The Herpetofauna of the Peruvian dry forest along the Andean valley of the Marañón River and its

264 Pages·2014·5.3 MB·English

by  Claudia Koch

The Herpetofauna of the Peruvian dry forest along
the Andean valley of the Marañón River and its
tributaries, with a focus on endemic iguanians,
geckos and tegus

Claudia Koch
The Herpetofauna of the Peruvian dry forest along
the Andean valley of the Marañón River and its
tributaries, with a focus on endemic iguanians,
geckos and tegus

Squamata: Iguanidae, Phyllodactylidae, Teiidae

Dissertation
zur
Erlangung des Doktorgrades (Dr. rer. nat.)
der
Mathematisch-Naturwissenschaftlichen Fakultät
der
Rheinischen Friedrich-Wilhelms-Universität Bonn
vorgelegt von
Claudia Koch
aus
Bonn
Bonn, 2013
Angefertigt mit Genehmigung der Mathematisch-
Naturwissenschaftlichen Fakultät der Rheinischen Friedrich-
Wilhelms-Universität Bonn

Erstgutachter: Prof. Dr. Wolfgang Böhme
Zweitgutachter: Prof. Dr. Wolfgang Wägele

Tag der Promotion: 13.02.2014
Erscheinungsjahr: 2014
“Even so, the loss of a few species may seem almost irrelevant compared to major
environmental problems […] There is one last reason for caring […] And it is simply
this: the world would be a poorer, darker, lonelier place without them.”
- MARK CARWARDINE -
(Source: Adams, D. & M. Carwardine (1990): Last Chance to See. Ballentine Books, 265 pp.)
CONTENT
CONTENT

CONTENT ............................................................................................................................. 1
ACKNOWLEDGEMENTS ...................................................................................................... 5
1. GENERAL INTRODUCTION .......................................................................................... 7
1.1 Objectives and Background ......................................................................................... 7
1.2 Investigation Area .......................................................................................................13
1.2.1 Physiography ........................................................................................................13
1.2.2 Climate .................................................................................................................14
1.2.3 Vegetation ............................................................................................................14
1.2.4 Fauna ...................................................................................................................16
2. BIODIVERSITY OF THE PERUVIAN DRY FOREST HERPETOFAUNA .......................17
2.1 Annotated checklist and key to the species of amphibians and reptiles inhabiting the
northern Peruvian dry forest along the andean valley of the Marañón River and its
tributaries ..........................................................................................................................18
Introduction ...................................................................................................................19
Investigation areas ........................................................................................................20
Materials, methods and fieldwork ..................................................................................28
Results ..........................................................................................................................30
Rhinella limensis (Werner, 1901) ...................................................................................39
Rhinella margaritifera (Laurenti, 1768) .........................................................................40
Rhinella marina (Linnaeus, 1758) ..................................................................................42
Rhinella poeppigii (Tschudi, 1845) ................................................................................44
Rulyrana mcdiarmidi (Cisneros-Heredia, Venegas, Rada, & Schulte, 2008) .................47
Nymphargus posadae (Ruiz-Carranza & Lynch, 1995) ..................................................48
Excidobates mysteriosus (Myers, 1982) ........................................................................49
Hyloxalus elachyhistus (Edwards, 1971) ......................................................................50
Hyloxalus insulatus (Duellman, 2004) ............................................................................51
Pristimantis lymani (Barbour & Noble, 1920) .................................................................54
Pristimantis incomptus (Lynch & Duellman, 1980) .........................................................55
Pristimantis percnopterus (Duellman & Pramuk, 1999) ..................................................56
Leptodactylus labrosus Jiménez de la Espada, 1875 ...................................................57
Leptodactylus sp. ..........................................................................................................58
Phyllodactylus delsolari Venegas, Townsend, Koch & Böhme, 2008 .............................60
Phyllodactylus interandinus Dixon & Huey, 1970 ...........................................................61
1
CONTENT
Phyllodactylus johnwrighti Dixon & Huey, 1970 .............................................................62
Phyllodactylus reissii Peters, 1862 ................................................................................63
Phyllodactylus thompsoni Venegas, Townsend, Koch & Böhme, 2008 ..........................65
Phyllopezus maranjonensis Koch, Venegas & Böhme, 2006 .........................................65
Gonatodes atricucullaris Noble, 1921 ............................................................................66
Pseudogonatodes barbouri (Noble, 1921) .....................................................................67
Polychrus jacquelinae Koch, Venegas, Garcia-Bravo, Böhme, 2011 .............................69
Polychrus peruvianus (Noble, 1924) ..............................................................................69
Microlophus stolzmanni (Steindachner, 1891) ...............................................................70
Stenocercus huancabambae Cadle, 1991 .....................................................................71
Varzea altamazonica (Miralles, Barrio-Amoros, Rivas & Chaparro-Auza, 2006) ............72
Ameiva aggerecusans Koch, Venegas, Rödder, Flecks & Böhme, in press ...................73
Ameiva concolor Ruthven, 1924 ....................................................................................74
Ameiva nodam Koch, Venegas, Rödder, Flecks & Böhme, in press ..............................74
Callopistes flavipunctatus (Duméril & Bibron, 1839) .....................................................75
Amphisbaena pericensis Noble, 1921 ...........................................................................78
Boa constrictor ortonii Cope, 1878 ................................................................................79
Micrurus peruvianus Schmidt, 1936...............................................................................80
Bothrops sp. .................................................................................................................80
Colubridae sp. ...............................................................................................................82
Chironius exoletus (Linnaeus, 1758) .............................................................................83
Clelia clelia (Daudin, 1803) ............................................................................................84
Leptodeira septentrionalis larcorum Schmidt & Walker, 1943 ........................................85
Mastigodryas reticulatus (Peters, 1863) ........................................................................87
Mastigodryas heathii (Cope, 1876) ................................................................................88
Mastigodryas boddaerti (Sentzen, 1796) .......................................................................89
Oxybelis aeneus (Wagler, 1824) ...................................................................................90
Sibynomorphus vagrans (Dunn, 1923) ..........................................................................92
Tantilla melanocephala (Linneaus, 1758) ......................................................................93
Epictia cf. rufidorsa (Taylor, 1940) .................................................................................96
Epictia sp. 1 ..................................................................................................................97
Epictia sp. 2 ..................................................................................................................98
Epictia sp. 3 ...................................................................................................................99
Epictia sp. 4 ............................................................................................................... 100
Epictia sp. 5 ................................................................................................................. 101
Discussion ................................................................................................................... 103
2
CONTENT
3. CONTRIBUTIONS TO THE KNOWLEDGE OF NEOTROPICAL PHYLLODACTYLID GECKOS .............. 106
3.1 Two New Sympatric Species of Leaf-Toed Geckos (Gekkonidae: Phyllodactylus) from
the Balsas Region of the Upper Marañón Valley, Peru .......................................................... 107
Materials and Methods ............................................................................................................. 109
Phyllodactylus thompsoni sp. nov. ......................................................................................... 110
Phyllodactylus delsolari sp. nov. ............................................................................................ 116
Discussion .................................................................................................................................. 122
3.2 Squamata, Phyllodactylidae, Phyllodactylus thompsoni Venegas, Townsend, Koch and
Böhme, 2008 and Phyllodactylus delsolari Venegas, Townsend, Koch and Böhme, 2008:
Latitudinal and altitudinal distribution extension and geographic distribution map ............. 126
3.3 Ecology of a gecko assemblage (Phyllodactylidae: Squamata) from northern Peru .. 135
Introduction ................................................................................................................................ 136
Materials and Methods ............................................................................................................. 137
Results ........................................................................................................................................ 139
Discussion .................................................................................................................................. 145
4. CONTRIBUTIONS TO THE GENUS POLYCHRUS ............................................................................ 150
4.1 A new bush anole (Iguanidae, Polychrotinae, Polychrus) from the upper Marañon
basin, Peru, with a redescription of Polychrus peruvianus (Noble, 1924) and additional
information on P. gutturosus Berthold, 1845 ............................................................................ 151
Introduction ................................................................................................................................ 152
Materials and Methods ............................................................................................................. 153
Polychrus jacquelinae sp. n. ................................................................................................... 154
Polychrus peruvianus (Noble, 1924)...................................................................................... 164
Polychrus gutturosus Berthold, 1845 ..................................................................................... 172
Discussion .................................................................................................................................. 177
5. CONTRIBUTIONS TO THE GENUS AMEIVA................................................................................... 182
5.1 Two new endemic species of Ameiva (Squamata: Teiidae) from the dry forest of
northwestern Peru and additional information on Ameiva concolor Ruthven, 1924 .......... 183
Introduction ................................................................................................................................ 184
Materials and Methods ................................................................................................ 186
Phylogenetic analyses ................................................................................................. 191
Ameiva nodam sp. nov. ............................................................................................... 191
Ameiva aggerecusans sp. nov. .................................................................................... 204
Ameiva concolor Ruthven 1924 ................................................................................... 215
Niche comparisons ...................................................................................................... 221
Discussion ................................................................................................................... 224
6. CONCLUSIVE SUMMARY .............................................................................................................. 228
3
CONTENT
7. REFERENCES ................................................................................................................................ 232
8. APPENDICES ................................................................................................................................. 247
8.1 Appendix of Chapter 2.1 ....................................................................................................... 248
8.2 Appendix of Chapter 3.1 ....................................................................................................... 254
8.3 Appendix of Chapter 4.1 ....................................................................................................... 255
8.4 Appendix of Chapter 5.1 ....................................................................................................... 255
9. PUBLICATIONS ................................................................................................................................. 260

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ACKNOWLEDGEMENTS
ACKNOWLEDGEMENTS
This dissertation would not have been possible without the help of many people and
institutions, some of which have already been acknowledged in the specific
publications, where their help was needed. Nevertheless there are still many more
people and institutions which I would like to thank:

Zu allererst möchte ich mich bei meiner Familie bedanken, die in mir schon in jungen
Jahren das Interesse an Biologie geweckt und mich einen respektvollen Umgang mit
der Natur gelehrt hat. Danke, dass Ihr immer für mich da wart, nie aufgehört habt an
mich zu glauben, mich all die Jahre unterstützt habt, mir so vieles ermöglicht und mir
den Rücken freigehalten habt, wenn es notwendig war!

I am indebted to Prof. Wolfgang Böhme and Prof. Wolfgang Wägele for supervising
my thesis. Prof. Wolfgang Böhme I further would like to thank for many fruitful
discussions and for being a professional guide throughout the years of my PhD
studies.

Thanks to my field assistants Alfredo Beraún, Antonio Garcia, Erick Hoyos, Jorge
Novoa, Manuel Palacios and Pablo Venegas for helping me to plan and coordinate
the field trips, for their support in the field, for the fun we had and for teaching me
Spanish slang.
I am further indebted to all the other people of CORBIDI, especially Wilfredo Ñáñez
Aizcorbe and Julio Rivera, who did not accompany me during my field trips but
helped me with Peruvian authorities and became good friends.

Stefan Ziemendorff, Sheila Falen and her family, Daniel Plenge, Fernando Balta,
Luciano Troyes, Napoleon Monsalve, the many people I met in Peru and who
became my friends and all the people who have offered us accommodation and who
cooked for us during the field trips I would like to thank for giving me a second home
and for making my time in Peru an unforgettable experience!
Dr. Andreas Schmitz kindly familiarized me with phylogenetic analyses and Dr.
Thomas Ziegler and Viola Hartmann showed me how to evert hemipenes of
preserved reptiles.
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ACKNOWLEDGEMENTS
I am indebted to Ulla Bott, Dr. Dennis Rödder, Dr. Johara Bourke, Dr. Stephanie de
Púry, Faraham Ahmadzadeh, Janina Aurich, Timo Hartmann, Flora Ilhow, Dr. André
Koch, Peter Geissler, Tran Thi Anh Dao, Dr. Truong Nguyen and Dr. Philipp Wagner
for their good company and fruitful conversations at the ZFMK and Morris Flecks also
for helping me with some data analyses.
I thank the DAAD, the AKG and the AKS for the financial support of my field trips.
Thanks to Melissa Koch for proofreading parts of my thesis.
My boss from infill kindly “allowed” me to stay away from work for the times of my
field work and my colleagues kindly took my projects off my hands during my stay
abroads.
I am grateful to my good friends for some diversion when it was urgently needed and
for their permanent mental support, their patience and understanding.
Finally I would like to thank everyone who did not permanently ask me, how long it
will still take until I am about to finish this thesis.
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