Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2): i-iii (e128).

Preface

Amphibian & Reptile Conservation

Special Angola and Africa Issue

William R. Branch

Curator Emeritus Herpetology, Bayworld, P.O. Box 13147, Humewood 6013, SOUTH AFRICA (Research Associate, Department of

Zoology, P.O. Box 77000, Nelson Mandela Metropolitan University, Port Elizabeth 6031, SOUTH AFRICA)

Citation: Branch WR. 2016. Preface ( Amphibian & Reptile Conservation Special Angola and Africa Issue). Amphibian & Reptile Conservation 10(2):

i-iii (el28).

4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided the original author

and the official and authorized publication sources are recognized and properly credited. The official and authorized publication credit sources, which

will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website <amphibian-reptile-conservation.

org>.

Published: 30 November 2016

Africa is a mega continent that was isolated for long pe¬

riods of its history. However, after the tectonic activity

and uplift that accompanied Gondwana’s breakup and

Africa’s birth c. 130 Ma the continent was relatively qui¬

escent for nearly 100 million years until the development

of the East African Rift system 31-30 mMa (Ring 2014).

Prior to this erosion prevailed and African landforms

underwent a complicated evolution. The mechanics and

consequences of both the accumulation of Kalahari sands

in the central basin, and the relatively recent cutback of

small coastal rivers through the fragmenting Great Es¬

carpment to drain the great palaeolakes of the interior,

are now slowly being pieced together (Senut et al. 2009;

Cotterill and De Wit 2013; Flugel et al. 2015; Neumann

and Bamford 2016). During this time of isolation Africa

developed many unique biological lineages and commu¬

nities, not the least of which are Afrotherian mammals

and the world’s most diverse assemblage of venomous

snakes.

At least 1,648 reptile species are known from main¬

land Africa (Uetz and Hosek 2016), but this is likely to

be a significant underestimate, given the high reptile di¬

versities of well-studied faunas such as those of Mexico

(>800 species) and Australia (>900 species). Even within

Africa many regions are rarely surveyed resulting in se¬

vere knowledge gaps. This is reflected in the number of

African reptile records in public databases, where for

Australia there are nearly 780 records per species, 475

for Mexico, but only 60 records per species for Africa.

Many areas, particularly in the Congo Basin, the Sahel

and the Horn of Africa, have zero records for many spe¬

cies (Tolley et al. 2016), and although there remain nu-

Amphib. Reptile Conserv.

merous additional locality records dispersed in museum

collections, they have not been consolidated and made

easily accessible. This hinders the study of the African

herpetofauna, preventing fuller understanding of its ori¬

gins and diversity, and therefore its conservation (Tolley

et al. 2016).

Although there have been numerous regional surveys

for reptiles south of the Equator, e.g., southern Africa

(Branch 1998), East Africa (Spawls et al. 2001), Zambia

(Broadley et al. 2003), Tanzania (Broadley and Howell

1991), etc., the last detailed review of Angolan herpetol¬

ogy was prepared over 120 years ago (Bocage 1895) and

consequently is now very out-of-date. The Reptile Da¬

tabase (Uetz and Hosek 2016) currently lists 253 reptile

species from Angola. However, this list, which is auto¬

matically generated, contains incorrect inclusions, e.g.,

Comsophis bonlengeri (Madagascar) and Micrurus bo-

gerti (Mexico), Agama finchi (East Africa), Trachylepis

quinquetaeniata (northeast Africa), etc. It also includes

species that have been fragmented by recent revisions,

and where the revived local species is included along

with the original species with which it was synonymized

and which no longer occurs in Angola, e.g., Chamae-

saura macropholis, Cordylus cordulus, Cordylus vittifer ,

Holaspis laevis, Leptotyphlops nigricans , Pachydactylus

serval , Pedioplanis undata, etc.

Southern Africa, the geographic region south of the

Cunene and the Zambezi rivers, has the most diversity

reptile fauna in Africa, both in terms of species and fam¬

ily richness (Branch 2006). South Africa, in particular,

has exceptional reptile diversity, with nearly 400 spe¬

cies and 44% endemicity (Branch 2014 and updates).

November 2016 | Volume 10 | Number 2 | e128

Branch

SPECIES ACCUMULATION CURVE

— Angola — Southern Africa

Figure 1 . Species accumulation curves for Angolan (red) and southern African (blue) reptiles.

Although alpha diversity for Namibia is lower, with 228

species and just 22% endemicity (Herrmann and Branch

2013 and updates), this is in part due to its smaller geo¬

graphic area (Namibia 0.82 million km 2 , South Africa

1.22 million km 2 ) and reduced habitat diversity. Angola,

however, is approximately the same size as South Af¬

rica (1.25 million km 2 ) with diverse habitats ranging

from the northern section of the world’s oldest desert,

the Namib, in the southwest, to lowland tropical rain

forest of the Congo Basin in the north. In addition, the

complex topography and geology of northern outliers

of the Great Escarpment and numerous central isolated

highlands create further habitat complexity. Despite this

the known Angolan reptile diversity (about 253) is only

slightly richer than that of adjacent Namibia and much

lower than that of South Africa. A species accumulation

curve for the description of reptiles from southern Af¬

rica has shown no decline during the last 150 years, and

now exceeds 600 taxa (Fig. 1). In contrast the curve for

Angola has shown very little increase during the last cen¬

tury (Fig. 1). Normally this would infer that the country’s

reptile fauna is well known and that few new species

remain to be discovered. However, recent surveys (e.g.,

Huntley 2009; Huntley and Francisco 2015; Ceriaco et

al. 2014, 2016) have uncovered numerous new species

(e.g., Conradie et al. 2012,2013; Stanley et al. 2016), and

it is evident that Angolan reptile diversity simply remains

poorly known. The articles presented in this special issue

detail new discoveries, provide updated checklists, and

taxonomic discussion, and begin to reveal the true extent

of Africa’s rich herpetofauna.

Literature Cited

Bates MF, Branch WR, Bauer AM, Burger M, Marais J,

Alexander GJ, de Villiers MS. 2014. Atlas and Red

List of the Reptiles of South Africa, Lesotho and Swa¬

ziland. South African National Biodiversity Institute,

Amphib. Reptile Conserv. ii

Pretoria. 487 p.

Bocage JVB. 1895. Herpetologie d Angola et du Congo.

Imprimerie Nationale, Fisbonne, Portugal. 203 p.

Branch WR. 1998. Field Guide to the Snakes and other

Reptiles of Southern Africa. Revised edition. Struik

Publishers, Cape Town, South Africa. 399 p.

Broadley DG, Doria CT, Wigge J. 2003. Snakes of Zam¬

bia. An Atlas and Field Guide. Edition Chimaira,

Frankfurt am Main, Germany. 280 p.

Broadley DG, Howell KM. 1991. A checklist of the rep¬

tiles of Tanzania, with synoptic keys. Syntarsus 1:

1-70.

Ceriaco FMP, Bauer AM, Blackburn DC, Favres ACFC.

2014. The herpetofauna of the Capanda Dam region,

Malanje, Angola. Herpetological Review 45(4): 667-

674.

Ceriaco FMP, de Sa SC, Bandeira S, Valerio H, Stanley

EF, Kuhn AF, Marques M, Vindum JV, Blackburn

DC, Bauer AM. 2016b. Herpetological survey of Iona

National Park and Namibe Regional Natural Park,

with a Synoptic list of the Amphibians and Reptiles of

Namibe Province, Southwestern Angola. Proceedings

of the California Academy of Sciences 63(2): 15-61.

Conradie W, Measey JG, Branch WR, Tolley KA. 2012.

Revised phylogeny of African sand lizards (Pedio-

planis), with the description of two new species from

south-eastern Angola. African Journal of Herpetology

61(2): 91-112.

Conradie W, Branch WR, Tolley KA. 2013. Fifty Shades

of Grey: Giving colour to the poorly known Ango¬

lan Ash reed frog (Hyperoliidae: Hyperolius cinere-

us ), with the description of a new species. Zootaxa

3635(3): 201-223.

Cotterill F, De Wit M. 2011. Geoecodynamics and the

Kalahari Epeirogeny: Finking its genomic record,

tree of life and palimpsest into a unified narrative of

landscape evolution. South African Journal Geology

114(3-4): 489-514.

Flugel TJ, Eckardt FD, Cotterill FPD. 2015. The Pres-

November 2016 | Volume 10 | Number 2 | e128

Preface

ent Day Drainage Patterns of the Congo River System

and their Neogene Evolution. Pp. 315-337 In: Editor,

M.J. de Wit et al. MJ. Geology and Resource Poten¬

tial of the Congo Basin. Regional Geology Reviews,

Springer-Verlag Berlin Heidelberg, Germany. 417 p.

Huntley B, Francisco P. (Editors) 2015. Avaliagao Ra-

pida da Biodiversidade da Regiao da Lagoa Carumbo.

Relatorio sobre a expedigao. - Rapid Biodiversity As¬

sessment of the Carumbo Lagoon area, Lunda Norte,

Angola. Expedition Report. Republica de Angola

Ministerio do Ambiente, Lunda Norte, Angola. 219 p.

Neumann FH, Bamford MK. 2015. Shaping of modern

southern African biomes: Neogene vegetation and

climate changes. Transactions of the Royal Society of

South Africa 70(3): 195-212.

Ring U. 2014. The East African Rift System. Austrian

Journal of Earth Sciences 107: 132-146.

Senut B, Pickford M, Seldalen L. 2009. Neogene deserti¬

fication of Africa. Comptes Rendus, Geoscience 341:

591-602.

Spawls S, Howell K, Drewes R, Ashe J. 2001. A Field

Guide to the Reptiles of East Africa. Princeton Uni¬

versity Press, Princeton, New Jersey. 544 p.

Stanley EL, Ceriaco, LMP, Bandeira S, Valerio H, Bates

MF, Branch WR. 2016. A review of Cordylus mach-

adoi (Squamata: Cordylidae) in southwestern Angola,

with the description of a new species from the Pro-

Namib desert. Zootaxa 4061(3): 201-226.

Tolley KA, Alexander GJ, Branch WR, Bowles P, Maritz

B. 2016. Conservation status and threats for African

reptiles. Biological Conservaton 204: 63-67.

Uetz, P, Hosek J. 2016. The Reptile Database. Available:

http://www.reptile-database.org [Accessed: 28 May

2016],

Amphib. Reptile Conserv.

November 2016 | Volume 10 | Number 2 | el28

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [Special Section]: 1-5 (e125).

SHORT COMMUNICATION

Senegal Flapshell Turtle (Cyclanorbis senegalensis) in

Ethiopia (Testudines: Trionychidae)

^omas Mazuch, 2 Vladimir Trailin, 3 Uwe Fritz, and *’ 3 Melita Vamberger

'Dfitec 65, 53305 Dfitec, CZECH REPUBLIC 2 Jana Masaiyka 1319, 50012 Hradec Kralove, CZECH REPUBLIC 3 Museum of Zoology?,

Senckenberg Dresden, A. B. Meyer Building, 01109 Dresden, GERMANY

Abstract .—Based on DNA sequences derived from two hatchlings from the Alwero river, Gambela

Region, Cyclanorbis senegalensis is recorded for the first time for western Ethiopia. Previously

published DNA sequences of C. senegalensis from Benin and Togo are slightly different,

suggesting phylogeographic structure.

Keywords. Africa, Alwero river, Cyclanorbinae, first record, Gambela Region, Reptilia

Citation: Mazuch T, Trailin V, Fritz U, and Vamberger M. 2016. Senegal Flapshell Turtle ( Cyclanorbis senegalensis) in Ethiopia (Testudines:

Trionychidae). Amphibian & Reptile Conservation 10(2) [Special Section]: 1-5 (e125).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any

medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and

authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official

journal website <amphibian-reptile-conservation.org>.

Received: 31 May 2016; Accepted: 5 July 2016; Published: 12 October 2016

Flapshell turtles (Cyclanorbinae) are a subfamily of

softshell turtles (Trionychidae) that include four species

in two genera ( Cycloderma , Cyclanorbis , each with

two species) in sub-Saharan Africa, and three species

of the genus Lissemys in the Indian subcontinent,

Myanmar, and perhaps Thailand (Branch 2008; Praschag

et al. 2011; van Dijk et al. 2014). The Asian species

constitute the sister group to the African taxa (Engstrom

et al. 2004). All cyclanorbines are characterized by the

presence of femoral and caudal flaps on the plastron,

together with the movable plastral forelobe allowing the

complete closure of the shell—a unique character among

softshell turtles.

The African species have been relatively little studied

and many records are historical (Gramentz 2008;

Broadley and Sachsse 2011; Baker et al. 2015; cf. also the

EMYSystem Database). The two Cyclanorbis species,

C. elegans and C. senegalensis , co-occur throughout

much of their distribution (Fig. 1), with C. senegalensis

having a much wider range. However, for non-specialists,

the two species are not easy to tell apart. They differ in

adult size and C. elegans has two, or less commonly,

four plastral callosities, while the smaller C. senegalensis

has up to nine callosities (Branch 2008; Baker et al.

2015). Yet, considering that the number of plastral

callosities may change during growth, it is obvious that

Correspondence. Email: *mvamberger@senckenberg.de

the two species can be easily confused and that some

records of either species may actually refer to the other.

In particular hatchlings and small juveniles are difficult

to determine.

Even though Ethiopia is included in the distribution

ranges of C. elegans and C. senegalensis by the IUCN Red

List of Threatened Species, neither species has ever been

recorded from that country (Largen and Spawls 2010).

However, the two species are known from the White

Nile system close to the Ethiopian border (EMYSystem

Database). Largen and Spawls (2010) suggested that

both C. elegans and C. senegalensis may occur in the

Baro river in Ethiopia because records exist downstream

in South Sudan, less than 30 km from the border. Also

Baker et al. (2015) speculated that C. elegans could occur

in westernmost Ethiopia, corresponding to the catchment

basins of the Baro and Akobo rivers, both tributaries of

the White Nile.

On 26 August 2014, during an excursion to Ethiopia,

one of us (V.T.) obtained two hatchlings of a flapshell

turtle near Ugudi village at the Alwero river (Gambela

Region, 7°58’55.83”N, SriCOG.lTE, 439 meters

above sea level). The Alwero river is a tributary of the

Baro river and the collection site is approximately 150

km distant from the closest known occurrence of the

species in South Sudan (Nasir; Siebenrock 1909; Largen

Amphib. Reptile Conserv.

October 2016 1 Volume 10 I Number 2 I el25

Mazuch et. al

Senegal

Sudan

Burkina Faso

Nigeria

Guinea

South Sudan

Central African Republic

Ghana,

& \ Cameroon

Ugand;

Gabon

Congo, DRC

Tanzania

Angola

Saudi Arabia

» I

) \ 1

Guinea

Bissau

Sierra Leone

Ethiopia

Somalia

Equatorial Guinea

' WCyclanorhis senegalensis (MTD D 49181,49182)

# Cyclanorbis senegalensis

□ Cyclanorbis elegans

500 1000

■ i Km

Fig. 1 . Distribution of Cyclanorbis elegans and C. senegalensis (localities from EMYSystem Database plus additional records

from Sierra Leone, see inset, and Ethiopia). Inset: Adult C. senegalensis from Sierra Leone, Rokel river at Robung village.

Photo: Bill Branch.

and Spawls 2010). The two specimens (Fig. 2) are now in

the collection of the Museum of Zoology, Senckenberg

Dresden (MTD D 49181,49182). The turtles were caught

by native children during daytime, in a temporary pool

close to the river. The species is well-known to locals. In

this region, late August is the climax of the rainy season

and lush vegetation at the collection site comprised dense

elephant grass. The straight line carapacial lengths of the

two specimens were 47.9 mm (MTD D 49181) and 40.6

mm (MTD D 49182).

Tissue samples of the ethanol-preserved hatchlings

were used for genetic characterization because species

identity was unclear. Laboratory procedures and

phylogenetic calculations are described in detail in the

Supporting Information of this article (Dryad Repository,

http://dx.doi.org/10.5061/dryad.g48pb). Sequences of

three mitochondrial DNA blocks (in total 2,429 bp)

that had previously been shown to be taxonomically

informative for softshell turtles (e.g., Engstrom et al.

2004; Fritz et al. 2010, 2014; Praschag et al. 2011)

were generated. Fragment 1 corresponded to 398 bp of

the 12S rRNA gene. The 1,140-bp-long fragment 2

represented the complete cytochrome b (cyt b )

gene. Fragment 3 contained 709 bp of the NADH

dehydrogenase subunit 4 gene (ND4) and 182 bp of

adjacent DNA coding for tRNAs. These individual DNA

Species

Provenance

Reference

12S

cyt b

ND4+tRNAs

Cyclanorbis elegans

Benin

Engstrom et al. (2004)

n/a

AY259570

AY259615

Cyclanorbis senegalensis

Benin

Praschag et al. (2011)

FR850553

FR850654

FR850604

Cyclanorbis senegalensis

Togo

Engstrom et al. (2004)

n/a

AY259569

AY259614

Cyclanorbis senegalensis

Ethiopia: Ugudi village, Gambela Region

This study (MTD D 49181)

LT595717

LT595719

LT595722

Cyclanorbis senegalensis

Ethiopia: Ugudi village, Gambela Region

This study (MTD D 49182)

LT595718

LT595720

LT595723

Cycloderma aubryi

Congo Brazzaville: Pointe Noire

Praschag et al. (2011)

FR850554

FR850655

FR850605

Cycloderma aubryi

Congo Brazzaville: Tchingoli

Praschag et al. (2011)

FR850555

FR850656

FR850606

Cycloderma aubryi

Gabon

Engstrom et al. (2004)

n/a

AY259566

AY259611

Cycloderma frenatum

Lake Malawi

Engstrom et al. (2004)

n/a

AY259565

AY259610

Lissemys ceylonensis

Sri Lanka: Colombo

Praschag et al. (2011)

FR850544

FR850645

FR850595

Table 1 . Used samples and GenBank sequences and their accession numbers.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | e125

Senegal flapshell turtle in Ethiopia

Fig. 2. Hatchlings of Cyclanorbis senegalensis from the vicinity of Ugudi village at the Alwero river, Gambela Region, western

Ethiopia. Left: MTD D 49181, right: MTD D 49182. Scale bars, 1 cm.

blocks were concatenated and merged for calculations

with previously published homologous data of all four

African flapshell turtle species. Sequences of Lissemys

ceylonensis were included as outgroup (Table 1).

Phylogenetic trees were then computed using Bayesian

Inference and Maximum Likelihood approaches as

implemented in mrbayes 3.2.1 (Ronquist et al. 2012) and

RAxML 7.2.8 (Stamatakis 2006). In addition, uncorrected

p distances between concatenated DNA sequences were

calculated using MEGA 5.05 (Tamura et al. 2011) and

the pairwise deletion option.

Both tree building methods yielded identical

topologies (Fig. 3), with the western Ethiopian samples

clustering with maximum support with C. senegalensis.

However, while the sequences of the Ethiopian samples

were completely identical, they differed slightly from

sequences for C. senegalensis from Benin and Togo.

When uncorrected p distances were compared, sequence

*/ 96

0 . 99/70

Lissemys ceylonensis

■ Cycloderma frenatum (Lake Malawi)

*/99

0 . 91/96

■Cycloderma aubryi (Gabon)

Cycloderma aubryi (Congo Brazzaville: Tchingoli)

Cycloderma aubryi (Congo Brazzaville: Point Noire)

.Cyclanorbis elegans (Benin)

MTD D 49181 (Ethiopia)

0 . 69/42

MTD D 49182 (Ethiopia)

—Cyclanorbis senegalensis (Togo)

-Cyclanorbis senegalensis (Benin)

0.04

Fig. 3. Maximum Likelihood (ML) tree for flapshell turtles (Cyclanorbinae) using 2,429 bp of mtDNA. Values at nodes are

Bayesian posterior probabilities and ML bootstrap support. Asterisks indicate maximum support under one or both

approaches. Note placement of Ethiopian samples (in red).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | e125

Mazuch et. al

divergences among the Ethiopian samples and the

C. senegalensis from Benin and Togo ranged between

1.5% and 1.9%, whereas divergences between C. elegans

and C. senegalensis differed by one order of magnitude

(15.2-15.4%; cf. Supporting Information). Thus, the

hatchlings from western Ethiopia are clearly referable

to C. senegalensis. The observed sequence divergences

within C. senegalensis suggest that phylogeographic

structuring exists and that further studies should be

conducted to examine this situation.

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PCH, van Dijk PP, Saumure RA, Buhlmann KA,

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Branch B. 2008. Tortoises, Terrapins & Turtles of Africa.

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Broadley DG, Sachsse W. 2011. Cycloderma frenatum

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PCH, van Dijk PP, Saumure RA, Buhlmann KA,

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EMYSystem Database. Available: http://emys.geo.orst.

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Engstrom TN, Shaffer HB, McCord WP. 2004. Multiple

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Hundsdorfer AK. 2010. The world’s economically

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P. 2014. Phylogeography of the Asian softshell turtle

Amy da cartilaginea (Boddaert, 1770): Evidence for

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GramentzD. 2008. African Flapshell Turtles, Cyclanorbis

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LargenM, Spawls S. 2010. The Amphibians and Reptiles

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Praschag P, Stuckas H, Packert M, Maran J, Fritz U. 2011.

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Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | e125

Senegal flapshell turtle in Ethiopia

Tomas Mazuch is a Czech amateur herpetologist. Since his teenage age he has dedicated his

life to the breeding of amphibians, reptiles, and invertebrates. During his studies of Veterinary

Sciences (not finished yet) he began to devote his research to herpetology and parasitology

of reptiles. His research focuses on taxonomy, systematics, and biogeography of amphibians

and reptiles of the Horn of Africa (mainly Somalia and Ethiopia). His main subjects of

study are geckos of the genus Hemidactylus from Eastern Africa. He is also interested in the

taxonomy of scorpions from North-Eastern Africa. He has authored or co-authored over 15

peer-reviewed papers and books on parasitology, systematics of scorpions, and herpetology,

including the book “Amphibians and Reptiles of Somaliland and Eastern Ethiopia, ...based on

two field trips in 2010/2011” (2013). He has co-authored the descriptions of six reptile and four

scorpion species.

Vladimir Trailin was born in Susice, a town in southern Bohemia in the Sumava Mountains.

The neighbourhood of the Sumava National Park formed his interest of nature. In 1978-1982,

he graduated from the High School of Mechanical Engineering, but remained continuously

interested in nature and natural history, especially entomology and herpetology. He travelled

many times to the Balkan Peninsula, also to Cuba, and for the first time to Africa in 1992.

Since then he has repeatedly visited Ethiopia, and together with colleagues has rediscovered

some poorly known snakes (e.g., Lamprophis erlangeri and Bids parviocula ), as well as the

first live individuals of Pseudoboodon boehmei. The species Cicindela trailini (Coleoptera),

Hottentotta trailini (Scorpiones), and Pandinus trailini (Scorpiones) are dedicated to him.

Uwe Fritz is director of the Senckenberg Natural History Collections at Dresden, Gemiany,

and extraordinary professor for zoology at the University of Leipzig. He has worked for many

years on the taxonomy, systematics, and phylogeography of turtles and tortoises, and also

studied to a lesser extent snakes and lizards. He is particularly interested in hybridization

patterns and gene flow in contact zones of distinct taxa. Uwe has authored or co-authored

numerous scientific articles, mainly in herpetology, and has also edited proceedings and

books, among them the two turtle volumes of the “Handbook of Amphibians and Reptiles

of Europe.” He is currently the President of the German Society for Biological Systematics,

and is responsible for the Senckenberg collections and the scientific journals edited by the

Senckenberg Society.

Melita Vamberger is a Slovenian herpetologist and evolutionary biologist. She studied

biology at the University Ljubljana, with the focus on the natural history of the European pond

turtle ( Emys orbicularis ). After her diploma she moved to Germany and studied for her Ph.D.

thesis (2014) at the University of Leipzig on the phylogeography and hybridization of two

closely related freshwater turtles ( Mauremys caspica and M. rivulata). Since then she has been

employed as a researcher at Senckenberg Dresden. Her main interests are studying processes of

speciation, gene flow, and evolution in different turtle taxa using genetic methods, with a focus

on the Western Palearctic and sub-Saharan Africa.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | e125

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [Special Section]: 6-36 (e126).

The herpetofauna of the Cubango, Cuito, and lower

Cuando river catchments of south-eastern Angola

12 *Werner Conradie, 2 Roger Bills, and 13 WilIiam R. Branch

1 Port Elizabeth Museum (Bayworld), P.O. Box 13147, Humewood 6013, SOUTH AFRICA 2 South African Institute for Aquatic Bio¬

diversity, P/Bag 1015, Grahamstown 6140, SOUTH AFRICA 3 Research Associate, Department of Zoology, P 0 Box 77000, Nelson

Mandela Metropolitan University, Port Elizabeth 6031, SOUTH AFRICA

Abstract.—Angola’s herpetofauna has been neglected for many years, but recent surveys have revealed

unknown diversity and a consequent increase in the number of species recorded for the country. Most historical

Angola surveys focused on the north-eastern and south-western parts of the country, with the south-east,

now comprising the Kuando-Kubango Province, neglected. To address this gap a series of rapid biodiversity

surveys of the upper Cubango-Okavango basin were conducted from 2012-2015. This report presents the

results of these surveys, together with a herpetological checklist of current and historical records for the

Angolan drainage of the Cubango, Cuito, and Cuando Rivers. In summary 111 species are known from the

region, comprising 38 snakes, 32 lizards, five chelonians, a single crocodile and 34 amphibians. The Cubango

is the most western catchment and has the greatest herpetofaunal diversity (54 species). This is a reflection

of both its easier access, and thus greatest number of historical records, and also the greater habitat and

topographical diversity associated with the rocky headwaters. As a result of these surveys, five new species

records were added to the Cubango catchment, 17 to the Cuito catchment, and nine to the Cuando catchment.

Seven of the records for the eastern catchments, including three for the Cuito and four for the Cuando, were

also new for Angola.

Keywords. Herpetofauna, Angola, Okavango, Cuito, Cubango, Cuando

Resumo.—A herpetofauna de Angola foi negligenciada durante muitos anos, mas varios levantamentos

realizados recentemente revelaram uma diversidade desconhecia e um consequente aumento no numero de

especies registadas para o pais. A maior parte dos levantamentos historicos realizados em Angola focaram-

se no nordeste e sudoeste do pais, sendo o sudeste, que agora abrange a provincia do Kuando-Kubango,

bastante negligenciado. Para preencher esta lacuna, foram realizados varios levantamentos de biodiversidade

na bacia superior do Cubango-Okavango entre 2012 e 2016. Neste relatorio sao apresentados os resultados

destes levantamentos, bem como uma lista de registos herpetologicos recentes e historicos para as bacias

dos rios Cubango, Cuito, e Cuando. Resumidamente, conhecem-se 111 especies para a regiao, incluindo 39

especies de cobras, 32 especies de lagartos, 5 especies de quelonios, apenas um crocodilo, e 34 especies

de anfibios. A bacia hidrografica do Cubango e a que esta localizada mais a oeste, e tern a maior diversidade

de herpetofauna (54 especies). Isto reflecte por um lado a sua maior acessibilidade, e consequentemente

um maior numero de registos historicos, e ainda a maior diversidade topografica e de habitat, associada as

nascentes rochosas. Como resultado destes levantamentos, foram adicionadas cinco novas especies a bacia

do Cubango, 17 a do Cuito, e nove a do Cuando. Sete dos registos para as bacias a este, incluindo trois para o

Cuito e quatro para o Cuando, sao tambem novos registos para Angola.

Palavras-chave. Herpetofauna, Angola, Okavango, Cuito, Cubango, Cuando

Citation: Conradie W, Bills R, and Branch WR. 2016. The herpetofauna of the Cubango, Cuito, and lower Cuando river catchments of south-eastern

Angola. Amphibian & Reptile Conservation 10(2) [Special Section]: 6-36 (el26).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any

medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and

authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal

website <amphibian-reptile-conservation.org>.

Received: 12 April 2016; Accepted: 31 May 2016; Published: 25 October 2016

Correspondence. *Email: werner@bayworld.co.za

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Introduction

Although neglected for many years, studies on the herpe-

tofauna of Angola have increased in recent years. Despite

this, the Angolan herpetofauna remains one of the most

poorly documented in Africa, particularly compared with

Namibia to the south (Herrmann and Branch 2013). The

only detailed synthesis occurred in the 19th century (Bo-

cage 1895), although Monard (1937a, b) presented sub¬

sequent updates. To complicate this neglect, most reports

dealing with the country’s herpetofauna, including the

early explorations upon which much of Bocage’s seminal

studies were based, were restricted to the western regions

of the country. Studies in the 20 th century, including those

of Schmidt (1933, 1936), Parker (1936), Mertens (1938),

Bogert (1940), FitzSimons (1959), Hellmich (1957a,

1957b), Poynton and Haacke (1993), Ruas (1996, 2002),

Haacke (2008), etc., did little to redress this geographical

bias. The catalogues of Laurent (1950,1954) and Tys van

den Audenaerde (1967), based on material sent to the au¬

thors from Museu do Dundo, listed numerous additional

species for extreme north-east Angola. Laurent (1964)

later presented a detailed report on additional material

from Museu do Dundo, as well as a collection by Barros

Machado from the south-west semiarid region of Angola

that included important new discoveries. South-eastern

Angola, which includes extensive wetland and miombo

habitats that are rare elsewhere in the country, remains

one of the most neglected regions in Angola. In part, this

neglect stemmed from the historical difficulties of access

to the flat wetlands of the region that drain south into the

Okavango Delta, a famous World Heritage Site. The dif¬

ficulty of access was further exacerbated by a protracted

civil war (1975-2002), with the destruction of the little

regional infrastructure that existed and the deployment of

extensive and poorly-documented mine fields.

Following the cessation of hostilities and the ongoing

redevelopment of regional infrastructure, modern biodi¬

versity surveys in the country have begun (e.g., Huntley

2009, Huntley and Francisco 2015; Brooks 2012, 2013;

Wild Bird Trust 2015; Ceriaco et al. 2016a, b). Some

have targeted areas that have never been scientifically

surveyed, and have led to the discovery and description

of new endemic species of amphibians (Conradie et al.

2012a, 2013) and reptiles (Conradie et al. 2012b, Stanley

et al. 2016). Others have resolved previous taxonomic

confusion (Channing et al. 2013; Channing and Baptista

2013; Ernst et al. 2015), or noted the addition of new

country records (Branch and Conradie 2013; Conradie

and Bourquin 2013; Ernst et al. 2014; Ceriaco et al. 2014,

2016a; Ernst et al. 2015). However, no formal herpeto-

logical surveys have previously been undertaken in the

Cubango, Cuito, and Cuando river catchments of south¬

east Angola. Previous material from the region included

only opportunistic collections and was mostly confined

to the western tributaries of the Cubango river basin

(Bocage 1895; Monard 1931, 1937a, b), with very few

records from the Cuito and Cuando river basin (Angel

1923). The only recent collection came from the Cuito-

Cuanavale area and added an additional five reptiles for

the region, mostly with fossorial habits and encountered

during excavations associated with military activity at

the time (Branch and McCartney 1992). To redress igno¬

rance of the biodiversity of the important wetland associ¬

ated with the Angolan drainage of the Okavango Delta

a number of international surveys have been initiated

(Brooks 2012, 2013; Wild Bird Trust 2016). All have in¬

cluded dedicated herpetological surveys, and the results

of these surveys and an updated checklist of the herpe¬

tofauna of south-eastern Angola based on both historical

and recent collections are presented here.

Methods

Surveys

The Okavango Delta is an internationally acclaimed

natural wonder and was recently ratified as a World Heri¬

tage Site in 2014. Although the Okavango Delta is well

protected within Botswana, there is a need to conserve

and assess biodiversity in the headwaters of the rivers

that drain south into the Okavango Delta and the associ¬

ated pans. To this end, a number of international biodi¬

versity surveys have recently been undertaken. They are

informed, in part, by the aims of the Strategic Action Pro¬

gramme of the Permanent Okavango River Basin Water

Commission (OKACOM), in accord with the Angolan

National Action Plan for the Sustainable Management

of the Cubango/Okavango River Basin (Okacom 2011).

As part of the Southern Africa Regional Environmen¬

tal Program (SAREP), in collaboration with the Ango¬

lan Ministry of Environment - Institute of Biodiversity

(MINAMB) and the Angolan Ministry of Agriculture -

National Institute of Fish Research (INIP), two surveys

were undertaken, including 1) May 2012, upper catch¬

ment of the Cubango-Okavango River basin (Brooks

2012); and 2) April 2013, lower Cuito and Cuando River

systems (Brooks 2013). The third and most recent survey

formed part of the National Geographic funded Okavan¬

go Wilderness Project (Wild Bird Trust 2015). It concen¬

trated on the Cuito River, from its source to its conflu¬

ence with the Cubango River. Collection sites of the

three surveys are listed in Table 1 and locality maps are

presented in Figures 1-3. Specimens from the watershed

of the Kwanza-Okavango-Zambezi Rivers, including the

source lakes of the Cuito and Cuanavale Rivers, will be

presented elsewhere (Baptista et al. in prep.) following

recent expeditions (2016) to the region.

Study area

The geographical scope of the three surveys is defined

by the Cubango-Okavango basin (Fig. 1). The area con¬

sists of two main rivers systems: the Cubango and the

October 2016 | Volume 10 | Number 2 | el 26

Amphib. Reptile Conserv.

Herpetofauna of river catchments in south-eastern Angola

16.0

18.0

20.0

22.0

Legend

• Sites 2012

Major Rivers

Cubango Basin

Cuito Basin

Cuando Basin

18.0

20.0

22.0

csi

16.0

100

200 km

NAMIBIA +

Moxico

Huambo

Huila

Cunene

ZAMBIA

Fig. 1. Map of the study area in south-eastern Angola, indicating surveyed sites for May 2012 survey.

Cuito Rivers (Fig. 1-3), both draining south-east into the

Okavango Delta. The Cubango River and its tributaries

lie to the west of the study area. They are underlain in

their headwaters by granite outcrops and characterized in

places by rocky substrates, rapids, and some waterfalls.

Typical habitats of the Cubango basin are shown in Fig.

4. In contrast, the Cuito River and its major tributary the

Cuanavale River lie to the east and have wide valleys,

with water courses that meander across deep Kalahari

sands, and are characterised by extensive wet grasslands,

peatlands, and ox-bow wetlands (typical habitats shown

in Fig. 5). The impeded drainage and high precipitation in

the rainy season cause temporarily waterlogged soils that

prevent the development of woodland but support humid

grassland borders with humic topsoils and dwarf shrubs

(Revennann et al. 2013). These areas act as “sponges”

that slowly release water into the Okavango system. In

their upper reaches, the surrounding hills are dominated

by various forms of woodland savannah, particularly

miombo. The lower Cuando River has a similar topog¬

raphy to the Cuito River, but with west to east drainage

lines which form a series of floodplains and pans (typical

habitats shown in Fig. 6).

Data collection

All surveys involved opportunistic visual encounters.

Diurnal searches involved active searches of specific

microhabitats, particularly beneath rocks and decaying

logs. Nocturnal surveys for amphibians were undertaken

in wetlands and surrounding woodland. Advertisement

calls were recorded in the field using either an Olympus

VN-3500PC Voice Recorder or NAGRA ARES-ML re¬

corder with an external Sony F-V4T Microphone to aid

in species identification and have been lodged in WC

personal audio library. Standard Y-shape trap arrays were

used in two surveys (May 2012 and April 2013), and each

trap array consisted of 3 x 10 m long and 50 cm high drift

fences positioned in a Y-shape, and with four pitfall traps

(one at the center and at each fence tip) and six one-way

funnel traps placed on adjacent sides of each ann.

Specimens retained for subsequent study were hu¬

manely euthanized by injecting reptiles and submerg¬

ing frogs in tricaine methanesulfonate (MS222) solution

(Conroy et al. 2009), after which they were formalin-

fixed for 48 hours and transferred to alcohol for long¬

term storage. Prior to fixing tissue samples (either liver

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Table 1. List of collection sites during the current field surveys. Site numbers corresponds with sites in Figure 1 (a.s.l. = above sea level).

May 2012 April-May 2013 May-June 2015

Site

Latitude (S)

Longitude (E)

Elevation

(a.s.l.)

Site

Latitude (S)

Longitude (E)

Elevation

(a.s.l.)

Site

Latitude (S)

Longitude (E)

Elevation

(a.s.l.)

16.89413 S

17.95766 E

1,763 m

17.87291 S

19.83333 E

1,055 m

16.88350 S

18.01180 E

1121 m

15.70452 S

17.45897 E

1,153 m

30a

17.50875 S

20.06594 E

1,069 m

14.58981 S

16.907389 E

1408 m

14.94277 S

17.71863 E

1,248 m

30b

17.51194 S

20.04305 E

1,091 m

14.70214 S

17.37806 E

1396 m

14.81913 S

17.67450 E

1,288 m

30c

17.51222 S

20.06027 E

1,079 m

14.68136 S

17.44530 E

1458 m

14.74628 S

17.66844 E

1,332 m

30d

17.51327 S

20.06111 E

1,078 m

14.66300 S

17.66550 E

1385 m

14.67155 S

17.73525 E

1,265 m

30e

17.51430 S

20.05527 E

1,082 m

14.59517 S

18.07111 E

1497 m

14.67458 S

17.73544 E

1,369 m

30f

17.52638 S

20.05825 E

1,075 m

14.58970 S

18.1711 E

1317 m

14.42966 S

17.82658 E

1,356 m

17.46777 S

20.03333 E

1,074 m

14.59333 S

18.2242 E

1491 m

14.43377 S

17.82957 E

1,359 m

17.04880 S

19.53333 E

1,086 m

53a

14.55600 S

18.40710 E

1305 m

14.43916 S

17.81491 E

1,359 m

33a

16.90980 S

19.30769 E

1,109 m

53b

14.56322 S

18.44394 E

1276 m

14.25705 S

17.77852 E

1,404 m

33b

16.92367 S

19.29675 E

1,110 m

14.46810 S

18.35488 E

1327 m

14.00269 S

17.40500 E

1,505 m

16.77988 S

19.11667 E

1,059 m

14.68478 S

18.67369 E

1289 m

13.71616 S

17.09661 E

1,538 m

16.62322 S

19.05352 E

1,155 m

15.06275 S

19.14322 E

1240 m

13.69413 S

17.06177 E

1,554 m

17.82305 S

22.61611 E

1,019 m

15.45969 S

18.76833 E

1224 m

12a

13.59333 S

16.87986 E

1,431 m

17.67833 S

22.61475 E

1,021 m

15.38206 S

19.06375 E

1185 m

12b

13.59638 S

16.87722 E

1,516 m

17.58830 S

22.65694 E

1,004 m

16.28392 S

18.84744 E

1107 m

13.28061 S

16.74722 E

1,607 m

17.46333 S

22.86638 E

995 m

16.92367 S

19.29675 E

1123 m

12.78555 S

16.75694 E

1,633 m

17.45786 S

22.91191 E

997 m

16.98919 s

19.40614 E

1079 m

12.87242 S

16.76742 E

1,692 m

41b

17.46777 S

23.07944 E

988 m

17.50875 S

20.06608 E

1080 m

16a

12.53072 S

16.69744 E

1,643 m

41a

17.46777 S

23.06667 E

988 m

17.93611 S

21.10269 E

1018m

16b

12.54222 S

16.67694 E

1,763 m

17.49611 S

23.13444 E

980 m

—

12.57008 S

16.49111 E

1,569 m

17.53500 S

23.18916 E

981 m

—

12.67105 S

16.11111 E

1,766 m

44b

17.56916 S

23.27305 E

982 m

—

14.70213 S

17.37772 E

1,375 m

44a

17.57333 S

23.26000 E

987 m

—

14.67175 S

17.15331 E

1,344 m

—

14.66586 S

17.07661 E

1,142 m

—

22a

14.66622 S

16.97842 E

1,380 m

—

22c

14.65386 S

16.93547 E

1,367 m

—

22b

14.66278 S

16.96081 E

1,367 m

—

22d

14.64991 S

16.90739 E

1,356 m

—

14.58972 S

18.17083 E

1,316 m

—

14.60622 S

18.46722 E

1,256 m

—

15.139194 S

19.14350 E

1,303 m

—

15.08686 S

19.14872 E

1,192 m

—

15.17127 S

19.19433 E

1,180 m

—

15.13486 S

19.19636 E

1185 m

—

or muscle) were preserved in 96 % ethanol for further

genetic analysis. Voucher specimens are held in the

herpetological collections of Port Elizabeth Museum

(PEM), South African Institute for Aquatic Biodiversity

(SAIAB), and the Instituto Superior de Ciencias da Edu-

cagao da Huila (ISCED), Lubango, Angola.

Relevant field guides (Broadley 1983; Branch 1998;

Channing 2001; Broadley et al. 2003; Du Preez and Car-

ruthers 2009) were used for species identification. No¬

menclature was based on established online databases

(amphibian, Frost 2015; reptiles, Uetz and Hosek 2015),

updated where appropriate. No regional conservation as¬

sessment has been undertaken for Angolan amphibians

and reptiles as yet. Where global conservation assess¬

ments are available (e.g., IUCN 2015) they are noted. En¬

demic (defined as species whose distribution is restricted

solely to Angola) and near-endemic species (>90% of

distribution within Angola) are noted.

The following relevant literature was consulted to

compile historical records for the study area: Bocage

(1895), Monard (1931, 1937a, b), Ahl (1931), Laurent

(1964), and Branch and McCartney (1998). Only PEM

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

16.0

18.0

20.0

22.0

Legend

• Sites 2013

— Major Rivers

Cubango Basin

Cuito Basin

Cuando Basin

18.0

20.0

22.0

csi

16.0

Moxico

Huila

ZAMBIA

100

200 km

Huambo

Cunene

+ NAMIBIA +

Fig. 2. Map of the study area in south-eastern Angola, indicating surveyed sites for April 2013 survey.

and SAIAB material were examined for this study, and

the current taxonomic identity of other historical records

may require verification. Where doubt exists as to their

possible current taxonomic status, this is noted in the

species accounts (below).

Results

Over 63 new locations were sampled in south-east An¬

gola, particularly within the Cubango, Cuito, and Cu¬

ando River catchments. A total of 70 species, includ¬

ing 29 amphibian and 41 reptile species, were recorded

during the three surveys. When collated with historical

literature (see above and Literature Cited) the known

herpetofauna for south-east Angola is increased to 111

species, of which 63% were collected during the surveys

reported here (Tables 2 and 3). Amphibians were better

represented (29 of 34 species, 85.3%) than reptiles (41

of 77 species, 53.2%). Small fossorial reptiles, which are

diverse but difficult to uncover in the Kalahari region, are

under-represented in these collections.

A number of specimens collected were difficult to as¬

sign to currently recognized species. Some comprised

only juveniles or tadpoles, and adult characteristics or

breeding calls were not available for comparison. In oth¬

ers, e.g., Ptychadena cf. mossambica and Ichnotropis

sp., specimens presented a mosaic of characters between

similar species and the present assignment is provisional,

pending ongoing studies on additional material. The spe¬

cies accounts below are arranged alphabetically by fam¬

ily, genus, and species and discuss specific aspects of the

specimens, as well as highlighting their importance or

novelty.

Species Accounts

Amphibia

Arthroleptidae

Leptopelis cf. anchietae (Bocage, 1873)

Anchieta’s Tree Frog

Material: PEM T578 (11); SAIAB 187423 (13). Com¬

ment: Only tadpoles were collected and are tentatively

assigned to Leptopelis anchietae based on tadpole mor¬

phology (Channing et al. 2012).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Table 2. The following are updated species lists for amphibians based on historical records as well as data from the new surveys within the

boundaries of the Cubango, Cuito, and Cuando river basin. Type of record: DR = New drainage record, CR = New country record, V = Voucher,

O = Observation, L = Literature record. Note that the taxonomy has been updated and original species citations may occur under other names.

Species

Cubango River

Cuito River

Cuando River

Reference

ARTHROLEPTIDAE

Leptopelis cf. anchietae (Bocage, 1873)

L,V

Monard 1937a

BREVIC1PITIDAE

Breviceps adspersus Peters, 1882

Bocage 1895; Monard 1937a

BUFONIDAE

Sclerophrys funerea (Bocage, 1866)

V DR

Monard 1937a

Sclerophrys guttural is (Power, 1927)

L, V

Monard 1937a

Sclerophrys lemairii (Boulenger, 1901)

V, DR

V DR

—

Sclerophryspusifla (Mertens, 1937)

—

Sclerophryspoweri (Hewitt, 1935)

—

V, CR

—

HEMISOTIDAE

Hemisus guineensis microps Laurent, 1972

Monard 1937a

HYPEROLIIDAE

Hyperolius angolensis Steindachner, 1867

L, V

Ahl 1931; Monard 1937a

Hyperolius benguellensis (Bocage, 1893)

—

Hyperolius bocagei Steindachner, 1867

—

Monard 1937a

Hyperolius cine re us Monard, 1937

V, DR

—

Hyperolius nasutus Gunther, 1865

L, V

—

Monard 1937a

Kassina kuvangensis (Monard, 1937)

L, V

—

Monard 1933, 1937a

Kassina Senegalensis (Dumeril and Bibron, 1841)

—

PHRYNOBATRACHIDAE

Phiynobatrachus mababiensis FitzSimons, 1932

Phiynobatrachus natal ensis (Smith, 1849)

L, V

—

Monard 1937a

Phiynobatrachus cf. parvulus (Boulenger, 1905)

—

PIPIDAE

Xenopus muelleri (Peters, 1844)

V, CR

Xenopus petersii Bocage, 1895

L, V

Monard 1937a

Xenopus poweri Hewitt 1927

—

V, CR

—

PTYCHADENIDAE

Hildebrandtia ornatissima (Bocage, 1879)

Bocage 1895; Monard 1937a

Ptychadena cf. grandisonae Laurent, 1954

—

Ptychadena guibei Laurent, 1964

—

Ptychadena mascareniensis (Dumeril and Bibron, 1841)

—

Ptychadena cf. mossambica (Peters, 1854)

—

V, CR?

—

Ptychadena oxyrhynchus (Smith, 1849)

L, V

Monard 1937a

Ptychadena subpunctata (Bocage, 1866)

—

V, DR

—

Ptychadena taenioscelis Laurent, 1954

V, DR

V DR

—

Ptychadena uzungwensis (Loveridge, 1932)

—

V DR

—

PYXICEPHALIDAE

Amietia angolensis (Bocage, 1866)

L, V

Bocage 1895; Monard 1937a

Tomopterna cf ciyptotis (Boulenger, 1907)

—

Tomopterna tuberculosa (Boulenger, 1882)

—

Bocage 1895; Monard 1937a

RANIDAE

Amnirana darlingi (Boulenger, 1902)

L, V

Monard 1937a

Total: 34

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

16.0

18.0

20.0

22.0

Legend

• Sites 2015

Major Rivers

Cubango Basin

Cuito Basin

Cuando Basin

18.0

20.0

22.0

csi

16.0

100

200 km

+ NAMIBIA +

Moxico

Huambo

Huila

Cunene

ZAMBIA

Fig. 3. Map of the study area in south-eastern Angola, indicating surveyed sites for May-June 2015 survey.

Bufonidae

Sclerophrys funerea (Bocage, 1866)

Somber Toad

Material: SAIAB 202005 (12°51 , 39.0 ,, S 18°17’

25.02”E). Comment: Monard (1937a) reported two

specimens from the Cubango system around the villages

of Kakindo [= Caiundo] and Mbale. Our specimen rep¬

resents the first record from the Cuito system. We follow

Ohler and Dubois (2016) in using the senior synonym

Sclerophrys Tschudi, 1838 for all African bufonids re¬

cently assigned to Amietophrynus (Frost et al. 2006).

Sclerophrys guttural is (Power, 1927)

Guttural Toad

Material: PEM A10412 (2), 10777 (29), 10817 (30a),

10834 (32), 10835 (32), 10842 (29), 10974 (35), 10976

(34), 10949 (44a), and 10950 (44); SAIAB 101001 (29),

101025 (30a), 190285 (30d), and 190300 (32). Com¬

ment: A common and widespread species recorded from

scattered localities across most of Angola (Laurent 1964;

Poynton and Haacke 1993; Ruas 1996; Ceriaco et al.

2014). Old historical records referring to Bufo regularis

need to be re-examined to confirm their identity and thus

fully understand the distribution of the various toad spe¬

cies in Angola (see Ruas 1996).

Sclerophrys lemairii (Boulenger, 1901)

Lemaire’s Toad

Material: PEM A10413 (7b), 10818-19 (31), 108120-

10829 (30d), and 11527(63); PEM T555 (7a); SAIAB

101019 (30a), 101031 (31), and 101032 (30d). Com¬

ments: This unusual bufonid (Fig. 7c) is adapted for

living in flood plains. Previously reported for the Oka¬

vango Delta (Botswana), adjacent floodplains in Zambia

(Poynton and Broadley 1985; Bittencourt-Silva 2014),

and Democratic Republic of the Congo (Boulenger 1901;

Laurent 1950, 1964; Schmidt and Inger 1959; Poynton

and Broadley 1985). Within Angola, it was previously

known from only five localities further north in Angola

(i.e., Muita-Laurent 1950; Cazombo, Chimboma, Cuilo,

Lake Caiundo - Laurent 1964). The new records collect¬

ed from just north Menongue and around M’Pupa Falls

represent the most southerly Angola records and are the

first for the Angolan Cubango and Cuito river systems.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Fig. 4. Cubango River Basin: A. upper Cacuchi River; B. middle Cacuchi River; C. vegetation around upper Cacuchi River; D.

dambo on upper Cacuchi River; bottom - rocky gorge on the Cuchi River (Site 46).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Fig. 5. Cuito River Basin: A. Cuito River at Cuito-Cuanavale; B. Cuito River south of Menongue; C. lower Cuito River near village

Rito; D. floodplain just north of Menongue; bottom - Source of the Cuito River surround by dry grassland (and base camp), wetlands

around the source lake, and miombo woodlands on higher ground.

They bridge the considerable gaps between previous

Angolan material and those of the Caprivi region (Chan-

ning 2001). Bittencourt-Silva (2014) reported dynamic

dichromatism in this species, and this was also noted

in males collected during the current surveys (Fig. 7d).

However, many features of life history (breeding, call,

and tadpoles) of the species remain unknown, and ad¬

ditional natural history data will be presented elsewhere

(Conradie in prep.).

Sclerophryspusilla (Mertens, 1937)

Eastern Flat-backed Toad

Material: PEM A10283 (6b), 10284 (6b), 10288 (6a),

10289 (6a), 10292 (3), 10322 (12a), 10374 (24), 10418

(20), 10466 (6a), 10467 (22d), 10813-10816 (30a),

11529 (62), 11531 (62), and 11598 (47); SAIAB 101021

(30a), 101006 (32), 101018 (41), 187406 (22d), 188214

(6d), 190247 (29), and 200501 (12a). Comment: Re¬

corded from mostly eastern Angola (Monard 1937a;

Poynton and Haacke 1993; Ruas 1996). Poynton et al.

(2016) revised S. pusilla for southern and eastern popu¬

lations of S. maculata , which is now restricted to West

Africa. For further comment see S. guttural is.

Sclerophrys poweri (Hewitt, 1935)

Power’s Toad

Material: SAIAB 101000 (29). Comment: Only one

specimen was collected on the Angola side of the Cuban¬

go River near Calai. This represents the first record for

Angola, but it is expected to be more widely distributed

in southern and eastern Angola. Many of the earlier re¬

cords of Bufo regularis may be assignable to this species

(see Ruas 1996).

Hyperoliidae

Hyperolius angolensis (Steindachner, 1867)

Angolan Reed Frog

Material: PEM A10417 (16), 10778 (29), 10779 (29),

10808 (30a), 10809 (30a), 10917 (43), 10951 (44a),

10952 (44a), 10973 (35), 11522 (45), and 11532 (62);

PEM T556 (6b), 557 (12a), 562 (28), 563(27), 566 (19),

569 (24), 697 (51), 698 (53b), 699 (54), 700 (63), 701

(57), 709 (55), and 710 (58); SAIAB 101002 (29), 101007

(32), 101015 (41b), 101016 (43), 101022 (30a), 101026

(44a), 101035 (30b), 101036(30b), 101039(35), 187409

(26), 187410 (10), 187414 (7c), 187422 (12a), 187432

(23), 187433 (28), 187437 (22a), 187439 (21), 188064

(19), 188073 (24a), 190262 (30d), 190292-190293 (35),

190397 (30a), and 200492 (12a). Comment: Frost

(2015) considers this species to be part of the unresolved

H. parallelus group which is widespread across Angola

and adjacent countries. There are regional color patterns,

with that in the study area conforming to that of H. ango¬

lensis (Schiotz 1999).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Fig. 6. Cuando River Basin: A. and B. floodplain south-east of Jamba, C. vegetation at site 37, D. pan at site 34.

Hyperolius benguellensis (Bocage, 1893)

Benguela Reed Frog

Material: PEM A10370-10373 (27), 10308 (11),

10398 (22b), 10399 (22b), 10400-1 (21), 10414 (7c),

10415-10416 (7c), 10431-10446 (12a), 10447-52 (5)

10810-10812 (30a), 10919 (43), and 11521 (45); PEM

T560 (6b) and 564 (8); SAIAB 101008 (32), 101023

(30a), 101036 (30d), 101040 (35), 187416 (7b), 187419

(8), 187421 (12a), 187440 (21), 188218 (12a), 190288

(30d), 200495 (12a), and 200497 (5). Comment: Part of

the H. nasutus super group, which has recently been re¬

evaluated (Channing et al. 2013). Only two species, i.e.,

H. nasutus and H. benguellensis , are known from south¬

ern Angola. We assign our specimens to H. benguellensis

based on call differences and the protruding sharp snout.

Genetic studies are ongoing.

Hyperolius cinereus Monard, 1937

Ashy Reed Frog

Material: PEM A10296-9 (5), 101300-10307 (12a),

10314-5 (12a), 1340 (18), 10342-3 (18), and 10350-4

(18); PEM T558 (11), 559 (4), and 565 (8); SAIAB

187417 (8) and 188069 (22d). Comment: Historically

only known from three localities in south-central An¬

gola, i.e., Caluquembe, Bimbe, and Entre Rios (Monard

1937a, Hellmich 1957b). This survey adds seven new lo¬

calities that extend the distribution of the species 350 km

east of the type locality (i.e., Caluquembe) and include

the first records for the Cubango-Okavango river system.

It is more widespread than previously known but remains

endemic to Angola in regions above 1,200 m above sea

level. These collections (Fig. 7a), and additional mate¬

rial collected around Lubango and also Lagoa Carumbo

in north-eastern Angola, led to the re-description of this

poorly known species and the description of a sister tax¬

on, H. raymondi, from northern Angola (Conradie et al.

2013).

Kassina kuvangensis (Monard, 1937)

Kuvango Kassina

Material: PEMT571 (10) and 572 (6b); SAIAB 187418

(8). Comment: Described from Vila-da-Ponte [=Ku-

vango] by Monard (1937a), this continued to be the only

known Angolan locality, although it was subsequently re¬

corded from five localities in adjacent Zambia (Channing

2001). No adults were obtained during our surveys, but

tadpoles (Fig. 7e) collected just east of the type locality

can be assigned to the species (Channing and Broadley

1992) and represent the first Angolan material since the

type description.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Fig. 7. Selective amphibians from south-eastern Angola. A. Hyperolius cinereus, B. Ptychadena cf. mossambica, C. Sclerophrys

lemairii (female), D. Sclerophrys lemairii (male), E. Kassina kuvangensis.

Kassina senegalensis (Dumeril and Bibron, 1841)

Bubbling Kassina

Material: PEM A10863-10873 (39); PEM T573 (24),

574 (6b), and 658 (39); SAIAB 101011 (39), 101014

(40). Comment: Reported from numerous localities

within Angola (Schmidt 1936; Monard 1937a; Laurent

1954, 1964; Poynton and Haacke 1996). The species is

widely distributed in sub-Saharan Africa with geograph¬

ic variation in coloration and morphology. The status of

the various subspecies proposed (e.g., Laurent 1957) re¬

quires a modem appraisal.

Phrynobatrachidae

Phrynobatrachns mababiensis FitzSimons, 1932

Mababe Puddle Frog

Material: PEM A10278 (6b), 10309 (11), 13010

(11), 10316 (12a), 10384-10397 (22d), 10402-3(21),

10426-10430 (19), 10453-10462 (3), 10780-1 (29),

10836-10841 (29), 10849-50 (38), 10892-10899 (39),

10925-10933 (43), 10943-4 (40), and 10953-10957

(44a). SAIAB 101041 (29), 101189 (35), 187403 (18),

187407 (22c), 187412 (6b), 187441 (21), 190301 (39),

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

and 200494 (3). Comment: We assign most of our dwarf

puddle frogs to P. mababiensis , although this remains

a taxonomically challenging group as multiple cryptic

taxa are embedded in the group (Zimkus et al. 2010). Al¬

though Channing (2001) illustrated multiple localities in

Angola, it has previous only been reported from Lagoa

Nutechite (Poynton and Haacke 1993).

Phrynobatrachus natalensis (Smith, 1849)

Natal Puddle Frog

Material: PEM A10286 (6b), 10287 (3), 10312 (12a),

10319-10321 (12a), 10358 (17), 10376-10382 (22d),

10404-10411 (21), 10423-10425 (19), and 10463-10465

(12a); PEM T577 (6b) and 576 (10); SAIAB 187405

(22d), 187408 (21), 187412 (6b), 188065 (19) 188065

(19), and 188215 (12a). Comment: Common and wide¬

spread across whole of Angola (Bocage 1866, 1879,

1895,1897; Boulenger 1905; Ferreira 1906; Parker 1936;

Schmidt 1936; Monard 1937a; Laurent 1950,1954, 1964;

Hellmich 1954; Poynton and Haacke 1993; Ruas 1996).

A taxonomical difficult group in which multiple cryptic

taxa are embedded (Zimkus et al. 2010).

Phrynobatrachus cf. parvulus (Boulenger, 1905)

Dwarf Puddle Frog

Material: PEM A10920-10924 (43). Comment: Report¬

ed from Angola by Boulenger (1905), Monard (1937a),

Parker (1936), Schmidt (1936), and Laurent (1964). It

is easily confused with P. mababiensis , although usually

larger and darker in coloration. We provisionally assign

our specimens to this taxon due to bars on jaw being pale

and not being confluent as in P. mababiensis (Channing

2001 ).

Pipidae

Xenopus muelleri (Peters, 1844)

Tropical Platanna

Material: PEM A10789 (29) and 10969-70 (44a); SA¬

IAB 101030 (44a). Comment: The taxonomic history of

th q Xenopus in Angola is confused. Prior to Bocages’ de¬

scription of X petersii he referred one specimen from

Dombe to X. muelleri (Bocage 1879). Later, in his de¬

scription of X petersii , he referred to the same Dombe

specimen as A petersii (Bocage 1895). By inference he

thus considered only X. petersii to be present in Angola,

although he recognized three varieties (Vars. A-C, see A

poweri for fuller discussion). Specimens collected west

of Calai on the Cubango River and the lower Cuando

River represents the first records of A. muelleri for Ango¬

la. Both A poweri and A muelleri were collected in sym-

patry in the Cuando River. The identifications are sup¬

ported by COl barcoding (Conradie unpublished data).

Xenopus petersii Bocage, 1895

Peter’s Platanna

Material: PEM A10293 (3), 12094-5(5), 10276-7 (6b),

and 11526 (53a); PEM T583 (10) and 584 (24); SAIAB

187443 (21). Comment: Reported from most of Angola

(Bocage 1879, 1895; Boulenger 1905; Ferreria 1906;

Parker 1936; Schmidt 1936; Monard 1937a; Hellmich

1957; Schmidt and Inger 1959; Laurent 1964; Loumont

1981; Poynton and Haacke 1993; Ruas 1996). Common

throughout the north-eastern catchments (Fig. 9b), it was

replaced in the Cuando River by A poweri.

Xenopus poweri Hewitt 1927

Power’s Platanna

Material: PEM Al0937-10942 (43), and 10971 (44a);

SAIAB 101029 (44a). Comment: Based on COl bar¬

coding genes, Xenopus from the Cuando River are ge¬

netically differentiated from those from the Cuito and

Cubango River systems (Conradie unpublished data).

Eastern populations from the Cuando River are there¬

fore provisionally assigned to A poweri Hewitt, 1927,

and western records from the Cuito and Cubango River

to A petersii Bocage, 1895. Although Schmidt and In¬

ger (1959) assigned A poweri to Bocage’s “Var. B” and

restricted A petersii to Bocage “Var. A,” preliminary

genetic findings do not support this, and most Angolan

material should be assigned to A petersii (Furman et al.

2015; Conradie and Evans work in progress). This in¬

cludes specimens from Cubal da Ganda (Laurent 1964)

and Huila (Schmidt and Inger 1959) referred by them to

A poweri. The survey records for A poweri are thus the

first for Angola. An early record from Cazombo (Laurent

1964) also falls within the newly proposed distribution of

A poweri by Furman et al. (2015), but requires verifica¬

tion. When Furman et al. (2015) validated the specific

status of A poweri (previously confused with A laevis),

they referred eastern A petersii material from the Oka¬

vango system in Botswana to the species.

Ptychadenidae

Ptychadena cf. grandisonae Laurent, 1954

Grandison’s Ridged Frog

Material: PEM All525 (53b). Comment: A grass frog

(Fig. 9c) collected on the edge of the upper Longa Riv¬

er floodplain is tentatively assigned to P. grandisonae.

This species has been described from northern Angola

(Muita, Laurent 1954) and reported elsewhere in Angola

(Laurent 1964; Poynton and Haacke 1993; Ruas 1996).

Although members of the genus Ptychadena are notori¬

ously difficult to identify in museum collections, recent

work has shown that species on a regional scale can be

distinguished by quantitative morphometries (Dehling

and Sinsch 2013).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Ptychadena guibei Laurent, 1964

Guibe’s Ridged Frog

Material: PEM A10934-10936 (43); SAIAB 101009

(32) and 190310 (44a). Comment: Previously only re¬

ported from four localities in eastern and northern An¬

gola: Muita (Laurent 1950, 1954), Cazombo (Laurent

1964), Dundo (Laurent 1964), and Cangandala (Ceriaco

et al. 2016). Common and widespread in adjacent Zam¬

bia and into the panhandle of the Okavango Delta (Chan-

ning 2001). The new material extends the distribution

into extreme southeast Angola, and about 150 km up the

lower Cuando River.

Ptychadena mascareniensis (Dumeril and Bibron, 1841)

Mascarene Ridged Frog

Material: PEMA10782-10785 (29), 10843-10848 (29),

10877 (39), 10945-6 (41a), 10958 (44a), and 11535-6

(63); SAIAB 101003 (29) and 190304 (43). Comment:

Recent mitochondrial DNA analysis suggests that P.

mascareniensis comprises a number of separate species

and that the name P. mascareniensis should be restricted

to the Madagascar, Seychelles and the Mascarene Islands

(Vences et al. 2004). The only name available for clades

from mainland Africa is P. nilotica for the Nile River

system (Dehling and Sinsch 2013). Further taxonomical

work is underway to determine the status of this species

in Angola (Ernst work in progress). Widespread in An¬

gola (Bocage 1867; Boulenger 1905; Ferreria 1906; Mo-

nard 1937a; Schmidt and Inger 1959; Ruas 1996).

Ptychadena cf. mossambica (Peters, 1854)

Broad-banded Ridged Frog

Material: PEM A10850-10862 (38), 10874-10876 (39),

10878-10880 (39), and 10918 (43); SAIAB 101010 (38)

and 101012 (39). Comment: A series of grass frogs (Fig.

7b) collected from the lower Cuando River near the vil¬

lage of Jamba are provisionally assigned to the Ptychade¬

na mossambica complex. Channing (1993) described

Ptychadena mapacha from Mapacha in the eastern

Caprivi, Namibia, close to our new collection, noting that

it was superficially similar to P. mossambica. Morpho¬

logically they differ in having a continuous paravertebral

fold from head to midbody in P. mossambica , that is in¬

terrupted in P. mapacha. Unfortunately, the current series

was collected during the dry season and no vocalization

was obtained for comparison with that recorded for P.

mapacha (Channing 1993). Genetic analysis is underway

on the P. mossambica complex to detennine the specific

identity of the new collection. Neither P. mossambica or

P. mapacha, which are both known from the Caprivi area

of Namibia, have been recorded from Angola.

Ptychadena oxyrhynchus (Smith, 1849)

Sharp-nosed Ridged Frog

Material: PEM A10282 (6b), 10881-10891 (39), 10947

(41a), and 10959-10963 (44a); SAIAB 101013 (39),

188217, 190306 (43), and 190309 (44a). Comment:

Common and widespread in Angola and adjacent coun¬

tries (Bocage 1866, 1895, 1897; Boulenger 1905; Fer¬

reira 1906; Parker 1936; Schmidt 1936; Monard 1937a;

Laurent 1950, 1954; Hellmich 1957).

Ptychadena subpnnctata (Bocage, 1866)

Spotted Ridged Frog

Material: PEM A10359-10368 (27), 10794-10807

(30a), 10948 (41a), 10900-10916 (43), 10964-10967

(44a), 11528 (63), and 11614-5 (56); SAIAB 101005

(33a), 101024 (30a), 101028 (43), 101033 (6b), 190260

(30f), 190307 (44a), 190350 (35), and 200502 (27).

Comment: Previously known from central and north¬

ern Angola (Bocage 1866; Schmidt 1936; Mertens 1938;

Laurent 1964; Ruas 1996), the new records fill the large

gap in the species’ range in southeast Angola.

Ptychadena taenioscelis Laurent, 1954

Small Ridged Frog

Material: PEM A10290-1 (3), 10279-81 (6b), 10285

(6b), 10419-22 (19), 10830-10833 (30d), 10972 (44b),

10975 (35), 11530 11523—4 (43b), (62), and 11610-1

(47); PEM T580-1 (10); SAIAB 101017 (43), 101034

(30d), and 188217 (6b). Comment: The species has been

previously recorded from northern (Laurent 1964) and

central Angola (Poynton and Haacke 1993), and the new

material extends its range into the grasslands of south¬

eastern Angola (Fig. 9a).

Ptychadena uzungwensis (Loveridge, 1932)

Uzungwe Ridged Frog

Material: PEM A10369 (27). Comment: In Angola it

was previously restricted to the east and central region

(Loveridge 1932; Monard 1937a; Laurent 1954, 1964;

Poynton and Haacke 1993; Ruas 1996), but the current

specimen extends the range southeast into Kuando-

Kubango Province.

Pyxicephalidae

Amietia angolensis (Bocage, 1866)

Angolan River Frog

Material: PEMA10311 (11), 10313 (12a), 10317-10318

(12a), and 10375 (22d); PEM T552 (11) and 553 (4); SA-

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

IAB 187112 (12a), 187402 (18), 187411 (14), 187413

(3), 187424 (13), 200499 (18), 200500 (12a), and

200503 (13). Comment: Common and widespread in

Angola (Bocage 1866, 1895; Boulenger 1905; Ferreira

1906; Parker 1936; Schmidt 1936; Mertens 1938; Mo-

nard 1937a; Laurent 1950, 1954, 1964; Hellmich 1954;

Poynton and Haacke 1993; Ruas 1996). A recent revision

(Channing and Baptista 2013) restricted A. angolensis to

Angola, albeit that few Angolan samples were available

for analysis. Subsequently, Larson et al. (2016) identi¬

fied several well-supported cryptic lineages of river frogs

previously assigned to Amietia angolensis in the Alber-

tine Rift region. It is, therefore, possible that cryptic taxa

of A. angolensis also occur in Angola, and that A. ango¬

lensis may also extend into adjacent western Zambia.

Tomopterna cf. cryptotis (Boulenger, 1907)

Cryptic Sand Frog

Material: PEM A10786-8 (29), 10790-3 (29) and

10968 (44a); SAIAB 101004 (29) and 187435 (22c).

Comment: Recorded from only a handful of localities

in Angola (Boulenger 1907; Poynton and Haacke 1993).

Species delineation in the genus is problematic and often

dependent upon vocalization and chromosome number.

We assign our material conservatively as specimens may

represent either T. tandyi, T. cryptotis or even further

cryptic diversity.

Ranidae

Amnirana darlingi (Boulenger, 1902)

Darling’s White-lipped Frog

Material: PEM 1711 (55); SAIAB 187436 (22a). Com¬

ment: Only tadpoles were obtained. Previously recorded

from central and southern Angola (Monard 1937a; Lau¬

rent 1964, Schmidt 1936; Schmidt and Inger 1959; Ruas

1996). Oliver et al. (2015) showed that African species

recently assigned to Hylarana are best placed in the ge¬

nus Amnirana.

Reptilia

Squamata

Serpentes

Colubridae

Crotaphopeltis hotamboeia (Laurenti, 1768)

White-lipped Snake

Material: PEM R20018 (12b); ANG (no number, 6a).

Comment: Common and widespread species in Angola

(Bocage 1895; Boulenger 1905; Ferreira 1906; Branch

and McCartney 1992; Laurent 1950, 1964; Parker 1936).

This species is widespread in sub-Saharan Africa. A phy¬

logenetic study is currently underway to investigation

different populations (Tolley pers. comm.).

Philothamnus hoplogaster (Gunther, 1863)

Green Water Snake

Material: photograph record (James Kydd: 14°08’56.0”S

18°48’36.0”E). Comment: This is the first record for the

Cuito River system. Only recorded from a handful of

other locations in Angola (Bocage 1887; Monard 1937b;

Laurent 1964; Thys van den Audenaerde 1967).

Philothamnus ornatus Bocage, 1872

Ornate Green Snake

Material: PEM R20013 (24). Comment: Philothamnus

ornata was described by Bocage (1872) from two speci¬

mens collected at Huila. Although subsequently recorded

from Zambia and Zimbabwe (Broadley et al. 2003), it

remained known in Angola from only few additional

collections, i.e., Bela Vista (Hellmich 1957), Benguela

(Boulenger 1905), Bie (Boulenger 1905), Caconda (Bo¬

cage 1895), Caluquembe (Monard 1937b), Cutatu (Mo¬

nard 1937b). Chimporo (Boulenger 1905), Cunene River

(Bocage 1895), and Huambo (Bogert 1940). Our collec¬

tion fills in the gap between the records from western

Angola and the Zambian and Zimbabwean populations.

Thelotornis capensis oatesi (Gunther, 1881)

Oates’ Twig Snake

Material: PEM R21484 (52). Comment: This subspe¬

cies has a large range from southern Angola to Malawi

and western Mozambique (Broadley and Wallach 2002),

but with relatively few Angolan records (Bocage 1895;

Parker 1936; Monard 1937; Bogert 1940; Laurent 1954,

1964; Thys van den Audenaerde 1967). When reviewing

the genus, Broadley (1979) noted few specimens from

the southern and eastern regions of the country, while

Laurent recorded sympatry between T. kirklandi and T.

c. oatseii at Dundo in northeast Angola. The new record

is only the second for the south-eastern Angola, follow¬

ing Monard’s (1937b) record from Vila-da-Ponte [=Cu-

vangu]. The present specimen helps fill the large gap in

records for the south-east of Angola. Although Broadley

(2001) distinguished T. c. oatesi by coloration of the top

of the head and ventral number (>160) these features are

variable. The status of T. c. oatesi as a valid taxon was

not resolved by a morphological and genetic analysis of

the Dispholidini (Eimermacher 2012) and remains prob¬

lematic.

Natricidae

Limnophis bangweolicus (Mertens, 1936)

Bangweola Swamp Snake

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Fig. 8. A selection of reptiles from south-eastern Angola. A. Natriciteres olivacea ; B. Linmophis bang\\>eolicus\ C. Tetradactylus

eJJenbergeri ; D. Zygaspis quadrifrons ; E. Typhlacontias rohani f; F. Acontias kgalagadi kgalagadr, G. Lubuya ivensii ; H. Pelusios

bechuanicus.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Fig. 9. Selective amphibians and reptiles from south-eastern Angola. A. Ptychadena taeniocelis; B.Xenopus peter sir, C. Ptychadena

cf. grandisonae: D. Ichnotropis spp; E. Causus cf. rasmusseni, F. Causus cf. rasmusseni ; G. Boaedon cf. angolensis; H. Trachylepis

cf. spilogaster.

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October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Material: PEM R20489 (44a). Comment: A specimen

(Fig. 8b) collected in the floodplain of the lower Cuan-

do River represents the second record for Angola. The

previous record was 700 km north at Calundo, Moxico

Province (Laurent 1964). The sister species Limnophis

bicolor is more widespread in western, central, and

northern Angola.

Natriciteres olivacea (Peters, 1854)

Olive Marsh Snake

Material: PEM R20501 (44a). Comment: A specimen

(Fig. 8a) collected in the floodplains of the lower Cuan-

do River represents a southern record of this species for

Angola, the other records occurring much further north,

i.e., Dondo (Hellmich (1957), Dundo and Muita (Laurent

1950, 1954), and Pungo Andongo (Bocage 1895; Bou-

lenger 1905). Other records from Bela Vista (Hellmich

1956) and Malanje (Bocage 1895) have subsequently

been referred to N. bipostocularis (Broadley 1966).

There is little variation in scalation among Natriciteres

species, and also no molecular phylogeny for the genus.

However, the Cuando specimen, despite its relative prox¬

imity to the Angolan N. bipostocularis records, still pos¬

sesses three postoculars and is conservatively referred to

N. olivacea.

Elapidae

Dendroaspis polylepis polylepis Gunther, 1864

Black Mamba

Material: Observation only (37). Comment: One speci¬

men was observed crossing the road and disappearing

into a tree. Only a handful of records exist for Angola

(Schmidt 1933; Monard 1937b; Bogert 1940; Baynham

2010), but it is believed to be more widespread in the

southern and central regions of Angola (Branch et al. in

prep.).

Naja ( Afronaja ) mossambica Peters, 1854

Mozambique Spitting Cobra

Material: PEM R20499 (30a) and 20500 (44b). Com¬

ment: The new records represent the second (M’Pupa

Falls) and third (near village Sashae) for Naja mossam¬

bica in Angola. The only other published Angolan record

(Maconjo, Broadley 1974) occurs in south-west Angola,

although other western records are known (Branch et al.

in prep.). This species is expected to be more widely dis¬

tributed in southern Angola.

Lamprophiidae

Boaedon cf. angolensis Bocage, 1895

Angola House Snake

Material: PEM R21846 (50). Comment: This unusual

pale yellow-green specimen (Fig. 9g) is referable to the

Boaedon lineatus-fuliginosus complex. A specimen of

similar coloration was photographed in the upper Cuito

River (James Kydd: 13 o 09’05.0”S 18°28’50.0 ,, E). Kel¬

ly et al. (2011) demonstrated complex cryptic diversity

with numerous deep lineages in house snakes of the B.

lineatus-fuliginosus complex. On morphology, Hughes

(1997) referred specimens from East and South Africa to

B. capensis Dumeril and Bibron, 1854, and this has been

adopted in the most subsequent literature (see Uetz 2016).

However, southern African specimens fall into a number

of well-defined clades (Kelly et al. 2011) that cannot all

be accommodated under B. capensis. The name Boaedon

lineatus var. angolensis Bocage, 1895 has not been con¬

sidered for western populations. Laurent (1956) noted

that populations from Angola to Katanga could be dis¬

tinguished morphologically, but took no taxonomic ac¬

tion and the availability of Bocage’s angolensis was not

discussed. Unfortunately, neither a type or type locality

was nominated for the taxon, and Bocage’s material was

also lost in the fire that destroyed the museum in Lisbon.

Psammophis mossambicus Peters, 1882

Olive Grass Snake

Material: PEM R20024 (16b) and 21827 (48); ANG

(23a). Comment: Species boundaries in the Psam¬

mophis sibilans complex remain problematic (Kelly et

al. 2008). Our specimens accord with P. mossambicus

(Broadley 2002). Historical Angolan records assigned to

P. brevirostris , P. leopardinus , P. mossambicus , P. phil-

lipsi , and P. sibilans need to be carefully re-examined,

and genetically assessed.

Psammophis subtaeniatus Peters, 1882

Yellow-bellied Sand Snake

Material: PEM R20506 (44a). Comment: Recorded by

Bocage (1896) and Monard (1937b). Broadley (2002)

mapped the known Angola material which clusters in the

south-west, as far north as Benguela. The current speci¬

men, the first from the Cuito drainage, fills the large gap

between populations in Caprivi region of Namibia and

Zimbabwe.

Psammophylax acutus (Gunther, 1888)

Sharp-nosed Skaapsteker

Material: PEM R20006 (23) and 21485 (51). Com¬

ment: Common and widespread species in Angola (Bo¬

cage 1895; Bogert 1940; Ferreira 1906; Laurent 1964

and Monard 1937b). The two from Longa River are the

first for south-east Angola. Transferred from Rhamphio-

phis by Kelly et al. (2008).

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October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Leptotyphlopidae

Leptotyphlops scutifrons complex (Peters, 1854)

Peter’s Thread Snake

Material: PEM R20020 (13). Comment: Many earlier

records from Angola attributed to L. scutifrons (e.g., Bo-

cage 1895, 186; Boulenger 1905; Werner 1917) were

referred to L. (= Namibiana) latifrons (Broadley and

Broadley 1999). Monard (1937b) referred seven Ango¬

lan specimens, including one from Cuvangu, to this spe¬

cies. These were not discussed by Broadley and Broadley

(1999), and although referred to L. scutifrons by Wallach

et al. (2014) their identity requires confirmation. Our

specimen is the second confirmed locality from Angola,

following Broadley and Broadley’s (1999) record of new

material from Chitau. We stress, however, that the sta¬

tus of the limited Angolan material, which involves large

disjunctions between the known records and from the

type locality (Tete, Mozambique), awaits further study,

especially as the L. scutifrons complex has been shown to

comprise numerous deep lineages that represent cryptic

species (Adalsteinsson et al. 2009).

Pythonidae

Python natalensis Smith, 1840

Southern Rock Python

Material: SARCA 153808 (1), 153811 (30a). Com¬

ment: Large pythons are known from multiple localities

in Angola (Bocage 1895; Monard 1937b; Bogert 1940;

Laurent 1954, 1964; Thys van den Audenaerde 1967;

Ceriaco et al. 2014, 2016). hollowing the revision of the

Python sebae complex, Broadley (1984) revived P. na¬

talensis for pythons from East and southern Africa. How¬

ever, scattered populations of P. sebae have subsequently

been recorded in East Africa (Spawls et al. 2002), and the

status of pythons in Angola, particularly in the northern

Congo drainage, should be re-evaluated. Python natalen¬

sis is known from adjacent Namibia and Zambia, and

pythons from southern Angola, including our material,

are referable to P. natalensis based on head coloration

and scalation.

Viperidae

Bitis arietans (Merrem, 1820)

Puff Adder

Material: PEM R20021 (9) and 21493 (61); road kill,

not kept (38). Site: 9, 38, 61. Comment: Widespread

species recorded throughout Angola, but recorded only

three times during the current surveys.

Causus cf. rasmusseni Broadley, 2014

Rasmussen’s Night Adder

Material: PEM R21488 (57). Comment: An adult fe¬

male from riverine habitat on the Rio Longa, and with

a diffuse color pattern (Fig. 9e, f) and only 132 ventrals.

It conforms in coloration and low ventral count with the

newly-described C. rasmusseni (Broadley 2014) but has

a lower subcaudal count (31) than that of the only female

(39) in the type series from north-west Zambia. As noted

by Rasmussen (2005), the status of night adders in Ango¬

la has been confused with many “ rhombeatus ” of earlier

authors now assigned to other species, e.g., C. maculatus

and C. bilineatus (Piri Dembos; Hellmich 1957a) and C.

resimus (Libolo-Luali, Hellmich 1957b). Ventral counts

are typically higher (143-145) for blotched females of

C. rhombeatus from the eastern half of Angola (Laurent

1964). A molecular phylogeny of Causus (Tolley et al. in

prep.) shows that the Rio Longa specimen is only weakly

differentiated from South African C. rhombeatus. How¬

ever, topotypic material is not available for C. rasmusse¬

ni and its taxonomic status thus remains equivocal. If the

specific validity of Rasmussen’s night adder is confirmed

the Rio Longa specimens would be the first Angolan re¬

cord for the species, and a southern range extension of

approximately 740 km from the Zambian series (Broad¬

ley 2014). If, however, it is only a plain color morph of

C. rhombeatus it would be the first record for south-east

Angola.

Sauria

Agamidae

Agama aculeata Merrem, 1820

Ground Agama

Material: PEM R20017 (15). Comment: Recorded

from central and southern Angola (Bocage 1895; Bou¬

lenger 1905; Monard 1937b; Laurent 1964), but is poorly

documented from the south-east.

Amphisbaenidae

Zygaspis quadrifrons (Peters, 1862)

Kalahari Round-headed Worm Lizard

Material: PEM R20524 (41b), 20525 (44b), 21491

(59), and 21492 (60). Comment: Along with southern

Africa, Angola has a very rich amphisbaenian fauna

and 11 species are known, with representatives of three

genera ( Dalophia , Monopeltis , and Zygaspis). Zygaspis

quadrifrons was recorded from Alto Chipata, north-west

Angola, by Laurent (1964), but this has subsequently

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

been referred to the new species Z nigi*a Broadley and

Gans, 1969. The only previous Angolan records are from

Kakindo [= Caiundo], Kuando-Kubango Province and

Chimporo [=Techimpolo], Cunene Province by Monard

(1937b). The new collections (Fig. 8d) from the Cuito

and lower Cuando River represent the first records for

both river systems and double the total number of Ango¬

lan locations.

Chamaeleonidae

Chamaeleo dilepis Leach, 1819

Flap-necked Chameleon

Material: SARCA153813 (30a). Comment: A common

and widespread species in Angola (Bocage 1895; Bou-

lenger 1905; Schmidt 1933; Parker 1936; Monard 1937b;

Laurent 1950,1954, 1964; Ceriaco et al. 2016). Although

Tilbury’s (2010) recent map of the species shows it lim¬

ited in Angola to the western and central regions, it is

probably more widespread. The gap in south-east Angola

is partly filled by the lower Cuito specimen.

Gekkonidae

Chondrodactylus cf. pulitzerae (Schmidt, 1933)

Angolan Thick-toed Gecko

Material: PEM R20477-20479 (30a), 21494-7 (62).

Comment: These geckos are widespread in the south¬

ern and western parts of Angola, reaching as far north as

Capanda Dam (Ceriaco et al. 2014). The Chondrodacty¬

lus turneri complex, of which pulitzerae was recently a

synonym, has been shown to have a number of distinct

lineages (Ceriaco et al. 2014), with the revival of C. pu¬

litzerae for most Angolan material. Species allocation of

our south-east Angolan material is provisional pending

molecular analysis.

Lygodactylus bradfieldi Hewitt, 1932

Bradfield’s Dwarf Day Gecko

Material: PEM R20492 (32), and 20490-1 (35). Com¬

ment: Pasteur (1965) revived Hewitt’s Namibian species

after FitzSimons (1943) had treated it as a subspecies

of L. capensis, and Loveridge (1944) had even placed

it in the synonymy of the same species. Jacobsen (2011)

affirmed features that distinguished the species and

confirmed its presence in sympatry with L. capensis in

western Limpopo Province, South Africa. Our material

confonns to the features highlighted by Jacobsen (2011),

and we therefore consider the survey records to represent

the first for the Cuito drainage, and possibly the first con¬

firmed records for Angola. Pasteur’s (1965) map did not

show L. bradfieldi extending into southern Angola, and

although he plotted six localities for L. capensis in south¬

west Angola, presumably based on records in Bocage

(1895) and Monard (1937b), he did not discuss voucher

material for any of his localities. Our specimens fill the

large gap for the L. capensis complex in south-east An¬

gola. Cryptic diversity within dwarf geckos is now well

established (e.g., Travers et al. 2014), and the status of

the isolated population of L. capensis in southwest An¬

gola (see above) and its relationship to L. bradfieldi in¬

vites further study.

Gerrhosauridae

Gerrhosaurus cf. nigrolineatus Hallowell, 1854

Black-lined Plated Lizard

Material: PEM R20004 (6a), 20481-2 (30a), 20483^1

(35), and 21826 (46). Comment: Bates et al. (2013) re¬

vived G. intermedins for eastern populations of G. ni¬

grolineatus, raised G. bulsii to specific status, and noted

the unresolved status of G. multilineatus in north-central

Angola. The status of these populations and others from

western Angola and relationships within the G. nigrolin¬

eatus complex in Angola is currently under investigation

(M. Bates, pers. comm.).

Tetradactylus ellenbergeri (Angel, 1922)

Ellenberger’s Seps

Material: PEM R20010 (23). Comment: A specimen

(Fig. 8c) collected along a dammed section of the Luass-

ingua River, represents only the fifth and most southern

record of this species for Angola. Wagner et al. (2012)

only listed one record from Angola, overlooking other

historical records, e.g., Caiundo as T. e. ellenbergeri (Lau¬

rent 1964), Dundo as T. e. boufengeri (Laurent 1964),

Cuando as T. africanus (Bocage 1895) and Tyiumbwe

as T. lundensis (Monard 1937b). Tetradactylus africanus

is now restricted to coastal regions of KwaZulu-Natal,

South Africa, and adjacent southern Mozambique, while

Laurent (1964) treated T. lundensis as a synonym of T.

ellenbergeri boulengeri. This is the first record for south¬

east Angola.

Lacertidae

Heliobolus lugubris (Smith, 1838)

Bushveld Lizard

Material: PEM R21500 (63). Comment: This represents

the first record for the south-eastern part of the country.

Previously only recorded in Angola from the south-west

(Monard 1937b).

Ichnotropis spp.

Rough-scaled Lizard

Material: PEM R20008-9 (25), 20486-8 (30b), 21490

(59), and 21843-5 (55). Comment: Branch and McCar-

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

thy (1992) referred an adult male (SVL 45 mm) from

Cuito-Cuanavale as the first record of I. capensis from

Angola. Our additional material comprises juveniles and

small adults (Fig. 9d, maximum SVL = 45 mm). Broad-

ley (1967) described I. grandiceps based on a very small

series {n = 3) from the Caprivi Strip. He differentiated

it from sympatric I. capensis based on higher midbody

scale counts (44-47), a broader head, five upper labials

anterior to the subocular, and an occipital scale that did

not protrude past the parietals. In our series the mid-body

scale count is low (36^12), upper labials are mostly four

(seven out of the 12 examined), and the occipital scale

protrudes past the parietals. The specimens thus display

a mixture of features intermediate between the two spe¬

cies. Although an adult male with bright yellow flanks

(indicative of I. capensis male breeding coloration) was

observed in the same area as juveniles (Conradie pers.

obs, it was not collected and scalation could not be de¬

termined). The genus Ichnotropis includes a number of

poorly-known northern species, and the lack of recent

material of taxa such as I. grandiceps, I. tanganicana,

I. microlepidota, I. bivittata pallida , etc., has precluded

a modern revision. These difficulties are compounded

by the annual reproductive strategies of some species

(Broadley 1979) that make collection of series of adults

and juveniles difficult. A revision of Angolan species,

and the description of new taxa, is in preparation.

Meroles squamulosus (Peters, 1854)

Rough-scaled Desert Lizard

Material: PEM R20493^1 (30e) and 20495 (35). Com¬

ment: Previously recorded from Angola only from

Kapelongo [=Capelongo], Huila Province (Monard

1937b), and Pereira de Ega [=Ondijiva], Current material

includes the third and fourth records for Angola. Previ¬

ously generically assigned to Ichnotropis , but recently

shown to be nested within Meroles (Edwards et al. 2013;

Engleder et al. 2013).

Scincidae

Acontias kgalagadi kgalagadi Lamb, Biswas and Bauer,

2010

Kalahari Legless Skink

Material: PEM R20474 (44a). Comment: Acontias k,

kgalagadi (previously Typhlosaurus lineatus lineatus

- see Lamb et al. 2010) occurs mostly in north-eastern

South Africa, extending to northern Namibia and Bo¬

tswana. Although Broadley (1968) reported a single re¬

cord of A.jappi from Gago Coutinho district in the south¬

east of Angola, there are no records of A. k. kgalagadi

from the country. This is also the first record (Fig. 8f) of

this species in Angola and north of the Okavango River

(Conradie and Bourquin 2013).

Lubuya ivensii (Bocage, 1879)

Iven’s Meadow Skink

Material: PEM R20005 (7c). Comment: A rarely sam¬

pled semi-aquatic skink, known from eight localities in

Angola, single localities in adjacent Zambia and Demo¬

cratic Republic of the Congo (Branch and Haagner 1993;

Wagner et al. 2012), and a recent isolated record from

Lavushi Manda National Park, Zambia (Broadley and

Willems 2015). It was previously listed as a near-endem¬

ic species to Angola, and can be considered as such pend¬

ing genetic assessment of the Lavushi Manda population.

The new record (Fig. 8g) from the floodplains just north

of Menongue is the first record for the Cubango-Okavan-

go system (previously only known to occur in the head¬

waters of the Zambezi, Kwanza, and Congo drainage

systems). Horton (1972) placed Mabnya ivensii into a

monotypic genus, Lubuya , that was recently revived due

to its basal position in a phylogeny of mabuyine skinks

(Metallinou et al. 2016) and it is unique matrotrophic re¬

production.

Mochlus sundevalli (Smith, 1849)

SundevalFs Writhing Skink

Material: PEM R20496 (39), 20497 (42), and 20498

(44a). Comment: Recorded by Bocage (1895), Laurent

(1964) and Monard 1937b. Previously placed in the gen¬

era Lygosoma Boulenger, 1895 and Riopa Smith, 1937,

both of which are now restricted to West Africa.

Panaspis maculicollis Jacobsen and Broadley, 2000.

Speckle-lipped Snake-eyed Skink

Material: PEM R20007 (43). Comment: Only a few re¬

cords of Panaspis exist for Angola (P. breviceps - Parker

1936; P. cabindae - Parker 1936; Bocage 1895; Ferreira

1906). A recent phylogeny of snake-eyed skinks (Medina

et al. 2016) placed Afroablepharus in the synonymy of

Panaspis and showed deep divergences for the basal taxa

P. breviceps and P. togoensis. The P. wahlbergi-mac-

ulicollis complex was also shown to contain numerous

cryptic taxa. When describing P. maculicollis Jacobsen

and Broadley (2000) recorded no specimens for Angola,

and the Cuando specimen of confirmed genetic mono-

phyly, represents the first for the country.

Trachylepis cf. spilogaster (Peters, 1882)

Kalahari Tree Skink

Material: PEM R20007 (20), 20016 (25), 20019 (12b),

20517 (30a), 20518 (35), 20519 (32), 21483 (49),

21486-7 (57), 21498-9 (62), 21828-21840 (47), and

21842 (6a). Comment: This form of the T. striata com¬

plex was elevated to specific status by Broadley (1969).

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

First recorded from southern Angola by Laurent (1964),

based on a single male from Serra do Moco, Luimbale,

Huambo. He noted the great disjunction between this

specimen and the species’ known distribution and cau¬

tioned that further investigation was necessary. His as¬

signment to spilogaster appeared to be influenced by the

tricarinate midbody scale count (34), supranasal and pre¬

frontal condition, and speckled ventrum. The new ma¬

terial basically conforms to the descriptions in Laurent

(1964) and Broadley (1969, 1977, 2000), in having su-

pranasals in broad contact and 35-37 tricarinate midbody

scale rows (31—40, mean 34.7; Broadley 1977). Howev¬

er, the prefrontal condition varies from broad contact to

widely separated (usually separated in spilogaster ), the

ventrum is never speckled although the throat is in adult

males, and adult males usually have a red-brown mid¬

dorsal band (Fig. 9h) that is not recorded in other popula¬

tions (Laurent 1964; Broadley 1969, 1977, 2000). The

Cuito population also has different habits to typical spi¬

logaster, being rarely found on trees although common

on the ground, in rock piles, and on village houses. Its

relationship with T. angolensis (Monard 1937b), and in¬

deed that taxon’s status with respect to earlier names (Eu-

prepes angolensis Bocage, 1872 and Sepsina angolensis

Bocage, 1866), remains problematic (see discussion in

Boulenger, 1887). The present series is the subject of a

more detailed morphological and genetic assessment.

Trachylepis varia (Peters, 1867)

Variable Skink

Material: PEM R20507 (30c), 20508-9 (35), 20510

(32), 20511-2 (37), 20513 (38), 20514 (39), 20515 (42),

20516 (44a), 21489 (59), 21825 (46), and 21841 (47).

Comment: Most common and widespread species of

skink in Angola, with a massive range from Ethiopia to

South Africa. The species is known to have both genetic

and reproductive diversity, and probably comprises a

complex of numerous cryptic taxa.

Trachylepis wahlbergi (Peters, 1869)

Wahlberg’s Skink

Material: PEM R20520-1 (43). Comment: The Tra¬

chylepis striata complex is widespread in southern and

eastern Africa, but relationships and taxonomic status of

the various subspecies of T. striata , elevated to specific

status by Broadley (2000), remain problematic. Subse¬

quently, Castiglia et al. (2006) showed that T. striata and

T. wahlbergii share the same karyotype and no significant

differentiation in 16S mtDNA gene. They argued that this

did not support Broadley’s (2000) action in treating both

taxa as distinct species. We retain the current taxonomic

arrangement but note that fuller studies on the T. striata

complex are required. First recorded from southeast An¬

gola from near Mavinga (Branch and McCarthy 1992),

but also known from numerous other localities in south¬

west Angola.

Typhlacontias rohani Angel, 1923

Rohan’s Legless Skink

Material: PEM R20522-3 (38), and 20526 (44a).

Comment: Angel (1923) described this species from

Lwankundu River, a western tributary of the Cuando

River. The only other record for Angola is from Chim-

poro, Cunene Province (Monard 1937b). The new col¬

lections (Fig. 8e) from the lower Cuando River represent

the third and fourth records for Angola. Haacke (1997)

reported the species to be common in north-eastern Na¬

mibia, the Caprivi Strip [=Zambezi Region], northern

Botswana and western Zimbabwe.

Varanidae

Varanns niloticus (Linnaeus, 1766)

Water Monitor

Material: observations only (3, 32, 43). Comment:

Common and widespread along major rivers in Angola.

Although only a single Angolan specimen was included

in a phylogeographic review of the V niloticus complex

(Dowell et al. 2016), it is likely that all Angolan water

monitors are preferable to V niloticus (of which V or-

natus is now a junior synonym) and not West African V

stellatus (Daudin, 1802), which was recently revalidated

(Dowell et al. 2016).

Crocodylia

Crocodylidae

Crocodylus niloticus Laurenti, 1768

Nile Crocodile

Material: SARCA 153812 (29) and other observational

records (35, 43, 44a, 45). Comment: During the 2015

Okavango Expedition and Expedition Okavango evi¬

dence was found of crocodiles breeding in the Cuito Riv¬

er (Shacks 2015), and adults and juveniles were observed

at the various sites.

Testudines

Pelomedusidae

Pelomedusa subrufa (Bonnaterre, 1789)

Marsh Terrapin

Material: PEM R20502-3 (40). Comment: Although

the previously monotypic Pelomedusa has been shown

to have high levels of cryptic diversity, with the descrip-

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

tion of six new species (Petzold et al. 2014), Angolan

populations remain assigned to P. subrufa.

Pelusios bechuanicus FitzSimons, 1932

Botswanan Hinged Terrapin

Material: PEM R20504 (44a). Comment: A localized

species associated with the Okavango Delta in Botswana.

A specimen (Fig. 8h) was caught in pushing flood waters

of the lower Cuando River. This new record represents

the fourth record for Angola. The other records for An¬

gola are from Chonga River, Moxico Province (Laurent

1964), Chimporo Marsh in the Cuanda drainage (Monard

1931) and Vila da Ponte (= Kubango) (Monard 1937b, as

P. subniger).

Testudinidae

Kinixys cf. belliana Gray, 1863

Bell’s Hinged Tortoise

Material: SARCA 153810 (13). Comment: The genus

Kinixys has been recently revised (Kindler et al. 2012),

with K. belliana now restricted to coastal Kenya and So¬

malia, and extending through Burundi to Angola. Kinix¬

ys spekii has previously been considered to have a wide

distribution in southeast Africa and to range into much

of Angola. Ceriaco et al. (2014) assign Capanda mate¬

rial to K spekii , but this was not confirmed by molecu¬

lar analysis. The only three Angola specimens sampled

in the recent revision of Kinixys (Kindler et al. 2012),

from Cuemba and Kuito, Bie Province, were all shown

to group with East African K belliana. The current speci¬

men is provisionally assigned to K belliana , pending

further molecular analysis of Angolan material and we

caution that the presence of K spekii in Angola requires

confirmation.

Stigniochelyspardalis (Bell, 1828)

Leopard Tortoise

Material: SARCA 153814 (44a). Comment: Only

known from a handful records from southwest Angola

and seems to be rare elsewhere in Angola (Bocage 1895;

Mertens 1937; Hellmich 1957; Monard 1937b).

Discussion

The surveys filled large gaps in the distributions of many

species (see Figures 1^1), and included 13 new amphib¬

ian catchment records for the various rivers (three for the

Cubango, seven for the Cuito, and three for the Cuando).

Four amphibian species, i.e., Sclerophrys poweri, Xeno-

pus muelleri, X. poweri , and Ptychadena cf. mossambica ,

and three reptile species, i.e., Causus cf. rasmusseni (if

valid), Acontias kgalagadi kgalagadi (see Conradie and

Bourquin 2013), and Panaspis maculicollis , were add¬

ed to the herpetofaunal list of taxa known to occur in

Angolan. A further 15 reptile catchment records for the

various rivers (two for the Cubango, nine for the Cuito,

and four for the Cuando) were also obtained. Amphibian

diversity comprises 10 families and 13 genera (Table 2),

while there are 19 reptile families and 52 genera (Table

3). Fuller discussion of the biogeographic insight and re¬

lationships of the present findings are deferred pending

analysis and further research on additional collections

from the source lakes of the Cuito, Cuanavale, and adja¬

cent rivers (Baptista et al. in prep.).

Other herpetofauna

Although the surveys considerably expanded knowledge

of the regional herpetofauna, further increases to herpe¬

tofaunal diversity in the region are likely. The following

species are considered very likely to be found in south¬

east Angola based on their presence in adjacent areas and

in similar habitats.

Amphibia

Poyntonophrymis fenoidheti - A small toad is known

from western and eastern Caprivi, but unrecorded from

south-east Angola (Channing 2001).

Poyntonophrymis kavangensis - A rare toad known from

scattered localities in the Caprivi region from Rundu to

Hwange, including a single record (23 km NW Pereira de

Eca) from southern Angola (Poynton and Haacke 1993).

Hildebrandtia ornata - Recorded from eastern and west¬

ern Caprivi (Channing 2001), but no localities known

from south-east Angola.

Pyxicephahis edulis - Recorded from Eastern Caprivi

(Channing 2001) and northern Namibia (Ondangwa,

Branch pers. obs.)

Reptilia

Afrotyphlops mucruso / schlegeli - The status of large

typhlopids in Angola previously assigned to Megatyph-

lops (Broadley and Wallach 2009) remains problematic.

Broadley and Wallach (2009) report no specimens from

south-east Angola, although they record three species

in the country: A. mucroso from adjacent western Zam¬

bia, with a single record of unstated provenance mapped

(Macanda, 10°50’S, 18°52’E) from north-eastern Ango¬

la; A. schlegelii from the Caprivi and northern Namibia,

extending along the escarpment into the Benguela Prov¬

ince; and A. anomalus in south-central Angola. Afro¬

typhlops schlegelii may, therefore, occur in upland areas

associated with sparse Miombo woodland bordering the

river catchments.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Table 3. The following are updated species lists for amphibians based on historical records as well as data from the new surveys

within the boundaries of the Cubango, Cuito, and Cuando river basin. Type of record: V = Voucher, O = Observation, L = Literature

record.

Species

Cubango

River

Cuito

River

Cuando

River

Reference

ORDER: SQUAMATA

SERPENTES - COLE BRIDAE

Crotaphopeltis hotamboeia (Laurenti, 1768)

L, V

—

Monard 1931, 1937b; Branch and McCartney 1992

Dasypeltis scabra scabrci (Linnaeus, 1758)

—

Monard 1937b

Dispholidits typiis typiis (Smith, 1829)

—

Bocage 1895; Monard 1931, 1937b; Branch and McCartney 1992

Limnophis bangweolicus (Mertens, 1936)

—

Limnophis bicolor Gunther, 1865

—

Monard 1937b

Natriciteres olivacea (Peters, 1854)

—

V, DR

—

Philothamnus heterolepidotus (Gunther, 1863)

—

Monard 1931, 1937b

Philothamnus lioplogaster (Gunther, 1863)

—

O, DR

—

Philothamnus irregularis (Leach, 1819)

—

Monard 1931, 1937b

Philothamnus ornatus (Bocage, 1872)

—

V, DR

—

Philothamnus semivariegatus (Smith, 1840)

—

Monard 1931, 1937b

Thelotomis capensis oatseii (Gunther, 1881)

—

Monard 1937b

ELAPIDAE

Dendroaspispolylepispolylepis (Gunther, 1864)

—

Elctpsoidea guntherii Bocage, 1866

—

Bocage 1895

Naja ( Uraeus ) anchietae (Bocage, 1879)

—

Monard 1931, 1937b

Naja (Boulengerina) melanoleuca Hallowell, 1857

—

Bocage 1895

Naja (Afronaja) mossambica Peters, 1854

—

V, DR

Naja (Afronaja) nigricollis Reinhardt, 1843

—

Monard 1931, 1937b

LAMPROPHIIDAE

Atractaspis congica congica Peters, 1877

—

Bocage 1895

Aparallactus capensis Smith, 1849

—

Branch and McCartney 1992

Boaedon cf. angolensis Bocage, 1895

—

Bocage 1895; Monard 1931, 1937b (as B. lineatus)

Gonionotophis capensis (Smith, 1847)

—

Monard 1931, 1937b; Branch and McCartney 1992

Lycophidion capense capense (Smith, 1831)

—

Bocage 1895

Lycophidion multimaculatum Boettger, 1888

—

Branch and McCartney 1992

Prosymna angolensis Boulenger, 1915

—

Monard 1931, 1937b

Psammophis jadae Peracca, 1896

—

Monard 1937b

Psammophis mossambicus Peters, 1882

—

Branch and McCartney 1992

Psammophis brevirostris Peters, 1881

—

Monard 1937b

Psammophis sibilans (Linnaeus, 1758)

—

Bocage 1895; Monard 1931, 1937b

Psammophis subtaeniatus Peters, 1882

—

V, DR

—

Pscimmophylax acutus (Gunther, 1888)

—

Psammophylax tritaeniatus (Gunther, 1868)

—

Monard 1931, 1937b

Pseudaspis cana (Linnaeus, 1758)

—

Bocage 1895

Xenocalamus mechowii inornatus Witte and Laurent, 1947

—

Branch and McCartney 1992

TYPHLOPIDAE

Afrotyphlops anomalus (Bocage, 1873)

—

Monard 1937b

LEPTOTYPHLOPIDAE

Leptotyphlops scutifrons scutifrons (Peters, 1854)

L, V

—

Monard 1937b

PYTHONIDAE

Python natalensis Smith, 1840

O, L

—

Monard 1931, 1937b

VIPERIDAE

Bids arietans (Merrem, 1820)

L, V

Monard 1931, 1937b; Branch and McCartney 1992

Causus cf. rasmusseni Broadley, 2014

V,CR

—

Monard 1931, 1937b

SAURIA - AGAMIDAE

Acanthocercus cyanocephalus (Falk, 1925)

—

Bocage 1895; Monard 1931,1937b

Agama aculeata Merrem, 1820

L, V

—

Bocage 1895; Monard 1931, 1937b

Agamaplaniceps Peters, 1862

—

Monard 1937b

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Conradie et al.

Table 3 (continued). The following are updated species lists for amphibians based on historical records as well as data from the

new surveys within the boundaries of the Cubango, Cuito, and Cuando river basin. Type of record: V = Voucher, O = Observation,

L = Literature record.

Species

Cubango

River

Cuito

River

Cuando

River

Reference

AMPHISBAENIDAE

Dalophia angolensis Gans, 1976

Monard 1937b

Dalophiapistillum (Boettger, 1895)

—

Monard 1937b; Branch and McCartney 1992

Zygaspis quadrifrons (Peters, 1862)

V, DR

—

Monopeltis anchietae (Bocage, 1873)

—

Monard 1931, 1937b

CHAMAELEONIDAE

Chamaeleo dilepis Leach, 1819

O, DR

Monard 1931, 1937b

CORDYLIDAE

Chamaesaura macrolepis (Cope, 1862)

Bocage 1895

GEKKONIDAE

Chondrodactylus cf. pul itzerae (Schmidt, 1933)

V, DR

Lygodactylus angolensis Bocage, 1896

—

Bocage 1896

Lygodactylns bradfieldi Hewitt, 1932

—

V, DR

—

Pacltydactyluspunctatus Petersl 855

—

Monard 1931, 1937b

GERRHOSAURIDAE

Gerrhosaurus cf. nigrolineatus Hallowell, 1854

L, V

Bocage 1895; Monard 1931, 1937b

Tetradactylus ellenbergeri (Angel, 1922)

—

V, DR

—

LACERTIDAE

Heliobolus htgubris (Smith, 1838)

Ichnotropis bivitata Bocage, 1866

—

Monard 1937b

Ichnotropis capensis (Smith, 1838)

—

Bocage 1895; Monard 1931, 1937b; Branch and McCartney 1992

Ichnotropis spp.

—

Meroles squamulosus (Peters, 1854)

—

SCINCIDAE

Acontias kgalagadi kgalagadi Lamb, Biswas and Bauer,

V CR

2010

Pcmaspis maculicollis Jacobson and Broadley, 2000

—

V, CR

—

Eumecia anchietae anchietae Bocage, 1870

—

Bocage 1895; Monard 1931, 1937b

Mochlus sundevalli (Smith, 1849)

—

Sepsina angolensis Bocage, 1866

—

Branch and McCartney 1992

Lubuya ivensii (Bocage, 1879)

V, DR

—

Trachylepis angolensis (?) (Monard, 1937)

—

Monard 1937b; Laurent, 1964

Trachylepis varia (Peters, 1867)

—

Trachylepis wahlbergi (Peters, 1869)

V, L

Bocage 1895; Branch and McCartney 1992

Trachylepis cf. spilogaster (Peters, 1882)

V, DR

—

Typhlacontias rohani Angel, 1923

—

V, L

Monard 1931, 1937b

VARANIDAE

Varanus albigularis (Daudin, 1802)

Monard 1937b

Varanus niloticus (Linnaeus, 1766)

L, O

Monard 1931

ORDER: CROCODYLIA

CROCODYLIDAE

Crocodylus niloticus Laurenti, 1768

L, O

Monard 1937b; Branch and McCartney 1992

ORDER: TESTUDINES

PELOMEDUSIDAE

Pelomedusa subrufa (Bonnaterre, 1789)

Peiusios bechuanicus FitzSimons, 1932

—

Monard 1937b

Pe/usios rhodesianus Hewitt, 1927

—

Monard 1937b

TESTUDINIDAE

Kinixys cf. belliana Gray, 1863

L, O

Bocage 1895; Monard 1931, 1937b

Stigmocheiyspardalis (Bell, 1828)

—

Total: 77 (38 snakes, 33 lizards, 1 crocodile, 5 chelonians)

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Herpetofauna of river catchments in south-eastern Angola

Atractaspis bibroni - There are few documented Ango¬

lan records for A. bibronii , and all are located in the west

or north: Benguela (Boulenger 1895), Catumbela and

Dombe (Bocage 1867), Cubal (Mertens 1938), Dundo

(Laurent 1964). Broadley (1991) in his revision of the

southern African populations of Atractaspis gives no

new records for south-east Angola, even though there are

numerous records for adjacent Namibia, Botswana, and

western Zambia.

Amblyodipsas polylepis - Recorded from the northern

Namibia, western Zambia (Broadley 1971) and western

Angola (Bocage 1895; Ferreira 1906), but not recorded

yet from southeast Angola.

Amblyodipsas ventrimaculata - Broadley (1971) noted

this fossorial snake was restricted to Kalahari sands in

northern Botswana, adjacent eastern Namibia, and west¬

ern Zambia. It is a specialist feeder on fossorial reptiles

and several suitable prey items (e.g., Zygaspis quadri-

frons and Typhlacontias rohani ) occur in south-east An¬

gola.

Xenocalamus b. bicolor - Although Broadley (1971)

records no Angolan material the species occurs in the

Caprivi area and adjacent western Zambia, and it is usu¬

ally associated with Kalahari sands.

Hemirhaggheris nototaenia - A dwarf arboreal snake

recorded from the western Caprivi and Okavango re¬

gion, east through Zambia to East Africa (Broadley and

Hughes 2000). Earlier records from southwest Angola

(Bocage 1895) have been referred to H. viperinus by

Broadley and Hughes (2000).

Psammophis jallae - Angel (1921) described Psammo-

phis rohani (type: MNHN 20-198; type locality Lumuna

River, a tributary of the Luina and Cuando Rivers) from

south-east Angola. It was synonymized with P. jallae

Peracca, 1886 by Broadley (2002). No other specimens

of P. jallae are known from the region.

Crotaphopeltis barotseensis - A semiaquatic snake, re¬

stricted to the Okavango and Kafue region (Rasmussen

1997), but presently unrecorded from Angola.

Elapsoidea semiannulata - Recorded in western Zambia,

northern Namibia, and west and central Angola (Broad¬

ley 1998), but no records from southeast Angolan are

known.

Zygaspis nigra - Known from three localities in east-cen¬

tral Angola (Gago Coutinho District, Alto Chicapa, Col-

anda, seven km E Vila Luso on Moxico road; Broadley

and Gans 1975) and adjacent populations in the Caprivi

and Zambia, west of the Zambezi River.

Dalophia longicauda - Described from northern Namibia

and known to extend through the Caprivi area to western

Zimbabwe (Broadley et al. 1976; Gans 2005). Popula¬

tions are therefore found on either side of the Okavango

River and it is expected to occur in southeast Angola.

Ichnotropis grandiceps - Since its description (Broadley

1967) from the Caprivi area, no new material has been

discovered. As noted for Ichnotropis sp. (above), the sta¬

tus of new Angolan material and the validity of I. grandi¬

ceps are under investigation.

Colopus wahlbergii - This small terrestrial gecko is

widespread in the Kalahari region, with a single, mar¬

ginal Angolan record (Angola-Namibia border, 18°E;

Haacke 1976). It has subsequently been recorded from

the Caprivi (Haacke 1998, Branch unpub. obs.), Zimba¬

bwe (Broadley and Spawls 1991) and extreme western

Zambia (Broadley and van Daele 2003), and is, there¬

fore, likely to occur in the sandy areas of the region.

Acknowledgments. —We thank the following interna¬

tional projects leaders for inviting us to collaborate: Chris

Brooks, organizer of the SAREP Aquatic Biodiversity

Survey of the upper Angola Catchments of the Cubango-

Okavango River Basin (May 2012) and the lower Cuito

and Cuando River basins (April 2013); Steve Boyes and

John Hilton of the Wild Bird Trust which administers the

Okavango Wilderness Project which took a team of bio¬

diversity experts to the source of the Cuito River down to

its confluence with the Cubango River (May-June 2015;

National Geographic Society grant number EC07 IS¬

IS). These projects were supported by and also involved

members of the Angolan Ministry of Environment Insti¬

tute of Biodiversity (MINAMB) and the Angola Ministry

of Agriculture National Institute of Fish Research (INIP),

who facilitated the export of voucher specimens for

analysis. Various colleagues are thanked for collecting

herpetological material during their fieldwork, including

Paul Skelton, Roger Bills, Maans Booysens, and others.

We especially thank our ISCED colleagues in Angola,

Pedro Vaz Pinto, Ninda Baptista (who also prepared the

Resumo), and Fernanda Lages, for their many insights

into their beautiful country, and for welcoming us with

such hospitality.

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Herpetofauna of river catchments in south-eastern Angola

Werner Conradie has ten years of experience in southern African herpetofauna, with his main research

interests focusing on the taxonomy, conservation, and ecology of amphibians and reptiles. He has published

numerous principal and collaborative scientific papers, and has served on a number of conservation and

scientific panels, including the Reptile Atlas Committee and Amphibian 1UCN Workshop. Werner has

represented his field on television and in numerous field guides and has participated in expeditions in various

countries including Namibia, Botswana, Zimbabwe, Mozambique, Angola, Malawi, Lesotho, and Zambia.

He is currently the Curator of Herpetology at the Port Elizabeth Museum (Bay world), South Africa.

Bill Branch (London-born) was employed as Curator of Herpetology at the Port Elizabeth Museum for over

30 years (1979-2011), and although now retired remains Curator Emeritus Herpetology. His herpetological

studies have concentrated mainly on the systematics, phylogenetic relationships, and conservation of African

reptiles, but he has been involved in numerous other studies on the reproduction and diet of African snakes.

He has published over 300 scientific articles, as well as numerous popular articles and books. The latter

include: South African Red Data Book of Reptiles and Amphibians (1988), Dangerous Snakes of Africa (1995,

with Steve Spawls), Field Guide to the Reptiles of Southern Africa (1998), Tortoises , Terrapins and Turtles of

Africa (2008), and Atlas and Red Data Book of the Reptiles of South Africa , Lesotho and Swaziland (multi-

authored, 2014), as well as smaller photographic guides. In 2004 he was the 4 th recipient of the “Exceptional

Contribution to Herpetology” award of the Herpetological Association of Africa. He has undertaken field

work in over 16 African countries, and described over 40 species, including geckos, lacertids, chameleons,

cordylids, tortoises, adders, and frogs.

Roger Bills is the Curator of Freshwater Fishes and the Collections Manager at the South African Institute for

Aquatic Biodiversity, Grahamstown, having been employed there since 1995. Research interests include the

■> taxonomy and systematics of southern African freshwater fishes particularly Zambezi and Lake Tanganyika

cichlids, mochokid, and claroteid catfishes. Recently he has developed a growing interest in southern

African amphibians and the continuing development of the SAIAB amphibian collection. He has extensive

expeditionary experience in southern and central Africa.

Amphib. Reptile Conserv.

October 2016 | Volume 10 | Number 2 | el 26

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [Special Section]: 37-67 (e131).

A survey of amphibians and reptiles in the

foothills of Mount Kupe, Cameroon

12 Daniel M. Portik, 34 Gregory F.M. Jongsma, 3 Marcel T. Kouete, 3 Lauren A. Scheinberg,

3 Brian Freiermuth, 56 Walter P. Tapondjou, and 34 David C. Blackburn

1 Museum of Vertebrate Zoology, University of California, Berkeley, 3101 Valley Life Sciences Building, Berkeley, California 94720,

USA 2 Department of Biology, University of Texas at Arlington, 501 S. Nedderman Drive, Box 19498, Arlington, Texas 76019-0498,

USA 3 California Academy of Sciences, San Francisco, California 94118, USA 4 Florida Museum of Natural History, University> of

Florida, Gainesville, Florida 32611, USA 5 Laboratory of Zoology’, Faculty of Science. University of Yaounde, PO Box 812 Yaounde,

Cameroon, AFRLCA 6 Department ofEcology and Evolutionary Biology, University of Kansas, 1450 Jayhawk Boulevard, Lawrence,

Kansas 66045, USA

Abstract.—We performed surveys at several lower elevation sites surrounding Mt. Kupe, a mountain at the

southern edge of the Cameroonian Highlands. This work resulted in the sampling of 48 species, including 38

amphibian and 10 reptile species. By combining our data with prior survey results from higher elevation zones,

we produce a checklist of 108 species for the greater Mt. Kupe region including 72 frog species, 21 lizard

species, and 15 species of snakes. Our work adds 30 species of frogs at lower elevations, many of which are

associated with breeding in pools or ponds that are absent from the slopes of Mt. Kupe. We provide taxonomic

accounts, including museum specimen data and associated molecular data, for all species encountered.

Finally, we compare the levels of biodiversity of Mt. Kupe to other regions, discuss biogeographic ties to other

montane systems, and note current conservation threats.

Keywords. Africa, Anura, biodiversity, biogeography, Cameroon, herpetofauna, lowland forest

Citation: Portik DM, Jongsma GFM, Kouete MT, Scheinberg LA, Freiermuth B, Tapondjou WP, Blackburn DC. 2016. A survey of amphibians and

reptiles in the foothills of Mount Kupe, Cameroon. Amphibian & Reptile Conservation 10(2) [Special Section]: 37-67 (el31).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any

medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and

authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation-, official journal

website <amphibian-reptile-conservation.org>.

Received: 10 August 2016; Accepted: 11 December 2016; Published:

Introduction

The Cameroon Volcanic Line (CVL), also referred to as

the Cameroonian Highlands (or la dorsal camerounaise),

is a series of insular-like reliefs including oceanic (An-

nobon, Sao Tome, Principe) and continental (Bioko) is¬

lands, and mainland volcanic ranges such as Mt. Camer¬

oon (4,085 m), Mt. Kupe (2,064 m), Manengouba (2,411

m), and Mt. Nlonako (1,822 m). The CVL continues

through the Bakossi Highlands and Bamenda Highlands,

including the Bamboutos Mountains (2,740 m), Mt. Lefo

(2,550 m), and Mt. Oku (3,011 m), ultimately ending at

the Adamawa (Adamaoua) Plateau, where Tchabal Mba-

bo (2,460 m) is located. Though these montane regions

exhibit both high species richness and endemism, beyond

larger summaries of Cameroonian reptiles and amphib¬

ians (Chirio and LeBreton 2007; Amiet 2012) detailed

Correspondence. Email: 1 daniel.portik@uta.edu (Corresponding c

December 2016

faunal reports for particular mountains are not common.

A growing number of surveys have provided baseline

checklists for reptiles or amphibians of particular sites,

including Mt. Cameroon, Mt. Nlonako, Mt. Kupe, Mt.

Oku, and Tchabal Mbabo (Amiet 1975; Hofer et al. 1999,

2000; Plath et al. 2004; Herrmann et al. 2005a, 2005b;

Herrmann et al. 2006; Gonwouo et al. 2007; Ineich et

al. 2015; Hirschfeld et al. 2016). Through this work, el-

evational distribution and biogeographic patterns have

emerged, though information for many regions is incom¬

plete or absent. Where elevational data are available, re¬

cent work demonstrates declines in amphibian popula¬

tions on several mountains, including Mt. Manengouba

and Mt. Oku (Hirschfeld et al. 2016). Inventories of the

herpetofauna of Mt. Nlonako, including sites ranging

from 400-1,700 m elevation, resulted in the documenta¬

tion of 93 amphibian and 89 reptile species, making it

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

1000m

1500m

Mt Manengouba

2000rn^ J f

QNkongsamba

Mt Nlonako

Mt Koupe

20 Kilometers

TCD

J7 7 -

NGA

CAF

COG

■ Wo.., <?

Fig. 1. Map of Cameroon with the location of several summits labeled, including Mt. Kupe, Mt. Manengouba, and Mt. Nlonako

(white triangles), and major towns (white squares). Localities sampled for this study are denoted by white circles. Elevation is

indicated by contour lines and shading, with lighter colors representing higher elevation.

one of the most species-rich mountains in tropical Africa

(Amiet 1975; Plath et al. 2004; Herrmann et al. 2005a,

2005b). Several distinctive anuran faunas converge in

this region (Amiet 1975), partially explaining the high

species richness. Recent survey and museum work across

the CVL continues to result in discovery of new species

(Bohme and Schmitz 1996; Amiet and Dowsett-Lemaire

2000; Boistel and Amiet 2001; Hernnann et al. 2004;

Rodel et al. 2004; Plath et al. 2006; Blackburn 2008;

Blackburn et al. 2009,2010; Zimkus and Gvozdlk 2013),

further demonstrating the need to better characterize the

herpetofauna in these regions.

Mount Kupe (also referred to as Mont Koupe) lies just

east of the Bakossi Mountains, south of Mt. Manengouba,

and southwest of Mt. Nlonako (Fig. 1). Mt. Kupe rises to

2,064 m, and is most comparable in size and elevation

to Mt. Nlonako. Several vegetation zones occur around

Mt. Kupe, including lowland forest (300-900 m) in the

foothills, submontane forest (900-1,800 m), transitional

montane forest (>1,800 m), and small grassy clearings at

the peak (>2,000 m) (Thomas 1986; Hofer et al. 1999,

2000). The primary lowland forest (below 900 m) has

been greatly reduced or degraded by logging and subsis¬

tence or commercial agriculture (Hofer et al. 1999,2000;

pers. obs.). Streams can be located up to 1,500 m on the

slopes of Mt. Kupe, but permanent ponds are only found

at lower elevations (<900 m) (Hofer et al. 1999, 2000).

Hofer et al. (1999, 2000) surveyed transects largely

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

through the submontane forest in an etfort to character¬

ize species turnover across an elevational gradient (900-

2,000 m). Their work resulted in the documentation of 33

amphibian, 15 lizard, and nine snake species. However,

this is a conservative estimate as no lowland forest sites

or sites containing permanent ponds or temporary pools

were surveyed.

During two periods in 2013 and 2014, we conducted

herpetological surveys in the foothills of Mt. Kupe, ad¬

jacent to the town of Manjo (Fig. 1) in a region known

as the Tombel Plain, that lies between the headwaters of

the Moungo and Dibombe rivers (Amiet 1975). Here, we

discuss the results of our survey work based on a collec¬

tion of 488 specimens. In an etfort to more fully char¬

acterize our results, representatives of all species were

DNA-barcoded using the 16S mitochondrial marker

(Table 1). We provide taxonomic accounts, including

museum specimen data and associated molecular data,

for all species encountered. We also make comparisons

to previous surveys at higher elevations on Mt. Kupe

(Hofer et al. 1999, 2000) and several low elevation sites

(Lala, 4.80°N, 9.76°E, 480 m; Mahole 4.82°N, 9.61°E,

300-350 m; Nlohe, 4.75°N, 9.75°E, 350 m) near Mt.

Kupe (Amiet 1975).

Methods

We conducted surveys at multiple sites in the foothills of

Mt. Kupe, accessed through the nearby town of Manjo.

Surveys were conducted by DMP, GFMJ, and MTK dur¬

ing July 11-14, 2013, and by DMP, GFMJ, LAS, BF,

WPT, and DCB during September 24-29, 2014, for a

total of ten days. A combination of sites was chosen to

maximize the habitats surveyed, and included permanent

ponds in semi-disturbed forest (Fig. 2A), fish ponds (Fig.

2B), large streams in disturbed habitat (Fig. 2C), small

streams in secondary forest (Fig. 2D), and anthropo¬

genic habitats (Fig. 2E). Though primary forest is intact

at lower elevations, a lack of accessibility, permissions,

and time precluded sampling at these sites. All localities

visited are lowland forests or lower strata of submontane

forests (between 470-1,010 m). The short time period

for surveys precluded the use of pitfall traps and other

systematic techniques, and all specimens were hand-cap¬

tured during diurnal and nocturnal visual searches. We

euthanized captured animals using MS-222, preserved

tissue samples in RNA Later (Ambion, Inc.), and pre¬

served whole specimens with 10% buffered formalin.

During this process we took relevant body size measure¬

ments for each specimen to the nearest 0.5 millimeters

(Reptiles—snout-vent length: SVL; tail length: TL; Am¬

phibians—snout-urostyle length: SUL). When possible,

we recorded the sex of specimens based on a variety

of observations. For amphibians this included: 1) male

secondary sexual characters such as nuptial pads, vocal

sacs, elongated fingers, gular glands, tympanic papillae,

2) the tracking of specimens found in amplexus, and 3)

the discovery of eggs in gravid females during tissue

collection. For reptiles this included: 1) male secondary

sexual characters such as coloration or preanal pores, and

2) eversion of hemipenes during the fixation process. In

the absence of such characters, we did not attempt to sex

specimens based on destructive dissection. In the species

accounts we only report the sex of adult specimens for

which we are confident, but do not distinguish between

the reasons for our inability to determine sex (i.e., juve¬

niles vs. lack of external characters). In the absence of

these data, we still include relevant size measurements

for all unsexed adult specimens, which is often useful

for distinguishing between closely related species. After

fixation, specimens were moved to 70% EtOH for long¬

term storage. All 488 specimen vouchers and associated

tissue samples are deposited at the California Academy

of Sciences (CAS).

We performed DNA barcoding on a subset of 92 spec¬

imens to confirm morphological identifications and/or

to generate sequence data for underrepresented groups.

Whole genomic DNA was extracted from liver samples

using a high-salt DNA extraction (Aljanabi and Martinez

1997). We obtained sequence data from the mitochon¬

drial marker 16S ribosomal RNA (16S) using primers

16SA and 16SB (Palumbi et al. 2001). Polymerase chain

reactions (PCRs) were carried out in 12.5 pi volumes

consisting of: 1.25 pi Roche lOx (500 mM Tris/HCl, 100

mM KC1, 50 mM (NH 4 ), S0 4 , 20 mM MgCl„ pH=8.3),

0.75 pi 25 mM MgCl 2 , 0.75 pi 2 mM DNTPs, 0.25 pi

10.0 pM forward primer, 0.25 pi 10.0 pM reverse primer,

8.40 pi H 2 0, 0.10 pi Taq, and 0.75 pi DNA. Amplifica¬

tion of 16S involved initial denaturation at 94 °C for four

minutes, followed by 35 cycles of 95 °C for 60 s, 51 °C

for 60 s, 72 °C for 90 s, and a final extension at 72 °C for

seven minutes. The PCR amplifications were visualized

on an agarose gel and cleaned using ExoSAP-IT (USB).

Gene products were sequenced using BigDye v3.1 on an

ABB730 (Applied Bio systems). All newly generated se¬

quences are deposited in GenBank (Accession numbers:

KX671711-802; Table 1).

Species Accounts

ANURA

Arthroleptidae

Arthroleptis poecilonotus Peters, 1863. Figure 3A, B;

23 specimens. 4.8148°N, 9.7691°E (475 m): (2014)

CAS 256686-89. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256711, 256809-10. 4.8498°N, 9.7718°E (513 m):

(2013) CAS 254029-39. 4.8551°N, 9.8188°E (560 m):

(2014) CAS 256824-27. 4.8247°N, 9.7702°E (494 m):

(2013) CAS 253867. The SUL of unsexed individuals

ranges from 12.5-28.5 mm (average: 20.6, n = 23).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Table 1. List of taxa, museum numbers, and GenBank numbers for 16S mitochondrial sequence data.

Genus

Species

Museum No.

Gen Bank No.

Anura Afrixalus

dorsalis

CAS253854

KX671711

Afrixalus

dorsalis

CAS253855

KX671712

Afrixalus

dorsalis

CAS256690

KX671713

Afrixalus

dorsalis

CAS256691

KX671714

Afrixalus

Jaevis

CAS254071

KX671715

Afrixalus

paradorsalis

CAS253947

KX671716

Afrixalus

paradorsalis

CAS253948

KX671717

Afrixalus

paradorsalis

CAS256750

KX671718

Afrixalus

paradorsalis

CAS256751

KX671719

Amietophrynus

regularis

CAS256712

KX671721

Amietophrynus

regularis

CAS256713

KX671722

Arthroleptis

poecilonotus

CAS253867

KX671723

Arthroleptis

poecilonotus

CAS254029

KX671724

Arthroleptis

poecilonotus

CAS256686

KX671725

Cardioglossa

elegans

CAS256915

KX671726

Cardioglossa

leucomystax

CAS256874

KX671727

Chiromantis

rufescens

CAS254025

KX671728

Conraua

goliath

CAS256830

KX671730

Conraua

robusta

CAS256822

KX671731

Hyperolius

bolifambae

CAS253883

KX671736

Hyperolius

bolifambae

CAS253884

KX671737

Hyperolius

bolifambae

CAS253885

KX671738

Hyperolius

camerunensis

CAS253935

KX671739

Hyperolius

camerunensis

CAS256916

KX671740

Hyperolius

concolor

CAS253869

KX671741

Hyperolius

concolor

CAS253874

KX671742

Hyperolius

dintelmanni

CAS253991

KX671743

Hyperolius

dintelmanni

CAS256693

KX671744

Hyperolius

fusciventris

CAS254005

KX671745

Hyperolius

fusciventris

CAS254006

KX671746

Hyperolius

ocellatus

CAS254057

KX671747

Hyperolius

ocellatus

CAS254058

KX671748

Kassina

decorata

CAS253990

KX671749

Leptopelis

aubryi

CAS253851

KX671750

Leptopelis

aubryi

CAS253852

KX671751

Leptopelis

aubryi

CAS256685

KX671752

Leptopelis

aubryioides

CAS253974

KX671753

Leptopelis

aubryioides

CAS256719

KX671754

Leptopelis

boulengeri

CAS253980

KX671755

Leptopelis

boulengeri

CAS256875

KX671756

Leptopelis

calcaratus

CAS253981

KX671757

Leptopelis

calcaratus

CAS253982

KX671758

Leptopelis

rufus

CAS254045

KX671759

Leptopelis

rufus

CAS254048

KX671760

Leptopelis

rufus

CAS256682

KX671761

Leptopelis

rufus

CAS256683

KX671762

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

Table 1 (continued). List of taxa, museum numbers, and GenBank numbers for 16S mitochondrial sequence data.

Genus

Species

Museum No.

Gen Bank No.

Leptopelis

rufus

CAS256882

KX671763

Nyctibates

corrugatus

CAS256829

KX671764

Petropedetes

cameronensis

CAS256887

KX671766

Petropedetes

cameronensis

CAS256888

KX671767

Petropedetes

euskircheni

CAS254066

KX671768

Petropedetes

euskircheni

CAS254067

KX671769

Phlyctimantis

leonardi

CAS253978

KX671770

Phlyctimantis

leonardi

CAS253979

KX671771

Phrynobatrachus

africanus

CAS253989

KX671772

Phrynobatrachus

africanus

CAS256872

KX671773

Phrynobatrachus

auritus

CAS256669

KX671774

Phrynobatrachus

auritus

CAS256670

KX671775

Phrynobatrachus

auritus

CAS256674

KX671776

Phrynobatrachus

cornutus

CAS256868

KX671777

Phrynobatrachus

cornutus

CAS256869

KX671778

Phrynobatrachus

cornutus

CAS256870

KX671779

Phrynobatrachus

cornutus

CAS256871

KX671780

Phrynobatrachus

cornutus

CAS256873

KX671781

Phrynobatrachus

sp.

CAS256714

KX671782

Phrynobatrachus

sp.

CAS256715

KX671783

Ptychadena

cf. aequiplicata

CAS254024

KX671789

Ptychadena

cf. aequiplicata

CAS254054

KX671790

Ptychadena

cf aequiplicata

CAS254055

KX671791

Ptychadena

cf aequiplicata

CAS254056

KX671792

Ptychadena

cf mascariensis

CAS254083

KX671785

Ptychadena

cf mascariensis

CAS254084

KX671786

Ptychadena

oxyrhynchus

CAS256820

KX671787

Ptychadena

oxyrhynchus

CAS256862

KX671788

Scotobleps

gabonicus

CAS254015

KX671793

Scotobleps

gabonicus

CAS256676

KX671794

Scotobleps

gabonicus

CAS256677

KX671795

Trichobatrachus

robustus

CAS254082

KX671802

Squamata

Agama

lebretoni

CAS253879

KX671720

Cnemaspis

spinicollis

CAS256886

KX671729

Dipsadoboa

duchesnii

CAS254086

KX671732

Grayia

inornata

CAS256859

KX671733

Hapsidophrys

smaragdinus

CAS256860

KX671734

Hapsidophrys

smaragdinus

CAS256861

KX671735

Panaspis

breviceps

CAS256855

KX671765

Poromera

fordii

CAS254085

KX671784

Trachylepis

affinis

CAS256707

KX671796

Trachylepis

affinis

CAS256856

KX671797

Trachylepis

affinis

CAS256857

KX671798

Trachylepis

maculilabris

CAS253881

KX671799

Trachylepis

maculilabris

CAS253882

KX671800

Trachylepis

maculilabris

CAS256792

KX671801

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 2 . Habitats surveyed during this fieldwork: (A) Main pond near plantation where a majority of hyperoliid species were caught;

(B) fish ponds and associated vegetation next to large stream close to village; (C) large stream adjacent to fish ponds; (D) smaller

streams located within secondary or primary forest; (E) disturbed habitat near plantation.

ArthroJeptis poecilonotus is a common and widespread

species, especially in degraded areas lacking closed can¬

opy forest. The Central African populations ranging from

Nigeria south to at least Republic of Congo represent a

distinct species from populations in West Africa (west of

Nigeria) that might represent one or more species (Rodel

and Bangoura 2004; Blackburn et al. 2010; Jackson and

Blackburn 2010). At higher elevations in the CVL, this

species is often replaced by a morphologically similar

but distantly related species, A. palava (Blackburn et al.

2010 ).

Cardioglossa elegans Boulenger, 1906. Figure 3C, D;

13 specimens. 4.8497°N, 9.7719°E (510 m): (2014)

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

CAS 256710. 4.8500°N, 9.7673°E (538 m): (2013) CAS

254064-65; (2014) CAS 256891, 256892-97, 256913-

15. Females average 35.9 mm SUL (35.0-36.5 mm, n =

5) and males average 27.4 mm SUL (23.0-28.5 mm, n

= 10). Individuals were caught on the ground or sitting

on leaves (< 0.75 m high) adjacent to a small stream.

Females were gravid. Cardioglossa elegans is widely

distributed in forests below 1,000 m with records from

on and near Mt. Kupe and nearby Mt. Nlonako (Amiet

1975, 1978; Hofer et al. 1999; Herrmann et al. 2005).

Cardioglossa leucomystax (Boulenger, 1903). Figure

3E; two specimens. 4.8406°N, 9.7793°E (491 m): (2014)

CAS 256874. 4.8498°N, 9.7718°E (514 m): (2013) CAS

254080. Two males measure 27.0 and 28.0 mm SUL. In¬

dividuals were found on the high banks of a larger sec¬

tion of stream in secondary forest. While not recorded at

sites around or above 900 m by Hofer et al. (1999), C.

leucoymystax is well known from forested sites below

1,000 m in this region (Amiet 1975).

Leptodactylodon ovatns Andersson, 1903. Figure 3F;

one specimen. 4.8498°N, 9.7718°E (513 m): (2013)

CAS 253933. Single male specimen measures 39.0 mm

SUL. This individual was found moving across leaf lit¬

ter between two permanent ponds during heavy rainfall.

This specimen agrees with the description of L. ovatus

orientals provided by Amiet (1971, 1980) based on the

rose coloration and more limited dark pigmentation on

its venter. While L. ovatus is known from forested sites

below 1,000 m in this region (Amiet 1975), the boundary

between the two subspecies ( ovatus and orientalis) is not

clear. This newly collected specimen is along the western

edge of the described distribution for L. ovatus orientalis

(Amiet 1980).

Leptopelis aubryi (Dumeril, 1865). Figure 3G; ten speci¬

mens. 4.8148°N, 9.7691°E (475 m): (2014) CAS 256685.

4.8498°N, 9.7718°E (513 m): (2013) CAS 253969-73,

253976. 4.8247°N, 9.7702°E (494 m): (2013) CAS

253851-53. Females average 37.8 mm SUL (30.5-51.5

mm, n = 5) and males average 35.8 mm SUL (31.0—44.0

mm, n = 3). This small Leptopelis most closely resembles

Leptopelis aubryioides and L. modestus , but lacks the

heel spurs possessed by the former and lacks the white

sub-ocular patch possessed by the latter (Amiet 2012).

Leptopelis aubryi is found in lowland forests in south¬

western and southern Cameroon (Amiet 2012), includ¬

ing near Manjo (Amiet 1975), and extends into mainland

Equatorial Guinea, Gabon, and the Republic of Congo

(Lasso et al. 2002; Burger et al. 2006; Jackson and Black¬

burn 2007; Pauwels and Rodel 2007).

Leptopelis aubryioides (Andersson, 1907). Figure

4A, B; eight specimens. 4.8498°N, 9.7718°E (513 m):

(2013) CAS 253974-75, 254053, 254061-63, 254088.

4.8560°N, 9.7480°E (1,010 m): (2014) CAS 256719.

Amphib. Reptile Conserv.

Females average 46.0 mm SUL (n = 2) and males av¬

erage 32.8 mm SUL (31.0-35.0 mm, n = 4). This spe¬

cies is easily distinguished by the presence of heel spurs.

Leptopelis calcaratus also has heel spurs; however, this

species is larger (males 35^42 mm, females 46-57 mm;

Schiotz 1999) and has a conspicuous white sub-ocular

patch (Amiet 2012). Among Cameroonian species of

Leptopelis, L. aubryioides is one of the most frequently

encountered lowland forest species (Amiet 2012).

Leptopelis boulengeri (Werner, 1898). Figure 4C, D; two

specimens. 4.8498°N, 9.7718°E (513 m): (2013) CAS

253980. 4.8560°N, 9.7480°E (1,010 m): (2014) CAS

256875. Female measures 70.5 mm SUL (CAS 256875).

Both the brown and green phases of this species were

found, which is identified by the presence of a conspicu¬

ous white spot (sub-ocular patch) below the eye and its

large body size (males 37-48 mm, females 60-81 mm;

Schiotz 1999). The brown phase exhibits a dorsal trian¬

gle with apex pointed forward (Fig. 4C), and both phases

often have black speckling on the flanks and a stripe

between the eyes. Leptopelis boulengeri is distributed

similarly to L. aubryioides and is frequently encountered

in lowland forests of Cameroon (including near Manjo;

Amiet 1975, 2012), mainland Equatorial Guinea (Lasso

et al. 2002), Gabon (Burger et al. 2006; Pauwels and

Rodel 2007), northern Republic of Congo (Capula et al.

2011), and may also extend west into southwestern Nige¬

ria (Onadeko and Rodel 2009; Onadeko 2016).

Leptopelis calcaratus (Boulenger, 1906). Figure 3H;

two specimens. 4.8498°N, 9.7718°E (513 m): (2013)

CAS 253981-82. Male specimen (CAS 253981) mea¬

sures 34.0 mm SUL, female specimen (CAS 253982)

measures 53.5 mm. Leptopelis calcaratus possesses heel

spurs, white sub-ocular patches, and usually has black

spots along its flanks. This forest species has a broad

elevational range (Hernnann et al. 2005a; Amiet 2012)

and, while not originally recorded near Manjo by Amiet

(1975), is now known to occur widely in this region of

Cameroon (Amiet 2012) and broadly across much of

Central Africa and extending west into southwestern Ni¬

geria (Onadeko 2016).

Leptopelis rufus Reichnow, 1874. Figure 4E, F; 31

specimens. 4.8148°N, 9.7691°E (475 m): (2014) CAS

256680-84. 4.8498°N, 9.7718°E (513 m): (2013) CAS

253968, 254042-52. 4.8551°N, 9.8188°E (560 m):

(2014) CAS 256823. 4.8560°N, 9.7480°E (1,010 m):

(2014) CAS 256717-18, 256806-08, 256876-83. Fe¬

males average 76.7 mm SUL (67.0-84.0 mm, n =9) and

males average 50.2 mm SUL (39.0-57.0 mm, n = 19).

This species is identified by its large size (males 45-55

mm, females 74-87 mm; Schiotz 1999), and is distin¬

guishable from L. boulengeri by having dark transverse

bars on the dorsum and the absence of a white eye spot.

Leptopelis rufus is distinguishable from L. millsoni by its

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 3. Representatives of arthroleptid frog taxa. (A ) Arthroleptis poecilonotus , male (CAS 256711); (B ) Arthroleptis poecilonotus

(CAS 254036); (C) Cardioglossa elegans, female (CAS 256893); (D) Cardioglossa elegans, male (CAS 256710); (E) Cardioglossa

leucomystax, male (CAS 254080); (F) Leptodactylodon ovatus (CAS 253933); (G) Leptopelis aubryi (CAS 253852); (H) Leptopelis

calcaratus (CAS 253981).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

smaller tympanum and pectoral glands in males and by

their much larger size for females. Specimens were often

collected from tree branches overhanging swiftly moving

streams, where males were heard calling. It is frequently

encountered in the lowland forests of Cameroon and Ga¬

bon up to -1,000 m, though it is not typically encoun¬

tered above 1,100 m (Amiet 1975, 2012; Herrmann et

al. 2005a; Burger al. 2006; Pauwels and Rodel 2007),

and is known from Mt. Kupe (Amiet 1975; Schmitz et

al. 1999).

Nyctibates corrugatus Boulenger, 1904. Figure 4G; one

specimen. 4.8551°N, 9.8188°E (560 m): (2014) CAS

256829. Female measures 55.0 mm SUL, and was gravid.

Nyctibates corrugatus is a common species at lower el¬

evations in the forests of southwestern Cameroon (Amiet

1973, 1975, 1978; Lawson 1993; Herrmann et al. 2005a)

and extending into mainland Equatorial Guinea (Lasso et

al. 2002). This specimen was collected 20 meters inland

from the large stream illustrated in Fig. 2C, in an agricul¬

tural plot of cocoa.

Scotobleps gabonicus Boulenger, 1900. Figure 4H; 19

specimens. 4.8148°N, 9.7691°E (475 m): (2014) CAS

256676-79. 4.8406°N, 9.7793°E (491 m): (2014) CAS

256863-67. 4.8497°N, 9.7719°E (510 m): (2014) CAS

256709. 4.8498°N, 9.7718°E (513 m): (2013) CAS

254015-22. 4.8551°N, 9.8188°E (560 m): (2014) CAS

256828. Confirmed males average 35.6 mm SUL (23.5-

44.0 mm, n = 7) and confirmed female specimen is 63.5

mm SUL. Specimens were generally collected in the

leaf litter adjacent to smaller streams, in semi-disturbed

habitat. The range of S. gabonicus extends from western

Nigeria, through Cameroon, and south through mainland

Equatorial Guinea, Gabon, and into the forested coastal

region of Democratic Republic of Congo (Laurent 1961;

Amiet 1975, 1978; Lasso et al. 2002; Burger et al. 2006;

Pauwels and Rodel 2007). Like Cardioglossa elegans, S.

gabonicus is common in the littoral plain at low eleva¬

tions in southwestern Cameroon (Amiet 1975, 1983).

Trichobatrachus robustus Boulenger, 1900. Figure 5A-

C; six specimens. 4.8339°N, 9.7783°E (490 m): (2013)

CAS 254069-70. 4.8498°N, 9.7718°E (513 m): (2013)

CAS 253934, 254082. 4.8506°N, 9.7671°E (541 m):

(2014) CAS 256884-85. Females average 79.5 mm SUL

(74.0 and 85.0 mm, n = 2) and males average 78.5 mm

SUL (72.5-90.0 mm, n = 3). Specimens were collected

in close proximity to swiftly moving streams in second¬

ary forest (Fig. 2D), but were also collected away from

the streams in burrows or moving during rainstonns. All

males that we collected exhibited developed “hairs” on

the legs and flanks. Trichobatrachus robustus is found in

the coastal Atlantic forests extending from Nigeria south

to northern Angola (Amiet 1978; Ernst et al. 2014) and is

known from low elevations both on and near Mt. Kupe

(Amiet 1975; Euskirchen et al. 1999; Hofer et al. 1999).

Amphib. Reptile Conserv.

Bufonidae

Sclerophrys regularis (Reuss, 1833). Three specimens.

4.8406°N, 9.7793°E (491 m): (2014) CAS 256712-13.

4.8498°N, 9.7718°E (513 m): (2013) CAS 254023. Male

specimens are 51.5 mm and 55.0 mm SUL (CAS 256713,

254023) and female is 66.5 mm SUL (CAS 256712).

Specimens were abundant in the most disturbed habitat

sampled (Fig. 2E), and were calling from puddles on the

main dirt road through a banana plantation.

Sclerophrys superciliaris (Boulenger, 1888). 4.8500°N,

9.7673°E: (2014) photo voucher MVZObs:Herp:20 (Fig.

5F). We found and photographed a large individual cor¬

responding to the subspecies S. superciliaris supercili¬

aris (Barej et al. 2011) perched on a large boulder under

a stand of bamboo that was adjacent to a swiftly moving

stream in secondary forest. This uncommon forest spe¬

cies generally occurs at low elevations from Nigeria to

Gabon (Barej et al. 2011). Previously found on Mt. Nlon-

ako (Herrmann et al. 2005a) and just to the northwest of

Mt. Kupe at Meked (Barej et al. 2011), we believe this to

be the first definitive record on Mt. Kupe.

Conrauidae

Conraua goliath (Boulenger, 1906). Figure 5D; single

specimen. 4.8551°N, 9.8188°E (560 m): (2014) CAS

256830. A juvenile specimen (measuring 51.5 mm SUL)

was collected on the bank of the large stream in Fig. 2C.

This specimen was identified in the field as C. robusta

but DNA sequence data reveals that it is a juvenile C.

goliath. Larger adults were spotted in large and more

slowly moving streams during surveys but not collected.

The largest adult encountered was spotted at night in a

culvert directly next to the highway where locals often

washed clothing. Conraua goliath is currently the only

anuran species afforded special protection by the Repub¬

lic of Cameroon, largely due to hunting of this species for

human consumption.

Conraua robusta Nieden, 1908. Figure 5E; two speci¬

mens. 4.8506°N, 9.7671°E (541 m): (2014) CAS 256822.

4.8551°N, 9.8188°E (560 m): (2014) CAS 256821. Juve¬

nile specimen (CAS 256821) is 56.5 mm SUL and un-

sexed adult (CAS 256822) is 130.0 mm SUL. Conraua

robusta occurs along fast-moving streams along a broad

elevational gradient (up to -1,850 m) and has long been

known from southwestern Cameroon (Amiet 1978), in¬

cluding near Nkongsamba and on Mt. Kupe (Lamotte

and Perret 1968; Amiet 1975, 1983; Euskirchen et al.

1999). The juvenile specimen was collected from a large

boulder within the stream in Fig. 2C, whereas the adult

was found in a small section of stream near its confluence

with a larger stream in the secondary forest illustrated

in Fig. 2D. Additional specimens were observed in this

stream but were not captured.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 4. Representatives of arthroleptid frog taxa. (A) Leptopelis aubryioides (CAS 256719); (B) Leptopelis aubryioides , female

(CAS 254061); (C) Leptopelis boulengeri (CAS 253980); (D) Leptopelis boulengeri (CAS 256875); (E) Leptopelis rufus (CAS

256684); (F) Leptopelis rufus (CAS 254046); (G) Nyctibates corrugatus (CAS 256829); (H) Scotobleps gabonicus (CAS 256678).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

Hyperoliidae

For this family our discussion of color patterns follows

usage of the terminology of Schiotz (1999), which in¬

cludes Phase J and Phase F. Phase J describes the ju¬

venile coloration, which is also typical of a majority of

adult males. Phase F describes the color pattern of adult

females, and in some species this can also occur in subset

of adult males. All females begin with Phase J coloration

and transition to Phase F coloration, whereas males can

either retain Phase J coloration or transition to Phase F

upon maturity. Here, we also report evidence for adult

males transitioning between these color phases, a phe¬

nomenon that has received little, if any, attention in the

literature. When possible, we quantify the percentage of

the adult males sampled in the population that exhibit

Phase J vs. Phase F coloration, as hyperoliid species vary

greatly in the composition of color morphs (DMP, pers.

obs.).

Afrixalus dorsalis (Peters, 1875). Figure 5G, H, and Fig.

6A; 29 specimens. 4.8148°N, 9.7691°E (475 m): (2014)

CAS 256690-92. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256747^19, 256819. 4.8498°N, 9.7718°E (513 m):

(2013) CAS 253955-61, 253965. 4.8551°N, 9.8188°E

(560 m): (2014) CAS 256842. 4.8247°N, 9.7702°E (494

m): (2013) CAS 253854-66. Females average 26.5 mm

SUL (24.0-28.5 mm, n = 4) and males average 25.4 mm

SUL (17.0-28.5 mm, n = 22). This species is superficial¬

ly similar to Afrixalus paradorsalis with regard to color

pattern, but can be distinguished by the smaller body size

of each sex. This lowland species was most abundant in

heavily disturbed habitat (ditches next to main highway),

but was also found in disturbed habitats close to second¬

ary forest. Amiet (1975, 2012) recorded A. dorsalis from

Nlohe at the southeastern extent of Mt. Kupe.

Afrixalus laevis (Ahl, 1930). Figure 6B-D; four speci¬

mens. 4.8498°N, 9.7718°E (513 m): (2013) CAS 253946,

254071-73. Female is 24.0 mm SUL and males average

20.2 mm SUL (19.5-21.0 mm, n = 3). Specimens were

found on the leaves of trees or palms (1-3 m high) over¬

hanging a small slowly moving stream. Though previ¬

ously emphasized by Amiet (2012), we also observed

stark differences in the coloration of individuals during

periods of nocturnal activity and inactivity during the

day. These differences are illustrated in the individuals

photographed at night after capture (Fig. 6A) and the fol¬

lowing morning (Fig. 6C, D). Afrixalus laevis is a wide¬

spread forest species previously recorded on both Mt.

Manengouba (Amiet 2012) and Mt. Kupe (Hofer et al.

1999).

Afrixalus paradorsalis Perret, 1960. Figure 6E, F; 25

specimens. 4.8497°N, 9.7719°E (510 m): (2014) CAS

256750-59, 256911-12. 4.8498°N, 9.7718°E (513 m):

(2013) CAS 253947-54, 253962-64, 253966-67. Fe¬

males average 33.3 mm SUL (31.0-36.0 mm, n = 5) and

males average 30.4 mm SUL (29.0-32.0 mm, n = 19).

Superficially similar to A. dorsalis , but distinguished by

the larger body size of each sex. Based on their dorsal

patterning, our records resemble A. paradorsalis para¬

dorsalis (Amiet 2012). We observed this widespread

lowland species breeding in permanent ponds (Fig. 2A)

or adjacent flooded grassy areas. Because of its frequent

confusion with A. dorsalis and A. equatorialis , the extent

of distribution of A. paradorsalis is not quite clear, but it

likely extends south into Equatorial Guinea and Gabon

(Lasso et al. 2002; Burger et al. 2006).

Hyperolius bolifambae Mertens, 1938. Figure 6G, H;

45 specimens. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256737^16, 256817-18, 256907-08. 4.8498°N,

9.7718°E (513 m): (2013) CAS 253883-912, 253977.

Females average 30.7 mm SUL (28.5-33.5 mm, n = 13)

and males average 24.8 mm SUL (23.0-26.5 mm, n =

32). This species occurred in high abundance at the per¬

manent ponds (Fig. 2A), and is easily distinguished from

all other sympatric Hyperolius species by the presence

of a heel spot. This species is sexually dichromatic, with

females (Fig. 6H) having bright red finger and toe tips, a

darker flank, a more vividly colored dorsum, and a con¬

spicuous ventral pattern with prominent white spots on

a black background. Males (Fig. 6G) sometimes display

the ventral spotted pattern, but never exhibit the black

background observed in the females. Rather, this color¬

ation ranges from whitish to dark grey. Hyperolius bo¬

lifambae is common at low elevations in southwestern

Cameroon as well as in far eastern Cameroon, though it

is rare in between (Amiet 2012). Amiet (1975) recorded

H. bolifambae from the nearby sites of Lala and Nhale,

as well as Mahole to the west of Mt. Kupe (Amiet 2012).

Hyperolius camerunensis Amiet, 2004. Figure 7A-D;

19 specimens. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256729-31, 256916. 4.8498°N, 9.7718°E (513 m):

(2013) CAS 253935, 253936^13, 254040-41, 254059-

60, 254078-79. Females average 29.0 mm SUL (24.5-

32.5 mm, n = 6) and males average 23.2 mm SUL (19.0-

25.0 mm, n = 13). This is a sexually dichromatic species.

Males resemble the pattern of male H. ocellatus by hav¬

ing pale dorsolateral stripes on a green background that is

sometimes speckled with a reddish coloration (Fig. 7A,

C), but they never display the pale triangle on the nose

that characterizes male H. ocellatus (see H. ocellatus

males in Fig. 9A, C). Females have a brownish-tan dor¬

sum that varies in the amount of red patterning outlined

by black and yellow, bright yellow ventral coloration,

and bright red on the thighs, fingers, toes, and webbing

(Fig. 7B, D). All specimens were found at shallow ponds.

Of the 13 males collected, a single male was found that

exhibited the typical female coloration (CAS 256916),

yielding a frequency of 7% for Phase F males. While re¬

corded on both Mt. Nlanoko and Mt. Manengouba (Her-

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 5. Representatives of arthroleptid, conrauid, bufonid, and hyperoliid frog taxa. (A) Trichobatrachus robustus. female (CAS

254069); (B) Trichobatrachus robustus, male (CAS 254070); (C) Trichobatrachus robustus, juvenile (CAS 253934); (D) Conraua

goliath, juvenile (CAS 256830); (E) Conraua rob ust a, juvenile (CAS 256821); (F) Sclerophrys superciliaris (photo voucher); (G)

Afrixalus doralis , female (CAS 256690); (H ) Afrixahis doralis, male (CAS 256691).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

rmann et al. 2005; Amiet 2012), this appears to be the

first record for 77 camerunensis at Mt. Kupe, and it is

approaching the southwestern extent of its range (Amiet

2012 ).

Hyperolius concolor (Hallowell, 1844). Figure 7E-H;

63 specimens. 4.8148°N, 9.7691°E (475 m): (2014)

CAS 256694-95. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256793-805, 256898-902, 256917-19. 4.8498°N,

9.7718°E (513 m): (2013) CAS 253913-32. 4.8551°N,

9.8188°E (560 m): (2014) CAS 256843-54. 4.8247°N,

9.7702°E (494 m): (2013) CAS 253868-75. Females av¬

erage 36.4 mm SUL (33.0^10.0 mm, n = 13) and males

average 28.3 mm SUL (24.5-33.5 mm, n = 46). This is

a sexually dichromatic species. Females have a bright

green dorsum, with yellow flanks and ventrum, and red

coloration occurs on the inside of the thighs and covers

the webbing, and sometimes discs, of the fingers and

toes (Fig. 7F). Males are generally dark to light brown

with patterning often involving a triangle between the

eyes with apex pointed backwards, with some amount of

banding or spots posteriorly (Fig. 7E). A Phase F male is

present, which is mainly green with some yellow ventral-

ly, but it does not exhibit any red coloration on the thighs,

or webbing or discs of the fingers and toes (Fig. 7G).

Of the 48 males collected, 11 were found with the typi¬

cal female coloration and two were in transition between

color morphs (Fig. 7H), yielding a frequency of 27% for

the Phase F male. This species was most abundant in the

grass-filled ditches of the banana plantation (Fig. 2E;

Amiet 2012), but some were collected at the ponds. This

lowland species is found in southwestern Cameroon and

was previously recorded at nearby Nhale (Amiet 1975).

Hyperolius dintelmanni (Lotters and Schmitz, 2004).

Figure 8A-C; 4 specimens. 4.8148°N, 9.7691 °E (475

m): (2014) CAS 256693. 4.8498°N, 9.7718°E (513

m): (2013) CAS 253991. 4.8551°N, 9.8188°E (560 m):

(2014) CAS 256840^11. Female measures 35.5 mm

SUL and males average 32.2 mm SUL (32.0-32.5 mm,

n = 3). This is a sexually dichromatic species. Phase J

males resemble the coloration of typical H. tubercula-

tus (Fig. 8A), though often with brighter green or yellow

markings on the dorsum, whereas females exhibit fine

reddish-pink points on a greyish-tan background, with

a pinkish-red ventral coloration (Fig. 8B). The Phase F

male displays a smaller number of reddish-pink points

on the dorsum, and bright reddish-pink coloration ven-

trally (Fig. 8C). Of the three males collected, only one

exhibited the Phase F coloration (Fig. 8C), yielding a

frequency of 33%, though our sample size is very small.

The female phase described for H. dintelmanni by Let¬

ters and Schmitz (2004) was not found at Mt. Kupe, and

they do not mention the Phase F we describe here. Our

measurements of males match those provided by Letters

and Schmitz (2004), but the female is substantially larger

(35.5 mm) than the described body size range for females

(27.3-29.8 mm). This species was collected mainly at the

fish ponds, but one male was found at the larger perma¬

nent ponds. Treated as a subspecies of H. tuberculatus

(Mocquard, 1897) by Amiet (2012), genetic analyses

have revealed that H. dintelmanni is a distinct lineage

that is sister to H. tuberculatus (Portik 2015; Bell et al.,

in prep). Previously recorded from the nearby Bakossi

Mountains, this is the first record of H. dintelmanni from

Mt. Kupe and at a much lower elevation (475-560 m)

than previously recorded (1,100-1,250 m; Lotters and

Schmitz 2004; Amiet 2012).

During July 17-18, 2013, DMP, GFMJ, and MTK

conducted additional survey work at the type locality for

this species (Edib, Cameroon: 4°57.578 N, 9°39.146 E,

-1,140 m elevation). Here, a series of 16 specimens of

H. dintelmanni were collected (three females, 13 males;

CAS 254135-37, 254156-68) matching the Phase J and

Phase F we have described for Mt. Kupe. Based on this

series, we also did not find any specimens that match the

females described by Lotters and Schmitz (2004). All of

the specimens from Edib are confirmed as genetically

identical to those collected at Mt. Kupe (Portik 2015;

Bell et al., in prep). The females collected at Edib are

also larger (31.3, 31.8, and 33.8 mm) than the body size

range described for the females described by Lotters and

Schmitz (2004) (27.3-29.8 mm). The range of the body

size for males we collected (n = 13, 29.6-32.5 mm, aver¬

age = 31.0 mm) matches the range described for this spe¬

cies (29.5-33.8 mm), and the Phase J coloration of these

males is consistent. However, the Phase F males collect¬

ed at Edib match the Phase F coloration we describe for

Mt. Kupe. The original description for the Phase F pre¬

sented by Lotters and Schmitz (2004) is not typical for

H. tuberculatus or any of its close relatives (77. viridifla-

vus and 77. marmoratus complexes), but this coloration

is characteristic of a clade recovered by Portik (2015)

consisting of 77. bolifambae , 77. camerunensis , 77. ocel-

latus , 77. riggenbachi, and the Hyperolius sp. presented

below. In these species, the females often have bright red

coloration on the thighs, fingers, and toes, yellow ventral

coloration, and complex patterning on the dorsum, which

is qualitatively very similar to the description and photo

presented by Lotters and Schmitz (2004).

Hyperoliids are notoriously difficult to identify due

to a conserved morphology and color variation result¬

ing from sexual dichromatism, but DNA barcoding is a

robust method for assigning species identifications. We

have collected a large series of 77. dintelmanni from two

localities (including the type locality), confirmed identi¬

fications through DNA barcoding, and found a consistent

Phase F that differs from the original description. Let¬

ters and Schmitz (2004) did not generate 16S data for the

type series of 77. dintelmanni , and assumed because the

male and female specimens were collected in close prox¬

imity they constitute a single species. Based on all the ev¬

idence presented, we propose that the females described

by Lotters and Schmitz (2004) are misidentified, and are

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 6. Representatives of hyperoliid frog taxa. (A) Afrixalus doralis, male (CAS 253854); (B) Afrixalus laevis, amplexus, night

coloration (CAS 254072-3); (C) Afrixalus laevis , male, day coloration (CAS 254073); (D) Afrixalus laevis , female, day coloration

(CAS 254072); (E) Afrixalusparadoralis, male (CAS 253954); (F) Afrixalus paradoralis , female (CAS 253953); (G) Hyperolius

bolifambae , male (CAS 256743); (H) Hyperolius bolifambae , female (CAS 256745).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

probably referable to the clade consisting of H. bolifam-

bae , H. camerunensis, H. ocellatus, H. riggenbachi, and

Hyperolins sp. We propose the following alternative hy¬

potheses to explain the females described by Lotters and

Schmitz (2004): 1) they represent a more variable Phase

F form of H. camerunensis, H. ocellatus, or H. riggen¬

bachi, though this is very doubtful, 2) they represent the

females of Hyperolius sp., for which we were unable to

collect females at Mt. Kupe, or 3) they represent an ad¬

ditional candidate species in this clade. Given H. dintel-

manni has been found at both sites, other Hyperolius oc¬

curring at Mt. Kupe may also have a similar geographic

distribution, including Hyperolius sp. Given the qualities

of color patterns in the H. viridiflavus/marmoratus group,

it is extremely unlikely the odd females at Edib represent

an additional Phase F of H. dintelmanni. Additional sur¬

vey work, coupled with DNA barcoding, will determine

which species these females belong to. The holotype of

H. dintelmanni is a Phase J male that is consistent with

males we collected, and we therefore maintain usage of

this species name for the lineage we have collected at Mt.

Kupe and Edib.

Hyperolius fusciventris Peters, 1876. Figure 8E-H; 81

specimens. 4.8148°N, 9.7691°E (475 m): (2014) CAS

256696-700. 4.8497°N, 9.7719°E (510 m): (2014) CAS

256720-24, 256760-91, 256811-15, 256903. 4.8498°N,

9.7718°E (513 m): (2013) CAS 253992-4004, 254005-

14, 254087. 4.8551°N, 9.8188°E (560 m): (2014) CAS

256831-37. 4.8247°N, 9.7702°E (494 m): (2013) CAS

253876-77. Females average 25.5 mm SUL (23.5-29.0

mm, n = 13) and males average 19.9 mm SUL (17.0—

22.0 mm, n = 65). This is a sexually dichromatic species.

Males are quite variable in coloration, ranging from plain

green background with narrow pale dorsolateral strips

starting behind the eye, to a green background heavily

speckled with brownish-black and broad yellow dorso¬

lateral stripes (Fig. 8E, G). Females have a green dor¬

sum with flanks with red, white, and sometimes yellow

vermiculations (Fig. 8E, F). These vermiculations often

continue to the canthus rostralis, and ventrum is unifonn

reddish-white. A Phase F male is present, which is simi¬

lar to the female form except in having no coloration on

the hands, feet, and thighs. Of the 65 males collected, one

exhibited the typical female coloration (Fig. 8H), yield¬

ing a frequency of 1.5% for the Phase F male. Hypero¬

lius fusciventris is a lowland species complex extending

across West Africa with its eastern extent in southwest¬

ern Cameroon (Amiet 2012). There are currently three

described subspecies, including H. f fusciventris, H. f

lamtoensis, and H. f burtoni, which occupy broad but al-

lopatric regions of Upper Guinea (Sierra Leone to west¬

ernmost Cote d’Ivoire, Cote d’Ivoire, and Ghana to Ni¬

geria, respectively), that are distinguishable on the basis

of female coloration. The form in Cameroon is actually

regarded as an undescribed subspecies in this complex,

H. f spp. (Schiotz 1999). The Cameroonian females are

distinguished by having a uniform reddish-white ven¬

trum, in contrast to the females of H. f burtoni (occur¬

ring in Nigeria), which are reported to have a white ven¬

trum with irregular black spots. Further molecular and

morphological work is required to clarify the taxonomic

status of this species complex, including the affinities of

the Cameroonian populations.

Hyperolius ocellatus Gunther, 1858. Figure 9A-D; 20

specimens. 4.8148°N, 9.7691°E (475 m): (2014) CAS

256701-06. 4.8497°N, 9.7719°E (510 m): (2014) CAS

256725-28, 256909-10. 4.8498°N, 9.7718°E (513

m): (2013) CAS 254057-58, 254074-77. 4.8551°N,

9.8188°E (560 m): (2014) CAS 256838-39. Females av¬

erage 30.2 mm SUL (27.5-32.0 mm, n = 9) and males

average 25.5 mm SUL (21.0-32.5 mm, n = 11). This is a

sexually dichromatic species. The male dorsum is green,

sometimes with red suffusion, and a conspicuous pale

yellow triangle is present on the nose along with dorso¬

lateral stripes (Fig. 9A, C). The females have a complex

color pattern. The dorsum is gray to brown with lightly

outlined fine black spots, and the flanks exhibit a white

background with large black vermiculations (Fig. 9A, B,

D). The thighs, fingers, and toes are bright red, and the

ventrum is bright yellow. No Phase F males have ever

been recorded for this species across its range, which

extends throughout western and southern Cameroon

through mainland Equatorial Guinea and Gabon (Lasso

et al. 2002; Burger et al. 2006; Amiet 2012). While previ¬

ously known from this region of Cameroon (Herrmann

et al. 2005; Amiet 2012), we believe this is the first re¬

cord on Mt. Kupe. Amiet (2012) described distinct color

patterns in females that correspond to allopatric regions,

but this variation was not assigned to distinct subspecies.

Molecular work is currently underway to investigate the

evolutionary relationships of populations of H. ocellatus

from throughout the known range (Bell et al., in prep).

Hyperolius sp. Figure 8D; 11 specimens. 4.8497°N,

9.7719°E (510 m): (2014) CAS 256732-36, 256816,

256904-06; 4.8498°N, 9.7718°E (513 m): (2013) CAS

253944—45. We only found males, which averaged 27.5

mm SUL (26.0-29.5 mm, n = 11). This species is quite

similar in coloration to males of H. camerunensis (Fig.

7A, C), the species to which we first attributed it in the

field. However, this undescribed species is larger and

non-overlapping in size (average 27.5 mm, range 26.0-

29.5 mm) when compared to H. camerunensis (average

23.2 mm, range 19.0-25.0 mm), and also has a distinc¬

tive red suffusion on the ventrum. This species overlaps

with H. riggenbachi in male body size (average 28.1,

25.5-30.8, n = 39; Portik 2015), is very similar in color¬

ation, and exceptionally difficult to distinguish. Based on

genetic data, it apparently occurs in direct sympatry with

H. riggenbachi in Bangoua (5.1763°N, 10.3487°E) (Por-

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 7. Representatives of hyperoliid frog taxa. (A) Hyperolius camerunensis, male (CAS 253939); (B) Hyperolius camerunensis,

female (CAS 253935); (C) Hyperolius camerunensis, male (CAS 253938); (D) Hyperolius camerunensis, female (CAS 255486);

(E) Hyperolius concolor, male (CAS 253873); (F) Hyperolius concolor, female (CAS 253928); (G) Hyperolius concolor, phase F

male (CAS 256898); (H) Hyperolius concolor, transitional male (CAS 256844).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

tik, unpubl. data). Further morphological and molecular

work is underway to identify this currently undescribed

species (Portik et al., in prep).

Kassina decorata (Angel, 1940). Figure 9E; one speci¬

men. 4.8498°N, 9.7718°E (513 m): (2013) CAS 253990.

This single female specimen (39.0 mm SUL) was col¬

lected near a flooded marsh at the edge of a plantation.

Amiet (2007,2012) discussed the considerable confusion

as to whether K. decorata is distinct from K. maculosa

(Sternfeld, 1917). This specimen is phenotypically simi¬

lar to Amiet’s (2007, 2012) description of K. decorata

with the dorsal spots coalesced into a single longitudinal

band as well as a supratympanic band extending postero-

ventrally around the tympanum (Fig. 9E). While the type

locality in the caldera of Mt. Manengouba is approxi¬

mately 1,950 m, Amiet (2007) reports specimens from

Santchou at 700 m. This specimen from near Manjo is

nearly genetically identical to specimens collected in the

caldera of Mt. Manengouba (Portik, unpubl. data) and

supports Amiet’s (2007) conclusion that phenotypically

distinct populations of K. decorata occur across a broad

elevational range.

Phlyctimantis leonardi (Boulenger, 1906). Figure 9F;

two specimens. 4.8498°N, 9.7718°E (513 m): (2013)

CAS 253978-79. Two male specimens measure 45.0

and 46.0 mm SUL. Both males were found calling in the

permanent ponds. Amiet (2007) records Phlyctimantis

from this region of Cameroon though the identity of

Cameroonian populations remains unclear because the

distinction between P. leondardi and P. boulengeri (Per-

ret 1986) remains confusing (Amiet 2007,2012; Gvozdik

and Kopecky 2012). In the two specimens from Manjo,

the skin is smooth with some finely colored tubercles,

resembling an intermediate form between P. leonardi

and P. boulengeri (Amiet 2012). Recent genetic data

demonstrate that these specimens of Phlyctimantis are

more closely related to P. leonardi of Gabon than they

are to P. boulengeri of Ghana (Portik 2015). We therefore

attribute this Cameroonian population to P. leonardii, but

acknowledge that both species might occur in Cameroon.

Petropedetidae

Petropedetes cameronensis Reichenow, 1874. Figure

9G; four specimens. 4.8500°N, 9.7673°E (538 m):

(2014) CAS 256887-90. A gravid female measures 42.0

mm SUL (CAS 256887), and other unsexed specimens

in the series measure 41.0 mm, 23.0 mm, and 23.5 mm

SUL, respectively. Specimens were found on a small

island surrounded by a fast flowing stream, alongside

Cardioglossa elegans. Petropedetes cameronensis oc¬

curs in lowland forests on Bioko Island and in western

Cameroon, including on Mt. Kupe where it extends up

to approxiately 1,200 m and co-occurs with several other

species of Petropedetes (Amiet 1975; Hofer et al. 1999;

Barej et al. 2010).

Petropedetes euskircheni Barej, Rodel, Gonwouo,

Pauwels, Bohme, and Schmitz, 2010. Figure 9H; three

specimens. 4.8339°N, 9.7783°E (490 m): (2013) CAS

254068; 4.8500°N, 9.7673°E: (2013) CAS 254066-67.

Two specimens exhibit tympanic papillae (a secondary

sexual characteristic of males) and measure 48.0 mm

(CAS 254067) and 60.0 mm (CAS 254068) SUL. The

specimen lacking tympanic papillae is 48.0 mm (CAS

254066). Many adults were found on the vertical moss-

covered rocks adjacent to a waterfall, and a small number

of individuals were found perched on large boulders im¬

mediately upstream of the waterfall zone. Petropedetes

euskircheni is known from -900-1,200 m on Mt. Kupe

(the type locality) and Mt. Nlonako, but our record ex¬

tends its elevational distribution down to -500 m.

Phrynobatrachidae

Phrynobatrachus africanus (Hallowell, 1858). Figure

10A, B; eight specimens. 4.8406°N, 9.7793°E (491 m):

(2014) CAS 256872. 4.8498°N, 9.7718 (513 m): (2013)

CAS 253983-89. Unsexed specimens average 24.1 mm

SUL (20.0-30.5 mm, n = 8). Phrynobatrachus africanus

is a common lowland forest species known from nearby

localities to the west (Amiet 1975) but not specifically on

Mt. Kupe (Hofer et al. 1999) and extending from Cam¬

eroon and Nigeria south through mainland Equatorial

Guinea and Gabon (Amiet 1975, 1978; Lasso et al. 2002;

Burger et al. 2006; Pauwels and Rodel 2007).

Phrynobatrachus auritus Boulenger, 1900. Figure

10C-E; seven specimens. 4.8148°N, 9.7691°E (475

m): (2014) CAS 256669-75. A gravid female is 35.0

mm SUL (CAS 256674), and males average 31.9 mm

SUL (28.5-31.0 mm, n = 4). This widespread species

is found in lowland forests from Nigeria and Cameroon

through mainland Equatorial Guinea, Gabon, and into

Democratic Republic of Congo (Amiet 1975; Lasso et

al. 2002; Burger et al. 2006; Pauwels and Rodel 2007;

Nagy et al. 2013).

Phrynobatrachus cornutus (Boulenger, 1906). Figure

10F; five specimens. 4.8406°N, 9.7793°E (491 m):

(2014) CAS 256868-71, 256873. A gravid female

specimen (CAS 256873) is 17.0 mm SUL, and unsexed

specimens average 12.6 mm (9.0-15.0 mm, n = 4). A

small lowland species that is found on Bioko Island

and extending through Cameroon and Central African

Republic and south into Gabon (Perret 1988; Burger et

al. 2006; Pauwels and Rodel 2006; Zimkus et al. 2010).

Phrynobatrachus cornutus is easily confused with P. cal-

caratus (Peters, 1863) but the Cameroonian populations

are genetically distinct from those that occur in more

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 8. Representatives of hyperoliid frog taxa. (A) Hyperohus dintelmanni, male (CAS 256841); (B) Hyperolius dintelmarmi ,

female (CAS 256693); (C) Hyperohus dintelmanni, male (CAS 253991); (D) Hyperolius sp., male (CAS 256733); (E) Hyperolius

fusciventris, amplexus (CAS 254007-08); (F) Hyperolius fusciventris, female (CAS 256815); (G) Hyperolius fusciventris, male

(CAS 256700); (H) Hyperolius fusciventris , phase F male (CAS 256699).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

western Africa and are referred to P. calcaratus (Zimkus

et al. 2010).

Phrynobatrachns sp. Figure 10G, H; three specimens.

4.8406°N, 9.7793°E (491 m): (2014) CAS 256714-16.

Two females measure 26.0 mm and 24.0 mm SUL (CAS

256714,256716), whereas the single male measures 21.5

mm SUL (CAS 256715). The specific identity of these

frogs remains unclear. Comparing the 16S DNA sequenc¬

es to those available in GenBank, the specimens are most

similar genetically (-6% uncorrected pairwise distance)

to a specimen of P. gutturosns (GenBank EU718724)

from Comoe National Park in Cote dTvoire.

Ptychadenidae

Ptychadena cf. aequiplicata. Four specimens. 4.8498°N,

9.7718°E (513 m): (2013) CAS 254024, 254054-56.

Three females were collected, measuring 65.0 mm

(CAS 254054), 50.0 mm (CAS 254055), and 65.0 mm

(CAS 254056) SUL. A single male was collected (CAS

254024) which is 51.0 mm SUL. These specimens were

found at the edge of a flooded marsh near a plantation.

DNA sequences of the mitochondrial 16S gene are genet¬

ically identical to GenBank sequences (e.g., AY517604)

identified as “alf. bibroni” by Dehling and Sinsch (2013).

The taxonomy of Ptychadena in Central Africa remains

confusing, including for species such as P. aequiplicata

and P. bibroni. These specimens are considerably larger

than noted for P. bibroni (34—48 mm SUL for males;

34-56 mm SUL for females) by Rodel (2000). We follow

Amiet (1975) and Herrmann et al. (2005) who identified

P. aequiplicata as present on and near Mt. Nlonako.

Ptychadena cf. mascareniensis . Two specimens.

4.8498°N, 9.7718°E (513 m): (2013) CAS 254083-84.

Both specimens are males, and measure 49.0 mm and

47.0 mm SUL, respectively. Both specimens were found

at the edge of a flooded marsh near a plantation. Geneti¬

cally, these two specimens are nearly identical to speci¬

mens (i.e., AF215408; AY517597) that belong to a clade

found in western and central Africa and referred to as “cf.

mascareniensis ” by Dehling and Sinsch (2013).

Ptychadena oxyrhynchus. Two specimens. 4.8406°N,

9.7793°E (491 m): (2014) CAS 256820, 256862. Both

specimens are males, and measure 57.0 mm and 56.0 mm

SUL, respectively. As currently recognized, this savanna

species is widespread across Africa (Rodel 2000; Chan-

ning 2001).

Rhacophoridae

Chiromantis rufescens Gunther, 1869. Figure 11 A, B;

Five specimens. 4.8497°N, 9.7719°E (510 m): (2014)

CAS 256708. 4.8498°N, 9.7718°E (513 m): (2013) CAS

254025-28. Four males were collected, each measuring

44.0 mm SUL (CAS 254025-28), along with a single

female measuring 72.0 mm SUL (CAS 256708). Foam

nests were observed on the vegetation pictured in Fig.

2A, and adults were found around the permanent ponds

near a plantation. This lowland forest species is wide¬

ly distributed across western and central Africa (Noble

1924; Lasso et al. 2002; Burger et al. 2006; Hillers and

Rodel 2007) including Cameroon (Amiet 1975, 1978,

1983; Hernnann et al. 2005) and it was previously docu¬

mented on Mt. Kupe by Hofer et al. (1999).

SQUAMATA- LIZARDS

Agamidae

Agama lebretoni Wagner, Barej and Schmitz, 2009.

Four specimens. 4.8551°N, 9.8188°E (560 m): (2014)

CAS 256858. 4.8247°N, 9.7702°E (494 m): (2013) CAS

253878-80. One adult female was collected (120.0 mm

SVL, 167.0 mm tail length; CAS 253879) along with

three juveniles ranging from 49.5 mm to 73.5 mm SVL.

They were abundant on the houses and other artificial

structures in the town of Manjo. While only recently

described, this species is widely distributed in western

Cameroon and also found on Bioko Island and in Gabon

(Chirio and LeBreton 2007; Wagner et al. 2009).

Gekkonidae

Cnemaspis spinicollis (Muller, 1907). Figure 11C; one

specimen. 4.8506°N, 9.7671°E (541 m): (2014) CAS

256886. An adult female (53.0 mm SVL, 57.0 mm tail

length) was collected while active at night on rocks adja¬

cent to a large stream. Cnemaspis spinicollis is found in

forests of western and central Africa (Bauer et al. 2006;

Chirio and LeBreton 2007) and previously documented

on Mt. Kupe (Hofer et al. 1999).

Lacertidae

Poromera fordii (Hallowell, 1857). Figure 11D; one

specimen. 4.8498°N, 9.7718°E (513 m): (2013) CAS

254085. An adult male of this colorful species was

collected while sleeping at night in the vegetation of a

permanent pond. It measures 46.0 mm SVL with a tail

length of 112.0 mm. This species occurs in forests across

a broad elevation range (near sea level to -1,300 m) in

Central Africa, including Cameroon, Gabon, Equatorial

Guinea (Lasso et al. 2002; Chirio and LeBreton 2007;

Pauwels et al. 2008). It was not recorded on Mt. Kupe by

Hofer et al. (1999).

Scincidae

Panaspis breviceps (Peters, 1873). Figure 11E; one spec¬

imen. 4.8551°N, 9.8188°E (560 m): (2014) CAS 256855.

Specimen measures 52.5 mm SVL with 81.5 mm tail

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 9. Representatives of hyperoliid and petropedetid frog taxa. (A) Hyperolius ocellatus, amplexus (CAS 254057-58); (B)

Hyperolius ocellatus, female (CAS 256838); (C) Hyperolius ocellatus, male (CAS 256704); (D) Hyperolius ocellatus, female (CAS

254076); (E) Kassina decorata (CAS 253990); (F) Phlyctimantis leonardi, male (CAS 253979); (G) Petropedetes cameronensis

(CAS 256887); (H) Petropedetes euskircheni, male (CAS 254068).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

length. This specimen was collected while active on the

ground near fish ponds. Panaspis breviceps is a common

lowland species found across Central Africa (Chirio and

LeBreton 2007; Pauwels et al. 2008). It was not recorded

on Mt. Kupe by Hofer et al. (1999).

Trachylepis affinis (Gray, 1838). Figure 1 IF; three speci¬

mens. 4.8148°N, 9.7691°E (475 m): (2014) CAS 256707.

4.8551°N, 9.8188°E (560 m): (2014) CAS 256856-57.

The three specimens differ in size: CAS 256856, 48.5

mm SVL + 90.5 mm tail length; CAS 256707, 61.0 mm

SVL +119.0 mm tail length; CAS 256857, 80.0 mm SVL

+ 125.0 mm tail length. The morphological identification

of these specimens was confirmed by a comparison of

16S data with that of Allen (2015). Specimens were col¬

lected on agricultural plantations near to either ponds or

small streams. A common species, T. affinis occurs across

a broad elevational range (sea level to -1,200 m) across

western and Central Africa (Chirio and LeBreton 2007),

though Hofer et al. (1999) previously recorded only one

individual on Mt. Kupe at 900 m.

Trachylepis maculilabris (Gray, 1845). Three specimens.

4.8497°N, 9.7719°E (510 m): (2014) CAS 256792.

4.8247°N, 9.7702°E (494 m): (2013) CAS 253881-82.

The three specimens differ in size: CAS 253881, 47.0

mm SVL + 80.0 mm tail length; CAS 253882, 52.5 mm

SVL + 102.0 mm tail length; CAS 256792, 89.0 mm SVL

+ 154.0 mm tail length. The morphological identification

of these specimens was confirmed by a comparison of

16S data with that of Allen (2015). Specimens were col¬

lected near roadside ditches (CAS 253881-82) or on ag¬

ricultural plantations (CAS 256792). Trachylepis maculi¬

labris is among the most widely distributed skink species

in Africa and occurs across an extremely broad eleva¬

tional range in Cameroon (sea level to >2,550 m), but

surprisingly was not previously recorded on Mt. Kupe by

Hofer et al. (1999).

Varanidae

Varanus niloticus (Linnaeus, 1766). 4.8506°N, 9.7671°E

(541 m): (2014) DMP 1698. A juvenile was collected at

night, and was found sleeping in a tree overhanging a

larger section of the stream in secondary forest illustrated

in Fig. 2D. This locality is within the described range

for V. ornatus , but based on a large molecular analysis

with extensive geographic sampling Dowell et al. (2016)

recently proposed V. ornatus be recognized as a synonym

of V. niloticus.

SQUAMATA- SNAKES

Colubridae

Dipsadoboa duchesnii (Boulenger, 1901). One specimen.

4.8498°N, 9.7718°E (513 m): (2013) CAS 254086. We

collected a male specimen (330.0 mm SVL + 90.0 mm

tail length) near a marsh on a plantation. A lowland spe¬

cies, D. duchesnii has a broad distribution in the forests

of Central Africa (Chirio and LeBreton 2007; Trape and

Balde 2014). Hofer et al. (1999) report both D. unicolor

and one unidentified Dipsadoboa from Mt. Kupe, but it

is unclear if the latter might correspond to D. duchesnii.

Grayia ornata (Bocage, 1866). Figure 11H; one speci¬

men. 4.8551°N, 9.8188°E (560 m): (2014) CAS 256859.

We collected a single specimen (415.0 mm SVL + 150.0

mm tail length) at a fish pond near a fast-flowing stream.

Being piscivorous, G. ornata is frequently encountered in

or near water in Central African lowland forests (Chirio

and LeBreton 2007), but was not recorded on Mt. Kupe

by Hofer et al. (1999).

Hapsidophrys smaragdinus (Schlegel, 1837). Figure

11G; two specimens. 4.8551°N, 9.8188°E (560 m):

(2014) CAS 256860-61. CAS 256860 is 667.0 mm in

total length (425.0 mm SVL + 242.0 mm tail length);

CAS 256861 is 402.0 mm in total length (267.0 mm SVL

+ 135.0 mm tail length). Both specimens were collected

near fish ponds on plantations near Manjo. A widespread

African snake, H. smaragdinus is common in forests but

also in plantations and near human habitation (Chirio and

LeBreton 2007).

Discussion

Our ten days of survey work in the foothills of Mt. Kupe

produced a total of 48 species, including 38 amphibian

and 10 reptile species (Table 2). The amphibians sampled

belong to eight families: Arthroleptidae (12 species),

Bufonidae (two species), Conrauidae (two species), Hy-

peroliidae (12 species), Petropedetidae (two species),

Phrynobatrachidae (four species), Ptychadenidae (three

species), and Rhacophoridae (one species). We encoun¬

tered seven lizard species in five families (Agamidae,

Gekkonidae, Lacertidae, Scincidae, Varanidae), and

three snake species in the family Colubridae. Although

our work was not conducted using a systematic sam¬

pling technique, we observed qualitative differences in

the relative abundances of species within these families.

We collected 471 amphibian specimens, and of this total,

304 specimens are hyperoliid frogs (64% of amphibian

specimens), 118 are arthroleptid frogs (25% of amphib¬

ian specimens), and 49 specimens are split among the

remaining six frog families (11% of amphibian speci¬

mens). Among the hyperoliid species we collected, sev¬

eral were abundant ( Afrixalus dorsalis , A. paradorsalis,

Hyperolius bolifambae , H. camerunensis, H. concolor,

H. fusciventris, H. ocellatus ) but a few species were rare¬

ly encountered ( Afrixalus laevis , Hyperolius dintelman-

ni , Kassina decorata, Phlyctimantis leonardi). In con¬

trast, only three arthroleptid species had relatively high

abundances ( Arthroleptis poecilonotus, Leptopelis rufus.

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Table 2. Checklist of the amphibians and reptiles of the greater Mt. Kupe region. Comparisons to Amiet (1975) are based on three

sites (Lala, 480 m; Mahole, 300-350 m; Nlohe, 350 m) and only include amphibians. Information from S & E comes from Schmitz

et al. (1999, 2000) and Euskirchen et al. (1999, 2000) for surveys on Mt. Kupe, though elevations for the records on Mt. Kupe are

not always noted.

Present

study

<1,000 m

Amiet (1975)

300-480 m

Hofer et al.

(1999)

900-1,500 m

Hofer et al.

(1999)

1,500-2,000 m

Hofer et al.

(1999)

elevation not

reported

S&E

Amphibia

Anura

Arthroleptidae (26)

Arthroleptis adelphus

Arthroleptis perreti

—

Arthroleptis poecilonotus

—

Arthroleptis variabilis

—

Astylosternus diadematus

—

Astylosternus laurenti

—

Astylosternus montanus

—

Astylosternus perreti

—

Cardioglossa elegans

—

Cardioglossa gracilis

—

Cardioglossa leucomystax

—

Cardioglossa nigromaculata

—

Cardioglossa venusta

—

Leptodactylodon bicolor

—

Leptodactylodon or n at us

—

Leptodactylodon ovatus

—

Leptopelis aubryi

—

Leptopelis aubryioides

—

Leptopelis boulengeri

—

Leptopelis brevirostris

—

Leptopelis calcaratus

—

Leptopelis modestus

—

Leptopelis rufus

—

Nyctibates corrugatus

—

Scotobleps gabonicus

—

Trichobatrachus robustus

—

850 m

Bufonidae (9)

Sclerophrys gracilipes

Sclerophrys regularis

—

Sclerophrys superciliaris

—

Sclerophrys tuberosa

—

Nectophryne afra

—

Nectophryne batesi

—

Werneria mertensiana

—

Werneria preussi

—

Wolterstorffina parvipalmata

—

Conrauidae (3)

Conraua crassipes

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

Table 2 (Continued). Checklist of the amphibians and reptiles of the greater Mt. Kupe region. Comparisons to Amiet (1975) are

based on three sites (Lala, 480 m; Mahole, 300-350 m; Nlohe, 350 m) and only include amphibians. Information from S & E comes

from Schmitz et al. (1999, 2000) and Euskirchen et al. (1999, 2000) for surveys on Mt. Kupe, though elevations for the records on

Mt. Kupe are not always noted.

Present

study

<1,000 m

Amiet (1975)

300-480 m

Hofer et al.

(1999)

900-1,500 m

Hofer et al.

(1999)

1,500-2,000 m

Hofer et al.

(1999)

elevation not

reported

S&E

Conraua goliath

—

Conraua robusta

—

Hyperoliidae (18)

Acanthixalus spin os us

945 m

Afrixalus dorsalis

—

Afrixalus lac tens

—

Afrixalus laevis

—

Afrixalus paradorsalis

—

Alexteroon obsetricans

—

Arlequinus krebsi

—

Hyperolius bolifambae

—

Hyperolius camerunensis

—

Hyperolius concolor

—

Hyperolius dintelmanni

—

Hyperolius fusciventris

—

Hyperolius guttulatus

—

Hyperolius ocellatus

—

Hyperolius sp.

—

Kassina decorata

—

Opisthothylax immaculatus

—

Phlyctimantis leonardi

Petropedetidae (5)

—

Petropedetes cameronensis

—

Petropedetes euskircheni

—

Petropedetes newtoni

—

Petropedetes parkeri

—

Petropedetes perreti

—

Phrynobatrachidae (6)

Phrynobatrachus africanus

Phrynobatrachus auritus

—

Phrynobatrachus cornutus

—

Phrynobatrachus cricogaster

—

Phrynobatrachus sp.

—

Phrynobatrachus werneri

Ptychadenidae (3)

—

Ptychadena cf. aequiplicata

—

Ptychadena cf. mascariensis

—

Ptychadena oxyrhynchus

Ranidae (1)

—

Amnirana albolabris

—

Rhacophoridae (1)

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Table 2 (Continued). Checklist of the amphibians and reptiles of the greater Mt. Kupe region. Comparisons to Amiet (1975) are

based on three sites (Eala, 480 m; Mahole, 300-350 m; Nlohe, 350 m) and only include amphibians. Information from S & E comes

from Schmitz et al. (1999, 2000) and Euskirchen et al. (1999, 2000) for surveys on Mt. Kupe, though elevations for the records on

Mt. Kupe are not always noted.

Present

study

<1,000 m

Amiet (1975)

300-480 m

Hofer et al.

(1999)

900-1,500 m

Hofer et al.

(1999)

1,500-2000 m

,Hofer et al.

(1999)

elevation not

reported

S&E

Chiromantis rufescens

—

Reptilia

Squamata (Lizards)

Agamidae (1)

Agama lebretoni

Chamaeleondidae (4)

Trioceros montium

Trioceros pfefferi

—

Trioceros quadricornis

—

Rhampholeon spectrum

—

Gekkonidae (4)

Cnemaspis koehleri

Cnemaspis spinicollis

—

Hemidactylus echinus

—

Hemidactylus fas ciat us

—

Lacertidae (2)

Adolf us africanus

Poromera fordii

—

Scincidae (9)

Lacertaspis chriswildi

Lacertaspis rohdei

—

Lepidothyris fernandi

—

Leptosiaphos sp. A

—

Leptosiaphos sp. B

—

Leptosiaphos sp. C

—

Panaspis breviceps

—

Trachylepis affinis

—

Trachylepis maculilabris

—

Varanidae (1)

Varan us nil otic us

—

Squamata (Snakes)

Boidae (1)

Calabaria reinhardti

Colubridae (6)

Dipsadoboa duchesnii

Dipsadoboa unicolor

—

Grayia inornata

—

Hapsidophrys smaragdinus

—

Rhamnophis aethiopissa

—

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

Table 2 (Continued). Checklist of the amphibians and reptiles of the greater Mt. Kupe region. Comparisons to Amiet (1975) are

based on three sites (Lala, 480 m; Mahole, 300-350 m; Nlohe, 350 m) and only include amphibians. Information from S & E comes

from Schmitz et al. (1999, 2000) and Euskirchen et al. (1999, 2000) for surveys on Mt. Kupe, though elevations for the records on

Mt. Kupe are not always noted.

Present

study

<1,000 m

Amiet (1975)

300-480 m

Hofer et al.

(1999)

900-1,500 m

Hofer et al.

(1999)

1,500-2,000 m

Hofer et al.

(1999)

elevation not

reported

S&E

Toxicodryas pulverulenta

Elapidae (2)

—

Dendroaspis jamesoni

—

Naja melanoleuca

Lamprophiidae (4)

—

Bothrolycus ater

—

Buhoma depressiceps

—

Chamaelycus fasciatus

—

Gonionotophis guirali

Viperidae (2)

—

Athens squamiger

—

Bids gabonica

—

850 m

Scotobleps gabonicus ), but we detected several arthro-

leptid species through the collection of only one or two

specimens ( Cardioglossa leucomystax , Leptodactylodon

ovatus, Leptopelis boulengeri, L. calcaratus, Nyctibates

corrugatus). By combining our survey results with those

of Hofer et al. (1999, 2000) as well as previous work by

Amiet (1975) and others, we have produced a checklist

of 108 species of reptiles and amphibians for the greater

Mt. Kupe region (Table 2). This includes 72 frog species,

21 lizard species, and 15 species of snakes.

Prior to our work, Hofer et al. (1999) perfonned tran¬

sects of higher elevation regions (900-2,000 m) and

some opportunistic lower elevation sampling of Mt.

Kupe. Across this elevation range, they documented a to¬

tal of 57 species, including 33 amphibian and 25 reptile

species (Table 2). Our survey work at lower elevations

uncovered an additional 30 species of frogs not detected

by Hofer et al. (1999), which can largely be explained by

the availability of suitable breeding habitat. The tempo¬

rary pools and permanent ponds present at lower eleva¬

tion sites are likely facilitating the occurrence of species

of Sclerophrys, Ptychadena, Phrynobatrachus , and Lep¬

topelis, as well as the hyperoliid frogs (A frixahis, Hyper-

olins, Kassina , Phlyctimantis). We collected twelve spe¬

cies of hyperoliids during our work, eleven of which were

not found previously at higher elevations by Hofer et al.

(1999). These particular species use lentic water systems

for breeding, features that are rare or absent from higher

elevation sites around Mt. Kupe due to increased slope.

The only amphibian species detected during all surveys

and across several elevational gradients are forest stream

(lotic) breeders ( Cardioglossa elegans, Leptopelis rufus,

Trichobatrachus robustus, Conraua robusta, Afrixalus

laevis, Petropedetes cameronensis ), suggesting this is

one of the few habitats present across elevation zones.

Several Cameroonian herpetological surveys have fo¬

cused on montane regions including Mt. Cameroon

(Gonwouo et al. 2007), Mt. Nlonako (Herrmann et al.

2005a, 2005b), Takamanda Forest Reserve (LeBreton et

al. 2003), Mt. Oku (Ineich et al. 2015), and Tchabal Mba-

bo (Herrmann et al. 2006). Surveys in lower elevation

regions have also been conducted, including in Korup

National Park (Lawson 1993) and Dja Reserve (LeBre¬

ton 1999). The focus of a majority of these studies has

been on reptiles, with few researchers reporting results

for both reptiles and amphibians. Though these surveys

each differ in collection technique, sampling effort, and

geographic scope, they do provide a baseline estimate for

herpetological diversity across several regions. Studies

providing data for both amphibians and reptiles include

survey work at Mt. Nlonako (182 total species), Korup

National Park (171 total species), Tchabal Mbabo (30 to¬

tal species), and our present summary of Mt. Kupe (108

total species).

In Cameroon the highest documented reptile diversity

occurs at Mt. Nlonako, with over 89 species reported

(Herrmann et al. 2005b), followed by 86 species for Mt.

Cameroon (Gonwouo et al. 2007), 83 species for Korup

National Park (Lawson 1993), 80 species for Dja Re¬

serve (LeBreton 1999), 71 species for Takamanda Forest

Reserve (LeBreton et al. 2003), 50 species for Mt. Oku

(Ineich et al. 2015), 36 species for Mt. Kupe (present

study), and 15 species for Tchabal Mbabo (Herrmann et

al. 2006). A large proportion of this diversity can be at¬

tributed to snakes, which are often difficult to encounter

during visual surveys. Our study and the study of Hofer

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Portik et al.

Fig. 10. Representatives of phrynobatrachid frog taxa. (A) Phrynobatrachus africanus (CAS 256872); (B) Phrynobatrachus

africanus (CAS 253989); (C) Phrynobatrachus auritus, male (CAS256670); (D) Phrynobatrachus auritus , male (CAS 256669);

(E) Phiynobatrachiis auritus , female (CAS 256674); (F) Phiynobatrachus cornutus (representative from series CAS 256868-71);

(G) Phrynobatrachus sp. (CAS 256714); (H) Phrynobatrachus sp. (CAS 256715).

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

et al. (1999) did not employ funnel or pit fall traps, and

as a result we believe the 15 species of snakes currently

documented for Mt. Kupe is not representative of the true

snake diversity. The lizard diversity of Mt. Kupe includes

at least 21 species, which is comparable to the diversity

detected at Mt. Oku (18 species; Ineich et al. 2015), Mt.

Cameroon (27 species; Gonwouo et al. 2007), and Mt.

Nlonako (22 species; Herrmann et al. 2005b). The high¬

est amphibian diversity in Cameroon also occurs at Mt.

Nlonako, with 93 amphibian species recorded (Herrmann

et al. 2005a), followed by 89 species for Mt. Manengou-

ba (Hirschfeld et al. 2016), 88 species for Korup National

Park (Lawson 1993), 72 species for Mt. Kupe (present

study), and 15 amphibian species for Tchabal Mbabo

(Herrmann et al. 2006). Overall, with at least 72 amphib¬

ian and 36 reptile species, the greater area of Mt. Kupe

is an important center of biodiversity in Cameroon and

is comparable in richness to other montane biodiversity

hotspots.

Given the close geographic proximity of Mt. Kupe to

Mt. Nlonako, we expect strong biogeographic ties be¬

tween these mountains, particularly of the lower eleva¬

tion fauna. The thorough sampling of Mt. Nlonako al¬

lows an initial biogeographic comparison of species we

have compiled for Mt. Kupe. We find that at least 50 am¬

phibians are found in both sites, along with 21 species of

reptiles, which represents 74% of the species document¬

ed for Mt. Kupe. Many of these shared reptile species are

found at lower elevations, though some are only present

at higher elevations (species of Leptosiaphos , Trioceros ,

Hemidactylus echinus). Of the 50 amphibians, close to

40 are species found at lower elevations (in the following

genera: Afrixalus , Chiromantis, Hyperolius, Leptopelis ,

Nectoprhyne, Nyctibates, Petropedetes, Phrynobatra-

chns, Ptychadena, Sclerophrys , Scotobleps , Trichobatra-

chus). There are fewer higher elevation species found in

common on both mountains, but include Afrixalus lac-

tens , Arthroleptis peered, Astylostermis montanus, Car-

dioglossa venusta, Leptodactylodon bicolor , Leptodac-

tylodon ornatus, Leptopelis modestus, Phrynobatrachus

cricogaster, Sclerophrys tuberosa , and Woltersorffina

parvipalmata. The close proximity of Mt. Kupe to other

mountains (Mt. Manengouba, Mt. Nlonako), combined

with historical fluctuations in climate, has produced an

island-like geographic range distribution for many of

these higher elevation taxa.

Acknowledgments. —The Cameroon Ministry of For¬

ests and Wildlife (MINFOF) and Ministry of Scientific

Research and Innovation (MINRESI) provided necessary

permits for conducting research and exportation. Field¬

work was supported by National Science Foundation

grant DEB #1202609 to DCB, and under the approval of

the Institutional Animal Care and Use Committee (2014-

2) at the California Academy of Sciences. We thank Jens

Vindum for cataloguing all specimens at the California

Academy of Sciences.

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Amphib. Reptile Conserv.

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Amphibians and reptiles in the foothills of Mount Kupe, Cameroon

Daniel M. Portik is currently a postdoctoral researcher at the University of Texas at Arlington, USA. He

received a B.S. from John Carroll University (2007), M.S. from Villanova University (2009), and Ph.D. at the

Museum of Vertebrate Zoology in the Integrative Biology Department at the University of California, Berkeley

(2015). His dissertation focused on the molecular systematics and morphological evolution of hyperoliid

frogs in tropical Africa, but he is broadly interested in biogeography, phylogeography, trait evolution, and the

natural history of a variety of African reptiles and amphibians.

Gregory F.M. Jongsma is a Ph.D. student at the Florida Museum of Natural History at the University

of Florida, USA. He received a B.S. from Acadia University (2010) and a M.S. from San Francisco State

University (2014). For his dissertation he is taking a comparative phylogeographic approach to explore the

diversification of frogs in Central Africa. During fieldwork he is often fueled by spaghetti omelettes and

Fighter.

Marcel T. Kouete is an amphibian biologist at the Cameroon Herpetology Conservation Biology Foundation

in Yaounde, Cameroon (Central Africa). He received a M.S. in aquaculture from the University of Liege,

Belgium in 2010 and a M.S. in Ecology, Evolution and Conservation from San Francisco State University

in 2015. He is starting a Ph.D. program in 2017 at the School of Natural Resources and Environment of

the University of Florida. His work will be assessing the impact of land use change on the biology and

morphology of amphibian communities in Cameroon.

Lauren A. Scheinberg is a Curatorial Assistant in the Department of Herpetology at the California Academy

of Sciences, USA. She received her B.S. from the University of California Santa Cruz (2006) and her M.S.

from San Francisco State University (2015). During fieldwork she thrives on powdered coffee and formalin

fumes.

Brian Freiermuth works in the museum and consulting fields, with a career focus on amphibians and reptiles.

He received his B.S. from California State University at Monterey Bay (2005) and a M.S. from San Francisco

State University (2015). He is driven by his desire to see the world and all the frogs and salamanders it has

to offer.

Walter P. Tapondjou is a graduate student in the department of Ecology and Evolutionary Biology at the

University of Kansas, USA. He received his B.S. in Animal Biology and his M.S. in Applied Zoology from

the University ofYaounde (in 2007 and 2011, respectively). His research is on intraspecific gene flow of the

mountain endemic chameleon from the Cameroon volcanic line.

David C. Blackburn is the Associate Curator of Herpetology at the Florida Museum of Natural History

at the University of Florida, USA. He received a BA from the University of Chicago (2001) and a Ph.D.

from Harvard University (2008). His research focuses on the diversity and evolution of frogs. His favorite

Cameroonian frogs include Leptodactylodon ovatus , Leptopelis brevirostris , and Trichobatrachus robustus.

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el 31

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [General Section]: 1-10 (e122).

The first female specimen of the poorly known Arfak Stout¬

tailed Snake, Calamophis sharonbrooksae Murphy, 2012

(Serpentes: Colubroidea: Homalopsidae), from the Vogelkop

Peninsula of Indonesian West New Guinea, with comments

on the taxonomic history of primitive homalopsids

Mark O’Shea and 2 Hinrich Kaiser

1 Faculty of Science and Engineering, University of Wolverhampton, Wulfruna Street, Wolverhampton, WV1 1LY, UNITED KINGDOM and West

Midland Safari Park, Bewdley, Worcestershire DY12 ILF, UNTIED KINGDOM 2 Department of Biolog) ', Victor Valley College, 18422 Bear Valley

Road, Victorville, California 92395, USA and Department of Vertebrate Zoology’, National Museum of Natural History, Smithsonian Institution,

Washington, DC 20013, USA

Abstract.—The recent resurrection of Calamophis Meyer, 1874, type species C. jobiensis, from the

synonymy of Brachyorrhos Kuhl in Schlegel, 1826, and the description of three new species of Cal¬

amophis, have changed concepts of homalopsid diversity in the Vogelkop Peninsula of West New

Guinea. Both Brachyorrhos and Calamophis are now accepted to comprise four species each and

are considered representatives of a unique fangless, nonvenomous, terrestrial to semi-fossorial,

homalopsid lineage. With the original and only specimen of C. jobiensis lost, the genus Calamophis

is now characterized by only six specimens (4 SS, 2 $$), comprising holotypes and paratypes of

the remaining three species; in each case the species is defined only by specimens of a single

sex. We here present the description of the first female specimen of C. sharonbrooksae, the larg¬

est specimen of the genus discovered so far, which exhibits a slightly longer body (96% of SVL

vs. 91%) and a higher ventral scale count (158 vs. 149 or 150) than the two males, combined with a

significantly shorter tail (4.4% of total length vs. 8.6%) and a lower subcaudal scale count (12 pairs

vs. 17 or 19 pairs). This is the first time both sexes of a Calamophis species have been available

for comparison. The specimen is also the first mainland Papuan Calamophis documented outside

the administrative boundaries of the Manokwari Residency, suggesting a wider distribution for the

genus than previously thought.

Keywords: Homalopsidae, Calamophis sharonbrooksae , Brachyorrhos , West Papua Province, Vogelkop Peninsula

Citation: O’Shea M and Kaiser H. 2016. The first female specimen of the poorly known Arfak Stout-tailed Snake, Calamophis sharonbrooksae Mur¬

phy, 2012 (Serpentes: Colubroidea: Homalopsidae), from the Vogelkop Peninsula of Indonesian West New Guinea, with comments on the taxonomic

history of primitive homalopsids. Amphibian & Reptile Conservation 10(2) [General Section]: 1-10 (e122).

cialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided

the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized publication

credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website <amphibian-

reptiie-conservation.org>.

Received: 15 March 2016; Accepted: 06 May 2016; Published: 26 August 2016

Introduction

Calamophis Meyer, 1874 was created as a monotypic

subgenus of Calamaria H. Boie in F. Boie, 1827 with

type species C. jobiensis Meyer, 1874, but only four

years later Calamophis was synonymized with Brachyor¬

rhos Kuhl in Schlegel, 1826 by Peters and Doria (1878).

Boulenger (1893) later synonymized B. jobiensis with

B. albus (Linnaeus, 1758). This resulted in a monotypic

Brachyorrhos, with no further mention of Calamophis

for over a century.

Correspondence. Emails: 1 oshea@markoshea.info (corresponding author); 2 hinrich.kaiser@vvc.edu

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

O’Shea and Kaiser

The phylogenetic position of Brachyorrhos :

Beyond any issue with Calamophis , the taxonomic posi¬

tion of Brachyorrhos within the Colubroidea (advanced

snakes) remained uncertain, and the genus appeared as

part of the Coronellae (Boie, 1827), Calamaridae (Gun¬

ther, 1858), Colubridae (Boulenger, 1893; Williams and

Wallach, 1989), and Natricidae (Dowling and Duellman,

1978). McDowell (1987) proposed that despite its ter¬

restrial and fossorial existence, combined with its lack

of fangs, Brachyorrhos belonged to the aquatic, rear-

fanged, piscivorous-carcinophagus Homalopsidae, with

which it shared viviparity and morphological similari¬

ties of the hemipenes, skull, and vertebrae. Although this

placement was generally accepted (Burbrink and Crother,

2011; Zaher et al., 2009), Brachyorrhos was considered

incertae sedis within the Homalopsidae (Lawson et al.,

2005; Murphy, 2007) until its phylogenetic position, as

the sister-taxon to all the aquatic, rear-fanged taxa, was

established by molecular analysis (Murphy et al., 2011).

Brachyorrhos is now viewed as representing a basal, ter-

restrial-fossorial, fangless, nonvenomous, vermivorous

lineage within the Homalopsidae (Murphy, 2012; Mur¬

phy and Voris, 2014; Wallach et al., 2014).

Calamophis or Brachyorrhos ? Finally, after more

than a century of what Murphy et al. (2012) referred to as

“a deeply entwined and confused nomenclatural history,”

Calamophis was resurrected (Murphy, 2012) for a group

of snakes from the Vogelkop (Bird’s Head) Peninsula

and Schouten Islands of northwestern New Guinea. The

snakes included sharing a series of pholidotic character¬

istics, including a single internasal, a single postocular,

and 19 dorsal scale rows that reduce to 17 anterior to

the vent. Brachyorrhos was reserved for snakes possess¬

ing paired intemasals, paired postoculars, and 19 scales

at midbody without reduction in number anterior to the

vent. Brachyorrhos sensu stricto was therefore confined

to the Moluccan Islands (Maluku and North Maluku

Provinces, Indonesia) and is no longer monotypic, owing

to the resurrection of two previously synonymized taxa

and the description of a new species (Murphy, 2012).

Brachyorrhos therefore comprises B. albus, a species

found on Seram, Ambon, Haruku, Nusa Laut, Saparua,

and Pulau Bisa; B. gastrotaenius (Bleeker, 1860) on

Burn; B. raffrayi (Sauvage, 1879) on Ternate; and B. wal-

lacei (Murphy et al., 2012) on Halmahera.

The distribution of Brachyorrhos : Murphy et al.

(2012) also provided tantalizing records for “unresolved

Brachyorrhos ” that may include potential second species

for both Seram and Buru. Brachyorrhos of undetermined

taxonomy have also been reported from Morotai (de

Jong 1928), Bacan (= Batjan) (Bleeker 1857; Boettger

1903), the Banda Islands (Boettger 1892; 1903), the Aru

Islands (Peters and Doria 1878), and Kofiau Island, in the

Raja Ampat Islands, Raja Ampat Regency, West Papua

Province, West New Guinea (Murphy et al., 2012). We

consider reports of Brachyorrhos for the Lesser Sunda

island of Timor (Boettger, 1903; Boulenger, 1893; de

Rooij 1917; fide Gunther 1858; Iskandar and Colijn,

2002; Welch 1988) doubtful, especially since we have

conducted an intensive herpetological survey of Timor-

Leste, comprising ten phases in 2009-2014 (Kaiser et al.,

2011; Kaiser et al., 2013; O’Shea et al., 2012; O’Shea

et al., 2015b; Sanchez et al., 2012). We also discount as

extremely unlikely the inclusion of Java in the range of

Brachyorrhos (Boettger, 1903; Boulenger, 1893; Schle-

gel, 1837), given the almost two centuries of herpetologi¬

cal collecting done there by individual collectors such as

Felix Kopstein (1893-1939) and Carel Pieter Johannes

de Haas (1895-1949), personnel from the Natuurkundige

Commissie voor Nederlands-Indie (Natural Sciences

Commission for the Dutch Indies), and subsequently,

Pusat Penlitian Biologi, Lembaga Ilimu Pengetahuan

Indonesia (Biological Research Centre, Indonesian In¬

stitute of Sciences), and the Museum Zoologicum Bo-

goriense.

A brief historical review of Brachyorrhos: It is

possible the inclusion of Java was due to the acciden¬

tal mixing of Javanese and non-Javanese specimens in

the collections of Heinrich Kuhl (1797-1821), Johan

Coenraad van Hasselt (1797-1823), and Heinrich Boie

(1794-1827), but see also comments under Sumatran re¬

ports below. All three collectors died in the Dutch East

Indies (Adler, 2007; 2012; 2014), while collecting speci¬

mens for the newly created Natuurkundige Commissie

voor Nederlands-Indie. Their collections were reported

upon by Hermann Schlegel (1804-1884), and Boie’s el¬

der brother Friedrich (1789-1870), both authors publish¬

ing new taxon names but ascribing them to those who

actually did the fieldwork (i.e., to Kuhl and H. Boie).

Schlegel (1826) listed the genus Brachyorrhos , as con¬

taining the species albus , dimidiatus Kuhl, kuhli Boie (=

brachyurus Kuhl), decussatus Kuhl, torquatus H. Boie,

schach H. Boie, badius H. Boie, and flammigerus H.

Boie. Friedrich Boie (1827) published an appraisal of

Versuch eines Systems derAmphibien (Merrem, 1820), in

which he included the genus Brachyorrhos Kuhl, in the

family Coronellae, and provided a numbered list of the

species in the genus that included albus , kuhlii , flammi¬

gerus, schach , badius , and torquatus. Both he and Schle¬

gel cited H. Boie’s unpublished manuscript Erpetologie

de Java as the source for many of the new species names,

but with H. Boie’s death in 1827 that manuscript was

never published; it therefore cannot be considered as a

vehicle to make species names available according to the

International Code of Zoological Nomenclature (ICZN,

1999; 2012). Today these taxa are cited as Boie (or H.

Boie) in Schlegel, 1826, or H. Boie in F. Boie, 1827

'Friedrich Boie introduced the species name kuhlii , which is an unjus¬

tified emendation of kuhli Schlegel.

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

First female specimen of Arfak Stout-tailed Snake, Calamophis sharonbrooksae

Kebar Valley

AMPAT

w&u

Ambuaki

Andai

SORONG

MANOKWARI

Mt Arfak

SORONG TELUK BINTUNI

SELATAN /

BIAK NUMFOR

► Jiu

YAPEN WAROPEN

Ansus

WAROPEN

Fig. 1. Distribution of Calamophis on the Vogelkop Peninsula, West Papua Province, and Schouten Islands, Papua Province, West

New Guinea. Titles in yellow italic font identify political entities (regencies) that are bordered by yellow lines. Titles in white font

label collection localities. Species are indicated by symbols, including C. sharonbrooksae (circle), C. ruuddelangi (downward tri¬

angle), C. katesandersae (diamond), and C.jobiensis (upward triangle). Scale = 200 km.

(David and Vogel, 1996). The fact that specimens from

many parts of the world had become mixed with the Ja¬

vanese specimens is clearly evident in the composition of

Brachyorrhos , as the forms badius, flammigerus, schach,

and torquatus are, in fact, South American taxa that are

now placed in the genus Atractus. Adler (2012:80) pro¬

vided additional examples of non-Asian snakes mixed

in with Javan specimens in Heinrich Boie’s collections.

The species described as Brachyorrhos kuhli , which was

earlier described as Coluber brachyurus by Kuhl (1820),

is actually a specimen of Atractus trilineatus {fide Hoog-

moed, 1982). This considerable confusion, caused by the

arrival at the home country institution of specimens from

far and wide, would present adequate grounds to doubt

that the specimen of B. albus originated from Java.

There have also been several reports of Brachyorrhos

from Sumatra, summarized in David and Vogel (1996),

who questioned its presence there. The author of these

reports was Bleeker (1857; 1858a; 1858b; 1860) who re¬

ported on specimens collected and donated by E.W.A.

Ludeking, from the west coast of Sumatra, and more

specifically from Padang and Again, both in West Su¬

matra Province. Unfortunately no supporting voucher

specimens appear to be extant (Murphy and Voris, 2013).

However, a new genus and species of fangless, semi-fos-

sorial homalopsid, Karnophis siantaris , was described

from a single specimen collected at Siantar, in North Su¬

matra Province (Murphy and Voris, 2013). This snake is

superficially similar in appearance to Brachyorrhos and

it is perfectly plausible that Ludeking’s “ Brachyorrhos ”

belong to this or a closely related taxon. The presence

of a Karnsophis- like homalopsid on at least one of the

Sunda Islands may provide an alternative explanation

for Javanese reports of Brachyorrhos (J.C. Murphy,

pers. comm.). We also discount reports of Brachyor¬

rhos from southeastern Kalimantan, Borneo, which were

questioned by de Rooij (1917), but accepted by de Jong

(1928).

The contributions of Murphy (2012): Murphy

(2012) expanded the resurrected genus Calamophis to

include three new species, each described from a holo-

type and a paratype, including C. katesandersae (MSNG * 2

56343-1 and 56343-2; $$) from Andai (0°54’58” S,

134°00’25” E), C. ruuddelangi (MNHM 5175, BPBM

3850; SS) from Ambuaki (0°46’00” S, 132°57’00” E)

and the Kebar Valley (0°43’21” S, 132°50’43” E) in

the Tamrau Mountains; and C. sharonbrooksae (MSNG

30193-1 and 30193-2; SS) from Mount Arfak (1°05’00”

S, 133°58’00” E). All these localities are located in the

northeast of the Vogelkop Peninsula (Fig. 1), in Manok-

wari Regency, West Papua Province, West New Guinea

(WNG). Calamophis jobiensis is known only from the

unsexed 3 holotype (MTKD 1026), collected at Ansus

on Yapen (formerly Jobi or Japen) Island (1°42’50” S,

135°49’03” E), in the Schouten Islands of Cenderawasih

(formerly Geelvink) Bay, in Yapen Waropen Regency,

Papua Province, WNG. This specimen is believed to

have been lost during the Allied bombing of Dresden

(13-15 February 1945). Iskandar and Colijn (2002) also

included Biak Island as a locality for B. jobiensis , but

Murphy et al. (2012) could locate no supporting speci¬

men. A Papua New Guinea record listed by Iskandar and

Colijn (2002) is also in error, as it was based on a speci¬

men of Mainophis robusta Macleay, 1877, a synonym of

the elapid Furina tristis (Gunther, 1858) fide Shea and

Sadlier (1999).

One of us (Mark O’Shea) recently received a loan

of Papuan snakes from the Naturhistoriska Riksmuseet,

Stockholm, Sweden (NRM, which included a specimen

Specimen numbers are preceded by the collection acronym, as listed

by Sabaj Perez MHE 2014. Standard Symbolic Codes for Institutional

Resource Collections in Herpetology and Ichthyology: An Online Ref¬

erence. Version 5.0 (22 September 2014). American Society of Ichthy¬

ologists and Herpetologists, Washington, DC, USA. [Online], Avail¬

able: http://www.asih.org [Accessed: 24 November 2014],

3 Given the low subcaudal scale count of MTKD 1026 provided in the

original description, it was most likely a female (see Table 1).

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

O’Shea and Kaiser

Fig. 2. Dorsal and ventral views of the first known female Calamophis sharonbrooksae (NRM 17803). Scale = 25 mm.

purporting to belong to the genus Brachyorrhos. Since

the most recent taxonomy indicates that it is Calamophis

and not Brachyorrhos that occurs on the Vogelkop Pen¬

insula of West New Guinea, we were able to identify this

specimen as a female specimen of C. sharonbrooksae.

This specimen is the third known exemplar of the spe¬

cies and the only known female specimen, collected ap¬

proximately 180 km west southwest of the type locality

at Mount Arfak. It is described herein.

Methods

Characters used in our evaluation and comparisons es¬

sentially conform to those used by Murphy (2012),

whose character nomenclature we follow to simplify

comparisons. Measurements of the specimen were tak¬

en to the nearest one mm and include snout-vent length

(SVL), tail length (TL), and total length (TTL). Scales

were counted following (Dowling 1951), with the ter¬

minal scale not included in the subcaudal count. Dorsal

scales were counted about one head length behind the

head, at midbody, and about one head length anterior to

the cloaca. Length measurements were obtained using a

non-elastic string and running it from the tip of the snout

along the ventral medial axis of the body to the cloaca,

with special attention to measurement accuracy (Natusch

and Shine 2012). Global Positioning System (GPS) co¬

ordinates were determined using individual museum

records, published species accounts, and Google Earth.

Results

Specimen identity: The specimen in question (NRM

17803) can be unequivocally assigned to Calamophis

sharonbrooksae based on the following pholidotic char¬

acteristics: (1) single internasal scale; (2) single post¬

ocular scale; (3) dorsal scale count of 19-19-17; (4) six

supralabials; (5) seven infralabials; (6) a single pair of

chin shields; (7) 158 ventral scales; and (8) 12 paired

subcaudals. Characters 1-3 distinguish Calamophis from

Brachyorrhos, Characters 4-8 distinguish C. sharon¬

brooksae from its congeners.

Interspecific variation in scale counts and scale

condition: The NRM specimen (NRM 17803) differs

from the males comprising the type series in exhibiting

expected higher ventral and lower subcaudal counts, and

also by the condition of the anterior temporal scales.

Calamophis sharonbrooksae is the only member of the

genus to possess paired anterior temporal scales, with the

lower scale larger than the upper scale, but this charac¬

ter is not uniformly represented across the three known

specimens. A pair of anterior temporals is present on both

sides of the head in the holotype, but the upper scale is

partially fused with the parietal on the left side; it is com¬

pletely fused, and therefore absent, on the right side in

the paratype. The female possesses a single anterior tem¬

poral on either side. Also, both parietals are not confined

to the dorsal plane, they both curve downwards onto the

lateral planes, their lowest points being in line with the

centre of the orbit, a situation most similar to the condi¬

tion on the right side of C. ruuddelangi as illustrated by

(Murphy, 2012: Fig. 3B). The left side of NRM 17803

also exhibits an additional anomalous scale (Fig. 3D, D'),

subequal in size to the postocular, sandwiched between

the postocular and the anterior temporal, and between

the parietal and the 4 th and 5 th supralabials. This scale,

which may have been formed by either a fragmentation

of the upper portion of the 5 th supralabial, by a horizontal

suture, or by the division of the anterior portion of the

anterior temporal, by a vertical suture, prevents contact

between the anterior temporal and the postocular. Clear¬

ly the condition of the anterior temporal is variable and

would therefore appear to be of little taxonomic value

in distinguishing between Calamophis species. Murphy

(2012) also used the tallest supralabial to differentiate

between C. katesandersae (5 th supralabial) and C. rn-

nddelangi and C. sharonbrooksae (4 th supralabial), but

August 2016 | Volume 10 | Number 2 | e122

Amphib. Reptile Conserv.

First female specimen of Arfak Stout-tailed Snake, Calamophis sharonbrooksae

Fig. 3. Detailed views of the head and tail of the first known female Calamophis sharonbrooksae (NRM 17803), presented as

both photographic and line-drawn illustrations for improved clarity. (A, A') Dorsal view of the head, illustrating rostral (R), single

internasal (IN), fused prefrontal-preocular (PF-PR), frontal (F), paired supraocular (SO), and parietals (P). (B, B') Ventral view of

the head, showing a single pair of chin shields (CS), seven infralabials (IL'-IL 7 ) mental (M), and the first ventral scute (V 1 ). (C,

C') Left lateral view of the head, additionally illustrating the undivided nasal (N), single postocular (PO), single anterior temporal

(AT), two posterior temporals (PT), and six supralabials (SL'-SL 6 ). (D, D') Right lateral view of the head, illustrating differences in

scalation compared to left side, three posterior temporals (PT), and small scale separating the postocular and anterior temporal (*).

(E, E') Ventral view of the tail, showing the final ventral (V 158 ), divided cloacal plate (CP), first paired subcaudal (SCI), and rounded

terminal scute (TS). Scale = 10 mm for Fig. 3A-D and 10 mm for Fig. 3E.

this character also fails with NRM 17803, which has the

5 th supralabial tallest on the right side, but the 4 th supra-

labial on the left side due to the presence of the small

anomalous scale. Assuming the situation on the left side

to be aberrant, then the tallest supralabial in NRM 17803

would conform to the type series.

History of NRM 17803: The female specimen of C.

sharonbrooksae (NRM 17803) was collected by the

Swedish zoologist Sten Bergman (1895-1975) on 6

March 1949 at Atinyu (1°26’04” S, 132 0 22’58” E), near

Lake Danau 4 , on the Vogelkop Peninsula, Sorong Selatan

Regency, West Papua Province, West New Guinea, at an

approximate elevation of 260 m (as determined using

Google Earth). This specimen also represents the west-

4 Danau is Bahasa Indonesian for “lake,” so the name of this locality is

effectively “Lake Lake.” This sort of name is not unusual in Indonesia:

a specimen of Brachyorrhos raffrayi was collected at Danau Laguna on

Ternate, a location that also translates as Lake Lake. Perhaps even more

interesting is the naming of larger areas. The country known as East

Timor in English and now called Timor-Leste was called Timor Timur

under Indonesian rule (1975-2001). Timur is Indonesian for east, leste

is Portuguese for east, so effectively this country has been called East

East since the 16 th Century.

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

O’Shea and Kaiser

Table 1: Meristic and morphometric data for Calamophis spp. Measurements are listed in mm and include snout-vent length (SVL),

tail length (TL), and total length (TTL).

Specimen

Sex

SVL

TTL

TL/SVL

TL/TTL

C. jobiensis 1

MTKD 1026

C. katesandersae

MSNG 56343-1

215

223

0.037

0.036

MSNG 56343-2

157

163

0.038

0.037

C. ruuddelangi

MHHN 5175

232

261

0.125

0.111

BPBM 3850

219

246

0.123

0.110

C. sharonbrooksae

MSNG 30193-1

288

314

0.090

0.083

MSNG 30193-2

290

318

0.097

0.088

NRM 17803

390

408

0.046

0.044

1 Meyer (1874) did not provide measurements for his holotype of Calamophis jobiensis.

2 The sex of this specimen was not reported in the original description and the specimen is now lost. However, given the low number of subcaudals,

we presume that it must have been a female.

ernmost record for the genus Calamophis and the first

mainland record outside Manokwari Regency, suggest¬

ing a much broader distribution for the genus in West

New Guinea, although it may possibly be confined to the

Vogelkop Peninsula and the Schouten Islands (see Dis¬

cussion).

Description of NRM 17803 Calamophis sharon-

brooksae : A female measuring 390 mm SVL +18 mm

TL = 408 mm TTL (Table 1).

Physique .—Moderately stout with a cylindrical body and

short tail (4.6% SVL, 4.4% TTL; Table 1) that terminates

bluntly, and a rounded head with laterally positioned nos¬

trils and eyes, with elliptical pupils.

Pholidosis (Body ).—19-19-17 dorsals, all smooth with a

high-gloss iridescence, imbricate, lacking apical pits; 158

ventrals; 12 paired subcaudals 5 ; divided cloacal plate

(Table 2).

Pholidosis (Head ).—Rostral wider than tall, visible from

above; nasals undivided with circular nares, separated by in-

temasal; intemasal single, pentagonal, in broad contact with

rostral; prefrontals paired, fused with prefrontal and loreal;

frontal shield-shaped, half again as deep as wide; parietals

paired, longer than wide, extending onto sides of head pos¬

terior to the postocular, lowest point level with the center

of the orbit; loreal fused with preocular-prefrontal; anterior

temporals on the right side comprising a single large scale

with narrow contact with the right postocular, on the left

side separated from the left postocular by a small anomalous

scale, subequal in size to the postocular; posterior temporals

comprising two on the right side, three on the left side;

supraocular single; preoculars fused with loreal and

prefrontal; postocular single; subocular absent; six su-

pralabials, the 3 rd and 4 th contacting the orbit; seven in¬

5 The subcaudal counts are 12 on the left side, 13 on the right, resulting

in 12 paired subcaudals.

fralabials, the l st -4 th contacting the chin shields; a single

pair of chin shields in broad contact, separated from 1 st

ventral scale by seven gular scales of gradually increas¬

ing size.

Coloration (after 66 years of preservation). —Dorsally

uniform dark chocolate brown body and head with cream

pigment confined to lower margins of the supralabials,

the infralabials, and the outer gular scales; chin shields,

enlarged 4 th infralabials, and gular scales between chin

shields and ventrals chocolate brown, as dorsum. Ven¬

tral scales dark brown with lighter lateral edges and dark

black margins on the ventral sutures, underside of tail

darker than body with black suturing on the subcaudals.

Discussion

With a length of 390 mm SVL +18 mm TL = 408 mm, the

female Calamophis sharonbrooksae from Atinyu (NRM

17803) is the largest known specimen of the genus Cal¬

amophis. The sole specimen of C. jobiensis is no longer

extant, and although Meyer (1874) provided scale count

data he did not provide SVL, TL, or TTL for this speci¬

men. The largest specimen of C. katesandersae is the

female holotype (MSNG 56343-1), with 215 mm SVL

+ 8 mm TL = 223 mm TTL, the largest C. ruuddelangi

is the male holotype (MNHM 5175) at 232 mm SVL +

29 mm TL = 261 mm TTL, and the previously largest C.

sharonbrooksae is the male paratype (MSNG 30193-2)

with 290 mm SVL + 28 mm TL = 318 mm TTL (Table 1),

a specimen probably not selected as the holotype due to

extensive damage, resembling fire or acid burns, on the

right side of the head.

Calamophis appears to occupy parts of western New

Guinea from which the widely distributed, and species-

rich vermivorous elapid genus Toxi'cocalamus, with 12

species known at last count (O’Shea et al., 2015a), ap¬

pears to be absent. Toxi cocalamus occurs throughout

mainland New Guinea, excluding the low-lying south-

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

First female specimen of Arfak Stout-tailed Snake, Calamophis sharonbrooksae

Table 2: Pholidotic characteristics for Calamophis spp.

Specimen

Dorsals

Ventrals

Cloacal

Plate

Subcau-

dals

Supralabials

(contact orbit)

Infral¬

abials

Post¬

oculars

Chin

Shields

Temporals

Posterior

Temporals

C. jobiensis

MTKD 1026

19-19-17 1

164

divided

10 paired

6 (3 rd + 4 th )

single

no data

“irregularly placed” 2

C. katesandersae

MSNG 56343-1

19-19-17

160

divided

9 paired

6 (3 rd + 4 th )

single

one pair

1-1

2-2

MSNG 56343-2

19-19-17

157

divided

9 paired

5p d ,2 d +3 a )

single

one pair

1-1

2-2

C. rmtddelangi

MHHN 5175

19-19-17

142

divided

21 paired

6 (3 rd + 4 th )

single

two pairs

1-1

no data

BPBM 3850

19-19-17

143

divided

23 paired

6 (3 rd + 4 th )

single

two pairs

1-1

no data

C. sharonbrooksae

MSNG 30193-1

19-19-17

150

divided

19 paired

6 (3 rd + 4 lh )

single

one pair

2-2

MSNG 30193-2

19-19-17

149

divided

17 paired

6 (3 rd + 4 th )

single

one pair

2-1

3-3

NRM 17803

19-19-17

158

divided

12 paired

6 (3 rd + 4 th )

single

one pair

1-1

3-2

‘Even though Meyer (1874) provided only the middorsal count, we presume that the dorsal pattern conforms to all of the species in the genus and

includes the posterior reduction in dorsal number.

2 Without a specimen it is not possible to determine with certainty why Meyer (1874) considered the pattern of temporal scales irregular. It may well

be that one side of the head included a different arrangement from the other, and that Meyer therefore did not consider it wise to include specifics of

the arrangement in his list of diagnostic features.

ern Trans-Fly of Western Province, Papua New Guinea

(PNG) and neighbouring Merauke Regency of Papua

Province, West New Guinea. Toxicocalamus also appears

to be absent from most of West Papua Province, WNG,

with the exception of the Onin or Fak Fak Peninsula,

from where four specimens of Apisthocalamus loennber-

gii Boulenger, 1908, currently in synonymy with Toxi¬

cocalamus loriae (Boulenger, 1898), were collected by

the English naturalist Antwerp Edgar Pratt (1852-1924),

and a single specimen of Toxicocalamus stanleyanus

Boulenger, 1917, was collected by the 1920 Dutch New

Guinea expedition (de Jong, 1927:306). Toxicocalamus

is also present on many of the islands to the southeast

of PNG (Goodenough, Fergusson, Normanby, Woodlark,

Misima, Sudest, and Rossel) and along the northeastern

and northern coasts of PNG (Karkar, Tarawai, Walis, and

Seleo), with a single specimen reportedly collected from

the large island of New Britain, but Toxicocalamus has

not been collected from the Schouten Islands of Cend-

erawasih (formerly Geelvink) Bay, despite being rela¬

tively well documented from northern and northwestern

mainland Papua Province, WNG. We therefore consider

it possible that Calamophis can only thrive in locations

not inhabited by its potentially more successful vermivo-

rous competitor Toxicocalamus , but this hypothesis does

require more investigation as other biogeographical fac¬

tors may be the cause for the apparent mutual exclusion

of the two genera.

Acknowledgments. —The authors would like to thank

Glenn Shea (University of Sydney, New South Wales,

Australia) for bringing the presence of this specimen to

the attention of Mark O’Shea, Sven Kullander and Bodil

Kajrup (Naturhistorika Riksmuseet, Stockholm, Sweden)

for facilitating the loan, and Andrew Black (University

of Wolverhampton, United Kingdom) for accepting the

loan and making laboratory facilities available to Mark

O’Shea. We also wish to extend our gratitude to John C.

Murphy for his invaluable comments on an earlier draft of

the manuscript and his generosity in sharing his thoughts

and opinions regarding the fangless homalopsids. Lastly,

the finished manuscript benefited from reviews by Ruud

de Lang, Gernot Vogel, and Harold Voris.

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O’Shea and Kaiser

Mark O’Shea is a British herpetologist with a specialist interest in the snakes of New Guinea. He wrote A Guide

to the Snakes of Papua New Guinea (1996) and is currently working on the second edition, expanded to encom¬

pass the entire New Guinea region, and he is also the author of four other books. Since 1986 he has made ten

expeditions to New Guinea to conduct herpetological fieldwork, capture medically important elapids for snake¬

bite research, or make films for Animal Planet or the BBC. He has worked in PNG for a variety of organisations

from Operation Raleigh to Oxford University’s Department of Clinical Medicine, Liverpool School of Tropical

Medicine, and the Australian Venom Research Unit, University of Melbourne. Mark also has considerable field

experience in other countries in Asia, Africa, and South America, and has been engaged in fieldwork projects since the 1980s. He

presented four seasons of the herpetological television series O’Shea’s Big Adventure, for Animal Planet and Discovery Channel,

and has made films with other companies and broadcasters. Mark was awarded the Millennium Award for Services to Zoology by

the British Chapter of the Explorers’ Club in 2000, and in 2001 was awarded a honorary Doctor of Sciences degree by his alma

mater, the University of Wolverhampton, for services to herpetology. He now teaches on the Animal Behaviour and Wildlife Con¬

servation course at the University but he also holds to post as Consultant Curator of Reptiles at West Midland Safari Park, in the

United Kingdom. Mark and Hinrich Kaiser (below) are also the leaders of the first comprehensive survey of the herpetofauna of

Timor-Leste, Asia’s newest country. With ten phases of the project completed since 2009, the team has recorded upwards of 70 spe¬

cies, with more than twenty of these new to science.

^ Hinrich Kaiser is a German-American herpetologist and educator with a research focus on biodiversity and con¬

servation of tropical environments. A passion for scuba diving with experiences in the arctic and the tropics led

Hinrich to study marine biology at McGill University and the University of Victoria in Canada. After an inspiring

semester learning about amphibians and reptiles in David Green’s herpetology class in the Redpath Museum, Hin¬

rich found his true calling and earned his PhD at McGill with a dissertation on the systematics and biogeography

of Lesser Antillean frogs. After a Boehringer Ingelheim postdoctoral fellowship at the University of Wurzburg,

Germany, Hinrich spent five years as Professor of Biology at La Sierra University, Riverside, California, USA,

before accepting his current position in the Department of Biology at Victor Valley College in Victorville, California, USA. Hinrich

also holds an appointment as Research Associate with the United States National Museum of Natural History, Smithsonian Insti¬

tution, Washington, D C., USA. He currently serves as an Editor-in-Chief of Herpetology Notes, but his interests in international

affairs and music also led him to memberships on the International Advisory Board of the Loundation for Post-Conflict Develop¬

ment, New York, and on the Advisory Council of the Baltimore Symphony Youth Orchestras. Hinrich also serves as a member of

the Executive Committee of the World Congress of Herpetology. His most recent publications have focused on the herpetofauna of

Timor-Leste and nearby areas of Wallacea, as well as on the defense of herpetological taxonomy against taxonomic vandalism. His

educational specialty is to expose community college students to biological, cultural, and historical experiences overseas, including

canopy walks in Brunei, cooking classes in Bali, tracking Komodo dragons on Rinca Island, homestays in Cuba, and surveying

Pacific atolls.

Amphib. Reptile Conserv.

August 2016 | Volume 10 | Number 2 | e122

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [General Section]: 11-17 (e123).

Notes on natural history and call descr ption of the

Critically Endangered Plectrohyla avia (Anura: Hylidae)

from Chiapas, Mexico

^esar L. Barrio-Amoros, 234 Christoph I. Griinwald, 345 Hector Franz-Chavez, 67 Angela Maria Mendoza and

8 Brandon Thomas La Forest

1 Doc Frog Expeditions, Uvita, COSTA RICA 2 Biencom Real Estate, Carretera Chapala -Jocotepec #57-1, C.P. 45920, Ajijic, Jalis¬

co, MEXICO 3 Herpetological Conservation International - Mesoamerica Division, 450 Jolina Way, Encinitas, California 92024,

USA 4 Biodiversa A.C., Avenida de la Ribera #203, C.P. 45900, Chapala, Jalisco, MEXICO 5 Centro Universitario de Ciencias

Biologicasy Agropecuarias, Carretera a Nogales Km. 15.5. LasAgujas, Nextipac, Zapopan, C.P. 45110, Jalisco, MEXICO 6 Depar-

tamento de Zoologia, Instituto de Biologia, UniversidadNacional Autonoma de Mexico, Apartado postal 70-153, 04510 Mexico

City, MEXICO 7 Grupo de Investigacidn en Ecologiay Conservacion Neotropical, Samanea Foundation, 760046 Cali, COLOMBIA

*815616 North 10th Place, Phoenix, Arizona 85022, USA

Keywords. Reproduction, vocalization, behavior, treefrog, Mesoamerica, Guatemala

Citation: Barrio-Amoros CL, Grunwald Cl, Franz-Chavez H, Mendoza AM, La Forest BT. 2016. Notes on natural history and call description of the

Critically Endangered Plectrohyla avia (Anura: Hylidae) from Chiapas, Mexico. Amphibian & Reptile Conservation 10(2) [General Section]: 11-17

(e123).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any

medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and

authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation; official journal

website <amphibian-reptile-conservation.org>.

Received: 25 May 2016; Accepted: 12 July 2016; Published: 12 September 2016

Plectrohyla avia Stuart, 1952 was described on the basis

of a single male from Granja Lorena, Quetzaltenango,

Guatemala (Stuart 1952). Since then, only a few

observations have been made of this species, expanding

the distribution from the vicinity of the El Triunfo

Biosphere Reserve in the Sierra Madre del Sur de

Chiapas southeast across the Volcan Tacana of Chiapas

and the Volcan Tajumulco of southwestern Guatemala

to the highlands of the Quetzaltenango District of

Guatemala (Duellman and Campbell 1992). Only general

information on morphology and distribution has been

published, and the natural history of this species remains

unknown (Duellman 2001; Kohler, 2010).

Plectrohyla avia is the largest member of its genus;

males reach 90.4 mm, and females 70.4 mm. None of

the large species of Plectrohyla (P. avia, P. teuchestes

Duellman and Campbell 1992; P. exquisita McCrannie

and Wilson, 1998; P. hartxvegi Duellman 1968) have been

tested molecularly (Faivovich 2005; Frost 2006; Pyron

and Wiens 2011; Duellman et al. 2016), and thus it is not

possible to establish relationships with the smaller species

of the genus. The recent split of Plectrohyla, describing

Sarcohyla for the bistincta group (Duellman et al. 2016)

and retaining Plectrohyla for the guatemalensis group,

helps to clarify the taxonomic panorama. All four species

of large Plectrohyla are considered Critically Endangered

by the IUCN (Acevedo and Smith 2004; Cruz et al. 2010;

Santos-Barrera et al. 2004, 2006). While they are likely

all closely related, P. avia stands out within the group as

it has a single prepollical spine (bifid on the others) and

very protruding teeth.

No detailed natural history observations on any of

the larger species of Plectrohyla (P. avia, P. teuchestes,

P. exquisita, and P. hartwegi ) have been published

(Duellman and Campbell 1992; Duellman 2001; Kohler

2010). We herein report observations on the reproductive

activity and natural history of P. avia in its habitat.

On May 5 th , 2016, we arrived at the small village of

Mirador Chiquihuites, Municipality of Union Juarez,

Chiapas, at ± 1700h, just as it was starting to rain (amongst

the first rains of the season, according to the locals).

Several Plectrohyla cf. sagorum Hartweg, 1941 were

calling from a small ravine below the road. The ravine

was located at 2015 m asl at the following coordinates:

15.095167°, -92.106669° datum = WGS84.

About 150 m upstream, within a narrow ravine

Correspondence. Email: cesarlba@yahoo.com (Corresponding author)

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e123

Barrio-Amoros et al.

Fig. 1. Aquatic axillary amplexus of Plectrohyla avia. (Right top). Photo by Cesar L. Barrio-Amoros. Fig. 2. Male of Plectrohyla

avia grasping the female’s head with its long and curved teeth. (Left top). Photo by Cesar L. Barrio-Amoros. Fig. 3. Amplectant

pair of Plectrohyla avia, two motionless males and eggs over the pool. (Bottom left). Photo by Cesar L. Barrio-Amoros. Fig. 4.

Reproductive male of Plectrohyla avia inside a hole on the waterfall side, from where it calls. (Bottom right). Photo by Cesar L.

Barrio-Amoros.

covered by dense canopy, we heard a low but distinctive

call (see below). Upon closer examination, we observed

five large green frogs in a small pool. The pool was

approximately one square meter in area and located at

the base of a three m high waterfall with a very light flow.

The five large green frogs (four adult males and one adult

female) were Plectrohyla avia. Upon discovery all five

animals were apart from each other and below the surface

of the water. Two males were noticeably larger than the

other two, and at one point engaged in what appeared to

be male-male combat. Two of us (CG and HF) saw one

male grasping the other and though not very clear (too

dark), we saw how the largest embraced the smaller and

forced it to escape. After this we filmed a short video

(https://youtu.be/aa20-BguqOY). Shortly after the

combat, the largest male embraced the only female

in an axillar amplexus (Fig. 1), while the other three

males remained inactive under water. This male was

significantly larger than the female, as has been reported

for some larg q Plectrohyla (Duellman and Campbell 1992;

Kohler 2010). During the first minutes the female moved

around the pool while in amplexus with the inactive male

(Fig. 2). The female appeared to be trying to escape from

the male who remained in strong amplexus. After some

minutes swimming across the pool, beneath the surface

of the water at all times, the female stopped on the edge

of the pool. The amplectant male remained motionless.

Another male began calling underwater (sec 16 of video;

the male was not seen calling directly but was the only

close male and it was under water; other underwater

calls were heard afterwards occasionally); immediately

thereafter the amplectant male began scratching the top

of the female’s head with the long teeth that protrude from

the upper jaw. The female tried to release herself from

the amplecting male and again began swimming, but

the male continued to move his head laterally scratching

the female’s head. While being passed by the swimming

female, the other males remained motionless (Fig. 3).

The pair again stopped at the other edge of the pool. The

rocky bed and walls of the pool were covered by large

eggs, all underwater, probably from earlier amplectant

pair(s). We did not see the female laying eggs.

Above the small pool with the five adults, we observed

three more adults. Two males were calling from within

small holes in the splash-zone of waterfalls varying from

one to four m in height (Fig. 4). Another adult male was

perched on the wet wall of the higher waterfall (Fig. 5).

Adult males called from within holes, crevices or beneath

the surface of the water.

Two kinds of eggs were observed in the pool. A

smaller one in dense quantities with pigmented pole,

and other bigger and unpigmented, more scattered and

in a much lower density. We cannot rely of which one

was laid by P. avia , as we did not see directly the female

laying eggs. Plectrohyla cf. sagorum was present on

the ravine as well (we saw several calling males and

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | el 23

Natural history and call description of Plectrohyla avia

Fig. 5. Active male of Plectrohyla avia on a waterfall wall, the

only male we saw outside the water or holes. Photo by Cesar

L. Barrio-Amoros.

one amplectant pair) and we cannot be sure to which

species the eggs belong. Only unpigmented eggs have

been reported for Plectrohyla (Duellman and Campbell

1992). An alternative explanation could be that the

larger unpigmented eggs belong to different stages of

development of the same species.

Vocalization

Recording specifications: The first call (Fig. 6) was

recorded with a Sony RX10 camera in HD 1080, and

sound extracted in a WAV file, 48 kHz of sample rate, 16

bit signed. The second call (Fig. 7) was recorded with a

Nikon d5100 camera and sound extracted in a WAV file,

44.1kHz of sample rate, 16 bit signed. Recordings were

analyzed in Raven Pro 1.5 Beta (Bioacoustics Research

Program 2013), with a Hann function window, FFT 1024

samples, and 50% overlap). A summary of spectral and

temporal features of vocalizations of Plectrohyla avia is

in Table 1, and details of each pulse of the calls are in

Table 2.

Vocalization description: The male advertisement

call of P. avia is a brief, rapid, trill. The first call

(Fig 6) consists in a series of seven consecutive pulses

at a mean frequency of 561.5 Hz (541-572 Hz). The

duration of the call is of 0.28 sec and duration between

pulses of 0.020 sec. The initial and final pulses show the

lesser intensity, lower than 80 dB, while the medium

pulses have amplitude above 85 dB (Table 2, Fig. 6A).

The second call (Fig. 7) has nine consecutive pulses at

a mean dominant frequency of 349.5 Hz (348-353 Hz).

This call is longer than the first one (0.405 sec) and we

are not sure if was emitted by the same individual. Each

pulse has a mean duration of 0.032 sec and the duration

between pulses is 0.011 sec. Like the first call, the first

and last pulse were lower in intensity (under 60 dB),

while the medium pulses presented amplitude above 62

dB (Table 2, Fig. 7B).

Discussion

The first exceptional thing about these observations is

that this is the first case of underwater breeding behavior

among hylids (with exception of Pseudinae, of which

two genera are highly adapted to aquatic life).

A second noteworthy observation is that combat

behavior has never been observed in Plectrohyla.

2 -

Fig. 6. Waveform (above) and spectrogram (below) of the first subaquatic call analyzed of Plectrohyla avia.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | el 23

Barrio-Amoros et al.

Fig. 7. Waveform (above) and spectrogram (below) of the second subaquatic call analyzed of Plectrohyla avia.

Dermal scratches, however, have been reported in some

species (like P. teuchestes and P. harwegi r; Duellman and

Campbell 1992), and these observations suggest that they

are caused by the prepollical spine used in some kind of

combat. While large but stout teeth have been reported

for P. exquisita, a very similar species (McCrannie and

Wilson 1998), the teeth in the upper jaw in a breeding-

condition male of P. avia (MZFC [Museo de Zoologia,

Facultad de Ciencias, Universidad Nacional Autonoma

de Mexico, Mexico] 29273) are extraordinarily long,

protruding from the upper jaw and curved (Fig. 8). This

may have an important function to promote the female

sexual reception or ovulation, as the male was seen

apparently using them to scratch the female’s head.

However, we did not find any trace of mental gland on the

reproductive male collected, and the function must not

be similar to that in plethodontid salamanders, in which

males also scratch female’s head with its premaxillary

teeth inoculating secretions from the mental glands

(Arnold 1977; Duellman and Trueb 1994). Another

possible explanation is that during the aquatic period,

the individuals eat freshwater hard-shelled insects

(Notonectidae, Hydrophilidae, and Gyrinidae). We did

not observe any expansion of the throat on the calling

male. Another striking feature was the presence of very

well developed lateral skin soft folds (Fig 9), that may

help in dermal respiration, as in high Andean frogs of the

family Telmatobiidae (Duellman and Trueb 1994).

All reproductive activity that we witnessed was from

1715h to 1830h, when it was still fight outside the dark

ravine. We collected one male and came back after dark

at 2200h, but not a single individual was observed in the

area.

In the same ravine, many Plectrohyla cf. sagorum

were heard and seen, including one amplectant pair. This

species is smaller (males up to 45.5 mm, females up to

51.9 mm; Kohler 2010) and no individual was observed

in the water; several males were calling from rocks facing

the ravine, and an amplectant pair was on the wall of a

small waterfall. In a nearby larger stream, with many P.

matudai (including recent metamorphs) but not P. avia ,

we found four Plectrohyla hartwegi, close to a four m

high waterfall, all were perched on stems and ferns in or

close to, the spray zone. This species has very flared lips

but no protruding teeth.

Fig. 8. Detail of the long and curved teeth protruding from the

upper maxilla of a preserved reproductive male MZFC 29273.

Photo by Chris I. Grunwald.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | el 23

Natural history and call description of Plectrohyla avia

Fig. 9. Lateral view of a reproductive male MZFC 29273, showing the lateral skin folds. Photo by Cesar L. Barrio-Amords.

Conservation status

In less than two hours we saw eight Plectrohyla avia in

a section of less than 20 m in the ravine. At the small

pool and inside the holes in the wall of the canyon were

many eggs attached to the rock or stems, some freshly

laid, some with white embryos visible, and already

some tadpoles recently hatched (Fig 10). We cannot be

absolutely sure that all those eggs belong to P. avia , as

we never saw females laying. This species, however, was

the dominant in that sector of the ravine and occupied

the lower pool (five individuals using the pool for

reproductive purposes), and though no adults were seen

in the upper pool (Fig. 10), two adult males were very

close. On the upper pool only unpigmented eggs were

seen, very likely belonging to P. avia.

Fig. 10. Pool full of eggs in different stages of Plectrohyla avia.

Photo by Cesar L. Barrio-Amords.

We queried local inhabitants about the frogs, and all

them recognized the species, and said they are common.

One boy told us that he enjoyed killing them. While the

IUCN (Santos-Barrera et al. 2006) and Stuart et al. (2008)

consider this species to be in the maximum category of

vulnerability (CR A3e), Johnson et al (2015) consider the

EVS score as 13 -of 20- (medium category). Probably

the species is more widely distributed and more abundant

than expected, but with a short period of activity at the

beginning of the rainy season. More information is

needed to establish a definitive category of conservation.

Acknowledgments. —All specimens which were

deposited in MZFC were deposited under permit #FAUT-

0093 issued to Dr. Adrian Nieto Montes de Oca. Permits

were issued by the Secretaria de Media Ambiente y

Recursos Naturales (SEMARNAT). We especially thank

Dr. Adrian Nieto Montes de Oca and the Universidad

Nacional Autonoma de Mexico - Museo de Zoologia de

la Facultad de Ciencias for his generous and unfaltering

support to further the understanding of Mexican

herpetofauna. Biodiversa A.C. and Herpetological

Conservation International provided important funding

for this project. Ray Morgan helped with the sound

extraction.

Literature Cited

Acevedo M, Smith E. 2004. Plectrohyla tenchestes. The

IUCN Red List of Threatened Species 2004: Available:

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e.T55887Al 1372355.http://dx.doi.org/10.2305/

IUCN.UK.2004.RLTS.T55887A 11372355.en

[Accessed: 12 May 2016],

Arnold SJ. 1977. The evolution of courtship behavior in

New World salamanders with comments on Old World

salamandrids. Pp. 141-183 In: Editors, Taylor DH,

Guttman. The Reproductive Biology’ of Amphibians.

Plenum Press, New York, New York, USA. 475 p.

Cruz G, Wilson LD, Castaneda F, Mendelson III J, Kolby

JE. 2010. Plectrohyla exquisita. The IUCN Red List

of Threatened Species 2010: e.T55874Al 1366549.

Available: http://dx.doi.org/10.2305/IUCN.UK.2010-

2.RLTS.T55874A11366549.en [Accessed: 12 May

2016],

Duellman WE, Campbell JA. 1992. Hylid frogs of the

genus Plectrohyla : Systmatics and phylogenetic

relationships. Miscellaneous Publications, Museum

of Zoology, University of Michigan 181: 1-32.

Duellman WE. 2001. The Hylid Frogs of Middle America.

Society for the Study of Amphibians and Reptiles,

Ithaca, New York, USA. 1,180 p.

Duellman WE, Marion AE, Hedges BS. 2016.

Phylogenetics, classification, and biogeography of the

treefrogs (Amphibia: Anura: Arboranae). Zootaxa.

4104: 1-109.

Duellman WE, Trueb L. 1994. Biology of Amphibians.

Johns Hopkins University Press, Baltimore, Maryland,

USA. 670 p.

Faivovich J, Haddad CFB, Garcia PC A, Frost DR,

Campbell JA, Wheeler WC. 2005. Systematic review

of the frog family Hylidae, with special reference

to Hylinae: a phylogenetic analysis and taxonomic

revision. Bulletin of the America?! Museum of Natural

History 294:1-240.

Frost, DR, Grant T, Faivovich J, Bain RH, Haas A,

Haddad CFB, de Sa RO, Channing A, Wilkinson M,

Donnellan SC, Raxworthy CJ, Campbell JA, Blotto

BL, Moler PE, Drewes RC, Nussbaum RA, Lynch JD,

Green DM, Wheeler WC. 2006. The Amphibian Tree

of Life. Bulletin of the American Museum of Natural

History 297: 1-370.

Johnson JD, V Mata-Silva, LD Wilson. 2015. A

conservation reassessment of the Central American

herpetofauna based on the EVS measure. Amphibian

& Reptile Conservation 9(2): 1-94 (el00).

Kohler G. 2011. The Amphibians of Central America.

Herpeton, Verlag Elke Kohler, Offenbach, Germany

(www.herpeton-verlag.de). 379 p.

McCranie JR, Wilson LD. 1998. Specific status of the

the Honduran frogs formerly referred to Plectohyla

teuchestes (Anura: Hylidae). Journal of Herpetology

32: 91-101.

Pyron, RA, Wiens JJ. 2011. A large-scale phylogeny of

Amphibia including over 2800 species, and a revised

classification of advanced frogs, salamanders, and

caecilians. Molecular Phylogenetics and Evolution

61: 543-583.

Santo s-Barrera G, Canseco-Marquez L, Acevedo

M. 2004. Plectrohyla hartwegi. The IUCN Red

List of Threatened Species 2004. Available:

e.T55877Al 1367812.http://dx.doi.org/10.2305/

IUCN.UK.2004.RLTS.T55877A11367812. en

[Accessed: 12 May 2016],

Santo s-Barrera G, Canseco-Marquez L, Acevedo M,

Munoz A, Mendelson III J. 2006. Plectrohyla avia. The

IUCN Red List of Threatened Species 2006: Available:

e.T55871Al 1365383.http://dx.doi.org/10.2305/

IUCN.UK.2006. RLTS.T55871 All 365383.en

[Accessed: 12 May 2016]

Stuart LC. 1952.Some new amphibians from Guatemala.

Proceedings of the Biological Society of Washington

65: 1-12.

Cesar L. Barrio Amoros is an anthropologist who has worked with herpetofauna in Spain, Venezuela,

Peru, Costa Rica, and Mexico. His research interests include the biogeography and systematics of

Amphibia and Reptilia, with emphasis in Dendrobatoidea and Terrarana from Venezuela, especially

of the Guiana Shield. Now a resident of Costa Rica, he is a free-lance investigator and photographer.

Cesar has authored or co-authored more than 200 papers, including the description of 50 new species of

amphibians and reptiles.

Christoph I. Grunwald is a German-Mexican herpetologist who for the last 15 years has led field

expeditions to study herpetofauna. A specialist in Mexican biogeography, he has been involved with the

discovery of over 100 range extensions and dozens of state records for amphibians and reptiles from

around the country. Christoph specializes in rattlesnakes and pitvipers, and many important discoveries

have involved this group. Currently he is leading research expeditions on Mexican rattleless pitvipers

and direct-developing frogs. A co-founder of Biodiversa, A.C., an anti-extinction non-profit organization,

Chris currently is developing a system of “micro-reserves,” aimed at preserving the most vulnerable,

high-endemism localities in Mexico.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | el 23

Natural history and call description of Plectrohyla avia

Hector Franz-Chavez was bom in Guadalajara, Mexico, and has had a passion for herpetology since

childhood. He is a student of biology at the Universidad de Guadalajara (CUCBA), and his main interests

include biogeography, natural history, and ecology of the herpetofauna of Mexico. He also is an avid

nature photographer, and has collaborated in various herpetological inventories in different parts of

Jalisco, and currently is working on an inclusive project on Mexican direct-developing frogs and several

potentially new species of pitvipers. Hector has traveled extensively in the Sierra Madre Occidental and

the Sierra Madre del Sur, where he has collected numerous specimens of interest.

Angela M. Mendoza is a biologist from Universidad del Valle (Colombia) with a M.S. in biological

sciences and is currently a Ph.D. student at the Universidad Nacional Autonoma de Mexico (UNAM).

Her work has focused mainly in the application of molecular tools in solving questions in ecology and

conservation, with an emphasis in terrestrial vertebrates, mainly Neotropical amphibians.

Brandon La Forest was born in Phoenix, Arizona. He has been interested in herps, particularly vipers, as

far back as he can remember. He studied Ecology and Evolutionary Biology at the University of Arizona

in Tucson. A life long enthusiast, he enjoys traveling abroad to document reptiles and amphibians. Mexico

has always been a special place for him; where he spends over a month each year collaborating with

different biologist to catalog undocumented, undescribed, and under sampled reptiles and amphibians.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | el 23

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [General Section]: 18-26 (el24).

First population assessment of the endemic insular

Psychedelic Rock Gecko (Cnemaspis psychedelica) in

southern Vietnam with implications for conservation

^ai Ngoc Ngo, 2 Truong Quang Nguyen, 2 Tan Van Nguyen, 3 Frank Barsch, 4 ’ 5 Thomas Ziegler,

and 45 Mona van Schingen

1 Vietnam National Museum ofNature, Vietnam Academy of Science and Technology, 18 Hoang Ouoc Viet Road, Hanoi, VIETNAM

2 Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Hanoi,

VIETNAM 3 Federal Ministry for the Environment, Nature Conservation, Building and Nuclear Safety, Robert-Schuman-Platz 3,

53 1 75 Bonn, GERMANY A Department of Terrestrial Ecology, Institute of Zoology, University of Cologne, Ziilpicher Strafie 47b,

50674, Koln, GERMANY Cologne Zoo, Riehler Strafie 173, 50735, Cologne, GERMANY

Abstract.—The Psychedelic Rock Gecko (Cnemaspis psychedelica) was recently discovered on

Hon Khoai Island, Ca Mau Province, South Vietnam. Its striking coloration makes the species highly

desired on the international pet market. Although public access to the Island is generally prohibited,

a number of specimens were already illegally captured and internationally offered for extremely

high prices. In contrast, the current wild population size and the extent of human impacts on the

species remain unknown. The present study provides the first population size estimation using a

capture-recapture method and evaluation of potential threats to C. psychedelica in order to assess

its conservation status. While the wild population was found to be relatively stable and actively

reproducing at time, we simultaneously recorded increasing habitat destruction, which might

considerably affect the population of C. psychedelica. Thus, we herein provide recommendations

for in situ conservation. Furthermore, we report the first record of C. psychedelica from another

small offshore Island in Rach Gia Bay.

Keywords. Population size, conservation status, habitat degradation, threats, in-situ species management, endemism

Citation: Ngo HN, Nguyen TQ, Nguyen TV, Barsch F, Ziegler T, and van Schingen M. 2016. First population assessment of the endemic insular

Psychedelic Rock Gecko {Cnemaspis psychedelica) in southern Vietnam with implications for conservation. Amphibian & Reptile Conservation 10(2)

[General Section]: 18-26 (el 24).

4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any medium, provided the original author

and the official and authorized publication sources are recognized and properly credited. The official and authorized publication credit sources, which

will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website <amphibian-reptile-conservation.

org>.

Received: 31 March 2016; Accepted: 16 June 2016; Published: 30 September 2016

Introduction

The colorful Psychedelic Rock Gecko ( Cnemaspis psy¬

chedelica) was recently discovered on Hon Khoai Island

in Rach Gia Bay, Ngoc Hien District, Ca Mau Province,

southern Vietnam, where it was assumed to be endemic

(Grismer et al. 2010). The island, which is only about

eight km 2 in size, is one of the biggest of the total 92

islands in Rach Gia Bay. Currently, five Cnemaspis spe¬

cies are known from Vietnam, namely C. aurantiacopes,

C. boulengerii, C. caudanivea, C. psychedelica and C.

tucdnpensis , whereas two (C. boulengerii , C. psychedeli¬

ca) are endemic to islands in southern Vietnam (Grismer

Correspondence. Email: * mschinge@smail.imi-koeln.de

and Ngo 2007; Grismer et al. 2010). Cnemaspis psyche¬

delica is the most strikingly colored Cnemaspis species,

which makes it even more attractive for the pet trade. Al¬

though the public access to the island, which is managed

by the military, is prohibited, concrete evidence exists

for the repeated appearance of the species in the interna¬

tional trade (Altherr 2014; Auliya et al. 2016; Nguyen et

al. 2015). Island species are particularly vulnerable to ex¬

tinction due to human impacts, because over-collection

for the pet trade has frequently caused rapid local extir¬

pation of lizard species (e.g., Auliya et al. 2016; Stuart et

al. 2006; van Schingen et al. 2015).

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Ngo et al.

Fig. 1. Habitat of Cnemaspispsychedelica. A. Hon Khoai Island; B. Microhabitat on Hon Khoai; C. Hon Tuong Island; D. Deposited

eggs on Hon Tuong Island. Photos H.N. Ngo.

Basic knowledge on the current population status of

C. psychedelica , its ecology and potential threats are still

lacking, as it is the case for most lizard species in the

region. To better understand the threat level of a species,

population size estimations provide essential baseline in¬

formation and are thus crucial for wildlife management

strategies and the assessment of the conservation status

of populations and species (Ngo et al. 2016; Reed et al.

2003; Traill et al. 2007). The present study aims to pro¬

vide the first assessment of the population size of C. psy¬

chedelica as well as an evaluation of potential threats, in

particular human impacts, in order to assess its conserva¬

tion status and develop adequate conservation strategies.

We further investigated seasonal variation in population

size and structure by conducting field surveys both dur¬

ing the wet and dry seasons. In addition, we surveyed an¬

other, smaller offshore island in proximity to Hon Khoai

to investigate potential occurrence of the species to as¬

sess its distribution range.

Materials and Methods

Field surveys. Field surveys were conducted during

the wet season in November 2015 as well as during the

beginning of the dry season in January 2016 on Hon

Khoai Island, Rach Gia Bay, Ca Mau Province, south¬

ern Vietnam. In 2015, six transects (0.3 km to 1.1 km

in length) along granitic cliffs and boulders were repeat¬

edly surveyed three times each. In 2016, a further site

was studied in addition to previous sites. Furthermore,

another small island (Hon Tuong), situated in proximity

to Hon Khoai island was surveyed. Hon Tuong is about

250 m 2 in size and situated in about 2.3 km distance from

the ranger station on Hon Khoai Island. The access to

another small Island (Hon Sao), which is used to breed

monkeys, was not authorized. Surveys took place after

sunset from 19:30 to 23:30 h. Animals were captured by

hand and measured with a digital caliper to the nearest

0.1 mm. Lizards were individually marked with a per¬

manent pen and released on the same spot after taking

measurements (for details see Ngo et al. 2016, Fig. 2).

This marking technique has the advantage of being non-

invasive and inexpensive. Each spot was marked by a

label made of waterproof paper to identify exact posi¬

tions of captured and recaptured animals. Based on direct

observations during the field surveys on Hon Khoai, the

extent of habitat degradation and anthropogenic pres¬

sures on the island was assessed. Local forest rangers and

workers of the military were interviewed during random

oral conversations to determine the general cognizance,

perception, and use of the species in Vietnam. Due to pri¬

vacy rights, interviewees were kept anonymous.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Population assessment of the Psychedelic Rock Gecko in southern Vietnam

Fig. 2. A. Cnemaspispsychedelica on Hon Khoai Island; B. Marked C. psychedelica on likewise marked occurrence site; C. Group

of C. psychedelica at egg deposition site. Photos H.N. Ngo.

Data analyses. We estimated the population size of

C. psychedelica by applying a non-invasive “capture-re¬

capture” approach after Schltipmann and Kupfer (2009).

Estimated population sizes refer to investigated sites and

might not encompass the entire population of the species

on Hon Khoai and the surrounding small islands. Based

on snout-vent length (SVL) animals were categorized

into two age classes (SVL >58 mm = adult and SVL

<58 mm = subadult or juvenile), sexes, and in case of

females, into gravid and non-gravid specimens to assess

the population structure. To test for seasonal differences

in population structure a Chi 2 test with a = 0.05 was ap¬

plied with GraphPad Prism version 5.0 for Windows,

GraphPad Software, La Jolla, California, USA, www.

graphpad.com.

Results

New record. During the present study, C. psychedelica

was recorded for the first time from the small offshore

island Hon Tuong, in the vicinity of Hon Khoai Island

(Fig. 1C). In total 11 individuals were observed on this

island, of which two were juveniles. Animals were exclu¬

sively found within the center of the island, which was

covered by vegetation. In addition, some deposited eggs

were recorded, demonstrating a reproducing population

on Hon Tuong Island.

Population status. During the present study, individu¬

als of C. psychedelica were found along seven transects

on Hon Khoai Island as well as on the small offshore

island of Hon Tuong. A total of 267 different animals

were captured during the wet season, of which 196 were

adults, and 526 individuals (378 adults) were observed

during the dry season (Table 1). Based on four investi¬

gated sites, population size estimates revealed about 365

animals during the wet season and 576 estimated indi¬

viduals in the dry season. Including two further sites

(viz. six sites), we recorded a total population size of 732

individuals during the dry season on Hon Khoai Island.

The effective population size (considering only mature

individuals) was estimated approximately 507 animals

during the dry season (Table 1).

Regarding different surveyed transects, we found

higher numbers of individuals during the dry season

compared to the wet season. Only at site three, we found

fewer individuals during the dry season compared to

the wet season (Fig. 3). Contemporarily, we observed

a distinct increase in some disturbances, such as habi-

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Ngo et al.

” 70

>- 60 -

u i

£50

QJ-

£ 30 ^Hw

20 I^HKn HHBHHH

10 (*Ky : •' ':% -. < .|

Wet season Dry season

T1 T2 T3 T4 T5

Fig. 3. Seasonal variation in the population structure of

Cnemaspis psychedelica on Hon Khoai Island during the wet

and the dry season; n = 154 in the wet season, 410 in the dry

season, respectively; B. Seasonal variation in the densities of C.

psychedelica on Hon Khoai Island during the wet and the dry

season. Tl-6 = different habitat sites.

tat destruction or waste around caves at this site, which

probably had a negative impact on C. psychedelica. With

regard to seasonal variations, the mean density of C.

psychedelica along suitable habitat sites was estimated

to be around 120 individuals per km/transect during the

wet and 192 individuals per km/transect during the dry

season. While densities were generally higher during the

dry season, only at site three the calculated density was

lower during the dry season compared to the wet season

(Fig. 3). By comparing different habitat sites, the highest

density of C. psychedelica was observed during the dry

season at site five (277 individuals per km/transect; see

Fig 3). This site was characterized by undisturbed dense

forest, with closed canopy above the inhabited granite

outcrops, relatively far from the Hon Khoai Ranger Sta¬

tion and areas used by the military.

In addition to different numbers of encountered ani¬

mals during the wet and dry seasons, we further found

a significant seasonal difference in population structure

(Chi 2 = 7.98, df = 3 ,p< 0.05). Adult males accounted for

the highest portion of observed animals during the wet

season (47%), while adult females were most abundant

during the dry season (38%, see Fig. 3). Furthermore,

about one half of the observed females were found to be

gravid during the dry season, in contrast to only 13% dur¬

ing the wet season (Fig. 3).

Threats. A clear increase in habitat destruction was ob¬

served within only two months from November 2015 to

January 2016. A large road around the whole island is

being constructed, which already led to accelerated ero¬

sion within the adjacent forest (Fig. 4A-C). To flatten

the area for the road construction, granitic formations,

which provide the preferred habitat for C. psychedelica,

were blasted with dynamite (Fig. 4B). During daytime,

field surveys could not be conducted, because of frequent

bursting activities. For example, we observed deer killed

by these activities (Fig. 5C). Besides road construction,

three artificial ponds were being built for farming fish

as food resources and local use (Fig. 4D). Therefore, the

circumjacent forest was cleared. According to our inter¬

views, some ecotourism programs are planned on Hon

Khoai Island in the near future as well, which include the

alteration and deterioration of rock habitats for C. psy¬

chedelica. Furthermore, fishermen are temporarily living

on the island, and exclusively consume local resources

(Fig. 4E). For example, they set up traps to catch moni¬

tor lizards, which we observed on both Hon Khoai and

Hon Tuong islands (Fig. 4F). The fishermen set up their

camps within the preferred microhabitat by C. psyche¬

delica, under granite karst or within caves, consigning

a lot of waste in the surrounding forest. The recently in¬

troduced Long-tailed Macaques (Macaca fasciciriaris)-

known to feed on gekkonids-might represent another po¬

tential threat to C. psychedelica (Fig. 5A); however, the

impacts are not yet fully understood so far (Grismer et

al. 2010). In addition, C. psychedelica recently occurred

in the international pet trade, even though public access

□ Juvenile

□ Female (gravid)

□ Female {non gravid)

■ Male

Table 1. Estimated population sizes and total numbers of observed Cnemaspis psychedelica on Hon Khoai and Hon Tuong islands;

Est. to = estimated total individuals, Est. c = estimated effective size (mature individuals), Obs. tot = totally observed individuals, Obs. c

= number of observed mature individuals.

Site

Wet Season

Dry Season

EstJEst,)

Obs. tot (Obs. e )

Est.JEst,)

ObS.JObS.e)

1 (Hon Khoai)

6(5)

97(68)

74(56)

2 (Hon Khoai)

98(64)

63 (31)

157(101)

124(78)

3 (Hon Khoai)

99(94)

68(66)

72(49)

39(27)

4 (Hon Khoai)

69 (45)

53(35)

117(77)

101 (69)

5 (Hon Khoai)

99 (72)

77 (59)

133 (82)

83 (58)

6 (Hon Khoai)

—

156(130)

94(81)

7 (Hon Tuong)

—

11(9)

Total

365 (275)

267(196)

732 (507)

526 (378)

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Population assessment of the Psychedelic Rock Gecko in southern Vietnam

Fig. 4. Potential threats to Cnemaspis psychedelica. A. Erosion caused by opening of new ways; B. Forest opening and blasting of

granite karst formations; C. Building of new roads; D. Building of artificial ponds to store freshwater; E. Camp of a fisherman living

on the island; F. Trap for Monitor lizards. Photos H.N. Ngo.

to the island is generally prohibited (e.g., Altherr 2014;

Auliya et al. 2016; Nguyen et al. 2015).

Discussion

The colorful Psychedelic Rock Gecko was thought to be

endemic to Hon Khoai, one of the biggest islands in Rach

Gia Bay, while our new record from Hon Tuong Island

indicates a slightly wider distribution range and for the

first time its existence also on very small islands. Ngo et

al. (2016) also reported the occurrence of another insular

gecko from Vietnam, Goniurosaurus catbaensis, origi¬

nally thought to be endemic for Cat Ba Island, on very

small nearby offshore islands in the Ha Long Bay. These

Endings highlight the importance of such small islands

for gekkonids, which have long been overlooked. We as¬

sume that C. psychedelica probably occurs on other simi¬

lar small islands in the Rach Gia Archipelago (e.g., Hon

Sao, Hon Doi Moi, Hon Da Le), but still endemic to Rach

Gia Bay. Due to the small sizes of the islands and the lim¬

ited exchange between populations the genetic variabil¬

ity and thus the effective population size is assumed to

be quite low (Charlesworth 2009). We thus recommend

future studies on population genetics of C. psychedelica ,

which can shed more light into island ecology of tropical

lizards in general.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Ngo et al.

Fig. 5. A. Introduced macaque on Hon Khoai Island; B. Trapped Water Monitor Lizard for consumption; C. Deer killed by blasting

of granite outcrops; D. Naturally occurring but injured C. psychedelica. Photos H.N. Ngo.

The present population estimation suggests a stable

and actively reproducing population of C. psychedelica

at least on Hon Khoai Island. While densities of the spe¬

cies at untouched sites were found to generally have in¬

creased from wet to dry season, we observed a decrease

of the individual number in the areas, which were most

strongly affected by habitat degradation. We assume that

the current habitat destruction as well as the planned de¬

velopment of ecotourism will probably interfere with C.

psychedelica natural populations, because the species

was found to flee hastily in response to the presence of

humans. Touristic activities and the presence of humans

were already found to negatively affect other range re¬

stricted lizards such as Shinisaurus crocodilurus or Gon-

iurosaurus species in northern Vietnam (Ngo et al. 2016;

van Schingen et al. 2015). Regarding the small size of

Hon Khoai Island, the availability of remaining alterna¬

tive sites for C. psychedelica is limited. Since the species

was only discovered in 2010, long-term population data

is lacking and long-term consequences of habitat altera¬

tion are not yet investigated in detail.

Furthermore, the lizard’s striking coloration attracts

increasing interest in C. psychedelica in the international

pet trade. Even though public access to and the export

of wild fauna for commercial purposes from Hon Khoai

Island is prohibited and the collecting and caging of wild

animals must be permitted by authorized state bodies,

in accordance with the Law on Forest Protection and

Development No. 29/2004/QH11, unsustainable inter¬

national trade of C. psychedelica has been confirmed in

recent studies (Altherr 2014; Auliya et al. 2016; Nguyen

et al. 2015). Cnemaspispsychedelica is currently mainly

traded through internet platforms, where animals fetch

extremely high prices up to $3,500/ pair (Aulyia et al.

2016).

In addition to online trading, we also observed C. psy¬

chedelica on the world’s biggest reptile market in Hamm,

Germany in November 2014. The same dealer told us in

December 2015, that he was going to receive several new

pairs of C. psychedelica in the beginning of 2016. There

is already evidence for the existence of several indepen¬

dent traders offering C. psychedelica and an increasing

demand for the species, especially in Europe and the

US (e.g., Altherr 2014; Auliya et al. 2016; Nguyen et al.

2015). Even though the trade in the species as well as the

habitat destruction only started and the wild C. psyche¬

delica population still appears to remain stable, recent

examples on other lizard species underline the risk of

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Population assessment of the Psychedelic Rock Gecko in southern Vietnam

World Association ol

Zoos and Aquariums

W A Z A I United lor

Conservation*

DRAGON

CAPITAL

KOLNER ZOO

RuFFordl

*VAR

Federal Ministry lor the

Environment, Nature Conservation,

Building and Nuclear Safety

Chung tay bao ve loai

Tic ke duoi vang d dao Hon Khoai

;tion of the Psychedelic Rock Gecko on Hon Khoai Island

Protection

Ten khoa hoc:

Cnemaspis psychedelica Grismer, Ngo & Grismer, 2010

Mdi diftrc mo tii nam 2010

Hien chi ghi nhan of dao Hon Khoai

HSy chung tay bao v| loai T^c ke duoi vang

Bao ton sinh canh song:

- Hay bao ve rirng tren dao Hon Khoai

• Hay cung nhau phong chong chay rirng

- Hay bo rac vao thung de khong lam anh hu'dng den sinh

canh song ciia loai tac ke

Khong san bit tic ke va cac loai dong vat hoang da

Scientific name:

Cnemaspis psychedelica Grismer, Ngo & Grismer, 2010

A recently discovered species in 2010

Currently known only from Hon Khoai Island

Together we protect the Psychedelic Rock Gecko

Habitat protection:

- Protect forest of Hon Khoai Island

- Prevent forest fire

- Put the garbage into wastebasket

Don't collect geckos and other wildlife species

Fig. 6. Signboard handed over to the Forest Protection Department of Ca Mau Province to point to the threats and conservation

needs of the Psychedelic Rock Gecko in English and Vietnamese languages. Layout and text by Thai Do, Anna Rauhaus, Khoi Vu

Nguyen, Truong Quang Nguyen, and Thomas Ziegler. Photos T.O. Nguyen (habitat) and T. Ziegler (geckos).

local extinction in particular of range restricted and spe¬

cialist species. Cases of rapid local extirpations shortly

after discovery due to over-collection have been recorded

for other charismatic lizard species in the region, such as

Goniurosaurus luii and Shinisaurus crocodilurus (e.g.,

Auliya et al. 2016; Stuart et al. 2006; van Schingen et al.

2015) . Learning by experience, we recommend to initiate

immediate conservation actions, as long as access to the

island is still limited and wild populations are still intact.

As a first measure, a Psychedelic Rock Gecko breed¬

ing facility was recently built on the mainland of southern

Vietnam by Wildlife at Risk (WAR) in cooperation with

the Institute of Ecology and Biological Resources (IEBR)

and the Cologne Zoo, to establish an insurance popula¬

tion in captivity as basis for a potential future conserva¬

tion breeding program (Ziegler et al. 2015; Ziegler and

Nguyen 2015). In March 2015, a small breeding group

of C. psychedelica was transferred from Hon Khoai Is¬

land with relevant permits provided by the government

authorities to the Psychedelic Rock Gecko breeding fa¬

cility, where breeding has been succeeded (Ziegler et al.

2016) . In order to improve the conservation breeding

and long-term management of the species, comprehen¬

sive research on its ecology and habitat requirements is

currently being conducted by our team. This knowledge,

in concert with the present first population analysis and

evaluation of threats, will provide crucial baseline data

for further conservation measures. Recently, based on

our first research results, C. psychedelica was included

in the IUCN Red List of Threatened species, listed as En¬

dangered (Nguyen et al. 2016). To further enhance the

protection status and control the international trade in the

species, the listing of C. psychedelica on the Appendices

of the Convention on International Trade in Endangered

Species of Wild Tauna and Tlora (CITES) is strongly

recommended, since the rising international demand in

the species has posed increasing pressure on wild popu¬

lations.

To raise awareness, a campaign was recently initiated

by our team and signboards highlighting the conservation

needs of C. psychedelica were provided to the provincial

authorities to be shown on Hon Khoai Island (Tig. 6). Lo¬

cal rangers were equipped with camera and GPS devices,

and funds were raised to install big waste buckets along

forest paths to reduce pollution on the island. Moreover,

first discussions on the potential establishment of a na¬

ture reserve on Hon Khoai Island to facilitate long-term

habitat and species protection recently were held with the

Torest Protection Department of Ca Mau Province and

the joint planning of further activities for habitat and spe¬

cies protection took place in March 2016 (Ziegler and

Nguyen 2016).

Acknowledgments. —We thank the directorates of the

Torest Protection Department of Ca Mau Province for

their strong cooperation, the support of field work, and

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Ngo et al.

issuing relevant permits. We are grateful to H.V. Le and

B. T. Nguyen (FPD of Ca Mau Province), K.V. Nguyen

from Wildlife at Risk (WAR, HCM City), T. Pagel, and

C. Landsberg (Cologne Zoo), M.T. Nguyen (VNMN, Ha¬

noi), and T.H. Tran (IEBR, Hanoi), as well as A. Rauhaus

(Cologne Zoo), and M. Bonkowski (University of Co¬

logne) for their support of this research and conservation

work in Viet Nam. We thank M.D. Le (Hanoi), V.Q. Luu

(Hanoi), and U. Schepp (Bonn) for commenting on a first

draft of the manuscript. Field work on Hon Khoai was

supported by the Rufford Foundation (Grant No. 18631-

2) and the Federal Ministry for the Environment, Nature

Conservation, Building and Nuclear Safety (BMUB).

Cologne Zoo is partner of the World Association of Zoos

and Aquariums (WAZA): Conservation projects 07011,

07012 (Herpetodiversity Research, Amphibian and Rep¬

tilian Breeding and Rescue Stations).

Literature Cited

Altherr S. 2014. Stolen Wildlife - Why the EU needs to

tackle smuggling of nationally protected species. Re¬

port by Pro Wildlife, Munich, Germany. 29 p.

Auliya M, Altherr S, Ariano-Sanchez D, Baard EH,

Brown C, Cantu J-C, Gentile G, Gildenhuys P, Hen-

ningheim E, Hintzmann J, Kanari K, Krvavac M,

Lttink M, Lippert J, Luiselli L, Nilson G, Nguyen

TQ, Nijman V, Parham J, Pasachnik SA, Pedrono M,

Rauhaus A, Rueda D, Sachnez M-E, Schepp U, van

Schingen M, Scheeweiss N, Segniagbeto GH, Shep¬

herd C, Stoner S, Somaweera R, Sy E, Turkosan O,

Vinke S, Vinke T, Vya R, Williamson S, Ziegler T.

2016. Trade in live reptiles, its impact on wild popula¬

tions, and the role of the European market. Biological

Conservation Available: http://dx.doi.Org/l0.1016/j.

biocon.2016.05.017 [Accessed: 25 September 2016],

Charlesworth B. 2009. Effective population size and pat¬

terns of molecular evolution and variation. Nature Re¬

views 10: 195-205.

Grismer LL, Ngo, VT. 2007. Four new species of the

gekkonid genus Cnemaspis Strauch 1887 (Reptilia:

Squamata) from Southern Vietnam. Herpetologica

63(4): 482-500.

Grismer LL, Ngo VT, Grismer JL. 2010. A new colorful

new species of insular rock gecko ( Cnemaspis Strauch

1887) from southern Vietnam. Zootaxa 2352: 46-58.

Ngo HN, Ziegler T, Nguyen TQ, Pham CT, Nguyen TT,

Le MD, van Schingen M. 2016. First population as¬

sessment of two cryptic Tiger geckos ( Goniurosau-

rus) from northern Vietnam: Implications for con¬

servation. Amphibian & Reptile Conservation 10(1):

34^15.

Nguyen, TQ, Ngo HN, Pham CT, van Schingen M,

Nguyen KV, Rauhaus A, Ziegler T. 2015. Population

assessment, natural history and threat evaluation of

the Psychedelic rock gecko ( Cnemaspis psychedeli-

ca ). Part I: Trade analysis, literature survey, own data;

October 2015. Unpublished report for the Federal

Ministry for the Environment, Nature Conservation,

Building and Nuclear Safety, Division Species Protec¬

tion, Bonn, Germany and for the Species Programme,

UNEP World Conservation Monitoring Centre, Cam¬

bridge, UK, 1-18.

Nguyen TQ, Ngo HN, Ziegler T, van Schingen M. 2016.

Cnemaspis psychedelica. The IUCN Red List of

Threatened Species 2016: e.T97210381A97210384.

[Accessed: 05 September 2016],

Reed DH, O’Grady JJ, Brook BW, Ballou JD, Frankham

R. 2003. Estimates of minimum viable population

sizes for vertebrates and factors influencing those es¬

timates. Biological Conservation 113: 23-34.

Stuart BL, Rhodin AG, Grismer LL, Hansel T.

2006. Scientific description can imperil spe¬

cies. Science 312: 1137. http://dx.doi.org/10.1126/

science.312.5777.1137b

Traill LW, Bradshaw CJA, Brook BW. 2007. Minimum

viable population size: A meta-analysis of 30 years

of published estimates. Biological Conservation 139:

159-166.

van Schingen M, Schepp U, Pham CT, Nguyen TQ,

Ziegler T. 2015. Last chance to see? Threats to and

use of the Crocodile Lizard. Traffic Bulletin 27:19-26.

Ziegler T, Nguyen TQ. 2015. Neues von den Forschun-

gs- und Naturschutzprojekten in Vietnam und Laos.

Zeitschrift des Kolner Zoos 58(2): 79-108.

Ziegler T, Nguyen TQ. 2016. Aktuelle Projekte zur Er-

haltung des Psychedelischen Felsengeckos. ZGAP

(Zoologische Gesellschaft fur Arten- und Popula-

tionsschutz e. V.) Mitteilungen 32(1): 24-27.

Ziegler T, Rauhaus A, Nguyen TQ, Nguyen KV 2015.

Aufbau einer Erhaltungszuchtanlage fur Echsen in der

Hon Me Station von Wildlife at Risk in Siidvietnam.

ZGAP (Zoologische Gesellschaft fiir Arten- und Pop-

ulationsschutz e. V) Mitteilungen 31(1): 30-33.

Ziegler T, Rauhaus A, Nguyen KV, Nguyen TQ. 2016.

Building of a conservation breeding facility for the

Psychedelic Rock Gecko ( Cnemaspis psychedelica )

in southern Vietnam. Der Zoologische Garten N.F.

85: 224-239.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Population assessment of the Psychedelic Rock Gecko in southern Vietnam

r> ( 'i w

Hai Ngo Ngoc is a young scientist at the Vietnam National Museum of Nature since 2014. He has also

been involved with several research projects at the Institute of Ecology and Biological Resources (IEBR)

beginning in 2013. He received his M.Sc. degree in 2015 from the University of Science, Vietnam National

University, Hanoi. He has participated in numerous herpetological surveys in Vietnam and has important

experience in field research and conservation work. His focus is on the ecology, phylogeny, and conservation

of endemic and endangered reptile species of Vietnam.

Truong Quang Nguyen is a researcher at the Institute of Ecology and Biological Resources (IEBR), Vietnam

Academy of Science and Technology (VAST) and is a member of the Biodiversity and Nature Conservation

projects of the Cologne Zoo in Vietnam and Laos. He finished his doctoral degree in 2011 at the Zoological

Research Museum Alexander Koenig (ZFMK) and the University of Bonn, Germany (DAAD Fellow). From

2012 to 2014 he worked as a postdoctoral researcher in the Institute of Zoology, University of Cologne

(Alexander von Humboldt Fellow). He has conducted numerous field surveys and is co-author of eight books

and more than 200 papers relevant to the biodiversity research and conservation in Southeast Asia. His

research interests are systematics, ecology, and phylogeny of reptiles and amphibians from Southeast Asia.

Tan Van Nguyen is a young researcher at the Institute of Ecology and Biological Resources (IEBR), Vietnam

Academy of Science and Technology (VAST). He has participated in numerous herpetological surveys in

Vietnam and has experience in field research. He is interested in taxonomy, ecology, and conservation of

reptiles and amphibians in Vietnam.

Frank Barsch is a policy officer in the Division of Species Protection in the German Ministry for the

Environment, Nature Conservation, Building and Nuclear Safety (BMUB) working on national and

international aspects of species conservation and trade (e.g., in the context of CITES). Between 2004 and

2007 he worked for the German Development Services (DED) in the Department for Forestry and Range

Resources (Botswana) to improve the national management of natural plant resources. Between 2002

and 2004 he worked as a researcher and lecturer at the Institute for Ecology and Evolutionary Biology

(University of Bremen, Germany). He has furthermore extensive working experience with environmental

non-govemmental organizations (e.g., WWF).

Thomas Ziegler has been the Curator of the Aquarium/Terrarium Department of the Cologne Zoo since 2003

and is the coordinator of the Cologne Zoo’s Biodiversity and Nature Conservation Projects in Vietnam and

Laos. Thomas studied biology at the University Bonn (Germany), and completed his diploma and doctoral

thesis at the Zoological Research Museum Alexander Koenig in Bonn, with focus on zoological systematics

and amphibian and reptile diversity. He has been engaged with herpetodiversity research and conservation

in Vietnam since 1997. As a zoo curator and project coordinator he tries to combine in situ and ex situ

approaches, viz., to link zoo biological aspects with diversity research and conservation, both in the Cologne

Zoo, in rescue stations, and breeding facilities in Vietnam and in Indochina’s last remaining forests. Since

February 2009, he has been an Associate Professor at the Zoological Institute of Cologne University. Since

1994, Thomas has published 360 papers and books, mainly dealing with herpetodiversity.

Mona van Schingen is PhD. candidate at the Zoological Institute of the University of Cologne and the Cologne

Zoo, Germany. Since 2011, she has been investigating the herpetofauna of Vietnam, in the working group of

Thomas Ziegler and graduated in 2014 with her M.Sc. on the Crocodile lizard in Vietnam. She has conducted

diverse field excursions to Vietnam and is engaged in several research, conservation, and awareness projects

focusing on various species in Vietnam. Her current research focuses on ecology, population dynamics, and

conservation of endangered, specialist and range restricted lizard species in Vietnam.

Amphib. Reptile Conserv.

September 2016 | Volume 10 | Number 2 | e124

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [General Section]: 27-29 (el29).

Short Communication

A record of the Balkan Stripe-necked Terrapin,

Mauremys rivulata (Testudines: Geoemydidae)

from the Azov Sea Coast in the Crimea

^leg V. Kukushkin and 2 Daniel Jablonski

1 Department of Herpetology, Zoological Institute of Russian Academy of Sciences, Universitetskaya Emb. 1, 199034 Saint Pe¬

tersburg, RUSSIA 2 Department of Zoology, Comenius University in Bratislava, Mlynska dolina, Ilkovicova 6, 842 15 Bratislava,

SLOVAKIA

Keywords. Mauremys rivulata , first record, Crimea, Kerch peninsula, Azov Sea, overseas dispersal, occasional

relocation

Citation: Kukushkin 0 V, Jablonski D. 2016. A record of the Balkan Stripe-necked Terrapin, Mauremys rivulata (Testudines: Geomydidae) from the

Azov Sea Coast in Crimea. Amphibian & Reptile Conservation 10(2) [General Section]: 27-29 (el29).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any

medium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and

authorized publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal

website <amphibian-reptile-conservation.org>.

Received: 03 September 2016; Accepted: 7 November 2016; Published: 30 November 2016

The Crimean herpetofauna comprises such true Eastern-

Mediterranean species as Mediodactylus kotschyi and

Zamenis situla (Sillero et al. 2014). The occurrence of

these species in the Crimea is isolated and could sug¬

gest Late Pleistocene-Holocene range expansion across

the short-existing land bridge between the Anatolia, the

Crimea, and the Balkans or even ancient human-mediat¬

ed dispersal. At the same time, there are some other ther¬

mophilic species ( Triturns karelinii , Podarcis tauricus,

Pseudopus apodus, Dolichophis caspius, Elaphe sauro-

mates) that probably colonized the Crimea during former

interglacial epoch from the southern refugia (Kukushkin

2013a).

It is well known that the distribution range of the Eu¬

ropean Pond Turtle Emys orbicularis includes the Crimea

(Szczerbak 1966; Fritz et al. 2009). Here we report a re¬

cord of another pond turtle from the Azov Sea region of

the Crimea. During field work an adult female of Maure¬

mys rivulata (Valenciennes, 1833) was captured on Cape

Khroni in north-east of the Kerch Peninsula between the

villages Yurkino and Osoviny on June 19, 2016 (circa

11 a.m., Moscow time). The terrapin was found among

several individuals of E. orbicularis near the shore of a

small and shallow natural lake with Phragmites austra¬

lis (less than 1 m in depth and about 10 m in diameter).

The water body is located on the lower terrace of seaside

landslide, just a few meters above sea level (45.43267°N,

36.59960°E; Fig. 1A). There are outcrops and heaps of

limestone rocks on the abrasion-accumulative sea coast

below the lake (Fig. IB). In general, the locality remains

typical of habitats of M. rivulata within the area of its

natural distribution.

The water body is weakly streaming due to the cold-

water source. According to our observations, the local

herpetofauna comprises two species of amphibians ( Bn-

fates viridis, Pelophylax cf. bedriagae ) and six species

of reptiles (P. apodus , E. sauromates , D. caspius , Natrix

natrix, N. tessellata , and E. orbicularis). The local E.

orbicularis population is quite numerous, since at least

12 specimens have been observed. The Red-eared Slider

(Trachemys scripta elegans) is absent in this lake, al¬

though this alien species has been found in many points

of the Crimea including the city of Kerch. In May 2016

2-3 T. scripta adults and up to 19 E. orbicularis speci¬

mens were counted on 400-500 m along the Melek-

Chesme river enclosed in the concrete channel in the

central area of Kerch.

The coloration and pattern of M. rivulata is typical for

the species (Fig. 2A-C). The terrapin has a total straight-

line carapace length of 131.3 mm, body mass of 354.7 g,

and was healthy and strong.

The natural distribution range of M. rivulata is con¬

fined to the Eastern Mediterranean region, with the

northern distribution limit at 43°N in coastal Croatia and

at about 42°N in the Bulgarian Black Sea coast (Sindaco

and Jeremcenko 2008). The species is widespread along

Correspondence. Emails: 1 viper a_kuk@ukr.net (Corresponding author); 2 darnel.jablonski@balcanica.cz

Amphib. Reptile Conserv.

November 2016 | Volume 10 | Number 2 | el 29

Kukushkin and Jablonski

Fig. 1. M. rivulata capture locality at Kerch peninsula. Eastern Crimea: A. small natural lake; B. Azov sea coast opposite to the lake.

Fig. 2. M. rivulata from the Crimea: A. dorsal view; B. ventral view; C. details of the head coloration.

the Southeast European and West Asian sea coasts, in¬

cluding those of the Marmara Sea and the Bosporus (Tok

and £igek 2014). The records closest to the Crimea (with

the minimum distance of about 440 km) are known to the

west from Eregli (Kocaeli peninsula) in the Northwest¬

ern Anatolia (Fritz and Freytag 1993; Fritz et al. 2008).

Undoubtedly, our single record of this species does

not allow any inferences about the origin of the individ¬

ual. However, Kerch city is a large merchant port and

this circumstance increases the probability of an acciden¬

tal translocation. It should be noted that several turtles

identified as Caspian Stripe-necked Terrapins, Mauremys

caspica (Gmelin, 1774) were recorded in the Sevastopol

area in 1980s (Kukushkin 2013b). These specimens were

probably brought to the Crimea from the Transcaucasian

region, where M. caspica is fairly widespread. However,

in contemporary interpretation these records may be also

related to M. rivulata.

On the other hand, it is well known that M. rivulata

occurs in habitats along sea coasts and enters brackish

water (e.g., Broggi 2012). Moreover, this species is ca¬

pable of extensive transoceanic dispersal (Mantziou et al.

2004; Vamberger et al. 2014). Thus, we also can not ex¬

clude the possibility of natural overseas dispersal across

the Black Sea, taking advantage of surface sea currents

carrying water from the coast of Northwestern Anatolia

to the Southwestern Crimea similarly to Caretta caretta

or Chelonia mydas. These sea turtles were recorded on

the Caucasian coast of the Black Sea and even in the

Kerch Strait (Malandzia et al. 2012; Pestov and Kletnoy

2012). However, due to the single M. rivulata record and

considerable distance from the distribution range of the

species, the probability of transmarine migration is sup¬

posed to be low. In any case, our finding indicates that

monitoring of terrapin populations throughout the Black

Sea coasts would be beneficial for a better understanding

of overseas dispersal in M rivulata and reveal possible

shifts of northern boundary of its distribution.

Acknowledgments. —We are grateful to Pavel Ruch-

ko (Kerch) and Ilya Turbanov (Borok, Russia) for the

data on population density of the exotic turtle T. scripta

in Kerch. Our special regards are to Yuliya Krasylenko

(Kyiv, Ukraine) for the significant assistance in field re¬

search at Kerch peninsula during 2012-2016, and to Uwe

Fritz for his valuable comments and suggestions regard¬

ing the manuscript content.

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FF, Aloufi A A, Auer M, Rifai F, Silic T, Hundsdorfer

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AK. 2008. Go east: phylogeographies of Mauremys

caspica and M. rivulata — discordance of morphol¬

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21: 527-540.

Fritz U, Ayaz D, Hundsdorfer AK, Kotenko T, Guicking

D, Wink M, Tok CV, giqek K, Buschbom J. 2009. Mi¬

tochondrial diversity of European pond turtles (Emys

orbicularis ) in Anatolia and the Ponto-Caspian Re¬

gion: Multiple old refuges, hotspot of extant diversi¬

fication and critically endangered endemics. Organ¬

isms, Diversity & Evolution 9: 100-114.

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tofauna: a new vision of the problem. Pp. 22-25 In:

Commemorative zoological readings. Proceedings of

the International scientific conference devoted to 100 th

anniversary of SL Delyamure and 90 th anniversary of

SA Skryabin (December 5, 2013, Simferopol). Sim¬

feropol.

Kukushkin OV. 2013 b. Adventive herpetofauna of the

Ukraine. Pp. 25-27 In: Commemorative zoological

readings. Proceedings of the International scientific

conference devoted to 100 th anniversary of SL Dely¬

amure and 90th anniversary of SA Skryabin (Decem¬

ber 5, 2013, Simferopol). Simferopol. [In Russian],

Malandzia VI, Dbar RS, Solomko MO, Pestov MV. 2012.

Finding of the sea turtle Chelonia mydas in the eastern

part of the Black Sea. Current Studies in Herpetology

{Sovremennaya gerpetologiya) 12 (3/4): 155-157. [In

Russian],

Mantziou G, Rifai L. 2014. Mauremys rivulata (Valen¬

ciennes in Bory the Saint-Vincent 1833) - Western

Caspian Turtle, Balkan Terrapin. Pp. 080.1-080.9

In: Chelonian Research Monographs. No 5. Conser¬

vation Biology of Freshwater Turtles and Tortoises:

A Compilation Project of IUCN/SSC Tortoise and

Freshwater Turtle Specialist Group. Editors, Rhodin

AGJ, Pritchard PCH, van Dijk PP. et al. Chelonian

Research Foundation, Lunenburg, Massachusetts.

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Mylonas M. 2004. The inter- and intraspecific status

of Aegean Mauremys rivulata (Chelonia, Bataguri-

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Herpetological Journal 14: 35—45.

Pestov VV, Kletnoy MV. 2012. Finding of the sea turtle

Caretta caretta at the Russian coast of the Black Sea.

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petologiya) 12 (3/4): 158-159. [In Russian]

Sillero N, Campos J, Bonardi A, Corti C, Creemers R,

Crochet PA, Crnobmja-Isailovic J, Denoel M, Ficeto-

la GF, Gongalves J, Kuzmin S, Lymberakis S, de Pous

P, Rodriguez A, Sindaco R, Speybroeck G, Toxopeus

B, Vieites DR, Vences M. 2014. Updated distribution

and biogeography of amphibians and reptiles of Eu¬

rope. Amphibia-Reptilia 35: 1-31.

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Western Palearctic. 1. Annotated checklist and distri¬

butional atlas of the turtles, crocodiles, amphisbae-

nians and lizards of Europe, North Africa, Middle East

and Central Asia. Edizioni Belvedere, Latina. 579 p.

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Crimea (Herpetologia Taurica). Naukova Dumka,

Kiev, Ukraine. 239 p.

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Province of (/anakkale (Marmara Region, Turkey).

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P, Fritz U. 2014. Massive transoceanic gene flow in a

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mys rivulata). Zoologica Scripta 43: 313-322.

Oleg V. Kukushkin (born in Sevastopol, November 8, 1973) graduated from M.V. Fmnze Simferopol State

. Md & P pt University in 1991-1995. For the past 15 years he's been a research scientist of the Department of Biodiversity

Studies and Ecological Monitoring of T.l. Vyazemski Karadag Scientific Station - Nature Reserve (the Crimea).

He currentl y working on his Ph.D. thesis titled “Herpetofauna of the Crimean peninsula: Distribution patterns,

K. TES morphology, biology, genesis pathways, and protection” at the Department of Herpetology, Zoological Institute

R uss i an Academy of Sciences (St. Petersburg). He has published over 200 research and popular science

papers, mostly in the herpetological field. The recent monograph ( Pysanets E., Kukushkin O. 2016. Amphibians

of the Crimea. Kyiv: National Academy ofScienses of Ukraine, National Museum of Natural History. 320 p.) is

among the latest. He is a member of the A.M. Nikolsky Russian Herpetological Society and the Ukrainian Herpetological Society.

He is the official expert at the herpetological section of Rare and Endangered Species Commission (Russian Federation). Primary

research areas: Distribution, zoogeography, biology, ecology, systematics, and conservation of reptiles and amphibians of the

Crimean peninsula. His main scientific interests are . Regularities in the formation of distribution ranges of the Crimean herpetofauna

representatives; revealing of relationships of the Crimean amphibians and reptiles populations using molecular and genetic methods

in combination with current paleogeography data on the Black Sea region; genesis of the Crimean biota.

Daniel Jablonski is currently a Ph D. student of Zoology and Evolutionary Biology at the Comenius University

in Bratislava, Slovakia. He has been interested in amphibians and reptiles since early chilhood. His research

interests concern evolutionary and historical biogeography questions relating to the origin and distribution of

genetic diversity and its conservation in natural populations of amphibians and reptiles. His special focus is placed

in the Balkan Peninsula, one of the most important evolutionary areas in Europe. In parallel, it led to the the

establishment of the first online herpetofauna mapping of this area in the project: www.Balcanica.info. He loves

traveling and photography.

Amphib. Reptile Conserv.

November 2016 | Volume 10 | Number 2 | el29

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

10(2) [General Section]: 30-33 (el27).

Short Communication

Range extension and some morphological characteristics of

Ptychoglossus brevifrontalis, Boulenger, 1912 (Squamata:

Alopoglossidae) in Suriname

^awien Jairam and 2 Sabitrie Jairam-Doerga

1 Anton de Kom Universiteit van Suriname, National Zoological Collection of SURINAME 2 Anton de Kom Universiteit van Suriname, National

Herbarium of SURINAME

Abstract .—We herein document the range extension and morphological features of the second

specimen of Ptychoglossus brevifrontalis from Suriname, collected at the Brownsberg Nature Park,

extending the range of this species by approximately 372 km from the previous known record in

Suriname. Morphological features, apart from lamellae under fourth finger and scales around mid¬

body, fall within the range of characteristics recorded for this species.

Keywords. Herpetofauna, lizards, Brownsberg Nature Park, morphology, geography,

Citation: Jairam R, Jairam-Doerga S. 2016. Range extension and some morphological characteristics of Ptychoglossus brevifrontalis, Boulenger,

1912 (Squamata: Alopoglossidae) in Suriname. Amphibian & Reptile Conservation 10(2) [General Section]: 30-33 (e127).

NonCommercialNoDerivatives 4.0 International License, which permits unrestricted use for non-commercial and education purposes only, in any me¬

dium, provided the original author and the official and authorized publication sources are recognized and properly credited. The official and authorized

publication credit sources, which will be duly enforced, are as follows: official journal title Amphibian & Reptile Conservation ; official journal website

<amphibian-reptiie-conservation.org>.

Received: 10 June 2016; Accepted: 08 October 2016; Published: 31 December 2016

Introduction

Although the Herpetofauna of Suriname has been studied

to some extent (Hoogmoed 1973; Ouboter and Jairam

2012) new country records and or range extensions

are still documented. A recent example is the new

country record of Amapasaurns tetradactylus found

during a Rapid Assessment Program of Conservation

International at the Grensgebergte (Jairam and Jairam-

Doerga 2015). At present, the genus Ptychoglossus

comprises 15 valid species (Goicoechea et al. 2016). Of

these 15 species, P. brevifrontalis is known for having

the broadest geographic distribution (Peloso and Avila-

Pires 2010). A first revision of the genus was done by

Harris (1994) who reported a lower number of mid body

scales in the specimens from Peru, Bolivia, and the

single specimen from Suriname, in relation to specimens

from other localities. A subsequent study by Peloso and

Avila-Pires (2010) showed that P. brevifrontalis present

geographic variation in the number of scale counts, but

without a geographic pattern that could indicate specific

differentiation. The first specimen of P. brevifrontalis

in Suriname catalogue number BMNH 1939.1.1.75,

male, was collected in 1939 near the border with Brazil,

head of the Kutari River. The second individual of

P. brevifrontalis was collected at the Brownsberg Nature

Park (BNP) 01 November 2014 in a dried up depression

that is used by the park management to collect water

which is then pumped to the various tourist lodges in the

park. This paper highlights the range extension of the

lizard species Ptychoglossus brevifrontalis , previously

known from only one specimen (BMNH 1939.1.1.75),

collected in Suriname near the border with Brazil, head

of the Kutari river, and presents some morphological

information on the current collected specimen.

Material and Methods

Located in the north eastern part of Suriname, and

slightly northwest of the Brokopondo Reservoir, the

Brownsberg Nature Park (BNP) (4°56 , N 55°10’W) is

operated under the authority of the Foundation for Nature

Preservation in Suriname, also known as STINASU

(Lim et al. 2005). Located at a distance of approximately

100 km from the capital (De Dijn et al. 2007), BNP is the

only nature park in Suriname (Ouboter and Jairam 2012)

and one of the most visited nature locations, both by locals

and foreigners (ter Steege et al. 2004; Love et al. 2007).

Correspondence. Emails: 1 rawien_2000@yahoo.com (corresponding author); 2 jairam-doerga@uvs.edu

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | el27

Jairam and Jairam-Doerga

58°W 57°W 56°W 55°W 54°W

Occurence of Ptychoglossus brevifrontalis in Suriname

58°W 57°W 56°W 55°W 54°W

I Kilometers

Fig. 1. Map showing the occurrence of P. brevifrontalis in Suriname. The blue dot represents the first recorded specimen from

Suriname while the red triangle depicts the specimen collected at the BNP. Fig. 2. Lateral view of head of P. brevifrontalis showing

the labials. Fig. 3. Anal scales of P. brevifrontalis.

The different habitats found on the Brownsberg make

this an interesting place in terms of research purposes.

Some of these habitats are undisturbed streams and

forests, and some are streams disturbed by mining and

habitats of recently disturbed forests (De Dijn et al.

2007). The faunal diversity is high and represented by

approximately 116 species of mammals, 387 species of

birds, and 144 species belonging to the herpetofauna

(De Dijn et al. 2007). The collected specimen of

P. brevifrontalis extends the species range approximately

372 km in northeastern direction, measured from the last

known record of this species in Suriname. The specimen

was captured by hand during a survey conducted on a

trail going towards the Leo falls (Fig. 1). The specimen

was euthanized using lidocaine, subsequently placed

in four percent formaldehyde, and later transferred to

70% ethanol for long term preservation and storage at

the National Zoological Collection of Suriname. Species

identification was facilitated by examination of the

following traits: total length (from tip of the snout to the

tip of the tail), snout vent length (from the tip of the snout

to the cloaca), the number of supraoculars, supra labials,

infra labials (Fig. 2), chin shields, lamellae under fourth

finger, and fourth toe and anal scales (Hoogmoed 1973;

Avila-Pires 1995). Sex was detennined by the absence of

precloacal pores (Peloso and Avila-Pires 2010). Figure 3

gives an overview of the anal region.

Results

One juvenile female of P. brevifrontalis, voucher

number NZCS R679, was collected on 01 November

2014. The location where the specimen was collected

was covered with dried fallen leaves and consisted of

Fig. 4. Habitat where Ptychoglossus brevifrontalis was

collected at the BNP. Picture by A. Gangadin.

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | e127

Range extension of Ptychoglossus brevifrontalis in Suriname

Table 1 . A comparison of the P. brevifrontalis specimen collected at the BNP with the specimen from the head of the Kutari river

and the specimen described by Avila-Pires (1995).

Traits

NZCS R679

BMNH 1939.1.1.75

Avila-Pires (1995)

Total length (mm)

51.9

Snout vent length (mm)

25.4

60 (RMNH 26390)

Transverse dorsal scales

31-33

Transverse ventral scales

18-19

Scales around mid body

30-34

Supralabials

Infralabials

Lamellae left finger

10-12

Lamellae left toe

18-19

14-17

a muddy substrate interspersed with sand (Fig. 4). The

vegetation around the dry creek bed consists of high

forests with large tree species like Ceiba pentandra

(Malvaceae), Lecythis sp. (Lecythidaceae) and Couratari

sp. (Lecythidaceae), with some signs of disturbance. The

understory and herb layer is open with a few saplings

and seedlings. The herb layer in and around the creek

bed consists of some fern species, Piper sp. (Piperaceae)

and Centropogon sp. (Campanulaceae) (Fitzgerald

et al. 2002; ter Steege et al. 2004). Table one gives an

overview of the measurements and scale counts for

NZCS R679 and at the same time compares NZCS

R679 with the specimen from the head of the Kutari

river (BMNH 1939.1.1.75) and the specimen described

by Avila-Pires (1995). From the information presented

in the table and taking into account the morphological

variation described by Peloso and Avila-Pires (2010),

we tentatively conclude that the collected specimen

from the BNP falls within the range of features for

P. brevifrontalis. The only difference is noted in the

number of lamellae under the fourth finger and the scales

around mid body. A possible explanation for this might be

that scale counts were not made using the same method

or might be due to intraspecific variation, given that most

specimens in collections are from the western and central

part of the Amazon (Peloso and Avila-Pires 2010).

Discussion

The presence of this specimen at the Brownsberg Nature

Park in the north eastern part of Suriname is evidence

that the herpetofauna of Suriname is yet to be fully

investigated in order to attain a more comprehensive

overview of this group. A comparison of the single

specimen collected at the Brownsberg Nature Park

with the specimen described by Hoogmoed (1973)

on the basis of traits specified in table one shows no

significant differences that would suggest a different

taxon. Considering the distance of more than 300 km

between the two records in Suriname we are confident

that this species is present in more locations and that the

range of this species is underestimated. A more thorough

survey of the habitat requirements of this species coupled

with different sampling methods such as pitfall trapping

should elucidate the actual distribution of this species in

Suriname.

Acknowledgments —We would like to thank

STINASU for providing permits and allowing us to

conduct surveys at the Brownsberg Nature Park. A first

draft of this paper was reviewed by Teresa C.S. Avila-

Pires while Anielkoemar Gangadin kindly took pictures

of the location where the specimen was collected.

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Rawien Jairam is an assistant at the National Zoological Collection of Suriname. He has a

M.S. in Conservation Biology and has been interested in the herpetofauna of Suriname for

many years. Apart from general herpetology, he is specifically interested in taxonomy, species

descriptions, and distribution.

Sabitrie Jairam-Doerga has a M.S. in Conservation Biology and is working at the National

Herbarium Suriname at the Anton de Kom University of Suriname. She is involved in fieldwork,

both terrestrial and aquatic and has been an active participant on several joint expeditions to

the interior of Suriname.

Amphib. Reptile Conserv.

December 2016 | Volume 10 | Number 2 | e127