Silverstone, 1976

A REVISION OF THE POISON-ARROW FROGS
OF THE GENUS PHYLLOBATES BIBRON IN SAGRA
(FAMILY DENDROBATIDAE)
By PHILIP A. SILVERSTOi'E
NATURAL HISTORY
M US EU ~l
SCIEl'CE BULLETI!"
~7
OF LOS ANGELES COUNTY
• :-,:ovEMBER 25. 197b
A REVISIO:\, OF THE POISO:\'-ARRO\\' FROGS
OF THE GE:\' US PHYUOBATES BIBRO:-': IS SAGRA
( FA ~ lI lY DE:-':DROBATIDAE)
A REVISION OF THE POISON -ARROW FROGS
OF THE GENUS PHYLLOBATES BIBRON IN SAGRA
(FAMILY DENDROBATIDAE)
By PHILIP A. SILVERSTO NE
:-<ATU RAL HISTO RY
:\1U SElJ~l
OF LOS AV iELES COUI\TY
SCIE!\C E BULLETl I\ 17 • :-;OVBlB ER
~5.
1976
T AB LE OF CON TE NTS
AC,,"OWLEDGMESTS
~h T UI" LS ""D ~l ET HOOS
~ u-e um
abbrevialion$
PNlUJJA<1a BIB_
Ddinilion
,., S..u...1841
Di a~ nosi s
Descripelon
Conten t
Tad poles
Specie, group'
Life histo ry
Food
Summary of distribution
Key 10 the genera of Dendrobatidac
Key 10 the species o f Ph.l'lIoh/ll'-f
8 K."OW~ (ilt 0\Jl'
Phylloha tt!_, ,pec ies
PhylloM tt!s aurorut!nio
Phyllooott!Sbiro /or
Phylloha lt!S lugubri s
Phy/f oha 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 ralis
Phyllobau s tricohv
Ph,.llobau s zapar o
P O f!S (; 10l/P'
Phyllobat t!s bol Mu lllU
Phyllo oolt!S ingt!r;
Phyllohult' J pan'ulus
Phyllobal t!S pet ers!
Phyllohules plctu s
Phvltobates pulrhripeclu .f
PhyIJob/lles ,marulld;nus
T~lVmA I/JS (; IO\; P
Phyllobalt!s bault! ri
Ph.l'lIobalt!s triviu Ol,.s
U TElt U l,I RE CnE D
3
J
J
J
J
,,
,
7
7
9
9
9
20
20
21
2J
24
zs
27
27
aa
29
50
J2
JJ
as
""
J6
J7
aa
4J
44
"
"
"
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 .arm w 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 IP
pulchri~..,,,.,,/ . hololype. Scrra do J\avio . Brasil. 1'01 to ",.Ie. Painling by J. K lkanishi. staff an i' t. LACM
A RE VISlOS O F THE POISOS .A RROW FROGS O F THE GE:-iUS
C FA~lI L Y DESDROBATlDAE )'
PHYUOBATES BIBRON IX SAGRA
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' and
PII.d ' obu,<'J (dorsum bri,h ll) rolorcd and ..lein lO~irl, I'f",,,mabl}'
maintained. Fing.... Icn~'1h and
finj:C1" di'k size ~ 1I>Cd hc~ in 10di...in,"i"" ~lIJrob<l,... (fiN finlCT <hnft.... than 'ottOI>d. ""d fin' ....di.h
I'dal;"ely larj:C ) from PII) /Iob<1l<" Cfir>! finl!tt equal lO or IonI'C'" lhan >«und . and finrcr 4i<J;s n:1", r.d~
......11) in'lead of lhe uadilional chanc ier. absence lD..nJ,obu,<'J\ or pn:".'~ /.P."II,>b.I,... )of 1<'C lh. allhough
lhe IooCh charac1C1" i, ill u",f"l. U,in, finger k nglh and finger di..... 'ize a' a dcfinilion. fl\'e , pecic' (b<Jul..ri .
;"1("';. fJ<I" ,.. IIU, pic, and ";vi fl"''' ,') Iran ~fc ' from D..nJ,obu,<'J10 PIr,·lIobul... .
, pecic, It<> be oJe".;ribed b)
P. ) lIohal<" include' four , pecic, gr<>l1l"': (1) bi,'mo, group (Ph.•
C. W. My'cn and J, 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","'" " , and
CoIOlI<'ln"s . Phyll" l>aln u,ually ""ip<"il' Cal lea'l in """ 'pecic•• and I'l'ohahly in all ,pecic' . WilhoUi
amplc:1lud on lea"e,. Aflel' the ladp<' le, halch, a parenl , ,,,ric, IMm 'lIl il. had and rel, a"', lhem in waleI'.
whc~ Ihc ladpole, compkle thei, ,je"elnpment. In al lea... .........e , pecic" ,ilhe r the mak 1"''''11'1 01'lhe km alc
parenl may carry lhe ladpole, . Tl>c main food of Ph l/obm... i' 111>1, .
Ame, indians e ~ ltact blo....gun poi"'" from al k a" 1"'0 'P'""ic. (P. "" "'I",n;,, and P. mcolo, ).
P"'''''''' ..
H,,,,,,,...
I S 1 RO[)l",'C1IOS
A, p""viou,l y d i-.:u....-.ed (Silver'>lone 1975 ). lkrldru·
batjd 1Cnera are difficult to de fine . The ~enu, Ph).·//t>·
previou sl j- ..."" d ia~ n"""d by tbe prese nce o f
bright oo l"1>;ll co lorat ion and the pre-e nce of maxillary
a nd prema xillary teet h (Savage 1%8) . Because the
prese nce o f teeth "arie s 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 ge nu' nil the h'lSi. of
finger dis k size and the relative length of the first and
>C~" lIld finge rs. These finger c haracte 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 )'
lac k teeth) , ....h ich here are co nside red Phy //" rnll" s
rathe r than D.."drvbal es, I have tra nsferred t Selve r" one 19 75) thre-e addit ional spece-, (bun/rri. i" x.., i.
and /'<In ',,lu .l. ...·hich have teeth) 10 P.~-Jlobul..S on the
N , i, of both teeth and finger c harac ten.. I ha..... con,idc red a' CoIosI..,hus aU dull y colored toothed _pecic,.
although , i, species ( the [ .. _ ".ali. grnup) ...ith "right
co loratio n and basal toe lI'e bb ing . included he~ in
Ph...//oI><I,..s. rna)' be <If> do~ly retated to CoI<lsl..,hws
"" the)' a~ to the Ol:her ..peo:ics o f Ph,-lf obtJ' ..J .
"'u...
Th u, Ih i ~ revi,ion of Ph\'l/ol>a,..s include. tho se
specie - havin ~ a oombil'liltion of three c hariICten.: ( 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 partl y or e ntirely
brighlly co lored in life. In my op inion . the ~'(: nu~ Ihu\
de fined is mere natural than il ·...·a' unde r the prev iuu ~
definition . Ho we ver . delineat ion "f convi ncingly
munu phy lc lic ge ne ra of dc nd rob atid s will req u ire
co n, 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. Tr ux<l l, D . Wh i,c le r,
R.R . Snelling ( LACM ); J. M. Savage . P.H. Slar reu
ICRE): A ,H, Brame . Jr . II. McCu rd)' Xature Ce nter:
' EoI 10 10 U CO Mll rTTH FO. IHIS B ULU TI"
R OIlH T L. B .... y
W. RO" " I D Hn u
Roy W.
~k Ol """lD
WOl-'U S . PoL!».
JoH" W , W . I" HI
'RC'C~h
Auoc tolC in Ht'l"'l<Jlogy. :O>alutal HiSl<X) "'\/>Cum
of l.oo An",1n Counly. 900 E~ poJoilion 8 1vd .. ~ Angeln-.
Calif<>t1'l ;a 9(0)1 .
2
BULLETI N OF THE NATURAL HISTOR Y MUSEUM OF LOS ANGELES COUNTY
J.P. Bog art (l NHC ); C . F. Wa lke r. A.G . Kluge
CUM M Z ); R.G. Z.. -eitel. C .W. M yers ( 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. McDi armid. Univcr.;;I)' o f South flori da:
E.E . W ill iam ~. B. S hreve ( MeL); \\' .E. DI.IC'Jl man .
S.R. Echurds. 1.T . Collins (KU); C.l . McCoy. Jr. •
S . Ric hmond (eM); J.D . l ynch. University of Nc·
bravka; A,C . Thoresen , Andrews University; D, Travis.
Manhallan Beach: D.B. Wake . R.C. Stebbins (M VZ):
D.B . Ekken s. Lorna Lind a Un ivcNily: A . SIam:II .
Ca lirorni.. Sta te Vnwe n;!y at r.;or1 hr id ~e : M. Tye .
California Stale Un;w,rsity at Long Beach: R,f . IngeT,
H , M arx ( FM N H): J.A . Rivero , Un iversidad de Puerto
Rico , Maya guez: Inslil ulO Ev.mdro Chagas. Betem ;
Induslr;a e Comercio de M inerios. Serra do S avio:
R. Pradinaud and Dr. Rivieeez, Ca)'l"Tlne: TeKas InSll'\lmenls Co .: K . Adler. Un iversity of Noire D-a me; O.R .
da Cunha ( MPEG); W.C.A . Bokerm ann. Fundal;an
Parque Zon l,igiC<l de San Paulo ; MOMieur Ie Prd el de
Ia GUy'ane; C. serie and ....aff. Instilul p.,,<-Ieur de Ia
Gu)'ane Frarlf;ai<,('; Office de la Recherche Scienlifique
et Technique OUlre-Mer; Rdo . Hno. Nicetom Maria .
Rd" . Hno . Daniel (IL'\ ); F.e. Lehmann V.. Musco
Dep artam en la1 de Hi slllri a Natula l, Ca li ; K. v" n
Sneide rn. Mu"Co de Hi'l nr ia Natur al. Universidad
del Caec a . Popa) in ; C. Esco har. Cali; J.I. 8<lnero
H.• 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.• l nvtiturc de Desarrollo de los
Recuoos 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 . Lesc ure, J. Guire tMHNP); M.S
Hoo j!moe d ( RMN H); W.N . Pold e r . Alkma ar; B.
Hude ndid ( NMG); S ,H. W"-s ..cn. GOteborlls Etn(}gra fi' ka Mu-ee t; A .G .C . Grandison . E.S . Arno ld
( 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 Alto
de l Buey . J. Le-c ure pru vided a Ii.... of Ioca lil~ for
1.G 'JlCCimens . inciudinll ihe flf'! record of P h.\ # "ool<,J
p;rIUJ from mnch Gui..na . ......G .e. Gr.mdi....o J'N' idcd
coord inates for the tocalnies o f P . !>"Ii,·;UI'''.'. P. Br"me
drew 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:u m of Los
Anj!e les Co" nty. Pa intinj!' for the fmotispic<:es were
based on color slides by J.A. Pelcrs. R.W . McD iarmid.
J.D. Lynch . D.B . W..ke , A.C . Thore -e n. a nd me.
SlUdie- were p;>n iall)' suppon ed by a NSF Summer
hl lowship for GrlOdUalC Teac hin!! A",s i" an" (1966) .
by a NDEA Tille I V Felluw,h ip ( 19M· I%9 j, try Bio·
medical Sciences Support Gran t FR·070 12·02 # 570J
No . 17
from lhe :-.ralional Institules of Heallh to the Universily
of Southem California ( 1969) . and try lhe Los An~le,
CoulMy ~ use:um of r\at u... 1 Hislory Foundation ( 191 1).
M II Tf ~l lI t .s liND Mf.THOUS
All nlCil-surement, are 10 the nearest one·half millimelCr.
TlOdpole lIlages arc those of Gosncr (1960).
O' ioolo!!- ical , pecimen, were cleared wilh glycerin
and lwo percenl pola"ium hyd ro~ i de and were stained
wilh 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 i ne headinj! "stomach
conlenls. " 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 of
a ta xon from the names o f a ll authors CXCC pl the
ori!!inal aUlhor.
Under the headin!!- " doublful records" are localilie1.
that are separated by a major ~graph ic barrier from
all other rol leclinj! locali l~ for the specie«. or thai
lie far outvidc thc known e1evatio nal ra nge o f the
species . The<oc doubtful records "rc not included On
lhe 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
Sub species arc not used . bUI I have indicated to
whic h popt.a lation' subspccific names apply. because:
they are available as , peciflCor o;ubspcciflCep ilhets.
It is difflCUl1 10 define species of Phylloball"J beca use of intra.s pecific varialion . interspecific resem blance . and a paucity of kn"wn cases o f eympatry. To
define spec ics. I used pre'encc or absence of the follo.. in!!- char.JClers; toe .. ebbin g. Icclh. a calf·"P'>(. a
median do f'il l ...ripe. p;lirell ...npe s , and do rsa l or
ventral marbling: I aho used positio n and Icn!!-Ih o f
pairell ' tripes. snout -vent le ngth. , kin texture . and
color. When so me o f these cru.racters wcre not shared
by allop<llric populatioos . I Irnldc: <kcision' .." 10 con,peciflCily by wei!!- hling cert ain C....rac1C1'S lhal J co n. ille re d imponanl in Ihat panic ular case . bas ing
dec i, ion, on the known ran g.c "f variahili ty of the
cha racter in both popt.a lal io ns. and on thc pre-e cce
or "bsenc e uf morphol.>t: ically intermediate populalions . I h"ve bee n co nse rvaov e in dc scribt n!!- ne w
specie,.
The spec ie- of Phyll" ho l<' . are "rranj!ed herc in
four prov isional spcc ies j!m ups . T he ' e groups arc
intended 10 represent relationships , bul can do so o nl)·
imperfectl y h1x..u-e of the incomplelenc" of ",'ailable
data , Tn arran~ the , pecic- in groupe. I used prese nce
or "b<.ence of toe we bbing.. position of p~ i n:d '1ripe~ .
snout-ver n le ngth. and re-cmblancc of cnlor p attern.
/976
FROGS OF TH E GENUS PH YL LO BATE S
M u>cum
a~vi alion s
Th e foll owing ab breviat ions ind icat e tbe loc at ion
of preserv ed specimens:
AMNH
Amcrican !'>lus<:um of Natur al Hi, tury, Ncw
J
19Z6. PIt,.lIob<;tr. (pan). Soble : 9
1 9~6
lHordrobertn (part ). Sobk: 9.
19S7, Phyll ob.rlr . (pan) . DlIJIn; 77-7M
1968 Phyllohalr. Sevege: 747_7411.
Ym'
A~SP
BM
CAS
CAS·SU
CM
CRE
P..l~H
GO V
U.s
lAP
J~IS
IPB
K!'>l
KU
u..CM
LG
acz
MBUCV
MH!'OP
M"'G
MRHN
MSH
"tV Z
~ H MW
NMG
RMNH
TNHC
UMMZ
UPR-RP
USNM
wCAB
wcs
WED
ZS BS
AciOkmy ot :-;;Ilura] Scim;(,., Phil.xlph ia
Brili'" Mu-,<um (:-;aluraJHi>tory). Loodon
Cali fornia Aude my Qf Sc ien ~ •• San Fu.n.
cisco
Slanfor<l cQtlcetion al CAS
Cal'J'lc.gie Mu,,"um. Pill, "'ngh
Un;"..,n ity of Swthcm California........ Angcle>.
(~ Rican EJ.pcdil~)
Field Museum of :-;atural Hi>tory. Chicallo
G . On.~s V. collect ion 31 USSM
[n,tiluto de UI Salle. Bogola
J .A. !>clen collection at US:>i M
1.M. SanS" priute O>leOlogical coIl«tion aI
CRE
JP. Bog;onooll«tion at Th HC
KrakO'" Museum . Ja ~e ll o n i a n Universit y,
Krak"w
" Iuscu m o f :"O alural Hi. IO' )', Univeui ty of
Kan..a.. La" renc..
:"Oalur al Hi. toT) " l u!>C um of to. Angc l...
County. to. Angel....
J. i.e, cure private colleclion. Pari,
Museum of Conlparativ.. 7..oology. Harvard
Univen il) . Cambridge
Musco de Biolollia. Univenidad Central de
VenelllC'la.Car""'b
Mu..cum :"O alioMl d'Hi""ire :-;aluf!'lle. Pari,
Mu....u f'araense Emilio (;oeldi , Belem
[n"itut Roy al des Sdcnce . Natur ell ... de
BelgiQIIC' . Bru••els
M.S, Hoogmoed roIl«lion al RMS H
Museurn of Vmrbratr L.>loJy. Univcrnly of
California, Bend ..)
Naturhi'lnrisehe . Mu><:ul11. Vicnna
Naturhisloriska Museel, GOleborll
RijL;mu ,,"um van !"aluurlijke Hiuorie . u-idcn
l ln;,.......ily of Tu n . Au"-in
linivenily of Michigan. Mu,,"um of Z"O~I .
Ann Arbor
Unis'ersily of Pucn" R i~o, Rio Piedras
National :\lu....um of Natural Hi. tory. Wa,h.ngton
W.C.A . Bokermann private colleetion . Sio
. ".
W .e. Sllr rbroole ro lleclioo al US ~ M
W.E. D~ llm~n collectio" at KU
ltlo l o~isc he Sa mmlu nll de, Bay'et ischen
Staale>-. "' unich
PhylloOOIl" Bibron in Sagr a 11141
Poison-arrow Frogs
Frontispiece s 1·1[ and Figu res, [- 12
1M !.
11<41.
111112
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•.
Phyll oi", u .I. Dumo.'ril and Bibro n; 637.
D r ordToI><ltn (panl. Boolenga: 14~
1 81l~ . P/tJ-ih"' tn (pan) . Boulen~r. 19-1
IIIIUla.. Ph)'II0b0tn ( part ). Boulrnrer: 2Ot>.
19W Phl# oI><lrrJ ( part). Ba.rbour and :-;...ble: J98-402.
D efi nition. - Mu iUary 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 fing er
e4u al to or longer than second; toe webbing present or
absent ; in life . • Lin o f do Nim pattly o r emi n:Jy brightl ),
colored; pala tine bones absent ; omostcrnu m pre....m;
vertebrae usually ' not fused; tad poles ( Figs , 10-1 2)
with oral dis k laterally inde nted . number of demiclcs
not reduced . and anus dextral.
DiQgnm is.-Phyllooof.... d iffers fro m Dendrob«tt .
in havin, ue fiN fi nger eq ua l to or lo nge r than the
seco nd and in hav ing relatively ,mall fi nger d ish , In
Ph.l'lIob<lI..... the dish of the second 10 fourth lingers
usuall 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 fingers
usu a lly are al Ieas l 1"'lo;e Inc w idth o f lhe Finge rs .
Ph y /l"hrtfl'$ diffe rs from C "I(>$I(' l h ui in having the skin
of the dorsum partly or enti rely brightly colored in life ,
anti I pre cuma bly) in havin g ,Lin to xins .
D tJerip ti" n . -The snour.vem le ngth o f adu lts i.
IS 10 50 m m (Table I); female, avera ge lon ger than
mal es . T he skin is smooth or gra nula r Ma~iIIary and
pr em a xill a ry teet h a re present or absent; vomerine
teeth ar e absent. Adu ll ma les have voc a l slits and
u,u aU)' 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 l
slil_ and e xte rna l folds on the throat. The lip of the
snout 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'>lance
fro m the eye to rbe nos lril. The pocerodorsal por1ion
of the tympanum is conc ealed . Each digi lal disk beu..
on its dorsa l surface a pair of raised " scutes " separaled
b)' a media n groo ve: t he seco nd. thinf , and fourth fi nge r
d isks an: sm all 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 frin ges. and well-bing. A subartk ular
tu berc le occ urs on the vemral surface of eac h di gita l
joint. An oblong inner :.rnd roo nd outer tube rc le occur
on both the me tacarpu s and mc t:.rtar , us; .....meli me s a
median tubercle occurs on rbe ructamrsus. A ridgelikc
fold 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 ing
at 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 or
slig h.l ly 10 well devel oped . T he toe s are eitber weblc..
or bot....JJy ...ebbed l uwally between loe s II·III· I V).
In life , the ir is is blac l Of brt"" n (rar ely bronle)
and is neve r reticulated. Brighl co lors (red . orange,
•
~
~
TABH I
Snout-vern length (mm) of 732 adult specimen> of 20 Sf"'cie, of I'hyllohm cs. including , ample size (n). range. mean and sland.rd
error (;:±SEl. .00 ,t""dard devialion (SO)
All S[" cimen,
Sp..", ic,
,m ,h"Il\'i
' pecies"
aurmac"i"
b<".lcr;
hi"olor
f>oli,·j'IIjus
b<m/c"x cri
cW""'''''
fcm"",'i.
"'lie"
"
""
'"ro
.,
15
S
"
9
lu;:u/>rls b
p<1n'ulu.
"0
fN/cn,
38
p" 'W"
9
p,,'chri fNC/U.•
s m,,,,,~di,,,,,
/ricolor
"Mum u"
"
""
2
.W
129
";""'Ul'
n
zap"'"
"
Range
, :SE
SO
"
Range
16.5-21.0
19.2:0.20
40.0 :0,75
27.7 ±O ,54
37.3: 0,78
38.2± 0.35
24.8 ±091
17.7::: 0 .25
169±0.22
26.6 ±0.43
23.1 ± 1.J3
20,5 ±0,53
20,5 :0.14
26.6 : 0.39
23.8 : 0.27
24 .~ :0,62
26.5
23.7±0.29
39.8 ±O ,34
1.16
2.411
2.97
2.47
2.32
1.58
1.49
050
2""
2'"
1,75
u
16.5-19.0
36.0-40.5
23.5-3 1.5
35.0
32.5 -39.0
23.0
15.0 -20.0
16.0- 17.5
200 - 31.5
21.0
.~ 6 0_ 42 . 5
23.5_34.0
32,5_42.0
32.5- 42.0
23,0-2().0
150 -2 1.0
16 0 - 17,5
20.0 - 33.5
21.0_27 .0
1 9 .0 ~ 23. 5
17.5 _24,0
21.0 - 3 1.0
16 .~ -31.5
22.5 -27.0
26.5
19.0-26.5
31.5 - 49.5
23.0 _29 .0
26 .~ _JO.5
25.7 ± 0. ~4
27.9 ±O ,29
' '''
2.4 1
3.23
1.87
,
,
23
,
20
"S
ra
,,
17 .5 ~ 2 2.5
'"",
21.0-2 8.0
2
l.l\J
3.8 1
1.95
1.22
1 9 , 0 ~ 1 ':/ ,5
18
"","
1 () . ~ - 2 9. 5
22.5 -23.0
26.5
19.0-24.5
31.5~ 42 .0
23.0 -2 6,0
27.0 -30,5
i :SE
18, I : 0. 18
38.2: 0, 75
26.9 : 052
35.0
.>6.3 ± 0, 38
23.0
16.9:0,29
16.9 ±0.22
25.8 ±0.69
21,0
19.2 : 0.1 1
19,0 ±0. 15
24.8±O,49
2.' . 2 ± 040
22.6±0 12
SO
"
Ra nge
1':/,0- 2 1.0
42.0 - 42.5
26.5 _34 .0
, :SE
20.0 :0.17
42.2 ± 0. 12
30.4 ± 1.15
0,65
1,84
2.48
"1
9
32 . ~ -42.0
17.~ ±O , 83
1.70
"
"
,
3().5_42.0
25.5_26 ,0
39.8 :0.28
25.8
11I.5 : 0, 31
S
2
1.23
O. ~O
2.93
026
094
200
3.29
0.24
26.~
22.3 ±0. 42
37.4 :0,25
24,4±0.38
27.8 :0.56
~
~
Females
Males
25
S
12
"aa
S
as
1.59
2.26
SO
L'"
S
1.37
u
1 7 .0~ 21.0
230 -33. 5
220 - 27.0
205 -23.5
19.0_24.0
SO
0.75
0.27
''''
2.49
1.39
1.29
19.5-31.5
25.0 -2 7.0
27.3 ±0,51
23.8± 1.59
22.2 ±0.51
21.2 ± 0. 12
28,O±0.J7
24.4 :0.34
26.0 ± 0 .3~
3.10
0.79
21.5 -26.5
40.0 -49,5
26.5 -29.0
26.5 -30.5
24.4 ±O, JO
43.(): 0 ,33
27.7±0.44
28.0: 0,34
1.48
2.30
0.97
1.19
2~ ,O _31.0
2.57
2.75
1 15
L'"
L'"
"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 follo wing 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 males
22.5-26.0 (mean 25.0): 10 fem.1cs 26.0 -31 .0 (mea n 28.2)
~
~
~
~
~
-e
§.
~
~
§
~
8
~
~
~
1976
FROGS OF THE G£\'US PHYLLOBATES
yellow , green, blue, and inter media te hues) o' ually
form dorsolate ral or latera l stripes on a black or bro..... n
ground co lor. bot in some ,pecies, bright colo rs cover
the entire dorsum or form dorsal marblin g. The venter
us uall y is blac k with brig ht spot s o r marbling . In
prese rvative , the bright colo rs become: whi te , gray,
brown, or black, and the coke pallem may become
ind iSl ing u i~ habk from the ground color . The surfaces
of the mU!iCles arc sparsely to densely flec ked with
dark pigmen t. Pa latine bones arc abse nt. The o rbitosp henoid region is partially or com pletel )' os..sifiN.
The ornossemcm is present. The vertebrae usually are
not fused .
COil/till. -Phyllo/J<I/t's include s 20 species, occ urrin g from Costa Rica to Brasil (Figs. 13- 17); Ph.l'"oba It'S spec ies (to be dese ribed by C . W . Myers and
J . Daly) . P. on/hOlt)'i xoble 192 1. P . oumlot'n;o
(Boulenger 1913) . P. bout<,ri (Meli n 1941). P. bir%r
Bibron 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 . Sch midt
5
1857). P . fl(J",ulus (Bou lenger 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 ule nger 11199).P , Iril'iIlO Ilf.\' ( Spix 1824).
P , I·ill alu,. (Cope 1893). and P . Wptlru new species.
Table 2 shows all species and sub'flCcics desc ribed as
or referred 10 Phyllo/1(Jln .
TaJpo l<'s (Figs. 10 -12). -Tadpole measurements
are in Tab le 3_ The tadpoles illu, U;Ited herein arc
identical in the follo"'i ng characters: Tbe oral papillae
border Ille e ntire po'ter ior lip. but borde r onJy tile
lateral por nons of tile anterior lip. Tbe oral disk is
latera lly indented . The den ticle formula is ~. The
beak s are thin (moderately thick in P. ~iIlUl u_,) and
serrate ; lhe lo wer be ak is indented . The e ye s are don;aL
The spiriICle is sinistral. The anu' i~ <;Ie uraJ.
All kno..... n tadpo le, o f PII)'1I" OOI<'s have rounded
o ral pa pillae ucepl 1'11)'110001<'1 sp«'ies (10 be de scri bed), which has poinlCd papillae. All have one ro....
of papillae ucepl P . l ·i,faIUS. " 'hich h," two rows . All
hall: 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 subs pecies described as
Of
as I' lryIloINuu IlTlless OIheTwisoe
refcrnd to PJrylIoI>aln. All naJnC$ in til'<! column we re dieocribed
Subspecies aRc listed unde r lht ir spec"" ,
indic~ed .
NAM E
a1D8"J'O'!'"
albog""t1lU3
onrhotr.,;
" "rala (H)'/o)
a""",,.
a" roUlr ni" (lknd rob mu)
bau l" i IDr ndroba u.}
bralriciM
brtb<'; 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:rap is!
ca piwba
cNlh .
c"'ocw
(Hyliwhu)
evi4uU (HyliuJllS'
cyu i,lOlllltoidr.
d..M; (PrtnllorrapU'
clclOl'tS
rq lMlwritJl;. (I'nmM rapis!
upin(JJa;
r lU:nr mi s (Dr ndrooolr. )
lem" ",/jJ (ProJlhn ap i. )
ImlOra liJ (P",sllu:rap i.!
fr Utu ( Prollltrrapis)
A UTHOR ASO DATE
Bokel1lWln 196 7
BoulenCt'T 1903
Noble 1921
Wie<! 1821
Girard
1 8~~
Bouleng<:r 1913
Melin t94 1
Barbour and Dunn 1921
Nobk J92~
Bibron ill
s.a,n.
IM I
Jimenezde Ja Esp.da 1870
Boulenecr 1902
. . - 1909
FilZinget bt Sreindachner J864
Cope 1887
Wine t930
Test 1 9~ 6
Cope 1887
Bokenn...n J967
Bokenn an n 196 7
W. ~ la'1 1873
BouJenger 19 J2
Boulen,er 1912
Cope I Bn
Rivero 1961
Giinlher J808
B~Uf J908
Funkhou>eT 1956
FilZinger in Sleindachr.er t864
Bouleng er 1883
Barbour 190,S
PeracCI t 904
OPI....IOS OS STATU S
Col,,.,crl.... olo8"J'O'!...
Co/(mrlw,,/bo,llJlOt...
Phy llobatu " nrhotr.,i
nol Ph ),l/ohalrJ; sla1 u. uncenain
Drnd",b<" r . aUrolU,
Ph,,/Ioha'r~ " url>1"rni"
Ph,·/I0b0 tcs ban /cri
PI,,'IIQbat<,' /liI~..bris
CoiOSfrlw brill"......
Plryflobo u . lJic<>Io,
CoIour,hllJ bo<tvIgri
PIry.~lobotr'
bo/i-.r-...
Plryllobot cs bowl"ngr'i
pict...
P Iry.'l/oba ru pic ,...
Cro m>d"" ryllil' gaudic lwudi
Co losrn h" . bro mdic<>la
Co /oslrt h" J brlilrmr u.J
C<>Iosulh.,. c"pix aba
C<>I".UIN" c" ,ioca
S)TTItopIw. eNlI.., ...
COwlrl/.... chocot'mis
CdtW<',W roi/Qris
S,-rrltt:>p/tIIJ ryUigrwlllroidc.
Co/MIT'''''' <1_ i
nomen nudum
E /rlilIM rodoc T)-f... liI"iu rilO'lU
Plryl/obarrJ r JPinoJa i
Ph )'l/oha rr l pic'"'
Ph yUohulrJ Ir mo' '' /is
Ph yllobalU ho lillrnKcri
Co/m tclh ... I csrtu
PIry.~lobeur.
BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY
TULE
fI(Jw,pi(1(J IHyl"l'lni(J)
flolawr
fuliginos as (HyIMalus)
fusdcul(J
gl(Jnlfulosu<
grunulive"'ri< IHyllxulu<)
Iu,ltneli IlN~.drobales)
ltermlntlf'IPn>Sllterap;s)
It"igru.. (HylmralusJ
Itylaefi,rmi.
infrog uuulUS
ingeri -(Oe",ir obale. 1
incu inalls IProSlNrapis)
intermFllius
king.bu'}'i
laMalis IlNndmbm..s)
lari"".'us
limbalUS
lug"bri s (Oendmba'e»
m"ndela '"m
ma"'Nsi"n",
melanor,ltinu,
mertensi
nebl in" (ProSlltu"pi.J
nice/uri
nigerrim o fHylol
nigd....n"is (Hyl"plesial
nuhieola
"b"'urus (lNndrobarn)
ol/u.iaide, (Eupt'mpltlx)
polm",us
poMma".i.IHyloxulus)
porv ulus (O.."drobares)
peruen.".'
perU"lana.
pelt'rsi
piou (H)'laples ia)
pious g""yu"..nsls (Dendmba'eSI
prum
pulcNlIu", (Pltyllodrumus )
p"lehripeollJ
ranoide, (Dendrobale S)
ridens
""rosan8""
riveroi (Pro.INrapi. )
"hrevel (Pm.fllterupi,, )
smaragdinus
suhpunelulus (ProSlN rUpi, )
subpuru;u"us ....alesi
sylvalleus
IMnia/us
lal"mUlIf'a.. (Oendrub"'es)
1",,,,Ii.' (Pro.<llterapis)
,..""vitt,"us (De" droba'e, )
Iltomloni
lin'I",lu. vi"",u.' (Ch'mi ro""',,)
I,i,'olor (Pro-"hcropis )
Irili"" alus
rrinllmis
I,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
No .
27
2 (conlinued)
A. LUll 1925
Dunn 1931
J;rncne~ oe la E<pada 1870
Fit~inger 1860
F; t~;ngcr ill Stcindac hner 18(,9
Boul.nger 1919
Boulenger 1883
Boettger 1893
rowler 1913
Cope 1875
Boulenger 1898
Coe hran and Goin 1970
Cope 1868
Andersson 1945
Boulenger 1918
Cope 1874
Co pe 1863
Co pe 1862
O. S<hrnidl 1857
K . S<hmidl 1932
Me lin 1941
Benhold 1845
Cochran and Gain 1964
Test 1956
Noble 1923b
Spi~ 1824
A, LUI~ 1925
Dunn 1924
Durneri! and Bibron 1!W1
A. Lutz 1925
Werner 18'19
Dunn 1933
Boule~ ger 1882
Sleindachner 18(,9
Melin 1941
new,pec ies
Bibron in Tsc hudi 1838
Hea'wole. Solano. and Healwole 1965
Boulenger 1899a
J imenez de la Espada 1875
new spec ie,
Boulenger 1918
Cope IBM
Andersson 1945
Oonoso· Barms 1964
Rivero 1961
new ,pecies
Cope 1899
Coo;hrJn and Goin 1970
Barbour and Noble 1920
Ander,son 1945
Cope 1875
Wcrner 1916
Miranda Ribeiro 1926
Cochr an and Goi" 1970
Co pe 1893
Boulenger 18'i<1b
Bou len~'Cr 1883
Gannan 1887
Spi~ 1824
W, Pelers 1874
Cope 1861
Werner 1899
Hellmid, 194()
W. Peters 1870
Boulen~er 18'19b
Boulenger 1882
new'spo:cks
Phyllobale. pierus
ColoSl/rhus "uhieolu
Co/o.w,hus/uligi"osus
nomen nudum
PIt)""loemus olfe"i
Colaslnltus {X,lmalus
PIt)'Jloboln pieru,
ColoSlnhus I, ini'aris
£/tulneroducrvl". dia'lt"'U
Eleulhtrodo,r,'lu" It)''',../ ormi.
Colo'lel},"s in/ragu""'...,
Phvllobote> i'lger;
COIOSlNhu,- ingulntllis
ColoS/elh"s in/trmedius
Culo<""hu< killgs""ryl
PhyUobarn?; SlaIU' uneen ain
ColOSI..,ltus IUlillosus
S",in,hi/lu, limbalUS
PIt)'/Iolmlt s lugubris
Colost"lltu.\' I,;nitulis
Cmostetlt u, brun""u,
PIt\'/Iobaln birolor
Colosrn h" , merlt"si
Colostethus "tblina
PIt)'llobal"s birolor
Phylloba'es "iviltolu.,?
Eleu,NmdoCI)'lu. nlg';l'<'",rl,
Coloste,ltus nubkol"
Dmdrobaln; sralu, un..,nain
Cohwe,hus ol/e"ioid..
Colo.'lethus polmal""
Colo.,,,,,hu, ingui"ali .
I'hyllaholes poroul" .,
nol dendrobalid?
Colo.""hu.,.peruvionu .,·
I'hylJobates pete"i
I'hylloba,n pic,,,,
Phyllob<Ut's pierus
Col""'eIItU.' pralli
Colo ."",ltu.' puleJwllu.,
I'hyUolrcun p"lch,i!",elus
Colo ,lu,ltu.<ranoid"s
Eleulh"rod,/Crylus ride..'
Colmurhu.• riO{:o,I'ung"e
Col".<I(,ho.' ,i....",i
COI"Slelho• .shrewl
Phyll"ba,..s ." ntlr" !!di,,u.'
Colaslelh"s subpun"",u,COlm-INIt".' .uhpu''''IUlU.\'
Coloslelnus HJ""li,u.'·
Col''-''leIltus I;,eni",,,s
Colo",..,ltus lul""'I1',,,,,e
Colwlt/hu.' . "bp """'lIIu ,
Pltyllob<lIt s IrivlllulllS ~
Co loSlelhos l/rorn"",;
Plty/lob<ms "ill"'u"
Ph"lIoblllfS "i,,>lor
Col",,,,,,hu ., I,ilinearu."
Colrmerh",' trioiloli.'
Phyl/"I""e, "ivi"aru.'
Veod",b</les "uram , ?
Vend'obar" "o""IIOj'
C,,/m-ulhu s ,' obpunelm u,
Colo,'lerltu.' ""rgeli
S,·rrhophu,.- ve,ru,·" I", ,,-,
C"/,, .,'etho. ",," ehru/i.
Colosterhos ....hv"'pt'ri
Ph"/IobllleS wporo
1976
FROGS OF THE GE,\'US PHYLLO BATES
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 lhe
po...enoe ponion of the throaL
TiMJpoks of P1I.'"/fobol~J resembk mose o f 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 from
tbcee o f D~ndrob.II~J : t.-:lpo le1; of D~"Jrobuln lIa'e
th icke r !leah. lIa~e ( e ~cepl in llle mi" UIUJ group) a
med illn anus and an UIl iltdcnlC'd or.Il di'l . and ....}me·
time' ( in the hiJlrio"icUJ and pumi fi" group,) have
a reduced number of dent icks.
S/,<,ci~J ff, IOllp s. -The 'pecies of Ph.'"If(,h<lt~J fall
into t"..o well·defined groo ps based on pre...:nce o r
ab-eoc e of we webbing:
( I ) The ,il species wilh basal toe webbing are
included in the [emorata grou p: within lhis grou p
are three subgro ups:jf'",oru/i, and ~"f,,,m (w r)' similar
species 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" lm
in USl\·/-.1 specimens is an ani facl)).
(2) The 14 species without 100: webb ing arc divided
here into three species groups :
(a) The bicolor group t fi~e ,pe.·ies) seems to
he natu ral. Three allopatric species. ou roeuema, 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...:mbks
bico/Ol,
( b) The pi<"tlU group (se \ e n s ~ ie s) rna) be
onl) part ial]) natural. Pictw. and i"g~li (both with a
eal f-~pot) probabl y are dosel)' K'laled . and ,...n'ulu.
( which somenmes ha~ a cal f· ~pot ) resemble, the m.
P('lt rJi and pu lchriprcl'" share ~imil"r color panern~
and may· tIC clo-cly related : boli,·i"" ... (whose co lor
is unlnow'nl resemble~ them . ahhou¥h b"li';,mu. also
7
weblcss species of Col"i/ ('lhu•. The
re la tio ns h ips amon g 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:d
IOge: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 larg e: ~rc:cie:~
rese:mblin¥ each O(her in color pattern.
Lifr 1IisI0l)·. _ Dcndmbiltids have: a comp lex life hi~ ·
lOT) uniqLIC among anuranv . These frog~ often bella'".:
a~~~ivcly toward me:mber~ o f lheu- own speces: this
aggre"ion proh;ibly is territorial de fense: (Senfft 19.\6 ;
Tc:, 1 1954; Se xton 1960 ; Duellman 1966; Goodman
1971: 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 Durant
1(74 ),
Ther e i~ nu amplexus (accor ding 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) . Three
oviposiling 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 the
D('"Jrob<lItJ linooriwJ group; D_ aUlotllS eSe nHt
19 36 ; Wardenier 1973: Polder 1974c ). D . U:"'('II'
(Polder 1974a). D tinelOri... ( Polder 1974<:); Type 3:
mak I)i n¥ tranwcrsd) on female ' ~ hoICk. known in
COlmlrl1lllJ ,... Im,,'u. lliiddeckc: 1974) . In lhe o nl)
~p«ics of Ph.' //obulU for which dala are a\'ailab le
10 me. P. ..ill"I'... lhe o"ip''' iting posture is e~sc:nlially
Type 2. with a brief di~p1 aJ o f tbe Type J Jll""lure in
some: ovip"'itk>ns (Po lder 1976; D . TTiIVis . ~r.'l(>"ul
com municmitm ). In ~pecies lhal use the T) pe;;: posture ,
resemble~ ~ome
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 , tbe
lad p,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 maximum
total le ngth' for that ' tagc Data fur diffe",n t stages arc "' paraled b)' sem icolon'.
, pede'"
b",.lr"Krri
up;"" ,,,i
!rmora/j.
PCIr'"
pklll '
_ 'u"ai"",
Iri'~llUf'"
4(J (I ) 10 .~.
27.5: 42 (I ) I U. 26.S: 44 (I ) I 1.5. 16 .~
(91 3.5-4, 0. 10.5-1 1.0
(3) 4.0. 11.0
( I I) 3.5. IO.~: 29 (I ) 5.5 . "; 30 ( I) 7.0. -. 34 (2) 6.0- 1 .0. IH I'
(8) 4.5. 12.5: 28 tn 5.0 . 15,0
( I) 4 0. 11.0; 15 (9) 3.0 -4 5. 11.0_12 .0
(2) 5.0. 14,0; 15 (5) 5.0. 14,0- 15,5
6 ,5.10.5; 34 W 80 . 18.5; J7 (2) 90. 26.5-28,0; 38 (4) 8,5_9.0.
(196) 4 0-5 .~ . 12,5- 18.0; 29
26.0-30.5: 40 (I) 10.0. 31.5; " 2 12l 11.0. 24.5-31 .0; .u ru 13.5.2 1.5
25 (4) 5.0--5.5. 13.S": 26 (3) S.5. 1 4,~; 28 (3) 5.<)--8,5. 19.0-22.0"': 19 ( 1) 9.0. 23.5: J1 ( I) 12.0. 30.5;
.\S il l 11.0. 10,0
28
26
28
26
24
24
23
en
- An undc >crihed . P'C..... 10 be \k ..:rihtd b) C.W, M) CTs and J. DaI).
"Some or all >p:<" i"",n . un""''''II""bIe beuu~ ul ~ge
8
No . 17
BUULTf,'" OF THE ""ATU RAL HI STORY MUSEUM OF WS AS GEU S COUflo7Y
the m~le sil\ on lhe l~ )' i nll: ,itt both ~fore and after
oviposinon. hut SenlTt ( 19 .' 6 )
pro~eoJ
Ihal
,~nn
re-
le a"" ' 'c u rs only af ler u viJ1<" it io n.
Kno n oVlp'"it ing , ite, are
( u 'uall~llea"e'
(usually
anached. 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 and
Hendrickson 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
tScnffi 19J6: Polder 1974b .
y''' .
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 wor d " g uard ing"
less appro priate than lh~ word "l~ nd i n l!" fo r (hose
species [e.g .• D , auratu, and P . l'iIlUlU~ J in ....hich a n
adul t mm:1) ' v isils t~ egg' l)(Y\Oo a nd t~n. "llhough
in some spece-, of Col(m ttlu.s , lTW guard ing (d.m'i ng
. "' ''y pm:!.:l.lonl doo OI."Cur. :IIl"COmpilllicd b) ' " guard
relief 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'k
o f II paren !, ap pa rentl y ad her ing by t heir be lli es 10
muc us secreted by lhe adu lt (Slcbbin, and He nd rickso n
1959; Polder 19 74 b); af ter cerrying lhem one to seve n
d<lys. lhe JXlrenl dcpo;.it> the tadpolts in " ~mall . , lill
bod y o f ",'altr. ",here lhe) co mplete 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 d
o r deposilc'd simullaneou,ly ( R u lh~n and GlIige 1915;
Sentf 1936; Ste bbins and Hc ndrid ,o n 1959; D. Travis
cummlmimliOl1). Kappler's ubservation (qlK>l<.-.J
in Bocfenger 11I1I1Ib) tha I P. Iri "itl<UU~ oviposil' in
puddle~ lInd retrll/l .,porb u, lad po les 10 large r puddle>
proboobl) is incOll'eCt (Se nffl 19-'6 ; Stebbi", "nd H~n ·
d rit"kson 1959 ). TadpoJe-e"""iers ~ not (itS forme rly
be liev ed) u clu s j' e l) ma le . Fem llle tadpole -carrier s
hlIv'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 ld e r 1974b), D . pumilif! ( Kila'ako 1967), Ph.l" ·
1"i><Jlelf emora li I ( KU 1 11515. ~ ISH 389). and P .
p nul; 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. in
CQI<ljl~lh .... . a parent i ~ kno"'n 10 c arry 1"'0 to 25
laup"k, (Dun n 1941) . In lhis respect. Phyll"I>ttI~ .• ,
which i, know n to carry one I" 27 tadpole, (T a ble 4),
resembles C"I""fflhl.....
f ·""d. - R ,R. Snelling (LA CMl kknlif<Cd the ,lorn·
ar h contents o f 14 , pecimell\ o f 10 ~p«ies o f Ph..-I ·
lob<Ju. ( T " bl e 51. Th e ~ lo m"ch. eon t"i ne d 24 1
indi vid ual invenebra les; nearl y half thr invencbl'ate s
" ere anl~_ ~10>l o f thr spe cie . pre yed upon inha bi t
liller on the (ore,t n onr.
P" ld e r (19 76) nuled a differe nce between captive
Dnulro lmlf.l (five species) and Ph.l'!l oh<tlf ," (three
sJl<.-cic, l in prey size lind feeding behav ior . Dendmi><JIt'J
t al< on ly relat ively sm all prey (Senff! [ 19 36] also
noted that D_ <11""'11.' eals onl)' '"lhe ~mallt>l insects
and 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
1"",..,,,,,,1
TA BL~ 4
Tld pole-<:al'Tyi nt in P"yll<>bcut'~ . n: numhrr o f came... laken; Tadpok.: minimum.m.~im"m (mean. rounded to loweS! " 'link
numhel"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>
<inlloon.i
<>«;",'
b<Julr ngai
r<p'''''lU'
jr"""QiiJ
prlr"1
".rlUS
_
...,.ti .....
m .. "",,,,,
"Ulatus
,,
1
1
2
2
2
,
"•
,"'".,...
,
u4 - 9(6 )
2-3(2)
2- 11(6)
...
".
28
ie
28
C...,
.\.1 16. ~
.\.I41),j
M 19,0"
M 16,0- 17 .lJ(16~)
f 25.0- 31.5(28.3)
7
24.25
3-27(14)""
1_ 1:1(4)
28
.\.l 25,0; F 28_()(26 . ~ )
M 21.G-2 2.()(2U l
M 26 _ ~
M 3l.S -40.()(36_6'"
2~ ,26
10-1
l_llc4 )
4 -1116 r
"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)
26.211
24.2~
25 , G-26 . ()( 2S _ ~ t
/ 976
K~J /(J /h~
Ia
lb .
2a .
:lb.
la.
l b.
9
FROGS OF THE GENUS PH YLW BATES
eats both small and large prey (suc h a~ meal worm s.
and eart hworms longer tha n itselO. D. T ravis. ...ho
abo noIed the ge neric d iffere nce in prey size , suggeMed
to me that il is the result o f the absence of teeth in
Dendrobates and the presence of teeth in Phyffobal~s .
Ba sed on the spec ies th.il l Polde r o bserved, ho weve r .
tee th c annot be the sole e xpla natio n (I' . Iril·iuul US
lacks teeth) . nor can body size (P. villulu.< Is much
small er than D . a::. u r~ us). According to Polder , in
feeding behavior, Dend robates is a "seeker." hut
Phylloha/~s is a "sitter and watcher."
Sumlllur}" of distribution (Figs. 13-1 7l . - Sped es
o f Ph}"lIohat~s occ ur from Costa Ric a south ward to
Penl , Bolivia , and soutbern Brasil. Mosl species o f
the'-C diurnal. ler re slr ia l frog s inhabil wei tropical
jo rest. Reliable colJe,ctinllrecords are below 1800 m.
Only 1...·0 (I'. lugubr;s and P . l'itla/us) o f lhe 20
species live in Ce ntral America. and lhey are d,>,><: ly
related to a Cbocoan species ( I' . ourQra~lIio} . Ten
spec 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. zapa rov dwe ll in the
Ama zon ian regi o n. Fou r spec ies (I'. f~lIl .. ratii , P.
pin uJ. P. pulchrtpecna. and P. " il'illmu s) occ ur in
the Gu ianan region; all but one o f these (I'. putchnp{'c/UJ) also occur in the Amazon ian region . Fo ur
species (I' . aurotaenia, P. bicotor, P. fx}f/l~ngt'ri .
and P. eJpillo.' ui ) inh abit 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 tern
slope of tbe Andes in Peru. Two species IF. Iric"lor
and 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 PacifIC
Iowland~ and Andean fOOlhills o f south ...-estem Ecuador
and l"IOTlh...estern Peru.
Tbe dist ribution ...ithin polaical units is a~ fol lows:
Costa Rica: IWO ~pecies (I'. lugubris and P. "illatus);
Genera of
T.... uS
~ h "",,1 ( _ of 14 <pcei_ns of 10 opec:ies of
PII)'//obateJ . Colum n A: pm:enlare of 10IaI ....."ber
of prey individuals. Col umn B: pen:enu.s:e of
IOU.I number of "<>ma"h!..
Food Cal(llory
A( 4)
B('l)
AnlS
Tenn ilCs
Spiders
Bulin
H"mnpleran, and lrue bugs
49.4
78.6
17.0
21,4
7.,
42,8
Flies
Wasps
Inseel w...u
Sprinllail.
Gras.\hoppen
""""""""'"
14.S
,..
H
3n
21,4
21.4
7.'
7.'
7. '
7. '
'.0
L3
Doll"
0.'
9.'
0.'
Panama; one: species II'. /ugubris}; Colo mbia; eig ht
spec ie. (I' . a urola~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"r ali.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 s pecies (I' . m,liviallu.I and P. piNUS); Bra s il:
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,
I P.f~moralis and P. piNUJ}.
I have discussed the lOOgeogra phy of dendrobatid
frog s in an earlie r paper (Silverstone 19 75).
J)t'llclrob/llid(l~
Term inal d isk of thir d linger usually less than 1.5 lime s width of third linge r; dorsu m dully colored or
brightly co lored in life; maxillary and premaxillary teeth present or absent; tee webbing present or
absent ; first finger equal 10. aborter man. or longer than seco nd; palatine bones present o r absent:
omosremum present
2
Termi nal d isk o f third finger usuall)· IWO or more limes as wide as lhird linger ; dorsu m partly or entirely
brightly co lored in life; maxillary and premaxillary teeth absent ; toe webbing abse nl; firsl lin ger shorter
man second; palatine bones absenl; onjouernum presen t o r absen t
Dem/robares
Dorsum dully colored in life; maxillary and premaxillary teeth present; palatine bones present or
absent
.
Colostethus
Dorsum panl) or entirely bri~hlly colored in life; max illary and prema xillary teeth present Of" absent;
palatine bones absent
Phyllobatts
Basal webbing present between roes 1I·l lI-IV (Fig. 18, lefl)
Toe ,,'ebb ing absent (Fig. 18. righl )
.
.
10
BULLETI .'" OF THE NATURAL HI STORY M USEUM OF WS ANGELES COUf',7Y
2a (II).
~b .
31 (l b).
3b
4.1
co.
4b.
1'.'0.17
LiEli1latera l ~Iript complete tfrom groin 10 upper eyelid; Fig . 101: don-urn red or not red in life ... 4
Light latera l ..uipc: inoomplcte Of a~ nl. although com plete 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 o f t be An(\e, ,Fig" IA . SF . 7F)
P . n pi nma i ( pa~ 29 )
Adull snout-vent le ngth 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»
!-Ied ian dONlI ...ripe pre~nl ('OlTlelime, e xpanded 10 cove r ITlO'oI of back ;
~Icdian dorsal >triP<' a"'ent
.
ng'. SA.
D)
5
6
S. (4a).
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 Iengl h 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).
eb.
Do rsum red in life ..
Do rsum brown or black in life
7a (6b)
Adu ll snUU I' vent lenglh 1.'i.0-20.0 r um in males, 17.0 -21, 0 nun in km<tl<' s; skin of dors um lI suall ~
smooth (e xcept po sterior bm:k tubercular}, some times gra nular or with scatte red ruberc l...s; th roat
hl..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)
Adull snout-vent ....ngl h 20 .0-3 1..'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)
7b .
8a (l b).
8b
Lighl proximos 'cntral calf.spot present (Figs. 9C -F)
Lighr prox ill1O\~ntral calf. spot ab-e m
9a (Sa ) .
9b .
Lighl lateral stripe presem
Lighl late ral smpe ah!<nr ( Figs. 3<:. 9C)
.
P. t "p',m (page 33)
7
. ............ ..... .. .. . .. .............. 9
I.
II
.. 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 I a (8 b). Dorsolateral O)f lalerdl slripe pre-em
I I b.
Dorsolateral or lateral smpe a ~ nl
14
12
12a (lIb). Adu ll snout-vern length 32.5-42.5 mm
12b .
Adu lt snoul·v..-ntk'ngrh 17.5 -24.0 mm ( Fromispiece I; Figs. .'lB. 7D)
13
, .. P. p<lrl'ulul (page 36)
13a (12 0). Skin of back strongjy gran ular t granule s sometimes disappear in preservative): hack im mac ulate 01'
with sPOls and marbJin l,': : Cordille ra Azul of Peru {Frontispiece ll ; Fig. 4F ). Phyllol",rn species (page 2U)
13b.
Skin of back slllo01h; back immaculate ; Chocoan region u r Colombia Wig . olE) . P , bicolor { p;I~"- 2.1)
14a (l la) Dorsola teral Of lateral slripe co mplete (from base of Ihigh
14b.
Later al 'r ripe incom plete (Frontispiece l; Figs . .'iB. 7Dl
15a (14al . Ma\illary and prema\illary teeth prescnl .
15b.
Mu illa/) ' aoo pre ma \i llar) teeth ab~nl
gro in 10 upper eyelid : Fig, I B)
15
,
P. 1'0 " 'u/" $ (pa ge 31'1)
Of
.
""
100 (lSb). Aduh snouHem jength 3 1.5 -49. 5 mm: oo""wn wmetime, bears light median stripe; venter dark ...ith
lighl mar bling 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 ...ith
dar k marbl ing or spot s (in pre-efVali' e . venter may become entirel y black)
17
17. (I6b). Cnnspicuous light ()'e lJo... in life) prox i~ thigh·"JlOI pre....nl ( Frontispiece II; Fig _ 6E) . P .!",IC'IIripl'ctus (pa ge 43)
/ 976
17b .
FROGS OF THE.GV'US PHYLLOBATES
Li~ hl pnn imodoNl llhigh-~pol
/I
. .. P . boIMu"u.•
al><.em lFigs. X:.9Aj
(pa~c
35)
18a (l 5a). l ighl stripe dorsola teral ( i.e., posterior end o f stripe i, u" back ut ba-e of Ihigh ); C..ntrul Americ a or
Sou lh Amerka we- t o f I ~ Ande , ( f.'jg\ . ~A-D )
19
18b.
Li~hl «npe lall'f""l
po'>lerior end of -t ripe is in groin); Soulh Amerka ea-t (lf the Ande, I Figs . 2F.
ne..
M-C. F) . . .... ... .. ..... ....... . •.
21
19a ( 18a). Venter blad with lighl d<ll~; Ch,,,:oan region of Col ombia. west o f the Andes tPronnvpie cc I: Fig, .
.. P. <lurol,... ,,;U (pilge 211
-l.C-D.8 B)
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'ually
relatively narrow : light area' un dOT,;!1 surface of limb , go ld, yellu "" or ye llo w· gree n in life; Caribbean
dr..inage uf Co -ta Rica and f'-..nama (Figs. 4'\. 8A)
P. "('(Uhfi. (pa ge 24)
2Ob.
Adull MlOOI·\-enl lengt h 22 .5- 26 .0 mm in male,. 26 .0 -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 Rica
in Gol fo Dulce region (Frcnnv piccc I: Fig . 4B)
.. P , "i ll </lU,\ (page 25)
21a (l8b). Ad ult snou Hen lle ngth Ie" lh.an 32 mm
21b.
Adult ..notI1-\"('nl length greaier lha n 32 mm (Figs . 2F. 6A-0)
• . ..• P . hcr.u/tfi l Pi'gc 45)
er.
, . P. {'t'll'fJ ; (page 37)
2201 (21a ). Ve nter marb led or spotted (Fronnv picce II: Figs ,
9B )
22b .
V.. nter n'11 marbled UT sported IProntispicce II; Fig . 8C)
,., P . Wlll fux d;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),
ing complcle lighl lalcral stripe; (C)
P . UlfJUFO. paralyP<' . Saray.cu , Ecuador (KU 12(670). she.... ing \'CnlrolalcraJ stripe. Diagrammalic 3JId noI to ."'"~.
.!I".. .
12
BUU£TIN OF THE NATURAL HISTORY M USEUM OF WS AXGELES COU;"7Y
."io. 17
E
F1:Gun 2. Donal •• poet. tAl Ph)"lIobm u "lvill at..... PallCm I, ho lol)'pc , Rio Tefc . Bru il (redrawn fn,m Spi. 1824:p1atc 9. fi, I );
(8 ) P. rr;villar~J. Pallem 2. Sam. " abel, Pe", (USNM 166764); (C) P, 'rM JI"tuJ. Pauem 3. N• •""u (lehe rg1c. Suriname (RMNH
6413); (D) P. ',i "rtaru" . Pallcm I, Pampa Hermon , P"ni (AMNH 43 11 ~ ): (E) p, trMlla/u.•. Paltem I. Pampa Hcnno~. Peru
(AMNH 43 117); (F) P. ba..lm", 1!<,lolypc. Roque . Peni ( NMG 51 n, Diagrammalic and not to scale .
r
FIGURE. 3. Don.aI -- !'«t. (AI PJryllobau! . pKtld . Pancm 3 (DmJro#1GtrJ
,•• leckK)'pe). Rio Mamore . Brull (NH MV.'
19190:3); (DI P. pi rtMS. Panem 4 /Pt NJ'Obtzu ', h<JJtNli. 1tc1CM) pl'). Yurima'
PI:ni t BM 194 1.2 .15.17) ; (O P. u.~ri. ho\()(ypc.
iUt'nio near Rio fhe ado . Co lombia (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
(lAC M 61030) . Diagram lrlJlio; 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 . Cost a Rica (CR E 1 182); ( B) 'P ..in" "... Ri"""" <10 Osa, CoI.ta Rica
(C RE M); (C) P_""rot_"ia. narTOVl- ' lolripN fonn . PIa)a de Oro, Colombia (LACM 4)19 3) ; (D) P_" _ _ ,,ia. broad.miped
fo nn . Quebnda Bochorarna. Colombia (LACM • .' 8(0); (E) P_ bit:'oIt>r
Santa Cecilia. Colombia (USS M 137:J.661; (F) P II"IlDbal" ' pecie>. Diriooria. Peni (FMS H .s6272l . Diagnmmalic and nollO "' .
FIGI' U S. Don.al npecl_ (A I Ph) llohutr . uic-olor . kclOl)pe. """"=ir. EculIdol"(BM 1941. 2. 14 18); (Bl P. ptU""tII... . Pueno liM.
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.! S H 13140); t El P. botM"gui. Imbili. Coloml>ia( USS M 141433); IF) P. rlpi_i. ,-anoe. SanIODominJO de Jot,
Colortlc\M CCA S· SU IOnS)_
DilI~:r"mmaic and
noc k> Kale
14
BUllETIN OF THE NATU RAL HI STOR Y M USEUM OF LOS ANGEU;S COUNTY
No. 27
FrClJRl 6 . Dorsal (A - C. E-F) and ve ntr al {OJ . ' pel."I, (A -D)/'hyllo""te, ha"leri. Chula,., Peni (US :-JM 166903A. 166903C.
166903 8 . 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). Diagra mmatic 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 609 81); (D)P. 1"''''01••, PuC"O Lib",. Ecuador
(KU 122262); ( E) P. bou/rog,ri. Imbili . Colombia (USNM 147440); (F) p , esplow "i. par'l)'pe , Sa nlo Domingo de lu' Colorado. _
Ecuador (CAS·SU 10375), Diagram matic and nO! to "".Ie.
ts
FROGS OF lHE GEAUS PHYlLOBATES
1976
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 .
( ~ ~lf 36183); (O P . in~ri. p-aIJipe. Aoerrio ..... Rio Pe-w:ado. Colombia (US1"M 146849); ( D ) P.
Pal'
Rio Vi\Ol; . Pvll
pic"".
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 IlOIlO ocale
16
BUllETIN OF THE NATURAL HISTORY MUSEUM OF WS ANGELES COUNTY
No . 1 7
• 0-.
F1GllU. 10. Tad polt o f Phylloba ,~s " " alla . "C'n"al, 1••..-0.1. and do".,J ospe<:u.. SIIll" 29 . •aL~n from tQck o f Idul l
.fler JI'O""in, in Capllv;!)). RincOn de 0... C"".. Ri<. (e RE IIOl). ~ equ al. Oft<' m ill;_ I..-
( b'I1 l""~
G§..
c
o / .- ~
w..
fl(l ll ll.E II, T8dpob of PII.,l lo11<l,,,•• lalCnol"'~l. (All', """"r,,~ri. St. ll" ~K. from bai;k of ad ull mn"
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' fr male.SatQ C« ili•• Ecltldof tlW 112310):(0)1' _ ... &Jill ....
~ 2S. from bac k 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 bac k of ad ult male. h iarnal.. Boli\<ia(U MMZ 5894 7); Ie) P ,ri·
";IItmd, Sla,e 28 , from bao"kof Idl>h male. FrNerik .....dlem I V.nI ..... s ........... ( BM 1936 ,9 H.). Linn "'l ual one mil"","",
F ROGS OF THE GE/I'US PHYLLOBATES
J976
...
~
~
::..::::-
.@
,
•
17
@
,@
~ .~
~
~
" ~
~
FIGl:at 12. Tadpo"" of P~llobal~J • ........lhp.b Ofll... ""'"" 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'M MZ Sll947): (H) P. ,ri.inalld (BM 1936.9 .) ,6). u..... "'lual."... millimclC1'
r-
o
'.' ''"UTI'
v
••_
w,,~
~
••"" ,.,.
.. ,!....
a
•
•
'. """'0;~
. ". ~t~ .
+
r-
-t
18
No. 17
BUULT/N OF THE NATURAL Hl srORY M USEUM OF WS ANGELES COUNTY
·· -.. .... .".".
..
.
.-
::.-
,, ~
'. "
-,-
..
.'
+
+
·..
...
~
_.~
~
,
,
..
c-....... --L
+
1970
FROGS OF THE GEAUS PHYLLOBATES
/9
[J
.......
FlGl: u 16. Di..ni bulion of PIr. 'lIobu,r .
;,,~ri.
" ." 0 04'"
lUI "
P. pun ...I... . P. prtrm. P. ptJrltrif'«llU, and P. """'atJi",,•. Am abo\co 1(0) m
RUUET/N OF THE NATURAL HI STORYMUSEUM OF W S ANGELES COUNTY
20
NQ,2 7
o
Ii""
Flcicu. 18. Vcnual~.
rOOl or I'ltylJobt.'~I. """'*ina difference be,,,,ecn t-ally .. cbtlcd - . n-JII·lv (loft p_ ~_tIlu.
UCM 61011) .-d ,",cblru IOn (riJbl:; P. ....,.".......... LAOI,I980I. lirot eq...... o. millimelcr
bicolor ,roup
rIVe
spece
This group includes
.~~. lPlrylWbotu
s,
P. aurolfunia. P. bic(}/or . P. lu/C" br u, P. v;ttalltJ).
Definit ion: sec ut-venr length of aduh s sma ll 10
large (18. 5-4 2..5 rnm); lox. wcbk•• ; palmi smpes
(...hen presenl ) dorsolateral , nol lakral (i.e. , poslcrior
end of stri pe 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],
197 1.
I'hyllob<lu. bi'"<J/or. Cochran: 107, plale ~9 (~olor
pooto).
Phyl/<>bu/u bic% r, Liilin" 161- 174
Drfinirion .-TeClh present; toes webless: dorsum
red , red -orange, or orange . sometimes with black
~POIS and marbling , or don;um black with red. redorange . or oran ge SpolS and marbling; slripes absenl;
lighl proximoveraral calf,s pol absent; venter red, black,
or panl y red and panl y black.
Diagnosis .-Ph)'lIob<lt<,s speces is I. member of lbe
bkolor iJUup. differing from P. o_O<',,;a. P. lugu·
bru, and P . vi,lalWJ in IMBer size (Table 1) and in
lacking stripes, and from P. bicolor in llaving a strongly
granular ~um and in sometimes having a different
donal panem . 'The red. red-orange, or orange ground
color resembles lhal of P. biro/w . bUI somclimcs is
broken by black spots and mlllbling; in some ...pe<:i·
mens, the don;wn is p~inanlly black . and lhe red
is reduced 10 spots and marbling (in P. biro/or. lhe
dors um is smooth and uniform red·orange, orange . or
yellow·orange).
Description, -c-Measuremems are in Table I. The
skin is strongly granular on lhe back and the dorsal
surface of the hind limbs and smOO1h on lhe head. the
forelimbs. tbe venter. and the ventral surface of the
hind limb. (in USNM 166904A, lhe skin is smooth in
preservanve. bUIcolor s lidc~ taken by w .e. Shcrbrocke
show lhal the skin of the back and the dorsal ,urfaee
of lhe hind limbs was strongly granular in life). Maxlllary a nd premax ill ar y reerh are present. The fir'l
finger is very slighlly longer than lhe second . The
toes are ,..ebless.
In life (color slides by W.e. Sherbr ooke; venter
nO( visible), in USN ~1 l66904A (Pronnspiece
the
iris and tbe tympanum were black. 'The dorsum was
red wilh black spot s on lhe mid·hack and postel ior
back . The forelimbs were enlirel y red; lhc hind limbs
were partly red and panl y black.
m.
1976
FROGS OF THE GEl\BS PHYLLOBA TES
In preservative . in US ~ ~I 166904 A. the tympanum
i~ dark brown . The dorsum is ITllNl)' c ream. with a
'mall dark bro...n spot behind the right upper eyelid
lind seven ~all irrellul ar dark bro.. n spots scattered
over the back; the extreme po'>terloT back is dark bro...n .
The side of the head ilfId bod)' and the entire forelimb
and !'land are cream . Tbe hind limb is cream, exce-pI.
lhe di~ portion o f one thigh. the \'e-n1fal surfXC'
o f lhe thigh. the antelm'C'ntral surface of the calf , and
the ventr al surface of the fOOl . .. hich arC' dark brown
Thc C'lllIt'1l'lC' po'>te-rior belly is dark brown; the rest o f
the venter is cream with an indistinct lillht brown wash
on lhe tbroat and bell) ',
In preservative. in fl.1~H 56263·72 . the- tympanum
is dark brown. The head and beck arc: uniform cream
in four specime ns (except for a small dark bro wn area ,
some times preceded by one- or t wo bfllwn dot s, on the
e xtreme posterior back). Four other spec imens have
one to several sm all brown spots on tbe mid-back and
po slerior bad ; the brown area o n the e xtre me posterior
back of these speci mens sends fOl1ha pair of e xtensions
along the poste rior one· fourlh o r one·fiflh of the back .
In FMN H 56272 (Fig. 4 f) . e xte nsive dar k bro wn
miU"bling interrupts the 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' elo ngate
spols on the back and an exte nsive arca on Inc head
(broken by two small bro wn SpolS and by an im:gu lar
brown are a ext e-nding antenad from tM bac k). The
forelimb is entirely cream in a ll sped mens excepe
FM ~H 56268 and 56272 . in ...hich the ventral surface
o f the forelimb is panI)' brown. The dorsal surface of
tM hind limb is entirely' or mostly cream. excepe in
FM S H 56268 and 56272 , in which it is entire ly dark
becwn . TIle ventral surface: of tnc hind lim b is entirely
dark bro...n excejx for an elongate cream arca on the
calf of some spec;imC'ns . The ver ner of the eig ht lighlest
spec imens is cream. usua lly ....ith a partia l bro wn wash .
In FMNH 56272 , the venter is dark brown wilh irregular cre am areas alonl! us lateral portions . In FMN H
56268. the venter is entirel y dar k brown ,
Tadpot..s.-O n 8 Oc tober 1966, W .C . Sherbrooke
c aptured USNM 166904A , a 40 .5 mm male; Ihe frog
had six tadpo les (USNM 166904B ) on its back. When
preserved six weeks afler c;a plure, 1hC' ladpoles were
in Stages 40 , 4 2 , and 44 . Me a sure me nts are in
Ta ble 3.
In their native habitat, adults deposittadpoles in water
in small puddle s and d itches (LUling 197 1).
Life hislol')·. - Liilinl! (1971) witnessed intras pecific
a"ressive behavior (jerkin g the head upward seve ral
till'lC's, then allacling and biling the opponent) in c;aplive
specimC'ns. Luling also reported l!'Iat aI me tamorphosis,
young frop are uniform dark black.gray ; they take-over
three I\XII'lths to develop the full ad ult oolonltlon.
SIC",lOch COll fn u s {one spec;ill'lC'n ).-From FMSH
56265 : HYMES OPTE RA: Formicjdae: P~idol.. spp .
11
(27) , Ponerillltt: fragments, ~I)' rmecinae : fragll'lC'nts :
COLEO PT ERA : fragmen ts of beetles of families
Staph)linidae, Bostrkhldae, Curc ulionidae. and Ela rendae , a lso O IlC' unusual bee tle larva o f unkno wn
famil y; LEPIDO PTE RA: Soctuidae? (l lan'a): HEMI ·
PTERA: Mirnae~ fnogll'lC'nts
P,..dators .-Liiling (1971) sa'" a snake (C hi,o nild
sp .) from the Cordi llera Azul allack and seize a C;apl M
specimeno f Plr)'lIo bafcs speciC'S; the snake immed iately
released the frog , retreated . shook it-e lf for 0 .75 min.
kepi its mou th ... ide open for 6 min. and rubbe d lhe
sides o f its mou th agai nst branche s . Th is repul,itm o f a
predator (proba bly becacse of lhe unplea....nr, burnin g
ta,l e of the frog' s skin secreuoej, oomb ined ....ith Lu·
ling' s obse rvatilln lhat lhC' ffllp, in their native habilal,
~it in open areas and contras t co nepicuous fy wuh the
surrou ndin g 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 v auon
record s are from 1300 10 1600 m. Phyllob<l/cs species
is known only from the Co rd illera Azul , Per u, which
is the drainage divide between the Rios Ucayali and
Huallaga (tributaries oflhe Amazon). Liiling ( 1971) too k
this frog on forested slopes, on the grou nd. in clea ring s that were overgrown with grass and terrestrial
aro ids.
LOHl lilin . - PfR O, HIJUi. _ : km 189. F.D.O. Inti......
Tingo Marla'PllCallp a roael. Cord illera Azul ( USS M
I66904A_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 (FMS H ~26)-72)
CI..areJ alld stahl<'d ffWl.. , UlJ. - FMS H ~265
Phy/lobatrJ aurDlacniD (Bo ule-n goer 1913)
KOkoe
Fron ti,p iece l and Figures 4C-D, 8B
1913.
1957.
1963.
1966.
1968.
1970.
1970.
"wrOl~"ia Boulcnger; 10 29. pla~ 104,
fig. l, hololype : BM 194 7,2.1 5. 13, Peiia Lisa ,
Rio Condolo, ChocO, Colombia. 92 m
Phyll"bcl/ts a..'o".,,"u.. Dunn: 78.
Ph)lI<>b<urs bicalar . Marki and Wirkop: 330.
Plryllabarrs la/ina . ... , Latham: 683.
PIr)'lIabcll'n a..row rnia . Savage: 765_766.
Piryll ohlll 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).
P Ir.• flobalrs !rmartllif (part) . Cocbran and Goin : 40
(1Kl1 lhe plate no r ,he specimens eum i~d; onl y
Drrtdrohrurs QW""oc"itl and PJryftobtJlr J tll<l"f>ltlr'li<t
in the s)1'O"ymy are P . tl"""....Ilia).
Dr mJrobt>1rs
D cjinir;CHI . - Tee th prc:\enl: (OC'S
eble n ; dorsum
black . wmt times covered by yello -oeange or reoorange 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
do rsolatera! stripe; light prc xlmo veme at ,""If-s pot
"b-enl; venter black with blue or grttn doh.
Diagnosis. -PJIl-lloba,ts QllrOl~nio is a member
of lhf:,bko,," group. di ffering from PIr.I 'l/ob OlrJ lopeCies
and P. hie-o/ew in smaller sin (Table I) and in hllvinj!
slripeS, and from r . t"gubris and P. ,.;/1.. ' .... in I....king
ventra.lll\<lfbling.
Due- r;p/ion.-The name PIrJllo bul#:s ollrotarnio
cumolly 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 smoolh
on the \<C'ntcr;md tbe limbs. ~l a~ iIlary and premaxi llary
lttlh are pre...,nl . The fin;( finge r is longer than the
sewnd. The lOCI are wcl:>le",.
One form (Fromispiecc I and Figs. 4<:. liB) is slightly
smaller than the other, This fonn incl udes 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 gro und color of
the do rsum and of the side of the bo:.xly is black . A
relatively narrow complet e dor ...rlatcral stripe extends
fn,m the dOr!'al base of the Ihi~h ak,n~ tbe back. along
the outer borde r of lhe uppe r eyelid. and along the
camhus rostrahs to the tip of lhe snou l. where it meets
its fellow. lbe su-ipe is brig ht green or p,ale }e llow·
~ reen in speci me ns from lh e Se rrania de Bau do .
and yello w. li ghl or da rk yello ....· oranlle. or light
brmtonish gold in specimens from the upper San Juan
drain.a).~. l1le dorsal SurfllCe of the limbo is black ....ith
gold. orange. blue. or gree n dob; lhe venter and the
ventral SUrfllCe of the lim bs ~ blacl. ....ilh blue or
sreen dols. The doIs are a[...a), blue in .pecimens
from the SaTania de BaudO. The dots are 'p.an.e on the
venter and ITIOIl: numeruu.. on the limbs. In ~..:rva·
nve. t he snipes become !!ray. and Ihe dots o flen
disap 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: fou r males 31.0-31.5 (3 1. 1);
three females 33.0-3.l.0 (33.3 ); all seven spco.: imens
31.0-34.0 (3 2.D. In life. the color resembles thai
or the narrow-stripe d form , but the do rsolate ral stripe
( which is yello w -o range or rcd ·or ange) is broader.
T he dorsolateral stripes somet imes are nearly united
by a dOl'Sa l suffusion (Fig. 40) , whkh may also cover
the side of the !xxi }' and of the snou t and pan of the
dONiI surface of the limbs. The ventra l dOls oflen arc
more numerous than in the narrow- wiped form.
Rrt't'Jing call. _The ca ll of bo lh for ms of the
kLik,~ SOIlnds lhe s.ame to lhe human ear. a loud. bird lil.e. ",hi rring t",'iller. consis lin!! of rapid ly repea led
notes. The ..:rics is repealed after ilTt'gu lar inlervals
of a fe.... seconds lo.l5 seconds . The call duration in
seconds (mean in paremhe..:,) is: narn'''''·slriped fonn
R io Sapipi. Ii ve calls: .l,O-6.0 (5 .8). Quebrada
Boclloramio. four calls: 8.0-11 .0 (8 .1); bmad-<triped
No. 27
fomt : Q uebr..d" Bocbo rama. five calls: 3.5 -5.0 (4.0),
The Mkt~ usually catls ...hile coocealed benealh fallen
leaves or loS" but -.ometimes calls ...hile Sillin!! on lOp
of a fallen Ie"f,
Rt'l ul imu ll ips. - The inh.abilanh of P1a}a de Oro
lold me lhe 1\arT"O\Oo'·stnped kt>/w.t d....e ll, al k»ser ele ...·
lions Ih.an the broiod·slriped t~; a ChoOOlndi;on rotd
me. ho...ever. lh.at on the Quebrada Bochor.llmi (upper
San Juan d rail'Lilj!C). lhe J\OoO forms an: no! separ;ated
by- ele\-'ation. btU b} the quebrada itself. and lh.at lhe
narro...·s triped t.",,~ does occur on hill. on the left
hank. AI lhe Quebrada 8ocho r.una, I look the narrow ·
strip..-d ioto; only on the left bank al ISO m , and lhe
broad·sl riped toio; only on a hill (the Lorna de
Encamacitlfl) immediately opposite lhe presious 10e..l·
ily. on lhe r ight bank al 400-520 m, On the Ca mino
de Yupe in the Serrunia de Baud " (near the upper
Rio 0 p" l!ad\' in the weste rn Atratc dr..in..ge), where
the broad .slriped kill",,: is .. bsenr. the n"rro w·sl riped
k,ik",.' reache s al leasl 500 m
Should Ihe IWO fo r ms of k,ik",' pmve to be dist inct
,pecies. lhe n..me P . Ulm>we " i" ...ould apply to Ihe
narrow-srripcd fonn . J,W, Daly told me the two f"m ls
inter breed in capli,'ily. prodoc ing vi..ble offspring.
T he n..rro ...·s triped ioko; re semble s P. IUlluhr;s
,,00 P. ";/I"/lU of Centr,,1 Americ a. bul is sep;<raled
from them b} " di,tribulional gap in lhe Danen resion
o f Pan ..ma . Specimens of tbe broad·,lriped i"t,,;
...hose stripes are .. Imost joined by an orange dorsal
suffusion (Fi~. 40) re-emble P. bicoJor (...hich h.as
a uniform red -oran ge. or.ulge. or l'ello""-<>r,lOge coesum. lackin!! 'tripe~). bUI a"erage ,mailer: seven
specimen. 31.0-34.0 (32_1). ve rsu~ 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 al kaloid rhut is OI\C of
the 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 mumlIl"ls, BTX .·au...:s an increase in permeability " f mem ·
bmnc' to Na · , wilh con-cqucnt melllbmnc <lq,,, lari, ...
lion . cnhare ed spontaneou s rrunsmrtrcr release. lIlu'Cle
contrlli.' tuTl,', bloc kade of !!"Il~ lion ie lran, mi" ion. and
cardiac arrh)'lhlllias 4 Kayaalp et al. 197U: Alb"'lU<:"-luc
c laJ. 1970
Elhn<>z,,,,I<>KY, -The C hoco Indian, of lhe uppe r
S..II Ju..n dr"ina~~ of Colombi .. exlrdCt bk>wgun poi"'>n
fro m both form, of P , uur{1/<l~ni<l. "nd call both
/;0/;"; llhe first vo",el i, nasal: pronounced kohn~·
KWA Y). ( 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.
FROGS OF THE GEN US PHYLL OBATES
/976
Because lhe ti>lwi release, ih skin poi....111 only under
~ll·e~' . lhe India ns pierce its visce ra wilh " wooden
ske we r inse rted throu!!h lhe lI1<,luth . and so menm es
ho 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!!; the
poison 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 and
do no! e lllr.tel poison from lhe wlwi .
D iHrib"tion " nd h" b it,, ! tFig . 13j . _ Ele .'a l ion
record, 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: Pena Ul'lI, Rio Condolo
(BM 1947.2. 15.13); PI:.~ :. de Om, Rio S:.n Juan (lJSNM
1506n -42, I~ M MlIl; LACM 4~792 _94) ; 16_24 ~ m W Playa
de oro (US r-- M 146630_31, 147204_07); Rio San h an
(US NM 159401_~O, 1594.'1 -1I6); Quc hr. d. IJoehnra ma.
upper San J Uall drainage (LACM 4379 1. 43795-11(1); upper
Rio Napipi (LACM 4~782 - 1I9); Irail N:l....ccn Rio> Napipi and
Opo~ado (LAC...l 4~m); Camino de Yupe. bet.... ecn upper
Ri" Opo~:.d,i :.nd Domingooo drainage (LACM 7197M);
upper Rio B""y (LAC...I 6 1020-21); All" del B""y (LACM
71979_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 : EI
Paikin. 12 ~m from B""nllVCnlunI (USSM 1451(5 ); Io.. er
Rio Caling t US:";M 1451061. (Of the LACM 'I'f"imen , cited
abrM:. 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.
P h.l'fI"htJIl/i bie%r Bibron in Sal!l'il 1841
;\eara
Fillure 4E
8M 191 0.7.11.53. USNM 48865. 122104. 14510506. 146630-J I. 147204...(17. 1 4972~-27. ISObJ 2-4 2.
CAS 115151 ).
DI:/ i"it;on . -Teelh present; toe, .. eblecs: donum
uniform l"((I--onm~'e. ora nge . or yellow-orange : 'lripe~
li~ht proximo.-enlral calf.spot ab!<nt ; venter
black . oeange, or J!oId .
Di"XntlJ;s. -Ph.\'/ft>ba,t's bic%r i~ a member of the
b icolo . group , dirfe r in~ from P . aur otaenia , P .
!u/(ubri., and P. vtuosus in larger size (T able 1) and
in lackin~ 'lripc:~, and from P hyll ohatl/S specie, in
ha ..in~ a sme o rh back and in lading do rcal ,pol>
and m ar bl i n~ (Pln'//"1>a11:J species has a 'tron~l y
granular bad and corneumes has dorsal ~p<.lls and
ma rb ling).
DpcripliOl1. -s- Me usure rncn ts arc in Ta ble I (t hc
holot ypc i, a 39,0 mill male). The skin is entirely
smoot h. Maxillary and pre m a~ i Jlary reem arc pre, en !.
The f irst finger is s li~h lly lon ger than rbc secon d. The
toe, are ....... bl..".
In life. in specimens frum ncar Santa Ce<:ilia and
La Union on lhe: uppe r Rio San J uan (W as..cn 19S7.
-'ta nn 19 57. and my notes ). the ir i~ ""a~ black o r
redd ish brown . Tho: tympanum was panl)' black and
partly )'eU(lY>'-oran~ . In some specimens. the lip of
the snoul w"-' bhock Ohi, .....a' the reason for Benhold ' s
na me P. m" lan"rrhinu.•I. The dorsum wa s uniform
bri~hl or,lIIge o r yeljow-oeange. The ~ide of the bod y
wa~ !!old Ol" ~l low -oran~. The <Jonoa.1 surface of the
upper arm and lh i~h was gold. yellow-orange , or d irty
ye llo...' ...ilh a black wa~h . The doNI l surface of the
forearm and coM and lhe ventr"l surface of the limb>.
were blac k , ""jth o r ""ithout numerous ycllo... dOh .
The venter was uniform black or black " 'ith a golde n
wash (....'m c timcs imerr upeed by a black parch on the
absem:
threat) .
1841. Phyllo/'rrlt'S Nco/or Bibron , Sagf1l: plale 29 hi" holo ·
1) 'PC; M H ~P
1841.
1845.
18' M
11ISII
11I67.
11ItW
18112
11I82.
1920
1920.
1 92~b .
83ll. Cuba lin enur for Colom"ial.
PIo,H,""II~' bico/", . Dum.:ril and Bjbr,,,,: 6.'11 -1'>.'9.
Ployll"""tn m~/<III<",Io;nl<.' Bcrlhold: 43, I)'pe h::.li ly;
Popa~'iin,IC.uea,1 Colomhia [. 1760 m)
Phl/"!><IIe.!>icol",. Giinther: 'ltl
Ph /l"!><IIe.' me""'''''!>i''''.... Giinlhcr: III
P!>,-I/"hclle. mel", ,,,, !>i,,,,,,, Kcfe" tein; JS4 -~55
P!>."I/"/oatt'S e!>"cM n., {, Posada: 205-207, lype I,><:al·
ily: ClVI;O. Colombia
Ph,"I<>i",u" bico/", . Boulcngc,: 195
Ph."lI,matn mrl"""'hi""', Boulcnll"r: 1%.
Ph..-lI" butn bico/"'. Barhour and Sohle , 3eN
PIr.'I/.""',t'. "",lan". 10", ,,•. Barbour and Sobk: "1)2.
Pir:,'Jlulo«u. niu fQr; SOOle: 1-2. fig. I , holtlt~pc :
AM ~ H 140211. Antic. [to.. nl. Anlinquia. C...I.. ,,,bi a
1957.
1957.
1.13S7 mi.
Piol ilob<ll hi<:-oIOI'. Dunn; n .
PIn-/lohat _la_hilt",. DuM : 77.
1957.
1957.
Pir.,'llvI><un tti«fori_ Dunn; n .
/k nJmb<i,n rypogMpIt",. M;onn; 513 (color phoIo ).
liN'Wri",. wa....cn: 7M
bkolM t p;orl). Cock an a.nd (joia : ~5 -37.
6A---e tonly lht fol...... iIlg _ IIt>I P . bfcoi",:
1957. (k nJmba l
1910.
Pir) ih>bDI
plalC
In prese rvanvc , in specimens fro m San ta Cecil ia
and La UnKln, lhe dorsum and ,ide arc uniform Ii~hl
~ray ; the do rsa1 surface o f the limbs is uniform light
gray'. o r i> dar k gray with c lo \Cly "'I lighl gray doh.
The ventral su rfa,'e of the limbs i, dark gray ..... ith lighl
gray dotv. The venter i, uniform lighl ~ray c xc.. pl for
a dark gray p;ltch on the thro"l in ....'m e specime n"
Pre.scrv..d ,pe:cime n, from the Rio Sipi are en tirely
dark I>rown.
Nm",'"ci"',,rt' ,_Phyllobatl:~ rhoc,""ui.• Po sada
1869 (lhe de-eriplitln ""as rcpri nl..d in 1909 ) has heen
considered a synonym o f f)f'lId,."h<J,,, hiJlriOlri,'u,
Be rthojd . but the ca ll (a ,hrill. interrupted ",h i'II<').
the cok» ( '>r;ln~e·)c l k, ..... alll"'.: . toIuish I>htck bel"",,).
the ~nou t"e nl 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 . the
color pan..m usuilll) is sp' 1l:led (but sorn e lime , rescmbles tlla l o f r . bi""lor). the -nout·'ent Ie n~lh of 6 1
24
B ULLETIN OF THE NATURAL HISTORY MUSE UM OF LOS ANGE LES COUNTY
specimen, from the upper San J uan drainage averages
34.2 mm (maximum 37.0 mm), and the first finger
is shor ter than the second. Posada (1869 ) "aid that
the Indians extracted poison from P . chacoensis. The
Choco Indians today do nol extract pobon from D.
hisl rio nic u ,1 (Silverstone 1975). They do extract poison
from P. bir%r; they call this species neara , one of
the Indian names Posada ap plied to P. chocoensts
(Wassen 1957; a herpetologis t told Wassell to use the
name Dendrobotes ti nclOrius. but w assens neara is
P. bir%r). Posada sa id that P. chacoensis lac ked
teeth. but he may not have used a microscope . Ph}'l loba tes ch",;o",n, i,l Posada is a synonym of Ph yllo ba l<'s
bicolor,
Distrib Ulion and hahitat (Fig. 14),-Rel iable elevation records are from about 25 to 1525 m. P . bicolor
is reported from Cuba and from the footh ills of the
upper Atraic and San Juan drai nages. from the Rio
Raposo. from the Serrania de Baudo , from both slopes
of the Cordillera Occ idental. and from the west ern
slope and a central valley of the Cord illera Central
(all in Colombia), On ly the record, from the upper
Atrato and San J uan drainages and the Rio Raposo
have been included o n the map, because I distrust the
Other records.
Barbour and Ramsde n (1919:77, 113) noted that
the type locality of Cuba is incorrect , and that de la
Sagra shipped material from the Latin Ame rican mainland to Paris with his Cuban collections. They thought
that the type might have come from Mexico. P. bicolor
never has been collected in Mexico , nor in Cuba subsequent to de la Sagra's publication ( 1841). T he type
specimen resembles Colombian spec imens and probably
came from Colomb ia. Dunn (1957) and Luling (197 1)
thought that the holotype probably was collected in
Colomb ia 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 slope
of the Cordille ra Occ idental, 1357 m). I collected
near Andes and at eight sites in the nort he rn and
central Serrania de Baudo , but never found P. tncolor
there. K . von Sneidern, the collector of the " Serrania
de Baudo" specimen. has stated (see below) that he
has collected P . bicolor only at Santa Cecil ia and La
Union.
Specimens recorded from Popayan [1760 m, on the
western slope of the Co rdillera Ce ntra l] (USNM
137359 -62. 137364 -70) were co llected by K. von
Sneidero. According to von Sneidern (ill Jill. ), "All
the P. bicolor I have collected are from Santa Cecilia
[411 m] or La Union [451 m]. .. one hour by horseback. from . . . Santa Cecili a.
The frogs come
mostly from the hillsides between the two localities.
I have never seen any colo rful Ph y lloha/ n or D end w ·
bates near Popayan; they do not exist here." Berthold 's
1845 record from Popayan (P. m/'/ai10rrh;llll~ ) may be
only a shippiog point: this supposition is supported by
No . 27
his description in the same paper (and probably from the
same shipment) of another Chocoan species, Dendrob ates hiJlrio'l irll .<. from · ' Pa payan,"
The most t r u~t worthy Colombian locality record,
are in the western foothills of the Cordillera Occidental,
e.g .. Santa Cecilia on the upper Rio San Juan. The
highest of these records is the beudwaterx of the Rio
Sipi. which is 1525 m according to H.G.F . Spurrell's
field notes at BM.
uH:ali/ies.-COLOMBlA , RisartJlda: San1a Cecilia (FMNH
54187,541 89,54193 -94,54196,54204-05,54 207.542 12,
5422t , 54233; USN~ 147225-26; A:"SP 25755: LACM
60972,61050): vicinity of La Unionand SantaCecilia(USNM
137359-62, 137364_70). Chm:D: head ..... aters Rio Sipi (8 M
1915.10,2t ,46- 47); t20 km S Andagoya (USNM 139776-77);
Rio San Juan (USNM 144994); opper Rio Alrato (ANSP
25756: USN'" 53168, 53t70-71). Vall e: virology field
station, Rio Raposo (USNM 151381 ).
DOUBTFUL RECORDS.- CUBA (MHNP 838). CO·
LOMBIA. Callea : Quintana , 3200 m ( FMNH 54183):
Popayan (Berthold 1845): Atl/iaqu.;,,: Andes (AMNH
t4028 -29); Medellin ( AMNH 13540, 3(08 1); O weD:
Serrania de Baudo (ANSP 25490).
Cleared a nd stained malerial ,-USNM 13?360;
LACM 60972
lugubris (0. Schmidt 1857)
Love ly Po i, on-arrow Frog
Figures 4A. 8A
Phyllobale~
1857.
Dendrobates IIlgllbris 0 , Schmidt 12- 13, Ihe
1858
Dm drobates Jugubr;s O. Schmidt 250. plate 2.
syntypes; KM 101611347 (now lost), road from
B<>;; as del Toro to Vole"n Chiriqui, Panama
fig. 14.
1900 - 1902. Hyillp/esia tinCloria (part). Gunther; 207.
Phyllohares MllIridae Barbour and Dunn; 159,
1921.
holotype: MCZ 8022, Finea Victoria near u nt,
[limon,] Costa Rica
1924
1926.
1952
1968
Ph)'lIolmres lagllbris. Donn; 5.
Hylap/esi" linrtv,ia (part) . MirandaRibeiro; 224.
Phyl/obates Jugul>ris. Taylor; 640- 643. fig. lO.
Phyllobares lugub"s (pan). Savage: 763_766,
1970.
Dendrobates Jugu!>r;s (part), Cochran and croin'
19-21 (not the plate; in the synonymy, Den.
drohares lagahris O. Schmid1 and Ph)'lIobales
lugabris as used by Dunn and Taylor are P
IU!lubris; oflhe specimens examined,onl)' USNM
I t8231 is P. lugubrls).
fig. 4F.
Deflllilioll._Teelh present; toes webless: dors um
black; co mplete gold , yellow. yellow-orange, or orange
dorsolateral stripe: blue or white ventrolateral stripe;
lig ht pro ximoven tral calf-spot absent; venter black.
with blue, greenish, white. or silver marbling
Di/1g"o~is , - Phyl/oblllt'S lugllbris is a member of
the bicolor group, differing from Phy/{obare~ species
and P . bicolo r in smaller size and in having stripes,
from P , auro we'li/1 in having ventral marbling, and
from P, vi/WIIl,< in smaller size (Table I) and in having
narro wer dorso lateral stripes .
1976
,
FROGS OF THE GENUS PHYLW BATES
Description , -i- Measuremems are in Table I. The
skin is s lightly granular on the dor sum . 1!'Ie dorsal
"IIrface of the limbs. the belly . and the ventral Mlrface
o f 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 than
tile second. The roe s are weble".
In life. the iris is dar k brown . The ground color of
the en tire body is b lack. A co mplete gold. ye llow .
ye llow-ora nge. or oran ge dor solatera l anpe e ~ te nds
from the dorsal base of the thigh alon g the back.. a long
the outer border of the upper eyeli d. and along the
canthu s rostralis to the tip nf the snout. where it meets
its fellow . A go ld. cream. whiti sh . or b luish stripe
e ~ ten(h from the arm in...:rtion alon g the uppe r lip .
end ing ventral 10 the eye. The dooal '>Urf.ce of the
lim bo. bears e xtensive go ld. yellow , or yello w-g reen
marbling . The ventral surface of the limb" the belly .
and somenmes the throat rear blue. greenish. white.
or silver mar bling. e ~ tend in g o nto the side of the body
as a ve ntro lateral stripe .
An a berran t l>ptC imen from Turrial ba. Costa Riea
(T aylor 1952:b41 . 643). had the Mad. ground color
covered by a wa,h o f pink ish on the dorsal surfac e of
the thigh. of yellowish on the throat. and of yclhlW.
gree n on the belly ,
In pre-ervat ive. the ground co lor is black: the ,;.tripes
and marbling arc whue or gray .
8rrt'di"~ coli , - 1lIe ca ll is a " rasping em. lasting
four 1Usix 'c.'Collds. pau-e . repeat " (Savage 1% 8).
Re!m iorl.lhip.•, - Savalle ( 196 8) co nsidered P . viI·
WIU5. of the Pacific co ast o f Co!ila Rica . conspecjfic
with P . lugub' iJ. The cen tral mountain s of Cost a Rica
and western Panama separate the two form s. P . lugub'iJ
is smalle r (Table I). usually has narrow er stripes than
P . vil/IIIUJ, and has gold. yellow. or yello w-g reen
mottling on the limbs. instead of blue-green moultn g
;1$ in P . vil/ulUJ (Savage (968) . P. /ugub,is a nd P .
" i flut us resemble P . aU' Olaf!";O o f the C ltooeo, but
are separated from it by a d i,;.tributiona J l!ap in eastern
Panam a . P. OU' OI,"" io. P. lugubri J, and P . "itl at tU
may be conspecific, but it is best to reco gnize the
three as d istinct spec ies until more da ta are gathered ,
Noml!'nd uIU'I!' . -P. /u~ubris is not the DOldm/l(JII!'J
/ugubriJ of Cochran and Go in (1970) ; that spec ies is
D('ndmba/('J ITI1t1COIUJ (Cope 186 1); see SavlIl!e ( 1968:
765) .
Di sr,ibUli01/ und habilUI (F i" . 13). - Elevation records are from sea level to 650 m (the type locality is
higher. but see Savage 1968). P . /ugub,iJ inhabit, the
forests of the Car ibbean drainage of Cos ta Rica and
western Panam a .
Ltxa/iliu.-COSTA RICA. HU N ;': juno;tion Rim Sa·
rapiqu' ""d Puerto Viejo (CRE 65. 63_69); Fine. La Selva
ncar Puerto Viejo (CRE 512); 16 km N Puerto Viejo (MCZ
75323 ). 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
25
Turr'a lba ( KU 24908) . Li",,,,, : Los Diamantu (FMNH
101097-99: KU 24909, 370(8); Unt ( MCZ 8(22 ); Sllre1ka
tKU 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; KU
800 54 - 70); 4 ,8 - 5 km W Almirante (KU 94813-21 .
1156)2 - )). 116748- SO; AMSH 81450- 51); 10-12.8 km
SW Al miran~ (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!'a rl!'d and srai'll!'d marl!', ial . _JMS 1683.
Phyl/oh/lll"J ,';I/<I/tU (Co pe 189 3)
Golfodulcean Poison-arrow Frog
Frontispiece I and Figure 48
189l.
lNlldrobarn li...:Torias VillalllJ Cope: 340. I) 'pc
locality: Blleno, "'re., IPunlaren ...) Co. ta
Ri,a .
1900-1902 . Hylaplr>ia tinctor;" ( p;>rl). Gunther: 207.
19 3 1.
Phyilobulr J IlIgwh';J (part), Dunn: 392.
1942.
Dl!'IIdrobatr J linc/OTias Q"'anu ( part) . Lau~nl
u
1952.
rse
Dr"d ,oba/r. rilK"'orius villa/US. Tayloe-: 5&2 .
Taylor u.id Cope'. lype o;ould lMJI be found at
lhe USNM
Piryllo!><J lr< {ugub, ;< (pan ) Sava~e; 763-766
(lMJI the figure)
Dtfini/iml . -Tee th present: toes IIo-cble!>S; dorsum
black. some times with broken yellow medi an dorsa l
stripe; complete gold. ora nge, o r red-o range dorsolater al str ipe: wh ite or pale blue -green ven trolateral
stripe ; liglll proximoventral calf-spot absent ; venter
black with while or pale blue-green marbling _
DiognOJiJ. -Ph" l/obat('J ";UG'UJ i. a mem be r o f
the bi co!o' group , differ ing from P . lugub' ;J in larger
size (Table I) and in having broader stripes. from P .
auro tae nia in hav ing vent ral marb li ng , and from
Ph,.Uoba tl!'J species and P . bi color in smaller siz e
and in having str ipes .
Dt sc,ipt;01I. _Measurement s arc in Table 1. The
skin is slightly granular o n the do~m , the dorsal surface o f the limbs. the belly , and the ventral surface of
the thighs, and smooth on tile rest of the venter and
ventral surface o f tile limbs . Ma>:illar y and prema>:ilIar y
teeth arc prese nt. The first finger is longe r than the
seco nd. The tees are webless
In life . the dtmum is black , and somenmes bean; a
bro ken ye llow media n dorsal stripe (Front ispiece I).
A comp lete gold, orange. or red· orange dorsol atera l
stripe extends from the dorsa l base o f the thigh alo ng
the back . along the ou ter borde r of the upper e yelid,
and along lIM: canthus ~tralis to the tip of the SllOut.
BUUET/.V OF THE NATURAL HI STORY MUSEUM OF WS A....'G ELES COU/l.TY
...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 along
the upper lip. ending ventral to the eye . The dorsal
surface o f the hmbs bean numcrou~ blue-green dol~,
almo st obscuri ng the black. ground color. The venter
and the ventral surface o f the limbs are black with
white or pale blue - green marb ling . extendin g onto
tile side of the bod)' as a ventrolate ral stripe from the
groin to the arm insertion .
In pre§('rvlltive, tile color i. e ntirely black, except
for tile dooolatel'31 and upper lip stripes. ...-hich are
white , and the ventral marbling lind the dors.il "lIrface
o f thc limbs, which arc PUfllli.h gl'3).
Brt'tdin Kroll. -P. vittallU hi s t.....o ki nd ~ of ~ing
calls (D. Travis, J't'rSOfl<lI c"","" m ica/ion). One call is
a shon, high.pilChed chirpin g. con~iqi ng o f t...o 10
five notes, successbely dropping in pilCh; it is emitted
by the male when sitting Il('\t to the female or when
the female i. pursuing him. The other call is longer.
a high-pitched trill lasting two to three liC<:onds: it is
ermued by the male when he is tryin g to attract the
female 10 the sne of oviposition .
Tudpo /t' s (Fig s. 10, (28) .- Three IoLs havc been
talen from the bach of ildull males in Costa Rica in
Jul y and August ITable 4) . A«ording 10 the eRE
eata logLK'. one of the~ Iot~ (e RE 903 ) came from
small side pools. hut A. StlllTCtt. the collector , to ld
me the ladpoles came from an adult"s back. and were
aljo... ed to grow before pn'M"rvalion. Two other lots
(e RE 7234 and 3152. in Stagcs 25. 28. alll.! 37. taken
in Cosla Rica in June) an: not from tllc backs of adults
A. Starre tt told me he found tadpoles of this spec ies
in ...-ater in tree holes and falle n palm petioles, but
nol in streams or side pools .
Unlike tadpoles o f Dt'nJrobaln. tadpoles o f P.
•·;t/atu s iIl'C IlOt aggressive to"ard each other ( Polder
1976).
Meawremcnts Me in Tabk J_
R~IUlimuhips . - Savage ( I% S) co nsidered P . •';1·
1t/11U con. pecifrc ....ith P . lu![ubris tsee lhe rclationships
section in the P . /ug ubri5 account. above).
Lif~ hi5101",.. - Polder ( 1976) ....itnessed intra'p«ifrc
a~~re"i\'e behavior (jumping on the back and prc" inE
do wn) in P, vitnnus,
In captive anima ls (Polder 1976: D, Travis.I'","'ol/ll!
('omnumif'tllj"n). courts hip lasts from one to seven
day s before ovipovition . COIIn, hip includc ~ calling
by the male, circ ling, , haking of rbe limb~. j umping
on and , iuing on tllc eouni ng partner (do ne b). either
~ x). and puNling (lhe female fo llo....s lhe male). The
mak c hoo...:s tho: site of o..ipo<>ilion (u,u all) on a bromcliad leaf. -omc:times on a fern k a f. r.m:l) on tbe floor
of the terr ar ium) and attrXh tbe female 10 the ~ite by
calling.
There is no amplexus. The o"ipositing posture is
Type 2 (re"'mbling thaI of tllc D . lilll'l"ri". group );
while the female lays the egp. the male u,u ally rc-
/1.'0.27
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: in
one , their noses tOliChed; and in t....o. the y remained
apart, exccpe that, commencing 5 min after the start
of o viposition and la~ ti nE I min, the male assumed
the Type 3 posture: he lay on top of the female. almost
at a 90" angle to her bo4y ....ith his hind limh, projecting
off to one side, and rubbed his belly again,t her back.
The fe male cont inued to la)' eggs .. bile the male lay o n
her. The male' s cloaca did not approach eilher the e~s
or the female' s cloaca . This behlvior. ",'hich resembles
tha t o f Culo.Jlnhu5 pul"'<J11U (Liiddecke 1974 ). may
ha~e • stimulat ing funclion . lIiding the maintenallCC'
of a proper behavioral seqeence: fen ilization does not
occ 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 position) 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. tbe
male , its on the eggs ; fertilization prohlbly occurs
du ring this fir'>! visil. In four breeding c ycles that
Travis ....itnessed, the male al....ays visited the egg,
twice on the day of oviposition . His fU"Sl visit began
immediately after oviposition . and lasted 5 to 65 min.
After an inter·. a! of ahoot 30 min, he at:ain loat on the
eggs for 60 to 90 min. The male then desened the
eggs for one ....eek , During tile second ...eel; afte r
ovipositi" n. the male visited lhe eggs once a day and
sat o n them (perhaps to I!\\\isten tbem) for 5 min: the
third week, he visited the ej,\gsone to three time s a day.
Pold er ....itnessed tadpole -c arryin g once . Polder 's
male .....t o n the jelly·m"-,, and made stamping movements; all 13 larvee ...nggled o nto 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) inlo
water in a pl1s.tic dish. Jerking movements b)' thc:male
may ha~'t stimulated the larvae 10 Iea>e hi~ back .
Travis .... itnessed tadpolc-carr)"ing rwke . On the firsl
occasion , Travis's male took up his tildpoles on lhe
t.... emy -t hird day after u~ipl.,it i\\n (o ne .. eek after hatching). The male apparently stimulated the tadpoles to
wriggle \\nl\\ his back by shaking his posterior end in
three 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 fo ur
tadpole, and retained tbem 1w 0 days . After releasing
them into a , mall pond in the terrancm . he returned the
same day' to thc: leaf nil. picked up ,he remaining t...-o
tadpoles. and released Ihc:m the:next da)'. On tile SC'OOnd
O«llSion. the: male 100._ up his tadpoles on thc: sevenIccmh day after oviposinon (day o f 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
One of Polder's female s laid egp everyone 10 two
I>< 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 hatc hing is 13 10 17
days: lhe per iod fro m deposition 10 me tamorphosis is
40 days at 22 to 25°C.
Stomach co-runts (o ne spccimenl.- From C RE
7235 : H YM ENOPTER A : Fc rrnicid ae: Was",a""ia
aUrap,ln Clala ( Ro ge r) (I). Smilnislruma sp. ( 4 ).
Dolichodt'rus ~p . ( I l. Paratrt'china "p. (2 ). ROKu ia
sp. (2). Phd dolt' so. ( I ): COLEOPTERA: fragme nts;
DIPT ERA: MyeelOph ilidae ?: fragme nts ; COL LEM BO LA: Podur idae "!(I).
DiSt'ust's . _Polde r ( 1976) h s di scu s-e d d i-e a -e s
of Cllpli\.-c P. ,·illa/us.
Distribution and habitat (Fig . 13).-Elention rec·
ords arc from ]0 10 70 m. P. "illa/us inhabits the wet
forests of the Golfo Dulce region on rbc Pacific coast
of Costa Rica.
o.a
Localitit's.-eosTA RIC A . "'",ur"." RincOn ok
( M CZ 75322: CRE 705. 899. 3076. 3114. 6391. 72341:
Rio 1'0:......"""". 7.2 km S RinOOn de o.a (C RE 7235): 4.5
km W RincOn de Osa ( KU 102161-6)); Camp Seanlc. 6.4
km
~p
Rio Rin" ,n (e RE 757): Quehrada Bnruca, Palmar
No,-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 168 1-82.
/ emor<lfis 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 to
moderate ( 15.0 -33.5 mrn); ba sal web bin g betw een
toes lJ· Ill ·IV; pa ired st r ipe s late ra l (Fi gs . 3F, 5A.
D-F); light pm~ imoveotral calf_,pt,t abst ol.
Phyllobutt's "",Ito ")'i Soble 192 1
Anlho 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 presen t: toes basally webbed:
do rsum (in prcservanv e j da rk with light med ian dorsal
' tripe: com plete light lateral 'tripe: light ventro lateral
stripe ; Iighl pro~ imo,totral calf.spot abo;enl: venter dark
with light marbling or ligtll with datk marbling.
Diuf(nosis . -Ph-,'ffohal~s "Il/hon,vi i, a member of
rbc /t'm"roli s gro up. differing from P. 1x,,,lm!:t'r; in
having a light media n do rsal , tripe, [ro m P. opino"ai in
having a light med ian do rsal , tripe and in having a COIJ1·
27
plete light lateral str ipe (Fig . f B: inco m plete in P .
~Jpi"owi. Fig . IA). fromP . / t'm" rafis andP .zupuro in
having 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 (the
holotypc is a 21.0 mm female) . The skin is smoo th or
slig htly granula r on the do rsum (e :tCcpl o n the poste rior
bac k. where it is tuberculat) and smooth on the limbs
and the verner . Mu illary and premuillar)' leelh are
prtstnl. The firs t finge r i ~ longer t han the SC'Colld .
Basal webbing occ urs bet ween roes ]I·lII -I V.
The co lor in life is unknown , In preservative. the dotsum is dar k brown with a co mplete gray or wh ile media n dorsal sm pe. eapanding an teriotly 10 COvtT the
entire donal surface of the l>/lOUl; the l>/lOUt sometimes
bears dar k bro wn ~poIs. The side of tbe bod) is dart
brow n , A com plete gray or while lateral stripe extends
from the gro in to the upper eye lid , along the o uter
borde r o f tbe upper eyel id. and along the cant hus roo
strali~ 10 the tip of the soo ul. " here it mttl~ ib fellow
A gra~' or ...hite ventrolateral Slripe ex tends from the
gro ill 10 tbe f~limb in-e rtion. alxM: which it uniles
with lhe upper lip slripe: tbe latter jo ins the canthal
stripe at the lip o f the snout. The dor sal surface of the
forelimb usually is gray or white ; the anterior and peslerior surfaces o f lhe fore limb bear datk bro wn loegitudrnal stripes. The donal surfece of the hind lim b bears
altemaling gra)' and bro"' D bands: the lhi~ h bears a
...hite o l>lonll pro:timodors.al ~ptJl and loOmC: l ime~ { a~ in
the holol ypel a while lon~itu dinal po, tt'rodor~al , tripe .
The vent er so met imes is predominantly brow n. and
sometime, 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 gra y or whilt
marbl ing in some specimens. and while with hrown
marbling in other s.
In a '\Cries fro m Alamor , Ecuador . the venter is dark
bro...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. the
venIer and the ventral surf"cc of lhe limbs " re white .
except for a few brown spots on tbe throat. on rbe s ides
of the bell y, and on the distal portion of the Ihighs
In the boloeype (Fig . 7AI. also fro m Sabias, tbe ventral
marbhn~ is about even ly divided hetl><'een bro...n and
while .
Tadpoles. - O ne ku ha, been taken from the back of
an adult male in Peru in April (Tah le 4 ). The bead -body
of 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 f
the Quet>rada AngO'olOra. Pow Azul. Peni. lhe male
ca rried eight to 12 tadpolc:~ in a dark. fluid -filled "blis_
tcr" on his posterior do rsum (B. Barnell. letter of 25
September 1931 to H,W. Parker at B ~I) : since the tad-
BULLETIN OF THE NATURAL HIS TOR Y M USEUM OF LOS A/liGELES COUNTY
pules "were free in it. not in separate pits." the "blister" was not an unhatched egg O1a" . Thi, "hliste r" 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
/641 6: HY ME~O PTERA: Chaleidoidea ( I); ARAC HN IDA: P,<udu'iCurpionida (I).
From AM ;o..:H I647!/': COLEOPTE RA (2j .
Di .\nihurion "lUI Iwbi'", (Fig. 14).- Eleval ioo record, are from 153 to 1387 m. P. (Im },o "yi inhabils
southweste rn Ecuador and northwestern Peru, west of
the Andes. Salvia, ( I0611 m) and Pono veto (610 rn) arc
in the humid tropical zone . allhough mo st of the terrain
ncur Ponovclc is tree Ie" and arid ·appearing; Alamor
(I .'oll 7 01) i, on a rid!!e bet ween heav y subtropi cal forest
to the west and north and the humid tropical zo ne in the
valley s below (Ch apm 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 st
country. subject to ann ual droughts " al Gramadal: both
localities arc belween 153 and 305 m (B. Bamcn.Jcuers
of 25 -26 Septembe r 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 ts
alon g tbc western slope of the Andes (Ch apman 1926:
plate I).
Loca li /ies ._ ECUADOR. £ / 0 1'()/ Salvias (AMS H
13739--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•. on
Case.dem 1r.il (FMNH 98 40- 42, 9844_46 , 9848_49,
9853_56, 9859_60). Turnbe: : POlO Azul, Quebrada Angos.
lura, nonh of Rio Ttimbcl . one hour ride [by horse1 from
E<:uadi>r;an horder (BM 1931.10.7.1); Gramad.1, Hacienda
Cas;las ( si ~hl record by g, Barnell. leuer to H.W Parker
.1 BM)
CI/'(l1'<,d IItld s/uil/"" mmuia/. _A MN H 16416, 16478
Phyllobll/<'S houlcl/g.-ri ( Barbour 19(9)
Marbled Poison-arrow Frog
Figures I B, 5E, 7E
1905
1909.
1920.
1957.
1968.
1970.
Prosthnapi.l femo1'alis BarhoU': 101, 22 ,)'nlype"
MCZ 2422; USNM 52406, 118132 -33; BM
1947.2.1 3.92_ 93; UMMZ 48070, h la Gorgona .
rNari~o, ] Colombia
Pro.<therapis bow/ellgeri Baroour; 89. Replacement
name fur Prosrhen,pis femo1'alis Boroour 1905, preoccupied by Pros/herapis femo1'(jlis Boulenger 1883.
Phyllabaus bou/ellgaL Barbour and Noble; 402.
Phyllobale.< boulengeri , Dunn: 78
COIOSle/hus boulellgni . Savage: 757.
Phyllobales boulengeri . Cochran and Goin: 38-40.
plate 7A- F.
Deftnilio/!. _ Teeth present; toes b.,ally webbed;
dorsum brown. often with dar ker or lig hter spolS or
No.27
marbling; l'omp!ele yelh'" or red lateral 'tripe; light
proximovcmra! calf -spot absent : venier white w ith
black marb ling or black with white marbling .
Di<iglwJi' .-Phyllol)(II<,., l"'u lell8l'1'i is a member of
thefemowli., group , differing from P. wl/hollyi in lack ing a light med ian d"rsal stripe , from P , e.,pillo.I'ai in
having a brown dorsum in life (red in P . e.'pillo.,,,i ) and
in having a complete light lateral stripe ( Fig . 18 : incom .
plete in P . <"lpillo.,ai. Fig. IA) , from P. [emoralis in
having light markings on the throat {usually entirely
black in P . f"mowli.l) and in smaller size. from P. lri·
('( )/o r in lac~ ing a lig ht me<lian dorsal stripe 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. The
s ~in usually is ent irely ,nwoth [exce pt the posterior
back , whil'h is tubercular in 'orne spel'imen,j , In a few
specimens, the dors um is slightly gran ular. In one specimen (GOY 9231) , ee entire dors um bears scattered
dar k brown tubercles. Maxillary and premax illary teeth
are present. The first fi nger is longer than the second.
The th ird finger is swo llen in ma le, and not swollen in
females. Basal webbi ng occurs between toes lI·m·IV.
In life, in specimens from La G uayacana . Colombia
(my notes), the iris was blac ~. The dors um was brown.
T he side of the body was black with a complete yellow
lateral stripe , The upper lip lInu the proximodorsal
surface of the upper arm were wh itc. The dorsal surface of the thigh was black with yellow spots or two
yellow stripes. T he digits were banded bro wn and
white. The venter and the ventral surface of the limbs
were black with white marbling.
In specimens from the lower R io Ca lima, Colombia
( I. Cabrera. notes al USN M), the lateral «npe and
the dorsal thigh mar kings were red. The venter wa,
gree nish or white wilh blae ~ mar~i np.
In preservalive , Ihe dorsum is light brown with dark
brown spo ts and marb ling. T he side of the body is dark
brown . A <:omplele w hite lateral stripe exte nds from
the groin to the upper eyelid , A white stripe exte nds
from the forelimb insertion along the upper lip to the
tip of the snout, where it meeh ils fellow . In most
specimens, the dors al surface of the limbs i, orangish
brown w ith broken dark brown transverse bands. In
some specimens. the do rsal surface of the limb, is
dark brown with narrow wh ile transverse bands or an
undulatin g white longit ud inal stripe on the thigh , The
proximodorsal s urface of the upper arm is white. The
proximal portion of the postcrodorsa! sur face of the
th igh is dark brown with a white hook mar k. Dark
brow n longitudi nal stripes adorn the anter ior surface
of the upper arm, the posterior sur face of the upper arm
and forearm, and Ihe anterior surface of the thigh. The
venter and the ventral surface of the limb, are wh ite
with da rk brown mar bling ( Fig . 7E) or dar k brown
with white marbling , In a series from lmbili. Colombia.
/9 76
FROG S OF THE G£ \ 'I.JS PHYllO BAl ES
both cui".. p"uem~ occ ur ; 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 ~ a
white median longit udinal ' lrip;:. In lhe Imbili .,;:rie~.
the postcrove mral surface of the t high is darl brown
with whi le does: in mo-r ~ pce i mcn s of lhis 'C r ic~. the
re-t of lhe ventral surface of the lhigh i, while Wilh
darl brown spots. bomclime, mluced in number. e.g ..
Fi ~ . 7El or marbling . In one specimen from Imbili and
another from La Guay acana. the ventral ,urfa..-e of the
th i~ h is dan t.uwn .. ith t....o undulalin~ "hite Iongitu ·
diN I .lripe~
T<JJpo /n (Figs. llA. 12A l. -Two 10h 1\;1.(' been
talen from the Ncl. of adu ll male. in Colombia in
Februar) (Table ~ l . ~lea>urc menb 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" (D unn 192 4) . and ,a id (IX1 sed on unpubli,hed
nOle~ by E. R. Dunn ) lhal the nHlul hpans of lhoc lad·
pole, of P. boll/fIl lI..,i are umbelliform. resembling
tho>;: of the ladpo le, of C. ""bieola . Tadpo le' (U S ~ M
145248) from lhe bad of an adult ropotype of P .
fmu /ulg..,i . howeve r. have mou thparh ~ i m il ar to
those of ot her spccie~ of nnHobatt'3.' lhe moulhparts
are not umbcUiform (Fig. 12AI.
MOIt'rio l of " nerrl ai" Jla lu• . _ S pce i me n~ from
Tanda pi. Ecuador. 1460 m. on lhe ....eslern ~k>pe of
the Ande. ea.lWII lhcar.t of San to Dom ingo de 100 Colorados I KU 111529-61. 120f>65- 68 ). belong 10 lhe
f/!'mo ro/i.1 group; lhey resemble Colombian spec imens
of P. bm,/nl llt'fi. except lhal ihe lateral stripe is net
quil e complete. the ventral marbling is yello.... in life
(notes by J . D. Lynch al KU). and the dorsum ha) Iighl
spols . So me specimens IGO V 9 148 - 50. 9229-30)
from the Chocoan 1o.... 1ands of Ecuador (Rio Blanco and
San ~Ii~uel de I<:>s CoJorado') resemble the specimens
from l anda-pi. Pn;M.>on,ally. I h3ve referred a1lthe~
specimens 10 P. boul~ng~ri. but I have uc1udcd them
from lhe lmlp and Ioeal it~· Ii"-.
SlOmadl ronl~nlS (t....o specimens).-From US....M
145253 : HYME:-<O PT ERA: Form icida~ : S(>l~n"psis
sp . (.5). Ph~ ido/~ sp- (1). Iridom.r rmu sp . ( I) .
M)'coctpu rus . 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
""""'" 1909
Diuril'JlllilJtl and IuJmtOl (Fig . 15).-EJevalioo recceds ( not includin~ Tandapi) are from 10 to 1145 m . P
inhab its the dense . .. et fore sts of hla Gor·
gon.;( (aboul 32 k m off lhe PacifIC COO-~t o f Colo mbia)
and the wet southern Choco an reg ion from lhe lower
San Juan dra inage of Co lombia south to northw estern
boul~"grri
Ecuador . I . Cabfe~ found specimens in we I ~in forest
on the lo wer Rio C alima. Colombia: lhe)' j umped
rapidl )' and "ere difficult 10 seize .
u>c"lit;t's,-CO LOMBlA. S ari ii ,,: h la G"r~na (MCZ
2422; UMMZ 48070; 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'H
b2892 -93 ): Imbili. Rio Mira (USS M 147431_50): La
Gualac""a (LAOt blO34- H . stoss» JUJlin (LACM 60973.
61036-42) , Vlrlu: Camp CanOn de Colombia. lo.. er Rio
Cali.... CUS" M 149723-2 41; R"" kd cl ler ~ion Viroior.
f'.dd SIal..... Rio RajlOJO (US" M 1513'79: LAO! bl0431:
CMe ..... (B IS H u:m).
ECUAOOR.-E. ..........." Cac""bi CBM 911 3.1 31: Saft
J: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 tnOUlh
of Rio Yambi CGO V 9235_3b): N .",, ~ ah l o (GOV 9227);
Rio Silanchi (GOV 9599-%00); 18 km W Samo D"min~'<) de
10' Colorado,. on rood 10 Cl>one (GO V 9232 ) . ea relti:
P:uamha(BM 98,3, 1. 1- 2)
Clrllr ..d and s/(li"rJ ntlltt't'ial.-A~t ~ H 7494; USI'< M
1 452~3; LA CM b0973: from Tandapi: KU 111529.
~.pinfJSQi Fu nlltc:>U--e r 19.56
TUTqllOi.e-bellied Poi'OO-al"l'O\>- frot:
figures I A. 5F. 7F
Pll.,'lIobaln
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 prese nt: toes basall y we bbed;
dorwm red; incomplete tUTqllOi.,;: lateral slripe : light
pro~im",..emral ea lf·~pOI absent; venter darl bro .. n
with I UTq uoi~ marbl ing or spots.
DillXno.i3, --Ph.l·II0b0I~J ~3piNJJlli is a member of
thefrmorolit group. differing from P . onthoto}'i in !iKk·
ing II hghl median doNllolripe and in having afl eccmplctc li ~hl lateral stripe. from P. boul~n~ri in hlo\ing a
red 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~ng eri.
Fig. I B). from P. f .." w r" li" in having a red dot'l; om in
life ( bla~l or brown in P. f n ,w raliJ). 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 ng
a light median dorsal smpe, in hll'o'ing an in,,:omplele
light 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 peue m, blll differs;from P .
pan'ulltl in llaving basal loe ..-ellbing (P. pan'ulou !l.eh
tee ,,-dlbing).
Dri cription._Mcas uremen ls are in Table I (the
holoty pe is a 17 .0 mm male) , T he skin i. slighlly
30
BUUET1N OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUf,7Y
granu lar on the dorsum and smooth on the limbs and
t he venter. Maxi llary and premaxilla ry teeth are presen t.
The first finger is longer than the second. The third
finger is swollen in males and not swollen in females.
Basal webtung cc curs between toes II·III -I V.
In life, in the type series (Funkhouse r 1956), the
dorsum was dark red. The side of the body was black
with an incomple te turquoise diagonal lateral stripe .
Ph yllohat esfemorolis (Boulc nger 1883)
Brilliant -rhighe d Poiso n-a rrow Frog
Frontispiece 11 and Figures 3F. 8E
The dorsal surfac e of the thigh was black whh wrquoise
mark ings. The do rsal surface of the cal f was dar k
red . The venter and the ventral surface of the limbs
were dark bmwn with turquoise marblin g or spots .
In preservative , the dOl'mm and the side o f the body
are dark brown. An inco mple te light lateral str ipe
e xtends from the groin about one- fourth of the distance
betwee n the groin und the upper eyelid; in CAS-SU
10375, Ihi, stripe d i" o lves into spots (Fig . lA ). A
broken light stripe extends alo ng the upper lip, The
grou nd co lo r of the dors a l surface of the limbs i,
dark brow n . The proximodo rsal surface of the upper
ann and markings on the dorsal surface of the thigh
(a prox imoanter ior lon gitudinal mark , a prox imo po~terior hook-mark . and ill-defined transverse hands)
are light. The venter and the ventral surface of the
limbs arc dark brown with light marbling ( Funkllouser
1956;plate l , fig. 2) or spllts (Fig. 7F in this paper)
The throat usually bears a light longitudi nal median
stripe
Tadpoles (Figs. llB . 12C).- Two lots have been
taken from the hacks of adult males in Ecuador in
Octo ber {Table 4) . Measurements are in Table 3
Funkhouser ( 1956:77-78, plule l . fig. 3) described
the tadpoles: he said they have a median anus. This is
incorrect; the anus is dextral. as in all o ther known
tadpoles of Ph.1'll" lw le, .
Mot erial of uncertaill ,wow.<, -A 19.0 mm ma le
(CAS-SU 10340) fmm the Rio Suno . vapo. Ecuador,
400 m (east of the Andes ). resembles the types of
P. e.,pi'lO.wi (from west llf the Andes), e xce pt thut
CAS-SU 1034() has a large white spot on the throat.
This spot looks like an artifac t overlying the no rma l
white marhling. Funkhouser ( 195(1:77 ) did not devig nute CAS-SU 10340 as a type bec ause of its lllcalily.
I am not certain that CAS·SU 10340 belongs 1\) the
same species as the ty pes llf P. e.lp illo,\(li.- [ have
e xcluded CAS-SU 10340 from the map and locality
list.
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 the
wet Chocoan region west of the Andes in northwestern
Ecuador . It is diurnal and Jives in damp shaded areas of
the j ungle floor. The j uvenile paratypcs were take n on
the bank of a small river. The movements of specimens
of P. o pino.,,,i arc quick and "nervous." Thc fro!!s
seck shelter unde r terrestrial debri s, never in the wate r
( Punkbouscr 1956) .
No . 27
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); Santo
Domingo de I"" COlntadlls (CAS-SU 10375; CFW 4399 al
UMMZ)
1883. Proslha<lp;s f em"wlis Boulenger: 635. plale 57.
fig. 1. two .y otypcs: BM 1'147,2.14,21- 22, Yurimagua>. Huallaga River. [Lorero. ] Peru
1920.
1920
1935.
1952.
1952.
1957.
1970 .
Phylloh<Jtu inguinoli.< , Barbour and Noble: 401
Phyllobales /<·morc>lis. Barbour and Noble: 401.
Phyl/ob<Jtcs fmlO' <lii.<. Parker:506
Phyl/ohmcs f emom lis. B. LUll: pl,'e 2. figs. 7. 9 .
Phylld"acs j cmorolh . B. LUll and Kloss: 671-673
Phyll<>bnres femoralis. Dunn, 78 .
PhyllobolCs fe m<lrali, (part). Cochran and Goin:
40-4 2. 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 ye llow or gold lateral
stripe; wh ite, bluish while, llr silver vent rolateral stripe;
ligh t prox imoventral calf -spot abse nt; 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 of
lhcfemomli., group. differing from P, am/"'''.1'i in luckin£ a light median dorsal stripe and in larger size. from
P, boule"xeri in usually having an entirely black throat
(tbc throat has ligh t markings in P . bou/e>lgeri) and
in larger size, fm m P. e,'p illo.I<' I; in having a black or
brown dorsum in life (red in P. e.lp itw,mi), in having
a com plete light lateral str ipe (Fig. IB; incomplete in
P. espmosai. Fig. l A>, and in larger size . and from P.
trico lor in lacking a light median do rsal stripe and in having white hones (green in P. trico l" r) , P. jn/lorl/lis
close ly resembles P. w pttro in s ize ~ nd in having a short
tars al fold (e xtending along the d istal one-third of thc distance between thc inner meM arsul tubercle and the heel
fold). P . femondis differs from P . zaparo in having a
black or brown dors um in life (red in P. wptm», in
having a complete light lateral stripe (usually vestigial or
ab~ent in P. W p<lro), and in having a largc promincnt
lig,ht proximodorsal thigh-spot (small or absent in P.
zapurov . P. fmwra li., resembles and is sympattic with
P. pie/l<s , bill differs inlarger size (Table I), in usually
having the upper lip stripe meerln g me canthal stripe on
tbe tip of the snout, in lacking a light prox imllventral
calf-spot . in having the first finger longer than the second
/976
FROGS OF THE GES US PHYUOBATES
(.00...1«juallo!he seccod in P. piC/ILl). in having ba..al
toe ...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 . The
d in is granular on the dorsum and the dorsal surface
o f the limbs, and sm\1'l"th un the venter and the ventral
surface of the limbs. Muillary and pre m a ~ ilI ~ry teeth
are pre\Cn!. 'The fiN fi n~er is lon~er than the second
Basal webbing OCCIin between tues II-Ill-IV .
In lhe fo llow ing co mpo."ite desc ription (ootes by
J.D. Lynch at KU. W.R . Hey-er and P.A . Silverslone
aI LACl\.I)o f specimens from Santa Cecilia and limon ·
COl:ha. Ecuador . from 5 km north of Pueno Asis and
EI Pepino, Colom bia. from Teresinha and Serra. do
l'avio. Brasil. and from SaUl Ton ue. French Guiana,
a color llOIed only in specimens from a certain coonU)'
is follo....ed in paren the\Cs by the abbreviated name
of that country. In life , the iris was black with a gold
rin~ around the pupil. brown with red flecks (some
Ee .). or bronze (some Ee . ). The head and hack were
black or brown (Ee. ). The side of lhe OOdy was black.
A complete yet low or go ld lateral sn ipe e xtended
from the groin to the upper eye lid. along the outer
border of the upper eyeli d. and a lon g the c an thus
rostralis 10 the tip of the snout, ...here it mel il5 fel10",'. A ",hite (Co .. Br.), silver ( F.G .), ur gn:enis h
bronze (Ee.) ",ripe utended from the f~hmb insertion along the upper lip. u., uaIly meeting the Cantllal
stripe on the tip o f lhe "!lOU!. A white. bluish white.
or silver (F.G .) ventrolateral stripe exte nded an teriad
from one-half the groin -axilla distance up tn the entire
distance. 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 brown
or gray ....ith scane red black spots. The proximodorsal
surface 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 upper
lip s!ripe; (2) a "ripe on the ventral surface of the
upper arm twbere it was yello"" o r orange [F.G .))
and forearm (where il ....as .... hire): (3) a ....hne stripe
on the anterior surface \1' f the upper ann, separated
from the second stripe by a hlack area. and usually
meeting the second stripe on the forearm. The pro~imal
half and the posterior portion of the distal half nf the
dorsal surface o f the thigh were black; the black co kn
surrounded a co nspicuous oblique yellow or oran~
(F.G .) spol on the base of lhe thigh. The spol sometimes utended along the eeure length o f the thigh.
Sometimes a small 'P'1I of lhe ~ color lay distal
to !he Ihipa-spoI ( Fig. 3f). The venter and the ventral
surface o f lhe hind limbs were black. brown (some
Ee.). or grll)', and (e' c-epl for the Ih1Ollt) bore ... hite
or bluish ...hue marbling. In some spec ime ns. the
ventral ground colo r was ",'hite , and the black was
JI
reduced 10 small spols. In small juveniks, the ventral
areas lhal were marbled in Jd uhs were unifOllll gray.
In preselVative. yellow, o range, and gold become
whhe. The ground color is black or dark bro....n. Rarely
te.g .. in TNHC 36498 from the Rio Pachitea. Peru).
the belly is almost immaculate white. The light ventral
marbling never extends onto the throat. except in one
spr;x:imen (KU 14436 1 from Nevan, Peru, in the upper
Pachire a drainage ). Th is spec ime n re sembles mos t
specimens of P . f~m",uljs. e xcepl Ihat (in peeservative) the throa t is marb led with whlre . the belly is
",hite ",'ith a dark bro""n transverse band formed of
marbling , and the donum is light brown with dad
brmon marbling
8rr~diltg cull. -P. f~-.rQI;s "calls from lbe fOKSl
floor in the da ytime. a loud. slow mllSical trill lasting
four seconds, repealed at irregular intervals , spaced
usually at least 0 ne minute apart" (O. B, Wake and
R. Etheridge. field n" le s al CRE). In Suriname , males
call from the ground or from twigs and leaves up to
one meter above the ground; the call is high and prolonged. resembling a c ricket's chirpi ng (Hoogmoed
1971).
TuJpoIu ( Figs. II C. 12D). -T....o lois h<lve been
taken from the ba;ks of Jdull femalc:s in Eeuador in
J une: lIlld in Surirwne in July (fable 4 ). An<.M:her 101
(AMS H 70995. in Stages 29. 30. 34. and lWl Sformcd )
is not from the ba; k o f an adull. M.S . Hoogmocd found
tadpoles o f this species in a ...-ater-filled fallen palm
petiole in Suriname in February (Polder 1976). Measurement s are in Table 3.
M im ic r y . - P . f<'lIwrulis and P . picla,' may be
Muellerian mimics. They are syntopie, brigbtly colored .
probabl y poisono us. ar>d resemble eac h olher when
seen from above.
D;....us~s. -Polder (1973) has discussed occurrence
lIlld lrealmenl o f cramp o f the hind limbs lIlld fungal
infections of the eyes in caplivity·.
,"·omrltcl(1l"r~. - 1 desig nale B~I 1947 .2. 14. 21. a
26 .0 mm male from Yurimagu as. Huallaga River ,
[Lo re to.] Pe ru , c ollec ted by Dr. Hahnel, as the
leclOlylX' o f Pro.,~,upis f~nr"'(1lij Boulee gcr 1883.
Th is 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 the
lowl and forest s of Gu yana, Surinam e . a nt! French
Guiana and of the Amazon drainage o f Co lombia,
Ecuador. Peru. and Brasil. I look a specimen al 1745 h
in 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
"",,,,,,,,,-0:
U>culi/iu.--COW MBIA.
Umbria (FM)\H
~m 1'< Pucn" A. i. (LAC M b t044 _49); EI
b I78 9 - 1IO); S
31
BUUETI."I OF THE ,....ATURAL HI STORY M USEUM OF WS A....'GEUS COW..T r
C.,..
No. 27
Phy floba lu rricvlur (Boulen~ 1899)
Ptliinta'\Jllal Poiso n-arro'" Frog
Fronli. picce II and Figu,," SA . 8F
juraion PasIo-PIleno .... is and MCJC(q·P\M:no A,..
r.... d. ( L AC M 61031 -)2) .
~/li: Florcnd l ( U S !" ~l
~no.
1~72-73); MOftlia (AS SP 2537l}-71). M~I4 : Bl>tfIMo,l,J
(f"MSH 81810). A_ _
: 7_7.2 km :\>\.' l.c1icil ( LACM
61019 .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);
1899b I'ro' llv . api. Iri=1or Boulcngt'r: 4S5-456. four .y n·
'~ptl: B:4 1941. 2. 14, 16- 19. Porvenir, Boliv... .
Puenn Ore. Rio Aguan,,, (KU 122291); Lirnonco<: h. (LAC M
72629-41. 72676-82)
PERO.-Pasco: Chanl i n. , I<COlaz i n Valley ( LACM
72047-4 8. 72050-S6); P,," de AlOC ar, [.cola, ;" Valley (LACM
72049). H" linucu: Rio P~ hilca near Bo"!u c Nacional de
[pari... (TNHC 36477 , 364110..'6483, 36490. 364911 ); MonIC
Alegre, Rio PachilC<l(A MNH 430 12) Sa" Morl''': S"".",: he
near Juanjui. Rio Huall"g. (WCS 2780); upper Biabo Vallcy
(AM:'\H 42658). Lortl/. ,. junclion Rim San,iaso and MlIraioOn
1920
(AMNH 4 2450 ); 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. Rio
MoronI. (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 tAMSH
43183.43415); cceneeoce RiooCnnla~a;and Tapil;ho: tAMS H
42565. 43249. 43251); 1'IlIi'"" (A",I:-;H 42295. 4J614);
co nn ucnct' 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 ncar
Lelieia (LACM 60974-75 . 61029-30); Eirunepr:. Rio Jurua
( B. l Ull and Klo.. 1952); Bcnjamin Conslanl (CAS·SU
11873).
GUYA.\lA.-R""''' .. "i: '" Acarah) M.....nlain•• W Jl'e'A
R;." , ( KU 69692·95); KIIYII...-ini landing (AMNH 4m 2);
be ·ct'n upper Ku)u wini and Kas, ikaillU Riw... . uppt r
E ,qllibo drain.lIe CBM 19 36 .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.-S orrinam..: Param. ribo (RM:-< H unu O.
Maro .. ij,,~: PocIOt'gocd<1t' (RMNH uneal); P. loemell tRM:-<H
150(1); N.......u GebcrglC (RMNH 7030. 7725. 8212. 8356.
8647, 8929. 9144, 9626), Saromaua: La n~a Snela. Cop·
pename Rtvier (RMNH uncal). B r oJ:.op ondo : Brown.bc r~
(MSH 149) ; Ligo n.... Gran Rio. S Tapoe Wlilr. ·vallcn (RMNH
unclll). /'I'kbril: Fredcrik Willcm IV.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); Yllliollt, 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.pock
uo 1393-99); Saill ( La IOS2_57); R ;"ie- ~ Ouaqlli ( La
1446); Mon l o.Jbao ( La 1582); K. ....(illilolillooull IMIIO«
Gcni:ve 1225.2).
Cleared and Jlai~d malt'rial. -LACM 60914-75 .
Ecuadnr . ]769 m.
1'".,II,Jb<,ln ITicolo •. Barbour lind Soble; 401.
D/'ji1lilioll. -Teech. pres e nt: h H:S basally ....eb bed :
dorsum black or brown with yellow or whi le media n
do rsal stripe. sometimes expanded to cover most of
do rsum; com plete yellow o r white late ral stripe; light
proximove mral calf-spot absent: venter yellow or ", hile
with or without black or brown marbling , or venter
blac k or brown with yellow or white ,pots .
Dia gnosis . -Ph)-f1obales Iric-olD' is 1I member of
lhe /"n ll1ra!is group. differing from all lXher members
of 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 donal
snipe , from P . e.pinosai in having a Iighl median
dorsal stripe. in having a complete light lateral str ipe
( Fig. I B; incomplete in P . t')pilW.wi. Fig. I A), am!
in larger sil e, and from P. f~11I(>T{/lis and P. zaparo
in havin g a light media n dorsal stripe.
DnuiptiOlI of syntypes. - The snout-ver n length ill
millimete 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. Maxillary
and premaxillary teeth are prese nt. Tbe firsc finger is
longe r than lhe second. All roes are slight ly fringed
a nd basally ....e bbed; the ....e bbing is best developed
betw een loe s II -III·I V.
When recently prese rved. the dorswn llIld lhe side
of tbe body were black. emblazoned ....ith complete
lemo n yello .... later al and median dorsa l stri pes. The
base of the upper arm, the lum bar region. tbe posterodorsal surface of the thig h . and che ventral surface of
the calf were verm ilion red. The venter and lhe rest
of the limbs were lemon ye llow with black marb ling.
or b lack wh h lemo n yellow , pot, (Boulenger 1899b).
In pre serv ative , the ground co lor of the dors um
and o f the ,i de of the body is dark brown. A bro ad
lighc media n dors al stripe ex tend. from lhe veil! to
the snout , expa llding in from of lhe t'}'t's to cove r the
dorsal surface of the snout. This stri pe: narro ..... pos .em ly in two specimens. oontains a median row of
sma ll scattered dark spots in one: specimen. and bears.
a small Jl'O'Ierior break ill two specenens. A complete
light lateral .uipe extecds from the groin to the upper
eye lid and along lhe lateral border of the upper e )elid,
uniling Wilh the tight do rsal snow colo r in frolll o f l he
eye; tbe stri pe bears a small break on tbe left upper
1976
FROGS OF THE GESUS PHYllOBATES
e yelid.of one ~pecimen . 1lle d....k bro...n oon.ill l pound
color brolldly separate s. the median stripe from the
lateral Mripe in three specunen s: onl)' a narrow sliver
of ground color se parates the str ipes in the fourth
specimen (8 M 1947 .2.14.19 ). In this specimen and
one othe r specime n, the medi an and lateral stripes
fuse for a short distance behind the upper e ye lid.
iso lating the ground eolor o f the uppe r e ye lid as a
da rk spoe . A light stripe ex tends from the fore limb
insertion along the upper lip to the tip of the snout.
...here it meet s its fellow in some specimen s. The
dorsa l s urface of the forel imb is mostly light. The
doNII 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 dorsal
surface of the calf is dar\( with light specs. The venter
and the ventral surface o f the limbs are light wit h
d ark marb ling in three s pec ime ns, and dark with
mediu m-sized light spots in lhe fourth specimen ( BM
1947.2.14. 17).
D~scrip/ion of USNM matf'ri"J. - Thirty-seven specimens (US:'\ ~1 166897- 9(1) collec ted a few )"l:an ago
by l .A . Peters on the Cuenca-Macha la road in the
province o f Alua)'. Ecuador . on tbe Pacif IC slope of
the Andes (the ~ slope as the type local ity. aI an
elevation 300 to 4IXl m lower). resem ble (after preservation ) lhe type series . 1lle snout-vent length in millimeters o f 35 ad ults 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 o f the USl"M males have the third finger
slightly to strongly swollen . All the USNM spec imens
have basally web bed toes. as in the types . The color of
the muscles is very light. as in spec ies o f the genus
CoI(JS!f'/hus ,
The US :'\~1 specimens have light median doNtl and
complete lateral wi pes; in 3. fe...' specimens, the median
stripe is upandrd. covering lhe anle rior half o f the
do rsum and fusing with the a nterior half of the Ialeral
stripe . Th is fllSion ~mbles that o f one of the types.
( 8 M 1947.2. 14. 19). In most o f the USNM specimens.
the venter is d....k brown with light spots or marbling .
as in o ne o f the types (8M 1947.2. 14.17 ). In some
s pec imens. the venter i ~ immacu late ligh t. In o ne
specimen , the vente r is light wit h a trace o f dark
marbling, resem bling the venter of three of the types.
A h hou~h the USN~t spec imens resemble the types
in their color pattern. they differ in their color in life
( Frontis piece II). The US NM spec imens were dan
bro...n with white stripes and spol s. whereas the types
were black with yeuc w stripes and spxs and red flash
marks . 1bc USNM specimens have green bones; the
t}·PC specimens have 110I. bul cou ld have lost lhe pigment because o f their a~ . It is also possible that the
green color is an artifact. I allocate the USNM specimens to P. tricolor and include their meas urements in
Table I under that species .
3J
Stomach comems (one spccimen).-From USSM
16689 7: HYMES OPTE RA: FotmicidilC : ~I}' rmcci nac :
SoI~"''Psis ?
(2) .
NOIll~ncla/u", .- 1 designate B~l 1947 ,2.14 ,18. a
20 ,5 mm male from P orve nir, Bolivar. Ecuador. 1769
m. co llec ted by P ,O . S imons. as the lectoty pe of
Prfljth..,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 lhe
Pacific slope o f the Andes in southwestern Ecuador.
l .A . Peters collected specimen s eevt to a stream in a
dr)' area of high grass and pepper tree s I=SchinlU
or Capsic_?J.
Lo cu/itin.-ECUADOR
SoH.""
Po", enir tBM
19017.2.14.16- 19). Auoo.l': Rio Minn. 20 km W Santa Isabel.
km tOl on Cucnca·M.:h<lla ""'" ncar AbdOn Callieron. SW
Giron (US S M 166897-9 11. tbf>9(0): 2 km W Santi I. abo:!.
km 88 un Cuenca -Machata road (USKM 1669(1); 4 km S
Calavina. km 7 t on Cuenca.Machala ro. d (USNM 166899).
CII'<I",d ulld SI<li"f'd m<llf'rilll . _USNM t66897.
Ph)'Jwbat~1 ZlJptUO
new species
Sanguine Poison·arrow Frog
Fromispiett I and Figum. Ie. 8D
Holot)'Pf'. - KU 120669. collected 2 km west of
Canelos. Provincia de Paiiu ,u . Ecuador . 580 m, on
9 Jul)' 1968, by R .W. He nderson .
Pural}p n ( 17 ad ults . all from Ecu ador ). _K U
120670 . co llec ted at Sarayacu . Pastaza, 400 m, by
J .D. Lynch; UMMZ 12935 8. collected at the Alpa yacu. a small stream near Mera , Pastaza. ele vation
stated 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 Don
Tomb , 5 j m S Mo ntalvo. Pastaza. by G . H..ne ra and
R. Olalla; GO V 8932 . 89 34, collected at ~Iore t a .
below Montalvo. Rio Boboe aza. Pastaza . by. G . Herrera and R. Olal la; GO V 11935. collecled at the Rio
Villano . Pasraza . by unknown collector; GOV 9005 .
9007. collec ted at the co nfluence of the Rios Shyona
and Conambo. Pastaza . by R. Olalla; GO V 9190. collected at the Rio Ogl£lO. Pastaza. by R. Ola lla; GOV
9 212. collected at the Rio Conambo, Pesresa, by R.
O lalla; GOV 9380 . collected at the Rio Bobonaza E
Sarayac u, Pastaza. by R. Olalla; GO V 9 394 - 95, collected at Miazal. Moron a·San tiago. by M. Olalla ; GOV
9304, 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 Rio
Pucuec , S apo. 300 m . by' W. Clarle-~lac lnt )re .
Df'j i" ilio " . -Tee th prese nt; roe s ba~all)' webbed;
dors um red: lateral stripe usual ly ve.tigial or absenr:
white o r pale blue ventrolateral stripe; light proxilT}{'·
vent ral ca lf-s pot a bsent ; venter blac k with blue or
blue-gray marbling.
J4
B U~1N
OF THE.NATURAL HISTORY MUSEUM OF LOS A...'GEL£ S COUlWY
D iagnosis. - Ph)"lfoool n zaparo is • member of
the / ,,,,o,o/is group. differing from P. o",ho"~'i in
lackin g a light median dorsal stripe and in lv !!cr size
(Table I) . from P. boflle"ff ,ri ill having a red donum
in life (brown in P. boll/e/lgtri) and in larger size,
from P . n piflOs(); in larger size. and from P. tricolor
in 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 d lOlt
lana I fo ld. bUI differs from P. jtmoraiis in having I
red oc rsum in lift (black. or brown in P. fe moroUs),
in usually ha~ing the lighl lateral slripe (Le., the stripe
adjace nt 10 (he edge of tbe back) ves tigial or absent
(complete from groin to upper eyelid and prominent
in P. f tmorulis) , and in havi ng the: ligh l proximodorsal
lhillh -spo l small o r absent (large and prominent in
P.femoru/is). P. Zilpuro resemble s a sympatric species,
P. pun' ulus, in having a red dorsum in life, but dif fers
in having basa l toe webbing and in larger size .
Deseriplioll of ho IOl)'P<' . - The holotype is an adu lt
male , 27 .5 mm sno ut-vent length . The skin is strongly
granular on ee dorsum and the dorsal surfac e of the
limbs, 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 the
soo ut is subfruncate in dors.al bpe>C 1 and rounded in
laleral as pect. The canthus rostrali s Is rounded. The
Ioreal reg ion is slightly con cav e . The width o f lhe
upper eye lid is slightly jess than the interorb ital width
Tbe d iame ter o f the e)'C is !ilighdy grea ter than the
d istance from the eye to me l'lOiilril and twice the diamete r o f the tympan um. The posterodo l"§4l port ion
of me tympanum is concealed . 'Ibe finger d isks ;m
small relative to tbose of lhnJrobules . The fio.t finger
is longer than the seco ed. Tbe fingers are webless. 1be
larsa l fold is Slm ng l)' bent at its pro ximal e nd and
relati....ely short. extending along the dista l 33 percent
o 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 loe s !I· IlI· IV.
In life (notes by J.D. Lynch at KU ). the iris was
black with a bron ze ring around the pupi l. The dorsum
was br ick red , The side of lhe head and body was
black , The upper lip stripe was gree nish yellow . The
ventrolateral stripe was white . The dorsal surface of
the forelimbs was light brown with a few black spots .
The axilla was yello w-o range. The dors al surface of
the hind limbs was black with blue -gray mottling.
The throat and chest were black . The belly and the
ventral surface of the limbs were black . monied and
marbled with pale blue·gr ay .
In pre~ nl ati ....e. tlJf, grancjes of lite head and back
are EOkien-brown. separa ted by narrow black inter _
spaces . The side of the he ad and bod)' is black . A
.. hire .... entrolate ral snipe e xtends from lhe groin to
the ni lla. uniting donal to the Ioretimb insertion with
ue up per lip str ipe. " 'hich co ntinues 10 the oostril.
No . 27
The re is no light late ral stripe. The dors al surface
o f the limbs is gray . indi sti nctl y ban ded black . A
lon g-itud inal white srnpe extends along the vemrat
surface o f tbe forelim b from me axilla 10 the \\Tis!.
The postCfO\o'l:ntral surface of the thigh is gray w ith
white dots. The rest of lhe ventral surface of the limbs
and the venter are black . e xcept for \\'hile marbl ing
on the Ihigh and posterior belly.
Du criptio N of p<JratY/H's . -Measure me nts are in
Table I. T'hr snout is rounded in dooal a'flCCt in some
of 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 do rsum was red . There was a vest igia l copper
brown lateral stripe. The dorsal surface of the lim bs
was Hght brown with dark brown or black spots: the
limbs lacked bright spots. The axilla was orange . The
posterior surface of the thigh and calf was blue with
black reticulation. The throat was black . clouded with
blue. T he belly was black wit h pa le blue posterior
mottling. extendin g as a stripe along the lateral edge
o f the t:.::lIy til the axilla . In prescNative. the blue of
KU 1206 70 has become gray.
in preserv ative . lhe UMMZ paraty pes diff er from
the KU specime ns in It.aving a white spot on the proximodorsal surface o f the thigh and another covering
me axilla and the proximopo sterior and proximodon.al
surfaces of lhe upper arm. in ha ving me ventrolalcral
slripe incom plete (endi ng midway between the groin
and lhe uilJa) in UM MZ 129358 and no! visible in
UMMZ 90370. and in lack ing ventra! marb ling.
In prese rvalive, rl'lO'>l of the GOV para t) pes resemble
lhe ho lot )pc: . Two speci~ns ( GOV 9194 and 9 3(4)
have a promine nl .. hile spot cove ring the axilla and
dorsal base o f the upper arm. The dors.a1 surface o f the
thigh is either 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 and
prom inent ). The venlrolateral smpe is co mplete restending from the groin to the axilla. where it unites
with the upper lip stripe) or incom plete , Some speci mens have a prom inent or vestigial stripe on the canthu s
rostralis and upper eyelid . Five specimens (GOV 8929,
91XI5. 9 190, 9212, 9380) have a vestigial late ral stripe
adjacent to the edge of the back. In one specimen (GOV
9007) . the lateral stripe is com plete and prominent from
the groin to the tip of the snout; this condi tion diffe rs
from all otber parat}pe s and from the bolotypc. The
throat is enti rely dark: the belly and lhe ventr al surface
of lhe limbs bear light marb ling .
Distribution mul humtat (Fig. 15).-Elevation records are from about 230 to 1000 m. P . zaparo inhabits
the de nse forests o f the Pastaza and Xapo drainages of
Ecu ador , ea st o f the Andes; it is terres trial and di urnal
(lIOfes by J.D . L)'nch at KU) .
£ f)-molog)". - This species is named after the Ziparo
i ndians of ea' tem Ecuador.
/976
FROGS OF THE: GV,US PHYUOBAT'ES
pious group
This grou p includes seven species (Pltylloba/t'S boJiviamu, P.i n/,, ,ri , P . parv MfMS, P . pr /..rsi. P, pic/..s. p .
p..lch ri""C/.. l. P, .•maraxdin",I/.
Defi'li/lim : sno ut-vent len glh of ad ult s sma ll lO
moderat e ( 16.5-3 1.5 0101) ; toes we b tc ss: paired
stripe s (whe n 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
1920
sent . The venter and the ventral surface of the limbs an:
Iighl .. ilh bro w n marbling and spots; the ventral marlings are dsscrere in 89-90. btll cont inuous and closel) '
set in 91.
Nommcla/urr , - I design ate BM 1947.2 , 13.89 . a
26.0 mm female from San Carlos. Bolivia, 1200 m
elevation . collected by P.O . Simons. as the lectotype
of ProMherupi.• holill i<l""J Boulenger 190 2.
D;Jlriburion <l"d hobil<ll ( Fig. 15l.-Elevalion recORb are from 800 10 1200 m. P. boIivion..s inhabits
the Amal on d r'l1inage of Bolivia
~<llilks ._BOU VIA U Pc: San Cane-. Dc'" ~bpori.
about U "32·S.6TSS'W (BM 19047.2_1389_90) ; San ~o.
( B ~l 19-17.2.13.911.
about IS"OO'S. 6S"(l(l'W
P' o>lh.-rapi. boIi,·;o",.. Bouknge r: 397. lhftt .yn.
types: BM 1947.2.13.89-90. San Callos. [u Paz,I
Bolivia, 1200 m; BM 1947,2.13.91. San Er~ 'lo ,
( La Pal ,] Bolivia.800 m.
Ph.vl/ohalr. holivian,.., Barbourand Noble: 402.
D"fi"i/ion . - Teeth abse m; roe s webless: dorsum
dar k (in pre~n-al ive); 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 mem her of
tbe piC/..s group. differing from P. ing..ri in lacking
teeth. in having a comp lele Jigi'll lateral stripe (absen l
in P . inJ("';l, and in lacking a Iighl prox imovemral
calf- spot. from P , parvul".• in lad ing teet h and in
having a complele llght lateral slripe ( inc<.>mplcle in
P. pa,..,,,I,,.•). from P. P<'lrni in lacking teeth . from
P. p ier.... in hocking a light proximo ventral catf-spct.
from P. p"khriprc/us in lacki ng a lighr proxifllOdor,;al
thigh-spot . and from P. MtfuruJ(dinus in lacking teelh
and in having venlral marbling
D..scription. -MeasuremenlS are in Table I. The
skin is granuhll' on the dorsum. tubercular o n the
po~erior back and the donal !>Urf~ of the hind limbs.
and smooIh on the forelimbs. the verner, a nd lhe ventral
surface of the hind limbs. Ted h an: absen L The fin;l
finger is longer man tbe second. The toes are webless .
The co lor in life is unknown , In preservative , lhe dorsum and the side of the body are brown. A co mplete
white latera l 'lripe extend, from tbc groin to the upper
eye lid . along tbe outer borner o f the upper eyelid. and
sol1lClimes along lhe canthus rostralis to lhe lip of lhe
snout. where il meelS ilS fe llolli. A whne stripe extends
from the f.....-elimb insel1ion along lhe uppcT lip. ending
ventral 10 the nostril or connnuing around the snout 10
fIlCCI il> fellow . 1llt' ax illa and lhe proximovenlral and
proximoanlerior 'Ulface' of the upper ann an: .. hite.
Light malbJing covers lhe anterior . dorsal. and posterior
surfaces of the thigh. On the posterior surface of rbc
thigh. the marbling forms a lighl longitudinal undulating stripe in 89 -90. Thigh-sp ots and calf-spots are ab-
Phyf{l,b(l/t'S ingf"; (Cochran and Goin 1970 )
Brother Nicefom's P oison-arrow Frog
Figures 3C, 9C
1970.
Coehnn and Goin: 16_17. plale
2D- F. hoIoo:ype , CSS M 146ll46. A-no Dc'" Rio
~ado. C~...u. Colombia.
lH,.JroI>a~J in~ri
D..fini/ ion . -Tee lh presen t; toes ..-ebless; dorsum
darl ( in presen-ativt'); stripes absent; li/;ht proximo'o'e ntral ca lf·s pot present ; venter dark with Jigi'll marbl ing.
DioK" os ;s , -Phyf!o/>{I/<'J ingeri is a member of tbc
p;ctUJ grou p. differ ing fmm all other spec ies of lhe
gm 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 hind
limbs. and smoo th on rbe fore limbs. the venter , and the
ventral MJrface of the hind limbs. Max illll/)' and pre.
maxillary teeth are present. The finot finger i& longer
than 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 pt
for a white spoe on the ax illa and lhe proximodor;.al
surface o f 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 of
the calf (wmetimes ei\tending half the lenglh o f lhe
calO . and iII -defi.w ~ra)' marb ling on lhe belly (noloI) pel or on the emire ventcr and the venlral surface
of the limt>s(paralypes).
Rr //I/;oll.hip:> , _ P , ;" Ke d is c lose ly rela ted to P.
ptctus.
BULLETIN OF THE NATURAL HI STORY MUSE UM OF LOS ANGELES COUNTY
36
Di~trihuri"'l
and hahira' ( Fig. 16), -The only elcvation record is 4()') m. P. ingeri is know n only from
the type locality . in the Amazon drainll~c of Colombia.
uX'/Ililies.-eoLOMBlA , CllI/um i: Aserrfo. 20 km from
jUlIClion Rio. »c!IC.do and O neguaza. 5 km from righl bank
Rio 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. plale
11. 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: dorsum
red; incomplete blue lateral stripe: light proximovenrral
calf-spot usually absent; venter black with blue marbling or blue with black marbling.
Diagllosis . -Phyllob(l/<,s purvu/us is a member of
the pictus gro up. differ ing from all other species of
the group in having an incomple te light lateral stripe
(often disap pearing in preservative: the other speci es
have a complete light lateral stripe. e xcept P. illgeri ,
whic h lack s stripes), P . ptlfVU/US further differs from
P. 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 usually
lacking a light prcximoventral calf-spot. and from
P• .v"urugdinu.< in having ventral marbling in life.
Two specimens (LACM 72646 and 72655) of a series
of 26 P. parvulu.,' from Limoncocha have a light area.
yellow in life (notes by W,R. Heyer at LACM). on
the proximoventral surface of the calf. Un like the
distinct. white. rectangular or elli ptical calf-spot uf
preserved specimens of P. piCIIIS, the calf-spot in these
two specimens is indistinct. gray. and irregularly
shaped. Besides usually lack ing a calf-spot. specimens
of P . f}(lrvul us from Lsmoncoc ha differ from sympatric
specimens of P. pic/us as follows (the cond ition in
P. piclUJ is in parentheses): the lateral stripe is incomplete. often disappearing in preservative (the
lateral stripe is complete and dist inct in preservative);
the dorsum is red in life (black or brown): the dorsum
is strongly granular (slightly granular); the first finger
is longer than the second (about equal); maxillary and
premaxillary teeth are present (absent). In life , P .
punu/u.,' resembles a sympateic species, P. zaparo,
and an allopatric species. P. espinosai. which also
are red. P. ptlrv ulus d iffers from P. taparo in lacking
toe webbing and in smaller size (Table I). and from
No. 27
P. espinosai in lacking toe webbing (the toes are
basally webbed in P . tf,Ipuro and P . ".,pino.wi) .
Destr/plion . -Measurements arc in Table L The
skin is strongly ~ranular on the dorsum and the dorsal
surface of the hind limbs. and smooth on the forelimbs ,
the venter. and the ve ntral surface of the hind limbs.
Maxillary and premaxillary teeth are present. The first
finger is longer than the second. The toe s arc webless.
In life, in ,pecimens from Santa Cecilia and Limon eucha, Ecuador (notes by J.D. Lynch at KU and W.R
Heyer at LACM) , the d",.,mm was dark red (only the
numerous. c1o>ely set granules were red: between them
the skin was black). The ground color of the side of
the body was black. An incomplete Iighl blue lateral
stripe extended from the groin one -half to three-fou rths
of the groin-axilla distance . A li~ht blue stripe extended
from the forelimb insertion along the upper lip. ending
ventra l to the eye or nostril. A yellow axillary spot
extended onto the proximodorsal surface of the upper
arm. A small round or oolong yellow SPOl adorned the
proximodorsal surface of the thigh , The venter and
the ventral surface of the limos were marbled light
blue and black: the marbling e xtended onto the side
of the body . The blue and the black were 01 equal
extent in some specimens; in others, the blue formed
lhe ground color , Juvenile specimens from Santa Cecilia
were bluish black on the dorsum, and had less blue on
the Venter than did adults; the smallest juveniles lacked
yellow axillary and thigh-spots
In life. a topotype from Sarayacu. Ecuador (Frontispiece J. and notes by J.D. Lync h at KUj, resembled
specimens from Santa Cecilia, except that the proxi modorsal surface of the upper ann was light blue instead of yellow , and the proximodorsal thigh-spot
was absem. The types of P . parvulus from Sarayacu
and Cane!os also lack thigh-spots _
In preservat ive. the ground color is black or dark
brown. 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 LACM
specimens from Limoncocha, Ecuador). The ventral
marbling is indistinct and gray, and sometimes dis appears (i.e .. the venter becomes entirely black or
dark brown).
Nome ndaluT<, . - 1 designate BM 1947.2.30.89, a
22.5 mm male from Sarayacu, Ecuador , collected by
Mr. Buckley , as the lectotype of Dendrobates purvulus
Roulenger 1882.
Dinrlbutian and habila! ( Fig. 16).- Elcvation records arc from 330 to 1000 rn. P. parvutus inhabits the
forests of the Amazon drainage of Colombia. Ecuador,
and Peru, east of the Andes . Specimens from Santa
Cecilia were taken on the ground in fo rest (KU 11150714) and at night on the ground in a marsh (JDL 1226 at
KU); one specimen from Sarayacu was taken by day
next to a small creek in fo rest (KU 120664; notes by
1.0. Lynch at KU).
FROGS OF THE GElWS PHY LLO BATES
/976
Loculi/it's . -COLOMBIA. P..lI,_.nI: Sm l. R...... dl: 1m
Kofme•. Rio Guame. tO I SOb25-~8).
ECUAOOR._Sapo: M en ,UMMZ 9(371); Lo~<> (GO Y
8921); S .Iope Sumac<> (GOv 9382); ~ l m W I'I>crto Nap<>.
Rio Na p<> (JAP ~658); Rio Cocapioo.:-.cap<> drainage \CAS·SU
10337); 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 Asr io
(WED 35728); Puerto Lim . Rio A~uarico IKU 122252-(8);
Puerto O~ , Rio Agu;Uico (KU 122269). Pas/a.,.: Sarayacu
( BM 194 7.2 ,30.89 - 90 ; KU 120664); Cando. (B M
80 .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\lO
Puyo (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).
Cl uJrt'd and .lllI in ",J "'(IIl' ri"I , - LACM f!0980; KU
126309
PhylloboUJ pt'tt'ni new
spoc ie~
Peruvian Poison-arrow Frog
Pronnspjece II and Fi$ures 6 F. 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' . -U ST\M 166763. co llected at Santa Isabe l
(a village on the Rio Xevati. a tributary o r the Rio PiChis. Pachitc a d rainage . 3.5 km SE Puerto Bermudez. 80
km ENE Oxapampa: the specimen Wall taken s lighlly
upriwr Irom tho: village. accordin g to A . C. Th ere -e n.
in fill.), Depart amento de Pa'lCO. Peru. 4.58 m. on 16
lu ly l%ll. til G. R . Xconan .
ParatyPt'J (4 1 aduh~ and nine tadpoles , all from
Pc:ru). -KU 144348-50, 10popatal)pes. collected by
D,B. Ekkens: ~CM 924 14 and KU 144344-46. co lleered at
Pa ~co . by D . B. Ekkens; AMNH
43008. 4 3011 . 4 301 6 . 4 30 26 . 430 34 . collected at
Monte Alegre. Rio Pachite a , HuimU('(). by H. Ba"lcr;
AM NH 42 179 . 42 505~07 . 42.546 , collected at Achin amis a. Rio Hu allagu. 60 km SE Yurimag uas , San
Mart in. by H. Bassler; AM NH 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 rainage
above Contamana. Loreto. by H. Bassler; FM l\"H
56 248. COIJ«-led al Cerro Azu l, 24 km E Contamana.
Loreto . 500 m, b)' J.M . Schunke; MCZ 24443 (ad ult
and one tadpole). collecled90-138 lorn WSW Pucallpa.
Loreto. b)' W.F . Walker. Jr .; BM 1900.11 .27 .38 and
AM KH 17199. 17253-57. collected at Perene. J unin.
ROO m , by P.O . Simons (BM) and C. Schunk c (AMNH);
FMN H 36778 (adul1 and cightladpoles) and 36 779 - 85
(ad ul t ~). coll ec tcd in l he valle y of the R io Vi IOC.
Juoin. by F. W ylko wsk i; 8 M 190 7.5 .7.9 - 11, co l-
xevau.
37
J«-ted at La UniOn. Rio Huecamayo. Pueo. by .\tr.
Ockcnden .
Dtjini/ion .-Teelh present; toes webje ss: dorsum
brown. sometimes with small black ~POIS; com p lete
yellow 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 Irom P . boli>"illnUS and P . pulchriPt'CI.... in having teet h. from P . inilrri in having a
complete light latera l ~l ripe (absent in P. inxrri) and in
lack ing a li$ht prcximosentral calf-spot. rrom P. POl"
"ulll:l in having a complete ligh c late ral -.lripe (incom·
plete in P. pon'ulu s) and in having a brown (!,(mum in
liCe (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; ventr al marblin g .
Description of '10[01.1'(1<' . - The holu t),f'C is an adult
remale . 28 ,0 mm snout-ve nt length . The skin is moder.
alely to ~Irong l)' gra nular 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; vomerine
teeth are absent. The tip o f the: snout is )I,/blruocale
in don-a l aspect and rounded in laleral aspect. The
cant 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 is
1.2 times the distan ce frum the eye 10 thc nost ril and
twice the d iameter o f the tympanum. The posterodorsul
portion of the tym panum i.~ co ncealed. The ringer di sks
are small rela tive to lhoS(' or Dt'I/dmbtl/t's, T he first
fiegcr i~ kmge r than lhe second . The fingers are webless. The canal rold is ~l ighlly curved. e'lendin$ along
lhe di~tal b4 percent o r the disrarce bec...'ee n the inner
mc:t atan.altuberde and the heel fold . The rarsafrubercle
is absent. The toes are weble ss .
In life teeter slide by A.C . Thoresen; venter not
visible ), the do rsum was Iigh c brown. The side or the
body was hlack . A co mplete )'ellow late ral stripe
extended from the groi n to lhe upper c)'e lid, a long
the oute r border or the uppe r eyelid. and along the
ca nthus rcs tralis (whe re it was go ld) 10 the tip o r tbe
snOUI.where it mel its Ielksw. A white stripe extended
from lhe forel imb insertio n along the upper lip to tbe
lip of lhe: sno uc. ...'here il met che cantha! stripe. The
dorsal wrr....:e o r the: upper arm was gold . The dorsal
surface or the: forearm and hind limb " 'as lighl bn,)...n
with a fe.. ' small black ~poI s . The anterior surface
or the thig h bore a greenish was h .
In preserva tive. the dorsum and the sidc of the body
are 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 entire
anterior surrac e or thc snoul is gray , T he uppe r lip
~l ripe e\lends ove r lhe rore limb inser lio n i nto lhc
axilla. 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
of the limbs are purpl ish gray with black marbl ing
This ventral pattern extends onto the proximoanterior
portion of the dorsal surface of the thigh . where it
(<lOllS an inll isti ncl , pu!.
Nr>.27
at the National Museum of Natural H istory in Washinglon . D.C.
C"mred <I1ltl "w incd malerial. - A :>'l ~H 4.,016: FMNH
36783
Dnaipllon of p,mIlYp"s. - Measurements of adults
arc in Table I (four o f the paratypcs are juveniles),
Many paratypes have the second finge r almo st as long
a, the first. The KU paralypes have a long tarsal fol<1.
extending over the db tal70 to 80 percent of the distance
Pttytlobates pictus (Bibron in Tschud! 1838)
Spo t-legged Poison-arrow F rog
Figures 3A -B . D -E, 9 D-F
helwee n Ihe inn",r mCl3lar;al tubercle and the hcc! fold;
in these specimens. the tarsa l tubercle is slightly to
moderately developed.
ln life. in specimens from Nevati, Peru (notes by
O.B. Ekkens , in /ill .), the dors um was brown with
small black spots . The side of the body was black . The
latera l stripe was yellow -gree n; the upper li p str ipe
was lighte r and greener. The ve nter and lhe ventral
surface of the hind l im bs were blue wirh hlack
marbling.
In preservative. lhe sl ripe ~ and the ground co lor of
t he venter arc purplish gray to lig ht brown. The color
pattern of the para typcs differs from tha t of the ho!ol ype
a, follows : BM 1907 .5.7.9 has broa der dark ventral
marbling . BM 190 7.5 .7 .9 _11 ha ve broader la ter al
stripes A M1\ H 42507 and 421 79 have stripes so
broad that only a narro w. sometimes broke n. da rk
brown line remains bet ween lhe pu rplis h gray of the
stripe and that of the lowe r side o f the body . T he dar k
ventral marbling sometimes is reduced to dis crete
spots.
Tadl'0//',' ( Figs l IE . 12Fj ,-One tadpol e. in S tage
28. was taken between 9 and 14 Aug usl 193 9 from
the back of a 28 ,0 mm adult female (MCZ 24443) .
betwee n 90 and 138 kill we'l northwesl of Pu<:a llp<l.
Peru . Another lot (numbering eight). in Stage 26 .
was lake n on 28 October 1940 from the back of a
25.0 mm adult male (FMN H 3( 77 8). at 1600 h. after
a rain , in forest in the valley of the Rio v uoc, Per u;
the tadpoles lay w ith their heads median and their
tails lateral relative 10 the ad ult's hock. Measu reme nts
are in Table 3
Retationships, - p , petersi proba bly is closely related to P. h"lh'iall//< and P. p ulch rip"CIUJ.
Stomach COIlIelllS (two spec imens).- From FM NH
3678.~ : H YMENOPT E RA : Fon n ic idae: Ph/'idol/' sp.
( I ), Ponermac (I ): HE M IPT E RA: Tingidac": fragments . Fulgnridae?: fragments.
From AMN H 43016: H Y ~I I::'.t,;O PTER A : Fonnicidae:
Azteca sp. ( 1), Myrmecinae (ll; CO LEOPT ERA : Curculionidae (I ) ; IS O PT E RA : Na .l ulitermn? sp. ( I):
DlPT ERA: Dro sop hilid ae (9) ; ARAC HI\ IDA: Pisauridue (35) .
Distribution alld hahitat (Fig . J6) .-Elewtion .....~ ­
ords are from 274 to gOO m. P. pel('T"i inhabi ts the
for ests of the Amazon drainage of Pe ru. west to the
eastern foot hills of the Anlle s.
EtYllloloKy . -Thi, species is n.. mcd after the late
Jame s A. Pe ters , curator o f amphibians and reptiles
1838
184 1
1848.
1l\5l\
1864
1864
1864
1882.
1883.
1887.
1925
1926.
1926.
19 52.
1':152.
t955.
1957.
t':l6 1.
1965.
1970
1970
1970 .
Hy/aple.,ia pi, ·la Bibrnn. T....'hurJi: 21\. 71. two ,yo _
types MHNP 4910 , San ta Cr uz , [Santa Cruz.]
Bolivia.
UOIdrobatn piclus. Dumeril and Bibrnn: 656.
Drndro/mlr.pictus. Guichenot: 119_1 20.
Hy/aplesia picla , Gunther: 125_126.
DOIdrohale> '1igarimus (p<lrt) . Steindaehner: 257_
259. f,gs. 2. 2a-d
Drndroharr.f hr",..ra lu.f Fitzinger. Sieindaehner: 258259 . two syntypes )\;HMW 3818: 1_2, [probably
Sao Vicente. Meto Grosso,1 Brasi!.
Dendrohotes <'Ut'nemi, Filzin~,'er. Steindachner: 257258. plate 13, figs. 2, Za-d. four syntypcs: NHMW
19190:1-4, [probably Rio Mamore . Rondonia. ]
Bra,il
DOId",h mn Iri"ill<l/US (pal'\). B\lUlenj;er: 144- 145
Drndro/nur. hll l",r/i Boulenger: 1\36. plate 57. ji g.
4. se"en syntypes BM 1947.2 .15.14 _20 , Yuri maguas, Huallaga River, [LoreIO,) Peru.
Drndmbu les bra, ·,'mu. Cope: 53 _54 , hoIOl)'pe:
ANSP 134 14. Chapada I~ Ch a pad a dos Guimaraes,
according to BOKermann 19661. Mato Gros.'\{), Brasil
Hylapir.fia jlaw'p irla A. Lutz: 139. 'yntype: USNM
96986. Belo Horiw nte. Minas Gera;" Brasi!.
Dm"rofmlrs triviumu.< (1"'1'\ ). Miranda Ribeiro: 224
Dendrafm/C$ broauws . Miranda Ribeiro: 179- 180.
,nt-fig. 96, plale 20, figs. 3, Ja -b
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
D <,~drob<IICl pi('/us. B. Lutz and Klo", : 670-671.
Dr~drubm.... f/"",'p i'·III.,. Cochran: 8-':1 , piale I.
figs. G. H
D,'ndro/",l<'-< pi,.,u.,·. Dunn: 78
Dendroba/t>,f jla vapir-lu.•. Cochran: 107. pla'e 42
(c" l" r phOIO).
Dr~drolmlC$ piN" .< gua"U1", n, i.' Healwole. Solano .
and He. twole: J50- 3Si, hoiot}'pe: MBUCV 31 t2.
Alliplanicie de Nuria, & iivar. Veneluela. 100-25 0 m
Dr'ldrobd/cs h<lhaeii (pa"). Co<:hran .nd Goin: 17-1 9.
plale 5D-F (only FMNH 61789 is lIot P. pielu., ).
Deadmholr.• tril·iu"w.' (pan ). Cochran and Goin: 16
( no' the plale; only USNM 118681_82 are P. piclus ).
Phyllobau,f jemamli. ( part) . Cochran and Goin: 42
(11m lhe plale: only USN M 14681\3, 141\8 66. 147268
are P . pi,·/u. )
De/initiol1 . - Teet h usually absent. but present in
rome specimens; toes wcbless: dorsum blac k. brown.
or gray . sometimes w ith gray or bro ..vn spots or marbling. sometimes wi th paramedian rows of yellow
SpOIS: complete yello w. 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
39
FROGS OF THE GENUS PHY LLOBAT ES
or blue or white with black marbling (rare ly. verner
almost immaculate while ).
Diag"o,i.,. -Phyll"bal n piclu.l is a member of tbe
ph-tu, group. differing from all other species o f the
group except P . ingeri (and rare ly. P. parm!u,) in
having a lighl proximoventral ealf ·spot. P . piC/us dif.
fer s from P . ingeri in usually lack ing teeth and in
hav ing a eomple le light laleral stripe (ab'iCnl in P .
ingeri) , P. pinus further di ffers from P. parmlu< ;n
usually lackin g teeth , in having a black . brown. or
gray dors um in life (red in P. !nln'ulus ), and in having
a complete lig ht lateral stripe (incomplete in P. pan'ull,, ).
from P , pelusi in usua lly lacking teeth, and from
P. smllragdinu, in usually lacking teeth and in having
ventral mar bl ing . P . piC/us resemble s a sympamc
spe cies. P . femorali s . but differs in smaller size
(Ta ble I). in having lhe upper lip stnpc not meeting
the canlhal stri pe . in having a light proxiruovcntral
cal f-spot. in baving lhe firsl finger abo ul eq ual to
the second (P. femorali s has lhe fif'l finge r lo nger
than the second ), in lack ing webbi ng (P. femoralis has
basa l toe webbing ), and in usuall y lac king teeth
Dl'scril'/ir", . -Measurements are in Tables I and 6
The skin is slightly to strongly granular on the dorsum
and the dorsal surface of the limbs . and smoolh on
lhe venIer and the ven tra! surface of the limbs . Ma xillary and premaxillary teeth usually are absenl , bu t
are present in some specimens ( see below). T he firsl
linger usually is about equal to tbe secon d. but some limes i> sl ightly longer than tbc second. The roes are
web less
Although all popu laliuns of Phyllobales piClu., have
a light proxim oventral calf. spo l. this species varies
geographicall y (but not str ictly clinall y) in snout-vern
le ngth (Table 6) and color pallern . For co nve nience .
the variat ion in color pallern is described be low as
six patterns. which are not intended 10 represent taxa
Populalions whose members bear some of these pal.
terns have been described as species or subspecies:
so me of these populations. and some other po pcra lions lhal never have received a laxonom ic name . may
be spe cifica lly distinc t from P. pic/u s. Because of
lhis poss ibility . [ have indieat~..-l 10 which populations
published oame s apply .
Pallern I ( Figs. 3D. 9D; inclu des Dendrohate.•
bracelll" .• Cope 11\8 7 and Hylu"l esia /l av"" iela A.
LUI? 1925): Specimens bearing this palter n are known
from 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 is
characterized by a pair o f paramedian rows of ye llow
spots (Fig , 3D); the other pauerns sometimes have gra y
or brown dorsal SPOIS or marbli ng. but never ha ve
ye llow spots as in Pancrn I. The thigh-spot usuall y is
long (Ir is short in the holotvpe of D. bruemlu.' Co pe
from Cbapooa) . Frogs from Belo H",i?onl e (lhe lype
locality of Hylllp!e, iu jlIl,·"p;"la ) have a light ven Ier
heavily marbled with black (Fig. 9D) , The black ventral
marbling is slightly reduced in specimens from lhe most
westerly known popu lation with Pattern I (EI Porto n).
The marbling is greatly reduced in spe c imens from
J1<.,pulalions to the nurth (Cha vanlina. Cavakanl i) and
west (Chap;uJa ) uf Belo Hortz ontc. In specimens from
Chapada (the type locality o f Dendml"'lCS' brl'CCIl/!H
Cope). lhc venIe r is near ly enlirely light. with a few
black spots on lhe lateral ponion> of the belly; the dorsum is tbe same as in specimens from Belo Horizonte
ln life. in specimens from Belo Horizonle . Brasi l (lhe
type series of H . /lIW""i<'W: notes from B . Lut z 1952).
the iris was black. T he dorsum was black with ycllow
spots arranged in a pair of paramedian row s, The latera l
and upper lip stripe s were yellow. The do rsa l surface of
the limbs was black with yellow spots . T he thigh- spot
and the calf-sJ1<.,t were red-orange . The vent er 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: S UR[:-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,he a. 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
Pallern
Ran~e
re
'".1.'es
n
22.5-3),0
21.5- .'1.5
17.0-2 9,0
16.5-2 5.0
20.0-23. 5
Female,
M"I" ,
Rang"
27,5
2(>5
24.1
21,7
21.5
""",
",
n5 -29.5
21.5-29.0
17.0 _25.5
16,5-n5
20.0_2l.5
2~. (>
25.6
n3
19.8
20,8
,
u
'""
6
Range
28,0-31 .0
HO _31.5
19,5 -29.0
19,5-25.0
20.0-2.',j
29,4
27.7
25.7
22.5
22.3
4('
No. 27
BUUETI.V OF THE NATURAL H ISTORY MUSEUM OF LOS A.\'GELES CO UNTY
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 this
panern are known from
Vicente (, tate of MaiO
Grosso). Brasil. Sleindao;hner', ( 18M) co mbined Iocaljl~ li, t o f Drndrobates rucnemis and D. braccana fail,
to a,sign loc alitie s to e ach fonn. Steindacbners label
with 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 loc alities mentioned by
Steindacbncr. Sao vjceme is near Cope ' , locali ty of
Cbapada (B . LUll 1952), In thi, pattern. in preservenvc .
the dors um bears indistinct lil!hl spots not in rows (similar 10 the SP01s o f many specimens bearing Pattern 4 ).
The verner is lil!'hl. with a fe w black spots on ils lateral
portions (Fig . 9E ).
p...uem J tFig s. JA . 9 F; includes H.I"lapl~sia picto
Bibre>n i" T scbudi 1838 and Dt'lwrooott' s t'uc"t'mis
FilZinl!'er'I/ Sreindcchner 1864 ): Specimens bea ring
this patte rn arc kno wn from Bolivia (nort h. nonhw esl,
and west of EI Po rion. in the dep artments o f Santa
Cruz . Benl, and La Paz). Bras il Ithe terr itor y of
Rondonia. the Siale o f Para . and a l Urucum near
CClnlmoo on lhe Brastlian-Bol ivian border ). and Sunname . In Ihis pattern. tbe dorsum lach spots; the
thigh-spot is long ( Fig. 3A ) and sometimes broken.
and the venter is d ark with 1i!!lu marbli ng ( 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 were
while . The axilla . the thigh. spool. and the calf-spot were
red-orange. bordered with black . The j unction between
the rectang ular proxima l and the o blong dista l portion s
of lhe thigh,spll t was yello w. The venter was grayish
black with whitish spoe, and marbli ng. In specimens
from PosI Gon ini and Sipa li",'ini. Suriname (noieS by
M.S . Hoogmoed al RM:->H). the OOnum ....as black wjth
gray marblin g. The lateral str ipe:....as cream . The upper
lip stripe was while . T he ax illa. the postcrovemral surface of the upper arm. the thigh -spot, a nd the calf-s pot
were yellow . The venter and the ventral surface of lhe
lhigh were black ....ith blue . greenish white . or bluish
while marbl ing.
Pattern 4 (Fig. 3B : include s Dt'lldrobol~s hoh" d i
Boolenge r 1883): Specimens hearing Ih is pane rn are
known from most of Amazo nian
Ecuador. and
Colnm bia. and in Amazo nian Brasil in lhe lerrilo ry o f
Acre a nd the slale of Amazo nas. These relalively small
frogs (Table 6) lack rows of yello .... do rsal spools (al·
lhougb they 'Ome limes have indistincl gray dorsal
~s) and ha\Il.' a short Ihigh- spol (Fig . 3B); lhe venIer
is dark wilh lighl marbling (in pm.el'\-ative . the marbling is conspicuous on the belly and indistinct on the
throa t). In life , in s pccimen.~ from Tere~inha . Brasil.
Sao
Peru.
ncar Leticia (my notes}, tbc iris. the dorsum. and the
side o f the body were black . A complete yellow lateral
stripe ex tended from lhe groin 10 the upper eyelid. along
the o uter border of 111(0 uppe r eyelid. and along the:
can thus rostrali s 10 ee lip of the: snou l. where il mel ils
fellow . A while stripe eXlcnded from the forelimb insertion along the: upper lip. ending ventral to the eye or
nostril. Tbe dorsa l surface of the limbs was lighl bro....n.
exccpl rbe proximal ha lf and the posterio r plmion of the
dislal half of the lhigh. wbich were black; the black
color surrou.ndcd a soo n oblong orang e spot o n the
pro ximodorsal surface of the thigh. Blue spob oma meraed tbe posterior surface of the thigh . An orange
uillary spot united wilh an orange ventra l stripe e xte nding alo ng either the proximal half or the e ntire length of
the upper arm. The ventral surface of the forearm bore
white marbling. The ventral surface of the hind limb
bon: blue marbling and spots , except for the proximoventra l half o f the calf. which bon: a rectangu lar or
e lliptical oran goe spot. In one unusual specimen (LAC M
6 1006. one of 12 specimens from Teres inha). the calfspot was blue. The grou nd colo r o f the venter a nd ve ntral surface of the limbs was black , The thlOat bore blue
marbling and spots. The breast and belly bore ....bite
marbling. Spec imens from Pixuna , Brasil , near Maneus
(my OOIes). J'C'sembled the Teresinha specimens, except
lhal the iris WlLS black Wilha gold nng around lhe pupil.
the back was blac k ....ith gray spou. tbe lateral smpe
was gold, the dors al surface of the calf bore IWO da rk
brown tra nsverse bands. and the axilla and calf-spot
were yello w-orange . In two juveniles. the:dorsal surface
of the upper ann ....as orange·gold. and the dooal sur face of the forearm . ca lf. and larsu s was green ish gold .
Specimens from Santa Cecil ia and Sarayacu , Ecuador
(noles by J.D, Lynch at KU). resembled the Teresinha
specime ns. exce pt that the iris was bronze. the dorsu m
was black. brown ....ith black spots, or gray wilh black
spots , the lateral stripe 1'."3-' cream or yellow. the Uppel
lip siripe was blue-green in one specimen. the dor1;:al
surface of the calf and fore limb hore brown or bhd.
bands and spots. the axilla wa s yello...' or yello ...'·
orange . the thigh-spot and calf-spot were yello w or
orange. 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 this
patter n are known from paris of Amuon ian
(T sjcvemeni; color s lide by A.C . Thoresen: l\"evali: KV
144351-60. LACM 9241 3 ; Rio Pechitea near Bosque
Naciona l de Iparia: TNHC 3(4 69) . "These specimens
resemblc frog s bearin g Pallcrn 4 in s ize and color patlern (( he Ihigh ·spo t is shon ). bul differ in lhe following
characleristic s: the dorsum is Slmngly granular . leelh
are pre.cn t. lhe head and back hear small spols (some·
times panly fu~ logelher and somelime s arranged in
Ihree or four longilud inal rows) . lhe calf.spot ellends
onlo lhe disla l one- founh or o ne·lhird of tbe <lnr....1 sur·
face of thc thigh, and the entire ven tcr (includ ing lhe
Peru
/976
FROGS OF THE GE.,,"'US PHYLLOBA TES
Ihrtl;ll) and most of lhe ventral surface of the limbs are
lighl with dark marbling. Of a series o f \i~ specime n\ of
P . picfu, ( r.' HC 3646 1. 36469-70. 36476. 36482.
_\6.197) I..ken by J. P. Bogart at d ifferent ,Jate~ in 1967
and 1968 o n lhe Rio Pachitea neil!" tbe Bo"'lllC ~ac ional
de Iparia,
one (Th HC J6.1691 ha~ Pattern 5 and
five have Pattern 4; I do no( know if all tne..e 'J'«imens
came from thr ..arne deme . In life . in ~pecimen~ from
Nevali ( notes by D.B . Ekken ~ in li ll.) <UMl T~ io.'Cnle ni
(colo r slide by A.C. Tboresenl.
lho: dorMim "-a.~
orang ish brown .. ilh light olive bro" 'n ~pot:s (sometimes
arr anged in ro''''S). The lateral and upper lip Mri pe~ were
yellow-p-een or white . Tho: dorsa l surface of tho: limb"
..'as lighl ol ive brown. The a~ iII .., lho: thigh-spot, and
rbe calf- spot were yellow, yellow-ora nge. or orange.
The venter "'as blue with black marblin g ,
Pattern 6: This pattern is char acterized by a small
calf-s pot, and includes tbree subpauem s. Subpanern 6a
(includes Dem/m!l<Iln piaus KlUlYWWIlJis Heatwole ,
Solano , and Heatwole 1965): Specimens hearing this
subpattcrn are known front rfc sl"le uf Bolivar in Ve·
nezuela and from Guyana . The lighl ventr al marbling
is reduced to a few spots on lhe belly and tbi gh~. In life.
in a spec jrnen from Hecenda San Felip:: ncar Miamo ,
Ve nezuela (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 were
yello w. The tbigh-!opOl I..... red-orange. exce pt for a
yellow 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 bearin g this subpallem are
known from Guyana . Tho: thign..spol e ither is long and
unbroken or is broken into ... shon proximal ..pot and ...
long d is....1 )po(; the: venIer b...s ...ell -developed lighl
marbling .
Subpanem 60; (Fig. 3E): Spec imens bearing this subpattern are known from Gu yan a . The tbigh-spot i~ long
and unbrok en: the la te ral stripe: is rel ~ li vel y broa d
(Fig.3E).
Bru diflg coif. - T he c a ll of spec imens bea ri ng
Pattern I has been reported as "a piping cry, boowithoowlr" {Gans 1960, El Porton ), " 3 rather loud, very
persistent peep .' also a "trill' (B . LUll 19~2. Belo
Hortzor ne). and "beginning with a roll, p-r-r-r-, then
a soft whistle repealed hal f a dozen times and ~cry
high in pitch" (Coc hran 1955. Bclo Horizonte}.
T//dl'0lrs (Figs. I IF. 12G)._Two lOb have been
taken frum lbe backs o f adult males in Suriname in
June and in Bolivia in December (Table 4). A third
lot (MCZ 29825. number ing t..'o. in Stage 25) probably is from the back of an ad ult, altho ugh there i~ no
record of this, Two ot her 100s (U!-l!-IZ uncatalogued
and UMM Z 58949 . in Sta ges 38 and 42 . taken in
Boliv ia in August and Dece mber) are nol from tbe
backs o f ad ults. Me~remenb are in Table 3.
TOQfhlfd sP/fcimlffl s. - C.F. W...lker sbo 'ed me ...
specime n of P . pictus from tvoe , BoIivi (U M!-lZ
Peru.
Peru.
5894 1). from wh ich be had removed one of lhe malil.
lary bones . T in)' teeth were "isible on this bone. bUI
not o n the other maxilla..... hleh was lefl in place .
Thb )l.Igp:sted tholt m.tensibl.)' lOOlhles.s specimen) or
P. piaus hast: concealed teeth. I cjeared and !>Uined
four such specime ns: Patte rn 4: Tere vlnha. Bra s il
( LAC M 60977 , J~1S 1b80 ): Patte rn I : EI Porton,
Bolivia (LAC M 6(976): and Subpattern 60;: KUl\lpung
Rbe r. GU)';tn... CLACM 609711,: the maxillary and pre .
maxillill"}' bones lack teeth . Thus. at least !oOI'ne osscn sibly toothless specimens o f P. pic/ us do lad teesh .
In some case~ (e.g. • specime ns bearing Pattern 5,
fro m "' eval i, Pe ru ), all presc rved spec imens n f P .
pi, ·IN.< from a given locality have tee th, but leelh arc
not unifonnl y developed in all ; tee th are pro minent
and easy to see wilh a dissecting mic rosco pe in some
specimen s and tiny and difficult 10 see in olher slX'\: i·
mens . In other cases. some specimens from a given
locality have teeth. "nd other specimens from the same
lo..:ality lack teeth (at least. teelh are not visible in
the intact specimen s). The se data suggest that P . pictus
once wa s denli gerou s lik e most oth e r s pec ies of
PhJ llolwlts , thaI it is evolving toward a tooth less
co ndition. and Ihat this evolution ary trend has proceeded al diff erent rates in differe nt populations of
P. pic/ us .
Toot hed specim e ns of P . pi<'tus are known only
from Peni and Bolivia, from lho: following locali t~;
PERU: trail between Satipo and Puerto Ocopa ( ~ IL/.
24 433; LACM 60 979: the la tter was cle are d and
staine d) ; Lu is i...na ( MCZ 58 413); ~ eva li ( LAC M
92-tl 3; KU 14435 1-60). BOLIVI A: Santa Cruz (i n
the liCries BM 1904. 10.29.75-82 , some specimens
have tee th. and o thers lad tee th); B ue na\· i ~ta (B M
1927.8 .1 .49 - 5 1 have teeth; UMMZ 665 12: one specimen has reeur. anol her ""ilb the same museum number
lac ks teeth ; four u liler UMMZ speci mens and si ll
AMSH 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 specimen s Irom
the 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 cleare d and slained) lack visible teeth .
Swm//(·h ('on/nilS {two spccimens). _From LACM
60976: HYMENOPTERA : Fcrmicidac: Ph"idollf sp.
(2 ). Compo"o/uI sp, ( I ), Do rJffl Jrmu sp. (2). a lso
s ix unidennfiable ant) , Mymaeida e (I); COLE O PTERA : St"ph )'linidac (l J. Curculionidae (3,: HEMIP.
TER A: Aph id ida e (6 ): DIPTER A : Dro sophilidae:
Drosophila? (2).
From LACM 60918 : HYMENOPTERA: Formieidae :
D .-.Jichodlf rus? sp. (I ), Pltlfidol lf spp. ( 4): ISOPTER A:
....·~lI1illf~J (39) .
....omlfffCJo/"rlf . -J de ~jg nalc MHSP 49 10 (one of
1....'0 S}'nl)"pes bearing tho: same museum number. the
BUUETIN OF TH E NATURAL HI STORY M USEUM OF WS Al\'GELES COU.\ 7 Y
PilTill«loly J'C is a 24.0 mm femllle), a 22 .0 mm ma le
from SlInta Cruz, Santa C ruz. Bol ivia . lIS the JecMy J'C
o 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 from
Y wimll~U3s . Rio Hual laga. [Loreto,) PnU, coll«ted
by Dr. Hahnet. as the lectotype of lRnJ rob m t s Nlhnt /i
BouIenger 1883. I designate SHM W 3818 :1. a 2b.5
mm female. proba bly from Sao Vicente, MaIO Grosso.
Bra..i1, co llected by J. Nan erer , as lhe Iec l<ltrpe of
[Jtlldrobult. bcuccalU. Fiu in ~-.:r in SleindOIChner 11164
I <.bil;nate S H ~I W 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 G uyana .
S ur iname . Fren ch Gu ian a , rne G uianan regi on of
Venezuela . most of Ihc Amawn drain age of Colo mbia .
Ecu ador . Peru . Bo livia. and Bras il, and l he north ern
Parag u a y - p.aran~ drainage of Bolivill and Brllsil so ulh
10 about lifS lalil ude .
T he type loc ality' of P. pier" . (T K"hudi 1838) is
Santa Cruz [Bolivia] . bUI D umeril and Bibmn (1 84 1)
l'llid tha I lhe types CiIIlK' from Ch ile . G uichCoot (1848)
could not affirm tbe prese nce of this species in Chi le.
E .R . Dunn (in J;II .• quoted in B. L Ul l 1952 :( 11)
bel ieved thatl he types came from Bolivia . G uilX (1950)
listed the type locali ty' as Santa C ruz [Boliv ia ]. P .
piC-flU has llOl been report ed from Ch ile si~ Dumrnl
and Bibro n's wor k. P. picrUJ i\ a Iowlllnd Amazonian
'Jl« ies; it is unlikely th at il occ urs in Ch ile .
Loca !ifil'. :-cQW.\IBIA. .•/~fII: 42 ~m S. 22 Lm W Sm
Man in t MVZ 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 M ico. Ma; -..a
t USS M 144962_67); Cana Guapaya. Mllu rt' nli IUSl'oi M
14726'). A..... tona . : 7.2 Lm ,",W 1.elicia (LACM 6 10(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 li bre.
Rio Asuark o (KU 112287-90): L.,rclo (l AP 3'S8j, PU.IUU':
Sarayacu (KU 120671_72); Alpayacu ncar Mera (UMMl
<XJ:w,I)
PERO.-1Al'?tf1: Lago de Mirano. moulh Rio Napn(AMNH
43156. 43IS8_S9. 43162- 64. 43166- 67. 43172_76. 43178_
81. 4l2 ' I _' 2; UMMZ 89983); conOuence Rin. C...nlaya
and Tapiche (AMt"H 42773, 4299~ . 43, S7); divide helween
Tapid w: and Ul<>quinia drainascs (A M)';H 42474. 42691.
42698. 43372): Ollama near Qucbrada Ca. hi"")a (A.\IS H
42977- 80); Rio Pi>qui (AM!"H 4J2 7JJ; uppe' Rio Ila) .
C AM~ H 4.'-4(7): C hiboya tAMSH 42070); IQuilOt CAMS H
Rio Cu.haNlay (AMS H 42372.
42209); Pampa He
424~): Rio Tamal'a tAMS H 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. Rio
Hualla, a tB.\' 1947.2 . 15.14-20); Dumo Sanla Clara and
Campo SantaClara. nearOreliana tUSS .\t 127201. 127203--08.
127932). S... .U;Uf;. : Achinami.... Rio H..alla, a (AMS H
No,27
42375, 42WII- (9 ); Tocacht CA MS H 4272 1); S....andlC
near Juanjui. Rio Hllalla, a ( WCS 2757- S8). H" I " a, ..:
.\Ionl(c Alegrt . Rio f'xhi'ea (AMS H 43009. 4;\(36); Rio
PachilCa near Bosque Nac:ional de lparia (TNHC ~ I .
36469_70. J6476 . J6482 . J6497) . J ••;.: Irail bocl..-ccn
Salipo and J>uc:no Ocopa (MCZ 24433 ; LACM 1(0979);
Owo:Jwnayo tAMS H 42892-94 ). Pasco: Sewali (LAC.\1
92413; KU 14.1351--(0); TS~TC1Ii (color slide by A C.
~n). C.W" Luis.iana. Rio Apurimae IMCZ S8413J.
BOLlV IA. -S".,a Cral : Santa Cru 2 ( MHS P 49 10;
CM 2462-63 : BM 1904 10.29.75-112); Buenawista ( A.\l ~ll
22637_38. 3J9S9. J.I075. J9S62-63; 01 4l11_27 . 4387.
4416. 4423- 24. 4444; UM.\IZ ~35-40. 638811. 66507-12 .
66514. ~-45 . 66588. 66592; USS M 118701-03 : .\ICZ
128S9; BM 1927.8. 1.46-SlJ; Rio Coloralo (0 1 4308-09 .
4311-1 9): Rio Surulu (CM 3812); SarilOjaI (USSM 14(598):
EI Porron (CM 36155; CAS 97762_63; MCZ 29813-2 3•
29825; LACM 44399. 60976). Br"i ; Gua}'a' '' ·.\Ier;n. Rio
M~more tCM 2667; USt"M 123% 7; BM 1939.1.5 . 1): Ru·
rn:nal>ll<jue (UMMZ ~7SSO. SM940, S'94' ); Cavina. (UMMZ
58950): Ivon (UMMZ ~ 8 941, 5M949: .\ICZ 10096 _97);
belwcen Reye. ~ nd R u m: n ~ba<j 1lC (UMMZ 58944), III PUl:
lower Rio Bopi (UMMZ S7549): Huach; tUMMZ ~ 8945.
58951): Tumupasa tUMMZ 58942); upper Rio Bcni below
Huochi (UMMZ 5 M~43 ) : Co""ndo tUMMZ 5M946); hiama.
(UM.\l Z ~8947: USNM 147221: MCZ 15151, 15153); San
Fermin (Ande...."on 19\)(',).
BRASIL.-A.III" U'II" " Rio Tacbila near Leticia (LACM
60999): Tert sinha near 1.elicia (LAO l 60977. 6 1002_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); Rio
Mamo,,!; (l'oi H.\t..... 19190:1-4). A.cn; Tarau... i ( ~I S H
83248). G<iM. : f aunda OIiYer (US:-iM 149446--48); Cawal.
eanti (Miranda Ribeiro 19261: Rio Sill M i~ ue l (CAS·SU
11867_69). .
o: C","pada do. Guimarxs (KU
93139---40; A....SP 13414); Sill Vicmte Cr-c H.\"'· 3818:1_2);
So. so Sanliuimo? (Slcindac hner 11164); Chawanlina t S
Lw2 19S2); U"",um mounlai", ne. COOtIl>bi (CM 2431_36;
P..-ISH 9209_ 19: USSM 115974)_ 3/ i" ,.. G.....I;. ' Bdo
Horizonte t US:-iM %986; CAS 98099; KU 931 4 1- 42) ;
Lagoa Seen. f OOl of Serra do Curral (UM.\IZ (10023);
Aeaba Mundo on Serra do Cum l (B. Lutz 1952): Cai'll de
Arcia 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 (MBUC V 31( 2); Hacienda San Felipe
near 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 1925.11
4,8): jlllK'linn Mcamu and .\Ial.lllllni Rjw"" (AM!' H 18996.
18998); Kamal u... (AMS H 19(100); Kurupung Rjw"r(UM.\IZ
83S98 _60) ; LACM 6097 8); Mad reba Fall•• Kurupun,
River (UM.\IZ 85147); " . ano (U.\l.\tZ 83597); Paruima
.\I i~ ~io n (UMMZ 85145-46: US~ .\I 1I8681-82 ); CUlllni
RiVC1" 24- 32 Lm a ~ Banic-a (B.\I 1934. 11. 1,1-4) ; !ol .. opc
Fall•• CUluni Riwcr {BM 1934.11 . 1.51; Oko Mou ntain.
(P.>IS H 26724--26)
Sl:RISAME.-."kb rir, Sipaliwini (.\ISH 1lOt'>. 1131); Pn>l
Gonini. Cue"",ni RivieTt.\lSH 322. 350); Post Tip. Sc..
Riwer(R.\IS H 15896)
".'0G,.....
/ 976
FROGS OF THE GE...-US PHYLLO BATES
GUYAl"E FRA." CA'S E__ Empounou W>i.ki. I ~ km SSE ""
Maripa>oulIl (La 1023).
DOUBTFUL RECORDS .-CO LO~IBlA. Cllltlar: Villa
Maria (lJSS M 14Nlbl. (46866). BOU VlA. ClN1ttlba"' N:
Yungas do: COC hatwn N. 2200 m (UMMZ 66646).
Clear ..d and stained mm..rial, _ICClh pre",m: LACM
b0979; Pattern I: loACM b0916 ; Pane m 4: LAC~l 60977 .
JMS 1680;
P.~Uer n
6: LACM 60978 .
Phyll obQU J pukJrriputu s new species
Beautiful- breasted Poison -arrow Frog
Frontispiece II and Figure 6E
Holo fJpt' . -loACM 4 2297. co llec ted 1.1 Serra do
Navlo, Terr uorio 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 ( ni ne ad ulls) , - WCA B 18 22 1;
LACM 42296. abou t 120 m, I J uly 19MI; LAC M
42298 -301, about 120 m, 2 luly 1968; LACM 42302.
abo ut 160 to IIlO m, 3 Jul y 1961l: LACM 42303. about
100 m, 5 J uly 1968; LACM 42304 . Serra do Veado
(one of the hills at Serra do " lIvio ). abou t 290 to
3 10 m, 6 July 1968. (All LACM parIIt)~ were co llected b)' P.A. Silve~onc: and Sr. Fem lra.)
D..finilion . - Tet th abM:nt; IClC' S ",eblt ss; dors um
blac k; complete yello w later al \ !ripe; li!!:ht pro~ illlO­
ve nt ral ca U·s po t abse nt : ven te r blu e ",ilh hlack
marbl ing.
Diagnosis, -Ph.l'lIob<ll'" pr,lchripU:fUJ is a member
of the piclII _' group. dif fering from P. i" gt'ri in lac king
teeth, in having a co mpreie light lateral stripe (al'>sent
in P. i" g..,i ). and in lackin g a light proxim ovemra!
ca lf-spot, from P , fXJrvufus in lacking teeth. in having
a black oorsum in life ( red in P . p<lrvufus). and in
hn in!!: a complete li ~hl lateral ~tripe (incom pktc in
P . p<ln 'uIUsl. from P. piNUS in lack in~ a li!!:hl proximoventral calf-spol. and from P. )IJ'<lrQgdillUS in lacking
teeth and in ha\'i n~ ven tral marbli ng in life . P. pul ch ript' clUs resembles P . ~t..,si ( F i~. 9 8) a nd P ,
bolil'iQlluS (Fig. 9 A) in it. ventr a l patte rn o f da rk
marbli n~ on a lig ht gro und color. P. pu/chript'CIU.•
differ" from P. " n t'rsi in Ia.;:k ing teet h. P. pulchrip<'('/u.,
diffe rs from P, 1",/;,'i,mIlJ in having a conspicuo us
light proximod orsalthigh-spor (Fig. 6El. whic h is white
in preservative and bright yellow in life. In P. hdi ,'i,mUJ
(Fig. SC). tbe dorsal surface of the Ihigh bears light.
lonJ!:itud inal . someumes undulat ing mark ings .
lk~rip'i"" oIItoJOl)pt' . - The holol)'pe is an ..dull
female . 2 7.0 mm ~noul-ven l length. Thc , k in is
strong 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 o f the hind lim bs.
Teet h an: ab"Cnt. 1llc lip of the Mlout is sublrunca te
in dOThal aspect and rounded in late ral aspect. T he
canthus m stralis is rounded. The loreu l reg ion is
vertical. The width of the upper eye lid is three -fo urths
43
of the interorbital widlh The diameter o f the eye is
I A times the distance from the eye to the lIOloIril and
slightly more than twk e tbe diameter o f the I)"mpanu m.
The pos terodo rsa l portion of the ty mpa num is co ncealed. The finger dis ks are small relative to those of
D..nclm hll/ t'J . The first finger is about equa l in length
to the seco nd. T he f ingers are webt ess . The tarsal
fold is ill de fined and slightly curved , e xte nding along
the distal 4S perccnt of the distance between the inner
me ta la rsal luberc le a nd Ihc hc e l fold . Th e tarsa l
tuberc je is absent. The loes are ....ebless
In life (Frontispicct Ill. the iris, head . ~k. and
sides wert black . A com pk lc brigh t )c llow lateral
stripe e xte nded from the Jroin 10 the upper eyefid.
along the octer border of the upper cyclid. and w n!!:
tbe ca nthus rosrraiis 10 the lip of the soout. ",he n: il
mel its fello w. Anolher bright ye llow stripe extended
from the forelimb inse rtion along the uppe r lip, e nd ing
ventral to the nostril; the anterior e nd of the stripe WIlS
blue . A blue suffusion clouded the black limbs. The
proximoo or",,1 surface of the thigh bore a con~pic uous.
o blong. brighl yello w. black-bordered spot; a similar
spoe adorne d the axilla . e uendi n!!: omo the pmximo~ ior and prox imodon.al surfaces of the upper arm.
The venter and the ventral surface of the limbs were
med illm blue , beann!!:discrete black marb lin!!:,
In pre~rval rve . the grou nd color is black . Are as
that were ye tlo.... in life are whi te. The ca nthal port ion
of the lateral stripe . the blue an terior end of the uppe r
lip stripe. and the vent ral pattern have d isappe ared;
the venter is black , ellcept for a small while spot On
the left side of the posterior belly'.
Dt'Scripl; on 01 p<lrQf)pt's . - Measuremcnts of adu lts
are in Ta ble I (o ne parnt ype, LAC M 4230 1. is a
juve nilc fema lc). All parIIlYpeS lack teeth. In some.
lhe ca nlhus rosIra lis is near ly anj!ular . All have the
Ilrst finge r aboet equal in k ngth 10 the loCCond. In
some para l) pes. the tarsa l fold is "' cll defined, ulcnd ing along lbe distal 50 10 60 percent of the d istance
betwe e n the inner met atar sa l tube rcle and me beel
fold . and the tar\3 1 tubercle is lI'cli developed
In life . the paratype s resembled the bolotype. The
anterior end of the upper lip stripe usually was blue .
The prox imal spo t on the uppe r arm was yellow or
cream . Some spec imens had blue spots on the forearm
and the pos te ro do r sal s u rface o f t he th igh . T he
prox 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 42 300-01 , the upper lip
stripe con tinues ante riad 10 beneat h the nostril. In
some specimens. mosl of the dorsal surfece of the
upper arm is white . In LACM 42298 - 99 . the thigh·
spot exte nds o blique ly distoposteriad from the base
of the thigh. The ventral paue m of dan mar bling on
a light ground is distinct only in LACM 42300 . LACM
42 303 has a small whi te median spot on the posle·
rior belly .
BULLET/,V OF THE NATURAL HISTORY M USEUM OF LOS A.'l iGEU.S CO Ul'tTY
44
R~IQli(»/$hipJ .-P. pulchriprCIUS probably is closely
related 10 P . bvli"ianus and P . prf" rsi .
Sw much comems (one specimen) . _ From ~C~t
42302: HYME....OPTERA: FOIlTlid dao:: B/t'phuridana
b"<l~j{jtnsiJ
w hle. ( 2), M.w·ocr puru$ smil hl (Fo rd)
(II, Cypho myrma rimosus (Spino la) ( I), M.m n icocrypt" sp. (I), Myr micocrypl a ? ~ p. ( 41. P ht'ido lt'
~ . (8) , H, po ponr ra ~. ( 2). Wasma" " ;Q auroputICtUlQ
(Roge r) (5), also fragmen 15 of lints of three 10 four
othe r genera .
D istri bu ,;OII aJU1 ha bi tat (Fig. 16 ). _ Elevlll ion recerds an: from 100 10 3 10 m. P. pulchripulus is known
only from the type locality. in the Guianan region of
nonhem Brasil near the Rio Ama pari. I tributary of
the R io Ara guari, wh ic h flow s into the Atlantic Oc ean
north of the mo uth o f the Amazon .
I mol these agile diurnal frogs 110 tile f"re ~t floor .
betwee n the sca ttered brief f3 i n ~ howef~ of the opening
dry SC3,;on. One perched o n a fallen log two-thirds
of a meter above the ground : it dived to the ground
and hid when I approached.
Serra do N3Vio is a town and ma nganese mining:
complex surro unded by low. fore..tc:d hills, The forest
incl udes large-buttressed trees of grea t height and
d iamete r. Spiny palms guard the abendam melastomaceou s and ru biaceou s under gro wth . The ground
bears a heavy lear co ver: some portio ns of the ground
are poorly drained.
From 1954 10 1964. Serra do r-;avio had a mean
ann ual rainfall of 2402 mm. seven ty pe rce ne of it
during the ~ ingle we t seaso n from lanu ary 10 June .
The mea n annual temperature ..... as between 25 and
26°C. ee mean minimum was 22.2"C. lind the mean
muimum 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 in us new species
Emerald Poison-arrow Frog
Frontispiece II and Fig ure IIC
Ho l<>typl' . - LACM 6 4435 . co llected al Pa n de
Azucar . 39 km N ~ EOx apam pa . 10"15' 5 . 75° 14' W. in
the lscozaztn Valley (Ihe Rio Iscozazm is a Iributar)'
of the Rio Palca zu . in the Pechlte a d rainage). Depanamenlo de Pasco. Peril . 31lO m, o n 16 Augusl 196 1.
by R.E. Elheridge .
Parur),fWs (one adull and seven ladpoles). -LACM
6+436. jo poparatypes (seve n tad poles from the back
of the holol)'pel: LACM 6+434. collected at lscozazin
ainlrip. 47 km r-;NE O u pampa. urn ·s. 75°10·W.
in ttle lscozazin Valley. Pasco . Pe ril. 360 m, on 19
l uly 196 1. by 0 .8. Wake and R.E. Etheridge
N(}.17
D..fi nilion. - Tee th present: roes ....e bless: dorsum
black : co mplete gree n lateral stripe; light proximoventral cal f.~pot abseet; venter green (anteriorly) and
blue ( posteriorly). no l marbled.
Diagnosis . -Ph.~-IIobuJf'S sltUlragdinU$ is a member
of the piNus gro up. differing fro m P . boli,·jafllu
and P . p ukhrifWct u5 in having teeth and in lacking
ventral marbling. from P . ing..ri in having a complele
light la tera l strjpe (ab...:nl in P . ing..r i) . in lacking
a light pro ~ifno>.'C ntral cal f-spot, and in lacking vent ral
marb ling. from P. pan 'ulus in having a black dorsum
in life (red in P. pa n·uI U5). in having a complete light
later al stripe (incom plete in P . p<ln ·Wus). and in lacking
ventral marbling . from P. pet ersi in lack ing ventral
marbling. and from P . pit:rU5 in having teeth (usually
lacking in P. pi CIU5). in lack ing a light pro xim ove ntral
ca lf-spot. a nd 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 strongly
gr<tnul ar on the dor sum and the dorsal surface of the
limbs. and smooth on the venter and the ventral surface
of the limbs. Ma xilla ry and pre maxillary teet h art
present: vomerine teeth are absent. The tip of the snout
is subtruncate in dor",,1 aspect and rounded in I..teral
as pect. The canthus rostrahs is rounded. The lore al
region is slightly concave. The width of the upper eyelid is slig htly 1('S~ than the interor bital width . T he
diameter of lhe eye is slightly greater than the di~1aIlCe
from the eyt to the noslril a nd slightly II1Oll: than twice
the diameter of lhe tympanum . The posterodon.al portion of the tympanum i~ concealed. The finger d isks
are small relalM to tbose of D..lldrobaltS. The I irst
finger is abou l eq ual in le ngth to the second. The
fing ers are ....e bb s. The tarsal fold is slightly cu rved .
e Mending along the distal 50 percent of the dislance
bet....ee n the inne r me tatarsal tuber cle and the heel
fo ld. The tarsa l tubercle is slightly developed. The
toes are webless.
In life. the dorsum. the side of the body. and lhe
dorsal surface of the limbs were black. A bright green
co mple te la te ral stri pe ex tended from the g ro in 10
the upper eyelid . alo ng the o uter border of the upper
eyelid . and along the canthus rostralis to the tip of
the sno ut. where it met its fellow . Anoth e r brigh t
green stri pe e xtended from the forelim b insertion alnng
the upper lip. ending ventral to the nostril. The lo we r
lip. the breast, and the anterior t.....o-thirds of tilt belly
we re green . The ve ntral surface of the limbs and the
posterior one-Ihird of the belly w ere dark blue. The
throa t bore a pair of lateral black areas (perhaps a male
secondary sexual charac ter).
In preservanve , the holo type is entirel y black 10
dark gray. ex.ccpl for lighl gray lateral and upper lip
stripes .
Adull parat)·fW . -l n prese rvem..e. ttle adult parltype (LAC M 64434) resembles the bolory pe: it is
the same sex. and length .
FROGS OF THE G£SUS PHYL I.O BATES
1976
Tadpolet (Fi gs_ l ID. 1 2 E) .-~ ho\ol)pc earned
'>eVen ladpok<. (LAC M 644 36 ) 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 records are from 360 to 380 m. P . smaragJinll.• is kno....n
only from the type loca lities. in dense forest in the
Amazon drainag e of Pl:ru. east o f the Andes.
Etymology. - La tin: sma rafi:dinus (emera ld) re fers
to 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 le ngth of adults moderate
10 large (3 1.5 -49.5 mm); loes ....ebIeM; paired ~~
lateral (Figs. 2. 6A-C); lig ht 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 we bless: dor sum
black with gre en o r o ra nge spo ts or mar bling. o r
u ni form o ra nge: complete green o r o ran,e la te ral
stripe; Iighl pro :\imo \'entral cal f·spot absent; venter
blac k with bloc marblin,.
Diagnosis. -Ph.\'//oba/~J batsleri is a member of
lhe trivitta lllJ group . di lTering from P . tri,'itralll.1 in
having teeth .
D~sc ~iplion . - Meuure me nts of the bclorjpe (a
37.0 mm fe male) and referred spec imens are in Ta ble I.
The skin is strongly granular on the dors um and the
dorsal surface of the hind limbs. and smooth on the
forelimbs, the venter. and the ventral surfac e of the
hind limbs. Ma xiJla r) and premaxillary teeth are
present. The first finger is longer than the second.
The toes are we bless.
In life , in the holo type (Melin 194 1), the dorsu m
wil.s black ""ith a median row o f light green ~pols ( Fig.
2F). The side o f the body was black ....i th a co mplete
lighl green lateral suipe ex tendin g from the gro in 10
the upper eyelid, along the outer border o f the upper
eyelid. and alo ng the ca nlhus rostralis to the tip of
the snout.... here it met its fellow. Melin did 1101: mention the colo r o f lhe venter in life.
In preservative. the bolo type is dark bro....n. e xcepl
for the following white areas: a median do NiI row of
seven spots. a fe.... small spots on the righl s ide of
the beck. a complete lateral stripe on each side of the
45
bod)'. pan of the upper lip. the proxrrnodorsal surface
o f the upper arm, lrac~ o f light pigment on the donal
and 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 slope
of the Andes, in the drainage of the Rio Huallaga, 3S
k m so uth"'l utheast o f Mo yollamh a and 76 km we st·
northwest of Chazuta . I refer nine additiona l toothed
specimens to this species . Tbe se specimens are from
east of the type locali ty. from the drainage of the Rio
Hualla ga (Chazura. Pachiza , and the Rio Cainar acbc j.
Specimens from Pachiza differ from the bolorype
in having smaller. scattered dorsal ,pOls.
Some o f the spe<:imens from Chazuta have a color
pattern similar to tha t of the holotype: I do no! know
their color in life. At Chazura. W.C. Sherbrooke
collected lOOl hless specimens of P. trivunuus (....ilh
a pair of yellow lateral s.triJlC'o and a )'ClIollo median
d~ 1 stripe) together with three too thed speccneos
(....i th a pair of orange lateral stripes and no median
stripe). 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 specime ns (Fig. 6A-B)
had a blac k dorsum with oran ge marbling: the third
specimen (Fig. flC) had a uniform orange dorsum .
The venter in the toothed specimen s was black with
blue marbling (Fig. 60).
Ge neric oflm:ulion._ Mel in (1 941:65) desc ribe d
the boloty pe a.~ a Dend~ooo"s because he thought il
had no teeth. onl)' "tooth-like aspe nties." I e xamined
the holol)pe. It has teeth and small finge r dish; it
is a Phyf/obates.
S tomoch contents (one specimen)._ From A."-tN H
43402: HYMEl"O PTERA: Formicidae: Pheidofe .p.
( 21. CompOflOflls'! sp. ( I); O RT HO PTE RA: Telli·
gon iidae ( II: HEMIPTERA : fragments.
Dist~iburion and hobi tof (Fi g. 141.-Elevat ion reco rds are fmm 274 to 1097 m. p , bllss/eri inhabits the
Amazon drainage of Peru . from the easte rn foothills
of the Andes cast to the Rio Huallaga .
LOCilfiti H , -PERO. Son MOrf;n: Roque. 3~ km SSE
Moy"bamba (I'\MO ~1l); Ch:llUI~ (AMI'\H 4Z867. 42944:
USl'iM 166903A-C); Pachiza (AM ~ H 42321, 42333, 434(2);
Rio Ca;narachc tAMS H 42313).
Cfea~eJ und swi lled "I(l/e~iof. -A MS H 43402.
Ph)'ffoba fes trivinanu [Spix 1824)
Three-stri ped PoisorHul'Ow Frog
Figures 2A-E. 7C
1824
?1824
H,·/o ,ri'~II(l1O Spi1: 3~. pla....'I, fig. I. type locality:
Rio Tde , B nI~il
H,./o ";lIa~i"'(l Spi1: 36. p l ~le 'I, fig. 2. Iype
JoealiIY: 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
1826.
18.\0.
?18.\O
18).8
18~8.
111M
111M
1&82.
1926.
1926.
19~2 .
1\161.
1969.
1970 .
H ....~rrima belonged 10 1M sam: ~~s I 1La\~
noI "Cen tM I~ pe~ !he)' ..-ere ck"roye4 .. WoOd
War U. axordin!, 10 U. GlUber lin lill.)).
H,-kIpI'Wl";'~lIaUl , 50., in Schkp:l: 239
lHNlTt>ba". Im·iltalM•. Wal lct-: 202.
lHNlTt>ba"...i~rriIJIM' . War.lior: 202.
H,"""",Wlli~ . T..: hudi: 70-7 1.
H,.lapI,.;" Ii...."'"" (pan). GUnther: 125.
o,NlroDQU . "iK"rrUorIU ( part) . Ste ildal;hner. 259260.
lHNlrobm,. ti....1OTi1U ( put). Sa: Illda<:IIMr: 260 .
lHlldroD..,,. rrM llQ'1U (pan). Boult-np:r: 1 44_ 1 4~.
O,lIdrol>,,,,. trivilla'M.'. Miranda RiMir o: 18 1.
!ell·fi8. 98 .
On.droba,,. U lr(Jvit lallU Mir~nda Rib<oiro: 180.
!e\t· fi8. 'n . •yntype. in tM Mu~u Paulio.ta. type
Io<;alil.,,, 6 bidos. Brasil. and Rio JUllli. I Ei llllll:~.
"",«Jrdin8 to Bokermann 19t16.J Br•• il (according
to Bokermann 11966:3 41. O . 1, I'a"i",,, o. i, a
synonym of D . Iri.in",... ; I have not ,ee n the
_yntypes).
Orlfdmh<tln ,rMlldl...,. B . LUll ond Kin,,: N:>9
l >r lfd m lml r , I,il'illat...,. Cochran 107. plate 4 1
(cnl", phoIO).
Drndroh<tlr. "Mlltll... . Hoogmocd: plate I. fig. 4
(col", photo).
DrMrobaln " i, ·illal... (pan). C"",hran and (lo;>in '
14 -1 6 . plate ~G ~I (o nly US NM 1186 81 _ 82 .
MCZ 24443. ANSP 18188 are n<Il P. Iri,'illa"u).
Drfi ll i ti OlI . _ Teeth a ~nl; Illes web Je . , ; dorsum
bladr. I\-'ilb or I\-itnou ( yellow, yellow-green, or green
median donal stri pe (medi an stripe some lirTlCS bro ken
mto long segments or , bon spols); somet imes dorsum
bbd; with green miirbl inil; sorrenmes dorsum uniform
g ree n; complete yellow, ye tto .....·g re en. green. or
o r;IlIFC lateral srnpe: light proximoventral cal f.spot
absenl ; venter blac k ...-ith grren. blue -gree n . Of bl ue
'flOls or marbling.
D;oxn o.1;$. -Phyf1obol~$ Iri"i tl(JI" $ is a mem ber
o f lhe t ri" it/<JtUJ group. d iffering fro m P . boulrr;
in lacking teeth .
D o cript;oll . - Me asuremen ls are- in Tab le I. P.
/r;";/Illtu,' is the largest species of Ph.\' II"h<I/~ ' , T he
skin ts slightly to moderalely granu lar nn the dorsum
and the dnNtI surface of lhe hind limhs . and _nmotb
on the furelirnbs . lhe verner . and the ventral surface
o f the hind limbs. Te eth are absent, The fiN finger
is 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 of
Hyl<J /r;w'/Iu/a Spil 1824 ): Specimens beari ng thi s
pattern are known from Faro in Bra.il \\-'C.IWan:! alon g
lhe Amazo n to the Pengo de Ma n1<C'r iehe (Ihe pass
of 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 Ucayali
and its drainage at leaS! as far as the R io Tamaya
(SOUlhea'" of Yari nacocha). In lhis pal1cm. lhe ground
color of tbe doroIllm is blac k.. "There are three )'ellow.
yello ...-green, or gree n st r ipes (one media n dc r... 1
.'0'0.17
and one- pair Jateral). In some specimens. lhe median
corsal «npe 4 , one- or mo re sm all break. (Fig. 20).
Such spec imens are known from Santa J ulia ne.... Faro
in Bras il. Puerto ~ariilo in Co lom bia. and Morale s.
Pschiza. the uppe r R io Cllshah:l1ay. p...mp;.o Hen nusa
(lower CIl,habalay). and ~.5 km from Puerto Santa
Clara in Peni. A t 1IlO'>l of rbe-e localities. boIh ,peci
men, 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 f
tooth less specimens from Pampa He rmo sa whose patlerns inclu de transitio ns bet.....een an unbroken med ian
stripe and a medi an doNtI row of spot s . In life . in
the holot ype, from the R io Tefo1 (a southern tributary
of the Am azon east o f the R io Jurua) , Brasi l (Spix
1824:plate 9. fig. I). the dorsum was black with a
)'ellow median stripe (bro ken intcrorhitally}. T he s ide
of the body "la, black with a com plete yellow lateral
stripe . The do rsal surface of the limb, I\-'as black I\-'ilh
ye llo w spols and mar blin g . Spec imens from C ha7uta.
Peru (color slide s by W .C . Sberbrooke), resem bled
tbe holotype, but t he med ian slr ipe was unb rok en .
and lhe- Sl\oOUt and the-dONlJ surface of ue limbs were
e nt irely yellow. A specime n from Tinge Mari a . 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 specime n from 7.2 krn I'lOnh .
wc st o f Let icia . Colombia (my color photO) ). rese-ml>lCu
the bokxypc. but the strip:s were green.Tbe median
stripe 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 lim b was near l) cove red
by green marbling and str ipes. "The ventral .u rface
of lhe fo relimb wa.s blac k wilh a bl ue- stripe extending
tbe length of the uppe r arm. The vent ral surface of
the hind limb wa s blac k with blue spot s and marbl ing .
The vente r w as black w ith man y small blue spots
(Fig. 7C ).
Pattern 2 (Fi g, 28): Specimens bear ing this pattern
arc known defin itely (from well-preserved specimens
and from color note> and slides) from four locali ties'
Yartnacocha on the R io Urayali (60 km nonhnorth wes t of the junction o f the Rlns Pachitea and Uc ayal i).
the lowe r R io Pacbitea ncar the Bo sque Nac ional de
l part a . and lwo localitie, on the upper Pac hitea drainage . Pan de- Azocar and San ta Isabel. The pallem is
di fficu lt to d isc er n in pre-SCl"o'W spec imens; il may
occur in specimens fro m OIlter localhies in lhe Pach he e
draina~'C and along tbe Ucayali fro m the R io Tambo
III 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. Sa nta Isabel. and
/976
FROGS OF THE GENUS PHYLLOBATES
Pan de Aziicar, Peru (color photos by J .P. Bogart
and A.C . Thore se n: notes by D.B . Wake and R
Etheridge at CRE ), the dorsu m woo, uniform light
green. The side of the body was black with a complete
yellow lateral stripe e xtend ing from the groin to the
upper eyelid, where it merged with the green dorsum .
A white stripe extended from the forelimb insertion
along the upper lip, ending ventral to the ante rior
corner of the eye- The dorsal surface of the limbs was
mostly light yellow. green. In the specimens from Pan
de 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 b
green with a small amount of black marb ling a nd
spotting; in the other specimen , the dorsum is black
with green marbling.
Pattern J (Fig. 2C; probably includes the symype
of Dendrobate s tetraviuatus Miranda Ribeiro 1926
from Obidos): Spec imens bcaring this pattern are
known from Guyana, from Suriname, and from Bra, il
along the lower Amazon from Faro to the Atlantic.
The late ral stripes arc green , yellow-green, yellow.
or orange . The dorsum is black; there is no median
dorsal stripe . In life . in specimens from 5 km east of
Helem. Brasil (notes by M. Crump at KU), the dorsum
was black. There was no mcdian dorsal stripe. The
lateral and upper lip stripes were yellow-green. The
throat was black. The rest of the venter and the ventral
surfacc of the limbs were black with blue-green spots
a nd marbling. Specimens from Suri name (notes by
M.S. Hoogmocd at RMNH; Hoogmoed 1969:plate I,
fig. 4; Hcogmoed 1971: 1, 6 ) had a black dorsum and
lacked a median dor ' al stripe. In mosr specimens. the
latera! stripes were green , but in specimens from the
Nassau Gebe rgre in eastern Suriname , the stripe s
were yellow or light orange. Green spots ornamented
the dorsa l surface of the limb, : light blue spots adorned
the belly and the ventral surface of the limbs. In frogs
on the Lely Gebergtc . Suriname . a transition occurs
from yellow-green stripe, on one side of the plateau.
to yellow stripes, to bright orange stripes on the other
side of the plateau (Po lder 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 and
February; Polder (1976) heard them calling in August.
Tadpvi",~ (Figs. I IG , 12H). -Twe nly-five lots of
tadpole , bave been taken from the backs of adult males
in 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 a
small temporar y j ungle pool. Hcogruoed found tadpoles of this species in a water-filled fallcn palm petiole
in Suriname in February (Po lder 1976). Measurements
arc in Table 3.
47
Re/lllio'l!ihip!i._T oothlcss specimens of P. trivuuuus
<Ire symparnc with toothed specimens of P . ba.".lleri
in Peru. at Pachiza and Chazuta. As noted above, a
toothless specimen from Pampa Hennosa , PerU. (in tbe
Ucayall drainage ), has a median dorsal row of spots
(Fig. 2E). This pattem is very similar to that of the
toothed bolotype of P . h",sleri (Fig. 2F). Provisionally,
I am calling P. bassleri and P . tri virtatu .f distinct
species. The green-spotted specimens of P. bassieri
(incl ud ing the hotorypc) may merely be toothed P.
mvutatus. If so, Sherbroo kes orange specimens from
Chazuta (USNM 166903A-C) prObably belong to an
undescribed species.
Lif'" history . - Hoogmoed (197 1) and Polder ( 1'.I74a,
1976) observed wrestling arnung c<lptive specimens.
In captive specime ns, Polder (1976 ) observed that
eggs are laid on leaves, clutch size is eight to 22 (clutch
size is at least 27 in this species; see Table 4), tbe
period fro m fertilization to hatching is 14 to 17 days.
and the period from hatching to mctamorphosis is 41
to S4 days. Polder also noted tha t in rbeir native habitat
in Suriname , frogs of this species are extremely wary,
and their coloration is inconspicuo us: thus, he doubted
that their coloration has an aposemauc functio n.
Para!iires . - I found larval nematode, parasitizing
a specimen of P. mvlnatus (LACM 6098 1) from 7.2
km northwest of Leticia, Colombia. Seven nematodes
were free between the skin and the muscles (in the
tbighs and in the side of the body), and several more
were enclosed in c ysts attached to the thigh muscles.
Diu llsn . - Polder (1976) has discussed dise ases
in captive specimens of this species.
DiSlrihmimr and Iwhirlll (Fig. 15). _Elevation records are from 20 to 680 m. P. trrauaius inhabits the
lowland forests of Guyana, of Suriname , and of the
Amazon drainage of Colombia. Ecuado r. Pe ru. and
Brasil. .\1. Crump collected specimens near Belem.
Brasil, in the daytime on the fo rest floor , In Suriname ,
population density of P . trivillmus is greater beside
forest roads than in undisturbed forest; the frogs seem to
prefer fallen trees with their tangle of broken branches.
which provide call ing and hiding places (Polder 1976).
Localities . - COlO MBIA. Pat"",a)',,: junctionRim A~u;Jc,
and Mccaya. upper Ca~ ueta drainage (US!'lM 144992- 93).
Amazonas: Puerto Nariiio. Rio Ama1.Onas (ILS 3: USNM
146831-32): 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. 3649 t. 364(3);
Tingo ,I.-laria (USNM 129414; AM!'lH 53762; CAS 85152);
Parque Nacional de la Bella Durm;ente, 10 km by road S
Tingo Maria (USNM 166910); Ganzo Azul (FMNH 45t43.
45292): Monte Alegre. RIO ?achilea (AMNH 43021, 43025)
Sa" Mani,,: Tocache(USNM 166908-(9 ): Chazuta <US!'lM
166907; AMNH 42616. 42786-89. 42943}; Huayrapurina
near Chazula (AMNH 42952. 42954); Morales (AMNH
42012); PachiUl (AMNH 42326, 42g26); Achinamisa. Rio
Huallaga (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
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<hiyacu
E Bahapl.ll:rto (A~I S H 425751; Iquito. t~'S H 109839;
AMJ';H 422941; Cerro Azul. 24 lr.m E Contaman.i (FMS H
56249 _$0 ); Ptocallpa tP.d ..... H 5625 1); Yarina<o<;l.... Rio
UeayaJi t P.d ..... H 4~-O7 . ~5 2 -S6); Pam~ Henno......
Rio Cu..tlabuay (AMS H 4!OI 1. 420 13_ 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 ~' S H
42249.41468 _70. 42472-7 3. 42564. 42695. 4JO.17 . 43370.
43378); Puneua. Rio Tapic"" tAMS H 42873. 429151; jun<tion
Rim. C,mlay a and Tapiche (AMS H 429%); Rio Blanco
tA ~'S H 42985); Rio Tamara (AMNH 421101); Ca, hiboya
(A MSH 42044, 42976. 434H ); Oll,nla "" .. Quebrada
Ca.hihoya (AMS H 43212); Rean Re<l n.l..a", Suh, ya (AMNH
427(9); Rio Pi",!ui (A ~1:'m H192. 4319Q, 43552); Yuri·
magu.. (Bnuleng er 1883); n<:ar Tar apo lo (CoP" 1886 );
Peba. (Cop" 1886); E'liron . Rio Ampi ya~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 (USNM
1667641; PIon de Azuear. [",o ..li n Valley, 39 km l"NE
Oupampa (LACM 72035-46, 72057-61) . /"" ''': junclion
Rim.Tamho lllId lkayali (AMS H 43262).
BRASIL.......... lJlazo"... : Rio Tef<i (Spi> 1824); Td<i (MHNP
6242); Japuri . Rio Japurii (MP£ G 338_39); Benjamin Con.
, lanl (WCAB 133SO; KU 93153; CAS 11870-72; B. Luu
aJ>d K...... 19521; Rio l ",'ary. 30 lr.m ,jo,.'n>lrcamffOOl Benjamin Con!ltalll (CAS·SU 11879); hacoa ; l B. 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·SU
118tl6). "p o: 6bido> (Miranda Ribeiro 1926); Caotanh.aJ
Grande ""ar 6bidt» (MRHS 61. 6I B-6IE); 5.a.nla Julia
DCar F~ ro ( MRHS 4741; hiubni (ZS BS 99/ 1911; MCZ
121l8); llha de Mar~jO (MCZ 1S663); 19-.. apt·A~u ( BM
1904. 7.2 6 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 R i~er ( KU
b9696 _ 700); hud...-aler. h abu C re ~ k ( FMNH 30909);
Ku)u wini Landing (AMS H 46232); MaNd l (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 (AMNH
3'1119).
SU R INAME.-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 H 774SO;RMNH ISOI4. lS(17);
Afobida road , 10 lr.m from I'hodra t RMS H " neall , B"'....
,..,..: Bro""n.borlJ (RMS H 157041; railroa.l. km 121. near
q..-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 ~'S H 2631; Toek...eeeree Krcck
CRMS H "neat ); .... Sarama:ca Rn.-io-r. lr.m 91.5 (BM 1946.4
1.231;"pp:1" Stuin.amt: Rivitr ~ Sara Krcclr. (MCZ 10164,
No. 27
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....... (RMSH
lS022l; Mom ,., T"'I"X(RMSH 425. 534. (4 7)-
CI,arrJ and 51ai" rJ maltriol. -LAOI6098 1.
R U IJ)IE"
LIs ranas venenrn.as del genero PIt)-J/o oo1rJ (Fam ilia
Dendrobaudacj !>C encuemran desdc Co sa R ica hasta
el Pe ru . Boli via . )' e! Brasil. La mitad de: [a. e.pecies
.c eecuemran en la rellion amazonica. Las 20 especes
se 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, pulchri pa lu<. P .
.< muraXdin us); )' grupo /r jvilla l us (p. b<lJ./rri, P .
tr/ virrclfus).
Ame no nneme rode s las ranas de ndrobatid as . in
dentes se C'On,idc: raban Dtndrobalts. En esse revision
a1gunas e. pec ie. que Cll"ettn de diemes ~ considcran
Ph)'lIobal~s . As i es que Ph)'lIobal~s 00 se disl in8t1e
de DtruJrobcJlts por la presencia de dientes [aunque
la mayuria de la. especes de PIt)'II0b0 1t s t iene:dien,e.).
sino por el lamaOO de los discos )' la laf8ura de los
dc:dos. En Ph.l'ffobo1tS . los discos rerrmn ale s de los
ultimos Ires dedos de I. mano son relalivamente
peq uefios (menos de 1.5 veces mas ancho que et
dedo); d primer dedo de la mano es iguai a 0 mas largo
que e[ seg undo . En Drndrobo1ts . los diso;:ostermmales
de los ullimos ues dedos de la mano son rela livamcnte
grande> Cal meeos dos v«es mas aoc ho que el dodo);
d pri mer tkdo de la mano e. mas corte que e ! seg undo.
Las sig uienl e. e specic s a nterio rmente convlde radas
Dl!ndrobm ts se mcl uycn aqu i en Ph)'lIubol l!J; bassle ri,
ingeri , p<1Tvulu ,l . pklU.f, y trivusasu s. Las primer.."
Ires de e. las eSPl'cie s ne ne n dientes . Algunm eje mptares de P. pictus ricncn dicntes y la may oria carece
de diemes . P, Ir/vll/lITlI.' carece de diem.:s ,
El gen ero Phyl/ ob<l/tJ sc mannene distinto de au
parienl e . el genero Colostet hus , En Ph yll obmtJ . e l
corso ten elanimal vivo ) es negro ron listas de un co lor
sub jdo. 0 e. roratmeme de un color subido (pe r ejem plo.
rojo). mienl ras q ue en C%Slr1 hus. el dorso (en el
animal 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<.> en
la piel. 0 sea que Ia pie l de: PIt)·JloIJorrs uere veneoo .
) [a de ColoJttlluu no 10 liene . Hay especes quo: se
pareccn a am bo. generos; tal Vel 10. dos genet"O'l sc
junlarin en e! futuro ,
1976
FROGS OF THE GEt·...US PH YLLOB ATE S
AI menos dos e spccic s de Phyl/obmes se uufiz an
por los indigenas para envenenar los dardos que tlran
de bodoqueras . Los indigenes de Ia rribu choco, que
habita n la re gio 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 , y
buren las puntas de los dardos co ntra la piel de la rana.
Estc procedimiento, de scr !ro per Po sada (190 9) y
Wassen ( 1957), causa tanto dolor al pobre animal que
su veneno e xuda sobre la piel y se adhiere a un scrc o
espiral en los dardos . Las des especies de ranas que
tiencn la mala fortuna de unluarsc sc Uaman en la
lcngua e mbena 0 empera de los ind igenes " kokoC"
(Phy llobmo (lUrOllU!Ria ) y "near a' (Phyllobotes
hieolor ).
49
Las ranas del gencro Phyl/obmes son dium as, y la
mayor ia habita eI suclo sombrcado de la >elva humed a .
donde sc allmcnta princi palmcnte de hormigas pequcfras.
Po ne n sus hucvos en hojas . Despucs de salir de los
buevos, los renacuajos sobcn la espalda de un ad ulto
(al menos en algunas especies. e! adult o que llcva
los renaeuajos puede ser el padre 0 la madre ). EI adulto
Ius llcva a una peq ueria extension de agua (pm ejempto ,
el agua en e l pecfolo de una hoja de palma cafda}, donde
10 ' deja Los renaeuajos ccmplctan su desarroll o en
el ag ua
En 1<1 unica especic I P. \' il/ " IIIS) cup historia de
vida sc conoce bien, no hay amp lexus .
Se describcn en esta ohra cuatro especies nueva s
procedentes de Sudamerica, Ptrdtobares pete rsi, P ,
tndchripectus,
P. Jl1l11rugdillU.I' . y P. t " p" m
BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGEUS CO UNTY
50
1\'0.27
L ITElV.TURE C ITED
1912. Descriplion. of fit '" Nlrachians from the Andes
of South Amefita. preserved in the British Mu' eum.
Ann . Mag. Natur. Hist., Scr. 8. 10:185_191
AU llOl,,·UQI,'E. E.X., J .W. OUl' "'SD B, WnKOP. 197 1.
BatndlOto~in : c ~ mistry and pllannaoology. Science
172:995- 1002
1913. On a collection of batnll;hians and repciln INdc
by Dr. H.G .F. Spum:II, f .Z.S., in the Cboco, Colombi a
Proc. Zoo!. Soc , London 1913:1026-1038.
ANDERSSON , L. G . 1906 . On battlc hians from Bo livia ,
Argentina. and hru , coIlecled by Erland Non1enskiOld
1901-1902 and fl:ils Holmgren 1904- 1~ . An. . Zool.
;\(12):1- 19
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 .... frog
from lhe Andes of Colombi a. Proc. Zool. Soc. London
1919:79-81.
BUSSEU-. P. 1973. Vocaliz.alions in the territorial behav ior
of the froglhttd,.~. """,ilia . Copeia 1973:277- 284.
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 02
in O . Bangs. W. Brown, J . Thayer, and T. Barbour,
The v ertebrata o f Oorgona Island. Colombia Bull.
Mus. Compo ZooI. Hatvan:l46 :87- 102.
1908. Some eew repfiles and amphibians. Bull. Mus.
Compo Zoo!. Harvan:l5 1:315- 325.
1909. CllIT«tions regarding !he names of two mxntly
described 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 80 L'a . T . "'SD G .K . N OBLE. 1920. Some amphibian s
from 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 . ",s o 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 hene
Repilien aU. Neu- Gra.nada und Crusteceen au, Cltina
Nacltr. 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 "nol ada 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
brasildro (Anura, DendroNlidae). Rev . Brasil
BioI. 27:349- 353.
Bo L"LE"'Gu , G .A . 1882. Catalogue of the Batrachia Salicnlia
s , 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
.lIde,"
1888a. l'o'oce on the cl~siflCat ion of the Ranidae . Proc .
zcor. Soc. London 1888:204-206.
C H"'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..04 23_
.
~':' ~ 'l';;~i(f~~rc~i;~~.% ~~~
world. Doubleday and
COCH....S. O.M. ",so c.J . (joIN . 1964 . Description of a eew
frog,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 tropical
America and Asia , Proc. Acad. Nalur. Sc i, Pbiladelphia
12:368-374 .
1862. On "OInt ne...· and litlle kllO...n Atneric""l AnUlli .
Proc:. Aead. Natur. Sci. Philadelphia 14:151- 159.
1863 , O n h at;hy"ph" I". , Scophiop us, and oth er
American Barrachia. Proc. Acad . S atur. Sci. Philadelphia
15:43-54.
Fourth comribulion 10 the herpelO108Y of tropic,",
Amer ica: 3, Notes on neotropical batracllian•. Pnx: ,
Acad. Natur. Sci. Philadelpltia 18:129- 132.
1866.
1868. An namination of lhe reptilia and twra:; ltia cetained by lhe Orton nped ition to Ec,wlll' and lilt upper
Amazon....·ith notes 011 « hef spede•. Proc . Acad. Satul
Sc i. Phlladelpltia 20:96_ 140,
1874 , On some Batracltia and SemlIOgnathi brough t
from the upper Amuon by Profe ""'f On on . Proc.
Aead . !"alur. Sci. Philadelphia 26:120- 137.
1875. On lhe Balrachia and Reptili a of Costa Rica
J . Acad . I"atur. Sci. Philadelphia. Set. 2. 8:93- 154
18n. Tenth contribution 10 the: herpetology of tropical
America. Proc. Amer. Phil. Soc, 11:85-98
1888b. On the ··nur;in i '·hbits of D~,.J'obufn , u
oMen ed by A. Kapp ler . Ann . Mae . Nalu r. Hisl .,
Str .6. 1:454..0455.
1886 . Catalo gue of lite specits of batrac hians and
~ptiles contained in a colleclion made at ~bas. upper
Amaron . by John Hau•...-ell. Proc. Amer. Phil. Soc .
23:94-103.
1898. An acco unt 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.
Proc. Areer . Phil. Soc. 24;44-60.
1899a. Description. of fit\\' batrxhians in lhe collection
of lhe British Museum (Na lural History). Ann. Mag
Nalur. Hist.. Ser. 7. 3:273-277.
I 899b . De.cript:ion. of ne.... replik. and balrac hians
collected 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 Venezuela
Ann. Mag, l"alUr. Hisl. , Ser. 7, I I:4HI-484
1887. Syn0p'b of the Balfacbia and Reptilia oblained
by H ,H. Smillt. in the pnw ioce of MalO Grosso, BrIl i!.
1893. Second addition 10 t~ kl'lO'll'ledlJ:C of the Batrachia
and Reptili a of CO,l a Rica, Proc. Amer. Phil. Soc .
31:333 _347.
1899. Conlribuliom 10 lite hc-rpetoloJy of New Gr3nalb
and Argentina. Philadelphia Mus. Sci. Bull. 1:1-26
CM UM I', M.L. 1972. Terruo nahty and muting behavior in
De'ld'obul~S It' O'l UJif N IlS (Anura: Dendrobatidae), HerpelOlogica n :19S_198
poca. J.W. ",so P. DU.... S T. 1974. Coun .hi p beh....jor in
Colosfelh"s m/la,;. (Dendrobalidae) . Copeia 1974:988_
990.
/976
FROGS OF THE
ces..US
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.
PH YLLO BATES
51
des<:riplion o f 1"'0 ne ", fo rms . ACI. Bio i. Vene z.
4;349 _364 .
in dendrobalid
HU ut K"H , W. 1940 . Beilri~ zur Ke nnmi, der Gattun,
H ylou llu ( B r~ h}eep/l.lidx , Amp/l . ). Zoo!. An• .
131:11l-128
Du",ilIL, A.· M .·C . .. "0 G. Bl l ION. l SI!. Erpe tol o Ji e
genera le o u h isloi re n".lurelle com plele de. repi le. ,
Volume 8 . Libnline encyclo pedique de Rom , Paris
792 pp .
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 iden
44:133_ 141.
DUNN. E.R . 1924 . So me Panarn..nian Frogs. Dec... Pap .
Mus. Zoo!. Unlv. MIChigan 151: 1-17 .
1971. Dt.w.roba/~S , e ine farl:>enre iche: Ganung. Aqu,.,..
Terrar . Z. 24:1_7.
1931. New fm,s fronl Panama and Costa Rica. Occ....
Pap. Bm ton Soc . Natur. Hist , 5:385-401.
1972, Fro, ch<: . die ihre Kaulq uappen huckepack tragen.
Aquaricn M. g. 1972:28R-293.
1933, Amphibians and replile. from EI V.lle de Anton.
Panama, Dec..... Pap , Bo ston Soc . Natur. HiS!. 8:65- 79.
I"'STlTUTO R ~ C ION " 1 DE DUENV0 1VIMHlTO DO AMA' '''
( IRD A) "NO COl<SH HO N.. CtONAL DE GEOGI.. ft..
URG E) , 1966 . A,las do Amapa . Rio de Jane iro. 40 pp .
DUU~. ~~~ mw~ ~ior
1941. :-';OC.,s on DtNlrnbo'u '''''", ,,s. Copeia 1941:88 _
93 .
1944 . Notes in Colo mbun herpe to lon . 2. C.ldasi.a
2:391~.
1957 . l' COI:ropic.1 fro, ~ner ...: PrOJ'Iw,,,pi~ venus
Hd 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 in
Colosl~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....hen
N .lurfO~her an Sli u~thieren uoo Re plilien wlihrcOO
der Weltu m""g clu n,. Sr . M.je.lal Fre gaue Nova ra .
Sitzungsbe r. Akld, WI••. W'en 42:383-4 16 .
FoW1H, H, 1913. Amphibi ans and ll.'plile> Irem Ecuador,
Ve ne zue la , and Yuc atan. Proc. Acad. Natur . Sci.
Philadelphia 65 :15.1-176
FUNUIO USEI. J . W. 1956 . New frog' fro m Ecuad o r and
>oulh...'eslern Colo mbi., Zoologic a 41 :73- 80.
G ..N'. C. 1960, NOle. on a herpetol~kal ro llecl inii lrip
through dte !IOulhe:acern lowland s of Bolivl• . Annal.
Carne gie M ..... 35:283_3 14.
G...",.. s , S. 1837 . We>! IOOi. bauachia in the MlI'CUm of
Comparal ....., Zoology . Bull. Esse x lnsl. 19:13- 16,
GIUIlD, C. 11 8~51 1 8504. Abotnct o f 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«ropical
frog . Dmd1"Obalu K"' " .. /ifrnu. Copei.a 197 1:365_370.
Gos."~~b~Y~~ ~~·la-;.,:;m~,li;~edn~~~e :~r i~~~;~lic:ror:.
Herpetologica 16:183-1 90.
G UI8E, J . 1950. Cal,alo~ue de. I}P::' d 'arn phibiens du Mu...eum
Nalll'nal d' HIMOlre Nawrette. Imprimer ie Natlonal e.
Pari" 71 pp.
G Uln'Eso T, A. 11148. ,faun a c hilena:, 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 de
Historia Natural de Samiago, Chile.
GUNTHH, A.C. 18~ 8. Car.alo, ue of the Bauach i. Salienli.a
in lhe:coI1ec1ion of the Brilish M"""um. Brilislt M..." ,um.
lnndon. _vi + 160 pp.
1868. F...... acroun l o f specie> o f l. ilIe•• bamc hian,
addocl 10 the collection of the Brili:$b Mu.eu m. Prnc .
Zool- Soc. London 1868:4711-490.
JI",i" u. Dt: 1" E.S' AO.. , M. 18m . Faunae llCOIropicaiis 'J'CCies
q....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 de
1862 • 1865 por una romisiOn de ll3lunlisas enviada
por e l gobierroo cop;lilol . B.l racios . M ig~1 Gine>t• •
Madrid
K..YAu" S,O. , E.X. AuuQvuQI)!. Ao" O J.E . WU SJ('k
1970. GaPglionic and eardi"" ""lion. of b.alracholoxin.
Eu"'P" " J. Pharmacol . 12; 10-18.
KEFU STf-IN, W. 1867 . Ue ber ., inige neue ode. senene
Balf~h ier
aus Au>tralicn und cem tropischen America.
N""hr , Ges. Wi... GOuin,.,n 1867:34 1_.361.
KlT.. SAo ...O, J ,T . 19 67 . Ob'e rvali o ns on the biolog y o f
Dm ,I", "'" ,. pum iiiu Schmidt and D~"Jrt>""I" " ..raill.'
Giranl . M.S , Thesis. University of Southern Cal ifornia.
87 pp
L." THAM, M. 1966. Capluring ".lIa nge c reature. in Colombia
Nil . Geogr. Mag. 129:683-693 .
L.. URENT , R.. 1942. NOle su r Ie. proc oellen. flrm isterne .
I BaTr.eh 'a Anllt ...) . Bull . Mus . Ro y. 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 Leyden M!I'eUm2.5:83-94
L.VOOECU , H 1974 . Etholo, ische Unlersuc hungen zur
FonpflanlUn, von P lryl/ob<r, rj pal_I.... (Amphibi....
Ran id ae ). Ph .D. Dissert ation. Johanne. G UlCnberg.
Un_n.... zu Mainz. (Abstrae'l)
L.UUSG. K.H . 1971. De. F3rberfrmc h P1n~/ob"'~J bicol<>r
Bibron der O;"d illera Azul (Pt-11l ) . BOnner Zool. Beilr
22:16 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 co llec tion of
the Instiluto O,waldo Cruz. Me m. In.,1. as...·aldo Cruz
50: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. Grogr
Ma, . 111:497-524
MAnl, F. .."0 B. WlTl>or. 1963. TIle wcnom o f lhe CoIom,
bian arrow poison frog PIr, I/obalrs lHcoJor. E.1.perienlill
19:329-376.
~\lO~ ~~en~~B:::~~~~.
MEU" . D. 1,941. Con lributio n. 10 the kIlO. ledge o f lhe
Am~ubi ...o f Sourh Amenu. GOIel>or~ KWlg l. vee.... k.
Vincrll . Samh. Hand!. ser. B, 1(4): 1_71
HU TWOl-E. H.. H, 501""'0 .. "0 A. HE"no-UU . 1965. NOle.
on amp hibian. fm m Ille Ve nez ue lan G uayana. ....ilh
MIR..ND.. Rt Rt.llO , A . DE. 1926 . Nora. p. ra ",rvircm ao
.,..ud<> """ gymnol:>.amochim (A nura) h"a.i tem:... Arch
M... . Nac . Rio de Jane iro 27: 1-227 .
Ta ylor and Franc i•• U>ndon.
"
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' ICijlu ,
fTDS(:hes de. Oal1un~ P")'lIob<lr~J .us &:uador. Sal••
mandra' :81_&4.
Sr;HLUJE!- . H. 1826,: NOTice sur l' e.pelolo, ie de rue de
Java: par M . Bo,e(0II"f~ marluscnl). Bun. Sci . S alw .
Gcol . 9:233-240.
NOBU. G ,K . 1921. FM: new o.pccies of Salienlia from South
Amcrica. Amer . M",", S O'l'iI. 29:1_ 7.
SO<"IDT. K.P. 1932. Replib ~ am phibian> of the MII'IdcI
Venezuelan nped"'"". F.dd MIlS. Nll ur. HUI., Zoo!.
18:1S9_ 163.
19231. Ne.... b.lracllian s from the: trop ical resear ch
.rarion. Brili .h Guian• . Zoologic. 3:289-299
1923b. A new gekkOIlidliurd ... d a new bra<.: hyc<:phl.lid
fro, from Co lombi•. Amer _ Mu• . Novit . 88:1_3.
1926. The peo;1OnI 8in1~ of the bncb)'Cephalid
Amer . M U$, Nom. 23Q;1-14
OlItTT~~':::)I '::U'::~:::r?r.'f~:r~,,:,
_
hop.
(D... Erdbcer.
. 19S3 . Nech mals Dr ndrubaUJ rypogr<lp ltic",,: d• • Erd·
beerfroschchen . Aquai' . Terrar . Z . 6:260 -2b2 .
OEsE. , R . 1932.
l~iIUJ.
~ ZUehl des B.I,l!mlcige " (lHndrobd,u
BUillCf Aqu• . Tmarimlunde 4) ;196-200.
PUUI. H.W. 1935. The frop. Iiurds. IIld snal<n of Brili>ll
Guiana. Pnx. Zoo l. 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... Berlin
1171):641 -652.
. [1873)1 872. Oberdie _ SpU in Bnsilien cnammeltm
Balrachier <in kooil;l. N'l uralienhbinelS ZU Munchen
MOl1 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
SCH" IDT., K.P. A~lI R,F . [NOU . 19S1. Ampllibian s and
reptile 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>loJiKben
Cabinets !II Krak.u. SlIlun r ber . Akad. Wi••. Wien
24; IQ-U .
_ . US8 ~l icj;)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 of
lhe lerTLl0 'Y of a del>ilrobattdfrog. Prn."ltt,,,pi.. ,,mUdlJS
Ecology 41;107-1 IS
Sll.VEIlSro"'E. P.A. 1973. Ob",rv ll ions on the behilvior and
~J:,,=,:.~ ~~ ~~~:a"'~
29'1-301.
19 75. A revision of lhe poi!l(l/l-aml\V fro, s of the genus
lH nd' oltale.< Wagler. Natur. Hisl. Mus. u-.. Angeles
Co" Sci . Bull. 2 1:1_55.
SI'IJI. J .B. VON. 1824 Specie. nov~ R..... rwn 'lUli in ilwre
:~::":~2~ ~ili::ll!'~'~"~~~si~~
en ele
coIle, i! ee~rip>iI . Munich. 29 pp.
STU/ll~ S. R. A ~ D J. HE"'OUC" 50I<. 1919. Field .. udies of
amphibians in Co lombia, 500111 America. uev. Cali·
fornia Pub!. Zool. S6:497 _S40.
ill
ge~.n .
ST F.lNDAr;H~u .
&,~:: glndH~;~:""':: Ie;::,:~~
anden:
PoLDU . W.S . 1973 . Over
~erzorxin,
en
~OO"planlint
~;;:~g~~dh:'~a~~~e~H:~~;~~i~':Z:;:~'::_2e2"
1974a.
O~r ~erzorl;inll
19 7"b . Over
~rzor,in,
en
en
~OOr1plan'in ,
~oorrplanli n.
in
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!
Aquari wn 46:260-266 .
Pos'o~'ou 1~~~B~il.l~: ::eriZi'e~a:,,:,"';!~~o~~fr·
1909, EJ vere no de rana de los indio. del CItocO. Pages
78- 88 in C .A, Mo lina. ed. Esludios cie nlificos de l
doc1o. And~. Po!<ada con .I ,unoo Olro$ escrilOS su)'OS
sobrc d i~en .. lem• • . Imprenl. Of", ia!. Medellin.
Colombia
Rlvuo . J .A. 196 1. Salienli. of VenezlIl:l•. Bull. M... . Comp o
7.00 1. Harvaro 126:1- 207.
RUTHVE"', A .G ,AND H.T . GAIGE. 19l5 . The breedi ng
h.bi" of Pro..,ht,api .....bp .. ncl"I ... Cope . Oc c...
Pap . MIlS. Zoot Univ. Michi, ... IO:I- S.
SAGaA , R. l>E u . 1&41- 1843. Hisioirr physique. polilique .
ee Mlurelle de rile de Cu ba. Alias. ~plile • . Bertrand .
Pm., 31 plales
S~V AOE ,
J . M. 1968 . T ile dendrobatid
America. Copei. 1968;745_776.
fro~ s
of Cenlral
F, 1864 . Balrachologi sche Millheil ungen.
Verb. 7..001. BoI. W . Wien 14:239- 2118 .
1869 . Reise de.- o.1erreichi"c1oen Frepne Nov...... urn
die Etde in den Jahren 18S7, 1858. 18S9. Zoolo~scber
Theil. leo Bmd , 4:Amphibim . K.·K . Hof· und Sl....·
druckere i. Wien
TAYLO Il, E.H, 19S2 , A review of the frogs and toads of
eosla Rica, Univ. Kan.... Sci. Bull. 3S:S77- 942
T EST , F.H . 19S4. Soc ial alll re:. .ivcnes. in an .mphibi....
sceeee 120:140-1 "1.
19S6. Two _
dendrobalid frogs from nonhem Vene.
zuel•. Occas. Pap. Mu s, Zoo l. Univ. Michi,an S77;
,-,
1962, The lIi, hly developed counslli p of rhe fro g.
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. " "'....a r"... . and
parrial »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 . Ven l., van bel kweken mel
lHnd,oba,tJ ,,"'Qt..... L1«rta 31:167- 171.
WASSEl<, S. H, 19~7 . On Drnd,,,""u $.f.o, ..po i ,,~n material
amon g Emper.ii (ChoOOj" l"'aklllg Ind Ians III western
/ 976
FROGS OF t HE GE."'US PHYLLO BATES
CltJda~ . Col omb ia. Arm
I9SS- 1956:73_94
GOlebor,~ Elno" , Mu• .
WlI. "U. F. 18W . Ueh:r Rcpilien 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
53
W' EI>-S n.-"",W . .\I A Zo o 1821. Rei", na: h Bfa'i liell in
den 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. Repi k-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."10
ill Une mi~,ioo bioIojtiqllC beJjlC' aU Brbi!. "Olume 2
Bru>e l lc~
Accepted fur pcblic.uion Apr;1 1.\. 1976,