Terrestrial-Breeding Frogs (Strabomantidae) in PeruWILLIAM E.
DUELLMAN
andEDGAR LEHR
243 illustrations 41 maps 9 tables
I ~ :&I
Frogs Front Coyer Top left i>rrSiim"m ,s .amant! by C. Torres
- righ t Pristima ntis wiens! by E. R, Wild Buttom left
pristimantis percnopterus and rig ht Pristimantis schultei by P. J.
Venegas Back Coyer Top left Pr,Himantis modostichus by w. E.
Duellman - rig ht Pristimantis mendax by M. lundberg Buttom left
Pristimantis phoxocephalus and right Pristimantis exorisrus by P.
J. Venegas
The informal1on. results, etc. eootained in this book an: made
8'"ilable to t~ best of the authors' knowledge and have been
\erified. As tile poSSibility of errors can. hQWC'oer. nOl be fully
Qc1ud.,.-,!, these (\ala are published I'lihOllI any guarnms by Ihe
pubJish~ or authors. Both cannol be held respomiblc fOf lilly
possibly InCOll'eCI stalemenlS. -n All riilht.; rrsencd. in
pal1icular the righlS of mull1plicalion, distribution, and.or
11":I1I51atioli. No portlOll oflhis book may be reproduce1l by any
means (such as pnm, photostalic copy. microfil m. or any other
process) withoutthc explicit writtcn consent ofthc publishers. or
be processed, saved. or reproduced by means o f electronic
systems.
ISBN 978-3-86659-098-4
e
2009 NaTur uDd Ti er - Verlag GmbH
An dtr Klei mannbriieke 39141
0-48157 MilllSlcr WY:w.msveriag.de Layout: Ulrich Manthey,
Berlin Printed by: l.onnemannGmbll. Selm
(.onte ou
Contents
Preface ......................................................
............... ..Acknowledgments
....................................................... ..
56
Introduction
.....................................................
.......................................................................
.Historical Resume ......... ... ... .. .. .. ... ... .. .. ..
......... .... ................................ .. .........
........................... .
8 8
Classification
.............................................................................................................................
10 Reproductive Biology ...................................... ..
.. ... .. .... ...
......................................................... 12
Materials and Methods
.............................................. ....
..................................................... 15 The
Peruvian landscape ............ ................................
........................................................
16Physiography ............................ .. Climate ...........
.. Ecoregions ... .. .. Human Impact on the Environment
..........................................................................................
18... ........ ... .....................................
...................................... 21
22.............. ........ ... ... ..... ... ...................
.. ........ .... 32
Geologic and Climatic History
..................................................................................................
33
Characters of Strabomantid Frogs
..................................................................................
36Meristic Characters .. .... .. .. .. .. ... ..... ... ... .... ..
Morphological Characters ... ....... .. ... .. .. .. .. Coloration
.... ..... ...... ... ... .. .. ... .... ......... ... .. ..
...................................................................................
37 45
61 Trigeminal Nerve and Adductor Jaw Musculature
.................................................................
67 Advertisement Calls
..................................................................................................................
67 Keys to Identification
.....................................................................
.............. ......... ............... 68 Key to the
Strabomantids in the Cordillera Occidental and Pacific lowlands of
Peru ....... 68Key to the Strabomantids in the Cordillera Central
of Peru ............ .. .. .. ................................ 69
Key to the Strabomantids in the Northern Cordillera Oriental of
Peru .............................. 71 Key to the Strabomantids in
the Southern Cordillera Oriental of Peru ... ....... ......... ...
.. ....... 73 Key to the Strabomantid s in the Amazon Basin of Peru
........................................................ 74
Accounts of the Genera and Species
.............................................................................
76 Bryophryne ... .. .. ... ....
............................................................. ..
... .......... .... .................................. 76Hypo
dactylus ................ ....... .... ... ...... ... ..
..................................... .... .. ... .. .. .. ...
.......... ..... ...... ... .. ... Lynchius .. ..........
............. .. ................. .. ... .... ..... .. .. ..
............. .. .. .. ... ... .. .. .......
.................................. Noblella ....... .....
.............. .............................................
....... ... ................. ........................... ....
..... On'obates ...................... ..................... ...
......... ......... .................... .... ............... ....
......... .. ......... ....808690
97
Phrynopus ........................ ..
............................... ..... .... ..... ........ ...... ..
.. ..... ............... .... .............. .. 104 Pristimantis
.............................................................................................................................
.... 127 Pristimantis (Hypod/cryon)
........................................................................................
...... ... 128 Pristimantis (Pristimantis) .............
............ ...... ....... ................. ................ ..
........ ......... ......... 131 Pristimantis ( Yunganastes)
.................................................................................................
257 Psych rophry nef/a ... ... .. ... ...... ........ .....
......... ................... ..... ..... ... .. ......... ....
.......... .................... .. 259 5trab omantis
......................
.......................................................................................................
262
Biogeography ........................ ............ ... ..
................................... .......... ......... ....
....... .............. ... 265The An dean Cordilleras ......
............... ........ .... ............... ............. ..
............ ............................... .. . 266 Th e Arid low
lands ..... .... .....
.......................................................................................................
282 The Amazon Basin .............. ............. .. .............
.... ..... ..... ............. ..... .... .. ... .. ...
............ .. .... ....... .... 282
Conservation
............................................................................................................................
286 Future Research ............................ ..
........................................................ .... ...
...... .. ... ......... ..... . 291 Literature Cited ......... ..
................... .............. ..
....................................... .............. .........
..... ...... 293 Appendix I-Material
............................................................................................................
306 Appendix II-Localities ... .. .. .. ............ .. ... .....
.. .. ..... .... .. ........
............................................ ..... . 320 Appendix
III-Distribution Maps
................................................. .... .... ......
.. ................... 328 Addenda ...............
............................................. ...............
.............. .. ...... ...................................
370Bryophryne .........................................
............................................ " ....
..................................... 370 Pristimantis (Pris
timantis) ........ .. ..... .. .. ... .. .. .. ............. ..
.......................... .. .. .. ..............................
370
Additional new species of Pristimantis
...................................................................................
375Psychrophryn ella ...
..................................................
.................... ......... .............. ....
................... 375
Biogeography ............ .............
....................................................................................................
375 Conse rva tion .......... ............. ...............
............
............................................................................
375
Taxonomic Index
........................................................................
... .. ......... ............................... 376
AcknowledgmentsThroughout the (ourse of otIT iw.esligations we
ha\'f become indebted to many persons. Duellman is gratefollo his
fieldcompanions during many trips to Peru; lhese include Thomas J.
Berger, Bryant W. Buchanan, David C. Cannatelia, Fernando Cuadros,
Dana 1. Duellman, Thomas H. Frills, Javier leachea, David A.
Kilirian, VIctor R. Morales, Michael E. Morrison. Oscar
Ochoa, tHy O. Rodriguez, Antonio Salas, Rainer Schulte, John E.
Simmons, linda Trueb. John J. Wiens, and Erik R. Wild.Duellman's
fieldwork in Peru was supported at variolls times by the Natural
Histor)' Museum at the University of Kan sas and
granls from lhe National Geographic Society and National Science
Foundation. Likewise, Lehr is indebted to Cesar Aguilar.Josi
Boetlger, Mikael Lundberg. Elias Pooce, Rudolf \'011 May. Carola
Ramirez. and Daniel Rooriquez for tlleir aid in the field.
Also. he is grateful for logistic support including housing
provided by his family. Sonia Castello and Godalredo Ramirez, and
lIis friends Emilio Fuentes and Isabel Garcia. l.ehr is grateful to
his parents Brigitte and Erkll L{'hr for their support during his
absence from Cermany. His fieldwork in Peru was supported Ily
Bl0PAT,the Gt':rman Research F oundation, and the Natural History
State Collections Dresden. For lhe loan of specimens and/or
provision of working space in their re~pecti\e C'OIitions. we thank
Cesar Aguilar and JesUs C6rdova, Musco de Historia Natural
Unil'ersidad Naciona! M de San \[arcos: Christopher Austin.
Louisiana Stale ayor University Museum of Zoology: W olfgang Bohme.
Zoologisches Forsc:hungsin!lilut und \Iuseum Ale~ander Koenis; Juan
Carlos Chaparro, Musco de Hisloria Natural Universidad San Anloinio
Abad. Cusco: James R. Dixon, Texas Cooperative Wildlife Collection;
Darrel R. frost and Charles W. M}~rs, American Museum of Natura!
History: W. Ronald Heyer, National Museum of Natural History:
Maureen Ke;J.rney and Alan Resetar, Field Mus!.'um of Natural
Hislor)'. Gunther Kohler. forschungsinstitul und
NaltlrMuseumSenckenberg; Kenneth L. Kr)'S k and Max Nickerson.
Florida Stale .\Iuseum; Jonathan Losos and Jos~ Rosado, o M useum
of Comparative Zoology, Harvard University; David B. Wake. Museum
ofVertebr~te Zoology, University of California: arK! John W Wright,
Los Angeles Count)' Museum. . Lchr's postdoctoral research at the
Unil'ersityof Kansas [February 2005-January 2007) was supported by
a Feodorl.ynen Fellowship gi\~n by the Alexander von
l1umboltltFoundation. Lehr is grateful to Linda Trueb and Craig
Marlin for facilitating that fellowship. Thanks go to personnel in
the D ivision of Hrrprtology-Elisa Bonaccorsa. Rafe
Brown,AndrewCampbell, Rani Diaz, Eli Greenbaum,Juan Manuel
Guayasamin, Kyle Hesed. Chari!.'! Linkem. Oa\id McLeod. Jaime
O,lks. Jennifer Pramuk, Nick Rasmussen, Cameron Siler. John E.
Simmons, Jeet Sukumaran. lind Omar Torr!.'sCarvajal. For logistic
support in L.awrence, Lehr thanks M atthew Davis and Emilia
Barbosa. Lehr thanks the Museum of Comparative Zoology for an Ernst
Mayr Travel Grant and James Hanken and Jose R osado for their
support during his visit. H also thanks the American Museum of
~aturai History for a travel grant and Raoul Bain. Darrel e R.
Frost. Taran Grant, and Oal'jd K izirian for their support. Lehr is
grate(ulto tile Smithsonian Institntion for short term fel lowship
and W R . onald I!eyer. Roy W. McDiarmid. Ken neth Ti~be. .. nd
George Zug for their support during his visit, l.ehr also thanks
the Field Museum of Natural H istory for a travel gtan! and
\faureen Kearney. James Ladonski,Alan Resetar. and Harold Voris [or
their support. Agrant to Lehr from the German ReSl'arch roundauon
(OrC) helped to conduct research at KU in July 2007. and an ..
Iumni grant of Ihe A lexander \'On HumooJdtF aund'lIion gi\'en to
l.ehr in NOI-< ember 2007 aided our efforts to complete this
book. F pr0l-1ding information about speCimens in their
coJlt':c:ions. "'1.' thnl; IgnaCio De la Riva and Jose M. Padial,
Musco or Nacional de Cienclu :'>aturalu: Vrctor R. Morafes.
Museo d~ Historia ~atu ral Unh'ersidad Ricardo Palma; Goran Nilson.
Naturhistorisn \!U5eet GOll.'OOrg: Jasti P. Pombal. ~Iuseu
~aclo!lal Rio de Janeiro; and H ussam laher, Museu de Zoologia da
Uniwrsdade de 530 Pauio. Our owo colvr photographs of Iim;g fJc'g$
were augmented b)t the generosity of se-.'eI'al photogra phers.
including Cesar Aguilar, Cesar L BartioAmor6s..~e5Silndo Catenazzi,
Juan Carlos Chaparro, Luis AColoma. S. Blair Hedges, y. Hooker.
Mikael tundbtr~. CharR) W. M~els. Andres Sc~liJter, Roberto
Sindaco. Karen SiuTins, Claudia T orres. and Pablo J. Venegas. In
addition to Dueliman s pholo:; in the 01211il Archil'es of the
Natural History Museum at the University of KanS3ll, we used some
photos rontained th .. rrin Ibat ....'l're taken ~ William W.
Lamar. John O L . )'och, and Erik R. Wild. Most of the research
....as dl'lne in th~ Dnision ofH~ rpetolo~)' of the Museumof
Naturaillistory and Biodiversity Research Center. Universit)' of Ka
nsas. Personnelth.. rein. Especially Andri'1I' Campbell and John E.
SImmons, facilitated our work Ily packing and unpacking loans and
prQ"idinS neoessal) equipment. We are especially indebted IQLinda
Trueb for her efforts 10 improve our illustrations. sohing computer
problems. and critically reading parts of the manuscript.
Preface
PrefaceIn 1928 Thomas Barbour lamented "It seems wicked and
sinful in this dlly and generation to describe new species of
Syrrhophus.~ After all Eleullierodaclylus and its allies amounted
to 126 species in Nieden's (hecklist published in 1 By 1 the number
of specie~ had grown to 320 (Gorham, 1966) and to 523 in 1
(Duellman. 923. 953 992 1993). Now more than 800 specieS" are
recognized (Hedges el ai., 2008a) . who provided the lirst modern
comprehensive pbylogeny and classification 01 this group of [rogs
that they termed "Terrarana." All Peruvian Terrarana are members 01
the ramily Strabomantidae. Terrararla is an appropriate name for
ttlese frogs. because in the New World tropics they represent a
lineage of frogs that has evolved direct-delelopment of terrestrial
eggs. Their independence from aquatic habitats and absente of
aquatic larvae in meir llfe histories has to be considered a major
"success story," for terraranans hal'e undergone an explosive
adaptive radia, lion tllat has resulted in gnat differences in
size, body forms ranging from heavy-bodied toadJike creatures to
lithe leapers. Gnd diversity of habitats ranging from caves and
subterranean cavities to tbe rainforest canopy. Documenting and
interpreting :hlS diversification has been a challenge 10
biologists. among whom, in addition to us, Ignacio De la Riva, S.
Blair H edges, John D. Lynch. Jay M. Savage. and the late Albert
Schwartz have contributed significantfy to unraveling the
systematic relationships Glterrarana.ns. One of us, Duellnlan. was
introduced 10 the complexityof terraranans more than half a century
ago in Mexico. H became e -.ore and more fascina ted with the
diversity of these frogs as he carried oul intensil'e fieldwork in
Central America and itlrthwestern South America. Subsequent to his
introduction to the Perul'ian fauna in 1971. he continued fieldwork
in Peru Cltil 1994. More than a quarter of ceutur), from the time
of D uelhnan's first Peruvian excursion, Lehr met his first
terraranans '" the Cordi llera Orienta l in central Peru, a region
that he has visited many times since. In 2002, Lehr carried
Peruvian specimens of terrara nans and other laxillo The U
niversity of Kansas [or comparison with .spimens in lhe
herpetological collection. This was our first meeting, in which we
shared our personal knowledge of Peruvian ftoss. We soon realized
that our rcsp ecti~'e data were complementary and represented 11
wide atTay of undescribed species . . notable distributional
rec.ords. We decided to pool our data, seek additional information,
and to collaborale in preparing Ute material for publication. For
the past six years most of our efforts have been directed to this
project. Thi~ book is the "!'.sul! of those endeavors. Herein we
review the existing information on Peruvian terraranans with
respect to the morphology, coloration, and distri.oon of each of
the species. Throughout the course of our work, new species were
discovered in museum collections thnl -re studied mostly by L and
additional new tax~ were collected by Peruvian colleagues and
shared with us. These have ehr, !Iet'II described in an array of
papers published in the last six )ears. In this boox we endeavor to
provide detailed accounts of each of the described species and keys
to their identification, that uep.:eceded by a detailed description
of the taxonomic characters of strabomantid frogs. Distribution of
species is ascribed rwregiol1s defined in a section on the Peruvian
landscape and s)'nt!iesized in a seclion on biogeography of stra
bomantid .... 1$. which precedes lists of specimens. the localities
where they were collected. and distribution maps. We. like our
col~\IeS around lhe world, are well al~are of the declini ng
populations and app.irent extinction of mall)' species of frogs .
~se so little is known about populations of strabomanlid frogs in
Peru, our attempt to consider conservation efforts is rttremely
generalized. ItVlould be gratifying to assume that this is a
lasting cnmpelldlum of knowledge of these frogs in PerIJ, but it
certainly is tbe case. Instead, this book should be considered
snlely as what it is-a first step in what we visualize as a long
road of ~io ns, hopefully by Peruvian biologists, to\l'llrd a
meaningful s),nthesis of this fasci llating group of frogs in a
hiqhly ~ landscape.E/eulllerodocc)'lus and its ally
AcknowledgmentsThroughout the (ourse of otIT iw.esligations we
ha\'f become indebted to many persons. Duellman is gratefollo his
fieldcompanions during many trips to Peru; lhese include Thomas J.
Berger, Bryant W. Buchanan, David C. Cannatelia, Fernando Cuadros,
Dana 1. Duellman, Thomas H. Frills, Javier leachea, David A.
Kilirian, VIctor R. Morales, Michael E. Morrison. Oscar
Ochoa, tHy O. Rodriguez, Antonio Salas, Rainer Schulte, John E.
Simmons, linda Trueb. John J. Wiens, and Erik R. Wild.Duellman's
fieldwork in Peru was supported at variolls times by the Natural
Histor)' Museum at the University of Kan sas and
granls from lhe National Geographic Society and National Science
Foundation. Likewise, Lehr is indebted to Cesar Aguilar.Josi
Boetlger, Mikael Lundberg. Elias Pooce, Rudolf \'011 May. Carola
Ramirez. and Daniel Rooriquez for tlleir aid in the field.
Also. he is grateful for logistic support including housing
provided by his family. Sonia Castello and Godalredo Ramirez, and
lIis friends Emilio Fuentes and Isabel Garcia. l.ehr is grateful to
his parents Brigitte and Erkll L{'hr for their support during his
absence from Cermany. His fieldwork in Peru was supported Ily
Bl0PAT,the Gt':rman Research F oundation, and the Natural History
State Collections Dresden. For lhe loan of specimens and/or
provision of working space in their re~pecti\e C'OIitions. we thank
Cesar Aguilar and JesUs C6rdova, Musco de Historia Natural
Unil'ersidad Naciona! M de San \[arcos: Christopher Austin.
Louisiana Stale ayor University Museum of Zoology: W olfgang Bohme.
Zoologisches Forsc:hungsin!lilut und \Iuseum Ale~ander Koenis; Juan
Carlos Chaparro, Musco de Hisloria Natural Universidad San Anloinio
Abad. Cusco: James R. Dixon, Texas Cooperative Wildlife Collection;
Darrel R. frost and Charles W. M}~rs, American Museum of Natura!
History: W. Ronald Heyer, National Museum of Natural History:
Maureen Ke;J.rney and Alan Resetar, Field Mus!.'um of Natural
Hislor)'. Gunther Kohler. forschungsinstitul und
NaltlrMuseumSenckenberg; Kenneth L. Kr)'S k and Max Nickerson.
Florida Stale .\Iuseum; Jonathan Losos and Jos~ Rosado, o M useum
of Comparative Zoology, Harvard University; David B. Wake. Museum
ofVertebr~te Zoology, University of California: arK! John W Wright,
Los Angeles Count)' Museum. . Lchr's postdoctoral research at the
Unil'ersityof Kansas [February 2005-January 2007) was supported by
a Feodorl.ynen Fellowship gi\~n by the Alexander von
l1umboltltFoundation. Lehr is grateful to Linda Trueb and Craig
Marlin for facilitating that fellowship. Thanks go to personnel in
the D ivision of Hrrprtology-Elisa Bonaccorsa. Rafe
Brown,AndrewCampbell, Rani Diaz, Eli Greenbaum,Juan Manuel
Guayasamin, Kyle Hesed. Chari!.'! Linkem. Oa\id McLeod. Jaime
O,lks. Jennifer Pramuk, Nick Rasmussen, Cameron Siler. John E.
Simmons, Jeet Sukumaran. lind Omar Torr!.'sCarvajal. For logistic
support in L.awrence, Lehr thanks M atthew Davis and Emilia
Barbosa. Lehr thanks the Museum of Comparative Zoology for an Ernst
Mayr Travel Grant and James Hanken and Jose R osado for their
support during his visit. H also thanks the American Museum of
~aturai History for a travel grant and Raoul Bain. Darrel e R.
Frost. Taran Grant, and Oal'jd K izirian for their support. Lehr is
grate(ulto tile Smithsonian Institntion for short term fel lowship
and W R . onald I!eyer. Roy W. McDiarmid. Ken neth Ti~be. .. nd
George Zug for their support during his visit, l.ehr also thanks
the Field Museum of Natural H istory for a travel gtan! and
\faureen Kearney. James Ladonski,Alan Resetar. and Harold Voris [or
their support. Agrant to Lehr from the German ReSl'arch roundauon
(OrC) helped to conduct research at KU in July 2007. and an ..
Iumni grant of Ihe A lexander \'On HumooJdtF aund'lIion gi\'en to
l.ehr in NOI-< ember 2007 aided our efforts to complete this
book. F pr0l-1ding information about speCimens in their
coJlt':c:ions. "'1.' thnl; IgnaCio De la Riva and Jose M. Padial,
Musco or Nacional de Cienclu :'>aturalu: Vrctor R. Morafes.
Museo d~ Historia ~atu ral Unh'ersidad Ricardo Palma; Goran Nilson.
Naturhistorisn \!U5eet GOll.'OOrg: Jasti P. Pombal. ~Iuseu
~aclo!lal Rio de Janeiro; and H ussam laher, Museu de Zoologia da
Uniwrsdade de 530 Pauio. Our owo colvr photographs of Iim;g fJc'g$
were augmented b)t the generosity of se-.'eI'al photogra phers.
including Cesar Aguilar, Cesar L BartioAmor6s..~e5Silndo Catenazzi,
Juan Carlos Chaparro, Luis AColoma. S. Blair Hedges, y. Hooker.
Mikael tundbtr~. CharR) W. M~els. Andres Sc~liJter, Roberto
Sindaco. Karen SiuTins, Claudia T orres. and Pablo J. Venegas. In
addition to Dueliman s pholo:; in the 01211il Archil'es of the
Natural History Museum at the University of KanS3ll, we used some
photos rontained th .. rrin Ibat ....'l're taken ~ William W.
Lamar. John O L . )'och, and Erik R. Wild. Most of the research
....as dl'lne in th~ Dnision ofH~ rpetolo~)' of the Museumof
Naturaillistory and Biodiversity Research Center. Universit)' of Ka
nsas. Personnelth.. rein. Especially Andri'1I' Campbell and John E.
SImmons, facilitated our work Ily packing and unpacking loans and
prQ"idinS neoessal) equipment. We are especially indebted IQLinda
Trueb for her efforts 10 improve our illustrations. sohing computer
problems. and critically reading parts of the manuscript.
(a ili ng male Pri5timanti5 petrobardu5 photographed by Karen
Siu-Ting at Monte Seco, Cajamarca, Peru.
IntrodlKtlon
Introduction
~--many new genera aOO speeies from Ecuador; it was preceded by
two shari papers (1871 , 1812) in lI'hich other species lIere.
described. Altogether in these. publlcaliolts, Jimenez de la Espada
described three new genera inr.!ul1ing fOllr new species Ihat are
now known 10 occur in Peru-Cyclacephalus (= Prislimoflfis)
{acrimosus. fly/odes (""Pristimalltis) diadema IuS. Qreobnu!s
quixer/Sl$, and Prislimolltis goldi. The first straootn.lntid frog
recorded from Peru is H)'/odes (= Strabumantis) sulcatus described
by Cope in t864 ; the specimen was collected at Nauta on tile Rio
Amazonas (Departam ento de !.oreto) by a major Anlericatt
expedition led by James Orton. Nearly a decade passed until a
serond species was named from Peru; this was Phrynoplis peroanus
described b)' Wilhelm Peters io 1873 and based on a specimcn
obtained by 11m. Jelski at "Maraynioc in Peru" (Departamento de
Junin). The only other strabomanlid frog described in the 19t1l
Century and now known to occur ill Peru is Hy/odes (= hislimon/ls)
w-nigrum: it was named by Oscar Boettger in 1892 ilnd based on
specimens collected by C. f'. lehmann al ~ Zurucuchu" (Provincia de
Aluay), cuador, and placed in the Senckenbergischen
NaturforscheOllen Oesetlschaft 'In Germany. Thus by the beginning
01 the 20 th Century only nine species of strabomantids now known
10 occur ill Peru had been described. OUTlIIG the early pari of the
20 111 Century sCIotlral professional co!leclors, amons them G.
Ockenden, M. G, Palmer, and P. O. Simons, were sending specimens
from Ecuador and Peru to Ihe British Museum (Natural History),
where George A. Boulenger named scores of new species bel ....'een
1900 and 1920. including six species of Hylodts ('" Prislimontis)
and Pa/udicolo ('" Prislimantis) simo"sii. In the early 1910s
Thomas Barbour 01 H~TViIrd University in the United States
initiated ,tudies on amphibians and reptiles in western South
America_ An expedition 10 lIorthern Peru (Cajamarca and PiuTa) by
Han'ard Unil'ersit}' resulted in the discovery of
Eleulherodact)'/us (: Pristim(JJllis) cnjamorcensis and
EII!Ullierodacty/us ('" hislill/anlis) /ymanl described by Barbour
lind G. K. Noble in 1921. Barbour also collaborated with Emmet R.
DUlin in tile description of Eleulheforiac/ylu$ (Pristill/alllis)
altamo.zonicus from Nauta, Departamento de Loreto. Noble also
discovered specimens in the American Museum 01 Natural Histor)'
from "Juliaca,ft Peru. that in 1921 he named Smilhillus (=
Noblello) perullia"us. In the [930s Benjamin Shreve named two
species of S)"frhophus (= Pllf)'nopus) from Cascas, Departatnentn
de Junin, Peru, aud two species of l::leulherodaclylus (=
Pristimolllis) from Amazonian Ecuador that subsequently also were
found in Peru. In 1938 Hampton W Parker named /eutherodac/)'/us
(""
About half of all of the specics 01 frogs known to inllabit Peru
belong to the lami~' Strabomantidae. These are small to
medium-sized frogs with head an'''''''.~
"
1 00
* i ~~..
01
-
j
~ ~" li.sl ij~\) rk \\'as in Departamento de Lorf:Io:~ from
there contained diverse slrabomantid !lop ~ ~~ of two nell' species
described b!; Il-f,j/a:aJ; a.&3Jo..~b R. 'Iendelson [!J in
1995.
these frogs. The phylogenetic analyses were expanded and fo rmed
the basis for Ihe present classification (H edges et
al.,2008a).
Bj ttue lIlil-I=~' ~ ra..'lOUS Peruvian biologists wcre
studyl.Epll!w.~. In- 19S! lily O. RodriglJcz was the first
f"tranurodes.:ribe a nel' species of strabomantid frog; she
~.!OOtner In 199i "''ith Glenn flores. In 1 S. Blair 987 Hec~ marl
.. ilsma!l tollection of frogs from cloud forests in ~t"dr Pasco,
Peru. Remarkably, this small collee tiM mlaInrd ;even new species
ofPrislt"manris and three of ~"'1OPU{ described by Hedges in 1990
and by Duellman and Heo:.lges 1 200$. 2007, 200Sa). ft suggested
that the Andcs in central peru harbored a high diversity of
strabomanlid frogs. lehr initiated his field studies in Peru in
1997. Herpetological inrestigations along a trans-Andean transect
was tile tOpiC of his dissertation (lehr, 2002) . In 2000 he began
concentrating on the Cordillera Oriental in central Peru. His
fieldwork. together with various field associates from the Museo de
Historia Natural llniversidad Nacional M de ayor San Marcos (MllSM
Lima, principally Cesar Aguilar and ), M ikael Lllndberg, resulted
in the discovery of many new species of frogs of
thegeneraH)'podoctylus, Nob/el/a, Phrynopus, andPrisllinal[fis that
have been described by L and collabehr orators in 2002-2006.
Note....,orthy are ten new species 01 Pllrynopus which were named
by L and colleagues during ehr this time from central Peru ;
tissues of these were the basis for the first phylogeny of this
genus (Lehr et aI., 2005), whidl resulted in some new taxonomic
combi nations. Subsequent revision 01 Phf)l/opus let to further new
combinations and synonymys (LehT, 2006; Lehr et af.. 2006). Jose M.
Padial and Ignacio De la Ril'a and colleagues have worked primarily
on the frog fauna of Bolivia, but in collaboration with Juan Carlos
Chaparro in Cusco have made important discoveries of anurans in
central and southern Peru. The collaboration of Duellman and Lehr
on studies of strabomantid frogs in Peru began in 2003. Subsequemly
as a Humboldi Researchl'cllow, Lehr worked at the Natural History
MUseum and Biodiyersity R esearch Center at The llniversity of
Kansas (2005-2007) and traveled to major museums in tht United
Slates. Europe, and Peru, during which time w e studied hundrrds of
museum specimens and described many more sprcies from Peru (e.g.,
Duellman and Lehr, 2007: Lehr, 2007bJt thereb) bringing the number
of named strabomantid frogs known in PeTti 10 161 species, a
remarkable increase in the number of speCies during the last
halfcentury. While working wilh Lehr on the peruvian material,
Duellman COllaborated with S. Blair H edges and Matthew P. Heinicke
on the ph~10geny of "elcutherodactyline frogs.H Their preliminary
analysis of molecular data (H einicke el al.. 2007) resulted in the
disr.ol'ery of fou r distinct clades 01 it;
Classificationfrogs of the genus 'feulherodoC/ylus" have had a
confusing taxonomic history. During the latter part of the 1 1h and
9 early parI 01 the 20 th centu ries they were known as H ylodes. a
generic name proposed by Filzinger (1826) for the species Hyfa
ralloides Spix, 1824 (0: flyla nasus Lichtenstein, 1823); {his
species, now known as Hrlodes nasus (Hylodidae of frost et al.,
2006), inhabits the Atlantic Coastal forest in southeastern Brazil
(Heyer. 1 982). Stejneger (\904) deter mined that the generic name
Elelltherodaciyl(/s was apphca. ble to frogs that previously had
been referred erroneously to H ylodes: this dilemma also was add
ressed by Myers (1962), who supportrd Stl'jnrger"s conclusions,
among whidl is the. recognllion of Eleuth@rodaclyfus marlinicensis
Dumeril and Bibrnn. 184!. as the type species of the gen us. As
would be expected with an), widespread and variable group of frogs.
numerous generiCnames have been applied to species of
E/@utherodact...-Ius."' Most of these were synonymized with
Hylodrs" bl' BouJenger (1882), who placed /fy/odes ('"
Eleutherodaclylus) in the fami ly Cystignathidae, which was basrd
on Cyslignalhus Wagler, 1830, an objective synonym of Leptodacty/us
(see L ynch, 1971, lor an historical revie.....). Thus.
Efeuth(rodacty!us' and allied gellera have been placed in
lep(odactylidae; in some cases they were classified as
Eleutherooactylinae (Lutl, 19j4; Laurent, 1 980; Duellman, 2003) Qf
as the tribe Eleutherodactylini in the subfamily Telmatobiinae
(Lynch, 1971 ). foll owing Starrett's (1908) di5cOI'ery of
different pathways of the trigeminal nerve with the adductor jaw
musculature, Lynch (\986a) recognized a primarily Middle American
dade of 'Efeufherodacl),lus, ~ as the subgenus Craugoslor Cope
(1862): based on analysis of nuclear and mitochondrial genes.
Crawford and Smith (2005) elevated the clade to generiCstatus.
Asmall group of "eleutherodactyllnes' hay ing knoh-shaped terminal
phalanges and lacking toe pads in the Andes and upper Amazon Basin
had been recognized as the genus Ischnocnema, but Ca ramaschi and
Canedo (2001i) demonstrated that the type species of /schnocnema
(Leiuperus llf'mICO.WS R einhardt and Liitken, 1862) has Tshaped
termina l phalanges and toe pads and placed the species in
'feutherodactylus." They resurrected Oreoboles Jimenez de la
Espada, 1 for the Amazonian and Andearl 872. frogs formerly placed
in/sc/JnoclIma. Analyses of mitochondrial genes of few taxa b}
Ruvinsky and Maxson (\996) and Darsla nd Cannatella (2004)
indicated Ihat L eptodactylidae was not monophyletic; fu
rthermorc,
Classification
the latter authors shov..ed Bmcirycepholus tphippillJTl 10 be
Imbedd~d in Ihe Eleutherodactylldae. This led Dubois (2065) 10
place genera formerly in Eleulherodactylillilf! in the older fa
mily-group name, Brachycephallnae, coined by Giiuther iJ858). In a
lensthy work purported to be nThe Amphibian Tree of lire" Frosl 1'1
a1. (2006) constructed a new classification of Recent amphibians
based on varying numbers of genes of pitifully few taxa. They
placed alt eleutheroelactytlne g,enera in Brachycephalidae; most of
these placements .,,'ere inferred inasmuch as their data set
rorllalned only 17 species in eight of the 17 genera thaI they
recognized. The major advance in phylogeny and classification was
11)Heinicke et al. (2007), whose alla~"Sis of nuclear and
mitochondrial genes of 2i6 speties of "ele utherodaclylines~
revealed well-supported clades of biogeographicsignificance. rhu!!
they recognized the major MiddleAmeriC clade as lantar view of foot
(Fl '" loot le ngth). C. latera l view 0 1 head (EN '" eyenoslril
dist ance, ED " eye dia meter, TV s tympanum diameter),
A
-1
INO
8
--I1
~I-------
SVL
o ' 0 01, fJI.
1 1 1
IFL1 1 1
v
1 1
~-
/ j
I I I
~-
TY
c
' - , -- - - - )
Characters of Strabomantid Frogs
Snout-vent length (SVL) is the distance from the tip pf the
snout to the posterior end 01 the hody; this me-asurement normally
is made with a specimen flal on a horizontal surface. Tibia leng th
(=shank length) (n ) is the distance from the knee to the distal
end of the tibia. Foot le ngth (FL) is the distance from the
proximal margin of inner metatarsal tube rcle to Lip 01 foe IV.
Head length (HL) is the distance from angle of the jaw to the lip
of the snout; this is the cllord of head lengtll as defined by
Lynch and Duellman (1991), who estimated head length to be the
straight line distance parallel 10 body axis from angle of jaw to
point equal to the tip of snnut. Head w idth (IIW) is the greatest
widtll of the head, usually at the level of the angle 01 tile jaws.
Interorbital distance (IOD) is the breadth of Ihe braincase between
the orbits. Width of upper eyelid (EW) is the perpendicular dista
nce to the outer edge 01 the eyelid. lnternarial distance ClND) is
the straight line distance between the inner edges of the na rial
openings. Eye-nostril distance (EN) is the straight line distance
between anterior corner of orbit and posterior IlliIrgin of the
narial opening. Eye diameter (ED is the 1I0rilontailength ollhe
orbit, ) which is the length of visible eye. Tympanum diameter (TO)
is the horizontal distanr.e be!II-'een the peripheral borders of
the tympanic annulus.
108 8
-! E "w w ro
6
4
2 -
c. ~."2VE , 0 10 20 30
~.CJ.
EAI D~40Snout-vent length (mm )
Strahomantids are small to larse fross. Females attain longer
snout-vent lensths (SYL) than males of the same species: the amount
of sexua l dimorphism (females SVU male SYL) is highly variable.
Tile least amou nt 01 sexual dimorphism exists in smaller species,
such as in Nob/el/a, in which males are only slightly smaller than
fem ales. whereas in Siroboman{is sulcatus males are only about
one. half the size of females (Table I). Few data are avaHable on
the m~sses of strabomantids, but Duellman (2005) provided data on
five species in Amazonian Peru; this small data set reveals Ihat
mass increases proportionately with size and that differences in
mass and snl)utvent length are greater In la rger species than in
smaller ones (Fig. 36). Maximum known snout-vent lengtlls (SYLs) of
each species are given in Table 1. Interspecific differences in
body size are best made by comparing the maximum SYLs of females.
By so doing, it is obvious tha t Nob/elfa are among the smallest
slr~borllan!ids, with maximum SVI,s of femal es ranging from 12.4
mm in N. pygmaea to 20.2 mm in N, IYflclli. Among the terrestrial
strahomantids, the smallest Ps)"chrophryne/fa is P. boellgeri with
a maximum SVL of 18.4 mm, followed by 29.3 mm in 8ryophryne
cophiles and 43.3 mm in L YIICllills (/OIJomaculatus, whereas the
largest Phrynopus is P. kmlJleOrUm, which attains ~ maximum SVL of
56 .~ mm. These small sizes arc similar to those of the ter
restrial specit!s in tile Prislimantis orestes Group, in which the
smallest.P. seorsus, atta ins a maximum SYL of 20.9 mm and the
largest, P. simonsii, reaches 33.3 mm. Bowel'er, some members of
the Pristimaillis unis/rigailis Group are equally small; the ani),
known mature female of P. corona/us has a SVL of 1 mm, and females
of 5.3 the Amazonian leal litter inhabitant P. /fachyblepharis only
attain a SYL of 19.2 mm. In contrast, the SVI,s of adu lt fe males
01 Strabomalllis sulcatus, two spec ies of Oreobales, and eigh t
species of Pristimanris exceed 50 mill. Most species in the
Pristimanlisfig . 36. Relation~l"1lp betw een snou tvent len gtl"1
and mass ;n five specie of Pristimanti5 in Madre de Dios, Peru. So
lid circles are means of mate; open circles are mea n~ of femate ;
lin es afe ranges . A. p, ailamazoniws (3 males, 1 female). B. P.
feneHratu~ (1 9, 19). C. P. imitatri)f (9, t2) . D. P peruvianU$ .
(23. 12). E_ P. taltae (36, 24).
50
Meristic Charact ers
Ta bl e 1. Sele ct ed morpho logi cal f eatures an d ma ~i mum
known s no u t~e n t lengths (SVL) in mm of straboma nt id frogs i
n Peru. SO " se~ua l d imorp hi sm. SVL is max imu m lor each se~ .
Tympa num con dition: A " membra ne d iffere nti ated and ann ulus
prom inent , B '" membrane not diff erent iat ed bu t most of circ
umference of annulus v isible t hrough sk in, C = membrane not
differe nt iat ed and annu lus evid ent o nly ve ntrally. 0"
membrane and annul us absent. Toe (On dition : A " Toe II I = Toe
V. 8 = Toe V :> Toe III but not extendin g to penu ltimate
subarticular tu bercle 0 1 Toe IV. C '" Toe V: Toe HI. 0 " Toe V
< Toe II I.
Species
Tympanum
Fingers
Toes
SVL(male; fem ale)
SO1.05 1.29 1.16
Bryophryne B. bustamantei B. cophites B. nubilosus Hypodactylus
H. araiodactylus H. fallaciosus H. lucida H lundbergi H. nigro vi
ttarus Nob/ella N. duel/man i N. heyeri N.lynchi N. myrmecoides N.
peruana N. pygmaea Lynchius L. flavomaculatus L. nebulanastes L.
parkeri Oreobates o. crura lis o. granulosus o. lehri o. pereger o.
qu ixensis O. saxati/is Phrynopus P. auriculatus P. ayacucho P.
barthlenae P. bracki P. bufoides P. dagmarae P. heimorum P.
horstpauli P. juninensis P. kaun eorum P. kotosh
D
0D
111 111 1>11 1>11 1 size for body propo rti ons wit h lam
ple lizE> for se xual dimorphism In pare nt heses; ran ges (m ea
ns o f Individ ual specie s) are fo llowed by means fo r Ihe group
in p,enler is uniform Ian in P. orl/cucho. Ps),cllrophrynello
boeugeri also has a tympanum, but P/lryf/OPUS pemOflU,, is much
larger Ihan P. boettgeri (SVL 10 18.3 mm), which differs further
from P. peruanus bylacking nuptial padsand dentiger ous processes
of Ihe mmers. Distribution and ecology: Phrynopus perva /IUS is
only known from Ihe type locality in the puna at Maraynioc at an
elevation of 3825 m in the Vitoc Valleyin the Cordillera Oriental
in central Peru. Males and females were found inside bunches of
Peruvian feather Grass (Slipo icchu) next to a small stream in a
swamp)' area during the day; males were calling from inside bunches
of srass. - Remark s: More than a century passed between the
description of Phr)"f/opus peruanus ( Peters Ig74) until L)'nchs
(l975) redescription of the species based on two specimens (UM MZ
89477) from Maraynioc. Coloration in life remained unknown until
Lehr (2007a) described male characters, coloration in life for
males and females, and the habitat. Sexual dimorphism in dorsal
skin texture and colOfation exists in Phf)"IIOPUS peruonus. Males
bal"C tuberculate skin on the dorsum with the tubercles coalesced
into prominent ridges dorsally and dorsolalerally; females also
ha>,c a tuberculale dorsum, but the ridges are less prominent
than in males. The orange coloration apparent~ is restricled to
females. The only other Peruvian PhT)"nopus displaying sexual
dilTlOlp/lism in coloration is P. pe5(lmesi. Males of that species
M\"e)"ellowblotcht's on the venter and in the groin, whereas
females lack yellow blotches (Fig. 107). The specific name.
pUUOIIIIS. alludes to the countly of origin.
f ig . 107. Phrynopus pesimfesi. MUSM 19857, mate, 25.6 mm
5VL.
laguna Quimaccocha, Pasco, Peru. EL.
Phrynopus pesantes; Leht, Lundberg, and AguilarF"i!ure tnt. Map
11Phrynopus ~$QJ//t..; Lehr. LURdllerg, and Aguilar, 201)5:48&.
Ilotot}pe: MUS~1. adutl mate. from near Laguna QUimaccocha, to
33'DJr S. i5 5fOU" II'. 4390 rn. :1"8 km [ai rline) NN E
Huachon, Dep1lrlamenlO de Pasto. Peru.
Characteri stics: This species of Phrynopus is characterized by:
(I) skin on dorsum and flanks bearing loll', IIlherc1es; skin on
venter areolale; dorsolateral folds and discoidal fold absent; (2)
tympanic membrane not differentiated; tympanic annulus absent: (3)
snout short, rounded in dorsat view and in profile; (4) upper
eyelid lacking tubercles. narrower than 100; cranial crests absent;
(5) dentigerous processes of vomers absent; (6) vocal slits and
nuptial pads absent; (7)
Finger I slightly shorter than Finger II: tips of digits
rounded, not expanded; (8) fingers lacking lateral fringes; (9)
ulnar tubercles present; ( 10) heel and tarsu s lacking tubercles
and folds; (11) inner metatarsal tubercle ovoid, about 2x round.
outer metatarsaltuhercle; supernumerary plantar tubercles present;
(12) toes lacking lateral fringes; webbing absenl: Toe V slightly
longer than Toe!lJ; tips 01 digits rounded; (13) dorsum olil'eSTay
with gray and black flecks; venter dark brown with graymollling;
(24) SVL in males 19.6-25.5 10m. in fema les 21.9- 32.7 mm . -
Coloration : In life, the dorsum is dark olive-gray with small
black marks; a black canthal stripe is present. but an interorbital
bar. postorbital stripe, and labial bars arE' absent. The flanks
are paler than the dorsum, changin ~ from olive dorsally to pale
gray with black marks ventral ly. The throat is dark brown to black
with )'ellowish gret: blotches anteriorly and pale gray blotches
posteriorly: t ~"
Phrynopus
chest and belly are dark brown with small pale gray spots.
central Peru. where the frogs were under stones near small Da rk
yellow blotches are present in the groin and on the ponds. One
female contained 26 oyarian eggs. A male called ventral surfaces of
the forelimbs. chest at the base of the at night whell in captivily
with a female; 11 emitted a release forelimb. posterior part of the
belly and distal parts of the call when touched. thighs. The palmar
and plantar surfaces are dark brown R emarks: No olher species of
Peruvian PI/fynopus IS with the outer side of the inner palmar
tubercle and outer known from elevations of nearly4100 m. The
speiJll' ie annulus smooth, prominent, slightly higher than long,
its length slightl\' more Ihan 5096 length of eye; (3) snout long,
rounded in dorsal view, protruding weI! beyond margin of 10wI'r
lip, aClltely rounded in profile; (4) upper e)'elid I~cking
tubercles, narrower than 100: cranial crests absent; (5)
dentigerous processes of vomers prominent, oblique; (6) condition
of vocal slits and nuptial pads unknown; (7) Finger I
equal in length to II: discs on outer fingers expanded, trun
eate, about twice width of digi t proximal to pad: (8) fingers
lacking lateral fringes; (9) ulnar tubercles absent; (10) heel and
outer edge of tarsus Jacking tubercles; inner edge of tarsus
bearing low, e1l1ptical lubercle distally; ( 11) inner meta tarsal
tubercle elliptical. about 3)( subconical outer metatarsal
tubercle; supernumerary plantar tubercles absent; (12) toes lacking
~teral fring!'s; wehbing absent: Toe V slightly longer than Toe
III; discs sHahtl)' smaller than those on outer fingers; (1 dorsum
tan wilh small dark brown spots; venter 3) creamy ta n with diffuse
brown reticulations 011 belly and ven tral surfaces of thighs:
posterior surfaces of Ihighs brown; (14) SVL 29.1 nlln in one
female. - Coloration: The colors in life are unknown. In preserva
tive, the dorsum is tan with dark brown markings consisting of many
small, round or ovoid spots on the back and flanks, The canthal and
postorbital stripes are barely evident, as are the transverse mark
above the vent. small spots distal lyon the anterior surfaces of
t~e thighs. and narrow trans verse bars on the limbs. Abroad creamy
tan lahial strlpe is present. The posterior surfaces of Ihe thighs
are dull brown. The throat is creamy tan wilh minUle brown flecks;
the belly and ventral surfaces of the thighs are cream)' tan with
dif fuse, f3 int bt(lwn reticulations. and the venlral surfaces of
the tarsi are black. - Compa riso ns: Among the Perm'ian species in
the Prisliman/iS conspicillalus Group cllaracterized by having Toe
Vonly slightly Iooger than Toe III, P. cum!lrostris differs from
all others by having a long, a wedgl"sllaped snout; it is like P.
consp{cillatus and P. penwiollus in having a pale labial stripe,
bul it differs from lhose species by having brown retic lriations
on the \'en\er and hy lacking pale flecks on the posteTior surfaces
of the thighs. Prislimantis aL'icuporom and P.
Pristlmantis (Pristim.lntis)
sRytimoinos diffcr further from P. cuneiros tris by having an
inter{ltular fold and ".,eak-
Iy areolale skin on the ventcr in contrastto smooth skin
ventrall)'; P. buccinator differs further from P. cUlleiroslris by
having pink spots in the groin and 011 the hidden surfaces of the
thighs. Other Peruvian Prislimalltiswith smooth skin on the venter
include four large species (P cirriogasler. conrior, t;monf. and
w-nigrum; males to 46 mm, females to 73 mm). Among the smaller
species in the upper Amazon Basin and on the lower slopes of the
Andes-P. odl~
astolus, bipunctotus, conspiciflolUS, lUI/thanita mall/ini,
melobares. aoo peruL"ianus-the posterior surfaces of tile thighs
are dark brown with cream or red flecks, and three of these (P.
bipllncratus, ma/kim: and metabates) hal'e ba!XIllI'ebbing Uetween
the loes. The posterior surfaces of the thighs are uniform brown in
P.
Fig. 150. PrlstlmantiS danae, I(U 162309, female, 44.9 mm SVL, 4
km SW S~nta
a//)ertus, lenl'Strotus. maMoiaemus, stic ISilbet, CUKQ, Peru. W
ED. logaster; aoo L"ilarsi. Furthermore. P. IOJ/Ihanites has a
median pale streak on a dart throat and a pronl1' dorsolateral
folds weak; (2) tympanic membrane and tym.nent tubercle on the
heel. Distribution and ecology: This species is known only from the
Iype locality at 1700 III in humid mont~ne forest all the western
slopes of the Cordillera Colan, Depart~mento de AmalOnas, Peru.
Remarks: Among the strabomantid frogs tn Ecuador and Peru. the
long, I\'edge-shaped snout of this species is approached only by
the Inng, subacuminate snout of Craugastor {ongirostris in the
Chocoan region of Ecuador and exceeded by the Ilearly spatulate
snout in the small Pl1r),nopus fecliriorliYllchus in the Cordillera
oriental in cen tral Peru. The specific name is an adjettive from
the Latin cuneus meaning wedge and the Latin roslrum meaning snnut;
the name refers to the wedge-shaped snout (Duellman and
Pramuk,1999), panic anllulus prominent, round, ils length about 50"
length of eye; (3) snout long, sloping, rounded in dorsa! view and
in profile: (4) upper eyelid lackillg tubercles, slightly narrower
than 100; cranial crests absent; (5) dentigerous processes of
vomers triangular, prominent; (6) vocal slits present; nuptial pads
preselll on dorsal and dotsomedial surfaces of s....,oI len thumb;
(7) Pinger I soorter than Finger II ; discs truncale, 2.5x width of
digit proximal to disc; (8) fingers bearing weak lateral fringes:
(9) ulnar tubercles absent; (10) heel and tarsus lacking tubercles:
iuner tarsal fold on distal half of tarsus; (11) inner metatarsal
tubercle large, fl~t, elliptical, about 8>< minute subconical
outer metatarsal tubercl e; supernumerar~ plantar tubercles absent;
(12) toes bearing later~1 fringes; webbing basal: Toe V much longer
than Toe !II; discs smaller than Ihose on fingers; (13) dorsum
brown with dark brown chevrons: \'enter while with or wilhOlJt
black spots: hidden surfaces of thighs brown with cream flecks;
(14) SVL 27.3-33,8 mm in 44 males, 29.2-45.8 min in II females. -
Coloration : tuhfe, the dorsum is yellowish tan to dark reddish
brown with dark brown markings consisting of an interocular bar,
broad canthal stripe usually bordered abOl-'e b}' a narrow cream
line, postorbital spot, three or four chevrons on the body. and
narrow diagonal bars on the limbs, A pair of small black spots
usually is present in tile scapular region. The anterior and
posterior surfaces of the thighs are dark brown with small pale
yellow spots. The venter is pale yellow, and the iris is reddish
copper.
Pristimantis (Pristimantis) danae (Duellman)Figure 150. Map 20
Duettman, 1978a:422. Hotol)'pe: KU 162307, adult male. from the Rio
Cosnipala, ~ irm Iby road] SW of~nl~ tsabeta, 13'OS'S, 7t' t8'W,
t700 m. Depanamenlo de Cusco, Peru. Pnulma111s dunue-Heinlcke.
Due!!man. and /ledges, 2007:S1 Table 2.f."lemheroliacly/us
danae
Characteristics: A member of the Pristimanlis (Prislimolllis)
perulJianus Group having (I) sk in 011 dorsum shagreen; that on
venter arcolate; discoidal fold prominent;
Genera and Species
-Comparisons: Pristimantis danae and P. reichlei are essentially
indistinguishable morphologically, but the two species have
distinct molecular and call differences. The spots on the posterior
surfaces of the thighs tend to be pale yellowin P dallae, whereas
the spots are orange in P. reichlei; . the latter lacks
dorsolateral folds, which are present, but weak, in P. danae. Of
the members of the Pristimalllis callspicillalUs andperuvianus
groups that have areolate skin on the venter and pale spots or
flecks on the posterior su rfaces of the thighs, P. avieuporum,
peruvianus, and skydmainos differ from P. danae by having Pinger I
longer than Finger 11; P. skydmainos also differs by having an
interocular fol d and a finlike middoral tubercle. Prislimalllis
meridionalis differs from P. danae by having tubercles on the upper
eyelid, and P. tOllyrhynehus differs by having a conspicuous
conical tubercle on the heel. The slightly smaller P bipunetalils
is like many specimens of P. danae in having a pair of black spots
in the scapula r region, but P. bipullctatus differs from P. danae
by having a snout that is truncate in profile, small tubercle on
the heel, Fingers I and II equal in length, and brown mottling on
the throat, in contrast to a rounded snout in profile, no tubercles
on the heel, Finger I shorter thall Finger II, and no brown mouling
on the throat. Distribution and ecology: Priseimantis d(1!we occu
rs at elevations of 1180-1710 III on the Amazonian slopes of the
Andes from central P to central Bolivia. This species eru
inhabits humid lower montane forest, where individuals arc
commonly found on low vegetation at night. Remarks: Fifteen egg
clutches contained 28-40 eggs hav ing diameters 01 2.0-3.2 mm;
clutches at the highest elevation (2040 m in the Cosfiipata Valley,
Departamento de ) Cusco, contained fewer, larger eggs than those
from lower elevations (1240 and 1480 m) in the same valley
(Martinez and Rodriguez, 2008). The specific name is a patronym for
Dana T. Duellman. who at the age of four collected many of the
frogs comprising the type series.Pristimantis (Pristimantis) de/ius
(Duell man and Mendelson) Figure 151, Map 20 EleUl/ierodactytus
delius Duellman and ~lendeJson. 1995:352 . Hototype: MHNURP 100,
adult female, from San Jacinto. 02"1844.8S, 75'51'46.0", 183 m.
Departamento de Loreto, P eru. Pristimanris de/ius- H einide,
Duetlman, and Hedges, 2007:$t Tabte 2.
Characteristics: This member of the Pristimalllis (Pristimantis)
unistrigalUs Group has: (I) skin on dorsum smooth; that on venter
areolate; discoidal fold present; dor solateral folds absent; (2)
tympanic membrane and tympanic annulus prominent, round, its length
about 40% length of eye; (3) snout acutely rounded in dorsal view,
rounded in profile;
Prlsr;mifnris fPristjmanrisl
(4) upper eyelid lacking tubercles, narrower than
100; cranial crests al)sent; (5) dentigerous processes of vomers
absent; (6) condition of vocal slits and nuptial pads unknown; (7)
~'inger I shorter than Finger II; discs subtruncate; (8) fingers
bearing lateral keels; (9) ulnar tubercles present; (10) heel and
tarsus bearing tubertles; inner tarsal fold thick; (II) inner
metatarsal tubercle elliptical. about 5" round outer metatarsal
tubercle; supernumerary plantar tubercles absent; (12) toes bearing
lateral keels; webbing absent; Toe V much longer than Toe III;
discs smaller than those on fingers; (13) dorsum tan with three
longitudinal brown stripes; venter cream; (14) SYL 30.9 mm in
single female. Coloration : In life, the dorsum is yellowish tan
with pale brown stripes on the body alld transverse bars of the
limhs. The loreal region is dark brown, Flg. 151. Pristim26 mm) and
differ in coloration.
- Distribution and ecology: This species is known Irom
elevations of 1700 m in the Cordi!!era de Cucutu, 1550 m of the
weslern slopes of the Cordillera del C6ndor, and 1138 III on Ihe
eastern slopes or the Cordillera del C6ndor. The lasl locality is
Ihe only record for this species in Peru; the in(\i vidual was on a
leaf 2 m above the ground at night in humid 10,,"er monlane forest.
- Rema rks: The specific name is a patronym for Robert M. Peck,
wile collected many amphibians in Ecuador for Ihe Academy of
Natural Sciences of Philadelphia.Pristimantis (Prisfimantis)
percnopferus
(Duellman and Pramuk) Figure 183, Map 29 Jtulhtrodauylus
ptfCtIOpll!fU$ DucllllLln Ind Pr~mut. 1999:58. Itololypt; KU
217318. adult female, from Sanl~ Rosa (Ie iii Yunga, 0605"5,7S43'W.
IlOG m. Departamenlo de Ca~marca.l'fm.
PTislimtlnlisPf'ftIIOpltfUS-Heinicke, Ouellman. and Itwgu, 2007: 51
Table 2. - Characteristics: This member of the Pristimunils
(Prislimorltis) unislfigatus Group has: (I) skin on dorsum smoolh
with a few small tubercles: tM! on venter weakly are olate:
discoidal fold prominenl: dorsolateral folds absent; (2) tympanic
membrane and Iympanic annullls distinct. round, its length 40"
lenglh of eye; (3) snout moderalely long, sub acuminate in (\orsal
view. rottnded itt prolile: (4) upper eye lid with or withoullow
tubercles, narrower than JOD; cranial creSIS absent; (5)
denligerous processes 01 vemers absent;
Genera and Species
(6) vocal slits present; nuptial pads absent; (7) Finger I
shorter than Finger II ; discs on outer fingers nearl), truncate,
morc titan twice width of digit proximal to disc; (8) fingers
lacking distinct lateral fringes; (9) ulnar tubercles absent; (10)
heel and tarsus lacking tubercles; (II) inner metatarsal tubercle
elevated, elliptical, about Sx subconical outer metatarsal
tubercle; supernumerar)' plantar tubercles tow rounded; (12) toes
lacking lateral fringes; webbing absent; Toe V much longer than Toe
!II ; discs nearly as large as those on outer fingers; (13) dorsum
pale brown with dark brown ma rks in scapular region; venter tan
with dark brown flecks; (14) SVL 21.5- 23.2 mm in three males, 25.9
mm in single fema le. Coloration: Duellman and Pramuk (lg99)
reported Rainer Schulte'S observations on coloration in life of the
holotype as: "Dorsum pale grayish brown with reddish tint on head
and small black spots; ven ter whitish gray." Color photographs of
other specimens reveals a reddish tan dorsum with faintly darker
brown markings consisting of a broad interorbital bar, labial bars,
a broad X-shaped mark or chevron in the scapular region, behind
which are irregular marks, some of which
extend onto the flanks. and transverse bars on the limbs. An
orange tint is evident in the groin and on the proximal surfaces of
the thighs. The postorbital stripe is black, and the iris is pale
bronze with black flecks and a median horiZOlltal red dish brown
streak. Com pa risons: Pns/imantis percnoplerus superficially
resembles P. incompfUs, which is similar in size and also lacks
vomerine teeth (except in large females); the latter differs from P
percrwpterus by having a snout that is rounded in dorsal view,
discs on the finger rounded, the digits bearing lateral fringes,
and Ihe inner edge nf the tarSliS bearing one or two tubercles.
Furthermore, in P. ineomplUs the dor sal markings are more
extensive and contrasting with the ground color, and the venter is
brown, instead of whitish gray. Fourteen other species 01
Prlstiman/is in Peru lack vomerine teeth; in three others (P.
incomp/us, rhodoplichus, and sehullei) teeth are present only in
large females, and in P. eoladac/ylus, mor/ioe and oliuoeeus
vomerine teeth are present but buried in the buccal mucosa. Six of
these 21 species (P. alfabracus, corrugolUS, melonogaster,
po/aikas, simansii, and slicloboubonus) are robust-bodied members
of the
f ig. 183. Prinimantis percnoptfuUS. (ORB tO t 00494, mate,
Gokta, Amazonas, Pe ru. P. J. Ven e gas.
Prislimantis (Pristimantis)
Pristimalltis orestes Group that have small terminal discs on
the fingers, short hind limbs, and Toe V only slightly longer than
Toe III. Three of the remaining species (P. colodaetrills. imiwtn\
and martiae) lack a differentiated tympanum and tympanic annulus.
Six of the remaining species (P. il/camplus, laerimosas. mendax,
oliaaeells. perenoplerllS, and ScilUllls) have a dark interorbital
bar, which is absent in P: anemeras. All other species lacking
dentigerous processes of vomers or not hal'ing teeth (P. de/ills,
rhodoplichlls, sagftWIllS, and sehilltei) have one or more
tubercles on the heel (none in P: perenopterlls) . Distribution and
ecology; This species has a disjunct distribution in tbe Cordillera
del C6ndor at elel'ations of 1100-1300 m and in the northern part
of the Cordillera Central (1830-2400 01) in northern P eru. One
individual was found in an arboreal bromeliad by day in semiarid
forest, wbereas tbose specimens from the eastern slopes of the
Cordillera del C6ndor were on low vegetation at night in humid
montane forest. Remarks: According to Duellman and Pramuk (1999),
the specific name is the Greek noun, per/mop/eros, mean ing
vultllre; they applied the name in reference to Yullur Bf}'phllS,
the Andean condor, to which the Cordillera del Condor
refers.Pristimantis (Prist;mantis) peruvianus (Melin)Figure 184,
Map 29
Hytodesperuvimws Melin, t941:43. Uolotype: NlIMG 490.aduit
femaIe, [rom Roque. Departamento d~ San Martin.
Peru.E/eu/lierodact)"/us peruviUl/us-Gornam, 1966:91.
ffeUlhe,odacl.ltus cOflspicilfatus (pari) - L)'flCh. 1915a:J.
PriSlim(lfItis pem~'iaflus- Htinicke , Dueliman. and Uedges,
2007:Fig. 184. Pr'stimantis peruviiml.1~-A. KU 222027, male, 24.7
mm SVL. B. KU 2n024, male, 25.B mm SVl. both from 1.5km N Ten iente
l6pez, Loreto,
Peru. WED.
SI Table 2
-Characteristics; Tbis member 01 tbe Pristimamis (Pristimanlls)
peravianus Group has (I) skin 011 dorsum shagreen with scattered
tubercles, that on venter smoolh: discoidal fold prominent;
dorsolateral folds present; (2) tympanic membrane smooth; tympanic
annulus prominent, nearly round. its length about 50%length of eye;
(3) snout round ed to subacuminate in dorsal view, rounded in
profile; (4) upper eyelid lacking tubercles, broader than 100;
cranial crests absent; (5) dentigerous processes of vomers promi
nent, triangular; (6) males possessing vocal slits and white,
nOllspinous nuptial pads; (7) Finger I longer than Finger II; discs
expanded, about twice width of !1igit proximal to pad; (8) fingers
bearing lateral fringes; (9) ulnar tubercles absent; (l0) heel and
tarsus lacking tubercles except for small tubercle on inner edge of
tarsus; (\\) inner metatarsal tubercle oval. about6x subconical
outer metatarsal tubercle; supernumerary plantar tubercles few in
number; (12) toes
bearing lateral fringes; webbing absent; Toe V slightly longer
than Toe Ill; discs equal in size to those on outer fingers; (13)
dorsum usuall)' brown with darker brown chevrons; ven ter cream
with brown flecks. most dense on throat; posterior surfaces of
thighs brown with cream spots; (14) SVL 1 5.124.9 mill in males,
25.0-33.3 mm in females. Coloration: The dorsum is reddish brown,
olivetan, or less frequentl)' grayish tan, usually with darker
brOll'lI markings consisting of a narrow interorbital bar,
irregular middorsal marks or chevron'shaped marks on the back. and
narrow diagonal marks on the limbs. Black canthal and postorbital
stripes are incorporated into a black or dark brown facemask,
bordered below by a pale tan to creamy yellow or white labial
stripe that may be continuous onto the upper arm: labial bars are
diffuse or absent. Many individuals have a narrow cream or white
line extending from the tip of lhe snout along the outer edge of
the upper eyelid. The flanks are colored like the dorsum and
usually have diagonal brown marks tbat are continuous with the
chevrons on the back, except for a pale
Genera and Species
dorsolateral stripe. Tile posterior surlaces of tile thiglls are
dark brown or black with yellow-orange spots. Tile venter is white
with dark brown to black fleck s. small spots. or st reaks on the
throat. The ventral surfaces of the shanks are grayish tan with
faim yellow spots. The iris is pale bronze, usually with a median
horizontal reddish brown streak. Comparisons: Four PerUl'ian
species in the Prislimantis conspici/{arus and P. peruvianllSgroups
hal'e smooth skin on the belly and smal l pale spots or flecks on
the posterior slIrfaces of the thighs. Prislimantis buccinutor and
P. metabales have basalwcbbing between Toes 11 and V, and the
former differs from all Of the others by ha\~ng an orange-pink
groin and hidden surfaces of tile shanks. The remaining species. P
cOflspicillalUs and f peruviGlllls, are alike in lacking webbing
between the toes; they differ from one another in sublle
differences in coloration and in size~P pelvuial/Us is smaller
llian P. conspicil/utus. In P. pefVlJ ianuS a pale labial stripe is
present, and the venter is wliite, but dense brown flecks are
present on the throat and brown spots may be present on tlie bel
ly. especially in males; the spots on the posterior surfaces of the
thighs are compa ratively large and yellow to orange in color, and
the ventral surfaces of the sh~nks afC tan with pale yellow spots.
In contrast, in P. cOllspicilfarus the upper lip is gray or tan
wilh or without darker labial bars, and the throat and bell)' are
immaculate whitc, except fOT brown flecks on the throat in large
females; thl' spots on the posterior surfaces of the thighs are
small and orange or red, and tlleventral surfaces of the shanks
lack spots. The unrelated P lanthalliles is likef peruviGl/Us in
having a smooth venter bul dif fers iJ}' having conical tubercles
on the heels and a dark gray throat with a mediijn longitudinal
whlle stripe. Distribution and ecology: This species is "idely
distrihuted in the upper Amazon Basin in western Brazil, southern
Colombia. Ecuador. and Peru up to elevations of 2050 m on the
Amazoniall slopes of the Andes. It inhabits 10....'land Iropical
rainforest and humid montane fores!. By day indil'iduals are active
on the forest floor, and ~ome seek refu ge in bromeliads; at night
some individuals are on the forest floor, but most have been found
on low vegeta ti on. The advertisement ca ll consists of a series
of quickly repeated whistle like notes COucHman, 2!l!lS). Remarks:
The recognition of Pristimalllis per/JIJiarws as a species
distinctlrom f cOllspicillalus has fluctuated. Lynch ( 1975a.)
considered P.. peruvial/us to llc a synonym of P eon spicillatvs,
but Duellman and Tolt ( 1979). Lynch (1980), and Lynch and Ouellman
( 1980) recognized both species with some reservations. The
phylogenetic analyses of molecular data by Hedges el at. (2008a)
showed that P c(}lIspicillalll5 and P penJl}ialius are in different
clades, thereby substantiating the specific rc(:ognition of each.
Additional molecu ~ lar work by Padial and De la Riva (2008)
revealed tbe exis-
lenee of a cryptic species, P. reichlel. in southern Peru and
Bolivia. Possibly other cryptic species are now induded in P..
peruuitlllUs. For example, there is ~ noticeable differ ence in
size between individuals from elevations 01 14 1 01i40 m on the
Amazonian slopes of the Cordillera Oriental in ECllador and those
from the Amazonian towlands. especially fmm Departamento Madre de
Dios, Peru. Ten males from the Ecuadorian Andes have SVL
of29.2-35.8 (1< '" 31.8) mm s and 10 fema les have SVLsof
38.6~46.4 (:5< = 12.3) mm (Lynch and Duellman, 1980), in
contrast to 15.7- 24.9 ($! '" 21.7) in 25 males and 25.0-33.3 (x '=
29.7) in 12 females lrom Cusco Amazollico, Peru (Duell man, 2005) .
The SVLs of the former are more like tliose of P conspfcillatus
(25.0-30.0 mm in 22 males and 34.8-48.8 mm in t2 female s (Lym:h, 1
975a) than those of P pemuianus from the. Amazonian lowlands. The
ranges of the two species broadly overlap in northeast ern Peru,
but the)' have not been found in sympatty. although both species
hal'f' been col lected in the v1cinity of lingo Marfa. Departamento
de Hwnuco. Prislimantis conspiciflalus apparenlly is confined to
the lowlands, whereas P. pemvianus ascends the Amazonian slopes of
the Andes (0 elevations of 19tO m in the Cordillera Oriental in
Peru (Lynch and DueUman, 1980). 1700-1975 m in the Cordillera de
Cututu (Dueilltlan and Lynch, 1988) , 1500- 1830 Itl un the weslern
slopes of the Cordillera del C6ndor (Almendariz, 1997; LYflCh and
Duellman (1980), 665-17SU III on tile eastern slopes of the
Cordillera del Condor (Reynolds and lcochea, 1997), ~90- 1280 m in
the Serrania de Sira (Duellman and Toft , 1979). aod 950-1080 mon
the eastern slopes of the Cordillera Central in northern Peru
(Duellman and Prallluk. 1999). lIerein ....e report specimens from
modmtely high elevations farther south-124D-2050 m ill the
depattanlentos de Ayacucho. Junin, and Pasco. and 8001720 m in
Oep.1rtamento de Cusco. AI Cusco Amaz6nico. Departdmento de Madre
de Dios, four gravid females having SVLs of 25- 33 (x '" 29.7)
mmcon tained 26-32 (r "" 28.3) ovarian eggs with a diameter of 1.8
mm (Duell man, 2005). Although Melin (l 941) did not state the
derivation of the specific name, it obl'iousl), relers to the
country 01 origin, Peru.
Prisfimantis (Pristimantis) petrobardus {Duell man)
Frontispiece. Figure \85. Map 29 Eleuthe/'fJdaclJIus pt/robardus
Duellm~n, t99Ia:S. ttolotype: K 212292, adult male. irol"
a~proximatelJ' 2 kill lb)' roadl W U Huambos. 2500 m. Depar1amen10
d~ Cajalliarca. Peru. Pfls/imantis #Itommfus- Hcinicke. Duellman.
and H edges. 2007: SI Tabte 2. EleutJlerodactyll.lS
t~rtl/iClllaIlJS Duellman and Lehr. 2007:7. H olotype: FMNH
2323.}0. an adult male. from the Rio 7.ai'>a, ca. ont' S. 79'06
W t800 m. DeparLlmenlO de Cajaltlllrca, Peru. NewSynoll)'lll)'.
.
Pristimantis (f>ristimIJntis j
Characteri stics: This member of the Prislimantis (hislimanlis)
unislrigalus Group is characterized by: ( I) skin on dorsum smooth
with small irregularl), arranged pustules; that on I'enter
areolate; discoidal fold ~ident; dorsolateral folds absent; (2)
tympanic membrane and tympanic annulus distinct. round, its lensth
abo\lt 45% length of eye; (3) snout rounded in dorsal View and in
profile; (4) upper eyelid lacking tubercles, narrower thall 10D;
cranial crests absent: (5) dentigerous processes of vomers oblique,
prominent; (6) I'ocal slits present: nuptial pads present on dorsal
surface of thumb; (7) Finger I shorter than Finger II; discs broad;
(8) fingers bearing lateral fringes: (9) ulnar tuber c1es low,
diffuse; (1 heel bearing small tubercles: outer 0) edge of tarsus
lacking tubercles: inner edge of tarslIs bear ing elongate
tubercle: (II) inner melatarsallubercle ovoid, 3-4x conical outer
metatarsal tuberde; !upernumeraryplan tar tubercles small: (12)
toes bearing lateral fringes: web bing absent; Toe V much longer
than Toe Ill; discs as large as those on outer fingers; (13) dorsum
tan with irregu lar brown markings; venter cream, with or without
gra)' reticula tions; (1 SVL 27.0-39.5 !11m in males, 38.iJ-43.0 mm
in females. 4) Coloration : In life, the dorsum is )'tllowish tan
wilh oiivetan to brown markings on the dorsum 01 the head, bod)"
flanks, and limbs. The markings usually consist of a middorsal spot
midway between the level of the orbits and nostrils. an
interorbital bar, short canthal stripe, labial bars. a curved
postorbital bar, tll'O to four diagonal marks on the flank, and
slightly diagonal bars on the limbs. The marki~gs on the dar sum of
the body variously consist of an irregular line beginning at the
posterior edge of the upper eyelid curving medially at the
midJength of the body and thence posteriori), to a point above the
groin, an irregular chevronshaped mark postsacrally, or
interconnected, irregularly shaped marks. The vcnter is pale cream
to creamy )'Cllow, usually with minute black flecks on the chin
anterior to the yellow vocal sac; many individuals in the southern
part of the range h.we gray vermiculations on the chest and bell)'.
The iris is pale bronze with fine gray reticulations and a diffuse
median horizontal brown or red stripe. Variants in dorsal colora
tion include a narrow cream or yellow canthal stripe con tinuing
along the outer edge 01 the eyelid and then broadenin g inlo a
conspiClious rlorsolateral stripe that extends the full length of
the body. One individual has only faint dorsal markings: another
has a da rk brown darsolal eral stripe Irom just posterior to the
orbit 10 the sacrum and bordered below by a pale J'Cllow region
Ihat is broadest poslcrior~' and extends to the groin. One adult
male Ilas a cream middorsal slripe extending from the snout to the
venl and bordered by narrower, irregular black stripes.
Comparisons: Prisl/mantis pelrobardus is most similar to P.
cajamarcensis. unislrigalus, alld versicolor in having a
Fig.l as. Pris t/manto petrobardus-A. KU 212292, mate, 's 28.7
mm SVL, 2 ~m W H uambo~. C18, female, 32.0 mm SVl, Ab" QuilUl,
Cajamarca, Peru. WED.
culate skin on the dorsum, which is brown with darker brown
markings; they also have dentigerous processes of vomers. The
sympatric P. pOIaiJ/Os is like P. simonsii in lacking denlig erous
processes of I'omers and a tubercle on the heel, but it differs by
having ulnar tubercles coalesced into a ridge and a smooth dorsum
that is brown and by lacking pale spots in the groin and on the
hidden surfaces of the thiShs. Pn'stimalllis chimll also has
coalesced ulnar tubercles, but it differs from P. simOllsii in
cotoration and the presence of cranial crests. The coloration of P.
/Jidua is like that 01 P. pa/oikos; P. uidua also differs from P.
simonsli" by having the skin on the dorsum shagreen with scattered
tubercles and a weak dorsa lateral'old; P. corrogatlls differs from
all other members of the Pristimamis orestes Group by having
prominent conical tubercles Of! the upper eyelids. The on!)' other
Pri$limantis in Peru that are primarily black are two members of
the Prislimantisilnis/rigotus Group--P.IYlhrodes, an Amazonian
Genera and Species
species that has expanded digital discs, Toe V much longer than
Toes III. and red (white in preservative) blolches on the venter;
the other is P. coroliallJS thai also has a long Toe Vand has
narrow digits and orange-red markings on the flanks and anterior
surfaces of the thighs. Distribution and ecology: Pristimomis
simollsii inhabits humid puna above tree line at
elevalionsol30SO-3760 min the northern part olthe Cordillera
Occidental in Peru. Remarks: The specific name is a patronym for P.
O. Simons who collected many amphibians and repliles in Peru lor
the British Museum.
Pristimantis (Pristimantis) skydmainos(Flores and Rodriguez)
figure 208, Map ~ Eleulhero(}aclytus ~ydlOOinos Flores and
Rodrigue" 1997:388. lIolotype: MCZ SSJ04, adult female, from Cacha
tashu, 11"51'5, 71' \9'W, 380 m, Manu NadoJlilI Park, Departamento
de Madre de Dios, Peru. Eleul/Jerodaclylu5 karcharias flo res and
Rodr[guez, 1997:392. lIo10l~pe: MCZ 89075, immature female, from
Atva, 1000 m , between Chachapoyas and Baglla G rande. Departamento
de Amazonas. Pelu (synonymy fide Padial and De La Kiva.2005).
I'nulmantis skydmainru- Heinicte, Duellman, and Hedges, 2007: Sl
Table 2.Flg .20e. Pristimant;~~kydmainO$,
Ch aracteristics: This memtx:r of the Pristimantis
(Pristimalilis) conspicil/atus Group has: (\) skin on dorsum
shagreen, that on venter weakly areolate; discoidal fold prominent;
interocular fotd and dorsolateral folds present, fragmented or not:
a fin-shaped middorsaltuhercle; (2) tympan ic mcmbrane smooth;
tympanic annulus prominent, vertical Iy ovoid, its length about
twothirds length of eye; (3) snout moderately long, acutely rounded
in dorsal view, rounded in profile; (4) upper eyelid lading
tubertles, much narrower than 100; cranial cresls absent; (5)
dentigerous processes of vomers prominent, triangular; (6) vocal
slits and non spinous nuptial pads present; (7) ~'inger I slightly
longer than Finger II; discs on outer fingers expanded, nearly
truncate, about twice width of digit proximal to pad; (8) lingers
bearing lateral fringes: (9) ulnar tubercles absent; (10) heel
bearing minute subconical tubercles: outer edge of tarsus lacking
tubercles; inner edge of tarsus bearing distinct laid distally;
(11) inner metalarsaltubercle elevated, elliptical, 2-6x round
outer metatarsal tubercle: supernumerary plantar tubercles few,
indistinct; (12) toes bearing lateral fringes; webbing basal; Toe V
slightly longer than Toe 111; discs slightly smaller than those on
ouler lingers; (13) dorsum gra)'ish brown with brown or black
markings; venter cream with brown flecks on throat: ( I~ )
SVlI9.0-2S.5 mm in males, 25.6-32.4 mm in females.
AmnOrlas lodge, Madre de Dios. Pert!. W. leonard.
Pri5!imanli5 (Pri5timiOnris)
- Coloration: In life, the. dorsum and flanks are. orange brown
with pinkish tan dorsolateral folds. Faim brown marks and small
cream spots arc present on the dorsum and lIanks. Anarrow, dark
brown interorbital bar and diffuse pnslorbil. al stripe are
present. Labial bars and a canthal stripe are absent; instead the
side of the sead is covered in 11 dark brown facemask. Flores and
Rodriguez (1997) described the color paltern in preserved
specimens. The dorsum is gray L~h brown wilh dark brown to black
markings consisting of an interorbital bar, postorbital stripe,
spot at anterior cor ner of the eye, bare on forearm, spot or
stripe on the knee, two spots on the posteromedial surface of e~ch
shank, anal triangle, and bars on the hind limbs. Also Ihere is a
faint W shaped mark in the scapular region and 1-3 chevrons on the
back. The posterior surfaces of the thighs ate brown with minute
cream flecks. The Inner surfaces of Ihe tarsi and plao tar surfaces
are dark brown, whereas the venter is cream wilh fine dark broll'll
flecks en the threat. chest, and edges ef the limbs. - Compari
sens: Among the Peruvian species in the Pristimantis conspicillotus
Group charatlerized by having Toe V only slightly longer thatl Toe
III, P. ouieaporum, and P. sk),dmairlOs differ from all others by
having an interocu lar fold and weakly areolate ~kin on the venter
in contrast to smooth skin venlrally in Ihe other species; P.
meridiorlO/is also has areoiate skin on the venter, but it lacks an
interoc IIlar fold, whereas P. skydmainos differs from P.
auieupofllm by having a linlike middorsalluhercle. Pristimantis
cullei ro.llris has a wedgeshaped snout, broad pale labial stripe,
and many small dark spots on the dorsum in contrast 10 the rounded
snout, interorbila l bar. and single middorsal dark spol in p,
avicuporum; P. buccinator differs from P. skyd mainos by having
pillk spots In the grain and on the hidden surfaces of the thighs.
Remaining Peruvian members of Ihe Pris/imantis Clm spicilialUs and
pel"UlJiallllS groups include three large spe des (P. citriogaster.
cOlldor, alld {),malll; males to 46 mm, females to 73 mm) and many
smaller species (males to 38 mm, fema les to 51 mill). Of the
smaller species in the upper Amazon Basin and on the lower slopes
of the Andes - P. bipuncl{JW$, eOllspicf{/aws, danae, rna/kim;
met{Jhales. perIJ vianus, reichlei, skydmaillos, and
tall)'fh)'nehus-the pos terior surfaces 01 the thighs ijre dark
brown with cream or red flecks, and three of these (P. bipune/atus.
ma/kim; and metabOles) have basal webbing between the toes. TIle
poste rior surfaces of the thighs are uniform brown in p,
adisto/us, alberlUs, {enestratus, phorongabates. rhabdolaemu.l,
Sliclo gasrer, and /}/iarsi; the thighs have a longitudioal fed
stripe in P. sagirtulus. The utlrelated, P. /olllhaniles, has a
smooth ven ter, a median pale streak on a dark Ihroal, and a
prominent tubercle on the heel.
- Di stribution and ecology: Pristimnlltis sk),dmoill'
as occurs in the Amazonian lowlands in central and southern Peru
and extreme western Brazil, and on the lower Arnazoni~n slopes (up
to 750 m) of the C ordillera Oriental in Peru and Ecuador. At the
type localily males calted from low vegetation; the c~1l consists
of a sinsle chirp (Flores ~nd Rodriguez, 1 997). -Remarks : Gravid
females contained 11-25 ovarian eggs with a maximum diameter of 3.7
mm (Flores and R odrfguez,1997), who noted that the specifiC name
is Greek meaning angry and refers 10 Ihe furrowed brow expression
imparted by the interocular fold.
Pristimantis (Pristimantis) sternothylax (DueHman and
Wild)FiBure 209, Map 35 lilelliherodaclyills Slefllolhylnx Duellman
andWilcl, 1993: 17,Hololypt: KU 219793, adult mate, from 1 kw I by
road) ENECanchaque, 1 6 840 m, Departamento de Piura, Peru,
Prisrimun/is sterrwlhylax- Heinicxe , Dueliman. and Hedges. 2007:5t
Table2,
- Characteristics : This member of the Pristimantis
(PriSlim()nlis) unislrigalUs Group has: (I) skin on
dorsumshagreen, bearing few, low. round tubercles posteriorly and
laieraJly; thal on venIer areolate: discoidal fold evident; dor
solateral folds absent; (2) t}'mpanic membrane and tympanic annulus
distinct, round, its length abnltt one half length of eye; (3)
snout subacuminate in dorsal view, aCUlely rounded in pro file;
canthus rostralis acutely rounded; (4) upper eyelid bear ing low.
round tubercles, narrower than 100; cranial crests absent; (5)
dentigerolls processes of vemers prominent, oval; (6) l'OCal slits
presenl; nuptial pads present on dorsal surface of thumb; (7)
Finger I shorter than Fillger II; discs broad. trUIi' cate; (8)
I-'ilt!ler~ III and IV bearing narrow laleral frinses; (9) ulnar
tubercles absenl; (10) heel and tarsus lacking tubercles and folds;
inner edge of tarsus !laving a fold; (11) inner mela tarsal
elliptical, 3- 4x round outer metatarsal tubercle; plan tar
supernumerary tubercles diffuse; (12) toes be~ ring lal eral
fringes: webbing absellt; Toe V much longer Ihan Toe III; discs
smaller thall those on fingers; (13) dorsllm Ian with dark brown
markings; venter cream with minule brown necks; (14) SVL in males
18.3-29.1 mm, in females 28.3-36.7 mm. - Coloration: In life, the
dorsum [s brown to pale green ish tan with contrasting dark brown
marks conSisting of labi al bars, canthal and postorbital stripes,
Interorbital bar or trian.~ltlar mark with apex connected or not to
the pattern 00 the dorsum of the body-X, H or Wshaped marks in the
scapular regien, and diagonal or chevronshaped marks pes ~ terior
to Ihe scapular region. The limbs are distinctly barred lI'ilh dark
brown; the bars on the thighs extend onto the pos
Ge ne ra and Specie
Pristimantis (Pristimantis) stictoboubonus (Duellman, lehr and
Venegas)Figure 210, Map 35fJI!Ulherodac/yllJs $lic/obOlJ.bIJlIIIS
Duettman, Lehr ~nd Venega~,
200S:58. 1l0Iot}'pe: MUSM 24446. adult female, from Quintecocha,
3130 m, Deparlamcllto lie S~n Martin. Peril. PrislflllGiIlis
~rictoboubonlJ.f-Hcinicke, Ducllman, and Hedges. 2007:51 Table 2. -
Characteristics: This member of the Pristimantis (Pristimolllis)
orestes Group has: (I) skin on dorsum sh~greenFig.209. Pristimiln
lis sternothylil!l, KU 218793, ma le, 29.1mm SVL, 16 km ENE
Can(haque, Pi l.l ra, Peru. Ad apted from
Duel lman and Wild (1993).
leriM surfaces where they are separated by yellow to orange
interspaces. The I'enter is dull white, and the vocal sac is
yellow. The iris is a copper color. - Comparisons: The presence of
a subllcuminate snout in dorsal view and the absence of tubercles
on the heels distin guishes Pristimantis sternathy/ax from mosl
Peru~iall members of the genus. Pristimantis acuminalUS and P.
ran/anti dir[er b), having a uniformly green dorsum and the absence
of a tympanic membrane, whereas P. allemems differs by havIng a
rostf~ 1 papilla. Prislimantis tmdlyblepharis also differs b~
lacking a tympanic membrane, and P. I}ariabilis differs lly having
a large yellow spot bordered by black in the groin; P. wnisrum
differs by having a smooth venter ~nd the fifth Toe only slightly
longer than the third toe. in northern Peru P. S(erflOllly/ux and
P. cajullwcensis Me similar morphologically and in coloration.
However, P. stefl/othy/ax difJers from P. cajamarcensis in having a
larger tympanum (50%length of eye VS. 30%length of eye),
suhacuminate snout in dorsall'iew' (rounded in P. cajamarcellsis),
and a large vocal sac extending from near midlength of the throat
onto the chesl (small vocal sac restricted to throat in P.
cajamarcensis). Distribution and ecology: This species is known
from elevations of 1735-1 m on the western slopes of the 840
Cordillera de Huancabamtla and from elevations of 15002100 m on the
western slopes of the Cordillera Occidental in northern Peru. All
individuals have been found on low vegetation al night in hillnid
monlalle forest. The advertisemenL call is a single "tock~ repeated
at intervals of about 5 sec (DueJiman and Wild, 1993) . Remarks:
The vocal sac in males is large, single, and median; it begins near
midlength of the throat and extends well onto the chest. D ueJiman
and Wild (1993) noted that the specific Mme is derived from the
Gree~ nouns slllrrlQlI meaning chest and thY/l}x meaning sac; the
namewas used in reference to the position 01 the vocal sac
I!Xtending onto the clles!.
lacking tubercles and ridges, that on venter areolate; discoidal
fold nol evident: dorsolateral folds absent; (2) tympanic memo
br~ne smooth; tympanic annulus distinct, round, its length sli~hlly
more than half length of eye; (3) snoot short, rounded in dorsal
view, curved anterOllentrslly in profile; lips rounded; (4) upper
eyelid laCking tubercles, 4-1.\% laD; cranial crests' absent; (5)
denLigerous processes orvomcrs absent; (6) males lacking vocal
slits and nuptial pads; (1) finger I shoner than finger II; discs
on outer fingers narrow: (8) fingers bearins broad lateral fringes;
(9) ulnar tuberdes absent; (I0) heel and outer edge of tarsus
lacking tubercles; inner ta rsal fold tlarely evident distally;
(11) iuner metatarsal tubercle broadly elevated, subtrianguJar,
about lOx suocollical outer metatarsal tubercle; supernumerary
plantar tubercles present; (l2) toes bearing broad lateral
Irillges:Toe V slightly longer than Toe III; discs about same size
as those on lingers; (13) dorsum reddish tan with or lvilhoul
small, irregular brown markings; venter cream with brown flecks;
groin white with dark brown markings; ( 4) SVL 20.0-21.5 mm in two
males, 26.0 mm In one female. - Coloration : A culor photograph of
MUSM 24447 reveals that the dorsum is reddish tan, becoming pale
gray on the lIanks. The cantha l and postocular stripes are diffuse
brown, and the upper lip is creamy gra)'. The iris is dull bronze
wi th brown reticula tions and a broad, median , horizontal reddish
brown stripe. The groin is white in preservative, but it likely is
pink or orange in life. There is a diffuse pale tan labial stripe
in preserved specimens. - Compari sons: Among the species placed in
thePristimulitis orestes Group, P. slicloiJoubollus is onique in
having a large white spot in the groin with dark brown markings
therein and the anterior, posterior, and venlral surfaces 01 the
thighs d~r k brown with distinct white spots (nol small flecks).
Four species in the group have unirormly colored groins- orange in
P. corrugatus, gray in P. patrJikos, a yellow suffusion or not in
P. slino/lsii. and undifferentiated from the fl ank color in P.
vidua. Yellow, pink, or while spots are present on the posterior
surfaces of the thighs in P. melallogoster. oreSles, and
sill/ollbolioon; respectively. but ill these species the groin is
black with white or)"ellow spots. InP. cordoua!!, coalesced
tubercles form ridges dorsoiaterally on the bod~; the groin and
anterior sur-
PristimanfiS (Pri5fimanfis)
Fig ,210. iii 20.0 mm SVL. L 5000 m
""">-""'"
m
J()(J().4000 m
2000-3000 m 1000-2000 mQ- l000m200km
Map 8. Loca l it ie~ of known occurrence of four ~pecie~ of
Pilrynopu$ in Peru .
Appendix UI
ECUADOR
COLOMBIA
BRAZIL
P hors/paulip, kauneorumP. kotosh
P. n/co/eae
D D D
"-I
> 5000 m
' 000-5000 m3000-4000 m 2000-3000 m 10 5000 m4()()()-5O{)O
m
3000-4000 m 2000-3000 m 1000-2000 m 0-1000 m 200 km
M 5000 m
4000-5000 m
3000-4000 m2000-3000m
1000-2000 m0-1000 m 200 km
Map 17. Loca lit ies of known ocwrrence of f our spec ies of
PriHimamis in Peru.
Cistribvtiort Maps
ECUADOR
COLOMBIA
BRAZIL
P. carvalhoi
P. ceuthospilusP. chimu
P. citriogastar
o
D D D
~
> 5000 m .ooo-.ooom 300()-4000 m
20Q0-3000 m
1000-2000 m 0-1000 m200km
Map 18. Loulities of known occurrence of four s,recies of
PrluimimtiJ in Peru.
Appendillill
ECUADOR
COLOMBIA
!
:. P.
colodectylus
P. condor P. consplcillalusP.
.-
cordov8e
P. coronatuscosnipataeBRAZIL
P. corrugatus
D D D
'-1
>
5000 m
")0'),""""" m 3000- 5000 m' {)()()-)OO
m
3000-4000 m
2000-3000 m1000-2000 m (HOOOm
200,,"
M"p 20. Localities of known occurrence of six species of
P,istim"nris in Peru.
Appendix III
ECUADOR
,-,
COLOMBIA
/,\,
.,.,
,
""BRAZIL
...."r' r r
;.-
P. diadematus P. eurydactylus
P. exoris/us) ' ~
'. ,, _C'. ,, ~ < ,.',-e
~
5000 mm
,Q()()..O(l()()
3000-4000 m 2000---3000 m 1(01)..-2000 m
o-looo m
200km
M"p 22. localities of known occurrence of three species of
Pr;S/;manf;J in Peru.
Appendix III
ECUADOR
COLOMBIA
)
,
..BRAZIL
, -
P. imitatrix P. incomptus P. infragutt8/US P. /acrimosus
':...:. ,
o
o
o
0 0 0
~
> 5000m
'!lOO-6OOO m3000-- 5000 m
ooo--.ooom3()()0-4000 m 2001)-3000 m 1000-2000m 0-1000 m 200
km
Map 25. Localities of known oc(urren(e of four species of Pr;Hi
mIJntis in Peru.
tfl
Distribution
M"p~
ECUADOR
COLOMBIA
"
-~
!
"
~- -I
"-"
rc ,/
,
BRAZIL
P. meridionalis
P metabates P minululusp muscosus
P nephoph;lus
D D 0
~
"5000 m 4000-5000 m 3000-4000 m 2000-3000 m 1000-2000 m 0-1000 m
200 km
Map 26. Localitie s of known occurrence 01 five species 01
Pri$timanti5 in Peru.
-fie
Appendix III
ECUADOR
COLOMBIA
o
0 0 0
I:1l
"5000 m4000-5000 m300Cl-4000 m2()(>(hroOO m
1000-2000 m
0-1000 m
200 km
Milp 27. Localities of known occurrence of Prisrim,mtis
o(kendeni in Peru.
Di~tribution
Maps
ECUADOR
COLOMBIA
BRAZil
P. olivaceusP. omatus P. pardalinus
P. pa/aikos
P. peck!
o
D D D
~
> 5000 m
41XlO-5000 m30Q0-4000 m
20QO-3OOO m 1000-2000 m
0-1000 m
200 km
Map 28. Localities 01 known occurrence of fi ve species of
Priltimantis in Peru .
Appendi~
III
ECUADOR
COLOMBIA
~
,,. ,~~
'~
... .-.. BRAZIL
~
-,
" -= .'.: ~ ,,'' :
~
"t'
r -r
.
, >
P. percnop /erus P. peruvianus P
