 2 ee ibdidedpde i Le $4 ‘s tx EE SBR BE Ha LOREM BB Se O SRM eae ee we eee ta a (sees. rye emaeee cones 988 Staelin eeeeee C bd & # e e PEG REA BH EE i . : La a si a qrasesca ie Megsesses: eee eee s . ~ Saher DRS oe LRT BR -: 2ee8 2S SR ONG Se aa, ee Skee ee Sean Seerwed eae * ROMER SHREES kt One a SERS BS oe ee ae aa a 4 a * = a s * chet see ona e : * * > < * * * * > * & oe ett. pe tea i Ree epmperi rss bese ee ere dae of sectnee ieeaaeee © fetes seers: feet. Eaeaegeene* ighis openteee ls i eer es a breiies ppgte res teeee siegeaeels f lileeressiee 24% or epee! ee jake eet se a is Soy ae Photograph by E. R. Sanborn 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. “Blleaysny ‘VUINOg|ay[ 3B SUapIey [BoLdO[OOZ ayy UT “youRaq 0} YoUBIG WOAy duInt A[quIiU pues quII[D yyOq AeUT, SOOUVONVY GHUL HAIL VO “eh Old ; OOUVONVM ATUL LLANNAA “Zp OTA "yale q [Bo1sojoo7 yloX MAN AGVTTIVM GHMOANSNONNY FH Ol — = < - : , i : z Na - ' “e oe Te : : : ee - th FS hg PEG ey ‘ ee ee ‘ *yooy puly oyj UO sped surmoys ‘aj ddiu 07 peyoe}4e suno x snupyzauueg “JT ‘OOUVONVY ACYL LLANNGA “9h Ola ATVIOOSVHd GUTIVLHSNYd “sh “Old HM fg yan.bo,0YT fEn0gaT “WT '‘H “M fiq ydv..boz0Y4Q eA fanogaT *q* Photograph by W. H. D. LeSouey 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. “yIe@q [BoIsojooZ YAOK MAN SGIAVTTIVM GATIVI-HSNd “0S “Old ue tet me eon ete a Oat ey = Sie see Se Se eae ee wk a) mt a we a Kat eaes eas ee ieesetes see edt ae ete —o% oe OA AAA EN TL wee weet pe enea er ae ar ET ETeTy ays ik Bere “eet pe tetra” ; ee te a en “YAvq [Bo1LZojooZ yaoX MON “ooURqIN4SIP 489745118 94} UOdN AaYyjoUI ayy oJ poysna ynq ‘pauvom AT[NF SBM UIUIJDads BUNOA SY], “UMO.LS palyjz-ou0 [IQUN Yonod §,194,OU BY} UI a¥njor yYoos sooresuey SunoZ AGVTIVM-dNVMS MOVIE ‘TS ‘Dla udoquog "yy “gq Nq ydn.60,04q aie | rs My a er * sad ; Di Pe ee tae’ « ial “~ Te a * ; we as s Cy “yIBq [BoLso[007 YOK MON “UMOYBS A[1B9]O SI JOEJaLOJ BY} JO UOIZY[NdiueUW SNO1ezXep ay, AAVTIVM-dNVMS FHL 40 WIVN “2S “Old ULOQUDS “YY fig YdvLboj,0YT 441 abpd oop “9 “ONT ‘TT "JOA ‘D21H0]00Z “yAe@q [BIISolooZ yIOK MAN GVTIVM ATIOV (‘&$ “DIA 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 a ite 641 ABA Sa0Dq 9 “ONT ‘JT “JOA “p9160]00Z7 pur[susent) AGVTIVM AAAL AHL AO ANOH AHL FS “DIA 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 “BI10JDLA Ul YSN 9yy UI pjIM peydeisojoyd 1v9q eAljeU plo uy 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 “yaleg [eo1sojooz yaIox MaN ATOM TVICOSUVW -AIOM NVINVNSVL ‘F9 ‘DId 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. L6. abnd aon °9 “ON ‘TT "JOA “va1h0j007 “S[BWLUB SUIAT] JO SUOIZD9|[OD UL UBES ST Ajjuenbeszy pue AzIAIVdvO UL Spaeiq 4] “Mel JO Yj}SudI}s pue Iodula} VSBABS S}I WOT A[jaed pue ‘10jod 841 wWoaz A[jaed oureU S}I pdAlodeI sey [BUIIUB Suippiqioy aeyjes siyy Ajjuoreddy suepiey [BoIdO[OOZ VuUINOeT STIAGG NVINVWSVL JO ATINVA V “99 “Old 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.

  • 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 Paper Cloth _A Monograph of the Pheasants (Beebe), Vol. I—ready 62.50 Published in four royal quarto volumes. Illustrated ‘ with reproductions in colors of paintings by Thor- burn, Lodge, Knight, Fuertes and Jones, and many photogravures and maps. Four volumes, $250.00. Our Vanishing Wild Life (Hornaday)............. 1.65 Destruction of Our Birds and Mammals (Hornaday). .15 Notes on Mountain Sheep of North America TVS DEON Sie en SORE i OR oe aa RR PEROT Tart Ps BOER 40 DO ITO ROTOUE) soos ck e-Oek Wo see ea és 40 The Origin and Relationship, Large Mammals, North PIED Be AS TOILE Pos aie o's eos sos baa gow ha es 1.00 The Rocky Mountain Goat (Grant)... ......06....... 1.00 Tropical Wild Life (Beebe; Hartley; Howes)........ 3.00 Zoologica, Vol. I, Nos. 1-20 inclusive, set ............ 3.85 6.00 Zoologica, Vol. II, Nos. 1, 2, 3 and 4, 5, copy.......... sa POMotiPe VOL 1 INO. OcCOPY sss oy KR ee OS 75 Zoologica, Vol. II, Nos. 7-8 and 9, CONN CS eee 2b Zoopathologica, Vol. I, Nos. 1 to 3 CODY o 5.5 Woes oe 25 Bulletin: Bi-monthly. Single copies 20c. Yearly by mail 1.00 Bulletin: Nos. 24 to 60 inclusive, cloth bound........ 10.00 Official Guide, Zoological Park (Hornaday), boards. .40, postage 6c. Post Cards: 63 Subjects in colors, in sets of 21, set. .25, postage 2c. Souvenir Book: 48 pages, 78 four-color illustrations. .50, postage 8c. Animals in Art Stamps: 32 page album; 120 mounting spaces; 120 four-color animal-stamps—complete... .75, postage 6c. Wild Animal Stamp Primer: 96 pages; 49 animal stories, 50 mounting spaces and 50 four-color animal stamps—complete .......5.......-0.205-- .85, postage 7c. Panorama: Zoological Park, colored—flat or folded.. .10, postage 2c. Photo-Letter: 18 pictures, four colors—two letters... .10 Photogravures: Animals and views in the Zoological Park; 12 Subjects. Sepia—two prints............ 25 a Enlargements: 12 animal subjects—1l1 x 14 ‘prints: AAS aNd White, CACH SOs. goatee. fda wah ee Ws 5 eJe A 25) WuGtene-— Brown, 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 ‘eiajydouawAP ay} Suoure juasaid sem que uy ‘p ‘jdog ‘sjoasuT asay} Jo syed yyos pajysaz -1p Apaed ayy A[qeunsaid sem s9uI0F ayy, ‘sajjaeq Fo sjyivd prey ay} YAM 19q3950} ‘ajsed 314M B pautejuod INOF pue Safja—aq punoid uaye} pey om} ‘Ay ojeyng & uajea pey [eNprArpur auQ “¢z Avy] ‘apaday[iu e fo Juawsesz v Aq pajuasaidar a19M spodo| -diq ‘IT ‘Judy ‘sutewias yasur peynuaprun ay} Suowe sem vAie] ayl[-joseur eB pue ‘afj}aeq prjAydorpAy v uajwa pry Yysy yy, ‘snivqupvry & Jo aka ay} JO paysIsuod SUIVWAI UBadRIsNID 94], “PT YIryAl ‘930 ‘xafign yf FO Sate ‘Spljauue [[eWS a19M spod —-012YD 7 [[Y ‘saywos yons Sursvaq Ajt[Lso] sty} Fo YSy JULpUnqe A[UO ay} ‘DuvsSoajJ0g e WOIZ Ajqeqoid ‘ajvos proua}d pajsasip Ajjied & punoj sem stiqap ay} Suowy ‘azis [[eUIS Jo SMuvquy’) B JO L[IYD ay} JO paj}stsuOd SUTBWAI UBIIEISNID BY], “T ‘uel 50 ae 50 20 +0 HI OZ trtitteeteeeee=* SLIgag paynuaprug a iti ‘lee eee eee eee eve eve eee +0 ese nite’ ve Sf Ronee ea ge Ce ate 4S) ae ne aaa CL Sy Vass OOT OOT OOL Us OUT 16 6Z 96 78 gL (ay BOC ROP RUA EEG | EGIL tb 0S vol ape 8b 0S £¢ IZ 0s It 8S OG ne ee 1OS8U Dey ene Pru) £0 Sian ayo tei 9T oid Sigs ZI OL SOO Ged Oued ee ee lo} COU SMArT I] Mpiy: 90 9¢ eee eee eee eee 4 10 +0 eee eee eee FDA ONS ESSE CIAO HBAS DOE enya 7G is) f) De 10 Oia) a eee ery eee eee eee eee 80 eee Ceoer eee ONesNet Che Lakes eee oe oa eT Og CLT Glo [ie ween oh eee eee oes eee eee eee eee eee Teo ee. 10 TL re To} ODI dae TO spi isAney 10 eee eee ene cece OCH Oho eon eee sete eee $0 i a see To) Op Tones Ea) O€ eee On on Oor 91 0S €¢ Ct +Z 6¢ LI 90 oh Ae Aas e2iemen.s Sse tence ieee Te TaN Og] OS) ynpy + eee cee ogi eee eee o aay th ee) .s een wileuts 10 Sea eT ORO STO gE Aeeay Z0 +1 eee eee eee ap 10 eee ‘orsue eee +0 eee a UOROD IO OG DEO OO DL OR Teh OOD E)p lt Md sg | 10 eee sips wae so neh eves apaye skate sue oats 80 cteedeconous ss: pqajdoraaiagdg [PAI] 10 ewe eee ere Cath eee eee eee +0 eee weiae eee aebepelaleleaia avellel sp whessham a laver eee ke epodojdiqg 10 eee eee ere eee eee ey eee Sn) +0 Denes 10 OS CHER SSIES aE GCs CN AOR hon oko RY GI 10 Cervu cor sow as aite tants ayiene S0 aie +0 sists at Sr eligi SEs “a ite 74s. ¢ cose = cea REN OLE UC) CO eee eve tbe ZL eee 60 tL eee eee 80 aye TEE La aiqvigba yy [vto 7, a ae = = = — 50 == = = 50 se SUTRMSY INET paynuepiuy + Cr inert aura 10 Sey ‘Oneite eens’ malta cn este ee FOES 3 I OC CM rey (Ge SAO NONA TT 20 nae A Ae it: or s. a ot ee ae ws Else l crasisie« digiennis shure aio aioe aeraaigy 87 87 9¢ Y4 th 9€ +7 0z +7 91 Ct Po aS Re TAs ie sade ZL, zl 8b tb 9¢ 7s 8b AS 88 9€ fp fae eae Gee Say UNE TA] -10AY ce eee eee 97 eee SE +6 Zs Og 272 Og ChE aS Le (1004006) yjsuaT [BpoWw 9S + 0 0 0 I 0 0 ¢ x4 ZI fl ‘ccc suaumoadg Ajdwigq fo raquinn 7re at Z Z +7 + ra + +Z GG 9 TOT ‘ccc c ccc suautsadg [ny fo raquinN S l + + LI €1 4 Il +I I I NaNV], FLV sjHoT, Dag “AON “JOM Jdeg) Ajnf ounf Aeyy dy “rey ‘qoy ‘uel $}]Ud}U0D YORWOYG JO UONL[NGe {, SLAD]ING SnjyOMas II “ON 219428] 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. ‘peynuaprun se paiajua asoy} Suowe juasaid a1aM syoasur JO spuly [e1aAeg “+ "Jdag “YyeUIO}s auUO UT pUNoF a1aM s}033euUI Inoy ‘Zt A[nf ‘sary propeyo oram BrajdouawiAyy ayy Jo soy “gt aunf ‘1addoysses3 [jews ajsurs e Aq pazuasaidal sem B19}d0yjI0Q aq], "$3]320q Jo sureural au} Ajqeqoid AraA os[e se JOISUT paynuapruN ay} Jo yon ‘UoReI90eUI juanbasqns woif parayns pey jnq ‘ainua paMmoj[eaas aq 0} ysnoua |[ewWS s[enprArpur Aq pazuasaidar a1aMm YoTYM JO OM} JaWIOF ay} ‘2pljawoskiyd pue xpriajelq ‘e2piqeieg : SYDBUIO}S AuBUl UT pUNOF a1aM $a]j}aaq [BII}S9119} JO Sat[rMey Surmojjof ayy, “sz AvP ‘epryey) pue ‘eprydshg ‘epryrydoipAyy : punoz sem Sal[ruey SurMojoF ay} Fo yova Fo aaeUasaidar ajduis YW ‘jue ue Uayea pry UaUTeds auC ‘ysyAeiod & JO Bay ay} JO pajsts “uoo sureuiat podeoap ayy, “TT [dy ‘ajjeeq pruieg e& punoz sem evaazdoajoo [eAre] Jo sureuas ay} Suowy ‘ds ‘xpwmi7 jo uautoads a[SuIs & JO pajSISUOD SUTBWAI UROSN][OU ay, ‘SIIqap ay2 BSuoure punoz sem ueiieurld paysasip Apaed yw ‘T ‘uel Z0 its Law Oe oF 7h aa rh: =0 ave ZI a ee ema an pannus ping) ae soe an Bove wk £5 Soe aes 51-8 Bap 55.0 FO, PRD R War ote 4a me eon ony a eleihe eee eee eee eee eee see 10 a) eee eee elja-lee\'eicg)-eifeelielelfeijatialetrel'siis| («tee leirelate Bpluyorry 86 Ai OOL OOL 66 96 OOL 86 £6 E8 Bib OOn Re ae Pere” SO AAS [IOUT €s beh +9 tL ey) OL OL 91 IS ce oF BE ig Flas ae ey ie eas wesuy peynuspruy) 80 +0 +0 +0 S0 og Z0 (Al) Gil eae see CC B1aj}douawi Ay yNpy L0 eee "AS +I eee oyetes eee eee S0 elvan eee £0 Hoo doo Oo amaptdls ho od buon viadiq ynpy a6 ado ais Soc O00 a 6 amb eves 10 DOK G0 60 ig 0-00 5G.c sizer) (eferraita ‘*eraidopiday [eaiey 0z x sags ae. 80 lz IZ eietis LL 8s siete stoke cI mechs her ii eet eeaatd03 [05 I] py z0 eae a's ake ro see oe aoc eee 80 co3 rAN siailelisiantis) iste SHS G00 +++ p1a3d0a]09 [CAI] ok sive o-Dio ‘0.0 Sd Jens yom 10 Go.a OO neha cols mC RuO 1 4 ORG ONG TrttssssspraydoyzO 3Npy + wae see eae eae se sae AN) eee see eee see ake Co) COU OTT Eee ay rAN) ooo Steve oo 0 ofan noo patie aiter's leno aia aie 9I SisualketekeyeUateh ey sixclickeh onesie B.193d092|g [BAIE'T + eee eee eee 10 eiene sae eee eee eee eee mo JOOOOOn0npt5006 “** ByRUOpO Jo yIdWIAN 90 one hess arate 5 oh cca) aivete eke 0-0 $Z shee v4 asiiswel tes Sa ao a8 ‘++ erajdo1amayd | [BAIE'T ae 50 a-aed aiveee awaits Abed avo ono ances none Lod 10 PPENENS SR DOE I MOIS Sra ists 03 eth ey a se 1G aa sae wae Sra eae 10 ood *hene amo CCR Oo oO Ay oaseo ce CRON Grd Oo Oe OMG “** Bpodeseg + Selle eee eee sae see eee eee 10 sae eee eee © (8) s/s see) ees) els .o\\a) se 6 elec Odea Gs epodojzya ae were TE San 10 FO pais ean ee Sion TOG Soy LD oO EAS Ono “ayy ajqvjaba 4 110 ae ao Tren aires morn 60 ayuike Siete au ao wei waive Su noo ae POM OLOM ELE “** BB[y snojuaweiy = oh oie wwe eee L0 10 2 20:9 CO eee eee eller elles Pe) OC Ch-cet OF TRG aio. Tee a SUIOTU LG — see ZS 8Z 0Z Zs +Z 0% 0Z Og een 0Z Sitelisiiel eile ei @) slis cet GO Gch) yisueT UWINUITUTJAY sose cee 9¢ 9¢ 9¢ 9¢ ZS 09 +9 2G aoe +9 siaiioyjaicettairs||s).% (=) /<\eb eens sees yisueT WINWIxE Ay SHAY Se 8€ FE NG a Tone arbres 0S 8 ae PSU ee en ate aS "1 * (aru) yisueT [epow OT 0 0 £ 0 c 0 I I 0 0 ¢ sees -suautoadg Ajdwiq jo 1aquiny +82 0 IT &¢ Sb 61 Or 9S 18 8 0 Te ‘litt suaupedg [[ng yo raquiny $ Z 4 t LT eT tC IT +I I I NaAV], FLV sje10y, ‘9q ‘AON ‘130 ydag = Atnf = aunf «seq ocidy «ore ‘qaq” ‘ue 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. ‘sulvis pues paurejuco suamiads omy, “p '39Q ‘v4dbo41dg SUM #S[e 9G} Jo SOP ‘pues a[qeiapisuod pajsadur pey suautoads 14 31q_ “Y ade ‘pakbosdg Ajqeqoid alam surewas [ese ayy fo awog “LT Ajn{ “urSt10 ajqejzadaa Jo A[jsour Ajqeqoid sem stigep e4t, 31331] AiaA uayea pey 2auQ ‘sy eunf [NJ AoA aiaM Ysy Aue ‘staqap afqejadaa Ajqeqoid sem yey YA JIS pue pues Pauley -uoo usutdads aU ‘VI4ay INDY SLM BA[e ayy Jo autos A[quqoig ‘A[aATIadSaI ‘SUIIOM 3914} PUB OM} PLY OA} ITY BAIL] JSUT ue pey auo ‘at, A1aA paurezuoo susuoeds Ino “€Z Avy ‘“YORUIO}S aUO UT SazTUL 19}BM Maz & Aq pazuasaidar a1aM BplILoy au, ‘SUIIOM dIzISeIed Jo sjsoy ey} A1aM YsSyY OMT, "[[BUIS AIQA 919M Ud}Ba SUIIOM ay} FO awos a[IyM ‘azjaeq prurtAydeys Be auo ‘poof ayy AiaA paurequoo aary “TT [dy ‘xafiqny Jo sate 1vou Ajqeqoid a1aMm spodoyzyD ayy, “poof FO [[NF Aj arey SeM QUO puke BAIL] JOasUT Ue aUO ‘3]331] ATaA play SuauIToads UaAeg “pT YTV “Paes psear oq 0} paivodde jeyM auo pue s}aSUT [BAIR] Uayey pey OM} ‘poof apy] A1aA paurej}uos suowoads Ary ‘“T “qaq ‘SWAIOM oIIseIed poaure}uOD Ysy 9ATYT “T ‘uel ee ee——ss=S$S09090a oo +0 mie aie Ste Ste ae +0 10 62 arma a SZ SLgaq paynuapruy) +. @ ‘ale a) s Oeacra see ® se eee eee ZA00 eee eee eee Sleuelietaaie) @ue) S29) 6..6.16;6"0 a nee ase, @1R) 8 (ee ce, Bplivoy 9F 9s OF 8& [0 OT cc 90 8P 69 08 08 eS ae Ewe Waser (Or 82 +I 0+ 8E 10 +0 +I +0 +f 9¢ 08 02 ee a ee ee aoes uy peuuepra 10 Sein eee ose Ginx ti a wie eee eee eee WOO Oe Il cm ET ydOpideq [esc +0 eve Bee io reece 3 Cen ZI 80 Z0 §1 Or eee eee Pee OA ee RCC meee) Cecio} B1ajdoajod yNpy 4- siete elaine! iS eee cas oe eee eee on aa +0 BOI 2. GIGS GIG 0-010 YO Fea ehyalasy (OLD) |i | 10 80 we eee ciate Oo o eee eee see cee eee Ol pelle! egw leis sielslecsl=ltriirisj ae B12}d002|q [BAIv'] =- faut. 1: eee One ees eee eve eee 10 eee eee eee eee eek BI BUOPEO) 70) UdwAN AN) +1 sis}.e aioe: O. ceo Cie a) eee eee eee £0 eee 0) aetate) ete) eid) 9: Susie piojdor1smayd | [BAIeT] se +9 eee tel) eee omeix ZO eee Z0 I¢ eee CZ TPIOIOS POO IA OOOO ots Ti aKorskore nfo) + a eB! eae Cie yet ere Percy 10 10 7400 Sis a eee Ruatie OG 2.05 OST OIE OOO TED COAYY pajusuisasul) ip care 09 Z29 66 BS I! 26 02 vers 02 TORE CIO UO oncrcnondspid 49410 J ajqvjaba 4 1210 L 70 = — a aE ae es ae ae aur 02 WT SUES JUL[g paynuepiuy 90 eee 10 10 €¢ 9Z eee 10 eee eee eee eye] GOO OOS OG ODO. torah CSSA CoRN oie 00s [Les 6g aoe 6S 19 99 8s IL 16 02 wa est FACIE eens weno at Yee wee os GIOTENE Or a3 82 (a3 as +Z (a3 4 82 91 Oi. nies Tein hg area se yysuay WNUWTUT A sane 8b 8b tt tb 8b AS zs 8b AS 8b wg hee sects oe UTS] AUB AL -19AY Tb BE'Ob cr Go we ce 9c eb. 8S 77 8E BE ge, Manas fee ee ae oe an) Ue TE PON SS 0 I 0 0 0 0 I 9 a3 /) L sree esses ssuaguoadg Ajdwgq fo JaquinN +87 L 91 6 872 Le 8Z 19 0+ LE S 92 ttt tt suauitadg [nq Fo Jaquinn S fi + + LI EI £2 Il +1 I I NaIAV], LV sjeqoy, ‘9q “AON ‘WO ‘3dag Ainf ounf Ae cady ‘re ‘434 ‘uel $]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. ‘9]399q puted auo Fo paysisuod sureuiai ev1a}doa ‘poof ayaa] AraA uayey pey aaiyy ‘védbosgy Aiqeqoid sem wxZ]e 241 JO awos ‘LI A[n{ ‘e2pruoumauyoy jo Sarsads auios yo uawdads ajZurs v auo pue ‘pnur pue pu ‘Ppooy WIA papuaystp Ajawiasyxa sem uautdads aug pasesur sjurjd ay) wosz [4ydorojyo ay) saseo | pue ‘xpraeqeievog ‘eprurAydejg peZtusoIII1 919M Sapiaaq Fo s Apnojo & UO “J “gq OE: OF SH: T Sulaq Burj}0a][/09 Jo aw oy} JO javd sO11a}sod ayy ur punoj sem Poof 94} FO soy “po peurej}uo0o euo puke ‘sjue ua}ea pey dary) ‘syduAu pryduo $I “G24 “Stay ay} Jo Aue UT JO ‘satias s,y}UOW sty) UL p © “9G EE 0) 40 SC sase -IdAY 97 $9¢ s[P30J, [OD [BAe] ayy ‘L “AON ‘yora suliom o1jIsered aaiyy play BIQAIS UT “ST aunt uNnOF 3q P[Noo Us}e—a poof pue 9ZIS UIIMjJ9q UOTL[ITIOD ON ‘SWIIOM J1VISBIed InoJ pey suo pue 9331] A13dA uazea pey inoq ‘sureid pUvsS aUIOS JayjoUR puk sUIIOM o1NISeIed OM} PI24 2uQ “4 0 OM) puke aft] AraA uaye} pey uauttdads auQ BS OUIOS OM} + SUIOM JI}ISvied paureyUOD aay ‘suautoads [ews Jo s]jeM Apog ury} ay} YySnosyy a[QISIA SEA ‘juasaid sem evioydoysudyy jo Uawtoads a[SuIs YW ‘#prjawosk1yg oNFUey 921 T, *9]3I] AtaA usyey pey uaurroads aug “¢z Av ‘A]JU<9a1 BuIpsez usaq you pey jy 3ey} s}saBens YoTyM | 9UT}Sa}UT 9yy OF 9]91] A1aA uayR} pey uO 3 uaye} pey sary) ‘suauoads usa}INOF Aq Udjee udaq pRYy apIyT AIdA “ABP ‘TL [iidy ‘earey JasuI peaynuaprun ue Ot ‘uel 7s Tos ae : Z0 ra Soe 69 Z0 3 : SOUP aD Obs “"" sriqaq paynuapruy bc . é 56.0 10 aie Be Meo o.5 o.0lho bine Dima are -"eprieoy 66 g& 49 ST Os €6 $0 9E 140) COL ee a A, POAT VIO 09 fe ch IS |! ce 6S 10 9 OL “fis OV oats © eS (parnmoprayy tae +0 nae ZO £0 . on +0 eee eee i ta a eso}dousuALy alnpiy, ZO Aer Salo aias ooo ssn da ib Bot a ee eee Dene Talcigy | EAE a L0 se IZ ahaha 91 O¢ +0 Slee +0 wiede oueie ee ee ee xo3 dooyocy a] Ay Z0 . cae a. ooen iets aise < a (eid Fejelie)aitaielie tele ed Dac B1a}doajog [BAIeT ae cote O00 co AG Chalke ave . 80 - OI sKemonetailatelieitel sleNekenene nei ews B193d092|g [PAIR] c 0G ; ; acre fs S0 aac sieve eles idence nichesacse) Iho Aur UAW e}JEUOpPC IZ . see . oe oe oe eee 90 eee tau i i pene toyCovouroud [BAIeT +0 on 6 . . . . 10 10 . 80 oo /te) le) (s| (eo) ai (ulin jolielie’'s)is) ele) vile ielleirelsietate Bpodojeyy tee a eee . se -- sae ee eae eee Teecean seca ee oe SUTOVAN pejuauidesuy 10 29 fae 68 St 80 t6 oe cee svone sere sretts sss sss daup ajqpjaba 4 10107 +0 ape TS ao 90 £0 ai sc Te : “ss sureuay JUe[g peynuapiuy se nag +- 80 LZ sae o. oa soe . Oa 5 OROEC OSS ceo oOo 0 td Dolo 2s|V Sno}UIWe ITY on 62 66 hy ST 50 £6 ono jou one eee a Bering 9¢ $7 +Z $Z AGS 8Z +7 0Z 0Z eee 8Z Geneve vn tic ewes vvuewesc yysueT UWINUWITUTTAY 96 89 cs 09 +9 9S NG 9S $8 ; Obie tines s~tiqcatele erties yysuaT DEE B IN SOS SU: ch 8E FE'ts 8E 9C Of'ts aS Pil ear ek ‘** (wu) YWsueT [epow 0 0 0 0 0 0 0 S 0z 0 To ovr" ****suampadg Adwgq fo raquiny Or 01 ras ra 9b It 82 eb gS 0 OF ‘" "tts suaumiroadg [[ny jo 1aquiny $ L b v LI eT tc IT tI I I Nadav], FLVd RCS ON aor adsG) Ain, “aun s Avy, “rd Ie | gqaq 0 *uel $}U9}U0D YILUIOJG Jo UONE[NGeT SNINULOD S104] 0 NJ A ON 2981 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. ‘giajdoajog Ajquqoid sem SUTeUIAI JOISUT payMuepruN day} Jo aWIOg “SUIIOM OIVISeIed paure}UOD aUO pue a[}}1[ A19A pauTe}UOD aUGQ “¢ aq «‘SUIvIS pUeS poauTe}UOD adI4T, “L “AON ‘SUIvId pues play 2214} pUe ‘WAIL] JOISUT payuapluN InoJ puke sw10M dI}ISvIed PI24 UO “+ WO ‘SuIeIS pues auo {poof ap}1] AraA auO !suUIOM dI}ISeIed paure}uOI XIG ‘“p ‘Jdag ‘sWIOM oI}ISeIed paute}UOD OM} puke pooy 3f}}1] AIBA PpaurejuOD xIg “ZT A[n{ “poof 2/331] A1aA pauteyuod suQ “ET euUN{ (‘2[qQu} ay} UT pa}SI] JON) ‘spruisap Maj AIOA B P2}OU JIM SUTRUII SNOIDBWIO}JeIP 94} Suowe :SUIOM dIVISeied paurejuOD aAYy {2/3}1] AIQA PauTejuOI BUG “EZ ALI ‘aha Jo] ay} UT puljq SeM YSY 9UQ “AIR JISUT Pay MuspruN OM} !SUIOM JIVISeIed paureyUOD VAY ‘a]}2aq O1}eNbe ue auO fapuy AraA paurejuos suaumdads jYysIq ‘T [Iudy ‘suleid puvs au0 pue SWIOM onIseied paurtejuod suamdeds OMY, “pT Ye ‘pljauue ue jOU Sem A[}UAPIAa YOTYM WAOM B SEM SIIgap ay} SuowWe PUL BAIL] ISU payljuaptuN paurejzuOD aaIyy, “T ‘uel aT = = 50 = a Se ae i eT aan sa SQag paynuapluy + eee ene 6 fmt yeee eee ‘tise eee eee eee eee 10 bys 8 m6 10 Ske, ©) eee BSG @ et Mie) a) 0.18 (8 0) es Bprluyoliy ZS 96 [2 Ze OL 98 00T th 8s 68 cole AL cccsttttssress ss surpuay jrasuy 10107 fb 8S Le EE 10 98 OO he +9 S¢ ee Co oosu pe payiappiay) 4. eee eee ese eee eee eee eee cipte eee eee 10 aoe ee Lo} OUOUT MT tag ND Ge ++ Deke eee Cec, eee eee eee eee +0 eee eee Ct GIG ROL SMG SONI EB ENN G feat al) oho AN) eee ews eee io reta eee et wens ees We eoe eee a eT 3 COD TC age |e ale ay +0 eee eee +0 60 eae eee We. +0 eee eee Z0 SI i Net aeA S Ba TdOS|OGed py 10 Ol eee eee eee eee stig. "e eee eee elecie Duels eee BEES NICO OE SEN (OE) (OK) [BAIeT Z0 +1 eee eee eee OAP eee eee 91 eee eee Bie. iw ee eae TO) OUT mneAS Ea) 60 +1 eee OO eee éMeite eee eee eee Bz eee II Cte ta Ce Se ad ee ee Tet Ch B19}d099] J [BAIvT > ee ° . eee ee eee eee . no ees ee 60 eS era dorouronagy [BAIVT +0 +0 Ce Cat ene rer) x. Oe see rere ci ‘eaule Ooo OL oe 6\ ia Cal Oe 6 6a et Oe pe. 6's) a8 6.5 ie) el te Bpodojzyy fe a 76 TG 06 70 ae 96 — aa ae ae aD 21qvI9vaA ORO] + see ee mene e sie 10 "ee eee “ene ee ‘eu © eee ee eee Se UIMALL Sy juLv|d peYisuspiuyy Al) eee Vr 10 a 6) 6 10 Chet) 10 sais eee eee eee eee Foes eacen dws 0 wie 2S |V snoyuaule|[ Ty a bea: ae if na “ii ate cc bee ees nee bee tence ee tee eee eeeeeeeee oes SUOIBIC] — 02 0z 0Z +7 8z +7 +Z 9T 9T 0z Neo eee ee yisueT wnwiulyy sase St 8b tb 8b tb tt 8b St Ge we ke Bb Es See es ee eS ay MERE -10AY ct 9Z BS BE BE eee woe Zo 7+ eee eae 7c ‘Cb wads ntaer el tie aenetehe of beh ca GUITTEL) yiduaT [Bpow 99 + I 9 0 + ¢ z £2 l I SI ‘ccc c cts suaumtdedg Ajdwgy fo taquinyy 161 L LE IZ 91 +7 $ 81 4 l I Tg “cccc ttt ts *suautadg [ny fo 1aquinny 5 l + + LI cI €Z Il +I I I NaAV], aLvd sje3oy, ‘99q “AON WO 3dag Ajnf aunf Aepwe cidy crew ‘qay ‘uel $JU9}U0D YORWIO}IG JO UL yR[NGe J, snspuoso shyjyoiuiyy . IA ‘ON 2981 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. ‘SNIIAQUNT & SEM SULIOM 3Y} JO BUM “S “VIG y ‘SUIRUIAaI JOISUT PayUepiuN 9y} BSuowe seM BAIR] YW ‘L ‘AON ‘pues pautrju0d sa147T, “ 3dag *pooy ay} FO ey} 0} [enba AyWueNb ur pues sem 919} SYDPUIO}S OM} UT “UISIIO ajquj}adaa jo Ajqeqoid sveM Sstiqap ay} Jo aUI0G “pooz af}}t] AIOA poureju0D aug “ZT Ajn{ *eiaydo -23]0) A[qeqoid seM sutewlas jOISUT payUapruN ay} Jo awWos pue UISsIIO afqev}aBaA Jo A[qeqoid seM Stiqap paymueprun ay} Fo aWOg ‘SUIIOM OVISeIed pautejUOI SYdeUIO}S I9yYI0 OMY, ‘s}SAD JI}ISeIEd A[quqord 919M YOTYM ‘satpog 9}BAO J9Yy}O [BIIAAS OS] 919M YORUIOJS des 94} UT “proutidAD ve A[quqoid ‘ysy [[eWIS aUICS Jo BAO dy} Aq pajyuasaidat a1aM SSsda YSyY yy, “ET oun ‘satdads pare ue 10 ‘xafiqn yz Ajqissod a1aM sprjauue ayy, ‘sWIIOM oNIseied pautejuoo susuoads OMY, “f] YOR iv 5 22 i ie eG. ho cams creme MONAT + eis bee ot eee Vien 60 Pcl Outta oe. babe ou stie Ph chats hss she tap cu slic ssn! of eve eat saceenese 22 BIA) ysty is oe LZ! wes 0S Z1 (Ns 0s 19 0S 4 oem oust] fut Ou 10 . . eae eae see eae eee 80 Byerte OPOL.O iQ see OCC. CC ee OO RCE OR Cat ey CAO. B1a}doajog yNpy Gif . . Bn (e eee UE (oJ) SO 902 see sae see see SEL OTT BITAT 91qv190a 4 1919. 10 vee wise cee eee SS vee mete Tee eee acho opetiou efolredatie)loleltMeiana sUIvWAaY JUL[g paynuapiuy —- sae see see see cee eae -- eee see see sae CPO CWT OSCE OO Ce CRT 2s|V SNO}JUIWIE[I 80 see see . Ls Il S0 9Z toe see see OOH. -wniGogod od SAT Siet* cB eee STUOTBIC] — 9¢ $Z cee 9¢ Zé 8Z +Z 0z $Z wee 8Z dS Odin ayotorao0 oixeite iceiaein=)=fe yjsueT WINUITUTIA] so5e +9 09 SZ tb 9¢ 8S 8+ ZS ZS eee +b eee ee eee Sikehin eit) wlele a yysueT WINUWITXB JA] -19AY sae a eee BE BE 0g 0g coe eee oie stelfem menelraiteWa\(elle) «MeN euelete “+ (mu) sysuay [epol 9T if I 0 I 0 c I I 9 0 ¢ veses ss -suauttoads Aydwy JO 1IqGUINN 6S G 6 if v 6 0¢ 6 £ c 0 0 sores sss ssuguiradg [[Ny FO Jaquinn S L b b LT el tc IT |! I I NaAV EE, ALvVd uO, weg “Aon 20 Jdeg Ajnf sunf AeyY ady ary qey «uel $]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 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 = ea eesleve 4% $f 2c. WaGtGNe——HTOW Ds # CAC sis ois bes Binh a(ieisheke) @ amido ws ereie elie 5 i 2c. *Can be furnished in 9th Annual Report—$1.50. . New York Aquarium Nature Series Sea nore Lite: (Mayer) Clown. ii. i. Sas oie Bodie eels ais adele 1.20 Cultivation of Fishes in Ponds (Townsend), paper...... LA Chameleons of the Sea (Townsend), paper............. 25 Northern Elephant Seal (Townsend), paper............ AS Porpoise in Captivity (Townsend), paper............... .25 Natural History of the Whale Shark (Gudger)......... 25 The Gaff-Topsail Catfish (Gudger), paper.............. iD Illustrated Guide to the Aquarium ( PORBCI boards... .50 Fishes in the Home (Ida M. Mellen).........0.....0008. 50 Aquarium Post Cards: Colored. Im sets...............- .25 For mail orders, add 5c postage, except ‘‘Sea Shore Life.” All publications of the Society for sale at the Zoological Park, 185th St. and Southern Boulevard, New York City. Aquarium publications for sale at the New York Aquarium, Battery Park, New York City. Corrected to July, 1922 OBJECTS OF THE SOCIETY A PUBLIC ZOOLOGICAL PARK A PUBLIC AQUARIUM THE PRESERVATION OF OUR NATIVE ANIMALS THE PROMOTION OF ZOOLOGY LOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY VOLUME II. NUMBER 15 THE FISHES OF SANDY HOOK BAY f By C. M. BREDER, JR. New York Aquarium EUR LIS: Hor. D B:¥ THE 5:.0°C: 1 Ee. g.¥ toe ZOOLOGICAL PARK, NEW: YORE AuGustT 15, 1922 New York Zoological Society General Office: 111 Broadway, New York City Offtrers 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 nf 19243 HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON, ROBERT S. BREWSTER, EDWARD S. HARKNESS, WILLIAM B. OSGOOD FIELD, WILLIAM WOODWARD, EDWIN THORNE, _PERcY A. ROCKEFELLER, JOHN E. BERWIND. Class nf 1924 3 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 ; Glass of 1925 Percy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE, C. LEDYARD BLAIR, ANTHONY R. KUSER, WATSON B. DICKERMAN, MORTIMER L. SCHIFF, FREDERIC C. WALCOTT, BEEKMAN WINTHROP, GEORGE C. CLARK, W. REDMOND CROSS. Sricntific 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 8S. CRANDALL, Curator of Birds; GEORGE S. HUNTINGTON, Prosector; GEORGE A. MACCALLUM, Pathologist; W. REID BLAIR, Veterinarian; ELWIN R. SANBORN, Photographer and Editor. Editorial Conunittee 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 15 THE FISHES OF SANDY HOOK BAY By C. M. BREDER, JR. New York Aquarium Pi oreoe tS HED BY ‘ber OC LE Py Pie sOOLOGICAL PARK, NEW YORK Aucust 15, 1922 AVd MOOH AGNVS HO dVW ‘621 ‘DIA /SOHL Sort Bdes tel : F- JSVUOHSNV So SNI3S sa eannod orca S3aMr4s ey Vie aunpva's a4AOS Sas tee ssn sene recctane eek ¥ P3y % a Vv \g wis HINES § yet ved o) ou @) pai x fa] paofuey yy Sa eed weurley a a, ; Poy Pa wast S pay wi Po anes 3 a aS 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. Publications oA Monograph of the Pheasants (Beebe) UiNinent Pipl S .(cloth) Four vols. $250.00 Four royal quarto volumes, illustrated with reproduc- tions in color from paintings by Thorburn, Lodge, Knight, Fuertes and Jones, and many photogravures and maps. Sold in sets only. Vols. I, II and III ready. Our Vanishing Wild Life (Hornaday)................. Destruction of Our Birds and Mammals (Hornaday).... Notes on Mountain Sheep of North America (Hornaday) UEC) eM CE 712) BOR CE eo a Rg ee A The Origin and Relationship, Large Mammals, North PATROL Gr HULTEL Niet ei Otadescye evan cressie ee ale so Will slate hark! deg The Rocky Mountain Goat (Grant)*......... Spree asa Heads and Horns Brochure (Hornaday), Nos. 1 and 2, eer et bate Me Je eNO ert Ser Bat DASE ra aC To ESE at i er Ct he et Ca A) Zoologica, Vol. I, Nos. 1-20 incl. (In parts)............ Zoologica, Vol. I, Nos. 1-20 incl. (bound)............ moolowica, Vol. My Nos. ‘1; 2,'S-4) 5, COPY o's 5 die c 0's ddielcse's MAO ORICR VOLS EE, NO G.NCOP Wels a scree ers sce sreutore ocdlae miate ars Zoologica, Vol II, No. 7-8-9, 10 and 11, copy............ Zoologica, Vol. III, Nos. 1-2, incl. (In one) .......... Zoologica, Vol. III, Nos. 3-11 incl. (In one)............ MOOOSICariVOls PEER INOM Doe nO eiish ere ie asst PATROL eae Zoopathologica, Vol. I, Nos. 1 to 5, copy................ 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Aquarium publications for sale at the New York Aquarium, Battery Park, New York City. Corrected to July, 1922 10.00 postage 6c. 2.00 postage 2c. sf 3c. postage 6c. 1.00 postage 2c. postage 2c. sf 2c. Ps 2c. 1.20 OBJECTS OF THE SOCIETY A PUBLIC ZOOLOGICAL PARK A PUBLIC AQUARIUM THE PRESERVATION OF OUR NATIVE ANIMALS THE PROMOTION OF ZOOLOGY 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 New York Zoological 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 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, WILLIAM WOODWARD, EDWIN THORNS, PERCY A. ROCKEFELLER, JOHN EK. BERWIND. 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 Glass nf 1925 Percy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE, C. LEDYARD BLAIR, ANTHONY R. KUSER, WATSON B. DICKERMAN, MORTIMER L. SCHIFF, FREDERIC C. WALCOTT, BEEKMAN WINTHROP, GEORGE C. CLARK, W. REDMOND CROSS. 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. REID BLAIR, Veterinarian; ELWIN R. SANBORN, Photographer and Editor. Editorial Commitier HENRY FAIRFIELD OSBORN, Chairman; WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. 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. Publications A Monograph of the Pheasants (Beebe).......... (cloth) Four vols. $250.00 Four royal quarto volumes, illustrated with reproduc- tions in color from paintings by Thorburn, Lodge, Knight, Fuertes and Jones, and many photogravures and maps. Sold in sets only. Vols. I, II and III ready. Our Vanishing Wild Life (Hornaday)................. Destruction of Our Birds and Mammals (Hornaday).... al Notes on Mountain Sheep of North America (Hornaday) _ .40 TES AGS TAT) CONE ETCH See a aa 40 The Origin and Relationship, Large Mammals, North SPLIT OE al B97 Di) Soa ie Ge Pee thesKocky Mountaim. Goat (Grant). ... 0.2... ceeecee ns Heads & Horns Brochure (Hornaday), Nos. 1, 2, copy.. 1.00 Paper Tropical Wild Life (Beebe; Hartley; Howes)........... Zeolorica, Vol. I,.Nos. 1-20 incl, (In parts)......... 0204 3.85 Zoologica, Vol. 1, Nos. 1-20:incl. (bound)....... 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 MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, GEORGE J. GOULD, OGDEN MILLS, LEWIS RUTHERFORD 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. DODGE, C. LEDYARD BLAIR, ANTHONY R. KUSER, WATSON B. DICKERMAN, MORTIMER L. SCHIFF, FREDERIC C. WALCOTT, BEEKMAN WINTHROP, GEORGE C. CLARK, JR., W. REDMOND CROSS, HENRY FAIRFIELD OSBORN, JR. Glass of 1926 HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F. DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON, ROBERT S. BREWSTER, EDWARD S. HARKNESS, WILLIAM B. OSGOOD FIELD, EDWIN THORNE, Percy A. ROCKEFELLER, JOHN E. BERWIND, IRVING K. TAYLOR. Srivntifc 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 8. CRANDALL, Curator of Birds; GEORGE 8. HUNTINGTON, Prosector; GEORGE A. MACCALLUM, Pathologist; W. REID Buarr, D. V. S., Assistant to Director; ELWIN R. SANBORN, Photographer and Editor. Editorial Committee HENRY FAIRFIELD OSBORN, Chairman; WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. Corrected to April, 1923. oe - . i bale : af toto 4 45 A ZT “ON “II ‘104 021601007 IOV INL 2IVY—OET “bry UIOQUBS YY UIATOE Aq ydeisojoyd WOTUVNOV MUOA MAN AH NI Gauvad SHHSIAGLINM ‘98t DTA 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. ‘IIA 91GB 99S OSTY ‘g0fF ‘d 99s ‘SuUBSIO x9S JO 983¥R1S 94} Surousap sszaief xy} 19 SuOTIBURTAXO IO WT, LE 9F PH FG G9 FG GPP IF 99E ZZE 89% ETZLEIT Beene ea S71 eee 7028 - eB . . ST 02 02 22 98 88 G9 The £96 S80 SIZ I6D 991 ZB LOT yo soReccan quowlasoul pue msue% GI L& LZ BZ FS GE GE OL BEE GOS BLZ 9G ZES EI PST F8 aie 2s (2661 ‘Swarm Teayg SI 61 ST &T S& VE GG BL 666 0846 G9 ZSG LIZ E8T 82l 0S LT& Zo6L ‘§ “URE CET < H ° oO be) G6 66 LE & G2 LT SV FO G6S ELS YES EIS I6L PLT TET LY aé 0ZE TZ6T ‘03 “90 = 6ZSFS co 46T LT 8 To Le 6 36 LOE 882 TLZ E96 GES GOS OFT FS War. 65 TZ6L ‘06 “99d = 82ShS Ol 0 t¢ v1 6 T&S Le GE Gea Pie IGT LLT 89T LET OLE Tz OP G&S TZ6T ‘GS “390 LBShS 86 vo IT &I &f OF €& 8h LIS E2S GIS GEL Y8I OFT SIT S9 -a9V 6 GLE TZ6L “8 “3dag gegPC cl i Té VE 16 VE 9E BE 806 FEL E9T GPT 82 FIT 8L OF OP? 026 TZ6T ‘IT “340g GZcrsG él VI ST Go 0G &E TE O09 YS GES LIS? SEI GLI SPT IIT TS dé 696 TZ6T “GZ “snVY Par 06 06 I¢ LI 92 86 16 GS 6IE 664 81% 19% SES LOS YIT 16 dae 686 TZ6T “€ “Sny EzZcre LT 8& 86 GT 61 LT €& 09 OLS GES FOS G8T Y9T 6HT YET 99 +d¢-P-8V 6 L8G TZ6T ‘§ “SNY Z3ZGhS SI G6 9T VG OF BT LI OT 60E L8G TLS LHS LIZ GET SBT 29 OP GG& T26L ‘86 Ane Tere e 8ST &T 96 8T 06 Gh GG 92 SHS SEZ 90% B8T 89T GIT LY dé 896 TZ6T ‘8a Ane OzZgPre 8 OT OT €& 96 ET 9 8h 8S 8ES 82S GOS BLT 99T OZT SL a? 96@ -T26T “8a Aine 6I1¢rE ST 81 9L 8T 86 VE 8h 8 962 8LZ 29% FHS OTIS SBT PET TS a? TTS T26T “16 AINE gicre 6L 16 06 8I 7S SE GS 18 16% OLS 0GZ SES 80Z SLT OST 69 OP? OTs T26r ‘Le Ane LIGhS 6&6 610 &@ GI 8I Te & GG 692 OFZ LIZ 86T O8T 6ST 9ET T8 asvé G86 TZ6T ‘Et Ane 91GPG 6L STI Go 8I GI VE GE TL 996 TFS GIS 10% G8T SST OZ 6P “dP-3V 6 GLzG TZ6T ‘Go eunf ¢TcPE L OT VI Go 62 6T LT 6H 02 S6T GLI LST 82t GOT 26 &F “dev é OTG T26T ‘8G ABI FIShS v ¥G LT VT LT 93 LS] GL PLS OSS SES 61S 20S YLT EFT OL ae 8LS TZ6T ‘83 “Ady gsIgp¢ 6&6 OT 8T 8T 0& SZ 06 696 LPS 666 IIé T8T 9ST 99 -d?rvsé 986 T26T ‘86 “IPI ZIShs 6&6 LG T& 92 OF TS 89 O86 €96 G2e 96T OST GET LO “-d8V6 608 Ie6l “P “BIN TISrS 0G ST G6 GG IZ €E 68 PIE 666 LL] GSS PES 10S SIL paj}BsooStAo{ VEE TZ6T ‘86 “G2d OLShS &— 9T L@ 0G OT FT OG 906 O6T E9T SPT LET EIT §9 -d8-2-TV 6 616 TZ6L “8G “URE 60S €T 8T oé ST 02 FS = LY VLE 996 VES GIS GET SLT 80T AP-s- TV 6 L8G OZ6T “0S “9d g0srE 6 €6 & LZ GZ OF 99 9S¢ S&S OTS E8T T9T TET G9 ae G9G OZ6T ‘IT ‘qd LOSHE 6 81 GI 0& 72 OT 8L 986 816 906 YLT SGT 9ET 8g ae SVG OZ6T ‘9% *20 90S7¢ OT 46 €6 Gé GF ET ar LTG O6T LOT GPT OTT LA eC a? L&S OZ6T ‘ST “SnV GOSS Oe eat et al OP 1 alee Ot Sle Sl pa ol Ne de a PT ~SUBBIC) FO a Tse Gg teq UN : 3381S “UU UT jo wunesnyy Pus xeS yysue'T al a ER! ‘OTCVIIBAB 919M YSYIIYM PlO-1eah-9uUO OU se ‘SoTBOS JO SJUIWIIINSVIW I9}OWIVIP UO pased SI ‘1I9gAQMOY ‘TIBIA ISI 9 JO YISud,T aSeaavav 89} ‘SJuUaMIIINSBveU Tene Aq peuluitojJop SB YUSFIYM UOINE 94e'T 8gzZ JO 1v9A YORI IOJ SJUSWIAIOUI [BNUUY ZSeIDAV PUR SYYSuI,T sSvVaaae 94} UVATS 9128 SUUINTOD 9} JO UL0}}JO 9Y} 1B SaSsVIIAV P9JB[NITVO 9Y} MOTI ‘IsIT JO AIBVAaA Yous AOJ San[VA paie[No[Vd asey} JO saSeaisav 94} pue ‘(‘0}9 “y “%y) SaTVOS WOAJ POUIUIIZ}Jap SB ‘WW UT S}JUSsUIeIOUIT YWMOIS Tenuue 9} ‘(910 “sy “yz MZ) SaTeOS 9Y} WoOTT PI}B[NOTVO SV IIIT JO LOJUIM YORVS JO pus 9} }B ‘WIU UT SYYSuUsT 9Y} ‘suvSIO x9S JO UOTIpuod pu xeS 9} “(TBpNed 0} yNOUS PsINSvsUT) YIP JO 9UIT} Je (MT) ‘WU UT YSueT, 9y} ‘YIVEepP JO 9}ep 9y ‘a9quInU wnoesnyT IW JO ‘Q 94} SATB YOIe RH Aeq-ul-jnq Woilj sS38S9 woiajy ‘gTgT ‘Aaenuer payoiey YSsyouuAA wWntaenby MIOX MON 23 AOJ SBSuUIMOYS—]T o[qeRy 388 (i it} Nid I \\ Ae AN} WSN aM i} ~ = 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 ‘ SAVE ces wh nae vm 1 : I) eae 3 VA ud) ‘ A) i 4 ae j } t i= et, > 7 Gay = SEL fy) 77. Se) 4 2 y #4 A ie JS Pha 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 Ball 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 } | y Ws) i I Ai : ; He i Uh ( |! ie i HH i) ah | ¥ << ANG ACA —— oe rUbhay NU \ ano Tg ZEA 2 eI ovon WD - 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 "UJMOIS LZ6T Z ‘OZ6L ‘T4dy 10 you OUTS » 9 Virdee 0°6 9°9T €8T TZ6T “GZ eunf TZET ‘GZ oune g Z0°L+ Z0'L 9ST 12% TZ6T ‘9g ABW g v:0— Lg eI 661 TZ6L ‘9% YoreyA Z Po ya eu GZL *GLT T26l Ogos [[s-uedQ STGP v 20+ VL as 681 TZoLe ch SICh = teehee Ie g og+ ragh 9T 12% TZ6r ‘64 oung Zz Faia 22'S g 0Zz TZ6T “9¢ [lady z Pot 0°6 GOT ESI T26T ‘G6 “2A T 498+ ¥9°8 9T G8T OZ6T “OE PQ Yoodd-2[qnoq gTGPgG g BOA 6% og GLT Oc6T ET seod Occr Tf - “eed g SiO ee Lg LOT OZ6T ‘9% “AON v aoa «8'S gg 102 0Z6T ‘9 "PO ¥YRG-peyootH = LOSHS peseg o4eq snjnuuy 1eaX 446 IVa Ysly a[BIG jo ysl Taquinny SI ase suIpod 48se'T apis IO U48 YMOI) JO yyeod [BPAOULO JO oule Ny uUnesnjy “AY YORY -otg vulg -jNO 7xVG Ul YWMoOIH = 44g 40 Jo 03" Jo oyeq "WFO'N yum uo peppy 0} peppy = [BulsaeyW YL JO PU So[vog Jo aseyUodJeg esvUusd.Ieg JOY Sue] 4¥ Salvag requin NV V SA/64 aseloAyY JO Y suey IO LA/8A 6A 40 QA ISVIOAVY 8A 40 LA ‘paseq o1@ SadvioAe oy} YoIyM UOodnN Salvos Fo Jequinu oy} pue o}¥p Sulpadeid 944 VdUIS poof; esejuooted YIMOIS [VULSIVU SI} Jo Jared 9Yy ‘Sn[NUUe 4S] 94} UL pepNjoUl JeyeWIeIpP AOT104SOd-o10},UB oY} UL (YAMOIS [eUIs1eW) SN[NUUe 4JSOUIEyNO dy} SPIS}NO YysUeT 9[VOS ay} JO oseyUsdIed oY} ‘uINIIeNbYy YOK MON oY} Ul pocvod YsyY -o}IYM FO S[VNPLAIPUL JO SoTVOS peylUseUL JO “WW UL YASUI [¥}O} odeIBAV OY} SoJep JUeTOYIp Je SUIMOYS—T]] qe I, 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 “AIVNUBL OJ ISNSny ‘Ivak YG p9e[dWoOdUI IOJ VBSvIBAWY-—oO ‘SUISOq JUDUIAIOUI IvaA YUIN—q ‘Hole 0} Jsnsny ‘1v9aA 1g pojJo[durooUl IOjJ 9SvI9AyW—e 2G°9 9°), 68 = oseIOAV T Ss 68 86 Ga6I ‘ES Arenue f T uy) G9 6S G26 ‘S Arenue ¢ I vs a8 SST TZ6T ‘03 equiedeqg 1! TL 3°9 &9 TZ6T ‘0% Aaquiesoqg T I 67 OST TS6I “SZ 19Q0}9O T GLE SOL og TZ6L ‘8T toquiezdeg T Lg SL S8r TZ6T ‘TT Jequieydag {! ae Lg TL TZ6I “GZ qysnsny iE ws 89 SL TE6T ‘8 ysnsny I 1g) SOL VéL TS6T ‘E ysnsny T eV qh 6g TZ6L ‘8% Ane T 0% VL ght TZ6T “82 Ang 1 0's S7 Cy TZ6L ‘8z Aqne I og gis 6g TZ61 ‘LS Ayng 4 Sy) SL 68 TZ6L ‘LZ Ane i! VL 91 80 TZ6L ‘ST Aue T 0°6 G9 8°6 TZ6T ‘SZ oun lt I VE 0g T8 TZ6L ‘8% Av E q9o'T 66 el TZ6L ‘82 [lady Bgrg G 68 es) TZ6I ‘83 Yore yy G g"8 6°01 TS6T ‘V7 yore] 3 v9 og T3261 “8g Adcenagey 3 29 8°8 TZ6T ‘8g Aacenuer G LY VL OZ6T “OE Tequiedeq G md 86 OZ6L ‘LT aAequieosq G TP a8 OZ6T ‘9% 40qG0390 G LY GTI OZ6T ‘ST ysnsny posed 21V AT9ATIOIAS9I so3e U9I010g weex Wwe 10 YI) FO Iv92X UL JO ivaxX yg jo y1ee9q jo 91eq Yoru MA uo yjsueyT 8,2OS UL Jus YYSueT e[eoOg UI JUsU YYSueT Y[VOg UI USUI solvog jo 19qWnN -910UT IV9K YG IO YI -910UT IV9K YI jo -910U[T AIVOK YY Jo po yeTduoouy jo 938 }uU90 89SebUIIINg LPA/SA e3eUI0IIg 9A/)JA -I9q 8A/6A 10 LA/8A ‘sosejuo010d Jo soles yore I0J VSvVIVAe 94} PUB ATaATJOOdSeI IVOA YS Pue YZ ‘419 94} JO YSsus] V9TVOS [B}0} 9Y} UT IvZA YG PUB u78 ‘UI, 94} JO JUSTIZIOUT OTVOS VY} Jo VSejUV0Ied 94} T PIG" JO YSY WnaeNnby YIOKX MON [[B OJ SuUIMOYS—T]IT I1qeL 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 (CMacGatlim) 45. 3s a ee 25 Camallanus and other Nematodes (MacCallum)................ B45) Studies in Helminthology (MacCallwm).........0......002...220.--...- 50 Postage, Zoopathologica, 5 cents Cloth All publications of the Society for sale at the Zoological Park, 185th Street and Southern Boulevard, New York City. ZOOLOGICA SCIENTIFIC CONTRIBUTIONS OF THE NEW YORK ZOOLOGICAL SOCIETY 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 New York Zoological 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 MADISON GRANT, WILLIAM WHITE NILES, FRANK K. STURGIS, OGDEN. Mitts, Lewis RUTHERFURD Morris, ARCHER M. HUNTINGTON, GEORGE D. Pratt, T. COLEMAN DUPONT, HENRY D. WHITON, EDWARD HATCH, JR., CORNELIUS R. AGNEW, HARRISON WILLIAMS Glass of 1925 Percy R. PYNE, GEORGE BIRD GRINNELL, CLEVELAND H. DODGE; CG. LEDYARD BLAIR, ANTHONY R. KuseR, MORTIMER L. ScHIFF, FREDERIC C. WALCOTT, BEEKMAN WINTHROP, GEORGE C. CLARK, JR., W. REDMOND CROSS, HENRY FAIRFIELD OSBORN, JR., ARTHUR A. FOWLER Class of 1926 HENRY FAIRFIELD OSBORN, LISPENARD STEWART, CHARLES F, DIETERICH, GEORGE F. BAKER, WM. PIERSON HAMILTON, RoBERT S. BREWSTER, EDWARD S. HARKNESS, WILLIAM B. OSGOOD FIELD, EDWIN THORNE, Percy A. ROCKEFELLER, JOHN E. BERWIND, IRVING K. TAYLOR. Scientific Stall 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. MAcCALuuM, Pathologist; W. Rerp Buair, D. V.8., Asszstant to Director; ELWIN R. SANBORN, Photographer and Editor. Editorial Committee HENRY FAIRFIELD OSBORN, Chairman; WILLIAM T. HORNADAY, CHARLES H. TOWNSEND. Corrected to August, 1923 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.