A REVISION OF THE POISON-ARROW FROGS

OF THE GENUS PHYLLOBATES BIBRON IN SAGRA

(FAMILY DENDROBATIDAE)

By PHILIP A. SILVERSTOi'E

NATURAL HISTORY MUSEU~l OF LOS ANGELES COUNTY

SCIEl'CE BULLETI!" ~ 7 • :-,:ovEMBER 25. 197b

A REVISIO:\, OF THE POISO:\'-ARRO\\' FROGSOF THE GE:\'US PHYUOBATES BIBRO:-': IS SAGRA

( FA~lIlY DE:-':DROBATIDAE)

A REVISION OF THE POISON-ARROW FROGS

OF THE GENUS PHYLLOBATES BIBRON IN SAGRA

(FAMILY DENDROBATIDAE)

By PHILIP A. SILVERSTONE

:-<ATU RAL HISTORY :\1USElJ~l OF LOS AV iELES COUI\TY

SCIE!\CE BULLETl I\ 17 • :-;OVBlB ER ~5. 1976

T AB LE OF CON TE NTS

AC,,"OWLEDGMESTS

~hTUI"LS ""D ~l ET HOOS

~u-eum abbrevialion$

PNlUJJA<1a BIB_ ,.,S..u...1841DdinilionDi a~nosi s

DescripelonConten tTad polesSpecie, group'Life historyFoodSummary of distributionKey 10 the genera of DendrobatidacKey 10 the species o f Ph.l'lIoh/ll'-f

8K."OW~ (ilt 0\Jl'

Phylloha tt!_, ,pec iesPhylloM tt!s aurorut!nioPhyllooott!Sbiro /orPhylloha lt!S lugubri sPhy/foha tt!s ,';IlOlll.S

F EMoifAUS (iltOliP

PhylJo M u s a'll ltwly';Ph)·fJobou s bol'lt!ngt!ri ••Phy/Jobatu t!spino so;Phy/Jobatt!Sjt!mo ralisPhyllobau s tricohvPh,.llobau s zapar o

P O f!S (; 10l/P'

Phyllobat t!s bol Mu lllUPhyllooolt!S ingt!r;Phyllohult' J pan'ulusPhyllobal t!S pet ers!Phyllohules plctu sPhvltobates pulrhripeclu .fPhyIJob/lles ,marulld;nus

T~lVmA I/JS (; IO\; P

Phyllobalt!s bault! riPh.l'lIobalt!s triviuOl,.s

U TElt U l,I RE CnED

3

JJJJ,,77,999

2020212J24zs

2727aa2950J2JJ

as

""J6J7aa4J44

"""50

FRONTI,PII:n: II. Left side. lOp10 bonom: Pilyll"b<i'e," , pe.:ie , . km I~<) . mad bclwc<:n Tingo :l4atia and Puca llpa. Cordille ra A1.UI.Peru; Emerald Poison·arrow Frog IP. .<n",ra~di"ujJ. holOl~pe. Pan de Azuear. Peru; Peruvi"" Poi,on .armw FrogiP. ~lerj;l. holo·type. Sanla lsa"" l, Peru. Righi . ide. 101'10 bon<>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(lrali'l. SaUlTonue. French Guiana; Beauliful.brea'led Poison·armw Frog IPpulchri~..,,, .,,/ . 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:-iUSPHYUOBATES BIBRON IX SAGRA C FA~lILY 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~ inhabil the A""""", d...u.a"e.

TIle di. linction bel.. eu Colo.,..'.... (dor",m d ull~' rolorcd and .....in lO~in. prc_umably ab.cnl' andPII.d 'obu,<'J (dorsum bri,h ll) rolorcd and ..lein lO~irl, I'f",,,mabl}' P"'''''''' .. maintained. Fing.... Icn~'1h andfinj:C1" di'k size~ 1I>Cd hc~in 10di...in,"i""~lIJrob<l,... (fiN finlCT <hnft.... than 'ottOI>d. ""d fin' ....di.hI'dal;"ely larj:C ) from PII) /Iob<1l<" Cfir>! finl!tt equal lOor IonI'C'" lhan >«und . and finrcr 4i<J;s n:1", r.d~

......11) in' lead of lhe uadilionalchanc ier. absence lD..nJ,obu,<'J\ or pn:".'~ /.P."II,>b.I,... )of 1<'Clh. allhoughlhe IooCh charac1C1" i, ill u",f" l. U,in, finger knglh and finger di..... 'ize a' a dcfinilion. fl\'e , pecic' (b<Jul..ri .;"1("';. fJ<I" ,.. IIU, pic, and ";vi fl"' '' ,') Iran~fc' fromD..nJ,obu,<'J10 PIr,·lIobul... .

P. ) lIohal<" include' four , pecic, gr<>l1l"': (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,pilloo<li . P.!, m'''"Ii.•. P. I, ico/" , . P . "Uf'tl'o ); (3) pi"/I' .' group ir. b<J/i,'i"","" P. i"x r , i .P, f'tl"""I ,,.<. P. 1'<'I<',.<i . P, pi"'", . P. p"l<-h, il'<'('/"". P . ""' '' ''' I(di" "s); (4) r,i";II<lI" .' group (/'. I><I.../rri .P, l,i,'ill"tuS ). Four new Soulh AmCl' ican ' I"-"Cic' are <kscrihed herein: I'h )'II<>I",, <'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","'" " , andCoIOlI<'ln"s . Phyll" l>aln u,ually ""ip<"il' Cal lea'l in """ 'pecic•• and I'l'ohahly in all ,pecic' . WilhoUiamplc: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" ,ilhe r the mak 1"''''11'1 01'lhe km alcparenl may carry lhe ladpole, . Tl>cmain food of Ph l/obm... i' 111>1, .

Ame, indians e ~ ltact blo....gun poi"'" from al ka" 1" '0 'P'""ic. (P. "" "'I",n;,, and P. mcolo, ).

I S 1 RO[)l",'C1IOS

A, p""viou,l y d i-.:u....-.ed (Silver'>lone 1975 ). lkrldru·batjd 1Cnera are difficult to define . The ~enu, Ph).·//t>·

"'u... previou sl j- ..."" dia~n"""d by tbe prese nce ofbright oo l"1>;ll co lorat ion and the pre-e nce of maxillaryand prema xillary teet h (Savage 1%8) . Because theprese nce of teeth "aries with ill o ne 'pecic, (pi ,'/iu)

a, well a' betw een 'evera l oth erwi se >'c ry , imilar'I',:e ie , (h,,f ; I·i<lllus·pel ,,, s' ·pl<lch"I'''·II<.I: l" IU/" " ·I';·1'111,,' 1<,,), I have redefined the genu' nil the h'lSi. offinger dis k size and the relative length of the first and>C~" lIld finge rs. These finger characte rs val')' ,""K"' ·da ruly with teet h in a ll hut fou r species (/.,,/il'imw.'.p iC/u.•. 1'" IC"h,ipt" cl"" . and Ir; l·iIlU'U I . ....hich u, uall )'lack teeth) , ....hich here are co nside red Phy//" rnll" srathe r than D.."drvbales, I have transferred t Selve r­" one 1975) thre-e addit ional spece-, (bun/rri. i" x.., i.and /'<In ',,lu .l. ...·hich have teeth) 10P.~-Jlobul..S on theN , i, of both teeth and finger charac ten.. I ha..... con­,idc red a' CoIosI..,hus aU dull y colored toothed _pecic,.although , i, species ( the [ .._ ".ali. grnup) ...ith "rightco loration and basal toe lI'e bbing . included he~ inPh...//oI><I,..s. rna)' be <If>do~ly retated to CoI<lsl..,hws"" the)' a~ to the Ol:her ..peo:ics of Ph,-lfobtJ' ..J .

Th u, Ih i ~ revi,ion of Ph\'l/ol>a,..s include. thosespecie - havin~ a oombil'liltion of three chariICten.: ( I ),mall fln~e r dish: (2) the fiN fin~"""C(juar to o r lon~er

than the ...,;ond; (3) the do r, um partly or entirelybrighlly co lored in life. In my op inion . the ~'(: nu~ Ihu\de fined is mere natural than il ·...·a' under the prev iuu ~

definition . Ho wever . delineat ion "f convi ncinglymunu phy lc l ic genera of dc nd rob atid s will req uirecon, idera hly more dala.

A (" Kro;OWI ,EI)(; MENTS

lthank the followi ng for <litJ or the 10<111' of specimen s:J .W . Wright , R ,L . Bel )' . F. Trux<l l , D . Whi,c le r,R.R. Snelling (LACM); J. M. Savage. P.H. SlarreuICRE): A,H, Brame . Jr . II. McCu rd)' Xature Ce nter:

' EoI10 10 U CO Mll rTTH FO. IHIS B ULU TI"R OIlH T L. B .... y

W. RO" " I D Hn uRoy W. ~kOl"""lD

WOl-'U S . PoL!».JoH" W , W . I" HI

'RC'C~h Auoc tolCin Ht'l"'l<Jlogy. :O>alutal HiSl<X) "'\/>Cumof l.oo An",1n Counly. 900 E~ poJoilion 8 1vd ..~ Angeln-.Calif<>t1'l;a 9(0)1 .

2 BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY No. 17

J.P. Bogart (lNHC ); C. F. Walker. A.G . KlugeCUM M Z); R.G. Z..-eitel. C.W. Myers ( A.M:" H ); J.E.BOblkc. E. V. Mal",. \( (AS SP); A.E. Leviton (CAS);l .A . PC'lers , W,R. Heyer, G. R. Zug . R.G. Tuck. R.Crom bie . S . CamJXIen-Ma in (USNM); w.e. Shcr­b....lke. R.W. McDiarmid. Univcr.;;I)' o f South flori da:E.E . Will iam~. B. Shreve ( MeL); \\' .E. DI.IC'Jl man .S.R. Echurds. 1.T . Collins (KU); C.l . McCoy. Jr. •S . Richmond (eM); J.D . l ynch. University of Nc·bravka; A,C . Thoresen, Andrews University; D, Travis.Manhallan Beach: D.B. Wake. R.C. Stebbins (MVZ):D.B . Ekken s. Lorna Lind a Un ivcNily: A . SIam:II .Calirorni.. Sta te Vnwen;!y at r.;or1 hr id ~e : M. Tye .California Stale Un;w,rsity at Long Beach: R,f . IngeT,H , M arx (FMN H): J.A . Rivero , Un iversidad de PuertoRico , Mayaguez: InslilulO Ev.mdro Chagas. Betem;Induslr;a e Comercio de Minerios. Serra do S avio:R. Pradinaud and Dr. Rivieeez, Ca)'l"Tlne: TeKas InSll'\l­menls Co.: K. Adler. University of Noire D-ame; O.R .da Cunha (MPEG); W.C.A . Bokermann. Fundal;anParque Zonl,igiC<l de San Paulo ; MOMieur Ie Prd el deIa GUy'ane; C. serie and ....aff. Instilul p.,,<-Ieur de Ia

Gu)'ane Frarlf;ai<,(' ; Office de la Recherche Scienlifiqueet Technique OUlre-Mer; Rdo. Hno. Nicetom Maria .Rd" . Hno. Daniel (IL'\ ); F.e. Lehmann V.. MuscoDep artam enla1 de Hislllri a Natula l, Ca li ; K. v" nSneide rn. Mu"Co de Hi'l nr ia Natur al. Universidaddel Caec a. Popa) in ; C. Escohar. Cali; J .I. 8<lneroH.• L.S . Espinal T.• R. Dial V.. H, Trapido. J.E.Ramos p.. Univer, idad del Valle . Cali; S.M. Frank)'V .• J. Hernandez C.• lnvtiturc de Desarrollo de losRecuoos Natu...les ReOO\ab les US DERENA). Bogota:C .I . !obrinke1le. Universid:>d de k>s .....ndes. ~ot;i;

B. Cboebe n , Centre Nalional de la Recherche Scien­nfiquc. Par is; J . Lescure, J. Guire tMHNP); M.SHoo j!moe d (RMN H); W.N . Pold e r . Alkma ar; B.Hudendid (NMG); S ,H. W"-s..cn. GOteborlls Etn(}­gra fi' ka Mu-ee t; A .G .C . Grandison. E.S . Arnold( B~ ) ; I . Ei.e h (NHMW); G.-F. de Wille IMRHN):w. Hellmi<; h (ZSBS); and I .H. Haffer. Te heran. J.H.Haffer gave advice on the mute of access to the Altodel Buey. J. Le-cure pruvided a Ii.... of Iocalil~ for1.G 'JlCCimens. inciudinll ihe flf'! record of P h.\# "ool<,Jp;rIUJ from mnch Gui..na . ..... .G .e. Gr.mdi....o J'N' idcdcoord inates for the tocalnies o f P. !>"Ii,·;UI'''.'. P. Br"medrew the tadpoles . J .T. Ktraeako drew thc master out­line for the color paac rn diaj!r"ms; I drew the patterns ."fhc, '>Uperb fmnlis pi«es arc the ...orl of J. "' akanishi." aff anisl of lhe SalUral History Muse:um of LosAnj!e les Co" nty. Pa intinj!' for the fmotispic<:es werebased on color slides by J.A. Pelcrs. R.W. McD iarmid.J.D. Lynch . D.B . W..ke , A.C . Thore-en. and me.SlUdie- were p;>n iall)' suppon ed by a NSF Summerhllowship for GrlOdUalC Teachin!! A",si" an" (1966) .by a NDEA Tille IV Felluw,h ip ( 19M· I%9 j, try Bio·medical Sciences Support Gran t FR·070 12·02 # 570J

from lhe :-.ral ional Institules of Heallh to the Universilyof Southem California ( 1969) . and try lhe Los An~le,

CoulMy ~ use:um of r\at u...1Hislory Foundation ( 191 1).

MII Tf ~l lI t .s liND Mf.THOUSAll nlCil-surement, are 10 the nearest one·half mil­

limelCr.TlOdpole lIlages arc those of Gosncr (1960).O' ioolo!!- ical , pecimen, were cleared wilh glycerin

and lwo percenl pola"ium hydro~ i de and were stainedwilh alizarin red S.

p."nermo and subpalleTn\ in the aocounb of P. piC/in

and P. " i' ; IIOI/(.. <Ire for cOll\-'Cnience of descriplion:lhey are m>( intended 10 represent lau

In the species ",:counts. under ine headinj! "stomachconlenls. " the museum number of a frog is ciled first.follo'A'ed by lhe names (afld numbers of indj,.·iduals )of the il1\1n1ehr.lle tu a found in its 'I<.>mach.

In the synon)·mies. a period separates the name ofa taxon from the names of a ll authors CXCC pl theori!!inal aUlhor.

Under the headin!!- " doublful records" are localilie1.that are separated by a major ~graph ic barrier fromall other rol leclinj! localil~ for the specie«. or thailie far outvidc thc known e1evatio nal range of thespecies . The<oc doubtful records "rc not included Onlhe range maps: lhey are included in lhe Iocali!)' Ii""10 brillj! stem 10 the auemjon of future collec tors.who may be ahle 10 verify them

Subspecies arc not used . bUI I have indicated to

which popt.a lation' subspccific names apply. because:they are available as , peciflCor o;ubspcciflCepilhets.

It is difflCUl1 10 define species of Phylloball"J be­cause of intra.specific varialion . interspecific resem ­blance. and a paucity of kn"wn cases o f eympatry. Todefine spec ics. I used pre'encc or absence of the fol­lo.. in!!- char.JClers; toe .. ebbin g. Icclh. a calf·"P'>(. amedian dof'il l ...ripe. p;lirell ...npe s , and do rsa l orventral marbling: I aho used positio n and Icn!!-Ih o fpairell ' tripes. snout -vent length. , kin texture . andcolor. When so me of these cru.racters wcre not sharedby allop<llric populatioos . I Irnldc: <kcision' .." 10 con­,peciflCily by wei!!- hling certain C....rac1C1'S lhal J con­. ille red imponanl in Ihat panic ular case . bas ingdeci, ion, on the known ran g.c "f variahili ty of thecharacter in both popt.a lal io ns. and on thc pre-e cceor "bsenc e uf morphol.>t: ically intermediate popula­l ions . I h"ve been co nservaove in dcscribt n!!- newspecie,.

The spec ie- of Phyll"ho l<' . are "rranj!ed herc infour prov isional spcc ies j!m ups. T he ' e groups arcintended 10 represent relationships , bul can do so o nl)·imperfectly h1x..u-e of the incomplelenc" of ", 'ailabledata , Tn arran~ the , pecic- in groupe. I used presenceor "b<.ence of toe we bbing.. position of p~ i n:d '1ripe~ .

snout-ver n length. and re-cmblancc of cnlor p attern.

/976 FROGS OF THE GENUS PHYLLO BATES J

KU

u..CM

IPBK!'>l

wCAB

19Z6. PIt,.lIob<;tr. (pan). Soble : 91 9~6 lHordrobertn (part ). Sobk: 9.19S7, Phyllob.rlr . (pan) . DlIJIn; 77-7M1968 Phyllohalr. Sevege: 747_7411.

Defi nition. - MuiUary and pr-ema~illM)' teet h pr-es·e nt or abse nt; di sh o f seco nd. th ird . and fou rth finge~

relatively small (Silverstone 1975:fig. I); first fingere4ual to or longer than second; toe webbing present or

absent ; in life . •Lin o f do Nim pattly o r emin:Jy brightl ),colored; pala tine bones absent ; omostcrnum pre....m;vertebrae usually ' not fused; tad poles (Figs , 10-1 2)with oral dis k laterally inde nted . number of demiclcsnot reduced. and anus dextral.

DiQgnm is.-Phyllooof.... d iffers fro m Dendrob«tt .in havin, ue fiN finger eq ua l to or lo nge r than theseco nd and in hav ing relatively ,mall finger d ish , In

Ph.l'lIob<lI..... the dish of the second 10 fourth lingersusuall y are Jess than I .S limes the widlh of the fing crs ,In Drndm oott • . lhe d isks o f the second 10 fourth fingersusu a lly are al Ieas l 1"'lo;e Inc width o f lhe Finge rs .Ph y /l"hrtfl'$ diffe rs from C"I(>$I(' l hui in having the skinof the dorsum partly or enti rely brightly colored in life ,

anti I pre cuma bly) in havin g ,Lin to xins .DtJerip ti" n . -The snour.vem length o f adu lts i.

IS 10 50 mm (Table I); female, avera ge lon ger thanmal es . The skin is smooth or gra nula r Ma~iIIary andprema xill a ry teeth a re present or absent; vomerineteeth are absent. Adu ll ma les have voc a l slits and

u,uaU)' have (at lea.' t in prese rvariv...) ;a pa ir o f lonil i·tudinal e xterna l fo ld, o n the throat ; fe ma les lacL voca lsli l_ and e xte rna l folds on the throat. The lip of thesnout is trunca te 10 slightly rounded in dorsal aspec t.The imercrbital distance is wider tha n tnc upper e yelid .

The diameter o r the e~e is greater than the di'>lancefro m the eye to rbe nos lr il. The pocerodorsal por1ionof the tympanum is conc ealed . Each digi lal disk beu..on its dorsa l surface a pair of raised " scutes" separaledb)' a median groo ve: the seco nd. thinf , and fourth finge rd isks an: small rela nve to lho<.eof D('nJ" >bultJ lSih·er·slol1(' 1975 :fig . I). The dis ks are not sexuall)' dimorphic ,T he first finger is elJual to or longer than rfe second.The fing et'!> lack fringes. and well-bing. A subartkular

tu berc le occ urs on the vemral surface of eac h di gita ljoint. An oblong inner :.rnd roo nd outer tube rc le occuron both the me tacarpu s and mc t:.rtar , us; .....meli me s amedian tubercle occurs on rbe ructamrsus. A ridgelikcfold e xte nds alo ng the d isla l po rtion of the larsu s; the

pro ximal end of lhe fold cu rves . loOtnelimes bend ingat rlil ht angle, 10 ue re'l o f !he fold . A tarsal tubercle

oflen projects fro m the prox imal end o f the tarsal fo ld ;wit hin a spectcs. the la rSiIi tubercle mu)' be ubscm orslig h.lly 10 well devel oped . T he toe s are eitber weblc..or bot ....JJy ...ebbed luwally between loe s II·III · IV).

In life , the ir is is blac l Of brt"" n (rarely bronle)and is neve r reticulated. Brighl co lors (red . orange,

Th e foll owing ab breviat ions ind icat e tbe loc at ionof preserv ed specimens:AMNH Amcrican !'>lus<:um of Natur al Hi, tury, Ncw

Ym'AciOkmyot :-;;Ilura] Scim;(,., Phil.xlphiaBrili'" Mu-,<um (:-;aluraJHi>tory). LoodonCali fornia Aude my Qf Sc ien~•• San Fu.n.ciscoSlanfor<l cQtlcetion al CASCal'J'lc.gie Mu,,"um. Pill, "'nghUn;"..,nity of Swthcm California........ Angcle>.(~ Rican EJ.pcdil~)

Field Museum of :-;atural Hi>tory. ChicalloG . On.~s V. collect ion 31 USSM[n,tiluto de UI Salle. BogolaJ .A. !>clen collection at US:>i M1.M. SanS" priute O>leOlogical coIl«tion aI

CREJP. Bog;onooll«tion at Th HCKrakO'" Museum . Ja~e ll o n i a n Universit y,Krak"w"Iuscu m o f :"O alural Hi. IO' )', Univeui ty ofKan..a. . La" renc..:"Oalur al Hi. toT) " lu!>C um of to. Angc l...County. to. Angel....J. i.e, cure private colleclion. Pari,Museum of Conlparativ.. 7..oology. HarvardUniven il) . CambridgeMusco de Biolollia. Univenidad Central deVenelllC'la.Car""'bMu..cum :"O alioMl d'Hi""ire :-;aluf!'lle. Pari,Mu....u f'araense Emilio (;oeldi , Belem[n"itut Roy al des Sdcnce. Natur ell ... deBelgiQIIC' . Bru••elsM.S, Hoogmoed roIl«lion al RMS HMuseurn of Vmrbratr L.>loJy. Univcrnly ofCalifornia, Bend ..)Naturhi'lnrisehe . Mu><:ul11. VicnnaNaturhisloriska Museel, GOleborllRijL;mu ,,"um van !"aluurlijke Hiuorie . u-idcnl ln;,.......ily of Tu n . Au"-inlinivenily of Michigan. Mu,,"um of Z"O~I .

Ann ArborUnis'ersily of Pucn" Ri~o, Rio PiedrasNational :\lu....um of Natural Hi. tory. Wa,h­.ngtonW.C.A . Bokermann private colleetion . Sio..".W.e. Sllr rbroole ro lleclioo al US~MW.E. D~llm~n collectio" at KUltlo l o~isc he Sammlu nll de, Bay'et ischenStaale>-. "'unich

A~SP

BMCAS

LGacz

MBUCV

P..l~H

GOVU.slAPJ~IS

CAS·SUCMCRE

MH!'OPM"'GMRHN

UPR-RPUSNM

wcsWEDZSBS

~HMW

NMGRMNHTNHCUMMZ

M u>cum a~vialion s

PhylloOOIl" Bibron in Sagr a 11141

Poison-arrow FrogsFrontispiece s 1·1[ and Figu res, [- 12

MSH"tVZ

1M !. Plt.I·lIo1><lln Bibruni" Sa, ,,.: platr 29 bi• . Type "J'C"'ie.PIt)'I1ohmts bieo /or Bib",n i" Sagra t841; plalc 29 bi• .

11<41. Phyll oi", u .I. Dumo.'ril and Bibro n; 637.111112 D r ordToI><ltn (panl. Boolenga: 14~

1 81l~ . P/tJ-ih"' tn (pan) . Boulen~r. 19-1IIIIUla.. Ph)'II0b0tn ( part ). Boulrnrer: 2Ot>.19W Phl# oI><lrrJ ( part). Ba.rbour and :-;...ble: J98-402.

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.rderror (;:±SEl. .00 ,t""dard devialion (SO)

All S[" cimen, Males Females

Sp..", ic, " Range , :SE SO " Range i :SE SO " Range , :SE 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- 2 1.0 20.0 :0.17 0.75' pecies" " .~6 0_42 . 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 0.27aurmac"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 2.49hi"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 1.39f>oli,·j'IIjus , 23,0-2().0 24.8 ±091 1.58 , 23.0 23.0 2 25.5_26 ,0 25.8b<m/c"x cri 15 150 -2 1.0 17.7::: 0 .25 1.49 " 15.0-20.0 16.9:0,29 1.23 " 1 7 .0~ 21.0 11I .5 : 0, 31 1.29

cW""''''' S 16 0 - 17,5 169±0.22 050 S 16.0- 17.5 16.9 ±0.22 O. ~O

fcm"",'i. " 20.0 - 33.5 26.6 ±0.43 2"" ra 200- 31.5 25.8 ±0.69 2.93 25 230 -33. 5 27.3 ±0,51 2.57

"'lie" 9 21.0_27 .0 23.1 ± 1.J3 2'" , 21.0 21,0 , 220- 27.0 23.8± 1.59 2.75lu;:u/>rls 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 15p<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 () . ~ - 29. 5 2.'.2 ± 040 3.29 aa 19.5-31.5 24.4 :0.34 3.10p,,'chri fNC/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~ 0.79sm,,,,,~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 0.97zap"'" " 26 .~ _JO.5 27.9 ±O ,29 1.22 , 27.0-30,5 27.8:0.56 1.37 u 26.5 -30.5 28.0: 0,34 1.19

"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- 2 1.0 (mean 19.8); nine females 21,0- 23,5 (n'can 22.8). P . ,·;,,"' u. ; njne males22.5-26.0 (mean 25.0): 10 fem.1cs 26.0 -31 .0 (mean 28.2)

•

~~~

~

~~

~~

~-e

§.~~

§

~8~

~

~

1976 FROGS OF THE G£\'US PHYLLOBATES 5

yellow , green, blue, and intermedia te hues) o' uallyform dorsolate ral or latera l stripes on a black or bro..... nground co lor. bot in some ,pecies, bright colo rs coverthe entire dorsum or form dorsal marblin g. The venterusuall y is black with brig ht spot s o r marbling . Inpreservative , the bright colo rs become: whi te , gray,brown, or black, and the coke pallem may becomeind iSl ing u i~ habk from the ground color . The surfacesof the mU!iCles arc sparsely to densely flec ked withdark pigment. Pa latine bones arc abse nt. The or­bitosp henoid region is partially or com pletel )' os..sifiN.The ornossemcm is present. The vertebrae usually arenot fused .

COil/till. -Phyllo/J<I /t's includes 20 species, occ ur­ring from Costa Rica to Brasil (Figs. 13- 17); Ph.l'"o­ba It'S spec ies (to be dese ribed by C .W . Myers andJ . Daly) . P. on/hOlt)'i xoble 192 1. P . oumlot'n;o

(Boulenger 1913) . P. bout<,ri (Meli n 1941). P. bir%rBibron in Sagra 184 1, P. holMan," (Boulenge r 1902).P. bout<,n~<,r; (Ba rbour 19(9). P . npinosoi Funk ­bouser 1956. P. frmo ralis (Bou jenger 1883). P. in~<,ri

(Cochr an and Go in 1970). P. /u[Jubr is (0 . Schmidt

1857). P. fl(J",ulus (Boulenger 1882). P . 1"<'1<,rs; IIC'"species. P. pi..ru., (Bibron in Tschudi 1838). P , pul_ehr ipt'(' /u.f new specie" P. smaragdm us new spec ies.P. mcotor(Bo ulenger 11199).P , Iril'iIlO Ilf.\' (Spix 1824).P, I·i llalu,. (Cope 1893). and P. Wptlru new species.Table 2 shows all species and sub'flCcics desc ribed asor referred 10 Phyllo/1(Jln .

TaJpo l<'s (Figs. 10-12). -Tadpole measurementsare in Tab le 3_ The tadpoles illu, U;Ited herein arcidentical in the follo"'i ng characters: Tbe oral papillaeborder Ille entire po'ter ior l ip. but borde r onJy tilelateral por nons of tile anterior lip . Tbe oral disk islatera lly indented . The den ticle formula is ~. Thebeak s are thin (moderately thick in P. ~iIlUl u_,) andserrate ; lhe lo wer beak is indented . The eyes are don;aLThe spiriICle is sinistral. The anu' i~ <;Ie uraJ.

All kno..... n tadpole, o f PII)'1I" OOI<'s have roundedo ral papillae ucepl 1'11)'110001<'1 sp«'ies (10 be de ­scribed), which has poinlCd papillae. All have one ro....of papillae ucepl P . l ·i,faIUS. " 'hich h," two rows . Allhall: the taiJ and fins mottled Jight and dark brown .e~cc pt P. \'illal"s. which is entire ly black . P. ont"'myi.

TABU 2

T'hc 100 .pee ie, and subspecies described as Of refcrnd to PJrylIoI>aln. All naJnC$ in til'<! column we re dieocribedas I' lryIloINuu IlTlless OIheTwisoe indic~ed . Subspecies aRc listed unde r lht ir spec"" ,

NAM E

a1D8"J'O'!'"albog""t1lU3onrhotr.,;" "rala (H)'/o)

a""",,.a" roUlrni" (lknd rob mu)bau l" i IDr ndrobau.}bralriciMbrtb<'; IH.,u-l...}

"""""bacGgr; lH_rIout/tu}bo/i>'ia1UU (I' rwtlorrapU!bo." rtrgr ri ( Prwrh<'ropis)br<r«" tllJ (Dr..,Jrobd tn)bro«al U3(Dr ndrobatu)brtuilirtr. l.brom r/ k ol"br"nnrUJ (ProSllu:rapis!capiwba

cNlh .c"'ocw (Hyliwhu)evi4uU (HyliuJllS'cyu i,lOlllltoidr.d..M; (PrtnllorrapU'clclOl'tSrq lMlwritJl;. (I'nmM rapis!upin(JJa;r lU:nrmi s (Drndrooolr. )lem" ",/jJ (ProJlhn api. )ImlOra liJ (P",sllu:rapi.!fr Utu ( Prollltrrapis)

AUTHOR ASO DATE

Bokel1lWln 196 7BoulenCt'T 1903Noble 1921Wie<! 1821Girard 1 8~~

Bouleng<:r 1913Melin t941Barbour and Dunn 1921Nobk J92~

Bibron ill s.a,n. IM IJimenezde Ja Esp.da 1870Boulenecr 1902..- 1909FilZinget bt Sreindachner J864Cope 1887Wine t930Test 1 9~6

Cope 1887Bokenn...n J967Bokenn an n 196 7W. ~la'1 1873BouJenger 19J2Boulen,er 1912Cope IBnRivero 1961Giinlher J808B~Uf J908Funkhou>eT 1956FilZinger in Sleindachr.er t864Bouleng er 1883Barbour 190,SPeracCI t904

OPI....IOS OS STATU S

Col,,.,crl.... olo8"J'O'!...Co/(mrlw,,/bo,llJlOt...Phy llobatu "nrhotr.,inol Ph),l/ohalrJ; sla1 u. uncenainDrnd",b<" r . aUrolU,Ph,,/Ioha'r~ " url>1"rni"Ph, ·/I0b0 tcs ban /criPI,,'IIQbat<,' /liI~..bris

CoiOSfrlw brill"......Plryflobo u . lJic<>Io,

CoIour,hllJ bo<tvIgriPIry.~lobotr' bo/i-.r-...Plryllobot cs bowl"ngr'iPIry.~lobeur. pict...P Iry.'l/oba ru pic ,...Cro m>d"" ryllil' gaudic lwudiCo losrn h" . bromdic<>laCo /oslrt h" J brlilrmru.JC<>Iosulh.,. c"pixabaC<>I".UIN" c" ,iocaS)TTItopIw. eNlI.., ...COwlrl/.... chocot'misCdtW<',W roi/QrisS,-rrltt:>p/tIIJ ryUigrwlllroidc.Co/MIT'''''' <1_inomen nudumE/rlilIM rodoc T)-f... liI"iurilO'lUPlryl/obarrJ r JPinoJa iPh )'l/oha rr l pic'"'Ph yUohulrJ Irmo' '' /isPh yllobalU ho lillrnKcriCo/m tclh ... I csrtu

BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY No . 27

fI(Jw,pi(1(J IHyl"l'lni(J)flolawrfuliginos as (HyIMalus)fusdcul(Jgl(Jnlfulosu<grunulive"'ri< IHyllxulu<)Iu,ltneli IlN~.drobales)ltermlntlf'IPn>Sllterap;s)It"igru.. (HylmralusJItylaefi,rmi.infroguuulUSingeri -(Oe",irobale. 1incu inalls IProSlNrapis)intermFlliusking.bu'}'ilaMalis IlNndmbm..s)lari"".'uslimbalUSlug"bri s (Oendmba'e»m"ndela '"mma"'Nsi"n",melanor,ltinu,mertensinebl in" (ProSlltu"pi.Jnice/urinigerrim o fHylolnigd....n"is (Hyl"plesialnuhieola"b"'urus (lNndrobarn)ol/u.iaide, (Eupt'mpltlx)polm",uspoMma".i.IHyloxulus)porvulus (O.."drobares)peruen.".'perU"lana.pelt'rsipiou (H)'laples ia)pious g""yu"..nsls (Dendmba'eSIprumpulcNlIu", (Pltyllodrumus )p"lehripeollJranoide, (Dendrobale S)ridens""rosan8""riveroi (Pro.INrapi. )"hrevel (Pm.fllterupi,, )smaragdinussuhpunelulus (ProSlN rUpi, )subpuru;u"us ....alesisylvalleusIMnia/uslal"mUlIf'a.. (Oendrub"'es)1",,,,Ii.' (Pro.<llterapis),..""vitt,"us (De"droba'e, )Iltomlonilin'I",lu. vi"",u.' (Ch'mi ro""',,)I,i,'olor (Pro-"hcropis )Irili""alusrrinllmisI,ivi"ol" (Hyla)" ivlu", ,,.'· ma,'ul",u" (Dendrob"'es)Irunealus""iobilis (P,o"lteropi$!vergeli fHylaxa/u, )""rTuculmu"...",bralis- (Plty/hlmmu,)....lty"'pt'rl (P,m,herapi,'),,'poro

TULE 2 (conlinued)A. LUll 1925Dunn 1931J;rncne~ oe la E<pada 1870Fit~inger 1860F;t~;ngcr ill Stcindac hner 18(,9Boul.nger 1919Boulenger 1883Boettger 1893rowler 1913Cope 1875Boulenger 1898Coe hran and Goin 1970Cope 1868Andersson 1945Boulenger 1918Cope 1874Co pe 1863Co pe 1862O. S<hrnidl 1857K. S<hmidl 1932Me lin 1941Benhold 1845Cochran and Gain 1964Test 1956Noble 1923bSpi~ 1824A, LUI~ 1925Dunn 1924Durneri! and Bibron 1!W1A. Lutz 1925Werner 18'19Dunn 1933Boule~ ger 1882Sleindachner 18(,9Melin 1941new,pec iesBibron in Tsc hudi 1838Hea'wole. Solano. and Healwole 1965Boulenger 1899aJ imenez de la Espada 1875new spec ie,Boulenger 1918Cope IBMAndersson 1945Oonoso· Barms 1964Rivero 1961new ,peciesCope 1899Coo;hrJn and Goin 1970Barbour and Noble 1920Ander,son 1945Cope 1875Wcrner 1916Miranda Ribeiro 1926Cochr an and Goi" 1970Co pe 1893Boulenger 18'i<1bBoulen~'Cr 1883Gannan 1887Spi~ 1824W, Pelers 1874Cope 1861Werner 1899Hellmid, 194()W. Peters 1870Boulen~er 18'19bBoulenger 1882new'spo:cks

Phyllobale. pierusColoSl/rhus "uhieoluCo/o.w,hus/uligi"osusnomen nudumPIt)""loemus olfe"iColaslnltus {X,lmalusPIt)'Jloboln pieru,ColoSlnhus I, ini'aris£/tulneroducrvl". dia'lt"'UEleulhtrodo,r,'lu" It)''',../ormi.Colo'lel},"s in/ragu""'...,Phvllobote> i'lger;COIOSlNhu,- ingulntllisColoS/elh"s in/trmediusCulo<""hu<killgs""rylPhyUobarn?; SlaIU' uneen ainColOSI..,ltus IUlillosusS",in,hi/lu, limbalUSPIt)'/Iolmlt s lugubrisColost"lltu.\' I,;nitulisCmostetlt u, brun""u,PIt\'/Iobaln birolorColosrn h" , merlt"siColostethus "tblinaPIt)'llobal"s birolorPhylloba'es "iviltolu.,?Eleu,NmdoCI)'lu. nlg';l'<'",rl,Coloste,ltus nubkol"Dmdrobaln; sralu, un..,nainCohwe,hus ol/e"ioid..Colo.'lethus polmal""Colo.,,,,,hu, ingui"ali .I'hyllaholes poroul" .,nol dendrobalid?Colo.""hu.,.peruvionu .,·I'hylJobates pete"iI'hylloba,n pic,,,,Phyllob<Ut's pierusCol""'eIItU.' pralliColo ."",ltu.' puleJwllu.,I'hyUolrcun p"lch,i!",elusColo ,lu,ltu.<ranoid"sEleulh"rod,/Crylus ride..'Colmurhu.• riO{:o,I'ung"eCol".<I(,ho.' ,i....",iCOI"Slelho• .shrewlPhyll"ba,..s ." ntlr" !!di,,u.'Colaslelh"s subpun"",u,­COlm-INIt".' .uhpu''''IUlU.\'Coloslelnus HJ""li,u.'·Col''-''leIltus I;,eni",,,sColo",..,ltus lul""'I1',,,,,eColwlt/hu.' . "bp """'lIIu ,Pltyllob<lIt s IrivlllulllS ~

ColoSlelhos l/rorn"",;Plty/lob<ms "ill"'u"Ph"lIoblllfS "i,,>lorCol",,,,,,hu ., I,ilinearu."Colrmerh",' trioiloli.'Phyl/"I""e, "ivi"aru.'Veod",b</les "uram , ?Vend'obar" "o""IIOj'C,,/m-ulhu s ,' obpunelmu,Colo,'lerltu.' ""rgeliS,·rrhophu,.- ve,ru,·" I", ,,-,C"/,, .,'etho. ",," ehru/i.Colosterhos ....hv"'pt'riPh"/IobllleS wporo

1976 FROGS OF THE GE,\'US PHYLLO BATES 7

P. bowl~"xai . P. f~m"ruli•• and P. tri"il/"tuJ have(in pre"':l'\'ali\e ) a dark brown Irans'erse band on lhepo...enoe ponionof the throaL

TiMJpoks of P1I.'"/fobol~J resembk mose of CoI"i/~'

11114. in 1Ia\'ing thin beals. a \k~lr.d " nus. "nd ll/l in.\knled ora l disl . Tedpole-, of PIr.'"lIob<ltn differ fromtbcee of D~ndrob.II~J : t.-:lpo le1; of D~"Jrobuln lIa'eth icke r !leah. lIa~e ( e ~cepl in llle mi" UIUJ group) amed illn anus and an UIl iltdcnlC'd or.Il di'l . and ....}me·time' ( in the hiJlrio"icUJ and pumi fi" group,) havea reduced number of dent icks.S/,<,ci~J ff, IOllps. -The 'pecies of Ph.'"If(,h<lt~J fall

into t"..o well·defined groo ps based on pre...:nce o rab-eoc e of we webbing:

( I ) The ,il species wilh basal toe webbing areincluded in the [emorata grou p: within lhis grou pare three subgro ups:jf'",oru/i, and ~"f,,,m (w r)' similarspecies of lbe Amamn d rainage); amhollyi . bou/en!:,., i.and I!"p;m).,,, i (, mall species o f the Pacific drainage);,md lric%r t whic h ha' green hone' [un)e,s Ihe c" lmin USl\·/-.1 specimens is an ani facl)).

(2) The 14 species without 100: webb ing arc dividedhere into three species groups :

(a) The bicolor group t fi~e ,pe.·ies) seems tohe natu ral. Three allopatric species. ouroeuema, futta·hli.•. " nd , ·iI/UlllS. are very similar. and some s~imen,

of "w"" "~,,i<l resemble bicoJor. There is some doubl;about the relalionship of a fiflh species ( unoJ,e \Cri~l.

hut it I ' included here because il ,uperflCiall) re...:mbksbico/Ol,

(b) The pi<"tlU group (se \ en s~ ies) rna) beonl) part ial]) natural. Pictw. and i"g~li (both with aeal f-~pot) probably are dosel)' K'laled . and ,...n'ulu.(which somenmes ha~ a cal f· ~pot ) resemble, them.P('lt rJi and pulchriprcl'" share ~imil"r color panern~

and may· tIC clo-cly related : boli,·i"" ... (whose co loris unlnow'nl resemble~ them . ahhou¥h b"li';,mu. also

resemble~ ~ome weblcss species of Col"i/ ('lhu•. There la tio nsh ips among the three subgroup~ i,, /{~ri ·

p<ln·UllU·pi.-t... . boJi.·iu"w. ·pn('rJi·pu lchri/'<'cl..... and.m" ' ugJi,,.... are obscure. and these:groups are indudc:dIOge:ther in the pinus group partl) for convenience.

(c) The trio·ilwl..J ¥roup ( two ~iesl proh;ibl)i~ nalural: huu/ui and tri, ·illalU. are large: ~rc:cie:~

rese:mblin¥ each O(her in color pattern.Lifr 1IisI0l)·. _ Dcndmbilt ids have: a comp lex life hi~ ·

lOT) uniqLIC among anuranv . These frog~ often bella'".:a~~~ivcly toward me:mber~ of lheu- own speces: thisaggre"ion proh;ibly is territorial de fense: (Senfft 19.\6 ;Tc:, 1 1954; Se xton 1960 ; Duellman 1966; Goodman1971: Ho" gmucd 1972; Crump 1972; Bunnell 197.1;Silverstone 197.1; Liiddec ke 1974; Polder 1974a . 1976;Durant and Dole 1975).

Ovipociuon is pre~eded by a co mplex cou rtshi p(Senffl I<BtI. 19,W; Te ' t 1% 2; Crump 1972; Silver ­'lone 197.1. 11l75; Polder 1974a. 1976: Dole and Durant1(74),

Ther e i~ nu amplexus (according to Polder [ 1',l74c],~I udr;fl: l ' s [1%9] o tl;,ervalion of clasping in a ,pecic~

of ColOJln huJ wa, cou rtship. not amplexus) . Threeoviposiling p''''lure~ ~rc l no",n: Type I : ,·enl·to-'enl.know n in lhe lHnJrobulU pumili" group: D. Il'''''u'lif('IUS tC rum p 19721. D . pumi/io (polder 197-k);T ype 2: male and female ...:parated. known in theD('"Jrob<lItJ linooriwJ group; D_ aUlotllS eSe nHt1936; Wardenier 1973: Polder 1974c ). D. U:"'('II'(Polder 1974a). D tinelOri... (Polder 1974<:); Type 3:mak I)i n¥ tranwcrsd) on female ' ~ hoICk. known inCOlmlrl1lllJ ,... Im,,'u. lliiddeckc: 1974) . In lhe o nl)~p«ics of Ph.' //obulU for which dala are a\'ailab le10 me. P. ..ill"I'... lhe o"ip''' iting posture is e~sc:nlially

Type 2. with a brief di~p1 aJ of tbe Type J Jll""lure insome: ovip"'itk>ns (Polder 1976; D . TTiIVis . ~r.'l(>"ul

communicmitm ). In ~pecies lhal use the T) pe;;: posture ,

T~ BI.£ 3

TlHJp'lc mc.'urcmeOls (mm) of nine specic. " f Ph."II" f><lIc,'. The first num~r lin b" ldface t}'J'>e ) after tile . peeifie epitbet j , tbeladp,k 'ta~e of G<I,ner (19llO) . Tbe se,:<>~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 maximumtotal length' for that ' tagc Data fur diffe",n t stages arc "'paraled b)' sem icolon'.

, pede'"b",.lr"Krriup;"" ,,,i!rmora/j.PCIr'"pklll '_'u"ai"",Iri'~llUf'"

4(J (I ) 10 .~. 27.5: 42 (I ) IU. 26.S: 44 (I ) I 1.5. 16 .~

28 (91 3.5-4, 0. 10.5-1 1.026 (3) 4.0. 11.028 ( I I) 3.5. IO.~: 29 (I ) 5.5 . "; 30 ( I) 7.0. -. 34 (2) 6.0- 1.0. IH I'26 (8) 4.5. 12.5: 28 tn 5.0 . 15,024 ( I) 4 0. 11.0; 15 (9) 3.0-4 5. 11.0_12.024 (2) 5.0. 14,0; 15 (5) 5.0. 14,0- 15,523 (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 13.5.21.525 (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 ""ATURAL HISTORY MUSEUM OF WS AS GEU S COUflo7Y No. 17

the m~le sil\ on lhe l~ )' i nll: ,itt both ~fore and afteroviposinon. hut SenlTt ( 19 .' 6 ) pro~eoJ Ihal ,~nn re-lea"" ' 'curs only af ler uviJ1<" it ion.

Kno n oVlp'"it ing , ite, are ( u 'uall~llea"e' (usuallyanached. MlfTIClimes fallen ) in D..",J",hal('S and Ph.'"l­lobatr5, and the ~rface of the wil (uOOerneath >tone .or in holes) in "OOle >jJ«ie s of CoioMerl" .., (Oe -e r 19.12;Senffl 1936; Dunn I~~ Oerner 1951; Stebbins andHendrickson 1959; SihcnolUne 1973; liiddede 1974;P(llde r 1974a . 197<k, 1976). Knmoll clutch ,in j ,

two 10 16 in D.." dro bute.<. seven III 27 in Phyll" bcllr.l .and 4 to 48 in Co {osl<"hu, (Dunn 1944: Der ner 1953;~I ud rack 1969; Polder 1974a. 1976: Ihi, paper); al­though each female lay. ej;l;' more than OJ1('¢' (,, >me.time. many lime') a y''''. tScnffi 19J6: Polder 1974b .1976). the [O(al an nuil e¥l! prod UCl ion per f~m,,1e i~

much Ie~~ !lun in "'altr -<l\oipo;.iting ;mU fiIllS . sug~ting

Il'1:llt dendrob;ll id ofhpring have a rela tivel y high '>Ur·vivlli rare.

Polde r (1976) noted lh al the word " guard ing"

less appro priate than lh~ word "l~nd i n l!" fo r (hosespecies [e.g .• D , auratu, and P. l'iIlUlU~ J in ....hich anadul t mm:1) ' visils t~ egg' l)(Y\Oo and t~n. "llhoughin some spece-, of Col(mttlu.s , lTW guard ing (d.m'i ng

. "' ''y pm:!.:l.lonl doo OI."Cur. :IIl"COmpilllicd b) ' " guardrelief ceremony and rem o,",,1 of mold y eggs b) mouth(M udrlOCk 1% 9 ).

After the ladpoles ha tch. lhe}' wrig i:1e onto the bac'kof II paren !, ap parentl y ad her ing by their be lli es 10

muc us secreted by lhe adu lt (Slcbbin, and Hend rickso n1959; Polder 1974b); af ter cerrying lhem one to seven

d<lys. lhe JXlrenl dcpo;.it> the tadpolts in " ~mall . , lillbod y of ",'altr. ",here lhe) complete t he ir de~lop­

menl o Transport and dcpo;. ition !>t)lTlCtimes occ ur in

~ag". i.e .• lhe ",hole brood is nOl nece '>ilTily carrie dor deposilc'd simullaneou,ly ( R u lh~n and GlIige 1915;Sentf 1936; Stebbins and Hcndrid ,o n 1959; D. Travis1"",..,,,,,,1 cummlmimliOl1). Kappler's ubservation (qlK>l<.-.Jin Bocfenger 11I1I1Ib) tha I P. Iri "itl<UU~ oviposil' inpuddle~ lInd retrll/l .,porb u, lad po les 10 large r puddle>proboobl) is incOll'eCt (Senffl 19-'6 ; Stebbi", "nd H~n ·

drit"kson 1959 ). TadpoJe-e"""iers ~ not (itS forme rlybe lieved) u clu s j' e l) ma le . Fem llle tadpole -carrier shlIv'e been found in seven specie,: C" lostelhlH , po(Mud md 1%9). C. inxui",,/i.• (K , D . Well, in lill .),C. wl" " uI/1("<le (e RE 9(1 ), De ndrol><tlf .• "Z/treIU(Po lder 1974b), D . pumilif! ( Kila'ako 1967), Ph.l" ·1"i><Jlelf emora li I ( KU 111515. ~ISH 389). and P .pnul; tM CZ 24.l.431_ In Dend,..,1'<l1<'J. a parem is

kno"n to carT) no moo: lha n three (Sen ff! 1936) tad.pole:, a t a time I u...all ) one or 1"'0 ) . In contract. inCQI<ljl~lh.... . a parent i ~ kno"'n 10 c arry 1"'0 to 25laup"k, (Dunn 1941) . In lhis respect. Phyll"I>ttI~.• ,which i, know n to carry one I" 27 tadpole, (T able 4),

resembles C"I""fflhl.... .f ·""d. - R ,R. Snelling (LA CMl kknlif<Cd the ,lorn·

ar h contents of 14 , pecimell\ o f 10 ~p«ies of Ph..-I·lob<Ju. ( T " bl e 51. Th e ~ lom"ch. eont"i ne d 24 1indi vidual invenebrales; nearl y half thr invencbl'ate s" ere anl~_ ~10>l of thr spe cie . pre yed upon inha bi tliller on the (ore,t nonr.

P" lder (19 76) nuled a differe nce between captiveDnulro lmlf.l (five species) and Ph.l'!l oh<tlf ," (threesJl<.-cic, l in prey size lind feeding behav ior . Dendmi><JIt'Jt al< on ly relat ively sm all prey (Senff! [ 1936] alsonoted that D_ <11""'11.' eals onl)' '" lhe ~mallt>l insectsand the ir Iar"ac; lhey were DOl " ble 10 lIWIa~e a ha lf.~rown meal,,'orm or a "nmolh lan-a·' ); PIn'lloha,'l

TABL~ 4

Tldpole-<:al'Tyint in P"yll<>bcut'~ . n: numhrr o f came... laken; Tadpok.: minimum.m.~im"m (mean. rounded to loweS! " 'linknumhel"l numhrr of tadpolc<per cartier; S11[!C: ""~ tGosner 1%0) of tildpo..le. "hen taken; Carrit1c 'u (.\.I: male,

f : fenulk l and minimum·mninwm (mt an) """"·\'t'1IIlength of e.m in millimt ten.

Specie> ,"'".,... .... C...,

<inlloon.i , u- " .\.1 16. ~

<>«;",' , , . .\.I41),jb<Julrngai 1 4- 9(6 ) 28 M 19,0"r<p'''''lU' 1 2-3(2) ie M 16,0- 17 .lJ(16~)

jr"""QiiJ 2 2- 11(6) 28 f 25.0- 31.5(28.3)prlr"1 2 l_llc4 ) 26.211 .\.l 25,0; F 28_()(26 . ~ )

".rlUS 2 4-1116 r 24.2~ M 21.G-2 2.()(2U l_ ...,.ti ..... , 7 24.25 M 26_ ~

m .. "",,,,, " 3-27(14)"" 28 M 3l.S -40.()(36_6'""Ulatus • 1_ 1:1(4) 2~ ,26 10-1 25 , G-26 . ()( 2S _ ~t

"f:, t;millod numhrr ej~ ht to 12: only' 1h",e we", l'"'>t:I'Vod' Sollle or all data "nkJ'KJ\\In .'G an. ( 1960) saw adult, carry in, e igh1 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<l b) C. W _ M},tn and J Dol)

/ 976 FROGS OF THE GENUS PH YLW BATES 9

eats both small and large prey (such a~ meal worm s.and earthworms longer than itselO. D. T ravis. ...hoabo noIed the generic d iffere nce in prey size , suggeMedto me that il is the result of the absence of teeth inDendrobates and the presence of teeth in Phyffobal~s .

Ba sed on the spec ies th.il l Polder observed, ho weve r .tee th cannot be the sole explanatio n (I' . Iril·iuul USlacks teeth) . nor can body size (P. villulu.< Is muchsmall er than D . a::. u r~ us). According to Polder , infeeding behavior, Dendrobates is a "seeker." hutPhylloha/~s is a "sitter and watcher."

Sumlllur}" of distribution (Figs. 13-1 7l . - Sped esof Ph}"lIohat~s occ ur from Costa Rica southward toPenl , Bolivia , and soutbern Brasil. Mosl species ofthe'-C diurnal. ler re slr ia l frog s inhabil wei tropicaljo rest. Reliable colJe,ctinllrecords are below 1800 m.

Only 1...·0 (I'. lugubr;s and P. l'itla/us) of lhe 20species live in Central America. and lhey are d,>,><: lyrelated to a Cbocoan species ( I' . ourQra~lIio} . Tenspec ic'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·dinus . P. trivinauu, and P. zaparov dwell in theAma zon ian regi on. Fou r spec ies (I'. f~lIl..ratii , P.pin uJ. P. pulchrtpecna. and P. " il'illmus) occ ur inthe Guianan region; all but one of these (I'. putchn­p{'c/UJ) also occur in the Amazonian region . Fourspecies (I' . aurotaenia, P. bicotor, P. fx}f/l~ngt'ri .

and P. eJpillo.'ui ) inhabit the rain-d renched Chocoan

region west of the Andes. One specie s (Ph)'l/oha/n' Iledes) occu pies moderate eleval ions on lhe eas ternslope of tbe Andes in Peru. Two species IF. Iric"lorand P. boulengu; VI " close relalive) occur on tbe...estern slope of the ATlIks al moderate eleslliions in~uador. One species (I'. amlwn)"i} li\"\", in the PacifICIowland~ and Andean fOOlhills of south...-estem Ecuadorand l"IOTlh ...estern Peru.

Tbe dist ribution ...ithin polaical units is a~ fol lows:Costa Rica: IWO ~pecies (I'. lugubris and P. "illatus);

T.... uS

~h "",,1(_ of 14 <pcei_ns of 10 opec:ies ofPII)'//obateJ . Colum n A: pm:enlare of 10IaI ....."ber

of prey individuals. Column B: pen:enu.s:e ofIOU.I number of "<>ma"h!..

Food Cal(llory A( 4) B('l)

AnlS 49.4 78.6Tenn ilCs 17.0 21,4Spiders 14.S 7.,Bulin H 42,8H"mnpleran, and lrue bugs ,.. 3nFlies '.0 21,4Wasps L3 21.4Inseel w...u Doll" 7.'Sprinllail. 0.' 7.'Gras.\hoppen 9.' 7. '

""""""""'" 0.' 7. '

Panama; one: species II'. /ugubris}; Colo mbia; eig htspec ie. (I' . aurola~nja. P. bieo/"r. P. boule ngeri ,P. f emorulis, P. ingtri, P. (1(m·ulus. P. pic/us. and

P. /r;"i/fu/ a.l); Ecuador: nine spec ies (I'. aruh"l1)'i.P. b(}ul~tI8~ri. P. {'.Ipin"sai. P.felll"rali.l, P. f4lrvulus.P. picllt," P. tricolor , P. trivntutus, and P. t.a/l('ro);Peni: nine species (Phylloba/t;'.1 species, P. an/hon)"i,P. b'l, s/e,,·. P. femoralts . P. purvalas. P. perer.li,P. pic"IIIJ. P. lI1lC1raxdillUJ . and P. "il'ilta/us ); Bolivia:IW O species (I' . m,liviallu.I and P. piNUS); Brasil:four species lP . femorali.l. P. pic/a.l. P. pufchripec /us.and P. Ir;l·ill,I/US); Venezue la: one: ~pecies (I' . pic/a.l};Guyana: lhree species II' . femoral;s . P. p;NUS. and P./ril·irtalus'; Suriname: three species II' . ftltlOralis. P.piNu .•. and P. Iri"illaIUS); French Guiana ; 1"'0 specje,

IP.f~moralis and P. piNUJ}.I have discussed the lOOgeogra phy of dendrobatid

frogs in an earlie r paper (Silverstone 1975).

K~J /(J /h~ Genera of J)t'llclrob/llid(l~

Ia Term inal d isk of thir d linger usually less than 1.5 limes width of third linge r; dorsu m dully colored orbrightly co lored in life; maxillary and premaxillary teeth present or absent; tee webbing present orabsent ; first finger equal 10. aborter man. or longer than second; palatine bones present o r absent:omosremum present 2

lb . Termi nal d isk of third finger usuall)· IWO or more limes as wide as lhird linger ; dorsu m partly or entirelybrightly co lored in life; maxillary and premaxillary teeth absent ; toe webbing absenl; firsl linger shorterman second; palatine bones absenl; onjouernum presen t o r absen t Dem/robares

2a . Dorsum dully colored in life; maxillary and premaxillary teeth present; palatine bones present orabsent . 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 ) .

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

Adull ' OOUI-venl leng th 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 )Adull snout-vent length moderate t l 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»

10

2a (II).

~b .

31 (l b).

3b

BULLETI .'" OF THE NATURAL HISTORY M USEUM OF WS ANGELES COUf',7Y 1'.'0.17

4.1co.4b.

S. (4a).

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

Ad...11 >oJlOIJ.I -~en\ lengrh 19.0 -245 mm in males. 21.5 -26.5 mm in females ; bones green (Fronl ispiece 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 ..eb. Do rsum brown or black in life

. P. t "p',m (page 33)7

7a (6b) Adu ll snUUI' vent lenglh 1.'i.0-20.0 r um in males, 17.0 -21, 0 nun in km<tl<' s; skin of dorsum lI suall ~

smooth (e xcept po sterior bm:k tubercular}, sometimes granular or with scatte red ruberc l...s; throathl..d; and white: Chocoa n reg ion of Colombia and F..c u<tdor. wesl of the And..-, (Figs. lB . .'iE.7E) P. ho"I"lIgu; (page 28)

7b . Adull snout-vent ....nglh 20 .0-31..'i mm in males. 23.0 - 33.0 mm in females; skin of dorsu m srrongl~

gT"dnular; throat usually enlird)' black: northern South Ame rica easl of ue Andes (Froraispicc e II;Figs . 3F. gEl P. 1m/om/i .• (page 30)

9a (Sa ) . Lighl lateral stripe presem9b . Lighl late ral smpe ah!<nr (Figs. 3<:. 9C)

8a (l b).8b

Lighl proximos 'cntral calf.spot present (Figs. 9C -F)Lighr prox ill1O\~ntral calf. spot ab-e m

. ............ ..... .. .. . .. .............. 9III.

.. P. int/ui (page 35)

lOa (93). Lighl lateral s,ripe iecompkte (Fig . IA); dorsum red in life (Fronlispiece I; Fig . .'iBl . .. P. P<J" '" I"I(page 36)

lOb. Light lateral suipe complele (Fig . IB) ; dorsum black or bro...n in life (Fip. 3A-B. D-E. 9D -F) . P.piC'11U (page 38)

I Ia (8 b). Dorsolateral O)f lalerdl slripe pre-emI Ib. Dorsolateral or lateral smpe a~nl

1412

12a (lIb). Adull snout-vern length 32.5-42.5 mm 1312b. Adu lt snoul·v..-ntk'ngrh 17.5 -24.0 mm (Fromispiece I; Figs. .'lB. 7D) , .. P. p<lrl'ulul (page 36)

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,': : Cordille ra Azul of Peru {Front ispiece 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) 1514b. 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. /r i ­vhuuus (page 45)

lOb. Adult snoul-vent length Ies, than 30 mm; corscm never bearv lighl median stripe: venter lighl ...ithdark marbl ing or spot s (in pre-efVali' e . venter may become entirel y black) 17

17. (I6b). Cnnspicuous light ()'e lJo... in life) proxi~ thigh·"JlOI pre....nl (Front ispiece II; Fig _6E) . P.!",I­C'IIripl'ctus (pa ge 43)

/ 976

17b .

FROGS OF THE.GV'US PHYLLOBATES

Li~hl pnnimodoNlllhigh-~pol al><.em lFigs. X:.9Aj

/I

. .. 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 orSou lh Amerka we-t o f I~ Ande , ( f.'jg\ . ~A-D) 19

18b. Li~hl «npelall'f""lne.. po'>lerior end of -t ripe is in groin); Soulh Amerka ea-t (lf the Ande, I Figs . 2F.M-C. F) . . .... ... .. ..... ....... . •. 21

19a ( 18a). Venter blad with lighl d<ll~; Ch,,,:oan region of Colombia. west o f the Andes tPronnvpiecc I: Fig, .-l.C-D.8 B) .. P. <lurol,... ,,;U (pilge 211

19b verner blad. with light marbling; Central America (Fig. 8A) 20

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'uallyrelatively narrow : light area' un dOT,;!1 surface of limb , go ld, yellu "" or ye llo w· gree n in life; Caribbeandr..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 -3 1.0 mm in females; dor'oOla1erai slrip: u'>llall}rdali\-ely wide: ligb t areas on do l'>al surfxe o f limn.. bl~-gl"C'C n in life ; f'-.ocifie dra inage of Costa Ricain Gol fo Dulce region (Frcnnvpiccc I: Fig . 4B) .. P , "i ll </lU,\ (page 25)

21a (l8b). Ad ult snou Henllength Ie" lh.an 32 mm21b. Adult ..notI1-\"('nl length greaier lha n 32 mm (Figs . 2F. 6A-0)

2201 (21a ). Ve nter marb led or spotted (Fronnv picce II: Figs , er. 9B)22b . V..nter n'11 marbled UT sported IProntispicce II; Fig . 8C)

• . ..• P . hcr.u/tfi lPi'gc 45)

, . P. {'t'll'fJ; (page 37),., P. Wlll fuxd;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 , boulmJl~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

EF1: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 (RMNH6413); (D) P. ',i "rtaru" . Pallcm I, Pampa Hermon , P"ni (AMNH 43 11 ~ ): (E) p, trMlla/u.•. Paltem I. Pampa Hcnno~. Peru(AMNH 43 117); (F) P. ba..lm", 1!<,lolypc. Roque . Peni (NMG 51 n, Diagrammalic and not to scale .

rFIGURE. 3. Don.aI -- !'«t. (AI PJryllobau! . pKtld . Pancm 3 (DmJro#1GtrJ ,•• leckK)'pe). Rio Mamore . Brull (NHMV.'19190:3); (DI P. pi rtMS. Panem 4 /PtNJ'Obtzu ', h<JJtNli. 1tc1CM)pl'). Yurima' PI:nitBM 1941.2 .15.17) ; (O P. u.~ri. ho\()(ypc.iUt'nio near Rio fheado . Colombia (USNM 146II4t»; (D) P. pi n ...., ""Uem I (H.,-/"pksiaj/uwJpkltl. lOp:ll)pc), Bdo Horizonte.Brasil (KU 93141); (E) P. pi(",... . Suhpalll:m 6<;, Kurupung Rivet, GUlana (U MMZ 835981; If) P fr_ro.li%. Te.., . inha, B~J

(lACM 61030) . DiagramlrlJl io; and no( 10 ...-ale.

/976 FROGS OF THE GEl'WS PHYLLO BATES IJ

F1GCU 4 D<:>fu,1 ... pect . (A) PIr.-llobJ lr . '"gltbri•• BamOO . 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.mipedfo 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. <lIt11oon,.;. ~)pe . Salvia>.Ec-'or (A)I.! SH 13140); t El P. botM"gui. Imbili. Coloml>ia(USSM 141433); IF) P. rlpi_i. ,-anoe. SanIODominJOde Jot,

Colortlc\M CCAS·SU IOnS)_ DilI~:r"mmaic 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. (Ai Phyll"i><Jtnont!Jlmyi. llololype, Salvia, . Ecuador (A MNH 13739); (BlP. <l1II1w,,)'i. paratype, Salvia"Ecuador (A MN H 13740); (C) I' , /t;v;r,,,,•.<. 7.2 km NW let;,:i •. Colo mbia (LAC M 60981); (D)P. 1"''''01••, PuC"OLib",. Ecuador(KU 122262); (E) P. bou/rog,ri. Imbili . Colombia (USNM 147440); (F) p , esplow "i. par'l)'pe , Sanlo Domingo de lu' Colorado. _Ecuador (CAS·SU 10375), Diagram matic and nO! to "".Ie.

1976 FROGS OF lHE GEAUS PHYlLOBATES ts

F1GlJU 8. Ventr~J aspect . (AI Plryllobal ts 1"1"l>rl•. Bambu. CO' la Rica (e RE 11 82); (B)P. <l"rol"~"i,,, nanuw .' triped form. Que_brad. Boehmamli. Co lombia (LA CM 4319J ); (Cl P. ,,,,,,mgdin,, s. hoIOl}pe. Pan llc Az!k·ar. Peru (LACM 6443'): (D) P. ~"f'<Iro .

paratype. Sarayacu. Ecuador (KU 12(670); (El P.!.",,,,,,,lis. Tere,inha. Bra'il (LACM 61030); (F) P. m·""I"r. le,·lotype. Porvenir.Ecuador ( BM 1947.2 .14, 18). Diagrammalie and not to scale.

FIGl-~ 9. Wllm,I~. (A ) P", I/obo'e. hoIi....""•• kelOl)pe. SMl Canoo. Bolivia (BM J941.2 . 13.89) ; (B) P. pr'e rsi. 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 Holi ron tc. BlaSiJ (WCAB 4218); t El 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 to­Iype). Rio Maroo"; . Bra.il (NHMW 191'XH ), Dialtrammatic and IlOIlOocale

16 BUllETIN OF THE NATURAL HISTORY MUSEUM OF WS ANGELES COUNTY No. 1 7

• 0-.

F1GllU. 10. Tadpolt of Phylloba,~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<l,,,•• lalCnol"'~l. (All', """"r,,~ri. St. ll" ~K. from bai;k of adull mn" w..Go'l""". Colom t>;.( US~'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. SLoll" 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\<ia(UMMZ 5894 7); Ie) P ,ri·";IItmd, Sla,e 28 , from bao"kof Idl>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', .. (FMS H 3671ll); (G) P. pin". (L'MMZ Sll947): (H) P. ,ri.inalld (BM 1936.9.) ,6). u..... "'lual."... millimclC1'

o ' . ' '' "UTI' ~••"" ,.,.

r- -t

r-v

a

••

••_ w,,~ .. ,!....

~ """'0;­'. .".~t~.

+

18 BUULT/N OF THE NATURAL Hl srORY MUSEUM OF WS ANGELES COUNTY

·-...-.... .".".· ::.- ,, ~

.. ' . " .

No. 17

+

-,- ..' .

+ .. ~ _.~...,· ~ , ..c- ....... --L

+

1970 FROGS OF THE GEAUS PHYLLOBATES

[J "." 0 04'" lUI "

/9

FlGl:u 16. Di..ni bulion 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 OF W S ANGELES COUNTY NQ,27

o

Flcicu. 18. Vcnual~. 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 ,roupThis group includes rIVe .~~. lPlrylWbotu speces,

P. aurolfunia. P. bic(}/or . P. lu/C" bru, P. v;ttalltJ).Definit ion: sec ut-venr length of aduh s sma ll 10

large (18.5-42..5 rnm); lox. wcbk•• ; palmi smpes(...hen presenl ) dorsolateral , nol lakral (i.e. , poslcriorend of stripe ts on bilck at base of th igh . not in groi n:Figs. 4A-D); light proxtmoventra t calf·spot a~nt .

Plryllobo/n spec ies(10 be described by C.W. Myers and J Daly)

Fromlspiece Jl and Fi~ure 4F

196], I'hyllob<lu. bi' "<J/or. Cochran: 107, plale ~9 (~olor

pooto).1971. Phyl/<>bu/u bic% r, Liilin" 161- 174

Drfinirion .-TeClh present; toes webless: dorsumred , red-orange, or orange . sometimes with black~POIS and marbling , or don;um black with red. red­orange . or orange SpolS and marbling; slripes absenl;lighl proximoveraral calf,s pol absent; venter red, black,or panly red and panl y black.

Diagnosis .-Ph)'lIob<lt<,s speces is I. member of lbebkolor iJUup. differing from P. o_O<',,;a. P. lugu·

bru, and P. vi,lalWJ in IMBer size (Table 1) and inlacking stripes, and from P. bicolor in llaving a stronglygranular ~um and in sometimes having a differentdonal panem . 'The red. red-orange, or orange groundcolor resembles lhal of P. biro/w . bUI somclimcs isbroken by black spots and mlllbling; in some ...pe<:i·mens, the don;wn is p~inanlly black . and lhe redis reduced 10 spots and marbling (in P. biro/or. lhedorsum is smooth and uniform red·orange, orange . oryellow·orange).

Description, -c-Measuremems are in Table I. Theskin is strongly granular on lhe back and the dorsalsurface of the hind limbs and smOO1h on lhe head. theforelimbs. tbe venter. and the ventral surface of thehind limb. (in USNM 166904A, lhe skin is smooth inpreservanve. bUIcolor s lidc~ taken by w .e. Shcrbrockeshow lhal the skin of the back and the dorsal ,urfaeeof lhe hind limbs was strongly granular in life). Maxll­lary a nd premax illary reerh are present. The fir' lfinger is very slighlly longer than lhe second . Thetoes are ,..ebless.

In life (color slides by W.e. Sherbrooke; venternO( visible), in USN~1 l66904A (Pronnspiece m. theiris and tbe tympanum were black. 'The dorsum wasred wilh black spots on lhe mid·hack and postel iorback . The forelimbs were enlirely red; lhc hind limbswere partly red and panly black.

1976 FROGS OF THE GEl\BS PHYLLOBA TES 11

In preservative . in US~~I 166904A. the tympanumi~ dark brown . The dorsum is ITllNl)' c ream. with a'mall dark bro...n spot behind the right upper eyelidlind seven ~all irrellular dark bro.. n spots scatteredover the back; the extreme po'>terloT back is dark bro...n .The side of the head ilfIdbod)' and the entire forelimband !'land are cream . Tbe hind limb is cream, exce-pI.lhedi~ portion o f one thigh. the \'e-n1fal surfXC'o f lhe thigh. the antelm'C'ntral surface of the calf , andthe ventr al surface of the fOOl . .. hich arC' dark brownThc C'lllIt'1l'lC' po'>te-rior belly is dark brown; the rest o fthe venter is cream with an indistinct lillht brown washon lhe tbroat and bell) ',

In preservative. in fl.1~H 56263·72 . the- tympanumis dark brown. The head and beck arc: uniform creamin four specime ns (except for a small dark brown area ,sometimes preceded by one- or two bfllwn dots, on thee xtreme posterior back). Four other spec imens haveone to several small brown spots on tbe mid-back andpo slerior bad ; the brown area o n the e xtreme posteriorback of these specimens sends fOl1ha pair of e xtensionsalong the posterior one· fourlh o r one·fiflh of the back .In FMN H 56272 (Fig. 4f) . e xte nsive dar k brownmiU"bling interruptsthe c ream ground co lor of the: back .In FMl"H 56268. the back is predominantly dark brown;the: cream oolor is con fined to small round OC' elongatespols on the back and an extensive arca on Inc head(broken by two small brown SpolS and by an im:gu larbrown area ext e-nding antenad from tM back). Theforelimb is entirely cream in a ll spedmens excepeFM~H 56268 and 56272 . in ...hich the ventral surfaceo f the forelimb is panI)' brown. The dorsal surface oftM hind limb is entirely' or mostly cream. excepe inFMS H 56268 and 56272 , in which it is entire ly darkbecwn . TIle ventral surface: of tnc hind limb is entirelydark bro...n excejx for an elongate cream arca on thecalf of some spec;imC'ns . The ver nerof the eig ht lighlestspec imens is cream. usua lly ....ith a partia l brown wash .In FMNH 56272 , the venter is dark brown wilh irregu­lar cream areas alonl! us lateral portions . In FMN H56268. the venter is entirel y dark brown,

Tadpot..s.-On 8 October 1966, W.C . Sherbrookecaptured USNM 166904A , a 40 .5 mm male; Ihe froghad six tadpo les (USNM 166904B ) on its back. Whenpreserved six weeks afler c;aplure, 1hC' ladpoles werein Stages 40 , 4 2 , and 44 . Measurements are inTa ble 3.

In their native habitat, adults deposittadpoles in waterin small puddle s and d itches (LUling 1971).

Life hislol')·. - Liilinl! (1971) witnessed intras pecifica"ressive behavior (jerking the head upward seve raltill'lC's, then allacling and biling the opponent) in c;aplivespecimC'ns. Luling also reported l!'IataI metamorphosis,young frop are uniform dark black.gray ; they take-overthree I\XII'lths to develop the full ad ult oolonltlon.

SIC",lOch COll fn u s {one spec;ill'lC'n).-From FMSH56265 : HYMES OPTE RA: Formicjdae: P~idol.. spp .

(27) , Ponerillltt: fragments, ~I)' rmecinae : fragll'lC'nts :COLEO PT ERA : fragmen ts of beetles of familiesStaph)linidae, Bostrkhldae, Curc ulionidae. and Ela ­rendae , a lso OIlC' unusual bee tle larva o f unkno wnfamily; LEPIDO PTERA: Soctuidae? (l lan'a): HEMI ·PTERA: Mirnae~ fnogll'lC'nts

P,..dators .-Liiling (1971) sa'" a snake (Chi,o nildsp .) from the Cordi llera Azul allack and seize a C;apl Mspecimeno f Plr)'lIo bafcs speciC'S; the snake immed iatelyreleased the frog, retreated . shook it-e lf for 0 .75 min.kepi its mouth ...ide open for 6 min. and rubbed lhesides o f its mouth against branche s . Th is repul,itm o f apredator (probably becacse of lhe unplea....nr, burnin gta,l e of the frog' s skin secreuoej, oombined ....ith Lu·ling' s obse rvatilln lhat lhC' ffllp, in their native habilal,~it in open areas and contras t co nepicuous fy wuh thesurrou nding vegetation and so il, , ul!llest~ that the bri~ hl

colora tion of this species has an aposemauc function .Distribution and habilal (Fig. 14 ). -c-Ele vauon

record s are from 1300 10 1600 m. Phyllob<l/cs speciesis known only from the Cord illera Azul , Peru, whichis the drainage divide between the Rios Ucayali andHuallaga (tributaries oflhe Amazon). Liiling ( 1971) tookthis frog on forested slopes, on the grou nd. in clea r­ings that were overgrown with grass and terrestrialaro ids.

LOHllilin . - PfR O, HIJUi. _ : km 189. F.D.O. Inti......Tingo Marla'PllCallp a roael. Cord illera Azul ( USS MI66904A_B); Divisoria , c~ of r1dgc "' here Tinro ~l aria·

Pu<aIlpa ro.l croo.ocs divide bcf,.« n Rm Uc:a)ali mil H... .llap (FMSH ~26)-72)

CI..areJ alld stahl<'d ffWl.. , UlJ. - FMSH ~265

Phy/lobatrJ aurDlacniD (Boule-ngoer 1913)KOkoe

Fron ti,p iece l and Figures 4C-D, 8B

1913. DrmJrobt>1rs "wrOl~"ia Boulcnger; 10 29. pla~ 104,fig. l, hololype: BM 1947,2.15. 13, Peiia Lisa ,Rio Condolo, ChocO, Colombia. 92 m

1957. Phyll"bcl/ts a..'o".,,"u.. Dunn: 78.1963. Ph)lI<>b<urs bicalar . Marki and Wirkop: 330.1966. Plryllabarrs la/ina . ... , Latham: 683.1968. PIr)'lIabcll'n a..row rnia . Savage: 765_766.1970. Piryllohlll r s bieol"r ( part) . Cochran and Goin ' 37

(no! the plate; o nly lhe follo", ing are P. awrOlumia:USNM 14510j-06. 146630 _3 1, 147204 -07.150632-42).

1970. PIr.• flobalrs !rmartllif (part) . Cocbran and Goin : 40(1Kl1 lhe plate nor ,he specimens eumi~d; onlyDrrtdrohrurs QW""oc"itl and PJryftobtJlr J tll<l"f>ltlr'li<tin the s)1'O"ymy are P. tl"""....Ilia).

Dcjinir;CHI . - Tee th prc:\enl: (OC'S eble n ; dorsumblack . wmttimes covered by yello -oeange or reo-orange suffusion: complete green . )'ello ..<green, yel­10.... )'ello ...-ceange, red-orange , or brownish gold

11 BUUET1N OF THE NATURAL HISTORY M USEUM OF W S ANGELES COUt.7Y No. 27

do rsolatera! stripe; light prc xlmovemeat ,""If-spot"b-enl; venter black with blue or grttn doh.

Diagnosis. -PJIl-lloba,ts QllrOl~nio is a memberof lhf:,bko,," group. di ffering from PIr.I'l/obOlrJ lopeCiesand P. hie-o/ew in smaller sin (Table I) and in hllvinj!slripeS, and from r . t"gubris and P. ,.;/1.. ' .... in I....kingventra.lll\<lfbling.

Due- r;p/ion.-The name PIrJllobul#:s ollrotarniocumolly is applied 10 '''''0 forms. Cumbincd mc~u~­

merus for the ' ....0 fon m are in Table I . In boIh forms.the skin is sJij;hll)" granu lar on the dor;um and smoolhon the \<C'ntcr;md tbe limbs. ~l a~ iIlary and premaxi llarylttlh are pre...,nl . The fin;( finge r is longer than thesewnd. The lOCI are wcl:>le",.

One form (Fromispiecc I and Figs. 4<:. liB) is slightlysmaller than the other, This fonn includes the holotypc ,a 28.5 mm male . The adult snout,vent length in milli­mete rs (mean in parentheses) is: 19 male s 23.5-28.5(26 .0) ; four females 26.5-30.0 (2t\.1); all 23 spcci­mens 23.5-30.0 (26.3). In life , the ground color ofthe dorsum and of the side of the bo:.xly is black . Arelatively narrow complet e dor ...rlatcral stripe extendsfn,m the dOr!'al base of the Ihi~h ak,n~ tbe back. alongthe outer borde r of lhe uppe r eyelid. and along thecamhus rostrahs to the tip of lhe snoul. where it meetsits fellow. lbe su-ipe is bright green or p,ale }e llow·~reen in specime ns from lhe Se rrania de Bau do .and yellow. li ghl or da rk yello ....·oranlle. or lightbrmtonish gold in specimens from the upper San Juandrain.a).~. l1le dorsal SurfllCe of the limbo is black ....ithgold. orange. blue. or gree n dob; lhe venter and theventral SUrfllCe of the limbs ~ blacl. ....ilh blue orsreen dols. The doIs are a[...a), blue in .pecimensfrom the SaTania de BaudO. The dots are 'p.an.eon theventer and ITIOIl: numeruu.. on the limbs. In ~..:rva·

nve. the snipes become !!ray. and Ihe dots o flendisap pear.

lbe other form ( Fil;l. .lD ) is large r than the narro w­striped form . The adult snout·venllenglh in millimeter!'(me"" in pare mbeses j is: four males 31.0-31.5 (3 1. 1);three females 33.0-3.l.0 (33.3 ); all seven spco.: imens31.0-34.0 (3 2.D. In life. the color resembles thaior the narrow-stripe d form, but the dorsolate ral stripe( which is yello w-o range or rcd ·or ange) is broader .The dorsolateral stripes sometimes are nearly unitedby a dOl'Sa l suffusion (Fig. 40), whkh may also coverthe side of the !xxi }' and of the snout and pan of thedONiI surface of the limbs. The ventra l dOls oflen arcmore numerous than in the narrow- wiped form.

Rrt't'Jing call. _The ca ll of bo lh for ms of thekLik,~ SOIlnds lhe s.ame to lhe human ear. a loud. bird ­lil.e. ",hi rring t",'iller. consis lin!! of rapid ly repea lednotes. The ..:rics is repealed after ilTt'gu lar inlervalsof a fe.... seconds lo.l5 seconds . The call duration inseconds (mean in paremhe..:,) is: narn'''''·slriped fonnR io Sapipi. Iive calls: .l,O-6.0 (5 .8). QuebradaBoclloramio. four calls: 8.0-11 .0 (8 .1); bmad-<triped

fomt : Quebr..d" Bocbo rama. five calls: 3.5 -5.0 (4.0),The Mkt~ usually catls ...hile coocealed benealh fallenleaves or loS" but -.ometimes calls ...hile Sillin!!on lOpof a fallen Ie"f,

Rt' lul imu ll ips. - The inh.abilanh of P1a}a de Orolold me lhe 1\arT"O\Oo'·stnped kt>/w.t d....ell, al k»ser ele ... ·lions Ih.an the broiod·slriped t~; a ChoOOlndi;on rotdme. ho...ever. lh.at on the Quebrada Bochor.llmi (upperSan Juan drail'Lilj!C). lhe J\OoO forms an: no! separ;atedby- ele\-'ation. btU b} the quebrada itself. and lh.at lhenarro...·s triped t.",,~ does occur on hill. on the lefthank. AI lhe Quebrada 8ocho r.una, I look the narrow ·strip..-d ioto; only on the left bank al ISO m, and lhebroad·sl riped toio; only on a hill (the Lorna deEncamacitlfl) immediately opposite lhe presious 10e..l·ily. on lhe r ight bank al 400-520 m, On the Caminode Yupe in the Serrunia de Baud " (near the upperRio 0 p" l!ad\' in the weste rn Atratc dr..in..ge), wherethe broad .slriped kill",,: is ..bsenr. the n"rro w·sl ripedk,ik",.' reache s al leasl 500 m

Should Ihe IWO for ms of k,ik",' pmve to be dist inct,pecies. lhe n..me P . Ulm>we " i" ...ould apply to Ihenarrow-srripcd fonn . J,W, Daly told me the two f"m lsinter breed in capli,'ily. prodoc ing vi..ble offspring.

The n..rro ... ·s triped ioko; resembles P. IUlluhr;s,,00 P. ";/I"/lU of Centr,,1 Americ a. bul is sep;<raledfrom them b} " di,tribulional gap in lhe Danen resiono f Pan ..ma . Specimens of tbe broad·,lriped i"t,,;...hose stripes are ..Imost joined by an orange dorsalsuffusion (Fi~. 40) re-emble P. bicoJor (...hich h.asa uniform red -oran ge. or.ulge. or l'ello""-<>r,lOge coe­sum. lackin!! 'tripe~). bUI a"erage ,mailer: sevenspecimen. 31.0-34.0 (32_1). versu~ 4-1 specimen.32.5-e.0 (.\S.2) inP. hi...,I,,,..

Sf"mudl nmft'<l1J (one specimen).-From LAC~1

.l3800: HYMES OPT ERA: rormicidae: C.l'phtmt.W.....:l:IOllllise<lpus We ber (5). Sirumig~"ys 'P. (I). Del'"W.,.1IkI -"n"'gnUlhu B,,' ...-n and Kempf ( I); ISO PTERA:fragments.

TO,lim. -The .kin of P. uurm<l<,,,iu .,,-'nlains batra­cboroxln (BTX). a ste roidal alkaloid rhut is OI\C ofthe most powerful non- protein poiso ns kno ...n; lbe' UOCUlaneou,s LD,.. for mice is t...-o microgram, per kiln·gram nf bo:.>dy weig ht (Tok uyallla et at. 1969). In mum­

lIl"ls, BTX .·au...:s an increase in permeability " f mem·bmnc' to Na · , wilh con-cqucnt melllbmnc <lq,,, lari, ...lion. cnhareed spontaneou s rrunsmrtrcr release. lIlu'Clecontrlli.' tuTl,', bloc kade of !!"Il~ lion ie lran, mi" ion. andcardiac arrh)'lhlllias 4Kayaalp et al. 197U: Alb"'lU<:"-lucc laJ. 1970

Elhn<>z,,,,I<>KY, -The Choco Indian, of lhe uppe rS..II Ju..n dr"ina~~ of Colombi .. exlrdCt bk>wgun poi"'>nfro m both form, of P , uur{1/<l~ni<l. "nd call both/;0/;"; llhe first vo",el i, nasal: pronounced kohn~·

KWAY). ( lbe~' abo ex tract poison from P. bic<nor .",'hieh lhe} call "",mU lbe Indian, find the it>};,'; b>imit..tinS ih call ..... histling ...hile I..ppin~ their cheeh.

/976 FROGS OF THE GENUS PHYLLOBATES

Because lhe ti>lwi release, ih skin poi....111 only under~ll·e~' . lhe India ns pierce its visce ra wilh " woodenske we r inse rted throu!!h lhe lI1<,luth . and somenm esho ld it <)Ver a fin: . The Indian, poison the tiP!' of blow.J!un dan , by beat in!! them on the back of ee fro!!; thepoison adhere~ to ,piral !!rno"C'~ in the liP!' of the dan,t Fma Iob IlIh9. 1909; W",..en 1957). The Cho..'6 Indian,of the Serra n ia de BaudO do JK){ usc blowguns anddo no! e lllr.tel poisonfrom lhe wlwi .

DiHrib"tion " nd h" bit,, ! tFig . 13j . _ Ele .'a l ionrecord, an: from 60 to 520 m . P. aurol~"iu inhabil'lhe wet fore ' h of the Chocoon region of Cok.mbia."'e,1 of the And.:,. in the AlralO and San Juan dr"ina~,

U 1(·"lilin,-eoW MBIA,ChtKfi: PenaUl'lI, Rio Condolo(BM 1947.2.15.13); PI:.~ :. de Om, Rio S:.n Juan (lJSNM1506n -42, I~ M MlIl; LACM 4~792_94); 16_24 ~m W Playade oro (US r-- M 146630_31, 147204_07); Rio San h an(USNM 159401_~O, 1594.'1 -1I6); Quc hr. d. IJoehnrama.upper San JUall drainage (LACM 43791. 43795-11(1); upperRio Napipi (LACM 4~782- 1I9); Irail N:l....ccn Rio> Napipi andOpo~ado (LAC...l 4~m); Camino de Yupe. bet....ecn upperRi" Opo~:.d,i :.nd Domingooo drainage (LACM 7197M);upper Rio B""y (LAC...I 61020-21); All" del B""y (LACM71979_112); Cmo DocordO ben.un Cucurrupi and N,IiIIl.....='.$;on Jwn drajna~ (CAS 1194-15); Quebrada Tap;trill. 20 l;m:.,; Pale' li",.. S:.n Juan drainage tCAS 1198ll6). V" III : EIPaikin. 12 ~m from B""nllVCnlunI (USSM 1451(5 ); Io.. erRio Caling tUS:";M 1451061. (Of the LACM 'I'f"imen , citedabrM:. only 43'795-80 1 are the: bro.I·>lripN form.)

C/I:au d and slaim'd mal t'rial. _narrow·$lripcd form:US~M 1S9447; broad·.uipe-d fonn : LAOl .~800.

Ph.l'fI"htJIl/i bie%r Bibron in Sal!l'il 1841;\eara

Fillure 4E

1841. Phyllo/'rrlt'S Nco/or Bibron , Sagf1l : plale 29 hi" holo ·1) 'PC; MH~P 83ll. Cuba lin enur for Colom"ial.

1841. PIo,H,""II~' bico/", . Dum.:ril and Bjbr,,,,: 6.'11 -1'>.'9 .1845. Ployll"""tn m~/<III<",Io;nl<.' Bcrlhold: 43, I)'pe h::.li ly;

Popa~'iin,IC.uea,1 Colomhia [. 1760 m)18' M Phl/"!><IIe.!>icol",. Giinther: 'ltl11ISII Ph /l"!><IIe.' me""'''''!>i''''.... Giinlhcr: III11I67. P!>,-I/"hclle. mel", ,,,, !>i,,,,,,, Kcfe" tein; JS4-~5511ItW P!>."I/"/oatt'S e!>"cM n.,{, Posada: 205-207, lype I,><:al·

ily: ClVI;O. Colombia18112 Ph,"I<>i",u" bico/", . Boulcngc,: 19511I82. Ph."lI,matn mrl"""'hi""', Boulcnll"r: 1%.1920 Ph..-lI" butn bico/"'. Barhour and Sohle , 3eN1920. PIr.'I/.""',t'. "",lan". 10", ,,•. Barbour and Sobk: "1)2.1 92~b . Pir:,'Jlulo«u. niu fQr; SOOle: 1-2. fig. I , holtlt~pc :

AM~H 140211. Antic. [to.. nl. Anlinquia. C...I..,,,bi a1.13S7 mi.

1957. Piol ilob<ll hi<:-oIOI'. Dunn; n .1957. PIn-/lohat _la_hilt",. DuM: 77.1957. Pir.,'llvI><un tti«fori_ Dunn; n .1957. /knJmb<i,n rypogMpIt",. M;onn; 513 (color phoIo ).1957. (knJmba l liN'Wri",. wa....cn: 7M1910. Pir) ih>bDI bkolM t p;orl). Cock an a.nd(joia : ~5-37.

plalC 6A---e tonly lht fol ......iIlg _ IIt>I P. bfcoi",:

8M 191 0.7.11.53. USNM 48865. 122104. 145105­06. 146630-J I. 147204...(17. 1 4972~-27. ISObJ2-4 2.CAS 115151 ).

DI:/i"it;on . -Teelh present; toe, .. eblecs: donumuniform l"((I--onm~'e. ora nge . or yellow-orange : 'lripe~

absem: li~ht proximo.-enlral calf.spot ab!<nt ; venterblack . oeange, or J!oId .

Di"XntlJ;s. -Ph.\'/ft>ba,t's bic%r i~ a member of thebicolo. group , dirfe rin~ from P . aur otaenia , P.!u/(ubri., and P. vtuosus in larger size (T able 1) andin lackin~ 'lripc:~, and from Phyllohatl/S specie, inha ..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~ly

granular bad and corneumes has dorsal ~p<.lls andmarbling).

DpcripliOl1.-s- Meusurerncn ts arc in Ta ble I (t hcholot ypc i, a 39,0 mill male). The skin is entirelysmoot h. Maxillary and prema~ i Jlary reem arc pre, en!.The f irst finger is s li~h lly lon ger than rbc secon d. Thetoe, are .......bl..".

In life . in specimens frum ncar Santa Ce<:ilia andLa Union on lhe: uppe r Rio San Juan (Was..cn 19S7.-'ta nn 19 57. and my notes ). the ir i~ ""a~ black o rredd ish brown . Tho: tympanum was panl)' black andpartly )'eU(lY>'-oran~ . In some specimens. the lip ofthe snoul w"-' bhock Ohi, .....a' the reason for Benhold ' sna me P. m"lan"rrhinu.•I. The dorsum wa s uniformbri~hl or,lIIge o r yeljow-oeange. The ~ide of the bod ywa~ !!old Ol" ~llow-oran~. The <Jonoa.1 surface of theupper arm and lh i~h was gold. yellow-orange , or d irtyye llo...' ... ilh a black wa~h . The doNI l surface of theforearm and coM and lhe ventr"l surface of the limb>.were black , ""jth o r ""ithout numerous ycl lo... dOh .The venter was uniform black or black " 'ith a golde nwash (....'mc timcs imerr upeed by a black parch on thethreat) .

In prese rvanvc , in specimens from Santa Cecil iaand La UnKln, lhe dorsum and ,ide arc uniform Ii~hl

~ray ; the do rsa1 surface o f the limbs is uniform lightgray'. o r i> dar k gray with clo \Cly "'I lighl gray doh.The ventral su rfa,'e of the limbs i, dark gray ..... ith lighlgray dotv. The venter i, uniform lighl ~ray c xc..pl fora dark gray p;ltch on the thro"l in ....'me specime n"

Pre.scrv..d ,pe:cimen, from the Rio Sipi are en tirelydark I>rown.Nm",'"ci"',,rt',_Phyllobatl:~ rhoc,""ui.• Po sada

1869 (lhe de-eriplitln ""as rcpri nl..d in 1909 ) has heenconsidered a synonym o f f)f'lId,."h<J,,, hiJlriOlri,'u,Berthojd . but the ca ll (a ,hrill . interrupted ",h i'II<').the cok» ( '>r;ln~e·)c l k, ..... alll"'.: . toIuish I>htck bel"",,).the ~nout"enl Ien~th (-10 111m). and tbe finger lenl!th(t he first fin~r is km~cr than lhe second in Posada ' ,figure) are rna racler islic of P. "irow. In D. hi.\Irw""ices. the ca ll is a ~rks of 1Joan<. ","' .!ike OOICs . thecolor pan..m usuilll) is sp'1l:led (but sorne lime, rescm­bles tlla l o f r . bi""lor) . the -nout·'ent Ien~lh of 6 1

24 BULLETIN OF THE NATURAL HISTORY MUSE UM OF LOS ANGELES COUNTY No . 27

specimen, from the upper San Juan drainage averages34.2 mm (maximum 37.0 mm), and the first fingeris shor ter than the second. Posada (1869 ) "aid thatthe Indians extracted poison from P. chacoensis. TheChoco Indians today do nol extract pobon from D.hisl rionicu ,1 (Silverstone 1975). They do extract poisonfrom P. bir%r; they call this species neara, one ofthe Indian names Posada applied to P. chocoensts(Wassen 1957; a herpetologis t told Wassell to use thename Dendrobotes tinclOrius. but w assens neara isP. bir%r). Posada sa id that P. chacoensis lackedteeth. but he may not have used a microscope . Ph}'l ­loba tes ch",;o",n, i,l Posada is a synonym of Phyllo ba l<'sbicolor,

Distrib Ulion and hahitat (Fig. 14),-Rel iable eleva­tion records are from about 25 to 1525 m. P. bicoloris reported from Cuba and from the foothills of theupper Atraic and San Juan drainages. from the RioRaposo. from the Serrania de Baudo, from both slopesof the Cordillera Occidental. and from the westernslope and a central valley of the Cord illera Central(all in Colombia) , On ly the record, from the upperAtrato and San Juan drainages and the Rio Raposohave been included on the map, because I distrust theOther records.

Barbour and Ramsden (1919:77, 113) noted thatthe type locality of Cuba is incorrect , and that de laSagra shipped material from the Latin American main­land to Paris with his Cuban collections. They thoughtthat the type might have come from Mexico. P. bicolornever has been collected in Mexico, nor in Cuba sub­sequent to de la Sagra's publication ( 1841). T he typespecimen resembles Colombian spec imens and probablycame from Colomb ia. Dunn (1957) and Luling (197 1)thought that the holotype probably was collected inColombia by von Humboldt.

I distrust the records from the Serran ia de Baudo[exact locality unspecified]. Medellin [Cordillera Cen­tral, 1538 ml. and the town of Andes [eastern slopeof the Cordille ra Occ idental, 1357 m). I collectednear Andes and at eight sites in the northern andcentral Serrania de Baudo, but never found P. tncolorthere. K . von Sneidern, the collector of the " Serraniade Baudo" specimen. has stated (see below) that hehas collected P . bicolor only at Santa Cecil ia and LaUnion.

Specimens recorded from Popayan [1760 m, on thewestern slope of the Cordillera Ce ntra l] (USNM137359-62. 137364 -70) were co llected by K. vonSneidero. According to von Sneidern (ill Jill. ), "Allthe P. bicolor I have collected are from Santa Cecilia[411 m] or La Union [451 m]. .. one hour by horse-back. from . . . Santa Cecilia. The frogs comemostly from the hillsides between the two localities.I have never seen any colo rful Phy lloha/ n or D end w ·bates near Popayan; they do not exist here." Berthold 's1845 record from Popayan (P. m/'/ai10rrh;llll~ ) may beonly a shippiog point: this supposition is supported by

his description in the same paper (and probably from thesame shipment) of another Chocoan species, Dendro­bates hiJlrio'l irll .<. from · ' Papayan,"

The most t r u~tworthy Colombian locality record,are in the western foothills of the Cordillera Occidental,e.g .. Santa Cecilia on the upper Rio San Juan. Thehighest of these records is the beudwaterx of the RioSipi. which is 1525 m according to H.G.F . Spurrell'sfield notes at BM.

uH:ali/ies.-COLOMBlA , RisartJlda: San1a Cecilia (FMNH54187,541 89,54193 -94,54196,54204-05,54207.54212,5422t , 54233; USN~ 147225-26; A:"SP 25755: LACM60972,61050): vicinityof La Unionand SantaCecilia(USNM137359-62, 137364_70). Chm:D: head .....aters Rio Sipi (8M1915.10,2t ,46- 47); t20 kmS Andagoya (USNM 139776-77);Rio San Juan (USNM 144994); opper Rio Alrato (ANSP25756: USN'" 53168, 53t70-71). Vall e: virology fieldstation, Rio Raposo (USNM 151381 ).

DOUBTFUL RECORDS.- CUBA (MHNP 838). CO·LOMBIA. Callea : Quintana , 3200 m (FMNH 54183):Popayan (Berthold 1845): Atl/iaqu.; ,,: Andes (AMNHt4028 -29); Medellin (AMNH 13540, 3(08 1); O weD:Serrania de Baudo (ANSP 25490).

Cleared a nd stained malerial ,-USNM 13?360;LACM 60972

Phyllobale~ lugubris (0. Schmidt 1857)Lovely Poi,on-arrow Frog

Figures 4A. 8A

1857. Dendrobates IIlgllbris 0 , Schmidt 12- 13, Ihesyntypes; KM 101611347 (now lost), road fromB<>;; as del Toro to Vole"n Chiriqui, Panama

1858 Dm drobates Jugubr;s O. Schmidt 250. plate 2.fig. 14.

1900- 1902. Hyillp/esia tinCloria (part). Gunther; 207.1921. Phyllohares MllIridae Barbour and Dunn; 159,

holotype: MCZ 8022, Finea Victoria near u nt,[limon,] Costa Rica

1924 Ph)'lIolmres lagllbris. Donn; 5.1926. Hylap/esi" linrtv,ia (part) . MirandaRibeiro; 224.1952 Phyl/obates Jugul>ris. Taylor; 640- 643. fig. lO.1968 Phyllobares lugub"s (pan). Savage: 763_766,

fig. 4F.1970. Dendrobates Jugu!>r;s (part), Cochran and croin'

19-21 (not the plate; in the synonymy, Den.drohares lagahris O. Schmid1 and Ph)'lIobaleslugabris as used by Dunn and Taylor are PIU!lubris; oflhe specimens examined,onl)' USNMI t8231 is P. lugubrls).

Deflllilioll._Teelh present; toes webless: dorsumblack; complete gold , yellow. yellow-orange, or orangedorsolateral stripe: blue or white ventrolateral stripe;light proximoventral calf-spot absent; venter black.with blue, greenish, white. or silver marblingDi/1g"o~is , - Phyl/oblllt'S lugllbris is a member of

the bicolor group, differing from Phy/{obare~ speciesand P. bicolor in smaller size and in having stripes,from P , auro we'li/1 in having ventral marbling, andfrom P, vi/WIIl,< in smaller size (Table I) and in havingnarrower dorso lateral stripes .

1976 FROGS OF THE GENUS PHYLW BATES 25

1952.

rse

189l.

,

Description , -i- Measuremems are in Table I. Theskin is s lightly granular on the dor sum . 1!'Ie dorsal"IIrface of the limbs. the belly . and the ventral Mlrfaceof the thigh~. llIld ~mooIh on the rest o f the venter and\'entra l surface of the limbs. Ma~illary and prema~i l ­

lary teeth are pre-e m. T he first finger is longer thantile second. The roe s are weble".

In life. the iris is dar k brown . The ground color ofthe en tire body is b lack. A complete gold. yellow .ye llow-ora nge. or oran ge dor solatera l anpe e ~ tends

from the dorsal base of the thigh along the back.. a longthe outer border of the upper eyelid. and along thecanthu s rostralis to the tip nf the snout. where it meetsits fellow . A go ld. cream. whitish . or b luish stripee ~ ten(h from the arm in...:rtion along the upper lip .end ing ventral 10 the eye. The dooal '>Urf.ce of thelimbo. bears extensive go ld. yellow , or yello w-greenmarbling . The ventral surface of the limb" the belly .and somenmes the throat rear blue. greenish. white.or silver marbling. e ~ tend in g o nto the side of the bodyas a ventro lateral stripe .

An a berran t l>ptCimen from Turrial ba. Costa Riea(Taylor 1952:b41 . 643). had the Mad. ground colorcovered by a wa,h of pink ish on the dorsal surface ofthe thigh. of yellowish on the throat. and of yclhlW.green on the belly ,

In pre-ervat ive. the ground color is black: the ,;.tripesand marbling arc whue or gray.

8rrt'di"~ coli , - 1lIe ca ll is a " rasping em. lastingfour 1Usix 'c.'Collds. pau-e . repeat " (Savage 1% 8).

Re!m iorl.lhip.•, - Savalle ( 1968) co nsidered P . viI·WIU5. of the Pacific coast o f Co!ila Rica . conspecjficwith P . lugub' iJ. The central mountain s of Cost a Ricaand western Panama separate the two forms. P. lugub'iJis smalle r (Table I). usually has narrow er stripes thanP . vil/IIIUJ, and has gold. yellow. or yello w-greenmottling on the limbs. instead of blue-green moultn g;1$ in P . vil/ulUJ (Savage (968) . P. /ugub,is and P .

" i flut us resemble P . aU' Olaf!";O of the C ltooeo, butare separated from it by a d i,;.tributiona J l!ap in easternPanam a. P. OU' OI,"" io. P. lugubriJ, and P . "itlat tUmay be conspecific, but it is best to reco gnize thethree as d istinct spec ies until more da ta are gathered ,

Noml!'nduIU'I!' . -P. /u~ubris is not the DOldm/l(JII!'J/ugubriJ of Cochran and Go in (1970) ; that spec ies isD('ndmba/('J ITI1t1COIUJ (Cope 1861); see SavlIl!e ( 1968:765) .

Di sr, ibUli01/ und habilUI (Fi" . 13).-Elevation rec­ords are from sea level to 650 m (the type locality ishigher. but see Savage 1968). P . /ugub,iJ inhabit, theforests of the Car ibbean drainage of Cos ta Rica andwestern Panam a.

Ltxa/iliu.-COSTA RICA. HU N ;': juno;tion Rim Sa·rapiqu' ""d Puerto Viejo (CRE 65. 63_69); Fine. La Selvancar Puerto Viejo (CRE 512); 16 km N Puerto Viejo (MCZ75323 ). Can agD: Turrialba (FMS H 174520; KU 24907,30414. 3~g-89, 370 t7; CRE )2 ); HacOenda La f1orene;"near TurTialba (CRE 7]95); Morehouse Finea. 6.4 km 5

Turr'a lba (KU 24908) . Li", ,,,, : Los Diamantu (FMNH101097-99: KU 24909, 370(8); Unt (MCZ 8(22 ); Sllre1katKU 36794-8(1); La C_ illa. Rio R~laZOn (ASSP 235S8.239M); Ta\.aJrwlca (M CZ 9902-(3); Baja Talamanea. BambU(CRE 7182-83); Pandora (CRE 7188·89); 4 km W Guapile.(CRE 850).

PANAMA,_ BoCOJ del Toro: Almiran!e (MCZ 9873-75 ;FMNH 16102, 83468_70 , 152016; USNM 118231; KU80054- 70); 4 ,8 - 5 km W Almirante (KU 94813-21 .1156)2 - )). 116748- SO; AMSH 81450- 51); 10-12.8 kmSW Almiran~ (FMS H 67971. 153659. 153756- 57. 153762;KU 94822- 26); Isla Eoeudo de Vera~u. (KU 115634- 38);lola de Colon (KU 94827- 30. 116753): Isla Bastimento.(KU \14831 _34): Cayo Nanc)' (KU \14835); Cayo de Agua(KU 94836_38. 116751); Isla Pope (KU 94852, 116752);Id a San Cristobal (KU 94853_ 54); Peninsula Valiente .Bluefield. (KU 94839-50); mainland NW lola sem Hill(KU 94851). CIlltHt: CeITO Santa Rowo (FMS H 67970).

CIl!'arl!'d and srai'll!'d marl!',ial ._JMS 1683.

Phyl/oh/lll"J ,';I/<I/tU (Co pe 1893)Golfodulcean Poison-arrow Frog

Frontispiece I and Figure 48

lNlldrobarn li...:Torias VillalllJ Cope: 340. I) 'pc

locality: Blleno, "'re., IPunlaren ...) Co. taRi,a .

1900-1902. Hylaplr>ia tinctor;" ( p;>rl). Gunther: 207.1931. Phyilobulr J IlIgwh';J (part), Dunn: 392.1942. Dl!'IIdrobatr J linc/OTias Q"'anu (part) . Lau~nl

uDr"d ,oba/r. rilK"'orius villa/US. Tayloe-: 5&2 .Taylor u.id Cope'. lype o;ould lMJI be found atlhe USNMPiryllo!><J lr< {ugub, ;< (pan) Sava~e; 763-766( lMJI the figure)

Dtfini/iml . -Teeth present: toes IIo-cble!>S; dorsumblack. some times with broken yellow medi an dorsa lstripe; complete gold. ora nge, o r red-o range dorso­later al str ipe: wh ite or pale blue -green ventrolateralstr ipe ; liglll proximoventral calf-spot absent ; venterblack with while or pale blue-green marbling_

DiognOJiJ. -Ph" l/obat('J ";UG'UJ i . a mem ber ofthe bico!o' group , differ ing from P . lugub' ;J in largersize (Table I) and in having broader stripes. from P .auro taenia in hav ing vent ral marbli ng , and fromPh,.Uoba tl!'J species and P . bicolor in smaller siz eand in having str ipes .

Dt sc,ipt;01I. _Measurement s arc in Table 1. Theskin is slightly granular o n the do~m, the dorsal sur­face of the limbs. the belly , and the ventral surface ofthe thighs, and smooth on tile rest of the venter andventral surface of tile limbs . Ma>:illary and prema>:ilIaryteeth arc present. The first finger is longe r than theseco nd. The tees are webless

In life . the dtmum is black , and somenmes bean; abro ken yellow media n dorsal stripe (Front ispiece I).A complete gold, orange. or red·orange dorsol atera lstripe extends from the dorsa l base of the thigh alo ngthe back . along the ou ter borde r of the upper eyelid,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..A ....hne 1>Iripc exte lllb from the arm in...:n ion alongthe upper lip. ending ventral to the eye . The dorsalsurface of the hmbs bean numcrou~ blue-green dol~,

almo st obscuring the black. ground color. The venterand the ventral surface of the limbs are black withwhite or pale blue -green marb ling . extending ontotile side of the bod)' as a ventrolate ral stripe from thegroin to the arm insertion .

In pre§('rvlltive, tile color i. entirely black, exceptfor tile dooolatel'31 and upper lip stripes. ...-hich arewhite , and the ventral marbling lind the dors.il "lIrfaceof thc limbs, which arc PUfllli.h gl'3).

Brt'tdin Kroll. -P. vittallU hi s t.....o ki nd ~ of ~ingcalls (D. Travis, J't'rSOfl<lIc"",""mica/ion). One call isa shon, high.pilChed chirpin g. con~iqing of t...o 10five notes , successbely dropping in pilCh; it is emittedby the male when sitting Il('\t to the female or whenthe female i. pursuing him. The other call is longer.a high-pitched trill lasting two to three liC<:onds: it isermued by the male when he is tryin g to attract thefemale 10 the sne of oviposition .

Tudpo /t's (Figs. 10, (28) .- Three IoLs havc beentalen from the bach of ildull males in Costa Rica inJuly and August ITable 4) . A«ording 10 the eREeata logLK' . one of the~ Iot~ (e RE 903 ) came fromsmall side pools. hut A. StlllTCtt. the collector , to ldme the ladpoles came from an adult"s back. and werealjo...ed to grow before pn'M"rvalion. Two other lots(e RE 7234 and 3152. in Stagcs 25. 28. alll.! 37. takenin Cosla Rica in June) an: not from tllc backs of adultsA. Starre tt told me he found tadpoles of this spec iesin ...-ater in tree holes and fallen palm petioles, butnol in streams or side pools .

Unlike tadpoles of Dt'nJrobaln. tadpoles of P.•·;t/atu s iIl'C IlOt aggressive to"ard each other (Polder1976).

Meawremcnts Me in Tabk J_R~IUlimuhips . - Savage ( I% S) considered P . •';1·

1t/11U con. pecifrc ....ith P. lu![ubris tsee lhe rclationshipssection in the P . /ugubri5 account. above).

Lif~ hi5101",.. - Polder ( 1976) .... itnessed intra'p«ifrca~~re"i\'e behavior (jumping on the back and prc" inEdown) in P, vitnnus,

In captive anima ls (Polder 1976: D, Travis.I'","'ol/ll!('omnumif'tllj"n). courts hip lasts from one to sevenday s before ovipovition . COIIn, hip includc ~ callingby the male, circ ling, , haking of rbe limb~. j umpingon and , iuing on tllc eouni ng partner (done b). either~x). and puNling (lhe female fo llo....s lhe male). Themak choo...:s tho: site of o..ipo<>ilion (u,u all) on a bro­

mcliad leaf. -omc:times on a fern k af. r.m:l) on tbe floorof the terrarium) and attrXh tbe female 10 the ~ite bycalling.

There is no amplexus. The o"ipositing posture isType 2 (re"'mbling thaI of tllc D. lilll'l"ri". group);while the female lays the egp. the male u,u ally rc-

mains §(,paratcd from her, fa.:ing her. D Tnovis .... it·nc.scd ~iJ; (Wi~itions; in t 'o of these oviposhioes.the male left and did not atch the female: in one.the male and female remained J to 6 mm apan: inone , their noses tOliChed; and in t....o. they remainedapart, exccpe that, commencing 5 min after the startof o viposition and la~ ti nE I min, the male assumedthe Type 3 posture: he lay on top of the female. almostat a 90"angle to her bo4y .... ith his hind limh, projectingoff to one side, and rubbed his belly again,t her back.The female cont inued to la)' eggs .. bile the male lay onher. The male' s cloaca did not approach eilher the e~s

or the female' s cloaca . This behlvior. ",'hich resemblestha t of Culo.Jlnhu5 pul"'<J11U (Liiddecke 1974). mayha~e • stimulat ing funclion . lIiding the maintenallCC'of a proper behavioral seqeence: fen ilization does notocc ur al this time. During oviposition . which las"15 to JO min, the female e ither turns (rom side 10

side (rem aining I min in eac h posit ion) or c ircles .appar ently 1O distribute the eggs . After nvipo xition,the female dese rts the eggs ami never visits them again.

Usually, as soon as the female leave, the eggs. tbemale , its on the eggs ; fertilization prohlbly occursduring this fir'>! visil. In four breeding cycles thatTravis ....itnessed, the male al....ays visited the egg,twice on the day of oviposition . His fU"Sl visit beganimmediately after oviposition . and lasted 5 to 65 min.After an inter·. a! of ahoot 30 min, he at:ain loat on theeggs for 60 to 90 min. The male then desened theeggs for one ....eek , During tile second ...eel; afte rovipositi"n. the male visited lhe eggs once a day andsat on them (perhaps to I!\\\isten tbem) for 5 min: thethird week, he visited the ej,\gsone to three times a day.

Polder .... itnessed tadpole -carrying once . Polder 'smale .....t on the jelly·m"-,, and made stamping move­ments; all 13 larvee ...nggled onto the mak's back.11lc maar carried the larvae three day's and then re­Iea<ocd them in stages (i.e .• not simultaneou,l y) inlowater in a pl1s.tic dish. Jerking movements b)' thc:malemay ha~'t stimulated the larvae 10 Iea>e hi~ back.

Travis .... itnessed tadpolc-carr)"ing rwke. On the firsloccasion , Travis's male took up his tildpoles on lhet....emy -third day after u~ipl.,it i\\n (one .. eek after hatch­ing). The male apparently stimulated the tadpoles towriggle \\nl\\ his back by shaking his posterior end inthree or four vhort bUNs (four or fi~e sha~es per burst).Tadp .le uptake la,ted 10 min. He carried thc: six tad­poles in ' ....0 "ages; in thc: first stage , he carried fourtadpole, and retained tbem 1w0 days . After releasingthem into a , mall pond in the terrancm . he returned thesame day' to thc: leaf nil . picked up ,he remaining t...-otadpoles. and released Ihc:m the:next da)'. On tile SC'OOndO«llSion. the: male 100._ up his tadpoles on thc: seven­Iccmh day after oviposinon (day of hatching unknown).He again carried tho:tildp"le, 10 tile pond in t...o stages(four tadpulcs carried tW\\days . the remaining four tad­p>les carr ied one day).

/97{, FROGS OF THE GEI'WS PHYLLOBATES 27

One of Polder's female s laid egp everyone 10 twoI>< ceh over a three-mo nth period (eac h of Travis' s IWO

females laid eglls every twc WC'Ch over a two-month

period) . Keown c1ulCh sue is seven 10 21. In ("a",ivity.lhe period from fenilizalion 10 hatching is 13 10 17days: lhe per iod from deposition 10 metamorphosis is40 days at 22 to 25°C.

Stomach co-runts (o ne spccimenl.- From C RE7235 : H YM ENOPTER A : Fc rrnicid ae: Was",a""iaaUrap,ln Clala ( Ro ge r) (I). Smilnislruma sp. ( 4 ).

Dolichodt'rus ~p . ( I l. Paratrt'china "p. (2 ). ROKu iasp. (2). Phd dolt' so. ( I ): COLEOPTERA: fragments;DIPT ERA: MyeelOph ilidae ?: fragme nts ; COL LEM ­BO LA: Podur idae"!(I).

DiSt'ust's . _Polder ( 1976) h s discu s-ed d i-ea -esof Cllpli\.-cP. ,·illa/us.

Distribution and habitat (Fig . 13).-Elention rec·ords arc from ]0 10 70 m. P. "illa/us inhabits the wetforests of the Golfo Dulce region on rbc Pacific coastof Costa Rica.

Localitit's.-eosTA RICA . "'",ur"." RincOn ok o.a(MCZ 75322: CRE 705. 899. 3076. 3114. 6391. 72341:Rio 1'0:......"""". 7.2 km S RinOOn de o.a (CRE 7235): 4.5km W RincOn de Osa(KU 102161-6)); Camp Seanlc. 6.4km ~p Rio Rin" ,n (e RE 757): Quehrada Bnruca, PalmarNo,-u, (KU 93924): 22 Ion E Palmar Norte (KU f>5296_) OH ):Rio 7..ap-'le. 8 km E Palmar None (KU 93925_30. 93934- 3.5):Botnoo Ain:s (CO(><' 1$93).

CktJrt'd and slai",d ",utt'rial. -JMS 1681-82.

/ emor<l fis group

This group includes si:t species (f'hyllobcltt's u",hon.\'i.P. bo ..l~nf(t'ri. P. npi"owi. P. /mwr"lis . P. trirolor .P. :apuro ).

D~ji"ilio,, ; )noul-vtnt length o f adu lts ~mall tomoderate ( 15.0 -33.5 mrn); basal web bing betw eentoes lJ· Ill ·IV; pa ired st r ipes late ra l (Figs . 3F, 5A.D-F); light pm~imoveotral calf_,pt,t abst ol.

Phyllobutt's "",Ito ")'i Soble 192 1Anlho n)'s Poiwn·arrow Frog

Figu res 5D. 7A-B

1921. PIt)'lIob<.ltt. <Jmltolty; Noble: 5-6 . fill 5. holot)pc :AM~H 1)739. S.ah'w. EI Om. &owdot'

D..ji"i/io" ._Teeth present: toes basally webbed:do rsum (in prcservanve j da rk with light med ian dorsal' tripe: com plete light lateral 'tripe: light ventro lateralstripe ; Iighl pro~imo,totral calf.spot abo;enl: venter darkwith light marbling or ligtll with datk marbling.

Diuf(nosis .-Ph-, 'ffohal~s "Il/hon,vi i, a member ofrbc /t'm"roli s gro up. differing from P. 1x,,,lm!:t'r; inhaving a light media n do rsal , tripe, [ro m P. opino"ai inhaving a light med ian do rsal , tripe and in having a COIJ1·

plete light lateral str ipe (Fig . f B: inco mplete in P.~Jpi"owi. Fig . IA). fromP . / t'm" rafis andP .zupuro inhaving ill light med iall~l §lIipe iIInd ill smaner size.and from P. lricolor in having while bones (~ll in P./firolor) and in smalle r size (Tab le I ).

Dt'scr iptirm. -c-Measuremems arc in T ablc I (theholotypc is a 21.0 mm female) . The skin is smoo th orslig htly granular on the do rsum (e:tCcpl o n the poste riorbac k. where it is tuberculat) and smooth on the limbsand the verner . Mu illary and premuillar)' leelh areprtstnl. The firs t finge r i ~ longer than the SC'Colld .Basal webbing occ urs between roes ]I·lII -IV.

The co lor in life is unknown , In preservative. the dot­sum is dark brown with a co mplete gray or wh ile me­dia n dorsal sm pe. eapanding an teriotly 10 COvtT theentire donal surface of the l>/lOUl; the l>/lOUt sometimesbears dark bro wn ~poIs. The side of tbe bod) is dartbrow n, A com plete gray or while lateral stripe extendsfrom the groin to the upper eye lid , along the o uter

border o f tbe upper eyel id. and along the cant hus roostrali~ 10 the tip of the soo ul. " here it mttl~ ib fellowA gra~' or ...hite ventrolateral Slripe ex tends from thegro ill 10 tbe f~limb in-e rtion. alxM: which it unileswith lhe upper lip slripe: tbe latter jo ins the canthalstripe at the lip o f the snout. The dorsal surface of theforelimb usually is gray or white ; the anterior and pes­lerior surfaces o f lhe fore limb bear datk brown loegi­tudrnal stripes. The donal surfece of the hind limb bearsaltemaling gra)' and bro"' D bands: the lhi~ h bears a...hite o l>lonll pro:timodors.al ~ptJl and loOmC: l ime~ { a~ inthe hololypel a while lon~itudinal po, tt'rodor~al , tripe .The venter so met imes is predominantly brow n. andsometime, is predominantly gray or ...hite; these varia­tions OCl:ur in tbe same populalion. For example, in a..eries from Huas imal. Pens, lilt venter and the ventral!oUrface of lhe limbb are dark hrown with gray or whiltmarbl ing in some specimens. and while with hrownmarbling in other s.

In a '\Cries from Alamor , Ecuador . the venter is darkbro...n with extensive while marb ling in some: speci ­mens: in o ther specimens. the marbling is reductd to!>mall "hile spols .

In a p;eralype (Fig. 7B) from Salvia' . Ecu ador. thevenIer and the ventral surf"cc of lhe limbs " re white .except for a few brown spots on tbe throat. on rbe s idesof the bell y, and on the distal portion of the IhighsIn the boloeype (Fig . 7AI. also from Sabias, tbe ventralmarbhn~ is about even ly divided hetl><'een bro...n andwhile .

Tadpoles. - One ku ha, been taken from the back ofan adult male in Peru in April (Tah le 4 ). The bead-bodyof each ladpo le is ':U mm long: the lai l i~ damag ed\I.·hell caf-(urtd among maidtnluir fem s on the ban" o fthe Quet>rada AngO'olOra. Pow Azul. Peni. lhe maleca rried eight to 12 tadpolc:~ in a dark. fluid -filled "blis_tcr" on his posterior do rsum (B. Barnell. letter of 25September 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 "blis­ter" was not an unhatched egg O1a". Thi, "hlister" i,unknown in other de ndroba ti,h , which <'arry their lad ­pole, without a proleclive ,t rU<:lure .

Sloma"h ""'lICIII,\ \ IWO spccimcns j.c--From A!',IN H/6416: HYME~OPTERA: Chaleidoidea ( I); ARAC H­NIDA: P,<udu'iCurpionida (I).

From AM ;o..:H I647!/':COLEOPTERA (2j .Di .\nihurion "lUI Iwbi'", (Fig. 14).-Eleval ioo rec­

ord, are from 153 to 1387 m. P. (Im },o"yi inhabilssouthweste rn Ecuador and northwestern Peru, west ofthe Andes. Salvia, ( I0611 m) and Pono veto (610 rn) arcin the humid tropical zone . allhough most of the terrainncur Ponovclc is tree Ie" and arid ·appearing; Alamor(I .'oll7 01) i, on a rid!!e between heavy subtropi cal forestto the west and north and the humid tropical zone in thevalley s below (Chapm an 1926). P , aw hollyi inhabi ts" fairly dense. very well-watered fores t" at Pow Azul .and " open cou ntry surrounded by much lighler fore stcountry. subject to ann ual droughts" al Gramadal: bothlocalities arc belween 153 and 305 m (B. Bamcn.Jcuersof 25 -26 September 193 1 [0 H.W. Parker at B~1 ) .

Apparemly . P. "whollyi is not found in ar id habitats,but in a nalTow tongue of hum id tropical zone fores tsalon g tbc western slope of the Andes (Chapman 1926:plate I) .

Loca li / ies ._ECUADOR. £ / 0 1'()/ Salvias (AMS H13739--4() ; Pomwelo (AMNH 13741-50 , 13951_59). Loja:Alamor{AMNH 16210,16348, 16411_91, 16493- 96, 16498­518; UMMZ 89982); Calamayo (GOV 9315. 9318- 19)

PERO._ Piura: near Hu. simal. Am()t.pe mountain• . onCase.dem 1r.il (FMNH 9840- 42, 9844_46 , 9848_49,9853_56, 9859_60). Turnbe:: POlO Azul, Quebrada Angos.lura, nonh of Rio Ttimbcl . one hour ride [by horse1 fromE<:uadi>r;an horder (BM 1931.10.7.1); Gramad.1, HaciendaCas;las ( si~hl record by g, Barnell. leuer to H.W Parker.1 BM)

CI/'(l1'<,d IItld s/uil/"" mmuia/. _AMN H 16416, 16478

Phyllobll/<'S houlcl/g.-ri (Barbour 19(9)

Marbled Poison-arrow FrogFigures I B, 5E, 7E

1905 Prosthnapi.l femo1'alis BarhoU': 101, 22 ,)'nlype"MCZ 2422; USNM 52406, 118132-33; BM1947.2.1 3.92_ 93; UMMZ 48070, h la Gorgona.rNari~o, ] Colombia

1909. Pro.<therapis bow/ellgeri Baroour; 89. Replacementname fur Prosrhen,pis femo1'alis Boroour 1905, pre­occupied by Pros/herapis femo1'(jlis Boulenger 1883.

1920. Phyllabaus bou/ellgaL Barbourand Noble; 402.1957. Phyllobale.< boulengeri , Dunn: 781968. COIOSle/hus boulellgni . Savage: 757.1970. Phyllobales boulengeri . Cochran and Goin: 38-40.

plate 7A- F.

Deftnilio/!. _ Teeth present; toes b.,ally webbed;dorsum brown. often with dar ker or lighter spolS or

marbling; l'omp!ele yelh'" or red lateral 'tripe; lightproximovcmra! calf -spot absent : venier white withblack marb ling or black with white marbling.

Di<iglwJi' .-Phyllol)(II<,., l"'u lell8l'1'i is a member ofthefemowli., group, differing from P. wl/hollyi in lack ­ing a light med ian d"rsal stripe , from P , e.,pillo.I'ai inhaving a brown dorsum in life (red in P. e.'pillo.,,,i ) andin having a complete light lateral stripe (Fig . 18 : incom .plete in P. <"lpillo.,ai. Fig. IA) , from P. [emoralis inhaving light markings on the throat {usually entirelyblack in P . f"mowli.l) and in smaller size. from P. lri ·('( )/o r in lac~ ing a light me<lian dorsal str ipe and in hav­ing white bones (green in P. tricotorr. and from P.lUrmro in having a brown dorsum in life (red in P.l.il f'<l w l and in smaller size .

Descrip tion, -c-Me asuremc ms are in Table I. Thes ~in usually is ent irely ,nwoth [exce pt the posteriorback , whil'h is tubercular in 'orne spel'imen,j , In a fewspecimens, the dors um is slightly gran ular. In one spec­imen (GOY 9231) , ee entire dors um bears scattereddar k brown tubercles. Maxillary and premax illary teethare present. The first finger is longer than the second.The th ird finger is swo llen in ma le, and not swollen infemales. Basal webbi ng occurs between toes lI·m·IV.

In life, in specimens from La Guayacana . Colombia(my notes), the iris was blac ~. The dors um was brown.The side of the body was black with a complete yellowlateral stripe , The upper lip lInu the proximodorsalsurface of the upper arm were whitc. The dorsal sur­face of the thigh was black with yellow spots or twoyellow stripes. T he digits were banded bro wn andwhite. The venter and the ventral surface of the limbswere black with white marbling.

In specimens from the lower Rio Calima, Colombia( I. Cabrera. notes al USN M), the lateral «npe andthe dorsal thigh markings were red. The venter wa,gree nish or white wilh blae ~ mar~inp.

In preservalive , Ihe dorsum is light brown with darkbrown spots and marb ling. The side of the body is darkbrown . A <:omplele white lateral stripe exte nds fromthe groin to the upper eyelid , A white str ipe exte ndsfrom the forelimb insertion along the upper lip to thetip of the snout, where it meeh ils fellow . In mostspecimens, the dors al surface of the limbs i, orangishbrown with broken dark brown transverse bands. Insome specimens. the do rsal surface of the limb, isdark brown with narrow while transverse bands or anundulating white longitud inal stripe on the thigh , Theproximodorsal s urface of the upper arm is white. Theproximal portion of the postcrodorsa! sur face of theth igh is dark brown with a white hook mark. Darkbrown longitudi nal stripes adorn the anter ior surfaceof the upper arm, the posterior sur face of the upper armand forearm, and Ihe anterior surface of the thigh. Theventer and the ventral surface of the limb, are whitewith da rk brown marbling (Fig . 7E) or dar k brownwith white marbling , In a series from lmbili. Colombia.

/976 FROGS OF THE G£ \ 'I.JS PHYllOBAl ES

both cui".. p"uem~ occur ; in ...:>fT"Ie of lhe.e 'P"imen~.

the veeeer i, .. hile .. ith ,mall clol.cJ~ set dllfl t.u.. n~pOb . The: throar ma~ be pmiominamJ~ "hite IFi~

7E) or predominantly dar l bro" n: it u,pally' h.a ~ awhite median longitudinal ' lrip;:. In lhe Imbili .,;:rie~.

the postcrove mral surface of the thigh is darl brownwith while does: in mo-r ~ pce imcn s of lhis 'C r ic~. the

re-t of lhe ventral surface of the lhigh i, while Wilhdarl brown spots. bomclime, mluced in number. e.g ..Fi ~ . 7El or marbling . In one specimen from Imbili andanother from La Guay acana. the ventral ,urfa..-e of theth i~ h is dan t.uwn .. ith t....o undulalin~ "hite Iongitu ·diNI .lripe~

T<JJpo/n (Figs. llA. 12Al. -Two 10h 1\;1.(' beentalen from the Ncl. of adull male. in Colombia inFebruar) (Table ~ l . ~lea>urcmenb arc in Table .t

Savage ( 1968:757) placed P . /x'ult' tI!:..,i in the genu,Cotostethu s, Hc considered il clo se ly rcla led 10 C.nub;"" I" (Dunn 1924) . and ,a id (IX1 sed on unpubli,hednOle~ by E. R. Dunn ) lhal the nHlulhpans of lhoc lad·pole, of P. boll/fIl lI..,i are umbelliform. resemblingtho>;: of the ladpo le, of C. ""bieola . Tadpo le' (US ~M

145248) from lhe bad of an adult ropotype of P.fmu/ulg..,i . however. have mou thparh ~ im il ar tothose of ot her spccie~ of nnHobatt'3.' lhe moulhpartsare not umbcUiform (Fig. 12AI.

MOIt'rio l of " nerrlai" Jla lu• ._Spce imen~ fromTandapi. Ecuador. 1460 m. on lhe ....eslern ~k>pe ofthe Ande. ea.lWII lhcar.t of San to Domingo de 100Colo­rados IKU 111529-61. 120f>65- 68 ). belong 10 lhef/!'mo ro/i.1 group; lhey resemble Colombian spec imensof 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) Iighlspols . Some specimens IGO V 9 148 - 50. 9229-30)from the Chocoan 1o....1ands of Ecuador (Rio Blanco andSan ~Ii~uel de I<:>s CoJorado') resemble the specimensfrom l anda-pi . Pn;M.>on,ally. I h3ve referred a1lthe~

specimens 10 P. boul~ng~ri. but I have uc1udcd themfrom lhe lmlp and Ioeal it~· Ii"-.

SlOmadl ronl~nlS (t....o specimens).-From US....M145253 : HYME:-<OPTERA: Formicida~ : S(>l~n"psis

sp . ( .5). Ph~ido/~ sp- (1). Iridom.rrmu sp . ( I) .

M)'coctpurus . mithi (Forel) (I) . My martd ae ( I):COL EOPT ERA (I ).

From KU 111529: HEMI PTERA : Aphidid ac (3)Nom~ndal..r~. _1 des ignale BM 1947 .2. 13.93. a

20 .0 mm fe male from Gorgona Island. Colombia.collected belllleen 19 June and 2 July 1904 by W ,W .Bro...e , Jr.• as the IeclOl)pe of ProslN rapis boulmxrri

""""'" 1909Diuril'JlllilJtl and IuJmtOl (Fig . 15).-EJevalioo rec­

ceds ( not includin~ Tandapi) are from 10 to 1145 m. Pboul~"grri inhab its the dense . .. et fore sts of hla Gor·gon.;( (aboul 32 km off lhe PacifIC COO-~t o f Colo mbia)and the wet southern Choco an reg ion from lhe lowerSan Juan dra inage of Co lombia south to northwestern

Ecuador . I . Cabfe~ found specimens in weI ~in foreston the lo wer Rio Calima. Colombia: lhe)' j umpedrapidl )' and "ere difficult 10 seize .

u>c"lit;t's,-COLOMBlA. S ari ii ,,: h la G"r~na (MCZ2422; UMMZ48070; IlM 1'147.2.13,92_93. 1926.1.20.123_125: USNM 5241lt>. 118232_33. 145248-3 00: AM."'H 7494,50970-72): Soolh G"'ion. (AMNH 62894_9M): GorJ(lnilla.a penin'ulo of 1,1a Gcqona. i", JalCd at hi~h lide IAM1s'Hb2892 -93 ): Imbili. Rio Mira (USS M 147431_50): LaGualac""a (LAOt blO34- H . stoss» JUJlin(LACM 60973.61036-42) , Vlrlu: Camp CanOn de Colombia. lo..er RioCali.... CUS" M 149723-2 41; R"" kd cl ler~ion Viroior.f'.dd SIal..... Rio RajlOJO (US" M 1513'79: LAO! bl0431:CMe ..... (B ISH u:m).

ECUAOOR.-E. ..........." Cac""bi CBM 911 3.1 31: SaftJ:l'Iier IBM 1901.3.29 44); PlImbclaJ (BM 1901,6 ,27.11);Tambo Alto. 10 km SW lila (GOV S47b. 84781: Quinindi(GOV 9231. 9391-92) . l'id .int:1uI: Rio Blanco Ileac tnOUlhof Rio Yambi CGO V 9235_3b): N.",,~ahlo (GOV 9227);Rio Silanchi (GOV 9599-%00); 18 km W Samo D"min~'<) de10' Colorado,. on rood 10 Cl>one (GOV 9232 ) . ea relt i:P:uamha(BM 98,3, 1.1- 2)

Clrllr ..d and s/(li"rJ ntlltt't'ial.-A~t~H 7494; USI'< M1 452~3; LACM b0973: from Tandapi: KU 111529.

Pll.,'lIobaln ~.pinfJSQi Funlltc:>U--e r 19.56TUTqllOi.e-bellied Poi'OO-al"l'O\>- frot:

figures IA. 5F. 7F

I~ I' I.yllob." t'J rs.plllOJtli Funkllooo"Ct: 7b- 78. pi"", I.fi~ •. 2_3. 110101)'"" , CAS·SU 1cr.'i77. Hacienda B·pi""'"'-. 9 km W Sant" Domingo de 10> Col<>nodos.Pkhineha. Ecuador. :105 m.

Drfi nili,m . - TeClh present: toes basally webbed;dorwm red; incomplete tUTqllOi.,;: lateral slripe : lightpro~im",..emral ealf·~pOI absent; venter darl bro .. nwith I UTq uoi~ marbl ing or spots.

DillXno.i3, --Ph.l·II0b0I~J ~3piNJJlli is a member ofthefrmorolit group. differing from P. onthoto}'i in !iKk·ing II hghl median doNllolripe and in having afl eccm­plctc l i ~hl lateral stripe. from P. boul~n~ri in hlo\ing ared don.um in life (brovo'n in P. bo,.l~n8~ri) and in hav·ing an incom plete li~ht laleral stripe (Fig . IA; the laler·al 'lripe i~ complele in P. Un/hOlI)'i and P. b<,ul~ngeri.Fig. I B). from P. f .." wr" li" in having a red dot'l; om inlife ( bla~l or brown in P. fn ,wraliJ). in having an in­com plete l i~ht 13teraJ ~tripe (complctc in P, Irnwmlis).and in smalle r siu (Table I) , fromP. Irico/or in liKli nga light median dorsal smpe, in hll'o'ing an in,,:omplelelight lalcnl wipe (com plo:te in P, /rico/or ). in having..'hire bones (gree n in P. rricolor). and in smalleT size ,and from P . zoparo in .maIler size. P. n piltlll<li r¢.,;:m­bin P. pan'ulllS in color and peuem, blll differs;from P .pan'ulltl in llaving basal loe ..-ellbing (P. pan'ulou !l.ehtee ,,-dlbing).

Dri cription._Mcas uremenls are in Table I (theholoty pe is a 17 .0 mm male) , The skin i. slighlly

30 BUUET1N OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUf,7Y No . 27

granular on the dorsum and smooth on the limbs andthe venter. Maxi llary and premaxilla ry teeth are presen t.The first finger is longer than the second. The thirdfinger is swollen in males and not swollen in females.Basal webtung cc curs between toes II·III -IV.

In life, in the type series (Funkhouse r 1956), thedorsum was dark red. The side of the body was blackwith an incomple te turquoise diagonal lateral stripe .The dorsal surface of the thigh was black whh wrquoisemark ings. The dorsal surface of the cal f was dar kred . The venter and the ventral surface of the limbswere dark bmwn with turquoise marbling or spots .

In preservative , the dOl'mm and the side of the bodyare dark brown. An incomple te light lateral str ipeextends from the groin about one- fourth of the distancebetween the groin und the upper eyelid; in CAS-SU10375, Ihi, stripe d i" o lves into spots (Fig . lA ). Abroken light stripe extends along the upper lip, Theground co lo r of the dorsa l surface of the limbs i,dark brow n . The proximodorsal surface of the upperann and markings on the dorsal surface of the thigh(a prox imoanter ior lon gitudinal mark , a prox imo­po~terior hook-mark . and ill-defined transverse hands)are light. The venter and the ventral surface of thelimbs arc dark brown with light marbling (Funkllouser1956;plate l , fig. 2) or spllts (Fig. 7F in this paper)The throat usually bears a light longitudinal medianstripe

Tadpoles (Figs. llB . 12C).- Two lots have beentaken from the hacks of adult males in Ecuador inOcto ber {Table 4) . Measurements are in Table 3

Funkhouser ( 1956:77-78, plule l . fig. 3) describedthe tadpoles: he said they have a median anus. This isincorrect; the anus is dextral. as in all o ther knowntadpoles of Ph.1'll" lw le, .

Mot erial of uncertaill ,wow.<, -A 19.0 mm male(CAS-SU 10340) fmm the Rio Suno . vapo. Ecuador,400 m (east of the Andes ). resembles the types ofP. e.,pi'lO.wi (from west llf the Andes), e xce pt thutCAS-SU 1034() has a large white spot on the throat.This spot looks like an artifac t overlying the norma lwhite marhling. Funkhouser ( 195(1:77) did not devig ­nute CAS-SU 10340 as a type because of its lllcalily.I am not certain that CAS·SU 10340 belongs 1\) thesame species as the ty pes llf P. e.lp illo,\(li.- [ haveexcluded CAS-SU 10340 from the map and localitylist.

Di.'trihll/iOIl W il l /I<I/>i/<I/ (Fig. 17). -c-Ele verio n rec­ords are from 305 to 500 m. P. espillo'lli inhabits thewet Chocoan region west of the Andes in northwesternEcuador . It is diurnal and Jives in damp shaded areas of

the j ungle floor. The j uvenile paratypcs were taken onthe bank of a small river. The movements of specimensof P. o pino.,,,i arc quick and "nervous." Thc fro!!sseck shelter unde r terrestrial debri s, never in the water(Punkbouscr 1956) .

Locali/i e.I. _ECUADOR. Pit:hincha: Hacienda Espinosa.9 km W Santo Domingo de los Colorado> on road to CJxme(CAS-SU 10577- 83. 10585- 86: USNM 145804); SantoDomingo de I""COlntadlls (CAS-SU 10375;CFW 4399 alUMMZ)

Phyllohat esfemorolis (Boulc nger 1883)Brilliant -rhighe d Poiso n-a rrow FrogFrontispiece 11 and Figures 3F. 8E

1883. Proslha<lp;s f em"wlis Boulenger: 635. plale 57.fig. 1. two .yotypcs: BM 1'147,2.14,21- 22, Yuri­magua>. Huallaga River. [Lorero. ] Peru

1920. Phylloh<Jtu inguinoli.< , Barbour and Noble: 4011920 Phyllobales /<·morc>lis. Barbour and Noble: 401.1935. Phyl/ob<Jtcs fmlO' <lii.<. Parker:5061952. Phyl/ohmcs f emom lis. B. LUll: pl,'e 2. figs. 7. 9 .1952. Phylld"acs j cmorolh . B. LUll and Kloss: 671-6731957. Phyll<>bnres femoralis. Dunn, 78 .1970. PhyllobolCs fe m<lrali, (part) . Cochran and Goin:

40-42. plale 8A-C (only lhe following are nol P.jemor" Ii.<: in tile ,)'nonymy: DClllJrohtl/CS " "rome ni"Boulenger. Phylloba/es o"r%cilia as used by' Dunn:in llle sp..,,,imen, examined: USI'M 127202, 127210_13, 146863- 66.1 47268. 151382).

1970 Dmdrobatc.< """"eli (partl, Cochran and Goin: 19(only R.INH 61789 i. P, frm,,,,"i,).

Deji"i/im ,. - Teeth present: toes basally webbed; dor­sum black or brown; complete yellow or gold lateralstripe; white, bluish while, llr silver vent rolateral stripe;light prox imoventral calf -spot absent; venter black ,brown, or dark gray with white or bluish white marbling(usually nOI extending onto throat).

Di<l/;'IO.I'i." - !' !lvl/" /><I/" " l<'III"I"IIIi,I' is ~ member oflhcfemomli., group. differing from P, am/"'''.1'i in luck­in£ a light median dorsal stripe and in larger size. fromP, boule"xeri in usually having an entirely black throat(tbc throat has ligh t markings in P . bou/e>lgeri) andin larger size, fmm P. e,'p illo.I'<I; in having a black orbrown dorsum in life (red in P. e.lp itw,mi), in havinga com plete light lateral str ipe (Fig. IB; incomplete inP. espmosai. Fig. l A>, and in larger size . and from P.tricolor in lacking a light mediandorsal stripe and in hav­ing white hones (green in P. trico l" r) , P. jn/lorl/lisclose ly resembles P. w pttro in s ize ~nd in having a shorttars al fold (e xtending along the d istal one-third of thc dis­tance between thc inner meM arsul tubercle and the heelfold). P. femondis differs from P. zaparo in having ablack or brown dorsum in life (red in P. wptm», inhaving a complete light lateral stripe (usually vestigial orab~ent in P. W p<lro), and in having a largc promincntlig,ht proximodorsal thigh-spot (small or absent in P.zapurov . P. fmwra li., resembles and is sympattic withP. pie/l<s , bill differs inlarger size (Table I), in usuallyhaving the upper lip stripe meerln g me canthal stripe ontbe tip of the snout, in lacking a light proximllventralcalf-spot . in having the first finger longer than the second

/976 FROGS OF THE GES US PHYUOBATES J I

(.00...1«juallo!he seccod in P. piC/ILl). in having ba..altoe ...ebbing (...ebbing is absenl in P. picllU).lIlld in hav­ing I« th (P. piCfUS asually lads t«lh).

Descripnon, - Mea'urement' are in Table I . Thed in is granular on the dorsum and the dorsal surfaceof the limbs, and sm\1'l"th un the venter and the ventralsurface of the limbs. Muillary and prema ~ ilI ~ry teethare pre\Cn!. 'The fiN fi n~er is lon~er than the secondBasal webbing OCCIin between tues II-Ill-IV .

In lhe fo llowing compo."ite desc ription (ootes byJ.D. Lynch at KU. W.R. Hey-er and P.A. SilversloneaI LACl\.I)of specimens from Santa Cecilia and limon ·COl:ha. Ecuador . from 5 km north of Pueno Asis andEI Pepino, Colom bia. from Teresinha and Serra. dol'avio. Brasil. and from SaUl Ton ue. French Guiana,a color llOIed only in specimens from a certain coonU)'is follo....ed in parenthe\Cs by the abbreviated nameof that country. In life , the iris was black with a goldrin~ around the pupil. brown with red flecks (someEe .). or bronze (some Ee . ). The head and hack wereblack or brown (Ee. ). The side of lhe OOdy was black.A complete yet low or go ld lateral sn ipe extendedfrom the groin to the upper eyelid. along the outerborder of the upper eyeli d. and alon g the c anthusrostralis 10 the tip of the snout, ...here it mel il5 fel­10",'. A ",hite (Co .. Br.), silver (F.G .), ur gn:enishbronze (Ee.) ",ripe utended from the f~hmb inser­tion along the upper lip. u., uaIly meeting the Cantllalstripe on the tip of lhe "!lOU!. A white. bluish white.or silver (F.G .) ventrolateral stripe extended anteriadfrom one-half the groin -axilla distance up tn the entiredistance. Ventral to this stripe ran a second ventre­lateral stripe (originating from the ventral marbling).The venlrolateral stripes sometimes ruptured into spots.Masl of the dorsal surface of the limbs was light brownor gray ....ith scanered black spots. The proximodorsalsurface of the upper arm was yellow or orange (F.G.).The a,illa ""as ~ello",' 01" OWlge (F.G.) . The lXiIIII')'color united ",'ilh three light str ipes: ( I) the upperlip s!ripe; (2) a "ripe on the ventral surface of theupper arm twbere it was yello"" or orange [F.G .))and forearm (where il ....as .... hire): (3) a ....hne stripeon the anterior surface \1' f the upper ann, separatedfrom the second stripe by a hlack area. and usuallymeeting the second stripe on the forearm. The pro~imal

half and the posterior portion of the distal half nf thedorsal surface of the thigh were black; the black co knsurrounded a conspicuous oblique yellow or oran~

(F.G .) spol on the base of lhe thigh. The spol some­times utended along the eeure length of the thigh.Sometimes a small 'P'1I of lhe~ color lay distalto !he Ihipa-spoI (Fig. 3f). The venter and the ventralsurface of lhe hind limbs were black. brown (someEe.). or grll)', and (e' c-epl for the Ih1Oll t) bore ...hiteor bluish ...hue marbling. In some specimens. theventral ground colo r was ",'hite , and the black was

reduced 10 small spols. In small juveniks, the ventralareas lhal were marbled in Jd uhs were unifOllll gray.

In preselVative. yellow, orange, and gold becomewhhe. The ground color is black or dark bro....n. Rarelyte.g .. in TNHC 36498 from the Rio Pachitea. Peru).the belly is almost immaculate white. The light ventralmarbling never extends onto the throat. except in onespr;x:imen (KU 14436 1 from Nevan, Peru, in the upperPachirea drainage). Th is spec imen resembles mos tspecimens of P. f~m",uljs. excepl Ihat ( in peeserva­tive) the throa t is marb led with whlre . the belly is",hite ",'ith a dark bro""n transverse band formed ofmarbling , and the donum is light brown with dadbrmon marbling8rr~diltg cull. -P. f~-.rQI;s "calls from lbe fOKSl

floor in the daytime. a loud. slow mllSical trill lastingfour seconds, repealed at irregular intervals , spacedusually at least 0 ne minute apart" (O. B, Wake andR. Etheridge. field n" les al CRE). In Suriname , malescall from the ground or from twigs and leaves up toone meter above the ground; the call is high and pro­longed. resembling a cricket's chirpi ng (Hoogmoed1971).

TuJpoIu (Figs. II C. 12D).-T....o lois h<lve beentaken from the ba;ks of Jdull femalc:s in Eeuador inJune: lIlld in Surirwne in July (fable 4). An<.M:her 101

(AMS H 70995. in Stages 29. 30. 34. and lWlSformcd )is not from the ba; k of an adull. M.S. Hoogmocd foundtadpoles of this species in a ...-ater-filled fallen palmpetiole in Suriname in February (Polder 1976). Meas­urements are in Table 3.

M im ic r y . - P . f<'lIwrulis and P. picla,' may beMuellerian mimics. They are syntopie, brigbtly colored .probabl y poisono us. ar>d resemble eac h olher whenseen from above.

D;....us~s. -Polder (1973) has discussed occurrencelIlld lrealmenl of cramp of the hind limbs lIlld fungalinfections of the eyes in caplivity·.

,"·omrltcl(1l"r~. -1 desig nale B~I 1947 .2. 14.21. a26.0 mm male from Yurimagu as. Huallaga River ,[Lo re to.] Peru , collec ted by Dr. Hahnel, as theleclOlylX' of Pro.,~,upis f~nr"'(1lij Boulee gcr 1883.This is the specime n illustrated in the original de­S<;rlptiun.

DiJI,ihutiof/ und Iwhi l<'l (F!):. 17). -Eltvation rec­ords are from 14 to 610 m. P . !;:mo rulis Inhabits thelowland forests of Gu yana, Surinam e . ant! FrenchGuiana and of the Amazon drainage of Colombia,Ecuador. Peru. and Brasil. I look a specimen al 1745 hin fores! al Saut Tortue . French Guiana. after a rain.on falk n Ic~s on the ground. J_ Lescwe (in lin .)found Ihi~ species in open microhabitats in primll')'or secondary forest in rnl'lCh Guiana

U>culi/iu.--COW MBIA. "",,,,,,,,,-0: Umbria (FM)\HbI789- 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,..r....d. ( L ACM 61031 -)2). C.,..~/li: Florcnd l ( US !" ~l

1~72-73); MOftlia (AS SP 2537l}-71). M~I4: Bl>tfIMo,l,J(f"MSH 81810). A_ _ : 7_7.2 km :\>\.' l.c1icil ( LACM61019 .61033)

ECUA DOR.-.\·" po: San ta cecut... t KU 104762_63 .1069~S. 109189-90. 109292. 109295, 109297-98. II ISIS­III. 112310, 122270. 122272. 122275-76. 122280 -8 6);Puenn Ore. Rio Aguan,,, (KU 122291); Lirnonco<:h. (LACM72629-41. 72676-82)

PERO.-Pasco: Chanl in. , I<COlaz in Valley (LACM72047-4 8. 72050-S6); P,," de AlOCar, [.cola, ;" Valley (LACM72049). H"linucu: Rio P~hilca near Bo"!u c Nacional de[pari... (TNHC 36477 , 364110..'6483, 36490. 364911 ); MonICAlegre, Rio PachilC<l(AMNH 430 12) Sa" Morl''': S"".",:henear Juanjui. Rio Huall"g. (WCS 2780); upper Biabo Vallcy(AM:'\H 42658). Lortl/. ,.junclion Rim San,iaso and MlIraioOn(AMNH 42450); Varina.cocha nur Puca.lIpa. . Rio Uu~·a.1i

(FM!"H 56236; CAS 85315); Ct'rTO Alili. 24 km E Con·llm.n" (F"I!"H 56239- 40); Vu,ima.lua.o. Rio Hua.lla.,.a.(BM 1947_2.14 .21-22); lIppt'r Ta.chsa.. HIIl.Chiyl.C lI. RioMoronI. (AM:-;H 42338); Pmlpa. Hmnosa. Rio Cu~a.y(AM:-;H 42104. 42354-55); OIbnla.l>C'ar Quebrad.o Ca..hibo)l(AMS H 42981); l.a.jo de Mirmo. moudl Rio /'iapo tAMSH43183.43415); cceneeoce RiooCnnla~a;and Tapil;ho: tAMSH42565. 43249. 43251); 1'IlIi'"" (A",I:-;H 42295. 4J614);connucnct' Rios Tama) 1 .nd Uca.~.li (AM:-;H 43426);Peru-Brasil fron.ic, (AMNH 42058. 42694). 1'.." ,,: L.UniOn. C.mbllya. Rio Huacam. )o (8 M 1907.5.7.8).

BRASIL.- I' ,,,,i: Saatarem, road 10 C""hoeira ok l>-alll.1o(MPEG 744); U' inga ncar Belem (LACM 42321); J"". re·Aeang• . Rio Tapajl>" IB, Lutz and Klos' (952); Rio Mapuer.(AMNH 53334). Am apa: Serra do Navio (LACM 42322).Amazonas: Estir"o do E<.j uador. Rio Javari (MPEG 634- 646);Rio Tachi,a ~ar Leticia (LACM 61011); Tel"l:, inha ncarLelieia (LACM 60974-75 . 61029-30); Eirunepr:. Rio Jurua(B. l Ull and Klo.. 1952); Bcnjamin Conslanl (CAS·SU11873).

GUYA.\lA.-R""''' .."i: '" Acarah) M.....nlain•• W Jl'e'AR;." , (KU 69692·95); KIIYII...-ini landing (AMNH 4m 2);be ·ct'n upper Ku)u wini and Kas, ikaillU Riw... . uppt rE ,qllibo drain.lIe CBM 1936.4 ,2 .7- 8); Onoro Rivt'r(_ Unoro o Rivt'f l (AMS H 44644. 44641. H689-91).,...... IN_ · Akyma(AMSH 13S05-{16); Wi<mar (AMl\l{45n4). .Ua:JUWtfi_I'OUIn>: Kanab<> (AMS H 10407, 13500.23124.39679-80.39687• .3969 1. 391Oll. 70991. 70993-96;USS"-l 118015); 0\:;0 Moum. in, (flol!"H 26710, 26760);Monbllili Cm:k , &..equibo drainagoe (BM 1930.10.10.2-9);junc.ion MCllIIlII. nd Mauroni Rivers (AM:-;H lll'W5).

SURIS AME.-Sorr inam..: Param. ribo (RM:-< H unu O.Maro .. ij,,~: PocIOt'gocd<1t'(RMNH uneal); P. loemell tRM:-<H150(1); N.......u GebcrglC (RMNH 7030. 7725. 8212. 8356.8647, 8929. 9144, 9626), Saromaua: Lan~a Snela. Cop·pename Rtvier (RMNH uncal). Br oJ:.opondo : Brown.bc r~

(MSH 149) ; Ligon.... Gran Rio. S Tapoe Wlilr. ·vallcn (RMNHunclll). /'I'kbril : Fredcrik WillcmIV.valien (BM 1936.9.3.5);Sip.oli ini (MSH 757. 769. 772, 798, 812. 829. 848. 859);Kay r Gebo:r~c (MSH .H4_7S. 598. 614. 629. 643. 656,672); Po.! Tigrie tMSH 367. 389); Po.! Gooini (MSH 321).

GUYA....E fR ASt AISE._ ncar Maripawula tMCZ44552­54); ncar La Gm't' (MC'Z 44555); Ylll iollt, uppct' R~

Camopi (MHS P 50-7); Siul Tonl,lC. Fkuve Approuagl,lC(LACM 42323- 24); Mon. AIKhi-Bacc. ( La 631); Eau...Clemenls ( La 1238); Trois S.UB. uppor flc:lM' Oy.pockuo 1393-99); Saill ( La IOS2_57); R;"ie-~ Ouaqlli (La1446); Mon l o.Jbao (La 1582); K. ....(illilolillooull IMIIO«Gcni:ve 1225.2).

Cleared and Jlai~d malt'rial. -LACM 60914-75 .

Phy floba lu rricvlur (Boulen~ 1899)Ptliinta'\Jllal Poison-arro'" Frog

Fronli. picce II and Figu,," SA . 8F

1899b I'ro' llv .api. Iri=1or Boulcngt'r: 4S5-456. four .y n·'~ptl: B:4 1941.2. 14, 16- 19. Porvenir, Boliv... .Ecuadnr . ]769 m.

1920 1'".,II,Jb<,ln ITicolo •. Barbourlind Soble; 401.

D/'ji1lilioll. -Teech. present: h H:S basally ....ebbed:dorsum black or brown with yellow or whi le media ndo rsal stripe. sometimes expanded to cover most ofdo rsum; com plete yellow o r white late ral stripe; lightproximovemral calf-spot absent: venter yellow or ",hilewith or without black or brown marbling, or venterblac k or brown with yellow or white ,pots .

Dia gnosis.-Ph)-f1obales Iric-olD' is 1I member oflhe /"n ll1ra!is group. differing from all lXher membersof che group in having green bones . P. lrico!o. fur­tber d iffers from P. anlhr",)'i in larger ,ize (Table I) .from P. boorlengeri in having a lighl median donalsnipe , from P. e.pinosai in having a Iighl mediandorsal stripe. in having a complete light lateral str ipe(Fig. I B; incomplete in P. t')pilW.wi. Fig. IA), am!in larger sil e, and from P. f~11I(>T{/lis and P. zaparoin havin g a light media n dorsal stripe.

DnuiptiOlI of syntypes. - The snout-ver n length illmillimete rs of the fou r symy pes, which are all adults ,is (mea n in parenthe ses): two ma les 20 .0- 20 .5 a O.3);two female, 22.0-22.5 (22 .3); four specimens 20,0 ­22.5 (2 1.3) . The skin is entire ly smooth. Maxillaryand premaxillary teeth are prese nt. Tbe firsc finger islonge r than lhe second. All roes are slight ly fringedand basally ....ebbed; the ....ebbing is best developedbetw een loes II -III·I V.

When recently prese rved. the dorswn llIld lhe sideof tbe body were black. emblazoned .... ith completelemo n yello .... lateral and median dorsa l stri pes. Thebase of the upper arm, the lum bar region. tbe postero­dorsal surface of the thigh . and che ventral surface ofthe calf were vermilion red. The venter and lhe restof the limbs were lemon ye llow with black marb ling.or b lack wh h lemon yellow , pot, (Boulenger 1899b).

In preserv ative , the ground co lor of the dors umand of the ,i de of the body is dark brown. A broadlighc median dors al stripe ex tend. from lhe veil! tothe snout , expallding in from of lhe t'}'t's to cover thedorsal surface of the snout. This stripe: narro ..... pos ­

.em ly in two specimens. oontains a median row ofsma ll scattered dark spots in one: specimen. and bears.a small Jl'O'Ierior break ill two specenens. A completelight lateral .uipe extecds from the groin to the uppereye lid and along lhe lateral border of the uppere)elid,uniling Wilh the tight do rsal snow colo r in frolll o f lheeye; tbe stri pe bears a small break on tbe left upper

1976 FROGS OF THE GESUS PHYllOBATES 3J

eyelid.of one ~pecimen . 1lle d....k bro...n oon.ill l poundcolor brolldly separates. the median stripe from thelateral Mripe in three specunen s: onl) ' a narrow sliverof ground color se parates the str ipes in the fourthspecimen (8 M 1947 .2.14.19 ). In this specimen andone other specimen, the medi an and lateral stripesfuse for a short distance behind the upper eyelid.iso lating the ground eolor of the uppe r eyelid as adark spoe . A light stripe extends from the forelimbinsertion along the upper lip to the tip of the snout....here it meet s its fellow in some specimen s. Thedorsa l surface of the forel imb is mostly light. ThedoNII surface of lhe thigh is dan. . with broad . light .sometime s broken anterior and postence borde!"' . meet·ing each other proximall y but 110I distally. Tbe dorsalsurface of the calf is dar\( with light specs. The venterand the ventral surface of the limbs are light wit hdark marb ling in three spec imens, and dark withmediu m-sized light spots in lhe fourth specimen (BM1947.2.14. 17).D~scrip/ion of USNM matf'ri"J. - Thirty-seven speci­

mens (US:'\ ~1 166897- 9(1) collec ted a few )"l:an agoby l .A . Peters on the Cuenca-Macha la road in theprovince of Alua)'. Ecuador . on tbe Pacif IC slope ofthe Andes (the~ slope as the type local ity. aI anelevation 300 to 4IXl m lower). resemble (after preser­vation ) lhe type series . 1lle snout-vent length in milli­meters of 35 adults is: 12 males 19.0-24.5 (22 .6);23 females 21.5-26.5 (24.6); 35 specimens 19.0- 26.5(23.9). Some of the USl"M males have the third fingerslightly to strongly swollen . All the USNM spec imenshave basally web bed toes. as in the types. The color ofthe muscles is very light. as in spec ies of the genusCoI(JS!f'/hus ,

The US:'\~1 specimens have light median doNtl andcomplete lateral wipes; in 3. fe...' specimens, the medianstripe is upandrd. covering lhe anle rior half of thedorsum and fusing with the anterior half of the Ialeralstripe . This fllSion ~mbles that of one of the types.(8 M 1947.2. 14. 19). In most of the USNM specimens.the venter is d....k brown with light spots or marbling .as in one of the types (8M 1947.2. 14.17). In somespecimens. the venter i ~ immaculate light. In onespecimen , the vente r is light with a trace of darkmarbling, resembling the venter of three of the types.Ahhou~h the USN~t spec imens resemble the types

in their color pattern. they differ in their color in life(Frontispiece II). The US NM spec imens were danbro...n with white stripes and spols. whereas the typeswere black with yeuc w stripes and spxs and red flashmarks . 1bc USNM specimens have green bones; thet}·PC specimens have 110I. bul cou ld have lost lhe pig­ment because of their a~ . It is also possible that thegreen color is an artifact. I allocate the USNM spec­imens to P. tricolor and include their meas urements inTable I under that species .

Stomach comems (one spccimen).-From USSM166897: HYMES OPTE RA: FotmicidilC : ~I}' rmcci nac :

SoI~"''Psis ? (2) .

NOIll~ncla/u",.-1 designate B~l 1947 ,2.14 ,18. a20 ,5 mm male from P orve nir, Bolivar. Ecuador. 1769m. co llec ted by P,O . S imons. as the lectoty pe ofPrfljth..,upis tricolor Boulcn ger 1899.

Di."ribwi,,,, <1m' Iw/>i/<II (Fig. 13).-Elevat iun rec­ord.s are from 1250 10 17tH m. P. /ricolor inhabits lhePacific slope of the Andes in southwestern Ecuador.l .A . Peters collected specimen s eevt to a stream in adr)' area of high grass and pepper tree s I=SchinlUor Capsic_?J.

Locu/itin.-ECUADOR SoH."" Po", enir tBM19017.2.14.16- 19). Auoo.l': Rio Minn. 20 km W Santa Isabel.km tOl on Cucnca·M.:h<lla ""'" ncar AbdOnCallieron. SWGiron (USSM 166897-9 11. tbf>9(0): 2 km W Santi I. abo:!.km 88 un Cuenca-Machata road (USKM 1669(1); 4 km SCalavina. km 7t on Cuenca.Machala ro. d (USNM 166899).

CII'<I",d ulld SI<li"f'd m<llf'rill l . _USNM t66897.

Ph)'Jwbat~1 ZlJptUO new speciesSanguine Poison·arrow Frog

Fromispiett I and Figum. Ie. 8D

Holot)'Pf'.-KU 120669. collected 2 km west ofCanelos. Provincia de Paiiu ,u . Ecuador . 580 m, on9 Jul)' 1968, by R.W. Henderson .

Pural}p n ( 17 ad ults . all from Ecu ador )._KU120670. co llec ted at Sarayacu . Pastaza, 400 m, byJ .D. Lynch; UMMZ 129358. collected at the Alpa ­yacu. a small stream near Mera , Pastaza. elevationstated to be 300 m. but note with specimen says ell.'·vation questionable [probably about 1000 m], by W.Clarke- Maclruy re; GO V 8929-30. collected al DonTomb , 5 j m S Montalvo. Pastaza. by G. H..ne ra andR. Olalla; GO V 8932 . 8934, collected at ~Iore t a .

below Montalvo. Rio Boboe aza. Pastaza . by. G. He­rrera and R. Olal la; GO V 11935. collecled at the RioVillano . Pasraza . by unknown collector; GOV 9005 .9007. collec ted at the confluence of the Rios Shyonaand Conambo. Pastaza . by R. Olalla; GO V 9190. col­lected at the Rio Ogl£lO . Pastaza. by R. Ola lla; GOV9212. collected at the Rio Conambo, Pesresa, by R.O lalla; GOV 9380. collected at the Rio Bobonaza ESarayac u, Pastaza. by R. Olalla; GO V 9394- 95, col­lected at Miazal. Morona·Santiago. by M. Olalla ; GOV9304, collected at l.on: to. Napo . by unkno...n collec tor;GOV 9 194, ccnectec at the Rio Suno. l"apo , by un­known colleo:tor; UMMZ 90370. collected al the RioPucuec , S apo. 300 m. by' W. Clarle-~lac lnt )re .

Df'ji" ilio" .-Teeth prese nt; roe s ba~all)' webbed;dorsum red: lateral stripe usual ly ve.tigial or absenr:white o r pale blue ventrolateral stripe; light proxilT}{'·vent ral ca lf-spot absent ; venter black with blue orblue-gray marbling.

J4 BU~1N OF THE.NATURAL HISTORY MUSEUM OF LOS A...'GEL£ S COUlWY No. 27

Diagnosis. - Ph)"lfoooln zaparo is • member ofthe / ,,,,o,o/is group. differing from P. o",ho"~'i inlackin g a light median dorsal stripe and in lv !!cr size(Table I) . from P. boflle"ff ,ri ill having a red donumin life (brown in P. boll/e/lgtri) and in larger size,from P. n piflOs(); in larger size. and from P. tricolorin lac king a lighl median don..J.l smpe and in having" 'hilt bones (gree n in P. (, i.olor) . P. :Pparo clo~ly

l'C'<oem bles P. femoro/is in size and in Iuving a dlOltlanaI fo ld. bUI differs from P. jtmoraiis in having Ired ocrsum in l ift (black. or brown in P. fe moroUs),in usually ha~ing the lighl lateral slripe (Le., the stripeadjace nt 10 (he edge of tbe back) ves tigial or absent(complete from groin to upper eyelid and prominentin P. f tmorulis) , and in havi ng the: ligh l proximodorsal

lhillh -spo l small o r absent (large and prominent inP.femoru/is) . P. Zilpuro resemble s a sympatric species,P. pun' ulus, in having a red dorsum in life, but dif fersin having basa l toe webbing and in larger size .

Deseriplioll of ho IOl)'P<' . - The holotype is an adu ltmale , 27.5 mm snout-vent length. The skin is stronglygranular on ee dorsum and the dorsal surface of thelimbs, and smooth on the venter and the ventral sur­face of lhe limbs . Max illary and premaxillary teeth;m present; vomerine teeth are absem. 1be lip of thesoo ut is subfruncate in dors.al bpe>C 1 and rounded inlaleral aspect. The canthus rostrali s Is rounded. TheIoreal reg ion is slightly con cave. The width of lheupper eyelid is slightly jess than the interorb ital widthTbe d iameter o f the e)'C is !ilighdy grea ter than thed istance from the eye to me l'lOiilril and twice the di­amete r o f the tympanum. The posterodo l"§4l port ionof me tympanum is concealed . 'Ibe finger d isks ;m

small relative to tbose of lhnJrobules . The fio.t fingeris longer than the seco ed. Tbe fingers are webless. 1belarsa l fold is Slm ngl)' bent at its pro ximal end andrelati....ely short. extending along the dista l 33 percento f lhe distance bet..-een the inner metatarsa l tubercle

and the heel fold . The tarsal tubercle is modera tely'developed . Basal webbing OI.'CUl)C between loes !I· IlI· IV.

In life (notes by J.D. Lynch at KU ). the iris wasblack with a bron ze ring around the pupi l. The dorsumwas br ick red , The side of lhe head and body wasblack, The upper lip stripe was gree nish yellow. Theventrolateral stripe was white . The dorsal surface ofthe forelimbs was light brown with a few black spots .The axilla was yellow-o range. The dors al surface ofthe hind limbs was black with blue -gray mottling.The throat and chest were black . The belly and theventral surface of the limbs were black . monied andmarbled with pale blue·gr ay.

In pre~nl ati ....e. tlJf, grancjes of lite head and backare EOkien-brown. separa ted by narrow black inter_spaces . The side of the head and bod)' is black. A.. hire ....entrolate ral snipe extends from lhe groin tothe nilla. uniting donal to the Ioretimb insertion withue upper lip str ipe. " 'hich continues 10 the oostril.

The re is no light late ral str ipe. The dors al surfaceof the limbs is gray. indi sti nctl y banded black . Along-itud inal white srnpe extends along the vemratsurface of tbe forelimb from me axilla 10 the \\Tis!.The postCfO\o'l:ntral surface of the thigh is gray w ithwhite dots. The rest of lhe ventral surface of the limbsand the venter are black . except for \\'hile marbl ingon the Ihigh and posterior belly.

Du criptioN of p<JratY/H's .-Measurements are inTable I. T'hr snout is rounded in dooal a'flCCt in someof the parat}pe s .

In life (notes by J.D. Lynch al KU). in KU 120670.the iris .. 'as black with a copper ring around the pupil.The dorsum was red . There was a vest igia l copperbrown lateral stripe. The dorsal surface of the limbswas Hght brown with dark brown or black spots: thelimbs lacked bright spots. The axilla was orange . The

posterior surface of the thigh and calf was blue withblack reticulation. The throat was black . clouded withblue. T he belly was black wit h pa le blue posteriormottling. extendin g as a stripe along the lateral edgeof the t:.::lIy til the axilla . In prescNative. the blue ofKU 120670 has become gray.

in preserv ative . lhe UMMZ paraty pes differ fromthe KU specime ns in It.aving a white spot on the proxi­modorsal surface of the thigh and another coveringme axilla and the proximopo sterior and proximodon.alsurfaces of lhe upper arm. in having me ventrolalcralslripe incomplete (endi ng midway between the groinand lhe uilJa) in UM MZ 129358 and no! visible inUMMZ 90370. and in lacking ventra! marb ling.

In prese rvalive, rl'lO'>l of the GOV para t) pes resemblelhe ho lot )pc: . Two speci~ns (GOV 9194 and 93(4)

have a promine nl .. hile spot cove ring the axilla and

dorsal base of the upper arm. The dors.a1 surface of thethigh is ei ther emire ly dark or banded with light and .dark ~ there is no lighl lhigh-spol . except in GOV 9007(s mall and vesligial) and 9194 and 9304 (small andprominent ). The venlrolateral smpe is complete res­tending from the groin to the axilla. where it uniteswith the upper lip stripe) or incom plete , Some speci ­mens have a prominent or vestigial stripe on the canthu srostralis and upper eyelid . Five specimens (GOV 8929,91XI5. 9 190, 9212, 9380) have a vestigial late ral stripeadjacent to the edge of the back. In one specimen (GOV9007) . the lateral stripe is com plete and prominent fromthe groin to the tip of the snout; this condi tion differsfrom all otber parat}pe s and from the bolotypc. Thethroat is enti rely dark: the belly and lhe ventral surfaceof lhe limbs bear light marb ling .

Distribution mul humtat (Fig. 15).-Elevation rec­ords are from about 230 to 1000 m. P. zaparo inhabitsthe dense forests o f the Pastaza and Xapo drainages ofEcuador , ea st o f the Andes; it is terres trial and diurnal(lIOfes by J.D. L)'nch at KU) .

£ f)-molog)". - This species is named after the Ziparoindians of ea' tem Ecuador.

/976 FROGS OF THE:GV,US PHYUOBAT'ES

pious group

This group includes seven species (Pltylloba/t'S boJiv­iamu, P.i n/,,,ri , P . parv MfMS, P . pr /..rsi. P, pic/..s. p .p..lchri""C/.. l. P, .•maraxdin",I/.

Defi'li/lim : snout-vent len glh of adult s sma ll lO

moderat e ( 16.5-3 1.5 0101) ; toes webtcss: pairedstripe s (when present) lateral (Figs . 3A -B . D-E.58-C. 6E-F); Iighl proximoventrat calf·spot present(Figs. 9C-Fl or abo;ent.

Pltyl/oba /es boIi>;a"u.1(Boulenger 19(2)Bolivian Pceon-errow Frog

Figures 5C . 9A

1902 P'o>lh.-rapi. boIi,·;o",.. Bouknger: 397. lhftt .yn.types: BM 1947.2.13.89-90. San Callos. [u Paz,IBolivia, 1200 m; BM 1947,2.13.91. San Er~ 'lo ,

(La Pal ,] Bolivia.800 m.1920 Ph.vl/ohalr. holivian,.., Barbourand Noble: 402.

D"fi"i/ion. - Teeth abse m; roes webless: dorsumdar k (in pre~n-alive); com plele lig ht la leral stripe;ligh l prcxisnovennal calf-spo t abse er; ven ter light....ith dark marbling and spots .

Diognosis.-Phylloba/"J boli. 'ionus is a memher oftbe piC/..s group. differing from P. ing..ri in lackingteeth. in having a comp lele Jigi'll lateral stripe (absen lin P . inJ("';l, and in lacking a Iighl prox imovemralcalf- spot. from P , parvul".• in lad ing teeth and inhaving a complele llght lateral slripe ( inc<.>mplcle inP. pa,..,,,I,,.•). from P. P<'lrni in lacking teeth . fromP. p ier.... in hocking a light proximo ventral catf-spct.from P. p"khriprc/us in lacki ng a lighr proxifllOdor,;althigh-spot . and from P. MtfuruJ(dinus in lacking teelhand in having venlral marbling

D..scription.-MeasuremenlS are in Table I. Theskin is granuhll' on the dorsum. tubercular o n thepo~erior back and the donal !>Urf~ of the hind limbs.and smooIh on the forelimbs. the verner, and lhe ventralsurface of the hind limbs. Ted h an: absen L The fin;lfinger is longer man tbe second. The toes are webless .

The color in life is unknown , In preservative , lhe dor­sum and the side of the body are brown. A completewhite latera l 'lripe extend, from tbc groin to the uppereye lid . along tbe outer borner o f the upper eyelid. andsol1lClimes along lhe canthus rostralis to lhe lip of lhesnout. where il meelS ilS fe llolli. A whne stripe extendsfrom the f.....-elimb insel1ion along lhe uppcT lip. endingventral 10 the nostril or connnuing around the snout 10fIlCCI il> fellow. 1llt' ax illa and lhe proximovenlral andproximoanlerior 'Ulface' of the upper ann an: .. hite .Light malbJing covers lhe anterior . dorsal. and posteriorsurfaces of the thigh. On the posterior surface of rbcthigh. the marbling forms a lighl longitudinal undulat­ing stripe in 89 -90. Thigh-sp ots and calf-spots are ab-

sent . The venter and the ventral surface of the limbs an:Iighl .. ilh brown marbling and spots; the ventral marl­ings are dsscrere in 89-90. btll cont inuous and closel) 'set in 91.

Nommcla/urr , - I designate BM 1947.2 ,13.89 . a26.0 mm female from San Carlos. Bolivia, 1200 melevation . collected by P.O . Simons. as the lectotypeof ProMherupi.• holill i<l""J Boulenger 1902.

D;Jlriburion <l"d hobil<ll (Fig. 15l.-Elevalion rec­ORb are from 800 10 1200 m. P. boIivion..s inhabitsthe Amal on d r'l1inage of Bolivia

~<llilks._BOUVIA U Pc: San Cane-. Dc'" ~bpori.

about U"32·S.6TSS'W (BM 19047.2_1389_90) ; San~o.about IS"OO'S. 6S"(l(l'W ( B ~l 19-17.2.13.911.

Phyf{l,b(l/t'S ingf"; (Cochran and Goin 1970)Brother Nicefom's P oison-arrow Frog

Figures 3C, 9C

1970. lH,.JroI>a~J in~ri Coehnn and Goin: 16_17. plale2D- F. hoIoo:ype , CSS M 146ll46. A-no Dc'" Rio~ado. C~...u. Colombia.

D..fini/ ion.-Tee lh present; toes ..-ebless; dorsumdarl ( in presen-ativt'); stripes absent; li/;ht proximo'o'en­tral ca lf·spot present ; venter dark with Jigi'll marbl ing.

DioK"os ;s,-Phyf!o/>{I/<'J ingeri is a member of tbcp;ctUJ group. differ ing fmm all other spec ies of lhegm up in lacking a light l"leml , tripe . and from all spe­cies o f the group except P. P;,·/..J (and rarely. P. per­vul.... ) in having a lighl prcx imovee tral ca lf·' p'lI. P.ing..r; furtber differs from P. boIiv;<lnus. P. pulch ripec­tUJ, and mosl specimens of P . piC'fUS in having teeth,and from P . Jm<lragd ;nus in having ventral marbl ing.

IJ.-scriptiOll. - Mea.suremenls are in Table I (rhe bel­Ol:) pe is a 27 .0 mm female). The skin is strongly gran u­lar on lhe don.um and the do~1 surface of the hindlimbs. and smoo th on rbe fore limbs. the venter , and theventral MJrface of the hind limbs. Max illll/)' and pre.maxillary teeth are present. The finot finger i& longerthan the second , The IOCS are weblcss.

The cllior in life is unknown (C .J. Goin, ill fill. ;Rdo. Hno. Niceforo Maria . ;n fill.). In preservative.stripes are absent. The I) pes are entirety hlack. exce ptfor a white spoe on the ax illa and lhe proximodor;.alsurface of the upper ann. a small round Of rtttangular.. hite spot 00 lhe proximodorsal sutfece of lhe Ihi ~h .

a large .. hile spot on lhe pnJi\irnoventrai surface ofthe calf (wmetimes ei\tending half the lenglh of lhecalO . and iII -defi.w ~ra)' marbling on lhe belly (nolo­I) pel or on the emire ventcr and the venlral surfaceof the limt>s(paralypes).

Rr //I/;oll.hip:> , _ P , ;" Ked is c lose ly rela ted to P.ptctus.

36 BULLETIN OF THE NATURAL HI STORY MUSEUM OF LOS ANGELES COUNTY No. 27

Di~trihuri"'l and hahira' (Fig. 16),-The only elc­vation record is 4()') m. P. ingeri is know n only fromthe type locality . in the Amazon drainll~c of Co­lombia.

uX'/Ililies.-eoLOMBlA , CllI/um i: Aserrfo. 20 km fromjUlIClion Rio. »c!IC.do and Oneguaza. 5 km from righl bankRio Pescado (US!' M 146846_49: lLS 43)

Ph.l'llohmes parvuius (Boulcnger 1882)Ruby Poison-arrow Frog

Frontispiece I and Figures SB. 70

1882 D~nd",hate., /",,,'ula,, goulenger: 145-146. plale11. fig. 6. lhree 'ynlypes: BM 1947,2.30.89-90.Snrap,u, (Pastaza.j Ecuador; BM 80 .12 .5.195.Canclos, tr-.'lal"] Ecuador.

Dcftnilio/l. -Tccth present; toes wcbless: dorsumred; incomplete blue lateral stripe: light proximovenrralcalf-spot usually absent; venter black with blue mar­bling or blue with black marbling.

Diagllosis .-Phyllob(l/<,s purvu/us is a member ofthe pictus group. differ ing from all other species ofthe group in having an incomple te light lateral stripe(often disap pearing in preservative: the other speci eshave a complete light lateral stripe. except P. illgeri ,which lacks stripes), P . ptlfVU/US further differs fromP. pctersi, P. pictus, P. pulchripecua. and P. smarug ­dinu.,' in having a red dorsum in life, fromP . !>o/iv/a"us,P. pulchripectus. and most spo;:cimens nf P. pi<'tU.I'

in having teeth. from P. ingrri and P. picsus in usuallylacking a light prcximoventral calf-spot. and fromP• .v"urugdinu.< in having ventral marbling in life.Two specimens (LACM 72646 and 72655) of a seriesof 26 P. parvulu.,' from Limoncocha have a light area.yellow in life (notes by W,R. Heyer at LACM). onthe proximoventral surface of the calf. Un like thedistinct. white. rectangular or elli ptical calf-spot ufpreserved specimens of P. piCIIIS, the calf-spot in thesetwo specimens is indistinct. gray. and irregularlyshaped. Besides usually lack ing a calf-spot. specimensof P. f}(lrvulus from Lsmoncoc ha differ from sympatricspecimens of P. pic/us as follows (the cond ition inP. piclUJ is in parentheses): the lateral stripe is in­complete. often disappearing in preservative (thelateral stripe is complete and dist inct in preservative);the dorsum is red in life (black or brown): the dorsumis strongly granular (slightly granular); the first fingeris longer than the second (about equal); maxillary andpremaxillary teeth are present (absent) . In life , P.punu/u.,' resembles a sympateic species, P. zaparo,and an allopatric species. P. espinosai. which alsoare red. P. ptlrv ulus d iffers from P. taparo in lackingtoe webbing and in smaller size (Table I). and from

P. espinosai in lacking toe webbing (the toes arebasally webbed in P . tf,Ipuro and P . ".,pino.wi) .

Destr/plion . -Measurements arc in Table L Theskin is strongly ~ranular on the dorsum and the dorsalsurface of the hind limbs. and smooth on the forelimbs ,the venter. and the ventral surface of the hind limbs.Maxillary and premaxillary teeth are present. The firstfinger is longer than the second. The toes arc webless.

In life, in ,pecimens from Santa Cecilia and Limon­eucha, Ecuador (notes by J.D. Lynch at KU and W.RHeyer at LACM) , the d",.,mm was dark red (only thenumerous. c1o>elyset granules were red: between themthe skin was black). The ground color of the side ofthe body was black. An incomplete Iighl blue lateralstripe extended from the groin one -half to three-fou rthsof the groin-axilla distance . A li~ht blue stripe extendedfrom the forelimb insertion along the upper lip. endingventral to the eye or nostril. A yellow axillary spotextended onto the proximodorsal surface of the upperarm. A small round or oolong yellow SPOl adorned theproximodorsal surface of the thigh , The venter andthe ventral surface of the limos were marbled lightblue and black: the marbling extended onto the sideof the body . The blue and the black were 01 equalextent in some specimens; in others, the blue formedlhe ground color , Juvenile specimens from Santa Ceciliawere bluish black on the dorsum, and had less blue onthe Venter than did adults; the smallest juveniles lackedyellow axillary and thigh-spots

In life. a topotype from Sarayacu. Ecuador (Frontis­piece J. and notes by J.D. Lynch at KUj, resembledspecimens from Santa Cecilia, except that the proxi ­modorsal surface of the upper ann was light blue in­stead of yellow , and the proximodorsal thigh-spotwas absem. The types of P. parvulus from Sarayacuand Cane!os also lack thigh-spots _

In preservat ive. the ground color is black or darkbrown. The axillary and thigh-spots usually are dis­tinct and white. The lateral stripe usually disappears(e.g .. it is visible in only six of 26 preserved LACMspecimens from Limoncocha, Ecuador). The ventralmarbling is indistinct and gray, and sometimes dis ­appears (i.e .. the venter becomes entirely black ordark brown).

Nome ndaluT<, . - 1 designate BM 1947.2.30.89, a22.5 mm male from Sarayacu, Ecuador , collected byMr. Buckley, as the lectotype of Dendrobates purvulusRoulenger 1882.

Dinrlbutian and habila! (Fig. 16).- Elcvation rec­ords arc from 330 to 1000 rn. P. parvutus inhabits theforests of the Amazon drainage of Colombia. Ecuador,and Peru, east of the Andes . Specimens from SantaCecilia were taken on the ground in fo rest (KU 111507­14) and at night on the ground in a marsh (JDL 1226 atKU); one specimen from Sarayacu was taken by daynext to a small creek in fo rest (KU 120664; notes by1.0. Lynch at KU).

/976 FROGS OF THE GElWS PHYLLO BATES 37

Loculi/it's . -COLOMBIA. P..lI,_.nI: Sm l. R......dl: 1mKofme•. Rio Guame. tO I SOb25-~8).

ECUAOOR._Sapo: Men ,UMMZ 9(371); Lo~<> (GO Y8921); S .Iope Sumac<> (GOv 9382); ~ l m W I'I>crto Nap<>.Rio Na p<> (JAP ~658); Rio Cocapioo.:-.cap<> drainage \CAS·SU10337); Santa Cecilia (KU 104773, 106953-54 , 109157_88.111507- 14, 11l684, 122225-5 1; l.ACM 60980. 61012-14);Limoncocha (LACM 7264/1_71; KU 99 t t8): Laso Asrio(WED 35728); Puerto Lim . Rio A~uarico IKU 122252-(8);Puerto O~ , Rio Agu;Uico (KU 122269). Pas/a.,.: Sarayacu(BM 1947.2 ,30.89 - 90; KU 120664); Cando. (BM80.12 .5 ,195) ; headwalers Rio Capahuar i (GOV 8925);head"'a,,," Rio Bobona:la tGOV &92 &; lA P 2124. 2ISS);~l<nla. l>ckM~~. Rio Bobonaza (GOV 8933); ron­Ouc:nce Rio> Sh}'OftI and Conambo (GOV 900b); Rio Co­nambo (ZSBS 221t962; GOV 9211; JAP 31166); Rio RUluno.Bobonua draina.., (GOV 9191-93; JAP 3909); 0.& lm I\lOPuyo (lAP 2(64). .U"TDn,,·S'''fli,,~..: 3.2 km E Su.:ua .'rai l from Suo:;ua 10 Rio UI"'IlO (lA P 2208. 2229); Millal(GO Y 9387); Rio Uu_hin N Arapirolo (GOV 8916)

PERO.- l.o,.,./o: Peha_ (BM 1915.3.9.12; CAS·SU 6332).CluJrt'd and .lllI in ",J "'(IIl' ri"I , - LACM f!0980; KU

126309

PhylloboUJ pt'tt'ni new spoc ie~

Peruvian Poison-arrow FrogPronnspjece II and Fi$ures 6F. 9B

1970. Dr..JTDbolt'l Ir;,-;/IO/". (I"'n). Cochran and Goin:16 (1'IOl lhe p1ale; onl) MCZ 24443 is P. pr/rrsi) .

HolOf)'pl' . -UST\M 166763. co llected at Santa Isabel(a village on the Rio Xevati. a tributary o r the Rio Pi­Chis. Pachitca d rainage. 3.5km SE Puerto Bermudez. 80km ENE Oxapampa: the specimen Wall taken s lighllyupriwr Irom tho:village. accordin g to A . C. Th ere -e n.in fill.), Departamento de Pa'lCO. Peru. 4.58 m. on 16lu ly l%ll. til G. R . Xconan .

ParatyPt'J (4 1 aduh~ and nine tadpoles , all fromPc:ru).-KU 144348-50, 10popatal)pes. collected byD,B. Ekkens: ~CM 924 14 and KU 144344-46. co l­leered at xevau. Pa~co . by D . B. Ekkens; AMNH43008. 43011 . 43016 . 43026 . 43034. collected atMonte Alegre. Rio Pachitea , HuimU('(). by H. Ba"lcr;AMNH 42 179. 42505~07 . 42.546, collected at Achin a­mis a. Rio Hu allagu. 60 km SE Yurimag uas , SanMart in. by H. Bassler; AMNH 42790 . 4294 5. co l­lected a t Chazuta. Rio Hua llaga. 75 km S Yunm aguas.San Mart in . 274 m. by H. Bassle r: AMNH 43.5.53 .43564 . collected at lhe Rio Pisqui. Ucayali d rainageabove Contamana. Loreto. by H. Bassler; FM l\"H56248. COIJ«-led al Cerro Azu l, 24 km E Contamana.Loreto . 500 m, b)' J.M . Schunke; MCZ 24443 (adultand one tadpole) . collecled90-138 lorn WSW Pucallpa.Loreto. b)' W.F. Walker. Jr .; BM 1900.11 .27 .38 andAMKH 17199. 17253-57. collected at Perene. Junin.ROO m , by P.O . Simons (BM) and C. Schunkc (AMNH);FMN H 36778 (adul1 and cightladpoles) and 36779- 85(ad ul t ~). coll ec tcd in lhe valle y of the R io ViIOC.Juoin. by F. Wylko wski; 8 M 190 7.5 .7.9 - 11, co l-

J«-ted at La UniOn. Rio Huecamayo. Pueo. by .\tr.Ockcnden .

Dtjini/ion .-Teelh present; toes webje ss: dorsumbrown. sometimes with small black ~POIS; com pleteyellow or yello w-gree n lateral stripe; lighl pro ximo­ventral caJr' SPOI absent: venter bluc with black mar­bling or spots

DiuIlIWl i.I.-Phylfobtl/t l petersi is a member o f rhe

pic/"s group. d iffering IromP . boli>"illnUS and P . pul­chriPt'CI.... in having teeth. from P. inilrri in having acomplete light latera l ~l ripe (absent in P. inxrri) and inlack ing a li$ht prcximosentral calf-spot. rrom P. POl""ulll:l in having a complete lighc late ral -.lripe ( incom·plete in P. pon'ulus) and in having a brown (!,(mum inliCe (red in P. pon·ulus) . from P. piClIl:l in Il;,ving teet h(u~uall )' lack ing inP. pic/us ) and in lacking a light prox·imoventral calf-s pot. and from P. smaru/:dillus in hav­int; ventral marblin g .

Description of '10[01.1'(1<' . - The holu t),f'C is an adultremale . 28 ,0 mm snout-ve nt length . The skin is moder.alely to ~Irong l)' granular on tbe dors um and the lloNiI

surfece of the hind limbs. and smooth on the rorelimbs.the venter. and lhe vencra! surface of lhe hind limbs .Maxillary and prema,illary teeth are prese nt; vomerineteeth are absent. The tip o f the: snout is )I,/blruocalein don-a l aspect and rounded in laleral aspect. Thecant hus rosrralis is rounded . The joreal region is slighll)'concave. The width of the upper cyelld is _Iightl) ' Ies~

than the intero rbital widih . The d iame ter or the eye is1.2 times the distan ce frum the eye 10 thc nost ril andtwice the d iameter o f the tympanum. The posterodorsulportion of the tympanum i.~ concealed. The ringer di sksare small rela tive to lhoS(' or Dt'I/dmbtl/t's, T he firstfiegcr i~ kmge r than lhe second . The fingers are web­less. The canal rold is ~l ighlly curved. e'lendin$ alonglhe di~tal b4 percent o r the disrarce bec...'ee n the innermc:tatan.altuberde and the heel fold . The rarsafrubercleis absent. The toes are weble ss .

In life teeter slide by A.C . Thoresen; venter notvisible ), the do rsum was Iighc brown. The side or thebody was hlack . A co mplete )'ellow late ral str ipeextended from the groi n to lhe upper c)'e lid, a longthe oute r border or the upper eyelid. and along theca nthus rcs tralis (whe re it was go ld) 10 the tip o r tbesnOUI.where it mel its Ielksw. A white stripe extendedfrom lhe forel imb insertio n along the upper lip to tbelip of lhe: snouc. ...'here il met che cantha! stripe. Thedorsal wrr....:e o r the: upper arm was gold . The dorsalsurface or the: forearm and hind limb " 'as lighl bn,)...nwith a fe.. ' small black ~poI s . The anterior surfaceor the thig h bore a greenish wash.

In preserva tive. the dorsum and the sidc of the bodyare blac k. The latera l and upper lip stripes are gray;they mee t eac h other o n the tip or che _nou\, Thc entireanterior surrac e or thc snoul is gray , T he uppe r lip~l ripe e\lends ove r lhe rore limb inser l io n into lhcaxilla. wh ich is gray. The vcneer and the vcnlral surface

38 BULLETI N OF THE NAT URAL HISTOR Y M USEUM OF LOS AN GELES COUNT Y Nr>.27

of the limbs are purpl ish gray with black marbl ingThis ventral pattern extends onto the proximoanteriorportion of the dorsal surface of the thigh. where it(<lOllSan inll isti ncl , pu!.

Dnaipllon of p,mIlYp"s. - Measurements of adultsarc in Table I (four of the paratypcs are juveniles),Many paratypes have the second finge r almost as longa, the first. The KU paralypes have a long tarsal fol<1.extending over the db tal70 to 80 percent of the distancehelwee n Ihe inn",r mCl3lar;al tubercle and the hcc! fold;in these specimens. the tarsa l tubercle is slightly tomoderately developed.

ln life. in specimens from Nevati, Peru (notes byO.B. Ekkens , in /ill .), the dors um was brown withsmall black spots . The side of the body was black . Thelatera l stripe was yellow-gree n; the upper lip str ipewas lighte r and greener. The venter and lhe ventralsurface of the hind l im bs were blue wirh hlackmarbling.

In preservative. lhe sl ripe ~ and the ground co lor ofthe venter arc purplish gray to lig ht brown. The colorpattern of the para typcs differs from that of the ho!ol ypea, follows : BM 1907 .5.7.9 has broa der dark ventralmarbling . BM 190 7.5 .7 .9 _11 have broader la ter alstripes A M1\ H 42507 and 421 79 have stripes sobroad that only a narro w. sometimes broke n. darkbrown line remains bet ween lhe purplish gray of thestripe and that of the lowe r side o f the body . The dar kventral marbling sometimes is reduced to dis cretespots.

Tadl'0//',' ( Figs l IE . 12Fj ,-One tadpol e. in S tage28. was taken between 9 and 14 Aug usl 1939 fromthe back of a 28 ,0 mm adult female (MCZ 24443) .between 90 and 138 kill we'lnorthwesl of Pu<:a llp<l.Peru . Another lot (numbering eight). in Stage 26 .was lake n on 28 October 1940 from the back of a25.0 mm adult male (FMN H 3( 778). at 1600 h. aftera rain , in forest in the valley of the Rio v uoc, Per u;the tadpoles lay with their heads median and theirtails lateral relative 10 the ad ult's hock. Measureme ntsare in Table 3

Retationships, - p , petersi proba bly is closely re­

lated to P. h"lh'iall//< and P. pulch rip"CIUJ.Stomach COIlIelllS (two spec imens) .- From FM NH

3678.~ : H YMENOPTERA: Fon n ic idae: Ph/'idol/' sp.( I ), Ponermac (I ): HEM IPT ERA: Tingidac": frag­ments . Fulgnridae?: fragments.

From AMN H 43016: HY~I I::'.t,;OPTERA : Fonnicidae:Azteca sp. ( 1), Myrmecinae (ll; CO LEOPTERA : Cur­culionidae (I ) ; ISO PT ERA : Na .lulitermn? sp. ( I):DlPTERA: Dro sop hilid ae (9) ; ARACHI\ IDA: Pisau­ridue (35) .

Distribution alld hahitat (Fig . J6) .-Elewtion .....~­

ords are from 274 to gOO m. P. pel('T"i inhabi ts thefor ests of the Amazon drainage of Peru. west to theeastern foot hills of the Anlle s.

EtYllloloKy . -Thi, species is n..mcd after the lateJame s A. Peters , curator o f amphibians and reptiles

at the National Museum of Natural H istory in Wash­inglon. D.C.

C"mred <I1ltl "w incd malerial.-A:>'l ~H 4.,016: FMNH36783

Pttytlobates pictus (Bibron in Tschud! 1838)Spo t-legged Poison-arrow Frog

Figures 3A -B . D-E, 9 D-F

1838 Hy/aple.,ia pi, ·la Bibrnn. T....'hurJi: 21\. 71. two ,yo _types MHNP 4910 , San ta Cruz, [Santa Cruz.]Bolivia.

184 1 UOIdrobatn piclus. Dumeril and Bibrnn: 656.1848. Drndro/mlr.pictus. Guichenot: 119_1 20.1l\5l\ Hy/aplesia picla , Gunther: 125_126.1864 DOIdrohale> '1igarimus (p<lrt) . Steindaehner: 257_

259. f,gs. 2. 2a-d1864 Drndroharr.f hr",..ra lu.f Fitzinger. Sieindaehner: 258­

259 . two syntypes )\;HMW 3818: 1_2, [probablySao Vicente. Meto Grosso,1 Brasi!.

1864 Dendrohotes <'Ut'nemi, Filzin~,'er. Steindachner: 257­258. plate 13, figs. 2, Za-d. four syntypcs: NHMW19190:1-4, [probably Rio Mamore . Rondonia. ]Bra,il

1882. DOId",hmn Iri"ill<l /US (pal'\). B\lUlenj;er: 144- 1451883. Drndro/nur. hll l",r/i Boulenger: 1\36. plate 57. jig.

4. se"en syntypes BM 1947.2 .15.14_20 , Yuri ­maguas, Huallaga River, [LoreIO,) Peru.

1887. Drndmbules bra, ·,'mu. Cope: 53 _54 , hoIOl)'pe:ANSP 13414. Chapada I~Chapada dos Guimaraes,according to BOKermann 19661. Mato Gros.'\{), Brasil

1925 Hylapir.fia jlaw'p irla A. Lutz: 139. 'yntype: USNM96986. Belo Horiw nte. Minas Gera;" Brasi!.

1926. Dm"rofmlrs triviumu.< (1"'1'\ ). Miranda Ribeiro: 2241926. Dendrafm/C$ broauws. Miranda Ribeiro: 179- 180.

,nt-fig. 96, plale 20, figs. 3, Ja -b1952. Drndroba lC$ piclu.' (including subs~cics bra,·,'alu,.

eU('l,emis . jluvopiclU". h"h~e1i. pi,",u.< ). B. Lutz:608-613 . piale I . plale 2, fig,. 1- 6. 8, plate 4

1':152. D<,~drob<IICl pi('/us. B. Lutz and Klo", : 670-671.t955. Dr~drubm.... f/"",'p i' ·III.,. Cochran: 8-':1 , piale I.

figs. G. H1957. D,'ndro/",l<'-< pi,.,u.,·. Dunn: 78t':l6 1. Dendroba/t>,f jla vapir-lu.•. Cochran: 107. pla'e 42

(c" l" r phOIO).1965. Dr~drolmlC$ piN" .< gua"U1",n, i.' Healwole. Solano .

and He. twole: J50- 3Si, hoiot}'pe: MBUCV 31 t2.Alliplaniciede Nuria, & iivar. Veneluela. 100-250 m

1970 Dr'ldrobd/cs h<lhaeii (pa"). Co<:hran .nd Goin: 17-1 9.plale 5D-F (only FMNH 61789 is lIot P. pielu., ).

1970 Deadmholr.• tril·iu"w.' (pan). Cochran and Goin: 16(no' the plale; only USNM 118681_82 are P. piclus ).

1970 . Phyllobau,f jemamli. ( part) . Cochran and Goin: 42(11m lhe plale: only USN M 14681\3 , 141\866. 147268are P. pi,·/u. )

De/initiol1 . - Teeth usually absent. but present inrome specimens; toes wcbless: dorsum blac k. brown.or gray . sometimes with gray or bro ..vn spots or mar­bling. sometimes wi th paramedian rows of yellowSpOIS: complete yellow. gold. yellow-green. white .N cream laleral stripe; yellow. yellow-orange. orange .red-orange. or (very rarely ) blue pro~imoventral calf ­~pot present: ven ter black with blue or white marbling.

!976 FROGS OF THE GENUS PHY LLOBATES 39

or blue or white with black marbling (rare ly. verneralmost immaculate while ).

Diag"o,i.,.-Phyll"bal n piclu.l is a member of tbeph-tu, group. differing from all other species o f thegroup except P . ingeri (and rare ly. P. parm!u,) inhaving a lighl proximoventral ealf ·spot. P. piC/us dif.fer s from P. ingeri in usually lack ing teeth and inhav ing a eomple le light laleral stripe (ab'iCnl in P.ingeri) , P. pinus further di ffers from P. parmlu< ;nusually lackin g teeth , in having a black . brown. orgray dors um in life (red in P. !nln'ulus ), and in havinga complete light lateral stripe (incomplete in P. pan'ull,, ).from P , pelusi in usua lly lacking teeth, and fromP. smllragdinu, in usually lacking teeth and in havingventral mar bl ing . P . piC/us resembles a sympamcspe cies. P . femorali s . but differs in smaller size(Ta ble I). in having lhe upper lip stnpc not meetingthe canlhal stripe . in having a light proxiruovcntralcal f-spot. in baving lhe firsl finger abo ul eq ual to

the second (P. femorali s has lhe fif' l finge r lo ngerthan the second ), in lack ing webbi ng (P. femoralis hasbasa l toe webbing ), and in usuall y lacking teeth

Dl'scril'/ir", . -Measurements are in Tables I and 6The skin is slightly to strongly granular on the dorsumand the dorsal surface of the limbs . and smoolh onlhe venIer and the ventra! surface of the limbs . Maxil­lary and premaxillary teeth usually are absenl , butare present in some specimens ( see below). The firsllinger usually is about equal to tbe secon d. but some ­

limes i> sl ightly longer than tbc second. The roes areweb less

Although all popu laliuns of Phyllobales piClu., havea light proxim oventral calf . spol. this species variesgeographically (but not str ictly clinall y) in snout-vernle ngth (Table 6) and color pallern . For co nve nience .the variat ion in color pallern is described be low assix patterns. which are not intended 10 represent taxaPopulalions whose members bear some of these pal.

terns have been described as species or subspecies:some of these populations. and some other popcra ­lions lhal never have received a laxonom ic name . maybe specifica lly distinc t from P. pic/u s. Because oflhis poss ibility . [ have indieat~..-l 10 which populationspublished oame s apply .

Pallern I ( Figs. 3D. 9D; inclu des Dendrohate.•bracelll" .• Cope 11\87 and Hylu"l esia /l av"" iela A.LUI? 1925): Specimens bearing this palter n are knownfrom S\>Ulheastern Brasil (slalcs of Minas Ger ais. Go·i,i.> . and f..lato Grosso ) and easlern Bolivia f EI Porion.18°10' S. 6O"08·W). T he sno ut-vent length of speer­mens bearing th is paUern averages longer than in speci­mens bearing otbcr pallerns (Tab le 6). Pattern I ischaracterized by a pair o f paramedian rows of ye llowspots (Fig , 3D); the other pauerns sometimes have grayor brown dorsal SPOIS or marbli ng. but never haveye llow spots as in Pancrn I. The thigh-spot usuall y islong (Ir is short in the holotvpe of D. bruemlu.' Co pefrom Cbapooa) . Frogs from Belo H",i?onl e (lhe lypelocality of Hylllp!e, iu jlIl,·"p;"la ) have a light ven Ierheavily marbled with black (Fig. 9D) , The black ventral

marbling is slightly reduced in specimens from lhe mostwesterly known popu lation with Pattern I (EI Porto n).The marbling is greatly reduced in spec imens fromJ1<.,pulalions to the nurth (Cha vanlina. Cavakanl i) andwest (Chap;uJa ) uf Belo Hortz ontc. In specimens fromChapada (the type locality o f Dendml"'lCS' brl'CCIl/!HCope). lhc venIe r is near ly enlirely light. with a fewblack spots on lhe lateral ponion> of the belly; the dor­sum is tbe same as in specimens from Belo Horizonteln life. in specimens from Belo Horizonle . Brasi l (lhetype series of H . /lIW""i<'W: notes from B. Lutz 1952).the iris was black. The dorsum was black with ycllowspots arranged in a pair of paramedian row s, The latera land upper lip stripe s were yellow. The dorsa l surface ofthe limbs was black with yellow spots . T he thigh- spotand the calf-sJ1<.,t were red-orange . The venter had a

T~a l . E (>

Snoul·", nt length (mm) of .dulls of Ph)'II<Jb<i1e< ph' I'" bearing five color pall"'"' . a,,-. ngcd in <>rdcr <>f de"rca, ing me.n' noul-\'C nl Icnglh. P~\1ern 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 ;" ion . Paller n J: BOLIVIA: Ru,rcnaha~ u" . [''<In.I"wet Rio B<"pi.

[, iamas. Sa"la Cru?. Buenavi, I", Yunps de C""h. b. mba: BRASIL: Rio M.,m,e.U,ucum: SUR[:-JA~lE, 1'",1Tigrie. P"Uern 4: COLO~lB IA: Sa" Marlin: ECUADOR:

Sam" Cc.;ilia. Lim"neocha. Puertn Libre. Sa,a~aca : PERU: Yurimag"a, .C.,hit,,, ,.•. 1<1";10'. Pampa Hern~"• • Tapi,hea. Sobr,,1, T""ache. Rk> 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 25.7es 16.5-2 5.0 21,7 " 16,5-n5 19.8 '" 19,5-25.0 22.5n 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 grou nd color with bl~ck

marbling.Pattern 2 (Fig. 'JE; includes Df'mlrobal~s bruc.."UIUS

Fillinger in Sleindadll1er 18(4) ; Specimen, bearing thispanern are known from Sao Vicente (, tate of MaiOGrosso). Brasil. Sleindao;hner', ( 18M) combined Iocal­jl~ l i , t o f Drndrobates rucnemis and D. braccana fail,to a,sign loc alitie s to each fonn. Steindacbners labelwith tile tWO syntypes of DeluJrub<lleJ braccutus says"Mato-grovso'"; thu, the syntypc, tN" HMW 38 18:1-2)probably an: from " San Vicente t Pmv. Marogrossol"(Sao Vicente) . one o f the localities mentioned bySteindacbncr. Sao vjceme is near Cope ' , locali ty ofCbapada (B. LUll 1952), In thi, pattern. in preservenvc .the dorsum bears indistinct lil!hl spots not in rows (sim­ilar 10 the SP01sof many specimens bearing Pattern 4).The verner is lil!'hl. with a few black spots on ils lateralportions (Fig . 9E).

p...uem J tFig s. JA . 9F; includes H.I"lapl~sia pictoBibre>n i" Tscbudi 1838 and Dt'lwrooott' s t'uc"t'misFilZinl!'er'I/ Sreindcchner 1864): Specimens bea ringthis patte rn arc known from Bolivia (north. nonhw esl,and west of EI Porion. in the dep artments of SantaCruz . Benl, and La Paz). Bras il Ithe terr itor y ofRondonia. the Siale of Para . and al Urucum nearCClnlmoo on lhe Brastlian-Bol ivian border ). and Sun­name . In Ihis pattern. tbe dorsum lach spots; thethigh-spot is long (Fig. 3A ) and sometimes broken.and the venter is dark with 1i!! lu marbling (Fig. 9F).In life. in specimens probably frum the Rio Mamore .Bracil (t he type serie s of D. t' u("11~mis; notes by J.:"ialkrer in Steindachner 1864). lhe ins and lhe dor.um....ere black . The laleral and upper lip snipes werewhile . The axilla . the thigh. spool . and the calf-spot werered-orange. bordered with black . The junction betweenthe rectangular proximal and the oblong dista l portion sof lhe thigh,spll t was yellow. The venter was grayishblack with whitish spoe, and marbli ng. In specimensfrom PosI Gonini and Sipa li",'ini . Suriname (noieS byM.S . Hoogmoed al RM:->H). the OOnum ....as black wjth

gray marblin g. The lateral str ipe:....as cream . The upperlip stripe was while . T he ax illa. the postcrovemral sur­face of the upper arm. the thigh -spot, and the calf-s potwere yellow . The venter and the ventral surface of lhelhigh were black ....ith blue . greenish white . or bluishwhile marbl ing.

Pattern 4 (Fig. 3B: include s Dt'lldrobol~s hoh" d iBoolenge r 1883): Specimens hearing Ihis pane rn areknown from most of Amazo nian Peru. Ecuador. andColnm bia. and in Amazo nian Brasil in lhe lerrilo ry ofAcre and the slale of Amazo nas. These relalively smallfrogs (Table 6) lack rows of yello .... dorsal spools (al·lhougb they 'Omelimes have indistincl gray dorsal~s) and ha\Il.' a short Ihigh-spol (Fig . 3B); lhe venIeris dark wilh lighl marbling (in pm.el'\-ative . the mar­bling is conspicuous on the belly and indistinct on thethroa t). In life , in s pccimen.~ from Tere~inha . Brasil.

ncar Leticia (my notes}, tbc iris . the dorsum. and theside of the body were black . A complete yellow lateralstripe extended from lhe groin 10 the upper eyelid. alongthe outer border of 111(0 uppe r eyelid. and along the:canthus rostrali s 10ee lip of the: snou l. where il mel ilsfellow. A while stripe eXlcnded from the forelimb in­sertion along the: upper lip. ending ventral to the eye ornostril. Tbe dorsa l surface of the limbs was lighl bro....n.exccpl rbe proximal half and the posterio r plmion of thedislal half of the lhigh. wbich were black; the blackcolor surrou.ndcd a soon oblong orang e spot on thepro ximodorsal surface of the thigh. Blue spob oma ­meraed tbe posterior surface of the thigh . An orangeuillary spot united wilh an orange ventra l stripe extend­ing alo ng either the proximal half or the entire length ofthe upper arm. The ventral surface of the forearm borewhite marbling. The ventral surface of the hind limbbon: blue marbling and spots , except for the proximo­ventral half of the calf. which bon: a rectangu lar ore lliptical orangoe spot. In one unusual specimen (LAC M6 1006. one of 12 specimens from Teres inha). the calf­spot was blue. The grou nd colo r of the venter and ven­tral surface of the limbs was black , The thlOat bore bluemarbling and spots. The breast and belly bore ....bitemarbling. Spec imens from Pixuna , Brasil , near Maneus(my OOIes). J'C'sembled the Teresinha specimens, exceptlhal the iris WlLS black Wilha gold nng around lhe pupil.the back was black ....ith gray spou. tbe lateral smpewas gold, the dorsal surface of the calf bore IWO darkbrown transverse bands. and the axilla and calf-spotwere yello w-orange . In two juveniles. the:dorsal surfaceof the upper ann ....as orange·gold. and the dooal sur ­face of the forearm . calf. and larsu s was green ish gold .Specimens from Santa Cecil ia and Sarayacu , Ecuador(noles by J.D, Lynch at KU). resembled the Teresinhaspecime ns. exce pt that the iris was bronze. the dorsu mwas black. brown ....ith black spots, or gray wilh blackspots , the lateral stripe 1'."3-' cream or yellow. the Uppellip siripe was blue-green in one specimen. the dor1;:alsurface of the calf and forelimb hore brown or bhd.bands and spots. the axilla was yello...' or yello ...'·orange . the thigh-spot and calf-spot were yello w ororange. and the ventra l marbling and spots were blue .

Pattern 5 (Specime ns hearing Ihi.s pattern may be spe­cifically distinct from P. pictus) : Spec imens bearing thispatter n are known from paris of Amuonian Peru (Tsjc­vemeni; color s lide by A.C . Thoresen: l\"evali: KV144351-60. LACM 9241 3; Rio Pechitea near BosqueNacional de Iparia: TNHC 3(4 69) . "These specimensresemblc frogs bearing Pallcrn 4 in s ize and color pat­lern (( he Ihigh ·spo t is shon ). bul differ in lhe followingcharacleristic s: the dorsum is Slmngly granular . leelhare pre.cn t. lhe head and back hear small spols (some·times panly fu~ logelher and somelime s arranged inIhree or four longilud inal rows) . lhe calf.spot ellendsonlo lhe disla l one- founh or one·lhird of tbe <lnr....1 sur·face of thc thigh, and the entire ventcr ( including lhe

/976 FROGS OF THE GE.,,"'US PHYLLOBA TES

Ihrtl;ll) and most of lhe ventral surface of the limbs arelighl with dark marbling. Of a series of \i~ specime n\ ofP. picfu, (r.' HC 3646 1. 36469-70. 36476. 36482._\6.197) I..ken by J. P. Bogart at d ifferent ,Jate~ in 1967and 1968 on lhe Rio Pachitea neil!" tbe Bo"'lllC~acional

de Iparia, Peru. one (ThHC J6.1691 ha~ Pattern 5 andfive have Pattern 4; I do no( know if all tne..e 'J'«imenscame from thr ..arne deme . In life . in ~pecimen~ fromNevali (notes by D.B . Ekken ~ in li ll.) <UMl T~io.'Cnle ni

(colo r slide by A.C. Tboresenl. Peru . lho: dorMim "-a.~

orang ish brown .. ilh light olive bro" 'n ~pot:s (sometimesarranged in ro''''S). The lateral and upper lip Mri pe~ wereyellow-p-een or white . Tho: dorsa l surface of tho: limb"..'as lighl ol ive brown. The a~ iII.. , lho: thigh-spot, andrbe calf- spot were yellow, yellow-ora nge. or orange.The venter "'as blue with black marbling,

Pattern 6: This pattern is characterized by a smallcalf-s pot, and includes tbree subpauems. Subpanern 6a(includes Dem/m!l<Iln piaus KlUlYWWIlJis Heatwole ,Solano , and Heatwole 1965): Specimens hearing thissubpattcrn are known front rfc sl"le uf Bolivar in Ve·nezuela and from Guyana . The lighl ventr al marblingis reduced to a few spots on lhe belly and tbigh~. In life.in a spec jrnen from Hecenda San Felip:: ncar Miamo,Venezuela (t he paratype of D . p. /(u<ly'lnlfnsis: lINe<;

from Heat..'ole, Solano , and Heatwole 1965), the dor­w m was black. The lateral and upper lip . tripes wereyello w. The tbigh-!opOl I..... red-orange. exce pt for ayellow junc tion between its proximal ano.l di~lal por­tions . The venter "as bluish black " 'ith lj:u y n'lou linlj:.

Subpanem 6b: Specimens bearing this subpallem areknown from Guyana. Tho: thign..spol e ither is long andunbroken or is broken into ... shon proximal ..pot and ...long dis....1 )po(; the: venIer b...s ...ell -developed lighlmarbling .

Subpanem 60; (Fig. 3E): Spec imens bearing this sub­pattern are known from Gu yan a . The tbigh-spot i~ longand unbroken: the la te ral stripe: is rel ~ li vel y broa d(Fig.3E).

Bru diflg coif. - T he call of spec imens bea ringPattern I has been reported as "a piping cry, boowit­hoowlr" {Gans 1960, El Porton ), " 3 rather loud, verypersistent peep .' also a "trill' (B . LUll 19~2. BeloHortzorne). and "beginning with a roll, p-r-r-r-, thena soft whistle repealed hal f a dozen times and ~cry

high in pitch" (Cochran 1955. Bclo Horizonte}.T//dl'0lrs (Figs. I IF. 12G)._Two lOb have been

taken frum lbe backs of adult males in Suriname inJune and in Bolivia in December (Table 4). A thirdlot (MCZ 29825. number ing t..'o. in Stage 25) prob­ably is from the back of an adult, altho ugh there i~ norecord of this , Two ot her 100s (U!-l!-IZ uncataloguedand UMMZ 58949 . in Sta ges 38 and 42 . taken inBoliv ia in August and Dece mber) are nol from tbebacks of adults. Me~remenb are in Table 3.

TOQfhlfd sP/fcimlffl s. - C.F. W...lker sbo 'ed me ...specimen of P. pictus from tvoe , BoIivi (U M!-lZ

5894 1). from which be had removed one of lhe malil.lary bones . T in)' teeth were "isible on this bone. bUInot on the other maxilla..... hleh was lefl in place .Thb )l.Igp:sted tholt m.tensibl.)' lOOlhles.s specimen) orP. piaus hast: concealed teeth. I cjeared and !>Uinedfour such specimens: Patte rn 4: Tere vlnha. Bras il(LAC M 60977 , J~1S 1b80 ): Patte rn I : EI Porton,Bolivia (LAC M 6(976): and Subpattern 60;: KUl\lpungRbe r. GU)';tn...CLACM 609711,: the maxillary and pre .maxillill"}' bones lack teeth . Thus. at least !oOI'ne osscn ­sibly toothless specimens of P. pic/ us do lad teesh .

In some case~ (e.g. • specimens bearing Pattern 5,from "' eval i, Peru ), all presc rved spec imens nf P.pi, ·IN.< from a given locality have tee th, but leelh arcnot unifonnly developed in all ; tee th are prominentand easy to see wilh a dissecting mic rosco pe in somespecimen s and tiny and difficult 10 see in olher slX'\: i·mens . In other cases. some specimens from a givenlocality have teeth. "nd other specimens from the samelo..:ality lack teeth (at least. teelh are not visible inthe intact specimens). These data suggest that P. pictusonce was denli gerou s like most oth er spec ies ofPhJ llolwlts , thaI it is evolving toward a tooth lesscondition. and Ihat this evolution ary trend has pro­ceeded al diff erent rates in different populations ofP. pic/ us .

Toot hed specim ens of P. pi<'tus are known onlyfrom Peni and Bolivia, from lho: following localit~;

PERU: trail between Satipo and Puerto Ocopa ( ~ IL/.

24433; LACM 60 979: the la tter was cle ared andstained) ; Luis i...na ( MCZ 58 413); ~evali (LAC M92-tl 3; KU 14435 1-60). BOLIVIA: Santa Cruz (i nthe liCries BM 1904. 10.29.75-82, some specimenshave tee th. and o thers lad tee th); Buena\· i ~ta (BM1927.8 .1 .49 - 5 1 have teeth; UMMZ 665 12: one speci­men has reeur. anolher ""ilb the same museum numberlacks teeth ; four uliler UMMZ specimens and si llAMSH specimen r-om Buenavisra lack visible teetb):Guayara Me,;n (BM9J9. 1.5.1l: Yungas de Cocha ·bamba (UMMl 6664<:; ); Ivoo (UMMZ 5894 1 has teeth:six UMMZ specimens and two MCZ specimens Iromthe same Incalit y lad visible teeth): Rurrenabaque(UM MZ 57550) , All ol her specimens of P, I'iclu _,'I examined (with a dissecting microscope : the spcci­mens ....ere not cleared and slained) lack visible teeth .

Swm//(·h ('on/nilS {two spccimens). _From LACM60976: HYMENOPTERA : Fcrmicidac: Ph"idollf sp.(2 ). Compo"o/uI sp, ( I ), Do rJfflJrmu sp. (2). a lsos ix unidennfiable ant) , Mymaeida e (I); COLE O P­TERA: St"ph )'linidac (l J. Curculionidae (3,: HEMIP.TER A: Aph id idae (6 ): DIPTERA: Drosophilidae:Drosophila? (2).

From LACM 60918 : HYMENOPTERA: Formieidae :D .-.Jichodlf rus? sp. (I ), Pltlf idol lf spp. (4): ISOPTER A:....·~lI1illf~J (39) .

....omlfffCJo/"rlf.-J de~jgnalc MHSP 49 10 (one of1....'0 S}'nl)"pes bearing tho: same museum number. the

BUUETIN OF THE NATURAL HI STORY MUSEUM OF WS Al\'GELES COU.\ 7 Y No,27

PilTill«lolyJ'C is a 24.0 mm femllle), a 22 .0 mm malefrom SlInta Cruz, Santa Cruz. Bol ivia . lIS the JecMy J'Co f Hy/apll', iu pic/u Bibron in T-ehlldi I IB8 . I desi~·

n..le B~I RR.I947.2.15 . 17. .. 21.0 mm female fromYwimll~U3s . Rio Hual laga. [Loreto,) PnU, coll«tedby Dr. Hahnet. as the lectotype of lRnJ robm t s Nlhnt /i

BouIenger 1883. I designate SHM W 3818 :1. a 2b.5mm female. proba bly from Sao Vicente, MaIOGrosso.Bra..i1, co llected by J. Nan erer , as lhe Iecl<ltrpe of[Jtlldrobult. bcuccalU. Fiu in ~-.:r in SleindOIChner 11164I <.bil;nate SH~IW 19 190: 3. a 2':!.5 mm mille . prob­abl}' from the Rio !-la more . [RondOnia.1 Bra-il. co l·jeered b} J, Neuerer. as lhe kctotype of DtnJmbult.

t",nl'mi. Fttzin ger in Stein dach ner 11\h4; thi, is lhe.sp::c imen illus trated by Steind ac hner (1864 : plate 13.figs. 2, 2a -<l).

DiJ/rib" tio" "nd h"biwf (Fig. 13). - Elevalion rec ­ord s are from six 1091 5 m. P. pieru. inhabits Guyana .S ur iname . Fren ch Gu ian a , rne Guianan region ofVenezuela . most of Ihc Amawn drain age of Colo mbia .Ecuador . Peru . Bolivia. and Bras il, and lhe northernParag u a y - p.aran~ drainage of Bolivill and Brllsil so ulh10 about lifS lalil ude .

The type loc ality' of P. pier" . (T K"hudi 1838) isSanta Cruz [Bolivia] . bUI Dumeril and Bibmn (184 1)l'llid tha I lhe types CiIIlK' from Chile . GuichCoot (1848)

could not affirm tbe prese nce of this species in Chi le.E .R . Dunn (in J;II .• quoted in B. L Ull 1952 :( 11)bel ieved thatlhe types came from Bolivia . GuilX (1950)listed the type locali ty' as Santa Cruz [Boliv ia ]. P.piC-flU has llOl been reported from Ch ile si~ Dumrnland Bibro n's work. P. picrUJ i\ a Iowlllnd Amazonian'Jl«ies; it is unlikely that il occ urs in Ch ile .

Loca !ifil'. :-cQW.\IBIA. .•/~fII: 42 ~m S. 22 Lm W SmMan in tMVZ 63120. 631~9); Rio Gua)abocro. ~h.; art'na

(USS .\t 1449SO-S8); Rio Guayabocro abowc EI Rcfu, io(US~M loU9S9-61); upper GUejar ond EI Mico. Ma; -..at USS M 144962_67); Cana Guapaya. Mllurt' nli IUSl'oi M14726'). A.....tona . : 7.2 Lm ,",W 1.elicia (LACM 610(8)

ECUADOR. - l\"apo: Sanla Ceci lia (KU 104764 _72.109291 .109293_94.109296. 111$()4. 111519_28. 122271.122273-7 4. 122277_79: .\ICZ S8414_ IS, S8411); limon·c"" ha (LACM 72672-7S; KU 99119- 20); Purrlll libre.Rio Asuark o (KU 112287-90): L.,rclo (lAP 3'S8j, PU.IUU':Sarayacu (KU 120671_72); Alpayacu ncar Mera (UMMl<XJ:w,I)

PERO.-1Al'?tf1: Lago de Mirano. moulh Rio Napn(AMNH43156. 43IS8_S9. 43162- 64. 43166- 67. 43172_76. 43178_81. 4l2 ' I _' 2; UMMZ 89983); conOuence Rin. C...nlayaand Tapiche (AMt"H 42773, 4299~ . 43, S7); divide helweenTapid w: and Ul<>quinia drainascs (AM)';H 42474. 42691.42698. 43372): Ollama near Qucbrada Ca. hi"")a (A.\IS H42977- 80); Rio Pi>qui (AM!"H 4J2 7JJ; uppe' Rio Ila) .C AM~H 4.'-4(7): C hiboya tAMSH 42070); IQuilOt CAMSH42209); Pampa He Rio Cu.haNlay (AMS H 42372.424~): Rio Tamal'a tAMSH 43360 ; Con'amaN (A.\t~H

42412); SobnI. Jo,o,-er Rio Tama)a (A.\ IS H 42881); Tapi.tlca.OJ'P'O"ilC Conwnani (AMS H 42930-32); Yurimasuao. RioHualla, a tB.\' 1947.2 .15.14-20); Dumo Sanla Clara andCampo SantaClara.nearOreliana tUSS .\t 127201. 127203--08.127932). S... .U;Uf;. : Achinami... . Rio H..alla, a (AMS H

42375, 42WII- (9 ); Tocacht CA MS H 4272 1); S....andlCnear Juanjui. Rio Hllalla, a ( WCS 2757- S8). H" I" a, ..:.\Ionl(c Alegrt . Rio f'xhi'ea (AMS H 43009. 4;\(36); RioPachilCa near Bosque Nac:ional de lparia (TNHC~I .

36469_70. J6476 . J6482. J6497) . J••;.: Irail bocl..-ccnSalipo and J>uc:no Ocopa (MCZ 24433; LACM 1(0979);Owo:Jwnayo tAMS H 42892-94 ). Pasco: Sewali (LAC.\192413; KU 14.1351--(0); TS~TC1Ii (color slide by A C.~n). C.W" Luis.iana. Rio Apurimae IMCZ S8413J.

BOLlV IA. -S".,a Cral : Santa Cru2 (MHS P 4910;CM 2462-63 : BM 1904 10.29.75-112); Buenawista ( A.\l~ll

22637_38. 3J9S9. J.I075. J9S62-63; 01 4l11_27 . 4387.4416. 4423- 24. 4444; UM.\IZ ~35-40. 638811. 66507-12 .66514. ~-45. 66588. 66592; USSM 118701-03: .\ICZ128S9; BM 1927.8. 1.46-SlJ; Rio Coloralo (0 1 4308-09 .4311-1 9): Rio Surulu (CM 3812); SarilOjaI (USSM 14(598):EI Porron (CM 36155; CAS 97762_63; MCZ 29813-2 3•29825; LACM 44399. 60976). Br"i ; Gua}'a' '' · .\Ier;n. RioM~more tCM 2667; USt"M 123% 7; BM 1939.1.5 .1): Ru·rn:nal>ll<jue (UMMZ ~7SSO. SM940, S'94' ); Cavina. (UMMZ58950): Ivon (UMMZ ~ 8941, 5M949: .\ICZ 10096 _97);belwcen Reye. ~nd R um:n~ba<j 1lC (UMMZ 58944), III PUl:lower Rio Bopi (UMMZ S7549): Huach; tUMMZ ~ 8945.

58951): Tumupasa tUMMZ 58942); upper Rio Bcni belowHuochi (UMMZ 5 M~43 ) : Co""ndo tUMMZ 5M946); hiama.(UM.\lZ ~8947: USNM 147221: MCZ 15151, 15153); SanFermin (Ande...."on 19\)(',).

BRASIL.-A.III"U'II"" Rio Tacbila near Leticia (LACM60999): Tert sinha near 1.elicia (LAO l 60977. 61002_10;AMl'oi H 7~57); Eirunepe. Rio Juru.i (B_ LUll and K in,"(952); hac"" i (B. LUlz and Klms 1952): Ben,iamin ConS/anl(CAS-SU 11874: B. Lulz and K..... 1952); laB" de JanOW).Pixuna channel neat Manaus (LACM 42293- 95). !,,,,.,;:Serra do E~rt (CAS 97569-70; CAS·SU 1I8St>. 11858.111l6()....65); MonICAk i rt (BM 96.6.29.18; KU 93IS1-52 );Jac"';· Acan, a. Rio TapajOs CB. L UIZ and Kloo. 1952).Ro.i...,.: Guajari . Mirim (f M.-.H 49770. 49844); RioMamo,,!; (l'oi H.\t..... 19190:1-4). A.cn; Tarau... i ( ~ISH

83248). G<iM. : f aunda OIiYer (US:-iM 149446--48); Cawal.eanti (Miranda Ribeiro 19261: Rio Sill Mi~ ue l (CAS·SU11867_69). .".'0G,.....o: C","pada do. Guimarxs (KU93139---40; A....SP 13414); Sill Vicmte Cr-c H.\"'· 3818:1_2);So. so Sanliuimo? (Slcindac hner 11164); Chawanlina t SLw2 19S2); U"",um mounlai", ne. COOtIl>bi (CM 2431_36;P..-ISH 9209_ 19: USSM 115974)_ 3/ i" ,.. G.....I;. ' BdoHorizonte tUS:-iM %986; CAS 98099; KU 93141- 42) ;Lagoa Seen. fOOl of Serra do Curral (UM.\IZ (10023);Aeaba Mundo on Serra do Cum l (B. Lutz 1952): Cai'll deArcia ncar Belo Horironle (B. LUll 1952); UIs"llnCe tIC~r

Scm do Cabral (B. LUll 1952)

VENEZUELA.- Bo/i."" 5 km E La, Chicharra' . Al ii ·planieie dc Nuria (MBUCV 31( 2); Hacienda San Felipenear Miamo (UPR·RP~.m.

GUYANA,_ M"l",,,,, i.P"ldro; Kanabo (A ....I~H 11667.23120-2 1. 23126_27. 24925. 39S7S_76. 39677_78• .19681_86. 39692-9 5. 3%9 8. 39702_04. 39706-07. 39709_17,70984_87; CM 4071. S437- 38: UM.\tZ 83596: BM 1925.114,8): jlllK'linn Mcamu and .\Ial.lllllni Rjw"" (AM!' H 18996.18998); Kamal u... (AMSH 19(100); Kurupung Rjw"r(UM.\IZ83S98 _60) ; LACM 60978); Mad reba Fall•• Kurupun,River (UM.\IZ 85147); " . ano (U.\l .\tZ 83597); Paruima.\I i~~ion (UMMZ 85145-46: US~ .\I 1I8681-82); CUlllniRiVC1" 24- 32 Lm a~ Banic-a (B.\I 1934.11.1,1-4) ; !ol .. opcFall•• CUluni Riwcr {BM 1934.11 .1.51; Oko Mountain.(P.>IS H 26724--26)

Sl:RISAME.-."kb rir, Sipaliwini (.\ISH 1lOt'>. 1131); Pn>lGonini. Cue"",ni RivieTt.\lSH 322. 350); Post Tip. Sc..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 ""Maripa>oulIl (La 1023).

DOUBTFUL RECORDS .-COLO~IBlA. Cllltlar: VillaMaria (lJSS M 14Nlbl. (46866). BOU VlA. ClN1ttlba"' N:Yungas do: COC hatwn N. 2200 m (UMMZ66646).

Clear ..d and stained mm..rial, _ICClh pre",m: LACMb0979; Pattern I: loACM b0916 ; Panem 4: LAC~l 60977 .JMS 1680; P.~Uern 6: LACM 60978 .

PhyllobQU J pukJrriputus new speciesBeautiful- breasted Poison-arrow Frog

Frontispiece II and Figure 6E

HolofJpt' . -loACM 42297. co llec ted 1.1 Serra doNavlo, Terruorio do Amapj , Brasi l , abou t 120 m.between 1330 and 1715 h on I lu ly 1968 . by P.A.Silverstone and Sr. Ferreira .

Topop Qf/l1rp" s (nine ad ulls) , - WCA B 18 22 1;LACM 42296. abou t 120 m, I July 19MI; LAC M42298-301, about 120 m, 2 luly 1968; LACM 42302.about 160 to IIlO m, 3 Jul y 1961l: LACM 42303. about100 m, 5 July 1968; LACM 42304 . Serra do Veado(one of the hills at Serra do " lIvio ). abou t 290 to3 10 m, 6 July 1968. (All LACM parIIt)~ were col­lected b)' P.A. Silve~onc: and Sr. Fem lra.)

D..finilion . - Tet th abM:nt; IClC' S ",eblt ss; dorsumblac k; complete yello w lateral \ !ripe; li!!:ht pro~ illlO­

ventral ca U·s po t abse nt : ven ter blu e ",ilh hlackmarbl ing.

Diagnosis, -Ph.l'lIob<ll'" pr,lchripU:fUJ is a memberof the piclII _' group. dif fering from P. i" gt'ri in lackingteeth, in having a compreie light lateral stripe (al'>sentin P. i"g..,i ). and in lackin g a light proxim ovemra!calf-spot, from P , fXJrvufus in lacking teeth. in havinga black oorsum in life ( red in P. p<lrvufus). and inhn in!!: a complete li ~hl lateral ~tripe (incom pktc inP . p<ln 'uIUsl. from P. piNUS in lack in~ a li!!:hl proximo­ventral calf-spol. and from P. )IJ'<lrQgdillUS in lackingteeth and in ha\'i n~ ventral marbling in life . P. pul ­ch ript'clUs resembles P. ~t..,si ( F i~. 98) and P ,bolil'iQlluS (Fig. 9 A) in it. ventr a l patte rn of da rkmarbli n~ on a lig ht ground color. P. pu/chript'CIU.•differ" from P. " n t'rsi in Ia.;:k ing teeth. P. pulchrip<'('/u.,diffe rs from P, 1",/;,'i,mIlJ in having a conspicuouslight proximod orsalthigh-spor (Fig. 6El. which is whitein preservative and bright yellow in life. In P. hdi ,'i,mUJ(Fig. SC). tbe dorsal surface of the Ihigh bears light.lonJ!:itudinal . someumes undulat ing markings.lk~rip'i"" oIItoJOl)pt' . - The holol)'pe is an ..dull

female . 27.0 mm ~noul-ven l length. Thc , k in isstrong I)' gm'lular on the donum and the donal sur­face of the hind limbs. and smoo th o n the fo relim bs,the venter. and the ventral surface of the hind limbs.Teet h an: ab"Cnt. 1llc lip of the Mlout is sublruncatein dOThal aspect and rounded in late ral aspect. T hecanthus m stralis is rounded. The loreu l reg ion isvertical. The width of the upper eye lid is three-fo urths

of the interorbital widlh The diameter o f the eye isI A t imes the distance from the eye to the lIOloIril andslightly more than twke tbe diameter o f the I)"mpanu m.The pos terodo rsa l portion of the tympa num is con­cealed. The finger dis ks are small relative to those ofD..nclm hll/ t'J . The first finger is about equa l in lengthto the seco nd. T he f ingers are webt ess . The tarsalfold is ill defined and slightly curved , extending alongthe distal 4S perccnt of the distance between the innerme ta la rsal luberc le and Ihc hce l fold . Th e tarsa ltubercje is absent. The loes are ....ebless

In life (Frontispicct Ill. the iris, head . ~k. andsides wert black . A com pklc brigh t )cllow lateralstripe e xte nded from the Jroin 10 the upper eyefid.along the octer border of the upper cyclid. and w n!!:tbe canthus rosrraiis 10 the lip of the soout. ",he n: ilmel its fellow. Anolher bright yellow stripe extendedfrom the forelimb insertion along the upper lip, end ingventral to the nostr il; the anterior end of the stripe WIlS

blue . A blue suffusion clouded the black limbs. Theproximoo or",,1 surface of the thigh bore a con~picuous.

o blong. brighl yellow. black-bordered spot; a similarspoe adorne d the axilla . euendi n!!: omo the pmximo­~ior and prox imodon.al surfaces of the upper arm.The venter and the ventral surface of the limbs weremed illm blue , beann!!:discrete black marblin!!: ,

In pre~rval rve . the grou nd color is black . Are asthat were ye tlo.... in life are whi te. The canthal port ionof the lateral stripe . the blue an terior end of the upperlip str ipe. and the vent ral pattern have d isappe ared;the venter is black , ellcept for a small while spot Onthe left side of the posterior belly'.

Dt'Scripl; on 01 p<lrQf)pt's . - Measuremcnts of adultsare in Ta ble I (one parnt ype, LAC M 4230 1. is ajuve nilc femalc). All parIIlYpeS lack teeth. In some.lhe ca nlhus rosIra lis is near ly anj!ular . All have theIlrst finge r aboet equal in k ngth 10 the loCCond. Insome para l) pes. the tarsa l fold is "' cll defined, u­lcnd ing along lbe distal 50 10 60 percent of the d istancebetwe en the inner met atar sa l tubercle and me beelfold . and the tar\31 tubercle is lI'cli developed

In life . the paratype s resembled the bolotype. Theanterior end of the upper lip stripe usually was blue .The prox imal spot on the upper arm was yellow orcream . Some spec imens had blue spots on the forearmand the pos te rodorsal su rface o f t he th igh . T heprox imal spol on the thigh II''''; round o r triangular .The black venlral marb ling wa.~ d iscrete or continuous.

In p-ese:rvatr.e. in LACM 42300-01 , the upper lipstripe con tinues ante riad 10 beneath the nostril. Insome specimens. mosl of the dorsal surfece of theupper arm is white . In LACM 42298 - 99. the thigh·spot extends o bliquely distoposteriad from the baseof the thigh. The ventral paue m of dan marbling ona light ground is distinct only in LACM 42300 . LACM42303 has a small whi te median spot on the posle·rior belly .

44 BULLET/,V OF THE NATURAL HISTORY MUSEUM OF LOS A.'liGEU.S COUl'tTY N(}.17

R~IQli(»/$hipJ .-P. pulchriprCIUS probably is closelyrelated 10 P. bvli"ianus and P. prf" rsi .

Sw much comems (one specimen) . _ From ~C~t

42302: HYME....OPTERA: FOIlTl id dao:: B/t'phuridanab"<l~j{jtnsiJ w hle. ( 2), M.w·ocr puru$ smilhl (Fo rd)

(II, Cypho myrma rimosus (Spino la) ( I), M.m nico­

crypt" sp. (I), Myr micocrypl a ? ~ p. (41. Pht'idolt'~. (8) , H, po ponrra ~. (2). Wasma" " ;Q auroputICtUlQ(Roger) (5), also fragmen15 of lints of three 10 fourothe r genera .

Distribu ,;OII aJU1 habi tat (Fig. 16). _ Elevll l ion rec­

erds an: from 100 10 3 10 m. P. pulchripulus is knownonly from the type locality. in the Guianan region ofnonhem Brasil near the Rio Ama pari. I tributary ofthe Rio Araguari, wh ic h flow s into the Atlantic Oceannorth of the mo uth o f the Amazon .

I mol these agile diurnal frogs 110 tile f"re ~t floor .betwee n the scattered brief f3 i n ~ howef~ of the openingdry SC3,;on. One perched on a fallen log two-thirdsof a meter above the ground : it dived to the groundand hid when I approached.

Serra do N3Vio is a town and ma nganese mining:complex surro unded by low. fore..tc:d hills , The forestincl udes large-buttressed trees of grea t height anddiameter. Spiny palms guard the abendam melasto­maceous and ru biaceous under gro wth . The groundbears a heavy lear cover: some portio ns of the groundare poorly drained.

From 1954 10 1964. Serra do r-;avio had a meanann ual rainfall of 2402 mm. seventy pe rce ne of itduring the ~ ingle we t seaso n from lanu ary 10 June .The mea n annual temperature .....as between 25 and26°C. ee mean minimum was 22.2"C. lind the meanmuimum was 32.a'C (Im.litulo Rq ;ional de Desen­volvimento do Amapii 19(6).

EtymoJogy. - Lalin: pukhu (beautiful) and pectus( brea~l) refer to the beautiful blue venter.

CII'"rI'J alt(J !wi","d m"I"';"'. -LACM 42:.'02.

PhylfQbatn smaragd inus new speciesEmerald Poison-arrow Frog

Frontispiece II and Figure IIC

Hol<>typl' . - LACM 64435 . co llected al Pan deAzucar . 39 km N~EOxapampa . 10"15'5 . 75°14' W. inthe lscozaztn Valley (Ihe Rio Iscozazm is a Iributar)'of the Rio Palcazu . in the Pechlte a drainage). De­panamenlo de Pasco. Peril . 31lO m, on 16 Augusl 196 1.by R.E. Elheridge .

Parur),fWs (one adull and seven ladpoles). -LACM6+436. jopoparatypes (seve n tadpoles from the backof the holol)'pel: LACM 6+434. collected at lscozazinainlrip. 47 km r-;NE O u pampa. urn ·s. 75°10·W.in ttle lscozazin Valley. Pasco . Peril. 360 m, on 19l uly 196 1. by 0 .8. Wake and R.E. Etheridge

D..fi nilion. - Teeth present: roes ....ebless: dorsumblack : co mplete green lateral stripe; light proximo­ventral cal f.~pot abseet; venter green (anteriorly) andblue (posteriorly). no l marbled.Diagnosis . -Ph.~-IIobuJf'S sltUlragdinU$ is a member

of the piNus gro up. differing fro m P. boli,·jaflluand P. pukhrifWct u5 in having teeth and in lackingventral marbling. from P. ing..ri in having a complelelight la tera l strjpe (ab...:nl in P. ing..r i) . in lackinga light pro ~ifno>.'C ntral cal f-spot, and in lacking vent ralmarbling. from P. pan 'ulus in having a black dorsumin life (red in P. pa n·uI U5). in having a complete lightlater al stripe (incomplete in P. p<ln ·Wus). and in lackingventral marbling . from P. pet ersi in lack ing ventralmarbling. and from P . pit:rU5 in having teeth (usuallylacking in P. piCIU5). in lack ing a light pro ximoventralca lf-spot. and in lacking: ventral marbling.

Descripuon of hO[Oly/'t' . _ The holotype is an ad ult

male. 26.5 mm snout-vent length. The skin is stronglygr<tnular on the dorsum and the dorsal surface of thelimbs. and smooth on the venter and the ventral surfaceof the limbs. Maxilla ry and premaxillary teet h artpresent: vomerine teeth are absent. The tip of the snoutis subtruncate in dor",,1 aspect and rounded in I..teralaspect. The canthus rostrahs is rounded. The lorealregion is slightly concave. The width of the upper eye­lid is slig htly 1('S~ than the interor bital width . Thediameter of lhe eye is slightly greater than the di~1aIlCe

from the eyt to the noslril and slightly II1Oll: than twicethe diameter of lhe tympanum. The posterodon.al por­tion of the tympanum i~ concealed. The finger disksare small relalM to tbose of D..lldrobaltS. The I irstfinger is abou l equal in length to the second. Thefingers are ....ebb s. The tarsal fold is slightly cu rved .eMending along the distal 50 percent of the dislancebet....ee n the inner metatarsal tubercle and the heelfold. The tarsa l tubercle is slightly developed. Thetoes are webless.

In life. the dorsum. the side of the body. and lhedorsal surface of the limbs were black. A bright greencomple te la teral stripe ex tended from the groin 10

the upper eyelid . along the outer border of the uppereyelid . and along the canthus rostralis to the tip ofthe snout. where it met its fellow . Anoth er brigh tgreen stripe extended from the forelim b insertion alnngthe upper lip. ending ventral to the nostril. The lowerlip. the breast, and the anterior t.....o-thirds of tilt bellywere green. The ventral surface of the limbs and theposterior one-Ihird of the belly were dark blue. Thethroa t bore a pair of lateral black areas (perhaps a malesecondary sexual charac ter).

In preservanve , the holo type is entirel y black 10dark gray. ex.ccpl for lighl gray lateral and upper lipstripes .

Adull parat)·fW .-l n preservem..e. ttle adult parl­type (LACM 64434) resembles the bolory pe: it isthe same sex.and length .

1976 FROGS OF THE G£SUS PHYL I.O BATES 45

Tadpolet (Figs_ l ID. 1 2E).-~ ho\ol)pc earned'>eVen ladpok<. (LAC M 64436) on hi, back. The lad­poles ...'ere in SliI~~ 24 (1....0 specimens) ;md 25 (fIVe

specime ns). ~lea.~uremenh an: in Table 3.Dist~ibutiOll and habitat (Fig. 16). _ EIC'IIlIlion rec­

ords are from 360 to 380 m. P. smaragJinll.• is kno....nonly from the type localities. in dense forest in theAmazon drainage of Pl:ru. east of the Andes.

Etymology. - Latin: smarafi:dinus (emerald) re fersto the green stripes and me green anterior venter.

l~ivl1llllUJ group

Th is group includes twc species. (f'hylfoiJ<,ft's bass·lu i. P. trivitla tus).

Defi nilion: snout-vent length of adults moderate10 large (3 1.5-49.5 mm); loes ....ebIeM; paired ~~lateral (Figs. 2. 6A-C); light proximoventral calf·spoI absent.

Phyl/oootes banlui (Mclin 194 1)Pleasing Poison-arrow Frog

Figures 2F. 6A-D

t941. DmdrOM/<'J ""'.,Iui Melin: 6~ -66, fi ll . 3f1, holo­type: NMOsn. R'>quc. ISanManln.! Peru(,t 097 mi.

Defi nition . -Tee th prese nt; toes webless: dor sumblack with green or ora nge spo ts or mar bling. o runi form orange: complete green or oran,e la te ralstripe; Iighl pro :\imo\'entral cal f·spot absent; venterblack with bloc marblin,.

Diagnosis. -Ph.\'//oba/~J batsleri is a member oflhe trivitta lllJ group . di lTering from P. tri,'itralll.1 inhaving teeth .D~sc~iplion . - Meuurements of the bclorjpe (a

37.0 mm female) and referred specimens are in Table I.The skin is strongly granular on the dorsum and thedorsal surface of the hind limbs. and smooth on theforelimbs, the venter. and the ventral surface of thehind limbs. MaxiJla r) and premaxillary teeth arepresent. The first finger is longer than the second.The toes are webless.

In life , in the holo type (Melin 194 1), the dorsumwil.s black ""ith a median row of light green ~pols (Fig.2F). The side of the body was black ....ith a completelighl green lateral suipe extendin g from the gro in 10the upper eyelid, along the outer border of the uppereyelid. and alo ng the ca nlhus rostralis to the tip ofthe snout.... here it met its fellow. Melin did 1101: men­tion the colo r of lhe venter in life.

In preservative. the bolo type is dark bro....n. exceplfor the following white areas: a median doNiI row ofseven spots. a fe.... small spots on the righl s ide ofthe beck. a complete lateral stripe on each side of the

bod)'. pan of the upper lip. the proxrrnodorsal surfaceof the upper arm, lrac~ of light pigment on the donaland ventral surfaces of the thigh. and the proximo­ventral half of the kft tarsus.

Refe~"d mOfuiaf. - Tbe holorype is from Roque.Per u. at 1097 m elevation on the Ama7.0nian slopeof the Andes, in the drainage of the Rio Huallaga, 3Skm south"'lutheast of Moyollamh a and 76 km west·northwest of Chazuta . I refer nine additiona l toothed

specimens to this species . Tbese specimens are fromeast of the type locali ty. from the drainage of the RioHualla ga (Chazura. Pachiza , and the Rio Cainar acbc j.

Specimens from Pachiza differ from the bolorypein having smaller. scattered dorsal ,pOls.

Some of the spe<:imens from Chazuta have a colorpattern similar to tha t of the holotype: I do no! knowtheir color in life. At Chazura. W.C. Sherbrookecollected lOOlhless specimens of P. trivunuus (....ilha pair of yellow lateral s.triJlC'o and a )'ClIollo mediand~1 stripe) together with three too thed speccneos(....ith a pair of orange lateral stripes and no medianstripe). The three toothe<! specimens (US !'\M 166903A­C) collected by Sberbrooke mal belong to an unde­scribed spec ies. but I provisionally assign them to P.basslrr i , Two of the toothed specimens (Fig. 6A-B)had a black dorsum with orange marbling: the thirdspecimen (Fig. flC) had a uniform orange dorsum.The venter in the toothed specimen s was black withblue marbling (Fig. 60).

Generic oflm:ulion._ Mel in (1 941:65) desc ribe dthe boloty pe a.~ a Dend~ooo"s because he thought ilhad no teeth. onl)' "tooth-like aspenties." I examinedthe holol)pe. It has teeth and small finge r dish; itis a Phyf/obates.

Stomoch contents (one specimen)._From A."-tN H43402: HYMEl"O PTERA: Formicidae: Pheidofe .p.(21. CompOflOflls'! sp. ( I); O RTHO PTE RA: Telli·goniidae ( II: HEMIPTERA : fragments.Dist~iburion and hobi tof (Fig. 141.-Elevat ion rec­

o rds are fmm 274 to 1097 m. p , bllss/eri inhabits theAmazon drainage of Peru. from the easte rn foothi llsof the Andes cast to the Rio Huallaga .

LOCilfiti H ,-PERO. Son MOrf;n: Roque. 3~ km SSEMoy"bamba (I'\MO ~1l); Ch:llUI~ (AMI'\H 4Z867. 42944:USl'iM 166903A-C); Pachiza (AM~H 42321, 42333, 434(2);Rio Ca;narachc tAMSH 42313).

Cfea~eJ und swi lled "I(l/e~iof. -AMS H 43402.

Ph)'ffoba fes trivinanu [Spix 1824)Three-stri ped PoisorHul'Ow Frog

Figures 2A-E. 7C

1824 H,·/o ,ri'~II(l1O Spi1: 3~. pla....'I, fig. I. type locality:Rio Tde, BnI~il

?1824 H,./o ";lIa~i"'(l Spi1: 36. p l ~le 'I, fig. 2. IypeJoealiIY: Ega [=Tefel. Braoil (according 10 W.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\~

noI "Cen tM I~ pe~ !he)' ..-ere ck"roye4 .. WoOdWar U. axordin!, 10 U. GlUber lin lill.)).

1826. H,-kIpI'Wl";'~lIaUl , 50., in Schkp:l: 23918.\0. lHNlTt>ba". Im·iltalM•. Wal lct-: 202.?18.\O lHNlTt>ba"...i~rriIJIM' . War.lior: 202.18).8 H,"""",Wlli~. T..: hudi: 70-7 1.18~8. H,.lapI,.;" Ii...."'"" (pan). GUnther: 125.111M o,NlroDQU. "iK"rrUorIU ( part) . Ste ildal;hner. 259-

260.111M lHNlrobm,. ti....1OTi1U ( put). Sa: Illda<:IIMr: 260 .1&82. lHlldroD..,,. rrMllQ'1U (pan). Boult-np:r: 1 44_ 1 4~.

1926. O,lIdrol>,,,,. trivilla'M.'. Miranda RiMir o: 181.!ell·fi8. 98 .

1926. On.droba,,. U lr(Jvit lallU Mir~nda Rib<oiro: 180.!e\t· fi8. 'n . •yntype. in tM Mu~u Paulio.ta . typeIo<;alil.,,, 6 bidos. Brasil. and Rio JUllli. I Ei llllll:~.

"",«Jrdin8 to Bokermann 19t16.J Br•• il (accordingto Bokermann 11966:3 41. O. 1, I'a"i",,, o. i, asynonym of D . Iri.in",... ; I have not ,ee n the_yntypes).

19~2 . Orlfdmh<tln ,rMlldl...,. B. LUll ond Kin,,: N:>91\161. l >r lfdm lml r , I,il'illat...,. Cochran 107. plate 41

(cnl", phoIO).1969. Drndroh<tlr. "Mlltll... . Hoogmocd: plate I. fig. 4

(col", photo).1970. DrMrobaln " i, ·illal... (pan). C"",hran and (lo;>in '

14 -1 6 . plate ~G~I (o nly USNM 118681 _ 82.MCZ 24443. ANSP 18188 are n<Il P. Iri,'illa"u).

Drfi ll i t iOlI . _ Teeth a~nl; Illes web Je . , ; dorsumbladr. I\-'ilb or I\-itnou ( yellow, yellow-green, or greenmedian donal stri pe (medi an stripe some lirTlCS bro kenmto long segments or , bon spols); somet imes dorsumbbd; with green miirbl inil; sorrenmes dorsum uniformg ree n; complete yellow, ye tto ..... ·green. green. oro r;IlIFC lateral srnpe: light proximoventral cal f.spotabsenl ; venter blac k ...-ith grren. blue -gree n . Of blue'flOls or marbling.

D;oxn o.1;$.-Phyf1obol~$ Iri"i tl(JI"$ is a mem bero f lhe t ri" it/<JtUJ group. d iffering fro m P. boulrr;in lacking teeth .

Do cript;oll . - Me asuremenls are- in Tab le I. P./r;";/Illtu,' is the largest species of Ph.\' II"h<I/~ ' , Theskin ts slightly to moderalely granu lar nn the dorsumand the dnNtI surface of lhe hind limhs . and _nmotbon the furelirnbs . lhe verner . and the ventral surfaceo f the hind limbs. Teeth are absent, The fiN fingeris longer than the second. The toes are- weble...

Phyll l.hll/ l'.1 trlvittatu s has three co lor pauerns:Pattern I (Figs. 2A . D- E. 7C; include s holotypc ofHyl<J /r;w'/Iu/a Spil 1824 ): Specimens bearing thi spattern are known from Faro in Bra.il \\-'C.IWan:! alon glhe Amazo n to the Pengo de Ma n1<C'riehe (Ihe passof the Amazon through the Andes) in Peru. and w Ulh­...an:! in Peru alon g the Huallaga and ib drainage al

leas t ~ far as Tin ge Mat ia a nd aJong the Ucayaliand its drainage at leaS! as far as the Rio Tamaya(SOUlhea'" of Yari nacocha). In lhis pal1cm. lhe groundcolor of tbe doroIllm is blac k.. "There are three )'ellow.yello ...-green, or gree n st r ipes (one median dc r ... 1

and one- pair Jateral). In some specimens. lhe mediancorsal «npe 4 , one- or mo re small break. (Fig. 20).Such spec imens are known from Santa J ulia ne.... Faroin Bras il. Puerto ~ariilo in Co lom bia. and Morale s.Pschiza. the uppe r Rio Cllshah:l1ay. p...mp;.o Hen nusa(lower CIl,habalay). and ~.5 km from Puerto SantaClara in Peni. At 1IlO'>l of rbe-e localities. boIh ,pecimen, with an untomlen median ...ripe and spedmen,I\-ith a broken median smpe have been conecred. One­spocimen (AM~H 43 117) from Pampa Hc-rmoSll. . P.;ni.has a med ian don;a.l row o f e ighl sma ll ,;pots (Fig. 2E)."The spec ime n lach teeth , and is pan of a 'Cries o ftooth less specimens from Pampa Hermo sa whose pat­lerns include transitio ns bet.....een an unbroken med ianstripe and a medi an doNtI row of spot s . In life . inthe holot ype, from the Rio Tefo1 (a southern tributaryof the Am azon east o f the Rio Jurua) , Brasi l (Spix1824:plate 9. fig. I). the dorsum was black with a)'ellow median stripe (bro ken intcrorhitally}. T he s ideof the body "la, black with a com plete yellow lateralstripe . The do rsal surface of the limb, I\-'as black I\-'ilhye llo w spols and marbling . Spec imens from C ha7uta.Peru (color slide s by W .C . Sberbrooke), resem bledtbe holotype, but the med ian slr ipe was unb rok en .and lhe- Sl\oOUt and the-dONlJ surface of ue limbs weree nt irely yellow. A specimen from Tinge Maria . Peru( notes by H.A . Allan:! al US r-;M). resembled lhe- nolo­type , but the- stripes were yellow. green. Tbe belly l\-itS

blac k wil h green 'poIs. A specimen from 7.2 krn I'lOnh .wc st o f Let icia . Colombia (my color photO) ). rese-ml>lCuthe bokxypc. but the strip:s were green.Tbe medianstripe was unbro ke n. indi" ioci . a nd darker than tbe:latera l stripe•. TIle- donal surtece of the fore-limb til-«a green stripe- eXlCnding the IC'n gth of the fo re limb .The- doNlI surface of the hind limb was near l) cove redby green marbling and str ipes. "The ventral .urfaceof lhe fo relimb wa.s blac k wilh a bl ue- stripe extendingtbe length of the uppe r arm. The vent ral surface ofthe hind limb was blac k with blue spot s and marbl ing .The vente r was black with man y small blue spots(Fig. 7C).

Pattern 2 (Fig, 28): Specimens bear ing this patternarc known defin itely (from well-preserved specimensand from color note> and slides) from four locali ties'Yartnacocha on the Rio Urayali (60 km nonhnorth­wes t of the junction o f the Rlns Pachitea and Ucayal i).the lowe r Rio Pacbitea ncar the Bosque Nac ional delparta . and lwo localitie, on the upper Pac hitea drain­

age . Pan de- Azocar and San ta Isabel. The pallem isdi fficu lt to d iscer n in pre-SCl"o'W specimens; il mayoccur in specimens fro m OIlter localhies in lhe Pach he edraina~'C and along tbe Ucayali fro m the Rio TamboIII Con tam ana . In this pattern. lhere IS a pair of Iateno!strip:s as in the OIncr patterns. bul the cooum is uni­form green . Panern 2 could have devejoped from Pal­tern I by e,pansion of the med ian stripe. In life.in spo:<:imens from the Rio Pachitea. Santa Isabel . and

/976 FROGS OF THE GENUS PHYLLOBATES 47

Pan de Aziicar, Peru (color photos by J .P. Bogartand A.C . Thore sen: notes by D.B . Wake and REtheridge at CRE), the dorsu m woo, uniform lightgreen. The side of the body was black with a completeyellow lateral stripe extending from the groin to theupper eyelid, where it merged with the green dorsum .A white stripe extended from the forelimb insertionalong the upper lip, ending ventral to the ante riorcorner of the eye- The dorsal surface of the limbs wasmostly light yellow.green. In the specimens from Pande Azucar, the venter was black with blue-green spots.Two specimens from the Rio Pachitea bear rnodifica­noes of thb pattern. In one specimen, the dorsum bgreen with a small amount of black marb ling andspotting; in the other specimen , the dorsum is blackwith green marbling.

Pattern J (Fig. 2C; probably includes the symypeof Dendrobate s tetraviuatus Miranda Ribeiro 1926from Obidos): Spec imens bcaring this pattern areknown from Guyana, from Suriname, and from Bra, ilalong the lower Amazon from Faro to the Atlantic.The lateral stripes arc green , yellow-green, yellow.or orange . The dorsum is black; there is no mediandorsal stripe . In life . in specimens from 5 km east ofHelem. Brasil (notes by M. Crump at KU), the dorsumwas black. There was no mcdian dorsal stripe. Thelateral and upper lip stripes were yellow-green. Thethroat was black. The rest of the venter and the ventralsurfacc of the limbs were black with blue-green spotsand marbling. Specimens from Suri name (notes byM.S. Hoogmocd at RMNH; Hoogmoed 1969:plate I,fig. 4; Hcogmoed 1971: 1, 6) had a black dorsum andlacked a median dor ' al stripe. In mosr specimens. thelatera! stripes were green , but in specimens from theNassau Gebe rgre in eastern Suriname , the stripe swere yellow or light orange. Green spots ornamentedthe dorsa l surface of the limb, : light blue spots adornedthe belly and the ventral surface of the limbs. In frogson the Lely Gebergtc . Suriname . a transition occursfrom yellow-green stripe, on one side of the plateau.to yellow stripes, to bright orange stripes on the otherside of the plateau (Polder 1976).

Bruding ("a i/.- In Suriname, Hoogrnoed (1971 )found males Calling ("a long-drawn-out tututuiutu . .to " ) from twigs, rocks . and fallen logs in January andFebruary; Polder (1976) heard them calling in August.

Tadpvi",~ (Figs. I IG, 12H).-Twenly-five lots oftadpole , bave been taken from the backs of adult malesin Colombia in December , in Peru. in January , April,and May, in Brasil in February and May. and in Suri­name in February to May and in October (Table 4).Anotber lot (LACM 72046. in Stage, 29. 34. 37, 38,40. 42, and 43. taken in Peru in Augu,t) is from asmall temporary jungle pool. Hcogruoed found tad­poles of this species in a water-filled fallcn palm petiolein Suriname in February (Polder 1976). Measurementsarc in Table 3.

Re/lllio'l!ihip!i._Toothlcss specimens of P. trivuuuus<Ire symparnc with toothed specimens of P. ba.".lleriin Peru. at Pachiza and Chazuta. As noted above, atoothless specimen from Pampa Hennosa , PerU. (in tbeUcayall drainage ), has a median dorsal row of spots(Fig. 2E). This pattem is very similar to that of thetoothed bolotype of P. h",sleri (Fig. 2F). Provisionally,I am calling P. bassleri and P. tri virtatu .f distinctspecies. The green-spotted specimens of P. bassieri(incl uding the hotorypc) may merely be toothed P.mvutatus. If so, Sherbrookes orange specimens fromChazuta (USNM 166903A-C) prObably belong to anundescribed species.

Lif'"history . - Hoogmoed (197 1) and Polder (1'.I74a,1976) observed wrestling arnung c<lptive specimens.

In captive specime ns, Polder (1976 ) observed thateggs are laid on leaves, clutch size is eight to 22 (clutchsize is at least 27 in this species; see Table 4), tbeperiod from fertilization to hatching is 14 to 17 days.and the period from hatching to mctamorphosis is 41to S4 days. Polder also noted that in rbeir native habitatin Suriname , frogs of this species are extremely wary,and their coloration is inconspicuo us: thus, he doubtedthat their coloration has an aposemauc function.

Para!iires . - I found larval nematode, parasitizinga specimen of P. mvlnatus (LACM 6098 1) from 7.2km northwest of Leticia, Colombia. Seven nematodeswere free between the skin and the muscles (in thetbighs and in the side of the body), and several morewere enclosed in cysts attached to the thigh muscles.

Diu llsn . - Polder (1976) has discussed dise asesin captive specimens of this species.

DiSlrihmimr and Iwhirlll (Fig. 15). _Elevation rec­ords are from 20 to 680 m. P. trrauaius inhabits thelowland forests of Guyana, of Suriname , and of theAmazon drainage of Colombia. Ecuado r. Peru. andBrasil. .\1. Crump collected specimens near Belem.Brasil, in the daytime on the forest floor , In Suriname ,population density of P. trivillmus is greater besideforest roads than in undisturbed forest; the frogs seem toprefer fallen trees with their tangle of broken branches.which provide call ing and hiding places (Polder 1976).

Localities . - COlO MBIA. Pat"",a)',,: junctionRimA~u;Jc,

and Mccaya. upper Ca~ ueta drainage (US!'lM 144992- 93).Amazonas: Puerto Nariiio. Rio Ama1.Onas (ILS 3: USNM146831-32): 7,2 krn NW Leticia(LACM 60981. 61015-17).

ECUADOR._ [unable to locate]: La Chanla(USNM 61761)PERO.-Hua"ueo: Rio Pachitea near BO><Jue Nacional

de lparia (TNHC 36467. 36484. 36489. 3649t. 364(3);Tingo ,I.-laria (USNM 129414; AM!'lH 53762; CAS 85152);Parque Nacional de la Bella Durm;ente, 10 km by road STingo Maria (USNM 166910); Ganzo Azul (FMNH 45t43.45292): Monte Alegre. RIO ?achilea (AMNH 43021, 43025)Sa" Mani,,: Tocache(USNM 166908-(9): Chazuta <US!'lM166907; AMNH 42616. 42786-89. 42943}; Huayrapurinanear Chazula (AMNH 42952. 42954); Morales (AMNH42012); PachiUl (AMNH 42326, 42g26); Achinamisa. RioHuallaga (AMNH 42t8 3-85. 42339-43. 42509. 42545,42576. 42589-90. 43204). Loreto: Pueno Santa Clara and

BUllETIN OF THE ,VATURAL HISTORY MUSEUM Of' WS AXG£LES COUl\7Y No. 27

Doma San.. Clara ncar Orellana tU SS M 127214 -18 );Chay....it. (BM 67.6.13.24-25 ); Balsapueno tFM~H SMIl;Cachipun10 Mar Balsapueno (AMS H 428761; Rio Ca<hiyacuE Bahapl.ll:rto (A~I SH 425751; Iquito. t~'S H 109839;AMJ';H 422941; Cerro Azul. 24 lr.m E Contaman.i (FMS H56249 _$0 ); Ptocallpa tP.d ..... H 5625 1); Yarina<o<;l.... RioUeayaJi tP.d ..... H 4~-O7. ~52 -S6); Pam~ Henno......Rio Cu..tlabuay (AMS H 4!OI 1. 42013_14. 42018. 42015.41145.42255-63.42360-70. 42375-76. 42457-~ . 42662.42~. 4282S. 42827. 4292.3. 43112_17. 43121-23 . 43125.4J263. 4J4(,()..OI) ; upper Rio Cu..tlabatay (AMS H 42310.4J2oW. 43431); Barranca. upp:1" Rio Maranoo (A..\ ' S H 43345);d;' i<!e bet"'.:.:n UlQquinia and Tupic"" d...i....;c, t A~'SH

42249.41468_70. 42472-7 3. 42564. 42695. 4JO.17. 43370.43378); Puneua. Rio Tapic"" tAMS H 42873. 429151; jun<tionRim. C,mlay a and Tapiche (AMS H 429%); Rio BlancotA~'SH 42985); Rio Tamara (AMNH 421101); Ca, hiboya(A MSH 42044, 42976. 434H); Oll,nla "" .. QuebradaCa.hihoya (AMS H 43212); Rean Re<l n.l..a", Suh, ya (AMNH427(9); Rio Pi",!ui (A~1:'m H192. 4319Q, 43552); Yuri·magu.. (Bnuleng er 1883); n<:ar Tarapo lo (CoP" 1886 );Peba. (Cop" 1886); E'liron . Rio Ampiya~u (CAS 93344);junclion Rio, Santiagu and Maran,'," (AMNH 42741), Pasco:Puerto Bermu dez (FMNH 35) 8); Nevati ( KU 144) 47);Sanla [", bol. Rio Nevati. 35 kIn SE Pucno BermUdez (USNM1667641; PIon de Azuear. [",o ..li n Valley, 39 km l"NEOupampa (LACM 72035-46, 72057-61) . /"" ''': junclionRim.Tamho lllId lkayali (AMS H 43262).

BRASIL.......... lJlazo"... : Rio Tef<i (Spi> 1824); Td<i (MHNP6242); Japuri . Rio Japurii (MP£G 338_39); Benjamin Con., lanl (WCAB 133SO; KU 93153; CAS 11870-72; B. LuuaJ>d K...... 19521; Rio l",'ary. 30 lr.m ,jo,.'n>lrcamffOOl Benja­min Con!ltalll (CAS·SU 11879); hacoa ; lB. Lull and KJo.l.I9SZ); Eirtrnept. Rio Jurui (Miranda Ribeiro 1926); M.....(K. Sc'-'tidl .-.d I n~r 19S1); Rio Amazonn tUM~lZ 62S53);Paranii do Jacan: . !iO\Ith of Io"'-er Rio 501,,no" ,C AS·SU118tl6). "po: 6bido> (Miranda Ribeiro 1926); Caotanh.aJGrande ""ar 6bidt» (MRHS 61. 6I B-6IE); 5.a.nla JuliaDCar F~ro ( MRHS 4741; hiubni (ZS BS 99/ 1911; MCZ121l8); llha de Mar~jO (MCZ 1S663); 19-..apt·A~u ( BM1904. 7.26 8); IP'EAS. 5 lr.m E BelCm (KU 127536-391;Uliog-a, 7 km E BelCm(KU 127.s40).

GUYANA._E... , lh"''''IlTa: Vr)'heid. LuSI ( BM 78.12. 13.1 9). 1..", D~ ...~rara : Wisma r (AMS H 45774 ).Rllpll"""I: N Acara hy ~' ou nl ai n •• \Ii Sew Ri~er ( KUb9696_ 700); hud...-aler. h abu Cre~ k ( FMNH 30909);Ku)uwini Landing (AMS H 46232); MaNdl (AMNH 46570);Paraboom (AMS H 46236), .'Jazon." i-Pol4ro: Oko M<Hmlain.(FMNH 26716-21); Kan ab<:> (AMNH 1349.1.2 4915.709118_89); Kamaku. a (AMN'H 21 401)~ Cama ria Road (AMNH3'1119).

SURINAME.-A'ickt r it : Frede r ik Willem I V'~ a ll e n

( FMNH ,10907 _0 8; BM 1936,4,2 ,1-2 . 1936,9, 3,6 -8);Sipaliwini Savanne (MSH 767); Kay..,r Gcborgle (MSH 575,6", 630, 645, 657, 670); Corantijn Rivier ....ar Washaho(RMNH uncal); Po.l Tigrie, New River (.\ISH 374•.190 );Pu,t Gt>nini, Coeroeni Rivier (MSH 321. 326, 353. 394);Wilhelmi na Geborgk. linker Coppename Rivier (RMNH"nefl l; Fallawatra Rivitr (Udlh de kude 19(4), S ..rUwmt:loden'IVan"", Kamp 8 (AMS H774SO;RMNH ISOI4. lS(17);Afobida road , 10 lr.m from I'hodra tRMS H " neall , B"'....,..,..: Bro""n.borlJ (RMS H 157041; railroa.l. km 121. nearq..-ry at fOOl of Bro.. noborJ (RMS H 9716. 9719. 9723);...ilroad , km 121. upper Mah.ami KInk (RMS H " ncaO;Brokopondo (RMS H unc~l ); Gotod P1~r IRMS H " neal);T~poo"'~lra . Gran Rio t ~'SH 2631; Toek...eeeree KrcckCRMS H "neat ); .... Sarama:ca Rn.-io-r. lr.m 91.5 (BM 1946.41.231;"pp:1"Stuin.amt: Rivitr~ Sara Krcclr. (MCZ 10164,

11030). S~.: filli li bbo camp (RMS H 6215). "'....~ij~: S...... Gebor8te (RMS H 6413. 6600--01. 6603, 6637.6642-43. 6906-01. ro3 1. 7372. 87661; Paloo....... (RMSHlS022l; Mom ,., T"'I"X(RMSH 425. 534. (4 7)-

CI,arrJ and 51ai"rJ maltriol. -LAOI6098 1.

RU IJ)IE"

LIs ranas venenrn.as del genero PIt)-J/ooo1rJ (Fam iliaDendrobaudacj !>C encuemran desdc Cosa Rica hastael Peru . Boli via . )' e! Brasil. La mitad de: [a. e.pecies.c eecuemran en la rellion amazonica. Las 20 especesse coloc an en cuatro grupos taxo nomic os: gru po birolor(PhyllobUl~.f especic. P , auroto enia. P. biml"r, P ,lugubri .,', P. vi l/ cI1I1J); grupof cmoralts (P. <1/1/hCIII)'/, P.boulengeri. P. 1!.'p i", ,,,,,I. P. femora/is, P, Iricolor . P .zuparos: grupo pil'Tu.• (P. bolivianus. P . ililler l, P .parvu/us . P , petersi. P . piaus. P, pulchripa lu<. P ..<muraXdin us); )' grupo /r jv illa lus (p. b<lJ./rri, P .tr/ virrclfus).

Ame no nneme rodes las ranas dendrobatid as . indentes se C'On,idc: raban Dtndrobalts. En esse revisiona1gunas e. pec ie. que Cll"ettn de diemes ~ considcranPh)'lIobal~s . As i es que Ph)'lIobal~s 00 se disl in8t1ede DtruJrobcJlts por la presencia de dientes [aunquela mayuria de la. especes de PIt)'II0b0 1t s t iene:dien,e.).sino por el lamaOO de los discos )' la laf8ura de losdc:dos. En Ph.l'ffobo1tS . los discos rerrmn ale s de losultimos Ires dedos de I. mano son relalivamentepequefios (menos de 1.5 veces mas ancho que etdedo); d primer dedo de la mano es iguai a 0 mas largoque e[ seg undo . En Drndrobo1ts . los diso;:ostermmalesde los ullimos ues dedos de la mano son rela livamcntegrande> Cal meeos dos v«es mas aoc ho que el dodo);

d pri mer tkdo de la mano e. mas corte que e ! segundo.Las sig uienl e. especic s anterio rmente convlde radasDl!ndrobm ts se mcluycn aqu i en Ph)'lIubol l!J; bassleri,ingeri , p<1Tvulu ,l . pklU.f, y trivusasu s. Las primer.."Ires de e. las eSPl'cie s ne nen dientes . Algunm ejem­ptares de P. pictus ricncn dicntes y la mayoria carecede diemes . P, Ir/vll/lITlI.' carece de diem.:s ,

El genero Phyl/ob<l/tJ sc mannene distinto de auparienle . el genero Colostet hus , En PhyllobmtJ . e lcorso ten elanimal vivo ) es negro ron listas de un co lorsubjdo. 0 e. roratmeme de un color subido (pe r ejem plo.rojo). mienl ras q ue en C%Slr1 hus. el dorso (en elanimal vivo) liene li.la. p;ilidas 0 grises, y 00 e. de:un color s.ubido, Probablernente la pre sencia de colore!'subjdcs se n:lac iona co n la presencia de venen<.> enla piel. 0 sea que Ia pie l de: PIt)·JloIJorrs uere veneoo .) [a de ColoJttlluu no 10 liene . Hay especes quo: separeccn a am bo. generos; tal Vel 10. dos genet"O'l scjunlarin en e! futuro ,

1976 FROGS OF THE GEt·...US PHYLLOB ATES 49

AI menos dos espccic s de Phyl/obmes se uufizanpor los indigenas para envenenar los dardos que tlrande bodoqueras . Los indigenes de Ia rribu choco, quehabita n la regio n lluviosa del Choc 6 en Colombia ,traspasan e l cuerpo de la rena venenosa con una varilla,en algunos cases teniendo la rana sobrc un fuego , yburen las puntas de los dardos contra la piel de la rana.Estc procedimiento, de scr !ro per Posada (1909) yWassen ( 1957), causa tanto dolor al pobre animal quesu veneno exuda sobre la piel y se adhiere a un scrc oespiral en los dardos . Las des especies de ranas quetiencn la mala fortuna de unluarsc sc Uaman en lalcngua embena 0 empera de los ind igenes " kokoC"(Phy llobmo (lUrOllU!Ria ) y "neara' (Phylloboteshieolor ).

Las ranas del gencro Phyl/obmes son dium as, y lamayor ia habita eI suclo sombrcado de la >elva humeda.donde sc allmcntaprinci palmcnte de hormigas pequcfras.Po nen sus hucvos en hojas . Despucs de salir de losbuevos, los renacuajos sobcn la espalda de un ad ulto(al menos en algunas especies. e! adult o que llcvalos renaeuajos puede ser el padre 0 la madre). EI adultoIus llcva a una pequeria extension de agua (pm ejempto ,el agua en e l pecfolo de una hoja de palma cafda}, donde10 ' deja Los renaeuajos ccmplctan su desarroll o enel agua

En 1<1 unica especic I P. \' i l/ " I IIS) cup historia devida sc conoce bien, no hay amp lexus .

Se describcn en esta ohra cuatro especies nueva sprocedentes de Sudamerica, Ptrdtobares pete rsi, 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

AUllOl,,·UQI,'E. E.X., J .W. OUl' "'SD B, WnKOP. 1971.BatndlOto~in : c~mistry and pllannaoology. Science172:995-1002

ANDERSSON , L. G. 1906 . On battlchians from Bo livia ,Argentina. and hru, coIlecled by Erland Non1enskiOld1901-1902 and fl:ils Holmgren 1904- 1~. An. . Zool.;\(12):1- 19

1945. Batrachians from East Ecuador collectW 1937,1938 by Wm. Clarke· Mxlntyre and Rolf Blombell.Ark . ZooI. 37N 2):I - 8S

BUlKWl, T. 1905. Repfilia and Amphibia. Pages 98-1 02in O. Bangs. W. Brown, J . Thayer, and T. Barbour,The vertebrata o f Oorgona Island. Colombia Bull.Mus. Compo ZooI. Hatvan:l46 :87- 102.

1908. Some eew repfiles and amphibians. Bull. Mus.Compo Zoo!. Harvan:l51:315- 325.

1909. CllIT«tions regarding !he names of two mxntlydescribed Amphibia Salientia . Proc. BioI. Soc, W3$ll.in~on 22:89

B"'''80,UR, T . "'1<0 E.R. DUNN. ,1921, Herpetological novel­lies. Proc . BIOI. Soc. Waslllngton 34; 157- 162.

BU 80L'a . T . "'SD G.K. NOBLE. 1920. Some amphibian sfrom north...-c"em hru. ",ith a ,"i>.ion of t~ &Cllelll

Ph>HobdfU and T~Jmafobi"•. Bull. Mus. Compo Zool.Harvatd 63:3%-427.

BUBOIiR. T . ",so C.T . R"'NSDEN. 19 19. The he~IO,Io\'of Cuba. ~cm . Mus. Compo ZooI. Harvard 47:71-.1 .

BERT HOLD, A.A, 1845. Ueber ...,~lt icdene neue coer !!C heneRepilien aU. Neu- Gra.nada und Crusteceen au, CltinaNacltr. Ge•. Wiss . Gotllni:C'n 1845:37-48

BoEnwl,O, 1893. Reptilien und B.atnll;ltier _ Yme zuet.Ber. Senckenberg, Nilturforx lt. Gn. 1893:3S-42.

B<lKERM"'NS. W.C.A. 1%6 . Li"a "nolada du localidade.lip<;> de anfibios bra, ileitos . Servilro de [)ocument....ilo,sse Paulo. 183 pp .

_. 1967. Nov... t.pc-des de "P")"JlObuf~s" do Its lt e.lIde," brasildro (Anura, DendroNlidae). Rev. BrasilBioI. 27:349- 353.

Bo L"LE"'Gu , G.A. 1882. Catalogue of the Batrachia Salicnlias , Ecaudata in lhe colleclion of lhe Brilisb Muoeum.2nd ed . British Mu.cu m. London. xvi + 'so3 pp.1883. On a collection of frog~ from Ye rtmag uas, Hua.

111:1;:6~~";;3:o~m Peru. Proc. ZooI. Soc. London

1888a. l'o'oce on the cl~siflCat ion of the Ranidae . Proc .zcor. Soc. London 1888:204-206.

1888b. On the ··nur;in i '·hbits of D~,.J'obufn , uoMen ed by A. Kapp ler . Ann . Mae . Nalu r. Hisl .,Str .6. 1:454..0455.

1898. An account of the rep il" and balrachian. cot.leered by Mr. W.F.H . Ro..enberg in wenem EcuadOl'.Proc. Zoo!' Soc. London 1898:107-126.

1899a. Description. of fit\\' batrxhians in lhe collectionof lhe British Museum (Nalural History). Ann. MagNalur. Hist.. Ser. 7. 3:273-277.

I899b . De.cript:ion. of ne.... replik. and balrac hianscollected by Mf . P.O. Simons in the Andc. of Ecuador.Ann, Mag.l'o'alur . Hbt.. SCI" . 7. 4:454-451.

:~~~:'o7;~'a~~I:::r~~~l~~d ~~~~~.J~:Ser. 7. 10:394-402.

1903. On some balrachians and reptile> from VenezuelaAnn. Mag, l"alUr. Hisl. , Ser. 7, I I :4HI-484

1912. Descriplion. of fit '" Nlrachians from the Andesof South Amefita. preserved in the British Mu' eum.Ann . Mag. Natur. Hist., Scr. 8. 10:185_191

1913. On a collection of batnll;hians and repciln INdcby Dr. H.G.F. Spum:II, f .Z.S., in the Cboco, ColombiaProc. Zoo!. Soc , London 1913:1026-1038.

1918. De"Cripcion. of rIC'" Soulh.American Nl rxltians.Ann. Mag. Natur. Hi.. ., Str . 9, 2:427..0433.

_. 1919. De"Criptions of I,", new lizards and a fit .... frogfrom lhe Andes of Colombi a. Proc. Zool. Soc. London1919:79-81.

BUSSEU-. P. 1973. Vocaliz.alions in the territorial behav iorof the froglhttd,.~. """,il ia . Copeia 1973:277- 284.

CH"'PM"',,". f,M . 1926, The di"ribulion of bird·li fe in Ecuador.Bull. Amer, Mus. Natur. Hi!>! . 55:1_784.

COCHUS, O.M . I95S. Frop of southeastern Brazil. Bull.U.S. Nat. Mus. 206:1..0423-

_ . ~':' ~ 'l';;~i(f~~rc~i;~~.% ~~~ world. Doubleday and

COCH....S. O.M. ",so c.J . (joIN . 1964 . Description of a eewfrog,of the r:enus P/ryJJoba~. from Colombia (Amphibia,Ranidac. Dendrobaunac), Stnckenherg Bioi. 45 :2.~5­

257 ,

COCHIt"'S. D.M. ",s o C.J. GotN. 1970. The frog-s of Colom·bia . Bull. U.S. Sal . Mu•. 288:1--tl55 .

COPf., E.D. (1861) 1860. De>eripfion. of rep llto from tropicalAmerica and Asia , Proc. Acad. Nalur. Sci, Pbiladelphia12:368-374 .

1862. On "OInt ne...· and litlle kllO...n Atneric""l AnUlli .Proc: . Aead. Natur. Sci. Philadelphia 14:151- 159.

1863 , On h at;hy"ph" I". , Scophiop us, and otherAmerican Barrachia. Proc. Acad . S atur. Sci. Philadelphia15:43-54.

1866. Fourth comribulion 10 the herpelO108Y of tropic,",Amer ica: 3, Notes on neotropical batracllian•. Pnx: ,Acad. Natur. Sci. Philadelpltia 18:129- 132.

1868. An namination of lhe reptilia and twra:; ltia ce­tained by lhe Orton npedition to Ec,wlll' and lilt upperAmazon.... ·ith notes 011 « hef spede•. Proc . Acad. SatulSci. Phlladelpltia 20:96_ 140,

1874 , On some Batracltia and SemlIOgnathi brough tfrom the upper Amuon by Profe""'f On on . Proc.Aead . !"alur. Sci. Philadelphia 26:120- 137.

1875. On lhe Balrachia and Reptilia of Costa RicaJ . Acad . I"atur. Sci. Philadelphia. Set. 2. 8:93- 154

18n. Tenth contribution 10 the: herpetology of tropicalAmerica. Proc. Amer. Phil. Soc, 11:85-98

1886 . Catalogue of lite specits of batrac hians and~ptiles contained in a collecl ion made at ~bas. upperAmaron . by John Hau•...-ell. Proc. Amer. Phil. Soc .23:94-103.

1887. Syn0p'b of the Balfacbia and Reptilia oblainedby H,H. Smillt. in the pnw ioce of MalO Grosso, BrIl i!.Proc. Areer . Phil. Soc. 24;44-60.

1893. Second addition 10 t~ kl'lO'll'ledlJ:C of the Batrachiaand Reptili a of CO,l a Rica, Proc. Amer. Phil. Soc .31:333_347.

1899. Conlribuliom 10 lite hc-rpetoloJy of New Gr3nalband Argentina. Philadelphia Mus. Sci. Bull. 1:1-26

CM UM I', M.L. 1972. Terruo nahty and muting behavior inDe'ld'obul~S It' O'lUJifN IlS (Anura: Dendrobatidae), Her­pelOlogica n :19S_198

poca. J.W. ",so P. DU....S T. 1974. Coun .hi p beh....jor inColosfelh"s m/la,;. (Dendrobalidae) . Copeia 1974:988_990.

/976 FROGS OF THE ces..US PHYLLO BATES 51

DoSO Sl>-S.. ...OS. R. 1964 A "" '"' Dl:nd robal idx froa:.P'o.u~,,,pfs ,ivrroe fro m V"!lC'z~bI . Carib . J. Sc:i.4:485-489.

DUU~.~~~mw~~ior in dendrobalid

Du",ilIL, A.· M.·C . .."0 G. Bl l ION. lSI!. Erpe toloJi egenera le ou hisloi re n".lurelle com plele de. repi le. ,Volume 8. Libnline encyclo pedique de Rom, Paris792 pp .

DUNN. E.R . 1924 . Some Panarn..nian Frogs. Dec... Pap .Mus. Zoo!. Unlv. MIChigan 151:1-17.

1931. New fm,s fronl Panama and Costa Rica. Occ....Pap. Bm ton Soc . Natur. Hist , 5:385-401.

1933, Amphibians and replile. from EI V.lle de Anton.Panama, Dec.....Pap , Bo ston Soc . Natur. HiS!. 8:65- 79.

1941. :-';OC.,s on DtNlrnbo'u '''''", ,,s. Copeia 1941:88_93 .

1944 . Notes in Colo mbun herpe to lon . 2. C.ldasi.a2:391~.

1957 . l' COI:ropic.1 fro, ~ner...: PrOJ'Iw,,,pi~ venusHd o=l..... ,""Itt rcman-.. on PIr.Y1lobtlIU . Cope'" 1<r.l7;n _78.

DU"''' T. P. .."0 J.W . 0 0 1.1:. 1915. Aggreu;"" bchzvlor inColosl~t"'u ( - p,.mlw,apfs) nJll"ns (An ur. :Drndro.balidac). Hcrpe toloa:ica 31:23-26.

FtTZISGEI. L.J . 1860 . Die Au.beule dcr o.lCrrcic: hi....henN.lurfO~her an Sli u~thieren uoo Replilien wlihrcOOder Weltu m""g clu n,. Sr . M.je.lal Fre gaue Nova ra .Sitzungsber. Akld, WI••. W'en 42:383-4 16.

FoW1H, H, 1913. Amphibians and ll.'plile> Irem Ecuador,Venezuela , and Yuc atan. Proc. Acad. Natur . Sci.Philadelphia 65 :15.1-176

FUNUIO USEI. J .W. 1956. New frog' fro m Ecuad or and>oulh...'eslern Colo mbi., Zoologica 41:73- 80.

G ..N'. C. 1960, NOle. on a herpetol~kal ro llecl inii lripthrough dte !IOulhe:acern lowlands of Bolivl• . Annal.Carne gie M ..... 35:283_3 14.

G...",.. s , S. 1837 . We>! IOOi. bauachia in the MlI'CUm ofComparal ....., Zoology . Bull. Esse x lnsl. 19:13- 16,

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!be U.S. N. A<lronollnc a! E.1.pednion 10 CI\lII. Proc.""al. ~l.Iur. Sci. Pltilale lphia. 7:226-227 .

Cio:Jor>M...... D.E . 1971. TerTilOrial behavior in • ne«ropicalfrog . Dmd1"Obalu K"'" ../ifrnu. Copei.a 197 1:365_370.

Gos."~~b~Y~~ ~~·la-;.,:;m~,li;~edn~~~e :~r i~~~;~li c:ror:.Herpetologica 16:183-1 90.

GUI8E, J . 1950. Cal,alo~ue de. I}P::' d 'arnphibiens du Mu...eumNalll'nal d' HIMOlre Nawrette. Imprimer ie Natlonal e.Pari" 71 pp.

GUln'Eso T, A. 11148. ,faun a chilena:, rcplite•. Pa,., . 5 - 136'" C. Gay, HMon a fislCa y politlCa de Chile , Zoolo,ia,volume 2. In the aUlhor'. house . Pari. , and Mw.ro deHistoria Natural de Samiago, Chile.

GUNTHH, A.C. 18~ 8. Car.alo,ue of the Bauach i. Salienli.ain lhe:coI1ec1ion of the Brilish M"""um. Brilislt M...",um.lnndon. _vi + 160 pp.1868. F...... acroun l of specie> of l. ilIe•• bamc hian,addocl 10 the collection of the Brili:$b Mu.eu m. Prnc .Zool- Soc. London 1868:4711-490.

~\lO~ ~~en~~B:::~~~~.Taylor and Franc i•• U>ndon.

HUTWOl-E. H.. H, 501""'0 .."0 A. HE"no-UU . 1965. NOle.on amphibian. fm m Ille Venezue lan Guayana. ....ilh

des<:riplion of 1"'0 ne", fo rms . ACI. Bio i. Vene z.4;349 _364 .

HU utK"H , W. 1940 . Beilri~ zur Kennmi, der Gattun,H ylou llu ( Br~h}eep/l.lidx , Amp/l . ). Zoo!. An• .131:11l-128

HooGWOED. M.S . 1969. !"Oles on lhe herpelOfaunaof Surinam :3. A ne'" . pedes of Dt.w.ro1J<Jt~J (Amphibia Salienti. ,Dr OOrobat idliC) from Surin~ , Zool, Meded . k iden44:133_ 141.

1971. Dt.w.roba/~S , eine farl:>enre iche: Ganung. Aqu,.,..Terrar . Z. 24:1_7.

1972, Fro, ch<: . die ihre Kaulquappen huckepack tragen.Aquaricn M. g. 1972:28R-293.

I"'STlTUTO R ~C ION" 1 DE DUENV0 1VIMHlTO DO AMA' '''( IRDA) "NO COl<SHHO N..CtONAL DE GEOGI.. ft..URG E) , 1966 . A,las do Amapa . Rio de Jane iro. 40 pp .

JI",i"u. Dt: 1" E.S' AO.. , M. 18m . Faunae llCOIropicaiis 'J'CCiesq....edam noOOum cogni l3e'. J. Sc:i. Ml.lh. Phys . Nalu.Acad . Real Sci. Usboa 3:57-68.

1815. Venebradm de l viaje aI PacifICO veriflcado de1862 • 1865 por una romisiOn de ll3lunlisas enviadapor el gobierroo cop;lilol . B.l racios . Mig~1 Gine>t• •Madrid

K..YAu" S,O. , E.X. AuuQvuQI)!. Ao" O J.E . WU SJ('k1970. GaPglionic and eardi"" ""lion. of b.alracholoxin.Eu"'P"" J. Pharmacol . 12;10-18.

KEFUSTf-IN, W. 1867. Ueber ., inige neue ode. seneneBalf~h ier aus Au>tralicn und cem tropischen America.N""hr , Ges. Wi... GOuin,.,n 1867:34 1_.361.

KlT..SAo ...O, J ,T . 1967 . Ob'e rvali ons on the biology ofDm ,I", "'" ,. pumiiiu Schmidt and D~"Jrt>""I" " ..raill.'Giranl . M.S , Thesis. University of Southern Cal ifornia.87 pp

L." THAM, M. 1966. Capluring ".lIange creature. in ColombiaNil . Geogr. Mag. 129:683-693.

L.. URENT, R.. 1942. NOle su r Ie. procoellen. flrmisterne .I BaTr.eh 'a Anllt ...) . Bull . Mus . Roy. Hlsl. Na lur .Belgique 18(4l ):I _20 .

L.lDTH ee JUlDf., T . W. V..N, 1904 . Rep:i.... and balrachian.from Surinam. SoIe:s from the LeydenM!I'eUm2.5:83-94

L.VOOECU , H 1974 . Etholo, ische Unlersuc hungen zurFonpflanlUn, von P lryl/ob<r,rj pal_I.... (Amphibi....Ranidae ). Ph .D. Dissert ation. Johanne. GUlCnberg.Un_n.... zu Mainz. (Abstrae'l)

L.UUSG. K.H . 1971. De. F3rberfrmc h P1n~/ob"'~J bicol<>rBibron der O;"d illera Azul (Pt-11l ) . BOnner Zool. Beilr22:16 1_ 174.

Lurz , A , 1925. Balr~icn. du Bri .il. Compl . R.,nd. Soc .BioI. Pari, 9.3:137- 139

LUTZ, B. 1952, Frogs in the Adolpha LUlz collec tion ofthe Instiluto O,waldo Cruz. Mem. In.,1. as...·aldo Cruz50:Ml8-613

LUTZ, B. "NO G , KLOSS. 19~2 . Short note. on some frop

~%: , t::" ~'&~a~C:: ~ng~_~~, vicariant forms.

M.."". W.M , 1957 . TIle wild animals in my life. !'Oat. GrogrMa, . 111:497-524

MAnl, F. .."0 B. WlTl>or. 1963. TIle wcnom of lhe CoIom,bian arrow poison frog PIr, I/obalrs lHcoJor. E.1.perienlill19:329-376.

MEU" . D. 1,941. Con lributio n. 10 the kIlO. ledge of lheAm~ubi...o f Sourh Amenu. GOIel>or~KWlgl. vee....k.Vincrll . Samh. Hand!. ser. B, 1(4):1_71

MIR..ND.. Rt Rt.llO , A . DE. 1926 . Nora. p. ra ",rvircm ao.,..ud<> """ gymnol:>.amochim (Anura) h"a.i tem:... ArchM... . 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~ Ke und ZUd ll einn Blau' ICi jlu ,fTDS(:hes de. Oal1un~ P")'lIob<lr~J .us &:uador. Sal••mandra' :81_&4.

NOBU. G ,K . 1921. FM: new o.pccies of Salienlia from SouthAmcrica. Amer . M",", S O'l'iI. 29:1_7.

19231. Ne.... b.lracllians from the: trop ical resear ch.rarion. Brili .h Guian• . Zoologic. 3:289-299

1923b. A new gekkOIlidliurd ...d a new bra<.: hyc<:phl.lidfro, from Colombi•. Amer _Mu• . Novit . 88:1_3.

1926. The peo;1OnI 8in1~ of the bncb)'Cephalid hop.Amer . MU$, Nom. 23Q;1-14

OlItTT~~':::)I '::U'::~:::r?r.'f~:r~,,:, (D... Erdbcer._ . 19S3 . Nechmals Dr ndrubaUJ rypogr<lpltic",,: d• • Erd·

beerfroschchen . Aquai' . Terrar . Z . 6:260 -2b2.

OEsE. , R . 1932. ~ ZUehl des B.I,l!mlcige" (lHndrobd,ul~iIUJ. BUillCf Aqu• . Tmarimlunde 4) ;196-200.

PUUI. H.W. 1935. The frop. Iiurds. IIld snal<n of Brili>llGuiana. Pnx. Zool. 50;. London 1935:505-530

Pu...oc .... M,G . 1904. Rcuili cd Amfibii, Boll. Mu•. Zoot.Anal. Compo Univ. Torino 19(465):1-4].

Puu s, W. 1870. Eine Milt!leilunJ ube. neue Amphihien(Hr"' idIXryl... . . . . ! des ko'''C bch wolol;~ben Mu.lOe~. M<XuL\her. KOni,1. Pl'cu..... Aka:!. Wi... Berlin1171):641 -652.

_ . [1873)1 872. Oberdie _ SpU in Bnsilien cnammeltmBalrachier <in kooil;l. N'l uralienhbinelS ZU MunchenMOl1 alsber. Konigl. Pn:u... Ahd. Wi... Berlin 1872:196_227 .

118741 1873. Ober ew neue SchildloJiitawt. CillO"""'''''''ElJrldlii und ein igc andere neue:oder ...,niF bek_Arnphibien. Morw .be r. KOnig!. Pn:u"'. Aka:!. Wi...Berlin 18"13:60 3-618

PoLDU . W.S . 1973 . Over ~erzorxin, en ~OO"planlint in

~;;:~g~~dh:'~a~~~e~H:~~;~~i~':Z:;:~'::_2e2" en ele

1974a. O~r ~erzorl;inll en ~OOr1plan'in , ill ge~.n .

&,~:: glndH~;~:""':: Ie;::,:~~ anden:

197"b . Over ~rzor,in, en ~oorrplanli n. in gevan .

~'::~!~ fJl~H::'"'~~':;'';,,''':: 3e;4~)~.1e .ndere

197":. ~ndrobalidae (4) . Ch-er ~erzorging en ~O<)".

plannn g In l;e~angen sch.p ~an Drndr ob<J'" al ll' rli.en enkde andere Drndrubalid~ . He!Aquarium 4S:122-

'"1976. lHNJ,obatr• • Pllyllobalr . en CoItnl rll.... . He!Aquariwn 46:260-266 .

Pos'o~'ou 1~~~B~il.l~ : ::eriZi'e~a:,,:,"';!~~o~~fr·1909, EJvere no de rana de los indio. del CItocO. Pages78-88 in C.A, Molina. ed. Esludios cie nlificos de ldoc1o. And~. Po!<ada con .I ,unoo Olro$ escrilOSsu)'OSsobrc d i~en.. lem• • . Imprenl. Of", ia!. Medellin.Colombia

Rlvuo . J .A. 196 1. Salienli. of VenezlIl:l•. Bull. M... . Comp o7.001. Harvaro 126:1- 207.

RUTHVE"', A .G ,AND H.T. GAIGE. 19l5 . The breedi ngh.bi" of Pro..,ht,api.....bp ..ncl"I ... Cope . Occ...Pap . MIlS. Zoot Univ. Michi, ... IO:I- S.

SAGaA, R. l>E u . 1&41- 1843. Hisioirr physique. polilique.ee Mlurelle de rile de Cuba. Alias. ~plile• . Bertrand .Pm., 31 plales

S~VAOE , J . M. 1968 . T ile dendrobatid fro~ s of CenlralAmerica. Copei. 1968;745_776.

Sr;HLUJE!- . H. 1826,: NOTice sur l' e.pelolo, ie de rue deJava: parM. Bo,e(0II"f~ marluscnl). Bun. Sci . Salw .Gcol . 9:233-240.

SO<"IDT. K.P. 1932. Replib~ amphibian> of the MII'IdcIVenezuelan nped"'"". F.dd MIlS. Nll ur. HUI., Zoo!.18:1S9_ 163.

SCH" IDT., K.P. A~lI R,F . [NOU . 19S1. Ampllibians andreptile s of the Hopkin ·Br . nner e ~pedltion Eo Brazrl.Fieldiana: Zool. 31:439-465.

SO<l4ll1T,O. 18S7. DiaJllO!ien,ne,," Ft-i\..:hedes nll>loJiKbenCabinets !II Krak.u. SlIlun r ber . Akad. Wi••. Wien24; IQ-U.

_. US8 ~licj;)C herpclOlo,icae mu-,.ei zoolOlici craco­viensi. : un,escll"'iinzlen Balrachiel . ~nkschf. Akad.wss. Wien 14:237-258

Sf!' .FT, W. 1936. Da> Bru!gn cllaft des Baum'lei~erfrosche.IDrndrobatrs. " ,.,al ... Girard ) in Gefangenschaft. Zool.GIIU1l Lelpll' 8:122_ 131.

_. 1939. Urn da ZUelllproblem ....... lHttti rolKJlr. " ",aI...G. Wochenschrift AqIW". Temrienblnde 36:7_9. 23-21.

S!XTO~ . Q,J , 1960. Some a'~·l' of the behavin. a nd oflhe lerTLl0 'Y of a del>ilrobattdfrog. Prn."ltt,,,pi.. ,,mUdlJSEcology 41;107-1 IS

Sll.VEIlSro"'E. P.A. 1973. Ob",rv ll ions on the behilvior and

~J:,,=,:..~~~~~~:a"'~29'1-301.

1975. A revision of lhe poi!l(l/l-aml\V fro, s of the genuslH nd' oltale.< Wagler. Natur. Hisl. Mus. u-.. AngelesCo" Sci . Bull. 2 1:1_55.

SI'IJI. J .B. VON. 1824 Specie. nov~ R..... rwn 'lUli in ilwre

:~::":~2~~ili::ll!'~'~"~~~si~~coIle, i! ee~rip>iI . Munich. 29 pp.

STU/ll~ S. R. A~D J. HE"'OUC" 50I<. 1919. Field .. udies ofamphibians in Colombia, 500111 America. uev. Cali·fornia Pub!. Zool. S6:497_S40.

ST F.lNDAr;H~u . F, 1864 . Balrachologi sche Millheilungen.Verb. 7..001 . BoI. W . Wien 14:239- 2118 .

1869 . Reise de.- o.1erreichi"c1oen Frepne Nov...... urndie Etde in den Jahren 18S7, 1858. 18S9. Zoolo~scberTheil. leo Bmd , 4:Amphibim . K.·K . Hof· und Sl....·druckere i. Wien

TAYLO Il, E.H, 19S2 , A review of the frogs and toads ofeosla Rica, Univ. Kan.... Sci. Bull. 3S:S77- 942

T EST, F.H . 19S4. Social alll re:..ivcnes. in an .mphibi... .sceeee 120:140-1 "1.19S6. Two _ dendrobalid frogs from nonhem Vene.zuel•. Occas. Pap. Mus, Zool. Univ. Michi,an S77;,-,1962, The lIi, hly developed counslli p of rhe frog.P,mllttropi.. /rinilll/u. Amer, Z001. 2:452. (AbslracTj

TQI;UY 'o"'A. T. , J. DA~Y ~ "'11 B. WnKoP. 1969. The """'lUre:of bau.d.olo~in. a Slel'OidaI alk.k>id from the Colom·bian IITO\V poi""" frog. Phyllobalr. ""'....ar"... . andparrial »T1Ibe.i. of b;mxholo~in and ilS .nalop and""""""". J. Arner. Chern. Soc. 9 1;3931- 39311 .

T!K'HUlIl, J .J . Vo "'. 1838, C las. ific ' lion der Balracllier .Pe!ilpiem: . Neucllatel. 99 pp.

W~Gl.n.. J . 1830. Nalijrliche. Sy.. em dc. Ampllibien miT

mrCll~,;,,~~~~~~iiu"hiere ond VO~1

W"'DE NIEI . R.l .A. 1973 . Venl., van bel kweken mellHnd,oba,tJ ,,"'Qt..... L1«rta 31:167- 171.

WASSEl<, S. H, 19~7 . On Drnd,,,""u $.f.o, ..po i ,,~n materialamong Emper.ii (ChoOOj" l"'aklllg IndIans III western

/ 976 FROGS OF t HE GE."'US PHYLLO BATES 53

CltJda~ . Col omb ia. Arm GOlebor,~ Elno" , Mu•.I9SS- 1956:73_94

WlI. "U. F. 18W . Ueh:rRcpilien unciBltlnChier -. Colum·biotn lIl'Id Trinidltd. Vem. ZooL BoI. 0.. •. Ww:n 49:,"'-'"1916 . s"mcrhn,en iibe. eini,e nieden: Wirl>ellien:de. Anden von Kolumbie n mi, Be-.<:hreit>un~n neue,Anen. Zool. Anz. 17:301_304

W' EI>-S n.-"",W . .\I A Zo o 1821. Rei", na: h Bfa'i liell inden 1;Oh~lI ISI S bi. 18 17. "Olume 2. Heinrich Brommer.F.-Hun . >vi;; .;. 3-16 PI'

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)CI dt-..:ri plion de ""pl llOlfVel~ e~~. Pa,e> 21 3_2."10ill Une mi~,ioo bioIojtiqllC beJjlC' aU Brbi!. "Olume 2Bru>e l lc~

Accepted fur pcblic.uion Apr;1 1.\. 1976,