A REVISION OF THE POISON-ARROW FROGS OF THE GENUS PHYLLOBATES BIBRON IN SAGRA (FAMILY DENDROBATIDAE)

By PHILIP A. SILVERSTOi'E

NATURAL HISTORY M US EU ~ l OF LOS ANGELES COUNTY SCIEl'CE BULLETI!" ~ 7 • :-,:ovEMBER 25. 197b A REVISIO:\, OF THE POISO:\'-ARRO\\' FROGS OF THE GE:\'US PHYUOBATES BIBRO:-': IS SAGRA ( FA ~ lI lY DE:-':DROBATIDAE)

A REVISION OF THE POISON-ARROW FROGS OF THE GENUS PHYLLOBATES BIBRON IN SAGRA (FAMILY DENDROBATIDAE)

By PHILIP A. SILVERSTONE

:-

AC,,"OWLEDGMESTS

~h T U I " LS ""D ~l ET HOOS ~ u-e um abbrevialion$ 3

PNlUJJA< 1a BIB_ ,.,S..u...1841 J Ddinilion J D i a ~ nos i s J Descripelon J Conten t , Tad poles , Specie, group' 7 Life history 7 Food , Summary of distribution 9 Key 10 the genera of Dendrobatidac 9 Key 10 the species of Ph.l'lIoh/ll'-f 9

8 K."OW ~ (ilt 0\Jl' 20 Phylloha tt!_, ,pec ies 20 PhylloM tt!s aurorut!nio 21 Phyllooott!Sbiro /or 2J Phylloha lt!S lugubris 24 Phy/foha tt!s ,';IlOlll.S zs

F EMoifAUS (iltOliP 27 PhylJoM u s a'llltwly'; 27 Ph)·fJobou s bol'lt!ngt!ri •• aa Phy/Jobatu t!spino so; 29 Phy/Jobatt!Sjt!mo ralis 50 Phyllobau s tricohv J2 Ph,.llobau s zapar o JJ

P O f!S (; 10l/P' as Phyllobatt!s bolMu lllU Phyllooolt!S ingt!r; " Phyllohult' J pan'ulus "J6 Phyllobalt!S peters! J7 Phyllohules plctu s aa Phvltobates pulrhripeclu .f 4J PhyIJob/lles ,marulld;nus 44

T~ lVm A I /JS (; IO\; P Phyllobalt!s bault! ri " Ph.l'lIobalt!s triviuOl,.s " "

U TElt U l,IRE CnED 50 FRONTI,PII:n: II. Left side. lOp10 bonom: Pilyll"bFrog IP. .m : Phanla,mal PoiSl>n·arrow P,,'g {P , /rico;"rl. Rio :l4ina•. 20 km W Sama habel. Ecuador, Brillianl-lhigbed Poi,on·",row Fmg fP .f~m(lra l i'l. SaUlTonue. French Guiana; Beauliful.brea'led Poison·armw Frog IP pulchri~.., ,, .,,/ . hololype. Scrra do J\avio . Brasil. 1'01 to ",.Ie. Painling by J. K lkanishi. staff an i' t. LACM A REVISlOS OF THE POISOS .ARROW FROGS OF THE GE:-iUS PHYUOBATES BIBRON IX SAGRA C FA~ lI L Y DESDROBATlDAE)'

By PHILIP A. SILVERSTOl"Ef

A. ST__~ : TIle 20 dumw. ~~illl. 1nO>lI)' focnl:~ellin, 'J'C"io of lhe anuran l:"nus P• .•1Iobu,... oecur from Casu Rica 108...-.;1; half of the sP'""ie~ inhabilthe A""""", d...u.a"e. TIle di. linction bel.. eu Colo.,..'.... (dor",m d ull~' rolorcd and .....in lO~in. prc_umably ab.cnl' and PII.d'obu,<'J(dorsum bri,h ll) rolorcd and ..lein lO~irl, I'f",,,mabl}' P"'''''''' .. maintained. Fing.... Ic n ~'1h and finj:C1" di'k size ~ 1I>Cd hc~ in 10di...in,"i"" ~lIJrobd. ""d fin' ....di.h I'dal;"ely larj:C) from PII) /Iob<1 l<" Cfir>! finl!tt equal lOor IonI'C'" lhan >«und . and finrcr 4ib.I,... )of 1<'Clh. allhough lhe IooCh charac1C1" i, ill u",f"l. U,in, finger knglh and finger di..... 'ize a' a dcfinilion. fl\'e , pecic' (bl1l"': (1) bi,'mo, group (Ph.•H,,,,,, ,... , pecic, It<> be oJe ".;ribed b) C. W. My'cn andJ, Dalyl. P. " w",'um i". P, biml,,,. P. /wJ(wbri" P. vi"",w,):(2)!, ,.,,,,,,li. gfOll p (/'. "",hm•.•'i. P. " "wlm g..,i. P. r,pillooI",, <'5 Il<'le,,1. P. f'"lch,i{'<'clU.'. P.."""'''Xdi"".,. and 1'. Z<'p""'. The life hi,tory of I'I" -/I,, l>ul ~ ,' (in",fa' '"' il ;, known) "''''' mhlr, Ihal of iI' relatiws Dend","'" " , and CoIOlI<'ln"s . Phyll" l>aln u,ually ""ip<"il' Cal lea'l in """ 'pecic•• and I'l'ohahly in all ,pecic' . WilhoUi amplc:1lud on lea"e,. Aflel'the ladp<' le, halch, a parenl , ,,,ric, IMm 'lIl il. had and rel, a"', lhem in waleI'. whc~ Ihc ladpole, compkle thei, ,je"elnpment. In al lea...... e , pecic" ,ilher the mak 1"''''11'1 01'lhe km alc parenl may carry lhe ladpole, . Tl>cmain food of Ph l/obm... i' 111>1, . Ame,indians e ~ ltac t blo....gun poi"'" from al ka" 1"'0 'P'""ic. (P. "" "'I",n;,, and P. mcolo,).

I S 1 RO[)l",'C1IOS Th u, Ih i ~ revi,ion of Ph\'l/ol>a,..s include. those specie - h avi n ~ a oombil'liltion of three chariICten.: ( I) A, p""viou,l y di-.:u....-.ed (Silver'>lone 1975). lkrldru· ,mall fln~e r dish: (2) the fiN fin~"""C(juar to or lon~e r batjd 1Cnera are difficult to define. The ~ enu , Ph).·//t>· than the ...,;ond; (3) the dor, um partly or entirely previou slj- ..."" d ia~ n"""d by tbe prese nce of "'u... brighlly co lored in life. In my op inion. the ~'(: n u~ Ihu\ bright oo l"1>;ll colorat ion and the pre-e nce of maxillary de fined is mere natural than il ·...·a' under the pre v iuu ~ and prema xillary teet h (Savage 1%8). Because the definition. However . delineat ion "f convi ncingly prese nce of teeth "aries with ill one 'pecic, (pi ,'/iu) munu phy lc lic genera of dc ndrob atid s will req uire a, well a' betw een 'evera l oth erwi se >'cry , imilar con, idera hly more dala. 'I',:e ie, (h,,f ;I·i

1'111,,' 1<,,), I have redefined the genu' nil the h'lSi. of A (" Kro;OWI ,EI)(; MENTS finger dis k size and the relative length of the first and lthank the followi ng for C ~" lIl d fingers. These finger characte rs val')' ,""K"' · J .W . Wright , R,L. Bel )' . F. Truxalutal HiSlCum Ph...//oI> do~l y retated to CoIt1'l;a 9(0)1 . 2 BULLETINOF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY No. 17

J.P. Bogart (lNHC);C. F. Walker. A.G . Kluge from lhe :-.ralional Institules of Heallh to the Universily CUMM Z); R.G.Z..-eitel. C.W. M yers ( A.M:" H ); J.E. of Southem California ( 1969) . and try lhe Los An~le , BOblkc. E. V. Mal",.\( (AS SP); A.E. Leviton (CAS); CoulMy ~ u se: um of r\at u...1Hislory Foundation ( 191 1). l .A. PC'lers, W,R. Heyer, G. R. Zug . R.G. Tuck. R. Crom bie . S. CamJXIen-Ma in (USNM); w.e. Shcr­ M II Tf ~l lI t .s liND Mf.THOUS b....lke. R.W. McDiarmid. Univcr.;;I)' of South flori da: All nlCil-surement, are 10 the nearest one·half mil­ E.E . W ill i a m ~ . B. Shreve ( MeL); \\' .E. DI.IC'Jl man . limelCr. S.R. Echurds. 1.T. Collins (KU); C.l . McCoy. Jr.• TlOdpole lIlagesarc those of Gosncr (1960). S . Richmond (eM); J.D. l ynch. University of Nc· O' ioolo!!- ical , pecimen, were cleared wilh glycerin bravka; A,C. Thoresen, Andrews University; D, Travis. and lwo percenl pola"ium h yd ro~ i de and were stained Manhallan Beach: D.B. Wake. R.C. Stebbins (MVZ): wilh alizarin red S. D.B . Ekkens. Lorna Linda UnivcNily: A . SIam:II . p."nermo and subpalleTn\ in the aocounb of P. piC/in Calirorni.. Sta te Vnwen;!y at r.; or1 h r id ~ e : M. Tye . and P. " i' ;IIOI/(.. ( intended 10 represent lau H , M arx ( FMN H): J.A . Rivero, Un iversidad de Puerto In the species ",:counts. under ine headinj! "stomach Rico, Mayaguez: InslilulO Ev.mdro Chagas. Betem; conlenls." the museum number of a frog is ciled first. Induslr;a e Comercio de Minerios. Serra do S avio: follo'A'ed by lhe names (afld numbers of indj,.·iduals) R. Pradinaud and Dr. Rivieeez, Ca)'l"Tlne: TeKas InSll'\l­ of the il1\1n1ehr.lle tu a found in its 'I<.>mach . menls Co.: K. Adler. University of Noire D-ame; O.R . In the synon)·mies. a period separates the name of da Cunha (MPEG); W.C.A. Bokermann. Fundal;an a taxon from the names of all authors CXCC pl the Parque Zonl,igiCd de k>s .....ndes. ~ot ;i; It is difflCUl1 10 define species of Phylloball"J be­ B. Cboebe n , Centre Nalional de la Recherche Scien­ cause of intra.specific varialion . interspecific resem ­ nfiquc. Par is; J. Lescure, J. Guire tMHNP); M.S blance. and a paucity of kn"wn cases of eympatry. To Hoo j!moe d (RMNH); W.N . Pold er . Alkma ar; B. define specics. I used pre'encc or absence of the fol­ Hudendid (NMG); S,H. W"-s..cn. GOteborlls Etn(}­ lo.. in!!- char.JClers; toe .. ebbin g. Icclh. a calf·"P'>(. a gra fi'ka Mu-eet; A.G .C . Grandison. E.S. Arnold median dof'il l ...ripe. p;lirell ...npe s, and dorsa l or ( B ~ ) ; I . Ei.e h (NHMW); G.-F. de Wille IMRHN): ventral marbling: I aho used position and Icn!!-Ih of w. Hellmi<; h (ZSBS); and I .H. Haffer. Te heran. J.H. pairell ' tripes. snout -vent length. , kin texture . and Haffer gave advice on the mute of access to the Alto color. When so me of these cru.racters wcre not shared del Buey. J. Le-cure pruvided a Ii....of Ioca li l~ for by allop"Ii,·;UI'''.'. P. Br"me deci, ion, on the known ran g.c "f variahili ty of the drew the tadpoles. J.T. Ktraeako drew thc master out­ character in both popt.a lal ions. and on thc pre-e cce line for the color paac rn diaj!r"ms; I drew the patterns. or "bsenc e uf morphol.>t: ically intermediate popula­ "fhc, '>Uperb fmnlis pi«es arc the ...orl of J. "' akanishi. lions . I h"ve been conservaove in dcscribt n!!- new " aff anisl of lhe SalUral History Muse:um of Los specie,. Anj!eles Co" nty. Paintinj!' for the fmotispic<:es were The specie- of Phyll"ho l<' . are "rranj!ed herc in based on color slides by J.A. Pelcrs. R.W. McD iarmid. four prov isional spcc ies j!m ups. T he' e groups arc J.D. Lynch . D.B . W..ke , A.C . Thore-en. and me. intended 10 represent relationships, bul can do so onl)· SlUdie- were p;>niall)' suppon ed by a NSF Summer imperfectly h1x..u-e of the incomplelenc" of ",'ailable hllowship for GrlOdUalC Teachin!! A",si" an" (1966) . data, Tn arra n~ the , pecic- in groupe. I used presence by a NDEA Tille IV Felluw,h ip ( 19M· I%9j, try Bio· or "b<.ence of toe webbing.. position of p~ i n: d '1ripe~ . medical Sciences Support Grant FR·070 12·02 # 570J snout-ver n length. and re-cmblancc of cnlor p attern. /976 FROGS OF THE GENUS PHYLLO BATES J

M u>cum a~v i alion s 19Z6. PIt,.lIob<;tr. (pan). Soble : 9 Th e foll owing ab breviations ind icat e tbe loc ation 1 9 ~6 lHordrobertn (part). Sobk: 9. 19S7, Phyllob.rlr . (pan). DlIJIn; 77-7M of preserv ed specimens: 1968 Phyllohalr. Sevege: 747_7411. AMNH Amcrican !'>lus<:um of Natur al Hi, tury, Ncw Ym' A ~ S P AciOkmyot :-;;Ilura] Scim;(,., Phil.xlphia pr-ema ~illM)' BM Brili'" Mu-,tory). Loodon Defi nition. - MuiUary and teet h pr-es· CAS Cali fornia Aude my Qf Sc ien ~•• San Fu.n. e nt or abse nt; dish o f seco nd. third . and fou rth finge~ cisco relatively small (Silverstone 1975:fig. I); first finger CAS·SU Slanfor. (~ Rican EJ.pcd i l ~ ) colored; palatine bones absent; omostcrnum pre....m; P..l~H Field Museum of :-;atural Hi>tory. Chicallo vertebrae usually' not fused; tad poles (Figs , 10-1 2) GOV G . O n.~ s V. collection 31 USSM with oral dis k laterally inde nted . number of demiclcs U.s [n,tiluto de UI Salle. Bogola not reduced. and anus dextral. lAP J.A. !>clen collection at US:>i M DiQgnm is.-Phyllooof.... d iffers fro m Dendrob«tt . J ~IS 1.M. SanS" priute O>leOlogical coIl«tion aI CRE in havin, ue fiN finger eq ua l to or longe r than the IPB JP. Bog;onooll«tion at Th HC seco nd and in hav ing relatively ,mall finger d ish , In K!'>l KrakO'" Museum . Ja ~ e ll o n i a n Universit y, Ph.l'lIobC um of to. Angc l... usu a lly are al Ieas l 1"'lo;e Inc width o f lhe Finge rs. County. to. Angel.... Ph y /l"hrtfl'$ diffe rs from C"I(>$I('l h ui in having the skin LG J. i.e, cure private colleclion. Pari, of the dorsum partly or enti rely brightly colored in life , acz Museum of Conlparativ.. 7..oology. Harvard Univen il). Cambridge anti Ipre cuma bly) in havin g ,Lin to xins. MBUCV Musco de Biolollia. Univenidad Central de DtJeripti" n . -The snour.vem length o f adu lts i. VenelllC'la.Car""'b IS 10 50 mm (Table I); female, avera ge lon ger than MH!'OP Mu..cum :"O alioMl d'Hi""ire :-;aluf!'lle. Pari, mal es . The skin is smooth or gra nular Ma~iIIa ry and Mu....u f'araense Emilio (;oeldi , Belem M"'G prema xilla ry teeth a re present or absent; vomerine MRHN [n"itut Royal des Sdcnce. Natur ell ... de BelgiQIIC' . Bru••els teeth are absent. Adu ll ma les have voc a l slits and MSH M.S, Hoogmoed roIl«lion al RMS H u,uaU)' have (at lea.' t in prese rvariv...) ;a pair o f lonil i· "tVZ Museurn of Vmrbratr L.>loJy. Univcrnly of tudinal e xterna l fo ld, o n the throat ; fe males lacL voca l California, Bend ..) slil_ and e xte rna l folds on the throat. The lip of the ~ H MW Naturhi'lnrisehe . Mu><:ul11. Vicnna NMG Naturhisloriska Museel, GOleborll snout is trunca te 10 slightly rounded in dorsal aspec t. RMNH RijL;mu,,"um van !"aluurlijke Hiuorie . u-idcn The imercrbital distance is wider tha n tnc upper e yelid . TNHC l ln;,...... ily of Tu n . Au"-in The diameter o r the e~e is greater than the di'>lance UMMZ linivenily of Michigan. Mu,,"um of Z"O~ I . fro m the eye to rbe nos lril. The pocerodorsal por1ion Ann Arbor of the tympanum is conc ealed. Each digi lal disk beu.. UPR-RP Unis'ersily of Pucn" R i~o, Rio Piedras USNM National :\lu....um of Natural Hi. tory. Wa,h­ on its dorsal surface a pair of raised " scutes" separaled .ngton b)' a median groo ve: the seco nd. thinf , and fourth finge r wCAB W.C.A . Bokermann private colleetion . Sio d isks an: small rela nve to lho<.eofD('nJ" >bultJ lSih·er· ". slol1(' 1975 :fig . I). The dis ks are not sexuall)' dimorphic , wcs W...e. Sllrrbroole ro lleclioo al US ~ M WED W.E. D~ ll m ~ n collectio" at KU T he first finger is elJual to or longer than rfe second. ZSBS ltlo l o~isc he Sammlu nll de, Bay'et ischen The fing et'!> lack fringes. and well-bing. A subartkular Staale>-. "'unich tubercle occ urs on the vemral surface of eac h di gita l joint. An oblong inner :.rnd roo nd outer tube rcle occur on both the metacarpu s and mc t:.rtar , us; .....meli me s a PhylloOOIl" Bibron in Sagr a 11141 median tubercle occurs on rbe ructamrsus. A ridgelikc Poison-arrow Frogs fold e xte nds alo ng the d isla l po rtion of the larsu s; the Frontispiece s 1·1[ and Figu res, [- 12 pro ximal end of lhe fold cu rves . loOtnelimes bend ing at rlilht angle, 10 ue re'l o f !he fold . A tarsal tubercle 1M !. Plt.I·lIo1>. In life , the iris is blac l Of brt"" n (rarely bronle) 19W Phl# oI>

TABH I ~ Snout-vern length (mm) of 732 adult specimen> of 20 Sf"'cie, of I'hyllohmcs. including , ample size (n). range. mean and sland.rd ~ error (;:±SEl. .00 ,t""dard devialion (SO) ~

All S[" cimen, Males Females ~

Sp..", ic, , :SE Range i :SE SO Range , :SE SO ~ " Range SO " " ~ ,m ,h"Il\'i 16.5-21.0 19.2:0.20 1.16 u 16.5-19.0 18, I : 0. 18 0,65 1':/,0- 21.0 20.0 :0.17 0.75 ' pecies" " .~ 6 0_ 4 2 . 5 40.0 :0,75 2.411 , 36.0-40.5 38.2: 0, 75 1,84 "S 42.0 - 42.5 42.2 ± 0.12 0.27 aurmac"i" " 23.5_34.0 27.7 ±O ,54 2.97 23 23.5-3 1.5 26.9 : 052 2.48 1 26.5 _34.0 30.4 ± 1.15 ~ b<".lcr; '"ro 32,5_42.0 37.3:0,78 2.47 , 35.0 35.0 9 32 . ~ -42.0 17.~ ±O , 83 ''''2.49 hi"olor .. 32.5- 42.0 38.2±0.35 2.32 20 32.5-39.0 .>6.3 ± 0, 38 1.70 3().5_42.0 39.8 :0.28 1.39 ~ f>oli,·j'IIjus , 23,0-2().0 24.8 ±091 1.58 , 23.0 23.0 "2 25.5_26 ,0 25.8 brls b 1 9 . 0~ 23. 5 20,5 ±0,53 1,75 , 1 9 , 0 ~ 1 ':/ ,5 19.2 : 0.1 1 026 S 205-23.5 22.2 ±0.51 1 15 p<1n'ulu. "0" 17.5_24,0 20,5 :0.14 18 17 .5 ~ 2 2.5 19,0 ±0. 15 094 12 19.0_24.0 21.2 ± 0.12 L'" ~ fN/cn, 38 21.0- 3 1.0 26.6 : 0.39 2'.4''' 1 21.0-2 8.0 24.8±O,49 200 2~ ,O _31.0 28,O±0.J7 L'" p" 'W" 16 .~ -31.5 23.8 : 0.27 3.23 " 1 () . ~ - 2 9. 5 2.'.2 ± 040 3.29 "aa 19.5-31.5 24.4 :0.34 3.10 ~ p,,'chrifNC/U.• "9" 22.5-27.0 24 .~ :0,62 1.87 '", 22.5 -23.0 22.6±0 12 0.24 S 25.0-2 7.0 26.0 ± 0 .3~ 0.79 s m,,,,,~di ,, ,,, 2 26.5 26.5 2 26.5 26.~ § /ricolor .W 19.0-26.5 23.7±0.29 l.l\J 19.0-24.5 22.3 ±0.42 1.59 as 21.5 -26.5 24.4 ±O, JO 1.48 "Mum u" 129 31.5 - 49.5 39.8 ±O ,34 3.8 1 " 31.5~ 42 .0 37.4 :0,25 2.26 SO 40.0 -49,5 43.(): 0 ,33 2.30 ";""'Ul' n 23.0 _29.0 25.7 ± 0. ~4 1.95 " 23.0 -2 6,0 24,4±0.38 L'" S 26.5 -29.0 27.7±0.44 0.97 26 .~ _JO.5 27.9 ±O ,29 , 27.0-30,5 1.37 u 26.5 -30.5 28.0:0,34 1.19 zap"'" " 1.22 " 27.8:0.56 ~ "An und.""ril>cd ' p.."ics. 10 l>c de""rjl>cd b)' C.W. M}'e", and J. Daly. "Sa""!.'" 0%8) ga"e the following IDC.'urcmenl': p , Iu ~ ubr;s: 10 males 18.5- 21.0 (mean 19.8); nine females 21,0- 23,5 (n'can 22.8). P . ,·;,,"' u. ; njne males 8 22.5-26.0 (mean 25.0): 10 fem.1cs 26.0 -31 .0 (mean 28.2) ~

~ ~ 1976 FROGS OF THE G£\'US PHYLLOBATES 5

yellow , green, blue, and intermedia te hues) o' ually 1857). P. fl(J",ulus (Boulenger 1882). P . 1"<'1<,rs; IIC'" form dorsolate ral or lateral stripes on a black or bro..... n species. P. pi..ru., (Bibron in Tschudi 1838). P , pul_ ground co lor. bot in some ,pecies, bright colo rs cover ehr ipt'(' /u.f new specie" P. smaragdm us new species. the entire dorsum or form dorsal marblin g. The venter P. mcotor(Bo ulenger 11199).P , Iril'iIlO Ilf.\' (Spix 1824). usually is black with brig ht spot s or marbling . In P, I·illalu,. (Cope 1893). and P. Wptlru new species. preservative, the bright colo rs become: white , gray, Table 2 shows all species and sub'flCcics desc ribed as brown, or black, and the coke pallem may become or referred 10 Phyllo/1(Jln . i nd i Sl i n g u i ~ ha bk from the ground color . The surfaces TaJpo l<'s (Figs. 10-12). -Tadpole measurements of the mU!iCles arc sparsely to densely flec ked with are in Tab le 3_ The tadpoles illu, U;Ited herein arc dark pigment. Palatine bones arc abse nt. The or­ identical in the follo"'i ng characters: Tbe oral papillae bitosp henoid region is partially or com pletel)' os..sifiN. border Ille entire po'terior lip. but borde r onJy tile The ornossemcm is present. The vertebrae usually are lateral por nons of tile anterior lip. Tbe oral disk is not fused. latera lly indented. The den ticle formula is ~. The COil/till. -Phyllo/J

TABU 2 T'hc 100 .peeie, and subspecies described as Of refcrnd to PJrylIoI>aln. All naJnC$ in til'

NAM E AUTHOR ASO DATE OPI....IOS OS STATU S

a1D8"J'O'!'" Bokel1lWln 196 7 Col,,.,crl.... olo8"J'O'!... albog""t1lU3 BoulenCt'T 1903 Co/(mrlw,,/bo,llJlOt... onrhotr.,; Noble 1921 Phyllobatu "nrhotr.,i " "rala (H)'/o) Wie1"rni" bau l" i IDr ndrobau.} Melin t941 Ph,·/I0b0 tcs ban /cri bralriciM Barbour and Dunn 1921 PI,,'IIQbat<,' /liI~..bris brtb<'; IH.,u-l...} Nobk J92~ CoiOSfrlw brill"...... Bibron ill s.a,n. IM I Plryflobo u . lJic<>Io, """""bacGgr; lH_rIout/tu} Jimenezde Ja Esp.da 1870 CoIour,hllJ bo'ia1UU (I' rwtlorrapU! Boulenecr 1902 PIry. ~lobot r ' bo/i-.r-... bo." rtrgr ri (Prwrh<'ropis) ..- 1909 Plryllobot cs bowl"ngr'i brd"" ryllil' gaudic lwudi brom r/k ol" Test 1 9~ 6 Co losrn h". bromdic<>la br"nnrUJ (ProSllu:rapis! Cope 1887 Co /oslrt h" J brlilrmru.J capiwba Bokenn...n J967 C<>Iosulh.,. c"pixaba Bokenn an n 196 7 C<>I".U IN" c" ,ioca cNlh . W. ~ la'1 1873 S)TTItopIw. eNlI.., ... c"'ocw (Hyliwhu) BouJenger 19J2 COwlrl/.... chocot'mis evi4uU (HyliuJllS' Boulen,er 1912 CdtW<',W roi/Qris cyui,lOlllltoidr. Cope IBn S,-rrltt:>p/tIIJ ryUigrwlllroidc. d..M; (PrtnllorrapU' Rivero 1961 Co/MIT'''''' <1_i clclOl'tS Giinlher J808 nomen nudum rq lMlwritJl;. (I'nmM rapis! B~Uf J908 E/rlilIM rodoc T)-f... liI"iurilO'lU upin(JJa; Funkhou>eT 1956 Plryl/obarrJ r JPinoJai rlU:nrmi s (Drndrooolr.) FilZinger in Sleindachr.er t864 Ph )'l/oha rrl pic'"' lem" ",/jJ (ProJlhn api.) Bouleng er 1883 PhyUohulrJ Irmo' '' /is ImlOraliJ (P",sllu:rapi.! Barbour 190,S PhyllobalU ho lillrnKcri fr Utu (Prollltrrapis) PeracCI t904 Co/m tclh ...I csrtu BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY No . 27

TULE 2 (conlinued) fI(Jw,pi(1(J IHyl"l'lni(J) A. LUll 1925 Phyllobale. pierus flolawr Dunn 1931 ColoSl/rhus "uhieolu fuliginos as (HyIMalus) J;rncne~ oe la ESllterap;s) Boettger 1893 ColoSlnhus I,ini'aris It"igru.. (HylmralusJ rowler 1913 £/tulneroducrvl". dia'lt"'U Itylaefi,rmi. Cope 1875 Eleulhtrodo,r,'lu" It)''',../ormi. infroguuulUS Boulenger 1898 Colo'lel},"s in/ragu""'..., ingeri -(Oe",irobale. 1 Coehran and Goin 1970 Phvllobote> i'lger; incuinalls IProSlNrapis) Cope 1868 COIOSlNhu,- ingulntllis intermFllius Andersson 1945 ColoS/elh"s in/trmedius king.bu'}'i Boulenger 1918 Culo<""hulor Irili""alus Bou len~'Cr 1883 Col",,,,,,hu ., I,ilinearu." rrinllmis Gannan 1887 Colrmerh",' trioiloli.' I,ivi"ol" (Hyla) Sp i~ 1824 Phyl/"I""e, "ivi"aru.' " ivlu", ,,.'· ma,'ul",u" (Dendrob"'es) W, Pelers 1874 Veod",b

P. bowl~"xai . P. f~m"ruli•• and P. tri"il/"tuJ have resemble~ ~o me weblcss species of Col"i/('lhu•. The (in pre"':l'\'ali\e) a dark brown Irans'erse band on lhe re lationships among the three subgroup~ i,, /{~ ri · po...enoe ponionof the throaL pns (Polder 1976; D . TTiIVis. ~r.'l(>"u l is unlnow'nl resemble~ them . ahhou¥h b"li';,mu. also communicmitm ). In ~pecie s lhal use the T) pe;;: posture ,

T~ BI.£ 3

TlHJp'lc mc.'urcmeOls (mm) of nine specic. " f Ph."II" f>

e ) after tile . peeifie epitbet j, tbe ladp,k ' ta~e of G~d number ( i ~ parcmlle",.) i' tile .a mple site ( n) , Follo....ing thi' arc the kno....n minimum and ma'imnm Ilead·b<>d)' Icnl1lb' for tbat ,tallC. a , o"' ma. and lbe kno.... n minimum .nd maximum total length' for that ' tagc Data fur diffe",n t stages arc "'paraled b)' semicolon'.

, pede'" 4(J (I ) 10 .~. 27.5: 42 (I ) I U. 26.S: 44 (I ) I 1.5. 16 .~ b",.lr"Krri 28 (91 3.5-4, 0. 10.5-1 1.0 up;"" ,,,i 26 (3) 4.0. 11.0 !rmora/j. 28 (II) 3.5. IO .~: 29 (I ) 5.5. "; 30 ( I) 7.0. -. 34 (2) 6.0- 1.0. IH I' PCIr'" 26 (8) 4.5. 12.5: 28 tn 5.0 . 15,0 pklll' 24 ( I) 4 0. 11.0; 15 (9) 3.0-4 5. 11.0_12.0 _'u"ai"", 24 (2) 5.0. 14,0; 15 (5) 5.0. 14,0-15,5 I ri'~llUf'" 23 (196) 4 0-5 .~ . 12,5-18.0; 29 en 6 ,5.10.5; 34 W 80. 18.5; J7 (2) 90. 26.5-28,0; 38 (4) 8,5_9.0. 26.0-30.5: 40 (I) 10.0. 31.5; "2 12l 11.0. 24.5-31 .0; .u ru 13.5.21.5 25 (4) 5.0--5.5. 13.S": 26 (3) S.5. 1 4,~; 28 (3) 5.<)--8,5. 19.0-22.0"':19 ( 1) 9.0. 23.5: J1 (I) 12.0. 30.5; .\S il l 11.0. 10,0

- An undc >crihed . P'C..... 10 be \k ..:rihtd b) C.W, M) CTs andJ. DaI). "Someor all >p:<" i"",n . un""''''II""bIe beuu~ ul ~ge 8 BUULTf,'" OF THE ""ATU RAL HISTORY MUSEUM OF WS AS GEU S COUflo7Y No. 17

the m~le sil\ on lhe l ~ )' i n ll: ,itt both ~fore and after ~ag". i.e .• lhe ",hole brood is nOl nece '>ilTily carrie d oviposinon. hut SenlTt ( 19.'6) p ro ~ eoJ Ihal ,~nn re- or deposilc'd simullaneou,ly ( R u l h~n and GlIige 1915; lea"" ' 'curs only afler uviJ1<" ition. Sentf 1936; Stebbins and Hcndrid ,o n 1959; D. Travis Kno n oVlp'"it ing , ite, are ( u 'uall~llea "e' (usually 1"",..,,,,,,1 cummlmimliOl1). Kappler's ubservation (qlK>l<.-.J anached. MlfTIClimes fallen ) in D..",J",hal('S and Ph.'"l­ in Bocfenger 11I1I1Ib) tha I P. I ri "itltone. pud dl e ~ lInd retrll/l.,porb u, lad po les 10 large r puddle> or in holes) in "OOle >jJ«ie s of CoioMerl".., (Oe -e r 19.12; proboobl) is incOll'eCt (Senffl 19-'6 ; Stebbi", "nd H~n · Senffl 1936; Dunn I~~ Oerner 1951; Stebbins and drit"kson 1959 ). TadpoJe-e"""iers ~ not (itS forme rly Hendrickson 1959; SihcnolUne 1973; liiddede 1974; be lieved) u clu sj' e l) ma le . Fem llle tadpole -carrier s P(llder 1974a . 197me. 1"i>Ur· CQIttI~ .• , Polde r (1976) noted lh al the word " guard ing" which i, know n to carry one I" 27 tadpole, (T able 4), less appro priate than lh~ word "l~ nd i n l!" for (hose resembles C"I""fflhl.... . species [e.g .• D , auratu, and P. l 'iIlUlU~ J in ....hich an f ·""d. - R ,R. Snelling (LA CMl kknlif

adul t mm:1) ' visils t~ egg' l)(Y\Oo and t~n. "llhough ar h contents of 14 , pecimell\ of 10 ~ p«ies of Ph..-I· in some spece-, of Col(m ttlu.s, lTW guard ing (d.m'ing lobl of thr spe cie . pre yed upon inha bit (M udrlOC k 1% 9 ). liller on the (ore,t nonr. After the ladpoles ha tch. lhe}' wrigi:1e onto the bac'k P" lder (19 76) nuled a difference between captive of II paren !, ap parentl y ad her ing by their be llies 10 Dnulro lmlf.l (five species) and Ph.l'!loh the tadpolts in " ~mall . , lill noted that D_ <11""'11.' eals onl)' '"lhe ~ mallt >l insects bod y of ",'altr. ",here lhe) co mplete t he ir de~l op­ and the ir Iar"ac; lhey were DOl " ble 10 lIWIa ~e a ha lf. menl o Transport and dcpo;. ition !>t)lTlCtimes occ ur in ~rown meal,,'orm or a "nmolh lan-a·'); PIn'lloha,'l

TA BL~ 4 Tldpole-<:al'Tyint in P "yll<>bcut'~ . n: numhrr of came... laken; Tadpok.: minimum .m .~im"m (mean. rounded to loweS! " 'link numhel"l numhrr of tadpolc ,"'".,...... C..., « ;",' , , ". .\.I41),j bt:I'Vod ' Sollle or all data "nkJ'KJ\\In . 'G an. (1960) saw adult, carry in, eigh1 to 10 ladpolc' "Mean of 21 lot. for whk h numbe", of uodpole> = k"" "'n 'Mean of 21 ..h,11 1I\ilIe. '..... " ...... ribed .pee;"•• '" be de..rnbe

eats both small and large prey (such a~ meal worm s. T.... uS

and earthworms longer than itselO. D. Travis. ...ho ~h "",,1(_ of 14 ma"h!.. Dendrobates and the presence of teeth in Phyffobal~ s . Ba sed on the species th.il l Polder observed, ho weve r. Food Cal(llory A( 4) B('l) tee th cannot be the sole explanatio n (I' . Iril·iuul US lacks teeth) . nor can body size (P. villulu.< Is much AnlS 49.4 78.6 small er than D . a::. u r~ us). According to Polder , in Tenn ilCs 17.0 21,4 Spiders 14.S 7., feeding behavior, Dendrobates is a "seeker." hut Bulin H 42,8 P hy lloha / ~s is a "sitter and watcher." H"mnpleran, and lrue bugs ,.. 3n Sumlllur}" of distribution (Figs. 13-1 7l .- Sped es Flies '.0 21,4 of Ph}"lIoha t~ s occ ur from Costa Rica southward to Wasps L3 21.4 Inseel w...u Doll" Penl , Bolivia, and soutbern Brasil. Mosl species of 7.' Sprinllail. 0.' 7.' the'-C diurnal. ler reslrial frog s inhabil wei tropical Gras.\hoppen 9.' 7. ' jo rest. Reliable colJe,ctinllrecords are below 1800 m. 0.' 7. ' Only 1...·0 (I'. lugubr;s and P. l'itla/us) of lhe 20 """"""""'" species live in Central America. and lhey are d,>,><: ly related to a Cbocoan species (I' . ourQra~lIio} . Ten specic's ir. bo:ss/ui, P. bo/i\·iaIlUS, P. f~moro/js . P. ingu;. P. pan·ulus. P. ,,"USI. P. pic/us. P. JltWrug· Panama; one: species II'. /ugubris}; Colo mbia; eig ht dinus. P. trivinauu, and P. zaparov dwell in the spec ie. (I' . a urola~nja. P. bieo/"r. P. boule ngeri , Amazon ian regi on. Fou r spec ies (I'. f~lIl ..ratii, P. P. f emorulis, P. ingtri, P. (1(m·ulus. P. pic/us. and pin uJ. P. pulchrtpecna. and P. " il'illmus) occ ur in P. /r;"i/fu/ a.l); Ecuador: nine spec ies (I'. aruh"l1)'i. the Guianan region; all but one of these (I'. putchn­ P. b(}ul~tI8 ~ ri . P. {'.Ipin"sai. P.felll"rali.l, P. f4lrvulus. p{'c/UJ) also occur in the Amazonian region . Four P. picllt," P. tricolor , P. trivntutus, and P. t.a/l('ro); species (I' . aurotaenia, P. bicotor, P. fx}f/l~ngt'ri . Peni: nine species (Phylloba/t;'.1 species, P. an/hon)"i, and P. eJpillo.'ui) inhabit the rain-drenched Chocoan P. b'l, s/e,,·. P. femoralts . P. purvalas. P. perer.li, region west of the Andes. One specie s (Ph)'l/oha/n P. pic"IIIJ. P. lI1lC1raxdillUJ . and P. "il'ilta/us ); Bolivia: ' Iledes) occu pies moderate elevalions on lhe eas tern IW O species (I' . m,liviallu.I and P. piNUS); Brasil: slope of tbe Andes in Peru. Two species IF. Iric"lor four species lP. femorali.l. P. pic/a.l. P. pufchripec/us. and P. boulengu; VI " close relalive) occur on tbe and P. Ir;l·ill,I/US); Venezue la: one: ~pecies (I' . pic/a.l}; ...estern slope of the ATlIks al moderate eleslliions in Guyana: lhree species II' . femoral;s . P. p;NUS. and P. ~uador . One species (I'. amlwn)"i} li\"\", in the PacifIC /ril·irtalus'; Suriname: three species II' . ftltlOralis. P. Iowland~ and Andean fOOlhills of south...-estem Ecuador piNu .•. and P. Iri"illaIUS); French Guiana ; 1"'0 specje, and l"IOTlh ...estern Peru. I P.f~moralis and P. piNUJ}. Tbe dist ribution ...ithin polaical units is a~ fol lows: I have discussed the lOOgeogra phy of dendrobatid Costa Rica: IWO ~pecies (I'. lugubris and P. "illatus); frogs in an earlie r paper (Silverstone 1975).

K~J /(J /h~ Genera of J)t'llclr ob/ll id(l ~

Ia Term inal disk of thir d linger usually less than 1.5 limes width of third linge r; dorsu m dully colored or brightly co lored in life; maxillary and premaxillary teeth present or absent; tee webbing present or absent ; first finger equal 10. aborter man. or longer than second; palatine bones present or absent: omosremum present 2 lb. Termi nal disk of third finger usuall)· IWO or more limes as wide as lhird linger ; dorsu m partly or entirely brightly co lored in life; maxillary and premaxillary teeth absent; toe webbing absenl; firsl linger shorter man second; palatine bones absenl; onjouernum present or absen t Dem/robares

2a . Dorsum dully colored in life; maxillary and premaxillary teeth present; palatine bones present or absent . Colostethus :lb. Dorsum panl) or entirely bri~hlly colored in life; maxillary and prema xillary teeth present Of" absent; palatine bones absent Phyllobatts

la. Basal webbing present between roes 1I·l lI-IV (Fig. 18, lefl) . l b. Toe ,,'ebb ing absent (Fig. 18. righl) . 10 BULLETI.'" OF THE NATURAL HISTORY M USEUM OF WS ANGELES COUf',7Y 1'.'0 .17

2a (II). LiEli1latera l ~I ript complete tfrom groin 10 upper eyelid; Fig. 101: don-urn red or not red in life ... 4 ~b . Light lateral ..uipc: inoomplcte Of a ~ n l. although complete ventrolateral stripe (from groin to uilla) may he pre~nl (Fig. , lAo 0 ; don-urn red in life _. •••••.••••• . 3

31 (l b). Adull ' OOUI-venl length rdatively ,mall (lrlO" n .pecimc:n' 16.0- 17.5 mm); Ctocoan region of Ecuador . " e-.t of tbe An(\e, ,Fig" IA . SF. 7F) P. n pi nma i ( pa~ 29 ) 3b Adull snout-vent length moderate tl llO"'n spccircen, 00 .0-«).0 mm): Amazonian region (If EcLl.ildor. (ll'>l oftht: Andt:, (Fronli,pie« I: Fig,_ IC, 80) P. zaparo (page 3»

4.1 co. !-Iedian dONlI ...ripe pre~nl ('OlTlelime, e xpanded 10 cove r ITlO'oI of back; ng'. SA. D) 5 4b. ~ Icdian dorsal >triP<' a"'ent . 6

S. (4a). Ad...11 >oJlOIJ.I -~en\ lengrh 19.0 -245 mm in males. 21.5 -26.5 mm in females ; bones green (Fronlispiece II; Figs . 5A. ISF) ...... ••. . P. mcotor (page 32) A"llIIt snOUI'vent Ienglh 16.5-1 9.0 mm in males. 19.0-2 1.0 mm in females; bones while (Figs. 50. 7A-B) . P. ,mrh""."; (page 27)

6a (4b). Do rsum red in life .. . P. t"p',m (page 33) eb. Do rsum brown or black in life 7

7a (6b) Adu ll snUU I' vent lenglh 1.'i.0-20.0 r um in males, 17.0 -21,0 nun in km

8a (l b). Lighl proximos'cntral calf.spot present (Figs. 9C -F) ...... 9 8b Lighr prox ill1O\~ntra l calf. spot ab-e m II 9a (Sa ). Lighl lateral stripe presem I. 9b . Lighl late ral smpe ah!

lOa (93). Lighl lateral s,ripe iecompkte (Fig . IA); dorsum red in life (Fronlispiece I; Fig. .'iBl . .. P. P

I Ia (8b). Dorsolateral O)f lalerdl slripe pre-em 14 I Ib. Dorsolateral or lateral smpe a ~ n l 12

12a (lIb). Adull snout-vern length 32.5-42.5 mm 13 12b. Adu lt snoul·v..-ntk'ngrh 17.5 -24.0 mm (Fromispiece I; Figs. .'lB. 7D) , .. P. p

13a (12 0). Skin of back strongjy gran ular t granule s sometimes disappear in preservative): hack immac ulate 01' with sPOls and marbJin l,': : Cordillera Azul of Peru {Frontispiece ll ; Fig. 4F ). Phyllol",rn species (page 2U) 13b. Skin of back slllo01h; back immaculate ; Chocoan region u r Colombia Wig. olE) . P, bicolor { p;I~"- 2.1)

14a (l la) Dorsola teral Of lateral slripe co mplete (from base of Ihigh Of groin 10 upper eyelid : Fig, IB) 15 14b. Later al 'r ripe incomplete (Frontispiece l; Figs. .'iB. 7Dl , P. 1'0" 'u/" $ (page 31'1)

15a (14al. Ma\illary and prema\illary teeth prescnl . 15b. Mu illa/) ' aoo pre ma \i llar) teeth ab~nl . " 100 (lSb). Aduh snouHem jength 3 1.5 -49. 5 mm: oo""wn wmetime, bears light median stripe; venter dark ...ith" lighl marbling or 'pOh (in prcsero·alj"-e, verner may become enlirely black: Figs. 2A-E. 7C ) .. P. /ri­ vhuuus (page 45) lOb. Adult snoul-vent length Ies, than 30 mm; corscm never bearv lighl median stripe: venter lighl ...ith dark marbl ing or spot s (in pre-efVali'e . venter may become entirel y black) 17

17. (I6b). Cnnspicuous light ()'e lJo... in life) prox i~ thigh·"JlOI pre....nl (Frontispiece II; Fig_6E) . P.!",I­ C'IIripl'ctus (pa ge 43) / 976 FROGS OF THE.GV'US PHYLLOBATES /I

17b . Li ~ h l pnn imodoNl llhigh-~pol al><.em lFigs. X:.9Aj . .. P. boIMu"u.• (pa~c 35)

18a (l 5a). l ighl stripe dorsolateral (i.e., posterior end o f stripe i, u" back ut ba-e of Ihigh ); C..ntrul Americ a or Sou lh Amerka we-t o f I ~ Ande , ( f.'jg\. ~A-D ) 19 18b. Li~hl «npelall'f""l ne.. po'>lerior end of -tripe is in groin); Soulh Amerka ea-t (lf the Ande, I Figs. 2F. M-C. F) ...... •. 21

19a ( 18a). Venter blad with lighl d

20a (19b). Adult sncut-veru lcngeh 18.5 -21,(1 mm in ma les, 20.5- 23 .5 mm in fema les; dorsotarcral , uipe u'ually relatively narrow : light area' un dOT,;!1 surface of limb , go ld, yellu"" or ye llo w· green in life; Caribbean dr..inage uf Co -ta Rica and f'-..nama (Figs. 4'\. 8A) P. "('(Uhfi. (pa ge 24) 2Ob. Adull MlOOI·\-enl lengt h 22.5- 26 .0 mm in male,. 26 .0 -31.0 mm in females; dor'oOla1erai slrip: u'>llall} rdali\-ely wide: ligb t areason do l'>al surfxe o f limn.. bl ~ -g l"C'C n in life ; f'-.ocifie dra inage of Costa Rica in Gol fo Dulce region (Frcnnvpiccc I: Fig . 4B) .. P , "ill

21a (l8b). Ad ult snou Henllength Ie" lh.an 32 mm 21b. Adult ..notI1-\"(' nl length greaier lha n 32 mm (Figs . 2F. 6A-0) • . ..• P . hcr.u/tfi lPi'gc 45)

2201 (21a ). Ve nter marbled or spotted (Fronnv picce II: Figs , er. 9B) ,. P. {'t'll'fJ; (page 37) 22b . V..nter n'11 marbled UT sported IProntispicce II; Fig . 8C) ,., P. Wlllfuxd;nu.• (J'l'Ige 44 )

F1GI'U I . Ltuenl aspeel. ( A) PIf}'llobdrfl ('spi".'",; , paralyI"' , Santo Domin", de los Colo~ . Ecuador (CAS·SU 10375), sho"' · ing iJlC<>mplelC Iighl laler~l suipe; (B) P , bo ulmJl~fi. 1mbili. Colomb;~ (USI\M 147440), .!I".... ing complcle lighl lalcral stripe; (C) P . UlfJUFO. paralyP<' . Saray.cu , Ecuador (KU 12(670). she.... ing \'CnlrolalcraJ stripe. Diagrammalic 3JId noI to ."'" ~. 12 BUU£TIN OF THE NATURAL HISTORY M USEUM OF WS AXGELES COU;"7Y ."io. 17

E F1:Gun 2. Donal ••poet. tAl Ph)"lIobmu "lvill at..... PallCm I, ho lol)'pc, Rio Tefc . Bru il (redrawn fn,m Spi. 1824:p1atc 9. fi, I ); (8 ) P. rr;villar ~ J . Pallem 2. Sam. " abel, Pe", (USNM 166764); (C) P, 'rM JI"tuJ. Pauem 3. N• •""u (lehe rg1c. Suriname (RMNH 6413); (D) P. ',i "rtaru" . Pallcm I, Pampa Hermon , P"ni (AMNH 43 11 ~ ) : (E) p, trMlla/u.•. Paltem I. Pampa Hcnn o ~. Peru (AMNH 43 117); (F) P. ba..lm", 1!<,lolypc. Roque. Peni(NMG 51 n, Diagrammalic and not to scale .

r FIGURE. 3. Don.aI -- !'«t. (AI PJryllobau!. pKtld . Pancm 3 (DmJro#1GtrJ ,•• leckK)'pe). Rio Mamore . Brull (NHMV.' 19190:3); (DIP. pirtMS. Panem 4 /PtNJ'Obtzu', h

F1GCU 4 D<:>fu,1 ... pect. (A) PIr.-llobJlr . '"gltbri•• Bam OO . Costa Rica (CRE 1182); ( B)'P ..in""... Ri"""" <10 Osa, CoI.ta Rica (CRE M); (C) P_""rot_"ia. narTOVl- ' lolripN fonn . PIa)a de Oro, Colombia (LACM 4)193); (D) P_"_ _ ,,ia. broad.miped fo nn . Quebnda Bochorarna. Colombia (LACM • .'8(0); (E) P_ bit:'oIt>r Santa Cecilia. Colombia (USS M 137:J.661; (F) P II"IlD- bal" ' pecie>. Diriooria. Peni (FMS H .s6272l. Diagnmmalic and nollO "' .

FIGI'U S. Don.al npecl_ (AI Ph) llohutr. uic-olor . kclOl)pe. """"=ir. EculIdol"(BM 1941.2. 14 18); (Bl P. ptU""tII... . PuenoliM. Ecuador tKU 1222(1 ); (C) P_bali.... " ... . kclOl)pe. San Carb. BoI;"ilI C8 M 1941_2_13891 ; (D) P. . Ec-'or (A)I.!S H 13140); t El P. botM"gui. Imbili. Coloml>ia(USSM 141433); IF) P. rlpi_i. ,-anoe. SanIODominJOde Jot, Colortlc\M CCAS·SU IOnS)_ D ilI~:r"mma ic and noc k> Kale 14 BUllETIN OF THE NATU RAL HISTOR Y MUSEUM OF LOS ANGEU;S COUNTY No. 27

FrClJRl 6 . Dorsal (A- C. E-F) and ventral {OJ . ' pel."I, (A -D)/'hyllo""te, ha"leri. Chula,., Peni (US :-JM 166903A. 166903C. 1669038 . 166903C); (El P, p ~l~h' ipeCius. poraty pe. Semo do Novio. Bra,i l (LACM 423(4); (F) P. pelc,",i. holOlype. Santa ".bel, Peru (USNM 16(763). Diagrammatic and not In scale

FIGURe 7. Vernral aspcel. (AiPhyll"i>

F1GlJU 8. Vent r~ J aspect . (AI Plryllobal ts 1"1"l>rl•. Bambu. CO' la Rica (e RE 11 82); (B)P.

F IG l-~ 9. Wllm,I~ . ( A ) P", I/obo'e. hoIi....""•• kelOl)pe. SMl Canoo. Bolivia (BM J941.2 . 13.89); (B) P. pr'ersi. pamype. Rio Vi\Ol; . Pvll ( ~ ~lf 36183); (O P. in~ri. p-aIJipe. Aoerrio ..... Rio Pe-w:ado. Colombia (US1"M 146849); (D) P. pic"". Pal' lCm 1 (Hylaplt w. flo""!'kro. IOpofype). Bd o Holiron tc. BlaSiJ (WCAB 4218); tEl P. pic"a. Parlem 2 lDt"Jro/>Q,u 1>ra«<11... Fitzinger in Slcindachncr. lccioI}pel. Silo Vicenlc. B,..il (1"HMW 38 18;1); (F) P. picr.... Pallem 3 (D~"JroOO'eHW:IfC "'is. ke to­ Iype). Rio Maroo"; . Bra.il (NHMW 191'XH ), Dialtrammatic and IlOIlOocale 16 BUllETIN OF THENATURAL HISTORY MUSEUM OF WS ANGELES COUNTY No. 1 7

• 0-.

F1GllU. 10. Tadpolt of P hylloba ,~s " " alla . "C'n"al, 1••..-0.1. and do".,J ospe<:u.. SIIll" 29. •aL~ n from tQck of Idul l ( b'I1 l"" ~ .fler JI'O""in, in Capllv;!)). RincOn de 0... C"".. Ri<. (e RE IIOl). ~ equ al. Oft<' mill;_ I..-

G§..

c

o / .- ~

fl(l ll ll.E II, T8dpob of PII.,l lo11;. ( U S ~' M 14 ~248 1 ; (B) p , ".pi",.",i, SlaIC 21>. from to.;k of . MI ...... 9 km ....,.. o r SmlO Domin"" de ...... ColorIodo>.Ecllad« (CAS·SU l~ 86I; (C) P.~_IiJ. SLo ll" 28. tmm.....kof odul' frmale.SatQ C« ili•• Ecltldof tlW 112310):(0)1' _ ... &Jill .... ~ 2S. from back of .Ju ~ male hoIoII)p'. PorI de Al Ui:•• Ptni (LAC M 644 36); (E) I' P'"";' Stare 26. from bad ofadull male. RiD VilOo: . PnU(P'-IS H 36nlll; (F) P , pin... . S ~. 25. from back of adult male. h iarnal.. Boli\h male. FrNerik .....dlem IV.nI..... s...... (BM 1936 ,9 H.). Linn "'l ual one mil"","", J976 FROGS OF THE GE/I'US PHYLLOBATES 17 @ ... ~~ ::..::::- • .@ ,@

, ~~ .~

"~~~

FIGl:at 12. Tadpo"" of P~llobal~J • ...... lhp.bOfll... ""'"" tadpo"" illu-"QIC'd in fig•. 10-11. (A) P. botd,,"g..' ; (US~M 145248): ( 8 ) P. ~illal ".•{CRE 9(3);(Cl P . ..spiltOJOi (CAS·SU I~) ; (Dl P. /t'mMali' IKU 112310);( E) P. _mgdi",.. (UCM (4436) : IF) P, ,wit',.. (FMSH 3671ll); (G) P. pin". (L'MMZ Sll947): (H) P. ,ri.inalld (BM 1936.9.) ,6). u..... "'lual."... millimclC1'

o '.' ''"UTI' ~ ••"",.,.

v ••_ w,,~ ..,!.... r- a • ~ """'0;­ • '. . ". ~t~ . +

r- -t 18 BUULT/N OF THE NATURAL Hl srORY M USEUM OF WS ANGELES COUNTY No. 17

·-...-.... .".". · ::.- ,, ~ .. '. " .

-,- . .' .

+ .. ~ _ . ~..., · ~ , .. c- ...... --L

+ + 1970 FROGS OF THE GEAUS PHYLLOBATES /9

[J "." 0 04'" lUI "

F...... lGl:u 16. Di..nibulion of PIr.'lIobu,r . ;,,~ri . P. pun ...I.... P. prtrm. P. ptJrltrif'«llU, and P. """'atJi",,•. Am abo\co1(0) m 20 RUUET/N OF THE NATURAL HISTORYMUSEUM OFW S ANGELES COUNTY NQ,27

o

Flcicu. 18. V cnu al~ . Ii"" rOOl or I'ltylJobt. '~I. """'*ina difference be,,,,ecn t-ally .. cbtlcd -. n-JII·lv (loft p _ ~_tIlu . UCM 61011) .-d ,",cblru IOn (riJbl:; P. ....,."...... LAOI,I980I. lirot eq...... o. millimelcr

bicolor ,roup bru, and P. vi,lalWJ in IMBer size (Table 1) and in This group includes rIVe . ~ ~. lPlrylWbotu speces, lacking stripes, and from P. bicolor in llaving a strongly ~um P. aurolfunia. P. bic(}/or. P. lu/C" br u, P. v;ttalltJ). granular and in sometimes having a different Definit ion: sec ut-venr length of aduh s sma ll 10 donal panem . 'The red. red-orange, or orange ground large (18.5-42..5 rnm); lox. wcbk•• ; palmi smpes color resembles lhal of P. biro/w . bUI somclimcs is (...hen presenl ) dorsolateral, nol lakral (i.e. , poslcrior broken by black spots and mlllbling; in some ...pe<:i· p~inanll y end of stripe ts on bilck at base of thigh . not in groi n: mens, the don;wn is black. and lhe red Figs. 4A-D); light proxtmoventrat calf·spot a~nt . is reduced 10 spots and marbling (in P. biro/or. lhe dorsum is smooth and uniform red·orange, orange. or yellow·orange). Description, -c-Measuremems are in Table I. The Plryllobo/n species skin is strongly granular on lhe back and the dorsal (10 be described by C.W. Myers and J Daly) surface of the hind limbs and smOO1h on lhe head. the Fromlspiece Jl and Fi ~ure 4F forelimbs. tbe venter. and the ventral surface of the hind limb. (in USNM 166904A, lhe skin is smooth in s lidc~ by Shcrbrocke 196], I'hyllobbu/u bic% r, Liilin" 161- 174 of lhe hind limbs was strongly granular in life). Maxll­ lary a nd premax illary reerh are present. The fir'l Drfinirion .-TeClh present; toes webless: dorsum finger is very slighlly longer than lhe second . The red , red-orange, or orange . sometimes with black toes are ,..ebless. ~POI S and marbling , or don;um black with red. red­ In life (color slides by W.e. Sherbrooke; venter orange. or orange SpolS and marbling; slripes absenl; nO( visible), in USN ~1 l66904A (Pronnspiece m. the lighl proximoveraral calf,s pol absent; venter red, black, iris and tbe tympanum were black. 'The dorsum was or panly red and panl y black. red wilh black spots on lhe mid·hack and postel ior Diagnosis.-Ph)'lIob

In preservative . in US ~ ~ I 166904A. the tympanum (27) , Ponerillltt: fragments, ~I )' rmeci nae : fragll'lC'nts : i~ dark brown . The dorsum is ITllNl)' c ream. with a COLEO PT ERA : fragments of beetles of families 'mall dark bro...n spot behind the right upper eyelid Staph)linidae, Bostrkhldae, Curc ulionidae. and Ela ­ lind seven ~all irrellular dark bro.. n spots scattered rendae , also OIlC' unusual beetle larva of unkno wn over the back; the extreme po'>terloT back is dark bro...n . family; LEPIDO PTERA: Soctuidae? (l lan'a): HEMI · The side of the head ilfIdbod)' and the entire forelimb PTERA: Mirn ae ~ fnogll'lC'nts and !'land are cream . Tbe hind limb is cream, exce-pI. P,..dators .-Liiling (1971) sa'" a snake (Chi,o nild lhe di~ portion of one thigh. the \'e-n1fal surfXC' sp.) from the Cordi llera Azul allack and seize a C;apl M of lhe thigh. the antelm'C'ntral surface of the calf, and specimeno f Plr)'lIo bafcs speciC'S; the snake immed iately the ventr al surface of the fOOl . .. hich arC' dark brown released the frog, retreated . shook it-elf for 0.75 min. Thc C'lllIt'1l'lC' po'>te-rior belly is dark brown; the rest o f kepi its mouth ...ide open for 6 min. and rubbed lhe the venter is cream with an indistinct lillht brown wash sides o f its mouth against branche s. Th is repul,itm of a on lhe tbroat and bell)', predator (probably becacse of lhe unplea....nr, burnin g In preservative. in fl.1~H 56263·72. the- tympanum ta,l e of the frog' s skin secreuoej, oombined ....ith Lu· is dark brown. The head and beck arc: uniform cream ling' s observatilln lhat lhC' ffllp, in their native habilal, in four specime ns (except for a small dark brown area, ~it in open areas and contras t co nepicuous fy wuh the sometimes preceded by one- or two bfllwn dots, on the surrou nding vegetation and soil, , u l!ll es t~ that the bri~ hl e xtreme posterior back). Four other spec imens have colora tion of this species has an aposemauc function . one to several small brown spots on tbe mid-back and Distribution and habilal (Fig. 14). -c-Ele vauon po slerior bad; the brown area o n the e xtreme posterior record s are from 1300 10 1600 m. Phyllob1rs "w rOl~"ia Boulcnger; 1029. pla ~ 104, 56268. the venter is entirel y dark brown, fig. l, hololype: BM 1947,2.15.13, Peiia Lisa , Tadpot..s.-On 8 October 1966, W.C . Sherbrooke Rio Condolo, ChocO, Colombia. 92 m captured USNM 166904A , a 40.5 mm male; Ihe frog 1957. Phyll"bcl/ts a..'o".,,"u.. Dunn: 78. had six tadpo les (USNM 166904B ) on its back. When 1963. Ph)lI<>bltlr'li/w.t d....ell, al k»ser ele...· ventra.lll\le",. broad·slriped toio; only on a hill (the Lorna de One form (Fromispiecc I and Figs. 4<:. liB) is slightly Encamacitlfl) immediately opposite lhe presious 10e..l· smaller than the other, This fonn includes the holotypc , ily. on lhe right bank al 400-520 m, On the Camino a 28.5 mm male . The adult snout,vent length in milli­ de Yupe in the Serrunia de Baud" (near the upper meters (mean in parentheses) is: 19 male s 23.5-28.5 Rio 0 p" l!ad\' in the western Atratc dr..in..ge), where (26 .0) ; four females 26.5-30.0 (2t\.1); all 23 spcci­ the broad .slriped kill",,: is ..bsenr. the n"rro w·sl riped mens 23.5-30.0 (26.3). In life , the ground color of k,ik",.' reache s al leasl 500 m the dorsum and of the side of the bo:.xly is black. A Should Ihe IWO for ms of k,ik",' pmve to be distinct relatively narrow complet e dor ...rlatcral stripe extends ,pecies. lhe n..me P . Ulm>we"i" ...ould apply to Ihe fn,m the dOr!'al base of the Ihi~h a k ,n~ tbe back. along narrow-srripcd fonn . J,W, Daly told me the two f"m ls the outer borde r of lhe uppe r eyelid. and along the inter breed in capli,'ily. prodoc ing vi..ble offspring. camhus rostrahs to the tip of lhe snoul. where it meets The n..rro...·s triped ioko; resembles P. IUlluhr;s its fellow. lbe su-ipe is bright green or p,ale }ellow· ,,00 P. ";/I"/lU of Centr,,1 Americ a. bul is sep;r,lOge coe­ ventral SUrfllCe of the limbs ~ blacl. ....ilh blue or sum. lackin!! 'tripe~) . bUI a"erage ,mailer: seven sreen dols. The doIs are a[...a), blue in .pecimens specimen. 31.0-34.0 (32_1). ve rsu~ 4-1 specimen. from the SaTania de BaudO. The dots are 'p.an.eon the 32.5-e.0 (.\S.2) inP. hi...,I,,,.. venter and ITIOIl: numeruu.. on the limbs. In ~..:rva· Sf"mudl nmft'dy weig ht (Tok uyallla et at. 1969). In mum­ The dorsolateral stripes sometimes are nearly united lIl"ls, BTX .·au...:s an increase in permeability " f mem· by a dOl'Sa l suffusion (Fig. 40), whkh may also cover bmnc' to Na· , wilh con-cqucnt melllbmnc z,,,,I<>KY, -The Choco Indian, of lhe uppe r lil.e. ",hi rring t",'iller. consis lin!! of rapid ly repea led S..II Ju..n d r"ina~~ of Colombi .. exlrdCt bk>wgun poi"'>n notes. The ..:rics is repealed after ilTt'gular inlervals fro m both form, of P, uur{1/};,'; b> Boclloramio. four calls: 8.0-11 .0 (8.1); bmad-

Because lhe ti>lwi release, ih skin poi....111 only under 8M 191 0.7.11.53. USNM 48865. 122104. 145105­ ~ ll·e~' . lhe India ns pierce its visce ra wilh " wooden 06. 146630-JI. 147204...(17. 1 49 72~- 27. ISObJ2-4 2. CAS 115151 ). ske we r inserted throu!!h lhe lI1<,luth . and somenm es hold it <)Ver a fin: . The Indian, poison the tiP!' of blow. J!un dan , by beatin!! them on the back of ee fro!!; the DI:/i"it;on. -Teelh present; toe, .. eblecs: donum poison ad he re~ to ,piral !!rno"C'~ in the liP!' of the dan, uniform l"((I--onm~'e. ora nge. or yellow-orange : 'lripe~ t Fma Iob IlIh9. 1909; W",..en 1957). The Cho..'6 Indian, absem: li~h t proximo.-enlral calf.spot ab!ba,t's bic%r i~ a member of the DiHrib"tion " nd h" bit,,! tFig. 13j . _ Ele.'alion bicolo. group , dirfe r in~ from P . aur otaenia, P. record, an: from 60 to 520 m . P. au rol~"iu inhabil' !u/(ubri., and P. vtuosus in larger size (T able 1) and lhe wet fore' h of the Chocoon region of Cok.mbia. in lackin~ ' lripc:~, and from Phyllohatl/S specie, in "'e,1 of the And.:,. in the AlralO and San Juan dr"ina~, ha..in~ a sme o rh back and in lading do rcal ,pol> and m ar bl i n~ (Pln'//"1>a11:J species has a 'tron~l y granular bad and corneumes has dorsal ~p<.lls and U1(·"lilin,-eoW MBIA,ChtKfi: Pena Ul'lI, Rio Condolo marbling). (BM 1947.2.15.13); PI :. ~ :. de Om, Rio S:.n Juan (lJSNM 1506n-42, I~ M MlIl; LACM 4~792 _9 4) ; 16_24 ~ m W Playa DpcripliOl1.-s- Meusurerncn ts arc in Ta ble I (t hc de oro (US r-- M 146630_31, 147204_07); Rio San h an holot ypc i, a 39,0 mill male). The skin is entirely (USNM 159401_~O, 1594.'1 -1I6); Quc hr. d. IJoehnrama. smooth. Maxillary and pr e m a ~ i Jl a r y reem arc pre, en!. upper San J Uall drainage (LACM 43791. 43795-11(1); upper The first finger is s li~ h lly lon ger than rbc second. The Rio Napipi (LACM 4~782 - 1I9); Irail N:l....ccn Rio> Napipi and Opo~ado (LAC...l 4~m); Camino de Yupe. bet....ecn upper toe, are ...... bl..". Ri" Opo~:.d,i :.nd Domingooo drainage (LACM 7197M); In life. in specimens frum ncar Santa Ce<:ilia and upper Rio B""y (LAC...I 61020-21); All" del B""y (LACM La Union on lhe: uppe r Rio San Juan (Was..cn 19S7. 71979_112); Cmo DocordO ben.un Cucurrupi and N,IiIIl.....='. -'ta nn 1957. and my notes ). the ir i~ ""a~ black o r $;on Jwn dra jna~ (CAS 1194-15); Quebrada Tap;trill. 20 l;m redd ish brown . Tho: tympanum was panl)' black and :.,; Pale' li",.. S:.n Juan drainage tCAS 1198ll6). V" III: EI Paikin. 12 ~m from B""nllVCnlunI (USSM 1451(5 ); Io.. er partly )' eU(lY>'-oran~ . In some specimens. the lip of Rio Caling tUS:";M 1451061. (Of the LACM 'I'f"imen , cited the snoul w"-' bhock Ohi, ..... a' the reason for Benhold' s abrM:. only 43'795-80 1 are the: bro.I·>lripN form.) name P. m"lan"rrhinu.•I. The dorsum wa s uniform C/I:au d and slaim'd mal t'rial. _narrow·$lripcd form: bri~h l or,lIIge o r yeljow-oeange. The ~ ide of the bod y US~M .~800 . 1S9447; broad·.uipe-d fonn : LAOl w a~ !!old Ol" ~ l lo w -ora n ~. The . Ph.l'fI"htJIl/i bie%r Bibron in Sal!l'il 1841 were black, ""jth o r ""ithout numerous ycllo... dOh . ;\eara The venter was uniform black or black " 'ith a golde n Fillure 4E wash (....'mc timcs imerr upeed by a black parch on the threat). 1841. Phyllo/'rrlt'S Nco/or Bibron , Sagf1l : plale 29 hi" holo· In prese rvanvc, in specimens from Santa Cecilia 1)'PC; M H ~P 83ll. Cuba lin enur for Colom"ial. and La UnKln, lhe dorsum and ,ide arc uniform Ii~hl 1841. PIo,H,""II~' bico/",. Dum.:riland Bjbr,,,,: 6.'11 -1'>.'9 . ~ray ; the do rsa1 surface o f the limbs is uniform light m~/ dar k gray with clo \Cly "'I lighl gray doh. Popa~' ii n,IC.uea,1 Colomhia [. 1760 m) 18' M Phl/"!>icol",. Giinther: 'ltl The ventral su rfa,'e of the limbs i, dark gray ..... ith lighl 11ISII Ph /l" !>i''''.... Giinlhcr: III gray dotv. The venter i, uniform lighl ~r ay c xc..pl for 11I67. P!>,-I/"hclle. mel",,,,, !> i,,,,,,, Kcfe" tein; JS 4 -~55 a dark gray p;ltch on the thro"l in ....'me specime n" 11ItW P!>."I/"/oatt'S e!>"cM n.,{, Posada: 205-207, lype I,><:al· Pre.scrv..d ,pe:cimen, from the Rio Sipi are en tirely ily: ClVI;O . Colombia dark I>rown. 18112 Ph,"I<>i",u" bico/",. Boulcngc,: 195 Nm",'"ci"',,r t' ,_P hyllobatl:~ 11I82. Ph."lI,matn mrl"""'hi""', Boulcnll"r: 1%. rhoc,""ui.• Posada 1920 Ph..-lI" butn bico/"'. Barhour and Sohle , 3eN 1869 (lhe de-eriplitln ""as rcpri nl..d in 1909 ) has heen 1920. PIr.'I/.""',t'. "",lan". 10", ,,•. Barbour and Sobk: "1)2. considered a synonym o f f)f'lId,."hr;l n ~e·)c l k , 1.13S7 mi. the cok» ..... alll"'.: . toIuish I>htck bel"",,). 1957. PiolilobbDI bkolM t p;orl). Cock an a.nd(joia: ~5 -37. color pan..m usuilll) is sp'1l:led (but sornelime, rescm­ plalC 6A---e tonly lht fol...... iIlg _ IIt>I P . bfcoi",: bles tllal o f r . bi""lor). the -nout·'ent Ie n~ l h of 6 1 24 BULLETIN OF THE NATURAL HISTORY MUSE UM OF LOSANGELES COUNTY No . 27 specimen, from the upper San Juan drainage averages his description in the same paper (and probably from the 34.2 mm (maximum 37.0 mm), and the first finger same shipment) of another Chocoan species, Dendro­ is shorter than the second. Posada (1869 ) "aid that bates hiJlrio'l irll .<. from ·' Papayan," the Indians extracted poison from P. chacoensis. The The most t r u~t wo rthy Colombian locality record, Choco Indians today do nol extract pobon from D. are in the western foothills of the Cordillera Occidental, hislrionicu ,1 (Silverstone 1975). They do extract poison e.g .. Santa Cecilia on the upper Rio San Juan. The from P. bir%r; they call this species neara, one of highest of these records is the beudwaterx of the Rio the Indian names Posada applied to P. chocoensts Sipi. which is 1525 m according to H.G.F . Spurrell's (Wassen 1957; a herpetologist told Wassell to use the field notes at BM. name Dendrobotes tinclOrius. but w assens neara is uH:ali/ies.-COLOMBlA , RisartJlda: San1a Cecilia (FMNH P. bir%r). Posada sa id that P. chacoensis lacked 54187,541 89,54193 -94,54196,54204-05,54207.54212, teeth. but he may not have used a microscope . Ph}'l­ 5422t , 54233; USN~ 147225-26; A:"SP 25755: LACM loba tes ch",;o",n, i,l Posada is a synonym of Phyllo ba l<' s 60972,61050): vicinityof La Unionand SantaCecilia(USNM bicolor, 137359-62, 137364 _70). Chm:D: head .....aters Rio Sipi (8M Distrib Ulion and hahitat (Fig. 14),-Reliable eleva­ 1915.10,2t ,46- 47); t20 kmS Andagoya (USNM 139776-77); Rio San Juan (USNM 144994); opper Rio Alrato (ANSP tion records are from about 25 to 1525 m. P. bicolor 25756: USN'" 53168, 53t70-71). Vall e: virology field is reported from Cuba and from the foothills of the station, Rio Raposo (USNM 151381 ). upper Atraic and San Juan drainages. from the Rio DOUBTFUL RECORDS.- CUBA (MHNP 838). CO· Raposo. from the Serrania de Baudo, from both slopes LOMBIA. Callea: Quintana , 3200 m (FMNH 54183): of the Cordillera Occidental. and from the western Popayan (Berthold 1845): Atl/iaqu.;,,: Andes (AMNH t4028 -29); Medellin (AMNH 13540, 3(08 1); O weD: slope and a central valley of the Cord illera Central Serrania de Baudo (ANSP 25490). (all in Colombia), Only the record, from the upper Cleared a nd stained malerial,-USNM 13?360; Atrato and San Juan drainages and the Rio Raposo LACM 60972 have been included on the map, because I distrust the Other records. P h y lloba le ~ lugubris (0. Schmidt 1857) Barbour and Ramsden (1919:77, 113) noted that Lovely Poi,on-arrow Frog the type locality of Cuba is incorrect, and that de la Figures 4A. 8A Sagra shipped material from the Latin American main­ land to Paris with his Cuban collections. They thought that the type might have come from Mexico. P. bicolor 1857. Dendrobates IIlgllbris 0 , Schmidt 12- 13, Ihe syntypes; KM 101611347 (now lost), road from never has been collected in Mexico, nor in Cuba sub­ B<>;; as del Toro to Vole"n Chiriqui, Panama sequent to de la Sagra's publication ( 1841). The type 1858 Dm drobates Jugubr;s O. Schmidt 250. plate 2. specimen resembles Colombian specimens and probably fig. 14. came from Colomb ia. Dunn (1957) and Luling (197 1) 1900- 1902. Hyillp/esia tinCloria (part). Gunther; 207. thought that the holotype probably was collected in 1921. Phyllohares MllIridae Barbour and Dunn; 159, holotype: MCZ 8022, Finea Victoria near u nt, Colombia by von Humboldt. [limon,] Costa Rica I distrust the records from the Serran ia de Baudo 1924 Ph)'lIolmres lagllbris. Donn; 5. [exact locality unspecified]. Medellin [Cordillera Cen­ 1926. Hylap/esi" linrtv,ia (part) . MirandaRibeiro; 224. tral, 1538 ml. and the town of Andes [eastern slope 1952 Phyl/obates Jugul>ri s. Taylor; 640- 643. fig. lO. 1968 Phyllobares lugub"s (pan). Savage: 763_766, of the Cordillera Occ idental, 1357 m). I collected fig. 4F. near Andes and at eight sites in the northern and 1970. Dendrobates Jugu!>r;s (part), Cochran and croin' central Serrania de Baudo, but never found P. tncolor 19-21 (not the plate; in the synonymy, Den. there. K . von Sneidern, the collector of the " Serrania drohares lagahris O. Schmid1 and Ph)'lIobales de Baudo" specimen. has stated (see below) that he lugabris as used by Dunn and Taylor are P IU!l ubris; oflhe specimens examined,onl)' USNM has collected P . bicolor only at Santa Cecil ia and La I t8231 is P. lugubrls). Union. Specimens recorded from Popayan [1760 m, on the western slope of the Cordillera Ce ntra l] (USNM Deflllilioll._Teelh present; toes webless: dorsum 137359-62. 137364 -70) were co llected by K. von black; complete gold, yellow. yellow-orange, or orange Sneidero. According to von Sneidern (ill Jill. ), "All dorsolateral stripe: blue or white ventrolateral stripe; the P. bicolor I have collected are from Santa Cecilia light proximoventral calf-spot absent; venter black. [411 m] or La Union [451 m]. .. one hour by horse- with blue, greenish, white. or silver marbling back. from . .. Santa Cecilia. The frogs come Di/1 g" o ~ is , - Phyl/oblllt'S lugllbris is a member of mostly from the hillsides between the two localities. the bicolor group, differing from Phy/{obare~ species I have never seen any colorful Phylloha/ n or Dend w · and P. bicolor in smaller size and in having stripes, bates near Popayan; they do not exist here." Berthold's from P , auro we'li/1 in having ventral marbling, and 1845 record from Popayan (P. m /' /ai1 0r rh ;llll ~ ) may be from P, vi/WIIl,< in smaller size (Table I) and in having only a shippiog point: this supposition is supported by narrower dorsolateral stripes. 1976 FROGS OF THE GENUS PHYLW BATES 25 Description , -i- Measuremems are in Table I. The Turr'a lba (KU 24908) . Li",,,,,: Los Diamantu (FMNH skin is slightly granular on the dor sum . 1!'Ie dorsal 101097-99: KU 24909, 370(8); Unt (MCZ 8(22 ); Sllre1ka tKU 36794-8(1); La C_ illa. Rio R~ laZOn (ASSP 235S8. "IIrface of the limbs. the belly . and the ventral Mlrface 239M); Ta\.aJrwlca (M CZ 9902-(3); Baja Talamanea. BambU of the th i g h~. llIld ~mooIh on the rest of the venter and (CRE 7182-83); Pandora(CRE 7188·89); 4 km W Guapile. \'entra l surface of the limbs. Ma~illary and prema~i l ­ (CRE 850). , lary teeth are pre-e m. T he first finger is longer than PANAMA,_ BoCOJ del Toro: Almiran!e (MCZ 9873-75 ; tile second. The roe s are weble". FMNH 16102, 83468_70, 152016; USNM 118231; KU 80054- 70); 4,8- 5 km W Almirante (KU 94813-21 . In life. the iris is dar k brown . The ground color of 1156)2 - )). 116748- SO; AMSH 81450- 51); 10-12.8 km the entire body is black. A complete gold. yellow . SW Al miran~ (FMS H 67971. 153659. 153756- 57. 153762; yellow-ora nge. or oran ge dor solatera l anpe e ~ t e nds KU 94822- 26); Isla Eoeudo de Vera~u. (KU 115634- 38); from the dorsal base of the thigh along the back.. along lola de Colon (KU 94827- 30. 116753): Isla Bastimento. (KU \14831 _34): Cayo Nanc)' (KU \14835); Cayo de Agua the outer border of the upper eyelid. and along the (KU 94836_38. 116751); Isla Pope (KU 94852, 116752); canthu s rostralis to the tip nf the snout. where it meets Id a San Cristobal (KU 94853_ 54); Peninsula Valiente. its fellow . A go ld. cream. whitish . or bluish stripe Bluefield. (KU 94839-50); mainland NW lola sem Hill e ~ te n(h from the arm in...:rtion along the upper lip. (KU 94851). CIlltHt: CeITO Santa Rowo (FMS H 67970). end ing ventral 10 the eye. The dooal '>Urf.ce of the CIl!'arl!'d and srai'll!'d marl!',ial . _JMS 1683. lim bo. bears extensive gold. yellow , or yello w-green marbling . The ventral surface of the limb" the belly . Phyl/oh/lll"J ,';I/ptC imen from Turrial ba. Costa Riea 189l. lNlldrobarn li...:Torias VillalllJ Cope: 340. I)'pc (Taylor 1952:b41 . 643). had the Mad. ground color locality: Blleno, "'re., IPunlaren...) Co. ta covered by a wa,h of pink ish on the dorsal surface of Ri,a . the thigh. of yellowish on the throat. and of yclhlW. 1900-1902. Hylaplr>ia tinctor;" ( p;>rl). Gunther: 207. green on the belly, 1931. Phyilobulr J IlIgwh';J (part), Dunn: 392. 1942. Dl!'IIdrobatr J linc/OTias Q"'anu (part). Lau ~ nl In pre-ervat ive. the ground color is black: the ,;.tripes u and marbling arc whue or gray. 1952. Dr"d ,oba/r. rilK"'orius villa/US . Tayloe-: 5&2 . 8 rrt' di"~ coli , - 1lIe ca ll is a " rasping em. lasting Taylor u.id Cope'. lype o;ould lMJI be found at four 1Usix 'c.'Collds. pau-e . repeat " (Savage 1% 8). lhe USNM Re!m iorl.lhip.•, - Savalle ( 1968) co nsidered P . viI· rse Piryllo!>S; dorsum is smaller (Table I). usually has narrow er stripes than black. some times with broken yellow medi an dorsa l P. vil/IIIUJ, and has gold. yellow. or yello w-green stripe; complete gold. ora nge, or red-o range dorso­ mottling on the limbs. instead of blue-green moultn g later al stripe: wh ite or pale blue -green ventrolateral ;1$ in P. vil/ulUJ (Savage (968) . P. /ugub,is and P . stripe ; liglll proximoventral calf-spot absent; venter "iflut us resemble P. aU' Olaf!";O of the C ltooeo, but black with while or pale blue-green marbling_ are separated from it by a di,;.tributiona J l!ap in eastern DiognOJiJ. -Ph" l/obat('J ";UG'UJ i. a mem ber of Panama. P. OU' OI,"" io. P. lugubriJ, and P . "itlattU the bico!o' group , differ ing from P. lugub' ;J in larger may be conspecific, but it is best to reco gnize the size (Table I) and in having broader stripes. from P. three as distinct spec ies until more data are gathered , auro taenia in hav ing vent ral marbli ng, and from Noml!'nduIU'I!' . -P. /u~ubris is not the DOldm/l(JII!'J Ph,.Uoba tl!'J species and P. bicolor in smaller siz e /ugubriJ of Cochran and Go in (1970) ; that spec ies is and in having stripes. D('ndmba/('J ITI1t1COIUJ (Cope 1861); see SavlIl!e ( 1968: Dt sc,ipt;01I. _Measurement s arc in Table 1. The 765) . skin is slightly granular on the do~m , the dorsal sur­ Di sr,ibUli01/ und habilUI (Fi" . 13).-Elevation rec­ face of the limbs. the belly , and the ventral surface of ords are from sea level to 650 m (the type locality is the thighs, and smooth on tile rest of the venter and higher. but see Savage 1968). P. /ugub,iJ inhabit, the ventral surface of tile limbs. Ma>:illary and prema>:ilIary forests of the Car ibbean drainage of Costa Rica and teeth arc present. The first finger is longe r than the western Panama. seco nd. The tees are webless In life. the dtmum is black , and somenmes bean; a Ltxa/iliu.-COSTA RICA. HUN ;': juno;tion Rim Sa· bro ken yellow media n dorsal stripe (Front ispiece I). rapiqu' ""d Puerto Viejo (CRE 65. 63_69); Fine. La Selva A complete gold, orange. or red·orange dorsol atera l ncar Puerto Viejo (CRE 512); 16 km N Puerto Viejo (MCZ stripe extends from the dorsa l base of the thigh along 75323 ). Can agD: Turrialba (FMSH 174520; KU 24907, 30414. 3~ g - 89 , 370t7; CRE )2 ); HacOenda La f1orene;" the back . along the outer borde r of the upper eyelid, near TurTialba (CRE 7]95); Morehouse Finea. 6.4 km 5 and along lIM: canthus ~tralis to the tip of the SllOut. BUUET/.V OF THE NATURAL HI STORY MUSEUM OF WS A....'GELES COU/l.TY /1.'0.27

...here it meet~ its fc:llo.....The side of the body i~ black.. mains §(,paratcd from her, fa.:ing her. D Tnovis .... it· A ....hne 1>Iripc extelllb from the arm in...:nion along nc.scd ~i J; (Wi~it ions ; in t 'o of these oviposhioes. the upper lip. ending ventral to the eye . The dorsal the male left and did not atch the female: in one. surface of the hmbs bean numcrou~ blue-green dol~, the male and female remained J to 6 mm apan: in almo st obscuring the black. ground color. The venter one , their noses tOliChed; and in t....o. they remained and the ventral surface of the limbs are black with apart, exccpe that, commencing 5 min after the start white or pale blue -green marb ling . extending onto of o viposition and la ~ ti nE I min, the male assumed tile side of the bod)' as a ventrolate ral stripe from the the Type 3 posture: he lay on top of the female. almost groin to the arm insertion . at a 90"angle to her bo4y .... ith his hind limh, projecting In pre§('rvlltive, tile color i. entirely black, except off to one side, and rubbed his belly again,t her back. for tile dooolatel'31 and upper lip stripes. ...-hich are The female continued to la)' eggs .. bile the male lay on white , and the ventral marbling lind the dors.il "lIrface her. The male' s cloaca did not approach eilher the e~ s of thc limbs, which arc PUfllli.h gl'3). or the female's cloaca . This behlvior. ",'hich resembles Brt'tdin Kroll. -P. vittallU his t.....o k i nd ~ of ~ing that of Culo.Jlnhu5 pul"'! visil. In four breeding cycles that July and August ITable 4). A«ording 10 the eRE Travis ....itnessed, the male al....ays visited the egg, eata logLK' . one of the~ Iot ~ (e RE 903 ) came from twice on the day of oviposition. His fU"Sl visit began small side pools. hut A. StlllTCtt. the collector , told immediately after oviposition. and lasted 5 to 65 min. me the ladpoles came from an adult"s back. and were After an inter·.a! of ahoot 30 min, he at:ain loat on the aljo...ed to grow before pn'M"rvalion. Two other lots eggs for 60 to 90 min. The male then desened the (e RE 7234 and 3152. in Stagcs 25. 28. alll.! 37. taken eggs for one ....eek , During tile second ...eel; after in Cosla Rica in June) an: not from tllc backs of adults ovipositi"n. the male visited lhe eggs once a day and A. Starre tt told me he found tadpoles of this species sat on them (perhaps to I!\\\isten tbem) for 5 min: the in ...-ater in tree holes and fallen palm petioles, but third week, he visited the ej,\gsone to three times a day. nol in streams or side pools. Polder .... itnessed tadpole -carrying once . Polder 's Unlike tadpoles of Dt'nJrobaln. tadpoles of P. male .....t on the jelly·m"-,, and made stamping move­ •·;t/atu s iIl'C IlOt aggressive to"ard each other (Polder ments; all 13 larvee ...nggled onto the mak's back. 1976). 11lc maar carried the larvae three day's and then re­ Meawremcnts Me in Tabk J_ Ieae hi~ back. section in the P . /ugubri5 account. above). Travis .... itnessed tadpolc-carr)"ing rwke. On the firsl Lif~ hi5101",.. - Polder ( 1976) .... itnessed intra'p«ifrc occasion , Travis's male took up his tildpoles on lhe a~~re"i\'e behavior (jumping on the back and prc" inE t....emy -third day after u~ipl.,it i\\n (one .. eek after hatch­ down) in P, vitnnus, ing). The male apparently stimulated the tadpoles to In captive animals (Polder 1976: D, Travis.I'","'ol/ll! wriggle \\nl\\ his back by shaking his posterior end in ('omnumif'tllj"n). courts hip lasts from one to seven three or four vhort bUNs (four or fi~ e s ha~es per burst). day s before ovipovition . COIIn, hip includc ~ calling Tadp .le uptake la,ted 10 min. He carried thc: six tad­ by the male, circ ling, , haking of rbe l imb ~ . jumping poles in '....0 "ages; in thc: first stage , he carried four on and , iuing on tllc eouni ng partner (done b). either tadpole, and retained tbem 1w0 days. After releasing ~ x). and puNling (lhe female follo....s lhe male). The them into a , mall pond in the terrancm . he returned the mak choo...:s tho: site of o..ipo<>ilion (u,u all) on a bro­ same day' to thc: leaf nil. picked up ,he remaining t...-o mcliad leaf. -omc:times on a fern k af. r.m:l) on tbe floor tadpoles. and released Ihc:m the:next da)'. On tile SC'OOnd of the terrarium) and attrXh tbe female 10 the ~ i t e by O«llSion. the: male 100._ up his tadpoles on thc: seven­ calling. Iccmh day after oviposinon (day of hatching unknown). There is no amplexus. The o"ipositing posture is He again carried tho:tildp"le, 10 tile pond in t...o stages Type 2 (re"'mbling thaI of tllc D. lilll'l"ri". group); (four tadpulcs carried tW\\days. the remaining four tad­ while the female lays the egp. the male u,u ally rc- p>les carr ied one day). /97{, FROGSOF THE GEI'WS PHYLLOBATES 27

One of Polder's female s laid egp everyone 10 two plete light lateral stripe (Fig . f B: inco mplete in P. I>< ceh over a three-mo nth period (eac h of Travis' s IWO ~ Jpi"owi. Fig. IA). fromP ./t'm" rafis andP .zupuro in females laid eglls every twc WC'Ch over a two-month having ill light med iall ~l §lIipe iIInd ill smaner size. period) . Keown c1ulCh sue is seven 10 21. In ("a",ivity. and from P. lricolor in having while bones ( ~ ll in P. lhe period from fenilizalion 10 hatching is 13 10 17 /firolor) and in smaller size (Tab le I). days: lhe period from deposition 10 metamorphosis is Dt'scriptirm. -c-Measuremems arc in T ablc I (the 40 days at 22 to 25°C. holotypc is a 21.0 mm female) . The skin is smoo th or Stomach co-runts (o ne spccimenl.-From C RE slig htly granular on the do rsum (e:tCcpl o n the poste rior 7235 : H YM ENOPTER A : Fc rrnicid ae: Was",a""ia back. where it is tuberculat) and smooth on the limbs aUrap,ln Clala ( Ro ge r) (I). Smilnislruma sp. ( 4 ). and the verner. Mu illary and premuillar)' leelh are Dolichodt'rus ~p . ( I l. Paratrt'china "p. (2 ). ROKu ia prtstnl. The first finge r i ~ longer than the SC'Colld . sp. (2). Phd dolt' so. ( I): COLEOPTERA: fragments; Basal webbing occ urs between roes ]I·lII -IV. DIPT ERA: MyeelOph ilidae ?: fragme nts; COL LEM ­ The co lor in life is unknown , In preservative. the dot­ BOLA: Podur idae"!(I). sum is dark brown with a co mplete gray or wh ile me­ DiSt'ust's. _Polder ( 1976) h s discu s-ed d i-ea-es dia n dorsal smpe. eapanding an teriotly 10 COvtT the of Cllpli\.-cP. ,·illa/us. entire donal surface of the l>/lOUl; the l>/lOUt sometimes Distribution and habitat (Fig . 13).-Elention rec· bears dark bro wn ~poIs. The side of tbe bod) is dart ords arc from ]0 10 70 m. P. "illa/us inhabits the wet brow n, A com plete gray or while lateral stripe extends forests of the Golfo Dulce region on rbc Pacific coast from the groin to the upper eyelid , along the o uter of Costa Rica. border o f tbe upper eyel id. and along the cant hus roo stra li~ 10 the tip of the soo ul. " here it mttl~ ib fellow Localitit's.-eosTA RICA . "'",ur"." RincOn ok o.a A gra~' or ...hite ventrolateral Slripe ex tends from the (MCZ 75322: CRE 705. 899. 3076. 3114. 6391. 72341: groill 10 tbe f~limb in-ertion. alxM: which it uniles Rio 1'0:...... """". 7.2 km S RinOOn de (CRE 7235): 4.5 o.a with lhe upper lip slripe: tbe latter jo ins the canthal km W RincOn de Osa(KU 102161-6)); Camp Seanlc. 6.4 km ~p Rio Rin" ,n (e RE 757): Quehrada Bnruca, Palmar stripe at the lip o f the snout. The dorsal surface of the No,-u, (KU 93924): 22 Ion EPalmar Norte (KU f>5296 _) OH ): forelimb usually is gray or white ; the anterior and pes­ Rio 7..ap-'le. 8 km E Palmar None (KU 93925_30. 93934- 3.5): lerior surfaces o f lhe fore limb bear datk brown loegi­ Botnoo Ain:s (CO(><' 1$93). tudrnal stripes. The donal surfece of the hind limb bears CktJrt'd and slai",d ",utt'rial. -JMS 1681-82. altemaling gra)' and bro"' D bands: the lhi~ h bears a ...hite o l>lonll pro:timodors.al ~ptJl and loOmC: l ime~ { a~ in the hololypel a while l on~itu din al po, tt' rodo r~al , tripe . /emormall "hile spols . Figu res 5D. 7A-B In a p;eralype (Fig. 7B) from Salvia' . Ecu ador. the venIer and the ventral surf"cc of lhe limbs "re white. except for a few brown spots on tbe throat. on rbe sides 1921. PIt)'lIob<.ltt. <'een bro...n and D..ji"i/io"._Teeth present: toes basally webbed: while . do rsum (in prcservanve j da rk with light med ian dorsal Tadpoles. - One ku ha, been taken from the back of ' tripe: com plete light lateral 'tripe: light ventro lateral an adult male in Peru in April (Tah le 4). The bead-body stripe; Iighl pro~ imo,tot ral calf.spot abo;enl: venter dark of each ladpo le is ':U mm long: the lail i ~ damag ed with light marbling or ligtll with datk marbling. \I.·hell caf-(urtd among maidtnluir fem s on the ban" of Diuf(nosis. -Ph-, ' ffoha l ~ s "Il/hon,vi i, a member of the Quet>rada AngO'olOra. Pow Azul. Peni. lhe male rbc /t'm"roli s group. differing from P. 1x,,,lm!:t'r; in ca rried eight to 12 tadpolc:~ in a dark. fluid -filled "blis_ having a light media n do rsal , tripe, [ro m P. opino"ai in tcr" on his posterior do rsum (B. Barnell. letter of 25 having a light med ian do rsal , tripe and in having a COIJ1· September 1931 to H,W. Parker at B ~I) : since the tad- BULLETIN OF THE NATURAL HIS TOR Y M USEUM OF LOS A/liGELES COUNTY No.27 pules "were free in it. not in separate pits." the "blis­ marbling; l'omp!ele yelh'" or red lateral 'tripe; light ter" was not an unhatched egg O1a". Thi, "hlister" i, proximovcmra! calf -spot absent: venier white with unknown in other de ndroba ti,h , which <'arry their lad­ black marb ling or black with white marbling. pole, without a proleclive ,t rU<:lure. Dir;an horder (BM 1931.10.7.1); Gramad.1, Hacienda Cas;las ( s i ~ h l record by g, Barnell. leuer to H.W Parker (I. Cabrera. notes al USN M), the lateral «npe and .1 BM) the dorsal thigh markings were red. The venter wa, CI/'(l1'<,d IItld s/uil/"" mmuia/. _AMNH 16416, 16478 gree nish or white wilh bla e ~ ma r ~i np. In preservalive , Ihe dorsum is light brown with dark brown spots and marb ling. The side of the body is dark Phyllobll/<'S houlcl/g.-ri (Barbour 19(9) brown . A <:omplele white lateral stripe exte nds from Marbled Poison-arrow Frog the groin to the upper eyelid , A white stripe exte nds Figures I B, 5E, 7E from the forelimb insertion along the upper lip to the tip of the snout, where it meeh ils fellow . In most 1905 Prosthnapi.l femo1'alis BarhoU': 101, 22 ,)'nlype" specimens, the dors al surface of the limbs i, orangish MCZ 2422; USNM 52406, 118132-33; BM brown with broken dark brown transverse bands. In 1947.2.1 3.92_ 93; UMMZ 48070, h la Gorgona. some specimens. the do rsal surface of the limb, is rNari~o, ] Colombia dark brown with narrow while transverse bands or an 1909. Pro.

both cui".. p"uem~ occur; in ...:>fT"Ie of lhe.e 'P"imen~. Ecuador . I . Cabfe~ found specimens in weI ~in forest the veeeer i, .. hile ..ith ,mall clol.cJ~ set dllfl t.u.. n on the lo wer Rio Calima. Colombia: lhe)' j umped ~pOb . The: throar ma~ be pmiominamJ~ "hite IFi~ rapidl)' and "ere difficult 10 seize. 7E) or predominantly dar l bro" n: it u,pally' h.a ~ a white median longitudinal ' lrip;:. In lhe Imbili .,;:rie~. u>c"lit;t's,-COLOMBlA. S ari ii ,,: h la G"r~na (MCZ the postcrove mral surface of the thigh is darl brown 2422; UMMZ48070; IlM 1'147.2.13,92_93. 1926.1.20.123_ 125: USNM 5241lt>. 118232_33. 145248-3 00: AM."'H 7494, ~ pce i mc n s 'C r ic~. with while does: in mo-r of lhis the 50970-72): Soolh G"'ion. (AMNH 62894_9M) : GorJ(lnilla. re-t of lhe ventral surface of the lhigh i, while Wilh a penin'ulo of 1,1a Gcqona. i", JalCd at h i~h lide IAM1s'H darl brown spots. bomclime, mluced in number. e.g .. b2892 -93): Imbili. Rio Mira (USS M 147431_50): La F i ~ . 7El or marbling . In one specimen from Imbili and Gualac""a (LAOt blO34-H. stoss» JUJlin(LACM 60973. 61036-42), Vlrlu: Camp CanOn de Colombia. lo..er Rio another from La Guay acana. the ventral ,urfa..-e of the Cali.... CUS" M 149723-2 41; R"" kd cller ~ion Viroior. t h i~ h undulalin~ is dan t.uwn .. ith t....o "hite Iongitu· f'.dd SIal..... Rio RajlOJO (US" M 1513'79: LAO! bl0431: diNI .lripe ~ CMe ..... (B ISH u:m). Tu rc m e n b arc in Table .t (GOV 9231. 9391-92) . l'id .int:1uI: Rio Blanco Ileac tnOUlh Savage ( 1968:757) placed P . /x'ult' tI!:..,i in the genu, of Rio Yambi CGO V 9235_3b): N . ",, ~ a h l o (GOV 9227); Cotostethu s, Hc considered il close ly rcla led 10 C. Rio Silanchi (GOV 9599-%00); 18 km W Samo D"min~'<) de nub;""I" (Dunn 1924). and ,a id (IX1 sed on unpubli,hed 10' Colorado,. on rood 10 Cl>one (GOV 9232). ea relti: P:uamha(BM 98,3,1.1- 2) nO le ~ by E. R. Dunn ) lhal the nHlulhpans of lhoc lad· Clrllr..d and s/(li"rJ ntlltt't'ial.-A~t ~ H 7494; USI'< M pole, of P. boll/fIl lI..,i are umbelliform. resembling 1 4 52~3; LA CM b0973: from Tandapi: KU 111529. tho>;: of the ladpole, of C. ""bieola . Tadpo le' (U S ~ M 145248) from lhe bad of an adult ropotype of P. fmu /ulg..,i. however. have mou thparh ~ i m il a r to Pll.,'lIobaln ~.pinfJSQi Funlltc:>U--e r 19.56 those of ot her spccie ~ of nnHobatt'3.' lhe moulhparts TUTqllOi.e-bellied Poi'OO-al"l'O\>- frot: are not umbcUiform (Fig. 12AI. figures IA. 5F. 7F MOIt'riol of " nerrlai" Jla lu• . _ S pce i me n~ from Tandapi. Ecuador. 1460 m. on lhe ....eslern ~k>pe of the Ande. ea.lWII lhcar.t of Santo Domingo de 100Colo­ I ~ I' I.yllob."t'J rs.plllOJtli Funkllooo"Ct: 7b- 78. pi"", I. fi~ •. 2_3. 110101)'"" , CAS·SU 1cr.'i77. Hacienda B· rados IKU 111529-61. 120f>65- 68 ). belong 10 lhe pi""'"'-. 9 km W Sant" Domingo de 10> Col<>nodos. f/!'moro/i.1 group; lhey resemble Colombian spec imens Pkhineha. Ecuador. :105 m. of P. bm,/nl llt'fi. except lhal ihe lateral stripe is net quile complete. the ventral marbling is yello.... in life (notes by J. D. Lynch al KU). and the dorsum ha) Iighl Drfi nili,m . - TeClh present: toes basally webbed; spols . Some specimens IGO V 9 148 - 50. 9229-30) dorwm red; incomplete tUTqllOi.,;: lateral slripe : light from the Chocoan 1o.... 1ands of Ecuador (Rio Blanco and pro~im",..emral ea lf·~pOI absent; venter darl bro.. n San ~Ii~uel de I<:>s CoJorado') resemble the specimens with I U Tq uoi ~ marbl ing or spots. from l anda-pi. Pn;M.>on,ally. I h3ve referred a1lthe~ DillXno.i3 , --Ph.l· II0b0I~J ~3piNJJlli is a member of specimens 10 P. boul~ng~ri. but I have uc1udcd them thefrmorolit group. differing from P. onthoto}'i in !iKk· from lhe lmlp and Ioeal it~· Ii"-. ing II hghl median doNllolripe and in having afl eccm­ SlOmadl ronl~nlS (t....o specimens).-From US....M plctc l i ~h l lateral stripe. from P. boul~n~ri in hlo\ing a 145253 : HYME:-l~n"psis red don.um in life (brovo' n in P. bo,.l~n8~ri) and in hav· sp. (.5). Ph ~ id o /~ sp- (1). Iridom.rrmu sp. ( I). ing an incom plete l i~ht la lera l stripe (Fig. IA; the laler· M)'coctpurus . mithi (Forel) (I). My martd ae ( I): al 'lripe i~ complele in P. Un/hOlI)'i and P. b<, ul~ngeri. COL EOPT ERA (I ). Fig. I B). from P. f.." wr" li" in having a red dot'l; om in From KU 111529: HEMI PTERA : Aphidid ac (3) life ( b la~ l or brown in P. fn ,wraliJ). in having an in­ Nom~ndal..r~. _ 1 des ignale BM 1947 .2. 13.93. a com plete l i~ h t 13teraJ ~t ri pe (complctc in P, Irnwmlis). 20 .0 mm fe male from Gorgona Island. Colombia. and in smalle r siu (Table I) , fromP. Irico/or in liKli ng collected belllleen 19 June and 2 July 1904 by W ,W . a light median dorsal smpe, in hll'o'ing an in,,:omplele Bro...e, Jr.• as the IeclOl)pe of ProslN rapis boulmxrri light lalcnl wipe (com plo:te in P, /rico/or). in having """"'" 1909 ..'hire bones (gree n in P. rricolor). and in smalleT size , Diuril'JlllilJtl and IuJmtOl (Fig . 15).-EJevalioo rec­ and from P . zoparo in .maIler size. P. n piltlll

on road to CJxme The first finger is longer than the second. The third (CAS-SU 10577- 83. 10585- 86: USNM 145804); Santo Domingo de I"" COlntadlls (CAS-SU 10375;CFW 4399 al finger is swollen in males and not swollen in females. UMMZ) Basal webtung cc curs between toes II·III-IV. In life, in the type series (Funkhouse r 1956), the dorsum was dark red. The side of the body was black Phyllohat esfemorolis (Boulc nger 1883) with an incomplete turquoise diagonal lateral stripe . Brilliant -rhighe d Poiso n-a rrow Frog The dorsal surface of the thigh was black whh wrquoise Frontispiece 11 and Figures 3F. 8E mark ings. The dorsal surface of the cal f was dar k red. The venter and the ventral surface of the limbs 1883. Proslha. Huallaga River. [Lorero. ] Peru are dark brown. An incomplete light lateral stripe 1920. Phyllohlis. Barbour and Noble: 401. extends from the groin about one-fourth of the distance 1935. Phyl/obbnres femoralis. Dunn, 78. 1970. PhyllobolCs fe mlgeri) and 400 m (east of the Andes). resembles the types of in larger size, fmm P. e,'p illo.I' , and in larger size . and from P. white marhling. Funkhouser ( 195(1:77) did not devig ­ tricolor in lacking a light mediandorsal stripe and in hav­ nute CAS-SU 10340 as a type because of its lllcalily. ing white hones (green in P. trico l"r) , P. jn/lorl/lis I am not certain that CAS·SU 10340 belongs 1\) the close ly resembles P. w pttro in size ~ nd in having a short same species as the ty pes llf P. e.lp illo,\(li.- [ have tars al fold (e xtending along the distal one-third of thc dis­ excluded CAS-SU 10340 from the map and locality tance between thc inner meM arsul tubercle and the heel list. fold). P. femondis differs from P. zaparo in having a Di.'trihll/iOIl W il l /Ii/

(.00...1«juallo!he seccod in P. piC/ILl). in having ba..al reduced 10 small spols. In small juveniks, the ventral toe ...ebbing (...ebbing is absenl in P. picllU).lIlld in hav­ areas lhal were marbled in Jd uhs were unifOllll gray. ing I« th (P. piCfUS asually lads t«lh). In preselVative. yellow, orange, and gold become Descripnon, - Mea'urement' are in Table I . The whhe. The ground color is black or dark bro....n. Rarely d in is granular on the dorsum and the dorsal surface te.g .. in TNHC 36498 from the Rio Pachitea. Peru). of the limbs, and sm\1'l"th un the venter and the ventral the belly is almost immaculate white. The light ventral surface of the limbs. Muillary and p re m a ~ ilI ~ r y teeth marbling never extends onto the throat. except in one are pre\Cn!. 'The fiN fi n~er is l on~er than the second spr;x:imen (KU 14436 1 from Nevan, Peru, in the upper Basal webbing OCCIin between tues II-Ill-IV . Pachirea drainage). Th is specimen resembles mos t In lhe following compo."ite desc ription (ootes by specimens of P. f~m",uljs. excepl Ihat (in peeserva­ J.D. Lynch at KU. W.R. Hey-er and P.A. Silverslone tive) the throa t is marb led with whlre . the belly is aI LACl\.I)of specimens from Santa Cecilia and limon· ",hite ",'ith a dark bro""n transverse band formed of COl:ha. Ecuador. from 5 km north of Pueno Asis and marbling , and the donum is light brown with dad EI Pepino, Colom bia. from Teresinha and Serra. do brmon marbling l'avio. Brasil. and from SaUl Ton ue. French Guiana, 8 rr~di lt g cull. -P. f~-.rQI;s "calls from lbe fOKSl a color llOIed only in specimens from a certain coonU)' floor in the daytime. a loud. slow mllSical trill lasting is follo....ed in parenthe\Cs by the abbreviated name four seconds, repealed at irregular intervals , spaced of that country. In life, the iris was black with a gold usually at least 0 ne minute apart" (O. B, Wake and rin~ around the pupil. brown with red flecks (some R. Etheridge. field n" les al CRE). In Suriname , males Ee .). or bronze (some Ee .). The head and hack were call from the ground or from twigs and leaves up to black or brown (Ee. ). The side of lhe OOdy was black. one meter above the ground; the call is high and pro­ A complete yetlow or go ld lateral sn ipe extended longed. resembling a cricket's chirpi ng (Hoogmoed from the groin to the upper eyelid. along the outer 1971). border of the upper eyeli d. and along the c anthus TuJpoIu (Figs. II C. 12D).-T....o lois hd resemble eac h olher when Masl of the dorsal surface of the limbs was light brown seen from above. or gray ....ith scanered black spots. The proximodorsal D;....us~s . -Polde r (1973) has discussed occurrence surface of the upper arm was yellow or orange (F.G.). lIlld lrealmenl of cramp of the hind limbs lIlld fungal The a,illa ""as ~ello",' 01" OWlge (F.G.) . The lXiIIII')' infections of the eyes in caplivity·. color united ",'ilh three light stripes: ( I) the upper ,"·om r lt cl(1 l " r~. - 1 desig nale B~ I 1947 .2. 14.21. a lip s!ripe; (2) a "ripe on the ventral surface of the 26.0 mm male from Yurimagu as. Huallaga River , upper arm twbere it was yello"" or orange [F.G .)) [Lo re to.] Peru, collected by Dr. Hahnel, as the and forearm (where il ....as .... hire): (3) a ....hne stripe leclOlylX' of Pro.,~,upis f~nr"'(1lij Boulee gcr 1883. on the anterior surface \1' f the upper ann, separated This is the specime n illustrated in the original de­ from the second stripe by a hlack area. and usually S<;rlptiun. meeting the second stripe on the forearm. The p ro~imal DiJI,ihutiof/ und Iwhi l<'l (F!):. 17).-Eltvation rec­ half and the posterior portion of the distal half nf the ords are from 14 to 610 m. P . !;:mo rulis Inhabits the dorsal surface of the thigh were black; the black co kn lowland forests of Guyana, Surinam e . ant! French surrounded a conspicuous oblique yellow or oran~ Guiana and of the Amazon drainage of Colombia, (F.G .) spol on the base of lhe thigh. The spol some­ Ecuador. Peru. and Brasil. I look a specimen al 1745 h times utended along the eeure length of the thigh. in fores! al Saut Tortue. French Guiana. after a rain. Sometimes a small 'P'1I of lhe ~ color lay distal on falk n Ic ~ s on the ground. J_ Lescwe (in lin .) to !he Ihipa-spoI (Fig. 3f). The venter and the ventral found Ihi~ species in open microhabitats in primll')' surface of lhe hind limbs were black. brown (some or secondary forest in rnl'lCh Guiana Ee.). or grll)', and (e' c-epl for the Ih1Oll t) bore ...hite or bluish ...hue marbling. In some specimens. the U>culi/iu.--COW MBIA. "",,,,,,,,,-0: Umbria (FM)\H ventral ground color was ",'hite, and the black was bI789- 1IO); S ~m 1'< Pucn" A. i. (LACM b t044 _49); EI 31 BUUETI."I OF THE ,....ATURAL HISTORY M USEUM OF WS A....'GEUS COW..Tr No. 27 ~no . juraion PasIo-PIleno ....is and MCJC(q·P\M:no A,.. Phy flobalu rricvlur (Boulen~ 1899) r....d. ( L ACM 61031 -)2). C.,.. ~/li : Florcnd l ( U S !" ~l Ptliinta'\Jllal Poison-arro'" Frog 1~72-73); MOftlia (ASSP 2537l}-71). M~I4 : Bl>tfIMo,l,J Fronli. picce II and Figu,," SA . 8F (f"MSH 81810). A_ _ : 7_7.2 km :\>\.' l.c1icil ( LACM 61019 .61033) ECUA DOR.-.\·"po: Santa cecut... t KU 104762_63. 1899b I'ro'llv .api. Iri=1or Boulcngt'r: 4S5-456. four .y n· 1069~S. 109189-90. 109292. 109295, 109297-98. II ISIS­ '~ptl: B:4 1941.2. 14, 16- 19. Porvenir, Boliv.... III. 112310, 122270. 122272. 122275-76. 122280 -8 6); Ecuadnr. ]769 m. Puenn Ore. Rio Aguan,,, (KU 122291); Lirnonco<: h. (LACM 1920 1'".,II,Jb<,ln ITicolo •. Barbourlind Soble; 401. 72629-41. 72676-82) PERO.-Pasco: Chanl in. , IC'ar Quebrad.o Ca..hibo)l lhe /"n ll1ra!is group. differing from all lXher members (AMS H 42981); l.a.jo de Mirmo. moudl Rio /'iapo tAMSH of che group in having green bones . P. lrico!o. fur­ 43183.43415); cceneeoce RiooCnnla~a;and Tapil;ho: tAMSH tber differs from P. anlhr",)'i in larger ,ize (Table I) . 42565. 43249. 43251); 1'IlIi'"" (A",I:-;H 42295. 4J614); from P. boorlengeri in having a lighl median donal connucnct' Rios Tama) 1 .nd Uca.~.li (AM:-;H 43426); Peru-Brasil fron.ic, (AMNH 42058. 42694). 1'.." ,,: L. snipe , from P. e.pinosai in having a Iighl median UniOn. C.mbllya. Rio Huacam. )o (8 M 1907.5.7.8). dorsal stripe. in having a complete light lateral str ipe BRASIL.- I',,,,i: Saatarem, road 10 C""hoeira ok l>-alll.1o (Fig. I B; incomplete in P. t')pilW.wi. Fig. IA), am! (MPEG 744); U' inga ncar Belem (LACM 42321); J"". re· in larger sil e, and from P. f~11I(>T{/lis and P. zaparo Aeang• . Rio Tapajl>" IB, Lutz and Klos' (952); Rio Mapuer. in havin g a light media n dorsal stripe. (AMNH 53334). Am apa: Serra do Navio (LACM 42322). Amazonas: Estir"o do E<.j uador. Rio Javari (MPEG 634- 646); DnuiptiOlI of syntypes. - The snout-ver n length ill Rio Tachi,a ~ ar Leticia (LACM 61011); Tel"l:, inha ncar millimeters of the four symy pes, which are all adults , Lelieia (LACM 60974-75 . 61029-30); Eirunepr:. Rio Jurua is (mea n in parenthe ses): two ma les 20.0- 20.5 a O.3); (B. l Ull and Klo.. 1952); Bcnjamin Conslanl (CAS·SU two female, 22.0-22.5 (22 .3); four specimens 20,0 ­ 11873). 22.5 (2 1.3) . The skin is entire ly smooth. Maxillary GUYA.\lA.-R""'''.."i: '" Acarah) M.....nlain•• W Jl'e'A R;." , (KU 69692·95); KIIYII...-ini landing (AMNH 4m 2); and premaxillary teeth are prese nt. Tbe firsc finger is be ·ct'n upper Ku)u wini and Kas, ikaillU Riw... . uppt r longe r than lhe second. All roes are slight ly fringed E ,qllibo drain.lIe CBM 1936.4,2.7- 8); Onoro Rivt'r and basally ....ebbed; the ....ebbing is best developed (_Unoro o Rivt'f l (AMS H 44644. 44641. H689-91). ,...... IN_ · Akyma(AMSH 13S05-{16); Wi: Kanab<> (AMS H 10407, 13500. When recently prese rved. the dorswn llIld lhe side 23124.39679-80.39687• .3969 1. 391Oll. 70991. 70993-96; of tbe body were black. emblazoned .... ith complete USS"-l 118015); 0\:;0 Moum. in, (flol!"H 26710, 26760); lemo n yello .... lateral and median dorsa l stripes. The Monbllili Cm:k , &..equibo drainagoe (BM 1930.10.10.2-9); base of the upper arm, the lum bar region. tbe postero­ junc.ion MCllIIlII. nd Mauroni Rivers (AM:-;H lll'W5). SURIS AME.-Sor rinam..: Param. ribo (RM:-< H unu O. dorsal surface of the thigh. and che ventral surface of Maro .. ij,,~: PocIOt'gocd<1t'(RMNH uneal); P.loemell tRM:-i/9(0): 2 km W Santi I. abo:!. dark marb ling in three spec imens, and dark with km 88 un Cuenca-Machata road (USKM 1669(1); 4 km S mediu m-sized light spots in lhe fourth specimen (BM Calavina. km 7t on Cuenca.Machala ro. d (USNM 166899). 1947.2.14. 17). CII'C 1 and rounded in modorsal surface of the thigh and another covering laleral aspect. The canthus rostrali s Is rounded. The me axilla and the proximopo sterior and proximodon.al Ioreal reg ion is slightly con cave. The width of lhe surfaces of lhe upper arm. in having me ventrolalcral upper eyelid is slightly jessthan the interorb ital width slripe incomplete (ending midway between the groin Tbe diameter of the e)'C is !ilighdy greater than the and lhe uilJa) in UM MZ 129358 and no! visible in distance from the eye to me l'lOiilril and twice the di­ UMMZ 90370. and in lacking ventra! marb ling. amete r of the tympanum. The posterodo l"§4l port ion In preservalive, rl'lO'>l of the GOV para t) pes resemble of me tympanum is concealed . 'Ibe finger disks ;m lhe holot )pc: . Two s peci~ns (GOV 9194 and 93(4) small relative to tbose of lhnJrobules . The fio.t finger have a promine nl .. hile spot cove ring the axilla and is longer than the seco ed. Tbe fingers are webless. 1be dorsal base of the upper arm. The dors.a1 surface of the larsa l fold is Slm ngl)' bent at its pro ximal end and thigh is either emire ly dark or banded with light and . relati....ely short. extending along the dista l 33 percent dark ~ there is no lighl lhigh-spol. except in GOV 9007 of lhe distance bet..-een the inner metatarsal tubercle (s mall and vesligial) and 9194 and 9304 (small and and the heel fold. The tarsal tubercle is modera tely' prominent). The venlrolateral smpe is complete res­ developed . Basal webbing OI.'CUl)C between loes !I·IlI·IV. tending from the groin to the axilla. where it unites In life (notes by J.D. Lynch at KU ). the iris was with the upper lip stripe) or incom plete , Some speci­ black with a bron ze ring around the pupi l. The dorsum mens have a prominent or vestigial stripe on the canthu s was br ick red ,The side of lhe head and body was rostralis and upper eyelid . Five specimens (GOV 8929, black, The upper lip stripe was gree nish yellow. The 91XI5. 9 190, 9212, 9380) have a vestigial late ral stripe ventrolateral stripe was white. The dorsal surface of adjacent to the edge of the back. In one specimen (GOV the forelimbs was light brown with a few black spots. 9007) . the lateral stripe is com plete and prominent from The axilla was yellow-orange. The dors al surface of the groin to the tip of the snout; this condi tion differs the hind limbs was black with blue -gray mottling. from all otber parat}pe s and from the bolotypc. The The throat and chest were black . The belly and the throat is entirely dark: the belly and lhe ventral surface ventral surface of the limbs were black. monied and of lhe limbs bear light marbling. marbled with pale blue·gr ay. Distribution mul humtat (Fig. 15).-Elevation rec­ In pre~ nl a ti ....e. tlJf, grancjes of lite head and back ords are from about 230 to 1000 m. P. zaparo inhabits are EOkien-brown. separa ted by narrow black inter_ the dense forests of the Pastaza and Xapo drainages of spaces . The side of the head and bod)' is black. A Ecuador , ea st of the Andes; it is terres trial and diurnal .. hire ....entrolateral snipe extends from lhe groin to (lIOfes by J.D. L)'nch at KU). the nilla. uniting donal to the Ioretimb insertion with £ f)-molog)". - This species is named after the Ziparo ue upper lip stripe. " 'hich continues 10 the oostril. indians of ea' tem Ecuador. /976 FROGS OF THE: GV,US PHYUOBAT'ES

pious group sent. The venter and the ventral surface of the limbs an: Iighl .. ilh brown marbling and spots; the ventral marl­ This group includes seven species (Pltylloba/t'S boJiv­ ings are dsscrere in 89-90. btll cont inuous and closel)' iamu, P.i n/,,,ri, P. parv MfMS , P. pr /..rsi. P, pic/..s. p . set in 91. p..lchri""C/..l. P, .•maraxdin",I/. Nommcla/urr , - I designate BM 1947.2 ,13.89 . a Defi'li/lim : snout-vent len glh of adults sma ll lO 26.0 mm female from San Carlos. Bolivia, 1200 m moderat e ( 16.5-3 1.5 0101); toes webtcss: paired elevation . collected by P.O . Simons. as the lectotype stripe s (when present) lateral (Figs. 3A -B. D-E. of ProMherupi.• holilli

Pltyl/oba /es boIi>;a"u.1(Boulenger 19(2) ~

1902 P'o>lh.-rapi. boIi,·;o",.. Bouknger: 397. lhftt .yn. types: BM 1947.2.13.89-90. San Callos. [u Paz,I Phyf{l,b(l/t'S ingf"; (Cochran and Goin 1970) Bolivia, 1200 m; BM 1947,2.13.91. San Er~ 'l o , (La Pal ,] Bolivia.800 m. Brother Nicefom's P oison-arrow Frog 1920 Ph.vl/ohalr. holivian,.., Barbourand Noble: 402. Figures 3C, 9C

1970. lH,.JroI>a~J in~ri Coehnn and Goin: 16_17. plale D"fi"i/ion. - Teeth abse m; roes webless: dorsum 2D- F. hoIoo:ype , CSS M 146ll46. A-no Dc'" Rio pre~n-al ive); dar k (in com plele light laleral stripe; ~ado. C~...u. Colombia. ligh l prcxisnovennal calf-spot abse er; ven ter light ....ith dark marbling and spots. Diognosis.-Phylloba/"J boli.'ionus is a memher of D..fini/ion .-Tee lh present; toes ..-ebless; dorsum tbe piC/..s group. differing from P. ing..ri in lacking darl (in presen-ativt'); stripes absent; li/;ht proximo'o'en­ teeth. in having a comp lele Jigi'll lateral stripe (absenl tral ca lf·spot present; venter dark with Jigi'll marbl ing. in P. inJ("';l, and in lacking a Iighl proximovemral DioK"os;s,-Phyf!o />{I/<'J ingeri is a member of tbc calf- spot. from P , parvul".• in lad ing teeth and in p;ctUJ group. differing fmm all other species of lhe having a complele llght lateral slripe (inc<.> mplcle in gm up in lacking a light l"leml , tripe . and from all spe­ P. pa,..,,,I,,.•). from P. P<'lrni in lacking teeth . from cies of the group except P. P;,·/..J (and rarely. P. per­ P. p ier.... in hocking a light proximo ventral catf-spct. vul.... ) in having a lighl prcx imoveetral ca lf·'p'lI. P. from P. p"khriprc/us in lacki ng a lighr proxifllOdor,;al ing..r; furtber differs from P. boIiv;Urf~ of the hind limbs. lar on lhe don.um and the do~1 surface of the hind and smooIh on the forelimbs. the verner, and lhe ventral limbs. and smoo th on rbe fore limbs. the venter , and the surface of the hind limbs. Ted h an: absen L The fin;l ventral MJrface of the hind limbs. Max illll/)' and pre. finger is longer man tbe second. The toes are webless. maxillary teeth are present. The finot finger i& longer The color in life is unknown , In preservative, lhe dor­ than the second , The IOCS are weblcss. sum and the side of the body are brown. A complete The cllior in life is unknown (C.J. Goin, ill fill. ; white latera l 'lripe extend, from tbc groin to the upper Rdo. Hno. Niceforo Maria. ;n fill.). In preservative. eyelid. along tbe outer borner of the upper eyelid. and stripes are absent. The I) pes are entirety hlack. exce pt sol1lClimes along lhe canthus rostralis to lhe lip of lhe for a white spoe on the axilla and lhe proximodor;.al snout. where il meelS ilS fellolli. A whne stripe extends surface of the upper ann. a small round Of rtttangular from the f.....-e limb insel1ion along lhe uppcT lip. ending .. hite spot 00 lhe proximodorsal sutfece of lhe I hi ~h . ventral 10 the nostril or connnuing around the snout 10 a large .. hile spot on lhe pnJi\irnoventrai surface of fIlCCI il> fellow. 1llt' axilla and lhe proximovenlral and the calf (wmetimes ei\tending half the lenglh of lhe proximoanlerior 'Ulface' of the upper ann an: .. hite. calO . and iII-defi.w ~ra)' marbling on lhe belly (nolo­ Light malbJing covers lhe anterior . dorsal. and posterior I) pel or on the emire ventcr and the venlral surface surfaces of the thigh. On the posterior surface of rbc of the limt>s(paralypes). thigh. the marbling forms a lighl longitudinal undulat­ Rr //I/;oll.hip:> ,_P , ;" Ked is closely related to P. ing stripe in 89 -90. Thigh-sp ots and calf-spots are ab- ptctus. 36 BULLETIN OF THE NATURAL HI STORY MUSEUM OF LOS ANGELES COUNTY No. 27 Di~trihuri"'l and hahira' (Fig. 16),-The only elc­ P. espinosai in lacking toe webbing (the toes are vation record is 4()') m. P. ingeri is know n only from basally webbed in P . tf,Ipuro and P. ".,pino.wi). the type locality. in the Amazon drainll~c of Co­ Destr/plion. -Measurements arc in Table L The lombia. skin is strongly ~ranular on the dorsum and the dorsal surface of the hind limbs. and smooth on the forelimbs , uX'/Ililies.-eoLOMBlA , CllI/umi: Aserrfo. 20 km from the venter. and the ventral surface of the hind limbs. jUlIClion Rio. »c!IC.do and Oneguaza. 5 km from righl bank Maxillary and premaxillary teeth are present. The first Rio Pescado (US!' M 146846_49: lLS 43) finger is longer than the second. The toes arc webless. In life, in ,pecimens from Santa Cecilia and Limon­ eucha, Ecuador (notes by J.D. Lynch at KU and W.R Heyer at LACM) , the d",.,mm was dark red (only the Ph.l'llohmes parvuius (Boulcnger 1882) numerous. c1o>elyset granules were red: between them Ruby Poison-arrow Frog the skin was black). The ground color of the side of Frontispiece I and Figures SB. 70 the body was black. An incomplete Iighl blue lateral stripe extended from the groin one -half to three-fou rths 1882 D~nd",hate ., /",,,'ula,, goulenger: 145-146. plale of the groin-axilla distance . A li~ht blue stripe extended 11. fig. 6. lhree 'ynlypes: BM 1947,2.30.89-90. from the forelimb insertion along the upper lip. ending Snrap,u, (Pastaza.j Ecuador; BM 80.12 .5.195. ventral to the eye or nostril. A yellow axillary spot Canclos, tr-.'lal"] Ecuador. extended onto the proximodorsal surface of the upper arm. A small round or oolong yellow SPOl adorned the Dcftnilio/l. -Tccth present; toes wcbless: dorsum proximodorsal surface of the thigh , The venter and red; incomplete blue lateral stripe: light proximovenrral the ventral surface of the limos were marbled light calf-spot usually absent; venter black with blue mar­ blue and black: the marbling extended onto the side bling or blue with black marbling. of the body . The blue and the black were 01 equal Diagllosis.-Phyllob(l/<,s purvu/us is a member of extent in some specimens; in others, the blue formed the pictus group. differ ing from all other species of lhe ground color ,Juvenile specimens from Santa Cecilia the group in having an incomple te light lateral stripe were bluish black on the dorsum, and had less blue on (often disap pearing in preservative: the other species the Venter than did adults; the smallest juveniles lacked have a complete light lateral stripe. except P. illgeri , yellow axillary and thigh-spots which lacks stripes), P . ptlfVU/US further differs from In life. a topotype from Sarayacu. Ecuador (Frontis­ P. pctersi, P. pictus, P. pulchripecua. and P. smarug ­ piece J. and notes by J.D. Lynch at KUj, resembled dinu.,' in having a red dorsum in life, fromP. !>o/iv/a"us, specimens from Santa Cecilia, except that the proxi­ P. pulchripectus. and most spo;:cimens nf P. pi<'tU.I' modorsal surface of the upper ann was light blue in­ in having teeth. from P. ingrri and P. picsus in usually stead of yellow, and the proximodorsal thigh-spot lacking a light prcximoventral calf-spot. and from was absem. The types of P. parvulus from Sarayacu P• .v"urugdinu.< in having ventral marbling in life. and Cane!os also lack thigh-spots _ Two specimens (LACM 72646 and 72655) of a series In preservat ive. the ground color is black or dark of 26 P. parvulu.,' from Limoncocha have a light area. brown. The axillary and thigh-spots usually are dis­ yellow in life (notes by W,R. Heyer at LACM). on tinct and white. The lateral stripe usually disappears the proximoventral surface of the calf. Un like the (e.g .. it is visible in only six of 26 preserved LACM distinct. white. rectangular or elli ptical calf-spot uf specimens from Limoncocha, Ecuador). The ventral preserved specimens of P. piCIIIS, the calf-spot in these marbling is indistinct and gray, and sometimes dis ­ two specimens is indistinct. gray. and irregularly appears (i.e .. the venter becomes entirely black or shaped. Besides usually lacking a calf-spot. specimens dark brown). of P. f}(lrvul us from Lsmoncocha differ from sympatric Nome ndaluT<, .- 1 designate BM 1947.2.30.89, a specimens of P. pic/us as follows (the cond ition in 22.5 mm male from Sarayacu, Ecuador , collected by P. piclUJ is in parentheses): the lateral stripe is in­ Mr. Buckley, as the lectotype of Dendrobates purvulus complete. often disappearing in preservative (the Roulenger 1882. lateral stripe is complete and dist inct in preservative); Dinrlbutian and habila! (Fig. 16).- Elcvation rec­ the dorsum is red in life (black or brown): the dorsum ords arc from 330 to 1000 rn. P. parvutus inhabits the is strongly granular (slightly granular); the first finger forests of the Amazon drainage of Colombia. Ecuador, is longer than the second (about equal); maxillary and and Peru, east of the Andes . Specimens from Santa premaxillary teeth are present (absent). In life , P. Cecilia were taken on the ground in forest (KU 111507­ punu/u.,' resembles a sympateic species, P. zaparo, 14) and at night on the ground in a marsh (JDL 1226 at and an allopatric species. P. espinosai. which also KU); one specimen from Sarayacu was taken by day are red. P. ptlrv ulus differs from P. taparo in lacking next to a small creek in forest (KU 120664; notes by toe webbing and in smaller size (Table I). and from 1.0. Lynch at KU). /976 FROGS OF THE GElWS PHYLLO BATES 37

Loculi/it's . -COLOMBIA. P..lI,_.nI: Sm l. R...... dl: 1m J«-ted at La UniOn. Rio Huecamayo. Pueo. by .\tr. Kofme•. Rio Guame. tO I SOb25-~8) . Ockcnden. Lo~<> ECUAOOR._Sapo: M en ,UMMZ 9(371); (GO Y Dtjini/ion .-Teelh present; toes webje ss: dorsum 8921); S .Iope Sumac<> (GOv 9382); ~ l m W I'I>crto Nap<>. Rio Na p<> (JAP ~658); Rio Cocapioo.:-.cap<> drainage \CAS·SU brown. sometimes with small black ~POIS; com plete 10337); Santa Cecilia (KU 104773, 106953-54 , 109157_88. yellow or yello w-gree n lateral stripe; lighl pro ximo­ 111507- 14, 11l684, 122225-51; l.ACM 60980. 61012-14); ventral caJr' SPOI absent: venter bluc with black mar­ Limoncocha (LACM 7264/1_71; KU 99 t t8): Laso Asrio bling or spots (WED 35728); Puerto Lim . Rio A~uarico IKU 122252-(8); Puerto O~ , Rio Agu;Uico (KU 122269). Pas/a.,.: Sarayacu DiuIlIWl i.I.-Phylfobtl/t l petersi is a member of rhe (BM 1947.2,30.89 - 90; KU 120664); Cando. (BM pic/"s group. differing IromP . boli>"illnUS and P . pul­ 80.12.5 ,195) ; headwalers Rio Capahuar i (GOV 8925); chriPt'CI.... in having teeth. from P. inilrri in having a head"'a,,," Rio Bobona:la tGOV &92 &; lA P 2124. 2ISS); complete light latera l ~ l r i pe (absent in P. inxrri) and in ~ lckM ~~ . Rio Bobonaza (GOV 8933); ron­ lack ing a li$ht prcximosentral calf-spot. rrom P. POl" Ouc:nce Rio> Sh}'OftI and Conambo (GOV 900b); Rio Co­ nambo (ZSBS 221t962; GOV 9211; JAP 31166); Rio RUluno. "ulll:l in having a complete lighc lateral -.lripe (incom· Bobonua draina.., (GOV 9191-93; JAP 3909); 0.& lm I\lO plete in P. pon'ulus) and in having a brown (!,(mum in Puyo (lAP 2(64). .U " TD n,, · S '''fli,,~..: 3.2 km E Su.:ua. liCe (red in P. pon·ulus) . from P. piClIl:l in Il;,ving teet h 'rai l from Suo:;ua 10 Rio UI"'IlO (lA P 2208. 2229); Millal (u ~u a ll )' lack ing inP. pic/us ) and in lacking a light prox· (GO Y 9387); Rio Uu_hin N Arapirolo (GOV 8916) PERO.- l.o,.,./o: Peha_ (BM 1915.3.9.12; CAS·SU 6332). imoventral calf-s pot. and from P. smaru/:dillus in hav­ CluJrt'd and .lllI in ",J "'(IIl'ri"I , - LACM f!0980; KU int; ventral marblin g . 126309 Description of '10[01.1'(1<'.- The holu t),f'C is an adult remale. 28,0 mm snout-ve nt length . The skin is moder. alely to ~Irong l)' granular on tbe dors um and the lloNiI PhylloboUJ pt'tt'ni new s poc i e~ Peruvian Poison-arrow Frog surfece of the hind limbs. and smooth on the rorelimbs. the venter. and lhe vencra! surface of lhe hind limbs . Pronnspjece II and Fi$ures 6F. 9B Maxillary and prema,illary teeth are prese nt; vomerine teeth are absent. The tip of the: snout is )I,/blruocale 1970. Dr..JTDbolt'l Ir;,-;/IO/". (I"'n). Cochran and Goin: in don-a l aspect and rounded in laleral aspect. The 16 (1'IOl lhe p1ale; onl) MCZ 24443 is P. pr/rrsi). cant hus rosrralis is rounded . The joreal region is slighll)' concave. The width of the upper cyelld is _Iightl)' Ies~ HolOf)'pl'. -UST\M 166763. co llected at Santa Isabel than the intero rbital widih . The diame ter or the eye is (a village on the Rio Xevati. a tributary or the Rio Pi­ 1.2 times the distan ce frum the eye 10 thc nost ril and Chis. Pachitca drainage. 3.5km SE Puerto Bermudez. 80 twice the diameter of the tympanum. The posterodorsul km ENE Oxapampa: the specimen Wall taken slighlly portion of the tympanum i.~ concealed. The ringer di sks upriwr Irom tho:village. accordin g to A. C. There -e n. are small rela tive to lhoS(' or Dt'I/dmbtl/t's, T he first in fill.), Departamento de Pa'lCO . Peru. 4.58 m. on 16 fiegcr i ~ kmge r than lhe second . The fingers are web­ lu ly l%ll. til G. R. Xconan . less. The canal rold is ~l ighlly curved. e'lendin$ along ParatyPt'J (4 1 aduh~ and nine tadpoles, all from lhe di~ta l b4 percent or the disrarce bec...'ee n the inner Pc:ru).-KU 144348-50, 10popatal)pes. collected by mc:tatan.altuberde and the heel fold . The rarsafrubercle D,B. Ekkens: ~CM 924 14 and KU 144344-46. col­ is absent. The toes are weble ss . leered at xevau. Pa ~ c o . by D . B. Ekkens; AMNH In life teeter slide by A.C . Thoresen; venter not 43008. 43011 . 43016. 43026. 43034. collected at visible ), the dorsum was Iighc brown. The side or the Monte Alegre. Rio Pachitea, HuimU('(). by H. Ba"lcr; body was hlack . A co mplete )'ellow late ral stripe AMNH 42 179. 42 505~07 . 42.546, collected at Achin a­ extended from the groi n to lhe upper c)'e lid, along mis a. Rio Hu allagu. 60 km SE Yurimaguas , San the oute r border or the upper eyelid. and along the Mart in. by H. Bassler; AMNH 42790 . 4294 5. col­ ca nthus rcs tralis (where it was gold) 10 the tip or tbe lected at Chazuta. Rio Huallaga. 75 km S Yunm aguas. snOUI.where it mel its Ielksw. A white stripe extended San Martin. 274 m. by H. Bassler: AMNH 43.5.53. from lhe forelimb insertio n along the upper lip to tbe 43564 . collected at lhe Rio Pisqui. Ucayali drainage lip of lhe: snouc. ...'here il met che cantha! stripe. The above Contamana. Loreto. by H. Bassler; FM l\"H dorsal wrr....:e or the: upper arm was gold . The dorsal 56248. COIJ«-led al Cerro Azul, 24 km E Contamana. surface or the: forearm and hind limb " 'as lighl bn,)...n Loreto . 500 m, b)' J.M . Schunke; MCZ 24443 (adult with a fe.. ' small black ~poI s . The anterior surface and one tadpole). collecled90-138 lorn WSW Pucallpa. or the thig h bore a greenish wash. Loreto. b)' W.F. Walker. Jr .; BM 1900.11 .27 .38 and In preserva tive. the dorsum and the sidc of the body AMKH 17199. 17253-57. collected at Perene. Junin. are blac k. The latera l and upper lip stripes are gray; ROO m , by P.O . Simons (BM) and C. Schunkc (AMNH); they mee t eac h other on the tip or che _nou\, Thc entire FMNH 36778 (adul1 and cightladpoles) and 36779- 85 anterior surrac e or thc snoul is gray , T he uppe r lip (a d ul t ~). coll ec tcd in lhe valle y of the R io ViIOC. ~l ripe e\lends over lhe rore limb inser lion into lhc Juoin. by F. Wylkowski; 8 M 190 7.5 .7.9- 11, co l- axilla. wh ich is gray. The vcneer and the vcnlral surface 38 BULLETI N OF THENAT URAL HISTOR Y M USEUM OF LOS AN GELES COUNT Y Nr>.27 of the limbs are purplish gray with black marbl ing at the National Museum of Natural History in Wash­ This ventral pattern extends onto the proximoanterior inglon. D.C. portion of the dorsal surface of the thigh. where it C"mred 'l ~H 4.,016: FMNH ( '1igarimus (p,f jla vapir-lu.•. Cochran: 107. pla'e 42 tails lateral relative 10 the ad ult's hock. Measureme nts (c"l" r phOIO). are in Table 3 1965. Dr~drolmlC$ piN" .< gua"U1",n, i.' Healwole. Solano . Retationships,- p , petersi proba bly is closely re­ and He. twole: J50- 3Si, hoiot}'pe: MBUCV 31 t2. Alliplaniciede Nuria, & iivar. Veneluela. 100-250 m lated to P. h"lh'iall//< and P. pulch rip"CIUJ. 1970 Dr'ldrobd/cs hamphibians and reptiles ~pot present: venter black with blue or white marbling. !976 FROGS OF THE GENUS PHY LLOBATES 39

or blue or white with black marbling (rare ly. verner terns have been described as species or subspecies: almost immaculate while ). some of these populations. and some other popcra ­ Diag"o,i.,.-Phyll"bal n piclu.l is a member of tbe lions lhal never have received a laxonom ic name . may ph-tu, group. differing from all other species o f the be specifically distinc t from P. pic/u s. Because of group except P. ingeri (and rare ly. P. parm!u,) in lhis poss ibility . [ have indieat~..-l 10 which populations having a lighl proximoventral ealf ·spot. P. piC/us dif. published oame s apply . fer s from P. ingeri in usually lack ing teeth and in Pallern I ( Figs. 3D. 9D; inclu des Dendrohate.• having a eomplele light laleral stripe (ab'iCnl in P. bracelll" .• Cope 11\87 and Hylu"l esia /l av"" iela A. ingeri) , P. pinus further di ffers from P. parmlu< ;n LUI? 1925): Specimens bearing this palter n are known usually lackin g teeth , in having a black . brown. or from S\>Ulheastern Brasil (slalcs of Minas Ger ais. Go· gray dors um in life (red in P. !nln'ulus), and in having i,i.> . and f..lato Grosso) and easlern Bolivia f EI Porion. a complete light lateral stripe (incomplete in P. pan'ull,, ). 18°10' S. 6O"08·W). T he sno ut-vent length of speer­ from P , pelusi in usua lly lacking teeth, and from mens bearing th is paUern averages longer than in speci­ P. smllragdinu, in usually lacking teeth and in having mens bearing otbcr pallerns (Tab le 6). Pattern I is ventral marbling . P . piC/us resembles a sympamc characterized by a pair o f paramedian rows of ye llow spe cies. P . femoralis . but differs in smaller size spots (Fig , 3D); the other pauerns sometimes have gray (Ta ble I). in having lhe upper lip stnpc not meeting or brown dorsal SPOIS or marbli ng. but never have the canlhal stripe . in having a light proxiruovcntral ye llow spots as in Pancrn I. The thigh-spot usuall y is cal f-spot. in baving lhe firsl finger abo ul eq ual to long (Ir is short in the holotvpe of D. bruemlu.' Co pe the second (P. femoralis has lhe fif'l finge r lo nger from Cbapooa) . Frogs from Belo H",i?onl e (lhe lype than the second ), in lack ing webbi ng (P. femoralis has locality of Hylllp!e, iu jlIl,·"p;"la ) have a light ven Ier basa l toe webbing ), and in usuall y lacking teeth heavily marbled with black (Fig. 9D) , The black ventral Dl'scril'/ir",. -Measurements are in Tables I and 6 marbling is slightly reduced in specimens from lhe most The skin is slightly to strongly granular on the dorsum westerly known popu lation with Pattern I (EI Porto n). and the dorsal surface of the limbs . and smoolh on The marbling is greatly reduced in spec imens from lhe venIer and the ventra! surface of the limbs . Maxil­ J1<.,pulalions to the nurth (Cha vanlina. Cavakanli) and lary and premaxillary teeth usually are absenl , but west (Chap;uJa ) uf Belo Hortz ontc. In specimens from are present in some specimens (see below). The firsl Chapada (the type locality of Dendml"'lCS' brl'CCIl/!H linger usually is about equal to tbe secon d. but some ­ Cope). lhc venIe r is near ly enlirely light. with a few limes i> slightly longer than tbc second. The roes are black spots on lhe lateral ponion> of the belly; the dor­ web less sum is tbe same as in specimens from Belo Horizonte Although all popu laliuns of Phyllobales piClu., have ln life. in specimens from Belo Horizonle . Brasi l (lhe a light proxim oventral calf. spol. this species varies type series of H . /lIW""i<'W: notes from B. Lutz 1952). geographically (but not strictly clinall y) in snout-vern the iris was black. The dorsum was black with ycllow le ngth (Table 6) and color pallern . For co nve nience . spots arranged in a pair of paramedian row s, The latera l the variat ion in color pallern is described be low as and upper lip stripe s were yellow. The dorsa l surface of six patterns. which are not intended 10 represent taxa the limbs was black with yellow spots . T he thigh- spot Populalions whose members bear some of these pal. and the calf-sJ1<.,t were red-orange. The venter had a

T ~ a l . E (> Snoul·", nt length (mm) of .dulls of Ph)'II rdcr <> f de"rca, ing me.n ' noul-\'C nl Icnglh. P ~ \1 e r n 1: BRASll,o Belo H,,,-i1-OnIC . Fa/cnda Oli"" . Ch"pada; BOLIVIA: EI POrl,;n Panun 6: GUYANA: Karlabo. junclion Mcamu and M. z.nm; Ri"ers. I"an o. Kurupung River. M. "krcha F.lls . Paru;ma ~l ;" io n . Paller n J: BOLIVIA: Ru,rcnaha~ u" . ['' P.chi,e,,: BRASIL Tarauaca. Pi'una . Rin Tachita. Terc, inha hltern 5: PERU, l"evali. Rio P-."hilea

All Specimens M"I", Female,

Pallern Ran~e Rang" Range

re 22.5-3),0 27,5 n5 -29.5 2~. (> , 28,0-31 .0 29,4 21.5- .'1.5 2(>5 " 21.5-29.0 25.6 u HO_31.5 27.7 '".1.' 17.0-2 9,0 24.1 " 17.0_25.5 n3 19,5 -29.0 25.7 es 16.5-2 5.0 21,7 ", 16,5-n5 19.8 " 19,5-25.0 22.5 n 20.0-23. 5 21.5 ", 20.0_2l.5 20,8 '"6 20.0-2.',j 22.3 4(' BUUETI.V OF THE NATURAL HISTORY MUSEUM OF LOS A.\'GELES COUNTY No. 27 greenish while or bluish white ground color with bl~ck ncar Leticia (my notes}, tbc iris. the dorsum. and the marbling. side of the body were black . A complete yellow lateral Pattern 2 (Fig. 'JE; includes Df'mlrobal~s bruc.."UIUS stripe extended from lhe groin 10 the upper eyelid. along Fillinger in Sleindadll1er 18(4) ; Specimen, bearing this the outer border of 111(0 upper eyelid. and along the: panern are known from Sao Vicente (, tate of MaiO canthus rostrali s 10ee lip of the: snoul. where il mel ils Grosso). Brasil. Sleindao;hner', ( 18M) combined Iocal­ fellow. A while stripe eXlcnded from the forelimb in­ jl~ li,t of Drndrobates rucnemis and D. braccana fail, sertion along the: upper lip. ending ventral to the eye or to a,sign loc alitie s to each fonn. Steindacbners label nostril. Tbe dorsa l surface of the limbs was lighl bro....n. with tile tWO syntypes of DeluJrubn i" Tscbudi 1838 and Dt'lwrooott' s t'uc"t'mis elliptical orangoe spot. In one unusual specimen (LAC M FilZinl!'er'I/ Sreindcchner 1864): Specimens bea ring 6 1006. one of 12 specimens from Teresinha). the calf­ this pattern arc known from Bolivia (north. nonhw esl, spot was blue. The ground color of the venter and ven­ and west of EIPorion. in the dep artments of Santa tral surface of the limbs was black , The thlOat bore blue Cruz. Benl, and La Paz). Bras il Ithe terr itor y of marbling and spots. The breast and belly bore ....bite Rondonia. the Siale of Para . and al Urucum near marbling. Spec imens from Pixuna, Brasil, near Maneus CClnlmoo on lhe Brastlian-Bol ivian border). and Sun­ (my OOIes) . J'C'sembled the Teresinha specimens, except name . In Ihis pattern. tbe dorsum lach spots; the lhal the iris WlLS black Wilha gold nng around lhe pupil. thigh-spot is long (Fig. 3A) and sometimes broken. the back was black ....ith gray spou. tbe lateral smpe and the venter is dark with 1i!! lu marbling (Fig. 9F). was gold, the dorsal surface of the calf bore IWO dark In life. in specimens probably frum the Rio Mamore. brown transverse bands. and the axilla and calf-spot Bracil (t he type serie s of D. t' u ("11~m is; notes by J. were yello w-orange . In two juveniles. the:dorsal surface :"ialkrer in Steindachner 1864). lhe ins and lhe dor.um of the upper ann ....as orange·gold. and the dooal sur­ ....ere black . The laleral and upper lip snipes were face of the forearm . calf. and larsu s was green ish gold . while . The axilla . the thigh. spool . and the calf-spot were Specimens from Santa Cecil ia and Sarayacu , Ecuador red-orange. bordered with black . The junction between (noles by J.D, Lynch at KU). resembled the Teresinha the rectangular proximal and the oblong dista l portion s specime ns. exce pt that the iris was bronze. the dorsu m of lhe thigh,spll t was yellow. The venter was grayish was black. brown ....ith black spots, or gray wilh black black with whitish spoe, and marbli ng. In specimens spots , the lateral stripe 1'."3-' cream or yellow. the Uppel from PosI Gonini and Sipali",'ini. Suriname (noieS by lip siripe was blue-green in one specimen. the dor1;:al M.S . Hoogmoed al RM:-> H). the OOnum ....as black wjth surface of the calf and forelimb hore brown or bhd. gray marblin g. The lateral stripe:....as cream . The upper bands and spots. the axilla was yello...' or yello...'· lip stripe was while . The ax illa. the postcrovemral sur­ orange . the thigh-spot and calf-spot were yello w or face of the upper arm. the thigh -spot, and the calf-spot orange. and the ventra l marbling and spots were blue . were yellow . The venter and the ventral surface of lhe Pattern 5 (Specime ns hearing Ihi.s pattern may be spe­ lhigh were black ....ith blue . greenish white. or bluish cifically distinct from P. pictus) : Spec imens bearing this while marbl ing. patter n are known from paris of Amuonian Peru (Tsjc­ Pattern 4 (Fig. 3B: include s Dt'lldrobol~s hoh" d i vemeni; color slide by A.C . Thoresen: l\"evali: KV Boolenge r 1883): Specimens hearing Ihis pane rn are 144351-60. LACM 9241 3; Rio Pechitea near Bosque known from most of Amazo nian Pe ru. Ecuador. and Nacional de Iparia: TNHC 3(4 69) . "These specimens Colnm bia. and in Amazo nian Brasil in lhe lerrilory of resemblc frogs bearing Pallcrn 4 in size and color pat­ Acre and the slale of Amazo nas. These relalively small lern (( he Ihigh ·spo t is shon ). bul differ in lhe following frogs (Table 6) lack rows of yello.... dorsal spools (al· characleristics: the dorsum is Slmngly granular. leelh lhougb they 'Omelimes have indistincl gray dorsal are pre.cn t. lhe head and back hear small spols (some· ~s ) and ha\Il.' a short Ihigh-spol (Fig . 3B); lhe venIer times panly fu~ logelher and somelime s arranged in is dark wilh lighl marbling (in pm.el'\-ative . the mar­ Ihree or four longiludinal rows). lhe calf.spot ellends bling is conspicuous on the belly and indistinct on the onlo lhe disla l one- founh or one·lhird of tbe

Ihrtl;ll) and most of lhe ventral surface of the limbs are 5894 1). from which be had removed one of lhe malil. lighl with dark marbling. Of a series of \ i~ specime n\ of lary bones . Tin)' teeth were "isible on this bone. bUI P. picfu, (r.' HC 3646 1. 36469-70. 36476. 36482. not on the other maxilla.....hleh was lefl in place . _\6.197) I..ken by J. P. Bogart at different ,Jate~ in 1967 Thb )l.Igp:sted tholt m.tensibl.)' lOOlhles.s specimen) or and 1968 on lhe Rio Pachitea neil!" tbe Bo"'lllC~ ac ion al P. piaus hast: concealed teeth. I cjeared and !>Uined de Iparia, Peru. one (ThHC J6.1691 h a ~ Pattern 5 and four such specimens: Patte rn 4: Tere vlnha. Brasil five have Pattern 4; I do no( know if all tne..e 'J'«imens (LAC M 60977 , J~1S 1b80 ): Pattern I : EI Porton, came from thr ..arne deme . In life. in ~pecimen~ from Bolivia (LAC M 6(976): and Subpattern 60;: KUl\lpung Nevali (notes by D.B . Ekk e n ~ in li ll.) uc:no Ocopa (MCZ 24433; LACM 1(0979); by Dr. Hahnet. as the lectotype of lRnJ robm t s Nlhnt/i Owo:Jwnayo tAMS H 42892-94 ). Pasco: Sewali (LAC.\1 BouIenger 1883. I designate SHM W 3818 :1. a 2b.5 92413; KU 14.1351--(0); TS~TC1Ii (color slide by A C. ~n) . C.W" Luis.iana. Rio Apurimae IMCZ S8413J. mm female. proba bly from Sao Vicente, MaIOGrosso. BOLlV IA. -S".,a Cral : Santa Cru2 (MHSP 4910; Bra..i1, co llected by J. Nanerer , as lhe Ieclll. 11858. believed thatlhe types came from Bolivia. GuilX (1950) 111l6()....65); MonICAk i rt (BM 96.6.29.18; KU 93IS1-52 ); listed the type locali ty' as Santa Cruz [Boliv ia]. P. Jac"';· Acan, a. Rio TapajOs CB. L UIZ and Kloo. 1952). piC-flU has llOl been reported from Ch ile si~ Dumrnl Ro.i...,.: Guajari . Mirim (f M.-.H 49770. 49844); Rio A.cn; ( ~ I S H and Bibro n's work. P. picrUJ i\ a Iowlllnd Amazonian Mamo,,!; (l'oi H.\t..... 19190:1-4). Tarau... i 83248). Gbi (CM 2431_36; (US~ M loU9S9-61); upper GUejar ond EI Mico. Ma; -..a P..-ISH 9209_ 19: USSM 115974)_ 3/ i" ,.. G.....I;.' Bdo t USS M 144962_67); Cana Guapaya. Mllurt'nli IUSl'oi M Horizonte tUS:-iM %986; CAS 98099; KU 93141- 42) ; 14726'). A.....tona .: 7.2 Lm ,",W 1.elicia (LACM 610(8) Lagoa Seen. fOOl of Serra do Curral (UM.\IZ (10023); ECUADOR. - l\"apo: Sanla Ceci lia (KU 104764 _72. Aeaba Mundo on Serra do Cum l (B. Lutz 1952): Cai'll de 109291 .109293_94.109296. 111$()4. 111519_28. 122271. Arcia ncar Belo Horironle (B. LUll 1952); UIs"llnCe tIC~r 122273-7 4. 122277_79: .\ICZ S8414_ IS, S8411); limon· Scm do Cabral (B. LUll 1952) c"" ha (LACM 72672-7S; KU 99119- 20); Purrlll libre. Rio Asuark o (KU 112287-90): L.,rclo (lAP 3'S8j, PU.IUU': VENEZUELA.- Bo/i."" 5 km E La, Chicharra'. Alii · Sarayacu (KU 120671_72); Alpayacu ncar Mera (UMMl planieie dc Nuria (MBUCV 31( 2); Hacienda San Felipe quinia drainascs (AM)';H 42474. 42691. 4,8): jlllK'linn Mcamu and .\Ial.lllllni Rjw" " (AM!' H 18996. 42698. 43372): Ollama near Qucbrada Ca. hi"")a (A.\IS H 18998); Kamal u... (AMSH 19(100); Kurupung Rjw"r(UM.\IZ 42977- 80); Rio Pi>qui (AM!"H 4J2 7JJ; uppe' Rio Ila) . 83S98 _60) ; LACM 60978); Mad reba Fall•• Kurupun, C AM~ H 4.'-4(7): C hiboya tAMSH 42070); IQuilOt CAMSH River (UM.\IZ 85147); " . ano (U.\l.\tZ 83597); Paruima 42209); Pampa He Rio Cu.haNlay (AMS H 42372. .\I i ~ ~ io n (UMMZ 85145-46: US~ .\I 1I8681-82); CUlllni 4 2 4 ~ ) : Rio Tamal'a tAMSH 43360 ; Con'amaN (A .\t ~ H RiVC1" 24- 32 Lm a ~ Banic-a (B.\I 1934.11.1,1-4) ; !ol.. opc 42412); SobnI. Jo,o,-er Rio Tama)a (A.\IS H 42881); Tapi.tlca. Fall•• CUluni Riwcr {BM 1934.11 .1.51; Oko Mountain. OJ'P'O"ilC Conwnani (AMS H 42930-32); Yurimasuao. Rio (P.>IS H 26724--26) Hualla, a tB.\' 1947.2 .15.14-20); Dumo Sanla Clara and Sl:RISAME.-."kb rir, Sipaliwini (.\ISH 1lOt'>. 1131); Pn>l Campo SantaClara.nearOreliana tUSS .\t 127201. 127203--08. Gonini. Cue"",ni RivieTt.\lSH 322. 350); Post Tip. Sc.. 127932). S... .U;Uf;. : Achinami.... Rio H..alla, a (AMS H Riwer(R.\IS H 15896) / 976 FROGS OF THE GE...-US PHYLLO BATES 43

GUYAl"E FRA." CA'S E__Empounou W>i.ki. I ~ km SSE "" of the interorbital widlh The diameter of the eye is Maripa>oulIl (La 1023). I A times the distance from the eye to the lIOloIril and RECORDS .-CO LO~IBlA. DOUBTFUL Cllltlar: Villa slightly more than twke tbe diameter of the I)"mpanum. Maria (lJSS M 14Nlbl. (46866). BOU VlA. ClN1ttlba"' N: Yungas do: COC hatwn N. 2200 m (UMMZ66646). The pos terodo rsa l portion of the tympa num is con­ Clear..d and stained mm..rial, _ICClh pre",m: LACM cealed. The finger dis ks are small relative to those of b0979; Pattern I: loACM b0916 ; Panem 4: LA C~l 60977 . D..nclm hll/ t'J. The first finger is about equa l in length JMS 1680; P.~ Uer n 6: LACM 60978 . to the seco nd. T he fingers are webt ess. The tarsal fold is ill defined and slightly curved , extending along the distal 4S perccnt of the distance between the inner PhyllobQU J pukJrriputu s new species me ta la rsal lubercle and Ihc hcel fold . The tarsal Beautiful- breasted Poison-arrow Frog tubercje is absent. The loes are ....ebless Frontispiece II and Figure 6E In life (Frontispicct Ill. the iris, head . ~k. and sides wert black . A com pklc bright )cllow lateral HolofJpt'. -loACM 42297. co llected 1.1 Serra do stripe e xte nded from the Jroin 10 the upper eyefid. Navlo, Terruorio do Amapj, Brasi l, abou t 120 m. along the octer border of the upper cyclid. and w n!!: between 1330 and 1715 h on I lu ly 1968 . by P.A. tbe canthus rosrraiis 10 the lip of the soout. ",he n: il Silverstone and Sr. Ferreira. mel its fello w. Anolher bright yellow stripe extended Topop Qf/l1rp"s (nine ad ulls) ,- WCA B 18 22 1; from the forelimb insertion along the upper lip, ending LACM 42296. abou t 120 m, I July 19MI; LAC M ventral to the nostril; the anterior end of the stripe WIlS 42298-301, about 120 m, 2 luly 1968; LACM 42302. blue. A blue suffusion clouded the black limbs. The about 160 to IIlO m, 3 Jul y 1961l: LACM 42303. about proximoo or",,1 surface of the thigh bore a con~p ic uo us. 100 m, 5 July 1968; LACM 42304 . Serra do Veado o blong. brighl yellow. black-bordered spot; a similar (one of the hills at Serra do " lIvio). abou t 290 to spoe adorne d the axilla . euendin!!: omo the pmximo­ 3 10 m, 6 July 1968. (All LACM parII t)~ were col­ ~ior and prox imodon.al surfaces of the upper arm. lected b)' P.A. Silve~onc: and Sr. Fem lra.) The venter and the ventral surface of the limbs were D..finilion . - Tet th abM:nt; IClC' S ",eblt ss; dorsum med illm blue, beann!!:discrete black marblin!!: , blac k; complete yello w lateral \!ripe; li!!:ht pro~ illlO­ In p re ~rv a l rv e . the grou nd color is black . Are as ventral ca U·s po t abse nt: ven ter blu e ",ilh hlack that were yetlo....in life are white. The canthal portion marbl ing. of the lateral stripe . the blue anterior end of the upper Diagnosis, -Ph.l'lIobsent the left side of the posterior belly'. in P. i"g..,i ). and in lackin g a light proximovemra! Dt'Scripl;on 01 p

R~ IQli(»/$hipJ .-P . pulchriprCIUS probably is closely D..fi nilion. - Teeth present: roes ....ebless: dorsum related 10 P. bvli"ianus and P. prf" rsi . black : co mplete green lateral stripe; light proximo­ Sw much comems (one specimen) ._ From ~C~t ventral c al f.~ pot abseet; venter green (anteriorly) and 42302: HYME....OPTERA: FOIlTl id dao:: B/t'phuridana blue (posteriorly). no l marbled. b".'C ntra l calf-spot, and in lacking vent ral Distribu ,;OII aJU1 habi tat (Fig. 16). _Elevlllion rec­ marbling. from P. pan 'ulus in having a black dorsum erds an: from 100 10 3 10 m. P. pulchripulus is known in life (red in P. pa n·uI U5). in having a complete light only from the type locality. in the Guianan region of lateral stripe (incomplete in P. pt ypl'. - LACM 64435 . co llected al Pan de the upper lip. ending ventral to the nostril. The lower Azucar . 39 km N ~ EOx a pam pa . 10"15'5 . 75°14' W. in lip. the breast, and the anterior t.....o-thirds of tilt belly the lscozaztn Valley (Ihe Rio Iscozazm is a Iributar)' were green. The ventral surface of the limbs and the of the Rio Palcazu. in the Pechlte a drainage). De­ posterior one-Ihird of the belly were dark blue. The panamenlo de Pasco. Peril. 31lO m, on 16 Augusl 196 1. throa t bore a pair of lateral black areas (perhaps a male by R.E. Elheridge. secondary sexual charac ter). Parur),fWs (one adull and seven ladpoles).-LACM In preservanve , the holo type is entirely black 10 6+436. jopoparatypes (seve n tadpoles from the back dark gray. ex.ccpl for lighl gray lateral and upper lip of the holol)'pel: LACM 6+434. collected at lscozazin stripes . ainlrip. 47 km r-;NE Ou pampa. urn ·s. 75°10·W. Adull parat)·fW .-l n preservem..e. ttle adult parl­ in ttle lscozazin Valley. Pasco . Peril. 360 m, on 19 type (LACM 64434) resembles the bolory pe: it is l uly 196 1. by 0 .8. Wake and R.E. Etheridge the same sex.and length. 1976 FROGSOF THE G£SUS PHYL I.O BATES 45

Tadpolet (Figs_ l ID. 1 2 E ) .-~ ho\ol)pc earned bod)'. pan of the upper lip. the proxrrnodorsal surface '>eVen ladpok<. (LAC M 64436) on hi, back. The lad­ of the upper arm, lrac~ of light pigment on the donal poles ...'ere in S liI~ ~ 24 (1....0 specimens) ;md 25 (fIVe and ventral surfaces of the thigh. and the proximo­ specime ns). ~le a.~uremenh an: in Table 3. ventral half of the kft tarsus. Dis t~ibu t iOll and habitat (Fig. 16). _ EIC'IIlIlion rec­ Refe~"d mOfuiaf. - Tbe holorype is from Roque. ords are from 360 to 380 m. P. smaragJinll.• is kno....n Per u. at 1097 m elevation on the Ama7.0nian slope only from the type localities. in dense forest in the of the Andes, in the drainage of the Rio Huallaga, 3S Amazon drainage of Pl:ru. east of the Andes. km south"'lutheast of Moyollamh a and 76 km west· Etymology. - Latin: smarafi: dinus (emerald) refers northwest of Chazuta . I refer nine additiona l toothed to the green stripes and me green anterior venter. specimens to this species . Tbese specimens are from east of the type locality. from the drainage of the Rio Hualla ga (Chazura. Pachiza , and the Rio Cainar acbc j. l~ i vl1llllUJ group Specimens from Pachiza differ from the bolorype in having smaller. scattered dorsal ,pOls. Th is group includes twc species. (f'hylfoiJ<,ft's bass· Some of the spe<:imens from Chazuta have a color lu i. P. trivitlatus). pattern similar to that of the holotype: I do no! know Defi nilion: snout-vent length of adults moderate their color in life. At Chazura. W.C. Sherbrooke ~~ 10 large (3 1.5-49.5 mm); loes ....ebIeM; paired collected lOOl hless specimens of P. trivunuus (....ilh lateral (Figs. 2. 6A-C); light proximoventral calf· a pair of yellow lateral s.triJlC'o and a )'ClIollo median spoI absent. d~ 1 stripe) together with three toothed speccneos (....ith a pair of orange lateral stripes and no median Phyl/oootes banlui (Mclin 194 1) stripe). The three toothequc . ISanManln.! Peru(,t 097 mi. The venter in the toothed specimen s was black with blue marbling (Fig. 60). Defi nition . -Tee th prese nt; toes webless: dor sum Generic oflm:ulion._Mel in (1 941:65) desc ribe d black with green or ora nge spo ts or mar bling. or the bolotype a.~ a Dend~ooo"s because he thought il uniform orange: complete green or oran,e la teral had no teeth. onl)' "tooth-like aspenties." I examined stripe; Iighl pro :\imo\'entral cal f·spot absent; venter the holol)pe. It has teeth and small finge r dish; it black with bloc marblin,. is a Phyf/obates. Diagnosis. -Ph.\' //oba/~J batsleri is a member of Stomoch contents (one specimen)._From A."-tN H lhe trivitta lllJ group . di lTering from P. tri,'itralll.1 in 43402: HYMEl"O PTERA: Formicidae: Pheidofe .p. having teeth . (21. CompOflOflls'! sp. ( I); O RTHO PTE RA: Telli· D~ s c ~ipl ion . - Meuurements of the bclorjpe (a goniidae ( II: HEMIPTERA : fragments. 37.0 mm female) and referred specimens are in Table I. Dis t~ iburion and hobitof (Fig. 141.-Elevation rec­ The skin is strongly granular on the dorsum and the ords are fmm 274 to 1097 m. p , bllss/eri inhabits the dorsal surface of the hind limbs. and smooth on the Amazon drainage of Peru. from the eastern foothills forelimbs, the venter. and the ventral surface of the of the Andes cast to the Rio Huallaga . hind limbs. MaxiJlar) and premaxillary teeth are present. The first finger is longer than the second. LOCilfitiH ,-PERO. Son MOrf;n: Roque. 3~ km SSE The toes are webless. Moy"bamba (I'\MO ~1l); Ch:llU I ~ (AMI'\H 4Z867. 42944: (AM ~ H In life, in the holotype (Melin 194 1), the dorsum USl'iM 166903A-C); Pachiza 42321, 42333, 434(2); Rio Ca;narachc tAMSH 42313). wil.s black ""ith a median row of light green ~pols (Fig. Cfea~eJ und swi lled "I(l/e ~iof. -AMS H 43402. 2F). The side of the body was black ....ith a complete lighl green lateral suipe extendin g from the gro in 10 Ph)'ffoba fes trivinanu [Spix 1824) the upper eyelid, along the outer border of the upper Three-stri ped PoisorHul'Ow Frog eyelid. and along the ca nlhus rostralis to the tip of Figures 2A-E. 7C the snout....here it met its fellow. Melin did 1101: men­ tion the colo r of lhe venter in life. In preservative. the bolotype is dark bro....n. excepl 1824 H,·/o ,ri'~II(l1O Spi1: 3~. pla....'I, fig. I. type locality: Rio Tde, B nI ~ il for the following white areas: a median doNiI row of ?1824 H,./o ";lI a~i "'(l Spi1: 36. p l ~ l e 'I, fig. 2. Iype seven spots. a fe.... small spots on the righl side of JoealiIY: Ega [=Tefel. Braoil (according 10 W. the beck. a complete lateral stripe on each side of the Pelers (t8 73:2131. the type. of H. l~iv ; IIa10 and BULLETIN OF THE NATURAL HISTORYM USEUM OF WS ANGEUS COW..TY .'0'0.17

H....~rrima belonged 10 1M sam: ~~s I 1La\~ and one- pair Jateral). In some specimens. lhe median noI "Cen tM I ~ pe~ !he)' ..-ere ck"roye4 .. WoOd corsal «npe 4 , one- or mo re small break. (Fig. 20). War U. axordin!, 10 U. GlUber lin lill.)). Such spec imens are known from Santa J ulia ne.... Faro 1826. H,-kIpI'Wl" ;'~ lIa Ul , 50., in Schkp:l: 239 ~ ari ilo 18.\0. lHNlTt>ba". Im·iltalM•. Wal lct-: 202. in Bras il. Puerto in Colom bia. and Morale s. ?18.\O lHNlTt>ba"...i~rriIJIM' . War.lio r: 202. Pschiza. the uppe r Rio Cllshah:l1ay. p...mp;.o Hen nusa 18).8 H,"""",Wlli~ . T..:hudi: 70-71. (lower CIl,habalay). and ~.5 km from Puerto Santa 18~8. Ii...."'"" (pan). GUnther: H,.lapI,.;" 125. Clara in Peni. At 1IlO'>l of rbe-e localities. boIh ,peci 111M o,NlroDQU. "iK"rrUorIU ( part) . Steildal;hner. 259- men, with untomlen median ...ripe and spedmen, 260. an 111M lHNlrobm,. ti....1OTi1U ( put). Sa: Illda<:IIMr: 260 . I\-ith a broken median smpe have been conecred. One­ 1&82. lHlldroD..,,. rrMllQ'1U (pan). Boult-np:r: 1 44_ 1 4~. spocimen (AM ~ H 43 117) from Pampa Hc-rmoSll. . P.;ni. 1926. O,lIdrol>,,,,. trivilla'M.'. Miranda RiMir o: 181. has a med ian don;a.l row o f e ighl small ,;pots (Fig. 2E). !ell·fi8. 98 . "The spec imen lach teeth , and is pan of a 'Cries o f 1926. On.droba,,. U lr(Jvit lallU Mir~nda Rib9 1824:plate 9. fig. I). the dorsum was black with a 1\161. l>r lfdm lmlr , I,il'illat...,. Cochran 107. plate 41 )'ellow median stripe (bro ken intcrorhitally}. T he s ide (cnl", phoIO). of the body "la, black with a com plete yellow lateral 1969. Drndrohin ' ye llo w spols and marbling . Spec imens from C ha7uta. 14 -1 6 . plate ~G ~I (o nly USNM 118681 _ 82. Peru (color slide s by W .C . Sberbrooke), resembled MCZ 24443. ANSP 18188 are nlCu bbd; with green miirbl inil; sorrenmes dorsum uniform the bokxypc. but the strip:s were green.Tbe median g ree n; complete yellow, ye tto ..... ·green. green. or stripe was unbro ke n. indi" ioci . a nd darker than tbe: o r;IlIFC lateral srnpe: light proximoventral cal f.spot latera l stripe•. TIle- donal surtece of the fore-limb til-« absenl; venter blac k ...-ith grren. blue -gree n. Of blue a green stripe- eXlCnding the IC'n gth of the fo relimb. 'flOls or marbling. The- doNlI surface of the hind limb was nearl) cove red D;oxn o.1;$. -P hyf1obol~$ Iri"i tl(JI"$ is a mem ber by green marbling and stripes. "The ventral .urface o f lhe tri"it/ and slides) from four locali ties' is longer than the second. The toes are- weble... Yartnacocha on the Rio Urayali (60 km nonhnorth­ Phyll l.hll/ l'.1 trlvittatus has three co lor pauerns: wes t of the junction o f the Rlns Pachitea and Ucayal i). Pattern I (Figs. 2A . D- E. 7C; include s holotypc of the lowe r Rio Pacbitea ncar the Bosque Nac ional de Hyl

Doma San.. Clara ncar Orellana tU SS M 127214 -18); 11030). S ~. : filli li bbo camp (RMS H 6215). "'.... Chay....it. (BM 67.6.13.24-25 ); Balsapueno tFM~H SMIl; ~ij~: S...... Gebor8te (RMS H 6413. 6600--01. 6603, 6637. Cachipun10 Mar Balsapueno (AMS H 428761; Rio CaC encuemran desdc Co sa Rica hasta Rim. C,mlay a and Tapiche (AMS H 429%); Rio Blanco tA ~ 'S H 42985); Rio Tamara (AMNH 421101); Ca, hiboya el Peru. Boli via . )' e! Brasil. La mitad de: [a. e.pecies (A MSH 42044, 42976. 434H); Oll,nla "" .. Quebrada .c eecuemran en la rellion amazonica. Las 20 especes Ca.hihoya (AMSH 43212); Rean Re 1824); Tdd K...... 19521; Rio l",'ary. 30 lr.m ,jo,.'n>lrcamffOOl Benja­ Ph)' lIobal~s . Asi es que Ph)'lIoba l~s 00 se disl in8t1e min Con!ltalll (CAS·SU 11879); hacoa ; lB. Lull and KJo.l. I9SZ); Eirtrnept. Rio Jurui (Miranda Ribeiro 1926); M..... de DtruJrobcJlts por la presencia de dientes [aunque (K. Sc'-'tidl .-.d I n ~ r 19S1); Rio Amazonn tUM~lZ 62S53); la mayuria de la. especes de PIt)'II0b01ts tiene:dien,e.). Paranii do Jacan: . !iO\Ith of Io"'-er Rio 501,,no" ,C AS·SU sino por el lamaOO de los discos )' la laf8ura de los 118tl6). "po: 6bido> (Miranda Ribeiro 1926); Caotanh.aJ dc:dos. En Ph.l'ffobo1tS . los discos rerrmn ales de los Grande ""ar 6bidt» (MRHS 61. 6I B-6IE); 5.a.nla Julia DCar F~ ro ( MRHS 4741; hiubni (ZS BS 99/ 1911; MCZ ultimos Ires dedos de I. mano son relalivamente 121l8); llha de Mar~jO (MCZ 1S663); 19-..apt·A~u ( BM pequefios (menos de 1.5 veces mas ancho que et 1904. 7.26 8); IP'EAS. 5 lr.m E BelCm (KU 127536-391; dedo); d primer dedo de la mano es iguai a 0 mas largo Uliog-a, 7 km E BelCm(KU 127.s40). que e[ seg undo . En Drndrobo1ts. los diso;:ostermmales GUYANA ._E... , lh"''''IlTa: Vr)'heid. LuSI ( BM 78. de los ullimos ues dedos de la mano son relalivamcnte 12. 13.1 9). 1..", D~...~rara : Wisma r (AMS H 45774 ). Rllpll"""I: N Acara hy ~' ou nl ai n •• \Ii Sew R i~ er ( KU grande> Cal meeos dos v«es mas aocho que el dodo); b9696_ 700); hud...-aler. h abu C re ~ k ( FMNH 30909); d pri mer tkdo de la mano e. mas corte que e! segundo. Ku)uwini Landing (AMS H 46232); MaNdl (AMNH 46570); Las sig uienl e. especic s anterio rmente convlde radas Paraboom (AMSH 46236), .'Jazon." i-Pol4ro: Oko M (AMNH 1349.1.2 4915.709118_ 89); Kamaku. a (AMN'H 21 401)~ Cama ria Road (AMNH ingeri, p<1Tvulu,l. pklU.f, y trivusasu s. Las primer.." 3'1119). Ires de e. las eSPl'cie s ne nen dientes . Algunm ejem­ SURINAME.-A'ickt rit : Frede rik Willem I V' ~ a ll e n ptares de P. pictus ricncn dicntes y la mayoria carece ( FMNH ,10907_0 8; BM 1936,4,2,1-2. 1936,9, 3,6 -8); de diemes. P, Ir/vll/lITlI.' carece de diem.:s , Sipaliwini Savanne (MSH 767); Kay..,r Gcborgle (MSH 575, El genero Phyl/obnini, Coeroeni Rivier (MSH 321. 326, 353. 394); corso ten elanimal vivo ) es negro ron listas de un co lor Wilhelmi na Geborgk. linker Coppename Rivier (RMNH subjdo. 0 e. roratmeme de un color subido (pe r ejem plo. "nefl l; Fallawatra Rivitr (Udlh de kude 19(4), S ..rUwmt: rojo). mienlras q ue en C%Slr1hus. el dorso (en el loden'IVan"", Kamp 8 (AMS H774SO;RMNH ISOI4. lS(17); animal vivo) liene li.la. p;ilidas 0 grises, y 00 e. Afobida road , 10 lr.m from I'hodra tRMS H " neall , B"'.... de: ,..,..: Bro""n.borlJ (RMS H 157041; railroa.l. km 121. near un color s.ubido, Probablernente la pre sencia de colore!' q..-ry at fOOl of Bro.. noborJ (RMSH 9716. 9719. 9723); subjdcs se n:lac iona co n la presencia de venen<.> en ...ilroad , km 121. upper Mah.ami KInk (RMS H " ncaO; la piel. 0 sea que Ia piel de: PIt)·JloIJorrs uere veneoo . Brokopondo (RMS H unc~l ) ; Gotod P1~r IRMS H " neal); ) [a de ColoJttlluu no 10 liene . Hay especes quo: se T~poo"'~lra . Gran Rio t ~ ' S H 2631; Toek...eeeree Krcck CRMS H "neat ); .... Sarama:ca Rn.- io-r. lr.m 91.5 (BM 1946.4 pareccn a am bo. generos; tal Vel 10. dos genet"O'l sc 1.231;"pp:1"Stuin.amt: Rivitr ~ Sara Krcclr. (MCZ 10164, junlarin en e! futuro , 1976 FROGS OF THE GEt·... US PHYLLOB ATES 49 AI menos dos espccic s de Phyl/obmes se uufizan Las ranas del gencro Phyl/obmes son dium as, y la por los indigenas para envenenar los dardos que tlran mayor ia habita eI suclo sombrcado de la >elva humeda. de bodoqueras . Los indigenes de Ia rribu choco, que donde sc allmcntaprinci palmcnte de hormigas pequcfras. habita n la region lluviosa del Choc 6 en Colombia, Ponen sus hucvos en hojas . Despucs de salir de los traspasan el cuerpo de la rena venenosa con una varilla, buevos, los renacuajos sobcn la espalda de un ad ulto en algunos cases teniendo la rana sobrc un fuego , y (al menos en algunas especies. e! adult o que llcva buren las puntas de los dardos contra la piel de la rana. los renaeuajos puede ser el padre 0 la madre). EI adulto Estc procedimiento, de scr!ro per Posada (1909) y Ius llcva a una pequeria extension de agua (pm ejempto, Wassen ( 1957), causa tanto dolor al pobre animal que el agua en el pecfolo de una hoja de palma cafda}, donde su veneno exuda sobre la piel y se adhiere a un scrc o 10 ' deja Los renaeuajos ccmplctan su desarrollo en espiral en los dardos . Las des especies de ranas que el agua tiencn la mala fortuna de unluarsc sc Uaman en la En 1<1 unica especic I P. \' il/ " IIIS) cup historia de lcngua embena 0 empera de los ind igenes " kokoC" vida sc conoce bien, no hay amp lexus. (Phy llobmo (lUrOllU!Ria ) y "neara' (Phyllobotes Se describcn en esta ohra cuatro especies nueva s hieolor ). procedentes de Sudamerica, Ptrdtobares petersi, P, tndchripectus, P. Jl1l11rugdillU.I' . y P. t" p" m 50 BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGEUS COUNTY 1\'0.27

L ITElV.TURE C ITED 1912. Descriplion. of fit '" Nlrachians from the Andes of South Amefita. preserved in the British Mu' eum. Ann. Mag. Natur. Hist., Scr. 8. 10:185_191 AUllOl,,·UQI,'E. E.X., J.W. OUl' "'SD B, WnKOP. 1971. 1913. On a collection of batnll;hians and repciln INdc BatndlOto~in : c ~ m is try and pllannaoology. Science by Dr. H.G.F. Spum:II, f .Z.S., in the Cboco, Colombia 172:995-1002 Proc. Zoo!. Soc , London 1913:1026-1038. ANDERSSON , L. G. 1906 . On battlchians from Bo livia, 1918. De"Cripcion. of rIC'" Soulh.American Nl rxltians. Argentina. and hru, coIlecled by Erland Non1enskiOld Ann. Mag. Natur. Hi.. ., Str . 9, 2:427..0433. 1901-1902 and fl:ils Holmgren 1904- 1~ . An.. Zool. _. 1919. De"Criptions of I,", new lizards and a fit .... frog ;\(12):1- 19 from lhe Andes of Colombi a. Proc. Zool. Soc. London 1945. Batrachians from East Ecuador collectW 1937, 1919:79-81. 1938 by Wm. Clarke· Mxlntyre and Rolf Blombell. BUSSEU-. P. 1973. Vocaliz.alions in the territorial behav ior Ark. ZooI. 37N 2):I - 8S of the froglhttd,.~. """,ilia . Copeia 1973:277- 284. BUlKWl, T. 1905. Repfilia and Amphibia. Pages 98-1 02 CH"'PM"',,". f,M . 1926, The di"ribulion of bird·li fe in Ecuador. in O. Bangs. W. Brown, J. Thayer, and T. Barbour, Bull. Amer, Mus. Natur. Hi!>! . 55:1_784. The vertebrata o f Oorgona Island. Colombia Bull. Mus. Compo ZooI. Hatvan:l46 :87- 102. COCHUS, O.M . I95S. Frop of southeastern Brazil. Bull. U.S. Nat. Mus. 206:1..04 23- 1908. Some eew repfiles and amphibians. Bull. Mus. Compo Zoo!. Harvan:l51:315- 325. _ . ~':' ~ ' l';;~ i(f~~rc~i;~ ~.% ~~~ world. Doubleday and 1909. CllIT«tions regarding !he names of two mxntly described Amphibia Salientia . Proc. BioI. Soc, W3$ll. COCH....S. O.M. ",so c.J. (joIN. 1964 . Description of a eew in~on 22:89 frog,of the r:enus P/ryJJoba~. from Colombia (Amphibia, Ranidac. Dendrobaunac), Stnckenherg Bioi. 4 5 : 2 .~ 5 ­ B"'''80,UR, T. "'1<0 E.R. DUNN. ,1921, Herpetological novel­ 257 , lies. Proc . BIOI. Soc. Waslllngton 34; 157- 162. COCHIt"'S. D.M. ",s o C.J. GotN. 1970. The frog-s of Colom· BU 80L'a. T . "'SD G.K. NOBLE. 1920. Some amphibian s bia. Bull. U.S. Sal . Mu•. 288:1--tl55 . from north...-c"em hru. ",ith a ,"i>.ion of t~ &Cllelll Ph>HobdfU and T~Jmafobi"•. Bull. Mus. Compo Zool. COPf., E.D. (1861) 1860. De>eripfion. of rep llto from tropical Harvatd 63:3%-427. America and Asia, Proc. Acad. Nalur. Sci, Pbiladelphia 12:368-374. BUBOIiR. T . ",so C.T . R"'NSDEN. 19 19. The he~IO,Io \' of Cuba. ~cm . Mus. Compo ZooI. Harvard 47:71-.1 . 1862. On "OInt ne...· and litlle kllO...n Atneric""l AnUlli . Proc: . Aead. Natur. Sci. Philadelphia 14:151- 159. BERT HOLD, A.A, 1845. Ueber ...,~lt icdene neue coer !!C hene Repilien aU. Neu- Gra.nada und Crusteceen au, Cltina 1863 , On h at;hy"ph"I". , Scophiop us, and other Nacltr. Ge•. Wiss. Gotllni:C'n 1845:37-48 American Barrachia. Proc. Acad . S atur. Sci. Philadelphia 15:43-54. BoEnwl,O, 1893. Reptilien und B.atnll;ltier _ Yme zuet. Ber. Senckenberg, Nilturforxlt. Gn. 1893:3S-42. 1866. Fourth comribulion 10 the herpelO108Y of tropic,", America: 3, Notes on neotropical batracllian•. Pnx: , B de anfibios bra, ileitos . Servilro de [)ocument....ilo, sse Paulo. 183 pp . 1868. An namination of lhe reptilia and twra:; ltia ce­ tained by lhe Orton npedition to Ec,wlll' and lilt upper " P ")" JlObu f~ s" _. 1967. Nov... t.pc-des de do Its lt e Amazon....·ith notes 011 « hef spede•. Proc . Acad. Satul .lIde," brasildro (Anura, DendroNlidae). Rev. Brasil Sci. Phlladelpltia 20:96_ 140, BioI. 27:349- 353. 1874 , On some Batracltia and SemlIOgnathi brought Bo L"LE"'Gu , G.A. 1882. Catalogue of the Batrachia Salicnlia from the upper Amuon by Profe ""'f On on . Proc. s, Ecaudata in lhe colleclion of lhe Brilisb Muoeum. Aead. !"alur. Sci. Philadelphia 26:120-137. 2nd ed . British Mu.cu m. London. xvi + 'so3 pp. 1875. On lhe Balrachia and Reptilia of Costa Rica 1883. On a collection of fro g ~ from Ye rtmag uas, Hua. J. Acad . I"atur. Sci. Philadelphia. Set. 2. 8:93-154 111:1; :6~~";;3:o~m Peru. Proc. ZooI. Soc. London 18n. Tenth contribution 10 the: herpetology of tropical 1888a. l'o'oce on the cl ~ s i flC a t io n of the Ranidae . Proc . America. Proc. Amer. Phil. Soc, 11:85-98 zcor. Soc. London 1888:204-206. 1886 . Catalogue of lite specits of batrac hians and 1888b. On the ··nur;in i '·hbits of D~ ,. J'o bu fn , u ~ ptile s contained in a colleclion made at ~bas . upper oMen ed by A. Kapp ler . Ann . Mae. Nalur. Hisl ., Amaron . by John Hau•...-ell. Proc. Amer. Phil. Soc . Str .6. 1:454..0455. 23:94-103. 1898. An account of the rep il" and balrachian. cot. 1887. Syn0p'b of the Balfacbia and Reptilia oblained leered by Mr. W.F.H. Ro..enberg in wenem EcuadOl'. by H,H. Smillt. in the pnw ioce of MalO Grosso, BrIli!. Proc. Zoo!' Soc. London 1898:107-126. Proc. Areer . Phil. Soc. 24;44-60.

1899a. Description. of fit\\' batrxhians in lhe collection 1893. Second addition 10 t~ kl'lO'll'ledlJ:C of the Batrachia of lhe British Museum (Nalural History). Ann. Mag and Reptili a of CO,la Rica, Proc. Amer. Phil. Soc . Nalur. Hist.. Ser. 7. 3:273-277. 31:333_347. I899b. De.cript:ion. of ne.... replik. and balrachians 1899. Conlribuliom 10 lite hc-rpetoloJy of New Gr3nalb collected by Mf . P.O. Simons in the Andc. of Ecuador. and Argentina. Philadelphia Mus. Sci. Bull. 1:1-26 Ann, Mag.l'o'alur. Hbt.. SCI" . 7. 4:454-451. CM UM I', M.L. 1972. Terruo nahty and muting behavior in De'ld'obu l~ S It' O'lUJifN IlS (Anura: Dendrobatidae), Her­ :~~~:'o7;~' a~~I::: r~~~ l~~d ~~~ ~~.J~: pelOlogica n :19S_198 Ser. 7. 10:394-402. poca. J.W. ",so P. DU....S T. 1974. Coun .hi p beh....jor in 1903. On some balrachians and reptile> from Venezuela Colosfelh"s m/la,;. (Dendrobalidae). Copeia 1974:988_ Ann. Mag, l"alUr. Hisl., Ser. 7, I I:4HI-484 990. /976 FROGS OF THE ces..US PHYLLO BATES 51

DoSO Sl>-S.....OS. R. 1964 A "" '"' Dl:nd robal idx froa:. des<:riplion of 1"'0 ne", forms . ACI. Bio i. Vene z. P'o.u~,,,pfs ,ivrroe fro m V"!lC'z~ bI . Carib . J. Sc:i. 4;349_364. 4:485-489. HU utK"H , W. 1940 . Bei l ri ~ zur Kennmi, der Gattun, DUU~. ~~~mw~ ~ior in dendrobalid H ylou llu ( B r~ h}ee p/l.lidx , Amp/l .). Zoo!. An• . 131:11l-128 Du",ilIL, A.· M.·C . .."0 G. Bl l ION. lSI!. Erpe toloJi e HooGWOED. M.S . 1969. !"Oles on lhe herpelOfaunaofSurinam : genera le ou hisloire n".lurelle com plele de. repile. , 3. A ne'" . pedes of Dt .w.ro1Jenre iche: Ganung. Aqu,.,.. Mus. Zoo!. Unlv. MIChigan 151:1-17. Terrar . Z. 24:1_7. 1931. New fm,s fronl Panama and Costa Rica. Occ.... 1972, Fro, ch<: . die ihre Kaulquappen huckepack tragen. Pap. Bm ton Soc. Natur. Hist , 5:385-401. Aquaricn M. g. 1972:28R-293. 1933, Amphibians and replile. from EI V.lle de Anton. I"'STlTUTO R ~ C I ON " 1 DE DUENV0 1VIMHlTO DO AMA' ''' Panama, Dec.....Pap, Bo ston Soc. Natur. HiS!. 8:65- 79. ( IRDA) "NO COlt• • Madrid DU"''' T. P. .."0 J.W . 0 0 1.1:. 1915. Aggreu;"" bchzvlor in Colosl~t" 'u ( - p,.mlw,apfs) nJll"ns (An ur.:Drndro. K..YAu" S,O. , E.X. AuuQvuQI)!. Ao" O J.E. WU SJ('k balidac). Hcrpe toloa:ica 31:23-26. 1970. GaPglionic and eardi"" ""lion. of b.alracholoxin. Eu"'P"" J. Pharmacol. 12;10-18. FtTZISGEI. L.J. 1860. Die Au.beule dcr o.lCrrcic: hi....hen N . lurfO~ her an Sli u~t hie re n uoo Replilien wlihrcOO KEFUSTf-IN, W. 1867. Ueber .,inige neue ode. senene der Weltu m""g clu n,. Sr. M.je.lal Fre gaue Nova ra . Balf~h ier aus Au>tralicn und cem tropischen America. Sitzungsber. Akld, WI••. W'en 42:383-416. N""hr , Ges. Wi... GOuin,.,n 1867:34 1_.361. FoW1H, H, 1913. Amphibians and ll.'plile> Irem Ecuador, KlT..SAo ...O, J ,T . 1967 . Ob'ervali ons on the biology of Venezuela , and Yuc atan. Proc. Acad. Natur . Sci. Dm ,I","'",. pumiiiu Schmidt and D~"Jrt>""I" "..raill.' Philadelphia 65 :15.1-176 Giranl. M.S, Thesis. University of Southern Cal ifornia. 87 pp FUNUIOUSEI. J .W. 1956. New frog' fro m Ecuad or and >oulh...'eslern Colo mbi., Zoologica 41:73- 80. L." THAM, M. 1966. Capluring ".lIange creature. in Colombia Nil . Geogr. Mag. 129:683-693. G ..N'. C. 1960, NOle. on a he rpet ol ~k a l ro llecl inii lrip through dte !IOulhe:acern lowlands of Bolivl• . Annal. L.. URENT , R..1942. NOle su r Ie. procoellen. flrmisterne . Carnegie M ..... 35:283_314. I BaTr.eh 'a Anllt...). Bull . Mus . Roy. Hlsl. Na lur . Belgique 18(4l ):I _20 . G...",..s , S. 1837. We>! IOOi. bauachia in the MlI'CUm of Comparal ....., Zoology. Bull. Esse x lnsl. 19:13- 16, L.lDTH ee JUlDf., T. W. V..N, 1904. Rep:i.... and balrachian. from Surinam. SoIe:sfrom the LeydenM!I'eUm2.5:83-94 GIUIlD, C. 11 8~51 1 8504. Abotnct of a tepee 10 liul. Jame' M. Gilli». U .S . ~ .• \l(X'lllhe ~ l'Iilc' collected during L.VOOECU, H 1974 . Etholo,ische Unlersuchungen zur !be U.S. N. AM ...... D.E . 1971. TerTilOrial behavior in • ne«ropical P1n~/ob"'~J frog . Dmd1"Obalu K"'" ../ifrnu. Copei.a 197 1:365_370. L.UUSG. K.H . 1971. De. F3rberfrmc h bicol<>r Bibron der O;"d illera Azul (Pt-11l ) . BOnner Zool. Beilr Gos." ~~ b~Y~~ ~~ ·la-;.,:;m~,li;~edn ~~~e :~r i ~~~; ~li c :ror:. 22:161_ 174. Herpetologica 16:183-1 90. Lurz , A , 1925. B a l r~ icn. du Bri .il. Compl. R.,nd. Soc. BioI. Pari, 9.3:137- 139 GUI8E, J . 1950. Cal ,al o~ue de. I}P::' d'arnphibiens du Mu...eum Nalll'nal d' HIMOlre Nawrette. Imprimerie Natlonal e. LUTZ, B. 1952, Frogs in the Adolpha LUlz collection of Pari" 71 pp. the Instiluto O,waldo Cruz. Mem. In.,1. as...·aldo Cruz 50:Ml8-613 GUln'Eso T, A. 11148 . ,faun a chilena:, rcplite•. Pa,., . 5- 136 '" C. Gay, HMon a fislCa y politlCa de Chile , Zoolo,ia, LUTZ, B. "NO G , KLOSS. 19~2 . Short note. on some frop volume 2. In the aUlhor'. house . Pari., and Mw.ro de ~%: , t ::" ~ '&~a~C:: ~ng~_~~, vicariant forms. Historia Natural de Samiago, Chile. GUNTHH, A.C. 18~ 8. Car.alo,ue of the Bauach i. Salienli.a M.."". W.M , 1957. TIle wild animals in my life. !'Oat. Grogr in lhe:coI1ec1ion of the Brilish M"""um. Brilislt M...",um. Ma, . 111:497-524 lnndon. _vi + 160 pp. MAnl, F. .."0 B. WlTl>or. 1963. TIle wcnom of lhe CoIom, 1868. F...... acroun l of specie> of l. ilIe•• bamc hian, bian arrow poison frog PIr, I/obalrs lHcoJor. E.1.perienlill addocl 10 the collection of the Brili:$b Mu.eu m. Prnc . 19:329-376. Zool- Soc. London 1868:4711-490. MEU" . D. 1,941. Con lributio n. 10 the kIlO. ledge of lhe Am~u bi...of Sourh Amenu. GOIel>or~ KWlgl. vee....k. ~\lO~ ~~en~~B:::~~~~ . Vincrll. Samh. Hand!. ser. B, 1(4):1_71 Taylor and Franc i•• U>ndon. MIR..ND.. Rt Rt.llO , A . DE. 1926 . Nora. p. ra ",rvircm ao HUTWOl-E. H.. H, 501""'0 .."0 A. HE"no-UU. 1965. NOle. .,..ud<> """ gymnol:>.amochim (Anura) h"a.i tem:... Arch on amphibian. fm m Ille Venezue lan Guayana. ....ilh M... . Nac . Rio de Janeiro 27:1-227. " BUUETIN OF THE NATURAL HI STORY M USEUM OF LOS ANGELES COUNTY Nr> _27 MUDlt"("K, W. 1% 9. Pf1 ~ K e und ZUdll einn Blau' ICijlu , Sr;HLUJE!- . H. 1826,: NOTice sur l' e.pelolo,ie de rue de fTDS(:hes de. Oal1un~ P")'lIob of the MII'IdcI Amcrica. Amer . M",", S O'l'iI. 29:1_7. Venezuelan nped"'"". F.dd MIlS. Nll ur. HUI., Zoo!. 18:1S9_163. 19231. Ne.... b.lracllians from the: trop ical resear ch .rarion. Brili .h Guian• . Zoologic. 3:289-299 SCH" IDT., K.P. A~lI R,F . [NOU . 19S1. Ampllibians and reptile s of the Hopkin ·Br . nner e ~pedltion Eo Brazrl. 1923b. A new gekkOIlidliurd ...d a new bra<.: hyc<:phl.lid fro, from Colombi•. Amer _Mu• . Novit. 88:1_3. Fieldiana: Zool. 31:439-465. SOloJiKben 1926. The peo;1OnI 8in1~ of the bncb)'Cephalid hop. Cabinets !II Krak.u. SlIlun r ber. Akad. Wi••. Wien Amer . MU$, Nom. 23Q;1-14 24; IQ-U. OlItTT~~':::)I '::U'::~:::r ?r.'f~:r~,,:, (D... Erdbcer. _. US8 ~l icj;)C herpclOlo,icae mu-,.ei zoolOlici craco­ viensi.: un,escll"'iinzlen Balrachiel . ~nk sch f . Akad. _ . 19S3 . Nechmals Dr ndrubaUJ rypogr Bru!gn cllaft des B aum'lei~ erf rosche . OEsE . , R . 1932. ~ ZUehl des B.I,l!mlcige" (lHndrobd,u IDrndrobatrs. " ,.,al ... Girard ) in Gefangenschaft. Zool. l~iIUJ . BUillCf Aqu• . Tmarimlunde 4) ;196-200. GIIU1l Lelpll' 8:122_ 131. PUUI. H.W. 1935. The frop. Iiurds. IIld snalll _. 1939. Urn da ZUelllproblem ...... lHtttirolKJlr. " ",aI... Guiana. Pnx. Zool. 50;. London 1935:505-530 G. Wochenschrift AqIW". Temrienblnde 36:7_9. 23-21. Pu...oc.... M,G . 1904. Rcuili cd Amfibii, Boll. Mu•. Zoot. S !X TO~ . Q,J , 1960. Some a'~·l ' of the behavin. a nd of Anal. Compo Univ. Torino 19(465):1-4]. lhe lerTLl0 'Y of a del>ilrobattdfrog. Prn."ltt,,,pi.. ,,mUdlJS Ecology 41;107-1 IS Puu s, W. 1870. Eine Milt!leilunJ ube. neue Amphihien (Hr"'idIXryl...... ! des ko'''C bch wolol; ~ben Mu. Sll.VEIlSro"'E. P.A. 1973. Ob",rv ll ions on the behilvior and lOe~. MiI . Munich. 29 pp. PoLDU . W.S . 1973. Over ~erzorxin , en ~ OO"plan l int in S TU/ll~ S . R. A ~ D J. HE"'OUC"50I<. 1919. Field .. udies of amphibians in Colombia, 500111 America. uev. Cali· ~;;:~g~~dh:'~a ~~~e~H:~~;~~i~':Z:;:~'::_2e2" en ele fornia Pub!. Zool. S6:497_S40. 1974a. O~ r ~ erz or l;in ll en ~ OOr1p lan'in , ill ge~ . n . ST F.lND Ar; H~u . F, 1864. Balrachologi sche Millheilungen. Verb. 7..001 . BoI. W . Wien 14:239- 2118 . &,~:: glndH~;~:""': : Ie;::,:~~ anden: 1869 . Reise de.- o.1erreichi"c1oen Frepne Nov...... urn 197"b . Over ~rzo r,in, en ~ oorrplanli n . in gevan . die Etde in den Jahren 18S7, 1858. 18S9. Zoolo~scber Theil. leo Bmd , 4:Amphibim . K.·K . Hof· und Sl....· ~'::~!~ fJl~H::'"'~~':;'';,,''':: 3e;4 ~)~ .1e .ndere druckerei. Wien 197":. ~ nd rob al i d a e (4) . Ch-er ~e rzorg ing en ~O<)" . TAYLO Il, E.H, 19S2, A review of the frogs and toads of plannn g In l;e ~angen sc h.p ~an Drndr obid from the Colom· sobrc d i ~e n.. lem• •. Imprenl. Of", ia!. Medellin. Colombia bian IITO\V poi""" frog. Phyllobalr. ""'....ar".... and parrial »T1Ibe.i. of b;mxholo ~in and ilS .nalop and Rlvuo . J.A. 196 1. Salienli. of VenezlIl:l•. Bull. M... . Comp o """"""". J. Arner. Chern. Soc. 9 1;3931- 39311 . 7.001. Harvaro 126:1- 207. T!K'HUlIl, J.J . Vo "'. 1838, C las. ific' lion der Balracllier . RUTHVE"', A.G ,AND H.T. GAIGE. 19l5. The breedi ng Pe!ilpiem:. Neucllatel. 99 pp. h.bi" of Pro..,ht,api.....bp..ncl"I ... Cope . Occ... W ~G l.n.. J. 1830. Nalijrliche. Sy.. em dc. Ampllibien miT Pap. MIlS. Zoot Univ. Michi, ... IO:I- S. mrCll~,;,,~~~~~~iiu" hiere ond VO~1 SAGaA , R. l>E u . 1&41- 1843. Hisioirr physique. polilique. ee Mlurelle de rile de Cuba. Alias. ~pl i le •. Bertrand . W"'DE NIEI . R.l .A. 1973 . Venl., van bel kweken mel Pm., 31 plales lHnd,oba,tJ ,,"'Qt..... L1«rta 31:167- 171. S~V A OE , J. M. 1968 . Tile dendrobatid fro~ s of Cenlral WASSEl<, S. H, 19~7 . On Drnd,,,""u $.f.o, ..po i ,,~ n material America. Copei. 1968;745_776. among Emper.ii (ChoOOj" l"'aklllg IndIans III western / 976 FROGS OF t HE GE."'US PHYLLO BATES 53

CltJ da~ . Colomb ia. Arm GOl ebor,~ Elno" , Mu•. W'EI>-S n.-"",W . .\IA Zo o 1821. Rei", na: h Bfa'i liell in I9SS- 1956:73_94 den 1 ;Oh~lI ISI S bi. 18 17. "Olume 2. Heinrich Brommer. WlI. "U. F. 18W . Ueh:rRcpilien unciBltlnChier -. Colum· F.-Hun . >vi;; .;. 3-16 PI' biotn lIl'Id Trinidltd. Vem. ZooL BoI. 0.. •. Ww:n 49: w n n , G.·F. 010 . 1930. LiMe do:1. Repik-s et 8.lI , .., io: n ~ ,"'-'" ,.xoll ( ~ au B,t> il par b. M;", ioo Ma>-urt (1922- 23) 1916 . s"mcrhn,en iibe. eini,e nieden: Wirl>ellien: CI dt-..:ri plion de "" pl llOlfVel~ e~~ . Pa,e> 21 3_2."10 de. Anden von Kolumbie n mi, Be-.<:hreit>un ~n neue, ill Une mi~,ioo bioIojtiqllC beJjlC' aU Brbi!. "Olume 2 Anen. Zool. Anz. 17:301_304 B ru> e l lc ~ Accepted fur pcblic.uion Apr;1 1.\. 1976,