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Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [Special Section]: 1-14 (el 06).

The anuran fauna of a West African urban area

v N’Goran Germain Kouame, 2 3 Caleb Ofori-Boateng, 2 3 Gilbert Baase Adum,

4 Germain Gourene, and 5t Mark-Oliver Rodel

1 Jean Lorougnon Guede University, Department of Biology and Animal Physiology, UFR-Environnement, Daloa, BP 150 COTE D’IVOIRE

2 Forestry Research Institute of Ghana, PO. Box 63, Fumesua, Kumasi GHANA 3 Department of Wildlife and Range Management, Faculty of

Renewable Natural Resources, Kwame Nkrumah University of Science and Technology, Kumasi GHANA 4 Nangui Abrogoua University, Laboratoire

d’Environnement el de Biologie Aquatique, UFR-SGE, 02 BP 801, Abidjan 02 COTE D’IVOIRE 5 Museum fur Naturkunde, Leibniz Institute for

Evolution and Biodiversity Science, Invalidenstrasse 43, 10115 Berlin GERMANY

Abstract . — Reported are the results of an amphibian survey in the district of Daloa and surroundings,

in central-western Ivory Coast. Spanning a three year period, we investigated two general areas,

each during the rainy and dry seasons. During 62 days of field work 30 anuran species were

recorded. The urban environment mainly contained widespread anuran species with preferences

for savannah-dominated landscapes and farmbush habitats. The recorded total anuran species

richness in the urban area exceeded the diversity in the savannah islands/forest mosaic bordering

the Daloa district. This indicates many savannah species may do well in urban situations. However,

this higher species richness was only due to one site that possessed particularly diverse amphibian

breeding sites, thus illustrating the necessity of maintaining suitable habitats for a wide-range of

species. One of the most surprising findings was Kassina schioetzi, a species usually difficult to

find in its natural habitat. In Daloa it seems to have successfully adapted to the urban conditions.

Although the anuran richness in the Daloa area was relatively low compared to other Ivorian humid

savannah areas, it supported an important part of the countries amphibian diversity. Nevertheless

the forest habitats, and specifically the forest islands bordering the Daloa district, should be

considered sensitive conservation areas.

Key words. Amphibians, conservation status, Cote dTvoire, Upper Guinea, urban ecology

Citation: Kouame NG, Ofori-Boateng C, Adum GB, Gourene G, Rodel MO. 2015. The anuran fauna of a West African urban area. Amphibian &

Reptile Conservation 9(2) [Special Section]: 1-14 (el 06).

NoDerivatives 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: 1 6 June 2015; Accepted: 29 October 201 5; Published: 1 6 November 2015

West Africa has been experiencing intensive urbaniza-

tion to such a point that human modified landscapes are

gradually taking over the majority of natural landscapes,

in particular native forests (Deikumah and Kudom 2010;

Bible 2013). Whereas various Ghanaian forests are pro-

tected and/or sustainably managed (Adum et al. 2013;

Ofori-Boateng et al. 2013), only very few Ivorian for-

est remnants receive sufficient protection and sustainable

management (e.g., see Mayaux et al. 2004). The Ivorian

population has exploded over the past four decades, tri-

pling from 6.7 million in the early 1970s to approxi-

mately 22 million people today (Bible 2013), has largely

accelerated an urbanization process causing massive en-

vironmental damage. The gradual disruption of forests,

has worsened in several forested areas of the Ivory Coast

during the prolonged political crisis in the first decade of

the 21 st century, has mainly stemmed from increase land

demand for agriculture and urbanization (Bible 2013;

Hansen et al. 2013).

The Haut-Sassandra region is traditionally an im-

portant trading center, particularly for cocoa production

in the Ivory Coast, has attracted 44.8% of national and

23.4% of foreign farmers (Assiri et al. 2009). During the

country’s 2010-2011 post election violence, Daloa, the

third largest city of the country and the regional capital

of the Haut-Sassandra region, became a refuge for peo-

ple from the northern, central, and western Ivory Coast,

resulting in a rapid urbanization process. As one result,

forests surrounding the city are increasingly fragmented.

To enhance the protection of biological diversity, the

Ivorian Ministry of Scientific Research has therefore re-

cently recommended the collection of scientific informa-

tion to update the biodiversity data of the Haut-Sassandra

region. As data for amphibians were still lacking, we sur-

Correspondence. Emails: *ngoran_kouame@yahoo.fr; t mo.roedel@mfn-berlin.de (corresponding authors).

Amphib. Reptile Conserv. 1 November 2015 | Volume 9 | Number 2 | el 06

Kouame et al.

veyed the amphibian fauna within the district of Daloa

and its surroundings, and herein report for the first time

an assessment of the species richness and composition of

the anuran fauna in a West African urban area.

Methods

Study area. Daloa is the third largest city in the

Ivory Coast and the regional capital of the Haut-

Sassandra region. It is situated in central- west-

ern Ivory Coast (06 o 53 , 01.8 ,, -06 o 94 , 97.8 ,, N;

006°25’65.3”-006°68'89.0” W), in the transition zone

between semi-deciduous forest and humid Guinea savan-

nah. The town is an important trading center, particularly

for cocoa. The region has a mean annual temperature

of 26.3 °C; the annual precipitation ranges from 1,200

to 1,700 mm. The climate includes a long rainy season

(April to June) with the highest precipitation peak in

June, a short dry season (July to August) alternating with

a short rainy season (September to October), and a long

dry season (November to March). The relative mean hu-

midity is 75% (Eldin 1971).

Description of the survey areas (Fig. 1). Our defini-

tion of an urban area follows McDonnell and Pickett

(1993) and Demographia (2008), i.e., taking into con-

sideration a minimum density of 400 humans/km 2 and

other factors such as density of buildings, roads and

other infrastructure. We surveyed two general areas: 1)

the district of Daloa (urban area), and 2) the savannah

islands/forest mosaic bordering the Daloa district (non-

urban aspect). Our surveys were covering a three year

period (see Appendix 1 for further details). We inves-

tigated four sites inside the urban area namely: Balou-

zon (Bal: 45 ha), Eveche (Eve: 50 ha), Gbokora (Gbo:

80 ha), and Tazibouo (Taz: 100 ha). As a comparison,

we surveyed Sapia (Sap: 150 ha) and Zaibo (Zai: 190

ha), two non-urban sites in the savannah islands/forest

mosaic adjacent to urban Daloa (see Appendix 1). The

Balouzon and Eveche areas were mainly characterized

by unpaved roads, a swampy area used for vegetable cul-

tivation, and a concentration of buildings. A large stream,

bordered by coconut trees and grasses, was used for fish-

ing activities. The vegetation in Gbokora was dominated

by grasses and a semi-deciduous forest interspersed by

a highway. Some swampy areas surveyed in this site

were being used for vegetable cultivation. This area was

noisy due to heavy traffic. A concentration of buildings

and streetlights were characteristic of the Gbokora site.

The Tazibouo site mainly consisted of unpaved roads,

buildings, semi-deciduous forest adjacent the Daloa Jean

Lorougnon Guede University, and some construction

sites. This site also comprised the Theological and Pas-

toral Institute of Daloa whose garden was dominated by

bamboo, other grasses, and several stands of different or-

Fig. 1 . Typical aspects of habitats of the urban Daloa; a = concentration of houses in a high population zone; b = degraded forest

on the periphery of the urban area; c = highway crossing degraded forest and farmbush; d = amphibian breeding pond in the urban

environment.

Amphib. Reptile Conserv.

2

November 2015 | Volume 9 | Number 2 | el 06

The anuran fauna of a West African urban area

namental plants. Some water bodies, i.e., two large per-

manent ponds situated near a roadside and bordered by

grasses, were encountered. The ponds served as a water

point for cattle. A few termite mounds were also present.

A swampy area investigated was used for rice and veg-

etable cultivations. Some parts of this area were light-

ened by streetlights at night. The sites Sapia and Zaibo,

adjacent to the district of Daloa represented non-urban

conditions. However, they had lost the majority of their

natural forest cover, resulting in an overall change from

a forest to a savannah-dominated landscape. Both sites

mainly consisted of farmbush, small farms, coffee, and

cocoa plantations. Some swamps that were part of these

two sites had been converted to rice fields. Small forest

islands were still encountered at Zaibo, but fewer forest

islands were left at the Sapia site.

Field work, sampling effort and vouchers. Amphib-

ians were mainly located opportunistically, during visual

and acoustic surveys of all available habitats by NGK.

Surveys were undertaken daily between 07:00-11:00

and 18:00-22:00 GMT over a total of 62 days (see Ap-

pendix 2) at all general survey areas. A hand-held GPS

receiver (Garmin 12XL) was used to record the geo-

graphical positions of all study sites. The searching tech-

niques used included acoustic surveying, visual scanning

of terrain and refuge examination (e.g., lifting logs and

rocks, peeling away barks, scraping through leaf litter,

looking around or within burrows, and termite mounds).

Amphibians encountered were not marked and repeated

sightings thus cannot be excluded. As we only include

presence/absence data and not abundances in our analy-

ses this seem to be of negligible importance.

Below we comment only on a few species being re-

markable concerning their distribution, taxonomy, biol-

ogy or threats, or being particularly typical for the urban

amphibian fauna. The nomenclature used herein follows

the taxonomy by Frost (2015). After capture, frogs were

identified to species level, measured, sexed, and if not

kept as vouchers, released in their respective habitats.

Snout-urostyle-length (SUL) was taken with a dial cali-

per (accuracy ±0.5 mm). Records of Xenopus muelleri

were based on visual observations only. For all other spe-

cies we deposited vouchers at the Jean Lorougnon Guede

University, Daloa, Ivory Coast (see Appendix 3). Frog

vouchers were euthanized humanly in a 1,1,1-Trichloro-

2-methyl-2-propanol hemihydrate (MS 222) solution and

thereafter preserved in 70% ethanol.

Statistics. We used the daily species lists to calculate

the sampling efficiency. We calculated the estimated spe-

cies richness with the Chao2 and Jack-knife 1 estimators

(software: Estimates, Colwell 2006). These estimators

are incidence based, calculating with the presence/ab-

sence data of the daily species lists (62 days of survey

work) for 30 anuran species. To avoid order effects we

accomplished 500 random runs of the daily species lists.

The Sprensen’s Similarity Index (/?) was used to deter-

mine the extent of similarity between the two main sur-

veyed areas (herein the district of Daloa and the savan-

nah islands/forest mosaic bordering the Daloa district; f

may vary from 0 to 1 (Sprensen 1948; Wolda 1981).

Results

Species richness and faunal similarities

In total we recorded 30 anuran species (Table 1). Acous-

tics indicated more than one Arthroleptis species live in

our area. So far, it is not possible to separate taxa from

Table 1. Anuran species recorded in the urban and non-urban areas of Daloa, with sites (see Appendices 1-3), general habitat prefer-

ence and distribution range. S = savannah, FB = farmbush (degraded forest and farmland), F = forest, A = Africa (occur also outside

West Africa), WA = West Africa (defined as the area west of the Cross River in Nigeria), UG = Upper Guinea (forest zone west of the

Dahomey Gap), E = endemic to Ivory Coast and eastern Guinea, * = taxon comprise complex of several species, ** = records on this

survey comprise several species (according to acoustics).

Family / Species

Site

Habitat

Distribution

S

FB

F

A

WA UG E

Arthroleptidae

Arthroleptis spp.**

Bal, Eve, Gbo, Sap, Taz, Zai

X

- X (?)

Leptopelis spiritusnoctis

Zai

X

—

X — —

L. viridis

Bal, Eve, Gbo, Taz

X

—

X

— — —

Bufonidae

Amietophrynus maculatus

Gbo, Sap, Taz, Zai

X

—

X

— — —

A. regularis

Taz

X

—

X

— — —

Dicroglossidae

Hoplobatrachus occipitalis

Hemisotidae

Bal, Eve, Gbo, Sap, Taz, Zai

X

—

X

— — —

Hemisus marmoratus

Taz, Zai

X

—

X

— — —

Hyperoliidae

Afrixalus dorsalis

Bal, Eve, Gbo, Sap, Taz, Zai

X

Amphib. Reptile Conserv.

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November 2015

| Volume 9 | Number 2 | el 06

Kouame et al.

Table 1 (continued). Anuran species recorded in the urban and non-urban areas of Daloa, with sites (see Appendices 1-3), general

habitat preference and distribution range. S = savannah, FB = farmbush (degraded forest and farmland), F = forest, A = Africa (occur

also outside West Africa), WA = West Africa (defined as the area west of the Cross River in Nigeria), UG = Upper Guinea (forest zone

west of the Dahomey Gap), E = endemic to Ivory Coast and eastern Guinea, * = taxon comprise complex of several species, ** =

records on this survey comprise several species (according to acoustics).

Family / Species

Site

Habitat

Distribution

Hyperoliidae (cont.)

S

FB

F

A

WA

UG

E

Hyperolius concolor concolor

Bal, Eve, Gbo, Sap, Taz, Zai

X

—

X

—

H. fusciventris fusciventris

Sap

X

—

X

—

H. guttulatus

Zai

X

—

X

—

H. nitidulus

Bal, Eve, Gbo, Sap, Taz

X

—

X

—

H. picturatus

Gbo, Sap, Taz, Zai

—

X

—

X

—

H. sp.

Taz

—

X

—

X

—

Kassina schioetzi

Taz

X

—

X

—

K. senegalensis

Sap, Taz

X

—

X

—

Phrynobatrachidae

Phrynobatrachus calcaratus *

Sap

X

P francisci

Bal, Taz

X

—

X

—

P gutturosus*

Sap, Taz, Zai

X

—

X

—

P latifrons

Bal, Eve, Gbo, Sap, Taz, Zai

X

—

X

—

Phrynomeridae

Phrynomantis microps

Taz

X

Pipidae

Xenopus muelleri

Taz

X

Ptychadenidae

Ptychadena bibroni

Bal, Eve, Gbo, Sap, Taz, Zai

X

Ptychadena mascareniensis *

Bal, Eve, Gbo, Sap, Taz, Zai

X

—

X

—

Ptychadena oxyrhynchus

Sap, Taz, Zai

X

—

X

—

Ptychadena tournieri

Sap, Taz

X

—

X

—

Ptychadena pumilio

Bal, Eve, Gbo, Sap, Taz, Zai

X

—

X

—

Ptychadena tellinii

Taz

X

—

X

—

Ranidae

Amnirana albolabris

Sap, Zai

X

A. galamensis

Taz

X

—

X

—

the Arthroleptis poecilonotus - complex based on mor-

phology. They can be distinguished by advertisement call

and genetic characters. However, assigning populations,

based on these characters, to available names (indistin-

guishable museum types without molecular data) is not

possible (for a short review of the taxonomic situation in

West African Arthroleptis spp. see Rodel and Bangoura

2004). We thus provisionally lumped all records of this

genus as Arthroleptis spp. A list of recorded anurans with

site records, known habitat preference and their distribu-

tion ranges is given in Table 1 .

Based on the daily species lists we calculated our

sampling efficiency. The Jack-knife 1 estimator calcu-

lated 33 anuran species, the Chao 2 estimator estimated

3 1 species for the study area. We hence recorded almost

the entire (94% and 99%, respectively) estimated species

Amphib. Reptile Conserv. 4

richness. More than one fifth of the encountered species

(seven spp., 23%; Table 1) depend on forest but tolerate

farmbush habitats (degraded forest). Nine species (30%)

are very closely associated with savannah habitats. Thir-

teen species (43%) exhibit a strong preference for savan-

nah and farmbush habitats and are normally not found

in forest. Four species (13%) do not occur outside West

Africa [defined as the area west of the Cross River in Ni-

geria; see Penner et al. (2011)], and are often restricted to

smaller parts of West Africa. Seven of all recorded spe-

cies (23%) occur only in the Upper Guinea forest zone

(forests west of the Dahomey Gap). The total number of

species recorded in the district of Daloa was 25, while the

species richness in the adjacent savannah/forest mosaic

was 21. However, the high species number for Daloa was

mainly due to one site (Tazibouo). When excluding this

November 2015 I Volume 9 I Number 2 I el 06

The anuran fauna of a West African urban area

Table 2. S0rensen’s similarity values for pairwise comparisons

of the anuran community between the six surveyed sites (see

text and Appendix 1).

Sites

Eve

Gbo

Taz

Sap

Zai

Bal

0.95

0.87

0.61

0.62

0.59

Eve

-

0.91

0.57

0.64

0.61

Gbo

-

0.65

0.67

0.71

Taz

-

0.65

0.63

Sap

-

0.71

site diversity was higher in the savannah/forest mosaic.

The number of species common to both areas was 16

(Sbrensen’s Similarity Index /?: 0.70). Within the district

of Daloa we recorded 11 species in Balouzon, 10 species

in Eveche, 12 in Gbokora, and 25 in Tazibouo. Within the

savannah/forest mosaic we recorded 18 and 16 species in

Sapia and Zaibo, respectively.

The results of the Sbrensen’s similarity for pairwise

comparisons in the six surveyed sites are presented in

Table 2. At least more than 50% of the recorded species

were similar between sites. The anuran fauna of Daloa

urban area was most similar to that of the Comoe Na-

tional Park, a savannah area in northern Ivory Coast (/?:

0.72). With 68% and 66% faunal similarity the Lamto

Faunal Reserve and the Marahoue National Park, which

are situated in the same vegetation zone as Daloa, were

very similar to the Daloa fauna. Other Ivorian protected

areas such as the Mont Peko and Mont Sangbe National

Parks comprise savannah and real rainforest zones and

thus consequently differed in their faunal composition,

compared to Daloa (Table 3).

Species accounts

Amietophrynus regularis (Reuss, 1833) - The genus

Amietophrynus currently encompasses 40 species of true

African toads [Frost 2015; although this list also con-

tains non-vaild taxa such as Amietophrynus chudeaui

(Chabanaud, 1919) see Rodel (2000)]. Amietophrynus

regularis has a wide distribution in Africa and inhabits

a broad range of habitats from moist and dry savannahs,

montane grassland, forest margins, and agricultural habi-

tats, as well as human settlements, often in association

with rivers (Rodel 2000; Channing and Howell 2006). In

our urban sites A. regularis (Fig. 2) seemed to reach its

highest abundances directly around human settlements.

At night, it was found in gardens, around houses, park-

ing areas, buildings, or below streetlights, preying mostly

on insects. During the day, it was found under rocks or

logs. The most imminent threat to the toad’s survival in

the city of Daloa is its exploitation for scientific courses

at the university. Every year several hundred individu-

als are collected by students and subsequently killed and

dissected in anatomy courses. This exploitation seems to

have reached a point where the species is becoming rare

in the city. However, concerning the entire range of the

species, it is very common and of Feast Concern (IUCN

2015).

Hoplobatrachus occipitalis (Gunther, 1859) is the

most commonly consumed frog species in West Africa.

The frog trade varies regionally from e.g., local scale in

Burkina Faso, to intensive cross-border trade in north-

ern Benin and Nigeria (Mohneke et al. 2009, 2010). The

consumption of H. occipitalis (Fig. 3a) has recently in-

creased to a considerable extent in the Ivory Coast where

this species is an important component of animal protein

in some local populations (NGK, unpubl. obs.). In Daloa,

the trade of H. occipitalis mainly took place on a local

scale at the different markets of the district. Usually a

batch of five adult specimens was sold for 500.00 FCFA

(app. 0.84 USD). Frog meat are sold fresh (Figs. 3b, c)

or dried (Fig. 3d). It is used in soups, stews, or sauces by

the local populations. The local price in Daloa markets

was mean to low compared to prices recorded in Burkina

Faso and Nigeria, respectively. According to Mohneke

et al. (2010), in Burkina Faso, the price for one frog de-

pended on its size and varied between 25.00 FCFA for

a small frog, up to 250.00 FCFA (0.05 USD and 0.50

USD) for a large one. In Nigeria, they reported one bag

containing at least 1,000 frogs cost 26.94-40.40 USD on

purchase and 40.40-67.34 USD at sale. In the urban area

of Daloa hard data on harvested frog numbers and re-

spective consequences for the local populations are lack-

ing. The local trade of H. occipitalis hence needs more

attention and detailed investigation.

Hyperolius concolor concolor (Hallowell, 1844)(Fig.

4) is a typical West African farmbush species living in

degraded forest of the forest zone and gallery forests in

the savannah zone (Schiotz 1967; Rodel 2000). It seemed

to do very well under urban condition and was hence

among the most widespread species recorded in the ur-

Table 3. Sprensen’s similarity value (ft) between the anuran fauna of the Daloa urban area and other Ivorian areas, and respective

species richness; twenty-five species were recorded in urban Daloa (this study); NP= national park; FR= faunal reserve.

Area

Species richness

Number of species common

with the Daloa urban area

/3-value (Sorensen)

Source

Comoe NP

33

21

0.72

Rodel and Spieler (2000)

Lamto FR

40

22

0.68

Adebaet al. (2010)

Marahoue NP

33

19

0.66

Rodel and Ernst (2003)

Mont Peko NP

33

11

0.38

Rodel and Ernst (2003)

Mont Sangbe NP

45

20

0.57

Rodel (2003)

Amphib. Reptile Conserv.

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November 2015 | Volume 9 | Number 2 | el 06

Kouame et al.

Fig. 2. Amietophrynus regularis female recorded in the garden of the Theological and Pasto-

ral Institute of Daloa.

ban sites of Daloa (Table 1); it was particularly abundant

among grasses near ponds. In the rainy season we record-

ed some, presumably migrating, individuals on windows,

balconies, and in houses.

Hyperolius sp. - The genus Hyperolius Rapp, 1842 is

one of the most diverse African anuran genera with cur-

rently approximately 28 species occurring in West Africa

(Schiptz 1967, 1999; Frost 2015). A major taxonomic

problem is many species of this genus are highly variable

(e.g., Schiptz 1999). On 15 September 2013, at around

07h00 GMT, we found a Hyperolius on humid ground in

the Tazibouo site, within the garden of the Theological

and Pastoral Institute, after it had rained heavily the night

before. Our individual lacked a vocal sac and gland and

hence is either female or juvenile (Fig. 5). It resembles

either a juvenile H. picturatus or a newly metamorphosed

individual phase J of H. sylvaticus ivorensis, which is

normally brownish to green with paired undelimited dor-

solateral stripes, and an hourglass pattern (Schiptz 1999).

The size of our reed frog was 20 SUL, thus exceeding the

size of freshly metamorphosed Hyperolius of most spe-

cies (compare e.g., Schiotz 1967; Rodel 2000). Its dorsal

surface was beige with a greenish grey hourglass pattern.

The iris was golden, the anterior and posterior sides of

pupil were red. The ventral surface was whitish. Without

having male specimens and advertisement calls available

it cannot be decided if this frog represents an undescribed

species or only an atypical, but known Hyperolius spe-

cies.

Kassina schioetzi Rodel, Grafe, Rudolf, and Ernst,

2002 was known so far from the Mont Peko National

Park, the Marahoue and Comoe National Parks, and the

Lamto Faunal Reserve, all situated in the Ivory Coast

Amphib. Reptile Conserv. 6

(Rodel et al. 2002; Rodel and Ernst 2003; Adeba et al.

2010). It lives along the savannah forest edge, reaching

into the savannah zone along rivers. The species may

also occur in Bia National Park, western Ghana, but a

voucher from there exhibited a mixture of characters

with K. cochranae (Hillers et al. 2009). Kassina schioetzi

is usually hard to find in all localities so far investigated

(see above and own experience of the authors). In the

district of Daloa (Tazibouo), some males were observed

calling at night from more exposed sites (Fig. 6). We also

encountered a small number of other males calling in a

bamboo patch within the Theological and Pastoral Insti-

tute, and at a grassy roadside in the vicinity of a large

pond. Our recorded males measured 32.1 ±1.6 (SUL, n

= 4), thus being within the known range of K. schioetzi

(Rodel et al. 2002).

Leptopelis viridis (Gunther, 1868) (Fig. 7) is one of

the most characteristic species inhabiting the West Af-

rican savannahs and the degraded areas of the former

rainforest belt. As a synanthropic species, it also lives in

villages (Schiotz 1967; Rodel 2000). It is one of the most

widespread anurans in the urban sites of Daloa. Leptope-

lis viridis was found around houses, and in gardens. The

majority of the recorded males were found at night call-

ing exposed on the ground between short grasses, which

is in contrast to the calling sites in natural habitats. There

the species calls, often from high perch sites, in bushes

and trees (Grafe et al. 2000; Rodel 2000).

Phrynomantis microps Peters, 1875 is a medium-

sized microhylid frog inhabiting the savannah regions of

West Africa (Hirschfeld and Rodel 2011) where it hides

in burrows or empty termite mounds during the day and

the dry season. The frog was also observed to occupy and

November 2015 I Volume 9 | Number 2 | el 06

The anuran fauna of a West African urban area

live essentially unharmed in the nest of the highly ag-

gressive ant species - Paltothyreus tarsatus (e.g., Rodel

and Braun 1999; Rodel et al. 2013). In Daloa, P. microps

was heard calling at night in tufts of grass around houses

after heavy rainfalls. In the garden of the Theological and

Pastoral Institute, a calling male was observed in associa-

tion with an Emperor Scorpion (Pandinus imperator ) in

a hole behind the wall of a building. The association of P.

microps with scorpions has also been reported by Rodel

and Braun (1999) and Rodel (2000). We captured another

male (Fig. 8) on 08 September 2013 around 22h00 GMT

at the edges of a wide roadside pond beside the Theologi-

cal and Pastoral Institute.

Xenopus muelleri (Peters, 1844) is an aquatic species

inhabiting the West African savannah ponds of highly

variable size during the rainy season and the edges of riv-

ers during the dry season (Rodel 2000). In the urban site

Tazibouo, the frog was observed to live in holes drilled

in the ground by the national company of water distribu-

tion. The depths of these holes varied from 0.7-1.20 m.

Discussion

Despite their importance to ecosystem functions

(Mohneke and Rodel 2009; Hocking and Babbitt 2014),

amphibians are still among the least studied vertebrates

particularly in urban and suburban areas in the tropics

(Hamer and McDonnell 2008; Pickett et al. 2001). Al-

most 85% of amphibian species threatened by urbaniza-

tion are encountered in the tropics (IUCN, Conservation

International and NatureServe 2006). Many factors are

known to negatively influence the herpetofauna inhabit-

ing big cities. Among these factors are habitat loss, habi-

tat fragmentation, isolation, pollution, over harvesting,

and road traffic (Hammer and McDonnell 2008; Perry et

al. 2008; Stuart et al. 2008; Deikumah and Kudom 2010;

Tonini et al. 2011). However, many species are able to

adapt to urban conditions and sometimes urban areas

may even surprise with the discovery of scientifically

new species (Newman et al. 2012; Feinberg et al. 2014;

Howlader et al. 2015). This also concerns the Ivorian

city of Abidjan where a monotypic genus Morerella cy-

anophthalma and a night-frog Astylosternus laticephalus

have recently been discovered and described (Rodel et

al. 2009, 2012).

With its geographic position in a transition zone be-

tween the semi-deciduous forest and humid savannah,

we expected the urban landscape of Daloa region to pro-

mote a diverse amphibian fauna. However, the overall

species richness (30 spp.) was lower compared to the

species richness recorded in western, central, and north-

ern Ivorian savannah areas, for instance the Mont Sangbe

National Park (45 species, Rodel 2003), Lamto Faunal

Reserve (40 species, Adeba et al. 2010), Marahoue and

Mont Peko National Parks (33 species for each park,

Rodel and Ernst 2003), or the Comoe National Park (33

Fig. 3. Hoplobatrachus occipitalis from the district of Daloa (a) and a woman trading this species on a local market (b); batches of

five adult specimens, fresh (b and c) or dried (d), were sold for 500.00 FCFA (app. 0.84 USD).

Amphib. Reptile Conserv. 7 November 2015 | Volume 9 | Number 2 | el 06

Kouame et al.

Fig. 4. A calling Hyperolius concolor concolor male recorded at the garden of the Theological and Pastoral

Institute of Daloa.

Fig. 5. Dorsolateral view of a juvenile Hyperolius sp. with uncertain taxonomic status from the urban Daloa.

species, Rodel and Spieler 2000). Compared to these and

other West African savannah areas with known amphib-

ian assemblages such as north-western Benin (Nago et al.

2006), east-central Guinea (Greenbaum and Carr 2005),

central-northern Guinea (Hillers et al. 2008a), or eastern

Ghana (Leache et al. 2006), the urban landscape of Daloa

ranks among the West African areas of medium to low

amphibian species richness. While we recorded few for-

est related species e.g., Amnirana albolabris , Leptopelis

spiritusnoctis, Phrynobatrachus calcaratus, and P. gut-

turosus (Rodel and Branch 2002; Assemian et al. 2006;

Kouame et al. 2014; Kpan et al. 2014; the latter two spe-

cies comprising out of cryptic species with savannah and

forest specialists), most of the recorded frogs were wide-

spread species with preferences for savannah-dominated

landscape and farmbush habitats. The six surveyed sites

all shared at least half of their species with all other sites.

We observed the highest species richness at the Tazibouo

site (25 spp.) which was the only urban site comprising

various suitable breeding habitats. For instance in the

Amphib. Reptile Conserv.

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November 2015 I Volume 9 | Number 2 | el 06

The anuran fauna of a West African urban area

Fig. 6. Kassina schioetzi from Daloa urban area; shown is a male calling from the ground (a), and its whitish venter and almost

circular gular gland (b) characteristic for the species.

garden of the Theological and Pastoral Institute, ephem-

eral and perennial wetlands, well-suited for the co-ex-

istence of species with different reproductive strategies

were present: e.g., very small temporary breeding sites

(. Phrynobatrachus spp.), larger, almost permanent breed-

ing sites {Afrixalus dorsalis , Hyperolius concolor, Kas-

sina schioetzi, K. senegalensis, Phrynomantis microps ),

and humid places for species with terrestrial direct de-

velopment ( Arthroleptis poecilonotus- group). This gar-

den also played an important role in providing refuge for

several other species in particular during the heat of the

day and the dry season.

In addition to the fact that many amphibian species

depend on different but complementary habitats (e.g.,

aquatic sites for the tadpoles, terrestrial site of the meta-

morphosed individuals), their populations are usually

structured as meta-populations (Pope et al. 2000; Marsh

and Trenham 2001). Urbanization and in particular frag-

mentation and isolation of habitats by roads and other

urban infrastructure is reducing the connectivity of popu-

lation networks (Vos and Chardon 1998). Hence, we ex-

pected to record lower amphibian diversity in the district

of Daloa than in the savannah-forest mosaic adjacent to

this district. Surprisingly, the total anuran richness in

the urban environment was higher than in the adjacent

savannah-forest mosaic. This result indicates many am-

phibian species may survive under urban situations, such

as in the district of Daloa. However, this high total spe-

cies richness was due to only one of four urban sites, i.e.,

the Theological and Pastoral Institute, comprising many

different habitat types and particularly diverse breeding

sites. The other urban sites actually had slightly lower

species richness than the non-urban sites. This illustrates

a high amphibian diversity in urban areas may be main-

tained and even exceed such as of nearby non-urban

areas; however, this can only be achieved by offering a

wide range of different habitats suitable for various am-

phibian species.

Apart from roads potentially reducing or ceasing gene

flow, amphibians further face direct threats in urban ar-

eas, in the form of the collection of anurans for anatomy

and food consumption. Compared to European towns

(e.g., Mollov 2005), however, there are still plenty of

habitats available to amphibians, the traffic is usually less

intense as many of the roads remain unpaved allowing

frogs to cross. In fact our non-urban sites were all within

a matrix of agricultural land and thus most likely prone

to a variety of pesticides which could be a higher threat

than the threats experienced in towns. The adaptability of

amphibians within the urban development seemed to be

species-specific and was highly variable even between

sites. For example some species such as Hyperolius

guttulatus, H. fusciventris fusciventris, Amnirana albo-

labris, Leptopelis spiritusnoctis, and Phrynobatrachus

calcaratus, encountered in the savannah-forest mosaic

outside of Daloa were never found in the urban sites. This

is most likely due to the fact that their specific habitats

are no longer present. For instance Hyperolius guttulatus

breeds almost exclusively in very large and deeper ponds

(Rodel 2000; Schiotz 1967, 1999); and Phrynobatrachus

calcaratus typically lives at rain forest edges or in gallery

forests in the savannah zone (Rodel 2000). Respective

habitat types for both latter species were not found in the

urban environment. It is known that in forested areas the

alteration of the microclimate, due to degradation of the

vegetation structure, causes a shift in species composi-

tion (Ernst and Rodel 2005, 2006; Hillers et al. 2008b;

Ofori-Boateng et al. 2013). Such effects might be even

worse in the usually more open habitats of urban areas.

Conclusion

The study is indicating that an unexpected high number

of anuran species seem to be able to survive in a current

African city. However, this is not the case for all species.

For those species the protection of natural forest and sa-

vannah ecosystems is very important. The forest habi-

tats, and specifically the forest “islands” bordering the

Daloa district, should thus be considered sensitive areas

and dispersal corridors need to be maintained. Within the

Amphib. Reptile Conserv.

9

November 2015 | Volume 9 | Number 2 | el 06

Kouame et al.

Fig. 7. Dorsolateral view of Leptopelis viridis, one of the most

widespread anurans from the Daloa urban area.

urban areas, the availability of a diverse set of habitats

is a prerequisite for the maintenance of high amphibian

species richness.

Acknowledgments. — We are indebted to Dago Gna-

kri, President of the Jean Lorougnon Guede University,

for providing authorization to undertake this survey. We

thank Daplex H. Ouenchist, Director of the Theological

and Pastoral Institute for permitting us to investigate the

garden of his institution. We are particularly grateful for

the support and collaboration from Chief Nanan Kra, el-

der of the Baoule-Ayetou from the Haut-Sassandra re-

gion.

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N’Goran Germain Kouame is an Ivorian herpetologist and biologist. He is lecturer at the Jean Lorougnon

University, Daloa, Ivory Coast and a member of the IUCN SSC Amphibian Specialist Group (ASG). He holds

a Diploma and a Ph.D. in natural sciences from the University of Abobo-Adjame, Abidjan, Ivory Coast, where

he used leaf-litter frogs ( Phrynobatrachus spp.) as models to determine the conservation status of the Banco

National Park, one of the rare remaining primary forests situated in the midst of a West African mega-city. His

current research interests focus on the taxonomy, ecology, distribution, and conservation of rare, threatened, and

new amphibian species in Ivory Coast.

i A Caleb Ofori-Boateng is a research scientist at the Forestry Research Institute of Ghana. He holds a Ph.D. in

wildlife management and a B.S. in Natural Resources Management from the Kwame Nkmmah University of

Science and Technology in Ghana. His research focuses on ecology, population genetics and conservation of

West African amphibians. Caleb is also the founder and Director of Help Conservation Ghana (Herp-Ghana), a

non-profit organization dedicated to amphibian and reptile conservation in West Africa.

fa# r -\ r ,

Gilbert Baase Adum is a research scientist based at Ghana’s premiere science university Kwame Nkrumah Uni-

versity of Science and Technology in Kumasi. He is also the co-founder and Executive Director of SAVE THE

FROGS! Ghana, West Africa’s first non-profit organization dedicated exclusively to amphibian conservation.

Through his work with SAVE THE FROGS! Ghana, Gilbert aims to help prevent the extinction of endangered

frogs, while spreading the message of frog conservation and environmental protection across the entire African

continent. He is currently working on a collaborative project at the Museum fur Naturkunde, Berlin, with the aim

of establishing knowledge about impacts of climate change on Ghanaian amphibians.

Amphib. Reptile Conserv.

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November 2015 | Volume 9 | Number 2 | el 06

The anuran fauna of a West African urban area

a Germain Gourene is the professor and founder of the “Laboratoire d’Environnement et de Biologie Aquatique”

at the Nangui Abrogoua University (ex-University of Abobo-Adjame, Abidjan, Ivory Coast). His research fo-

cuses on the systematics and taxonomy of fishes with emphasis on Africa; areas covered also include ecology

and aquaculture. In addition to his research interest on fishes Germain is interested in the conservation of aquatic

invertebrates and amphibians in the Banco National Park. Germain has served as Vice-President and President

of the University of Abobo-Adjame for 10 years. He is embarking on a political career and has been elected as

deputy of the locality of Kounahiri since 2012.

Mark-Oliver Rodel is the Curator of Herpetology and head of the department of “Diversity Dynamics” at the

Museum fur Naturkunde, Berlin, and teaches biodiversity at the Humboldt University, Berlin. Since his teenage

age he has dedicated his life to the study of amphibians and reptiles, mostly to those from Africa. Mark-Oliver

is the Chairman for West and Central Africa within the IUCN SSC Amphibian Specialist Group (ASG). With his

team he investigates the taxonomy, systematics, and biogeography of amphibians and reptiles, but in particular

uses amphibians as model organisms in order to understand the effect of environmental change on species and

ecosystems.

Appendix 1 . Geographic position and short description of study sites in the Daloa study area.

Site

Latitude (N)

Longitude (W)

Elevation (m a.s.l.)

Habitat description

Bal

N06°53’64.5”

W006°25’65.3”

259

Highway; grassy habitats; heavy traffic; dense human

population

Large stream bordered by coconut trees and grass; un-

Eve

N06°53’01.8”

W006°26’05.9”

261

paved roads; concentration of buildings; swampy area

used for vegetable cultivation; dense human population

Semi-deciduous forest patch; swampy area dominated

Gbo 1

N06°54’15.8”

W006°27’15.3”

265

by grassy vegetation; buildings; highway, heavy traffic;

streetlight; swampy area partly used for vegetable cultiva-

tion; dense human population

Gbo 2

N06°54’03.4”

W006°27’09.6”

275

Buildings; shrubby vegetation; unpaved roads; highway;

dense human population

Sap 1

N06°87’20.8”

W006°37’83.8”

239

Subsistence farming; rice field in a swampy area; high

grassy vegetation

Sap 2

N06°87’22.8”

W006°37’93.7”

260

Forests islands; cocoa plantation at edge of a rice field;

high grassy vegetation

Sap3

N06°87’37.1”

W006°38’09.7”

276

Stream crossing cocoa plantation; palm tree at edge of the

water body; plantain and coffee plantations

Sap 4

N06°86’83.8”

W006°38’99.3”

244

Swampy area; high grassy vegetation; rice field; humid

savanna; tracks

Sap 5

N06°86’83.8”

W006°37’61.5”

229

Rice field in a swampy area; coconut trees at edge

Semi deciduous forest patch close to the Jean Lorougnon

Guede University; garden of the Theological and Pastoral

Taz 1

N06°90’42.8”

W006°43’97.4”

274

Institute, dominated by bamboo, grasses and stands of dif-

ferent ornamental plants; two large wide ponds surrounded

by vegetation (Asteraceae); streetlight; unpaved roads,

concentration of buildings; dense human population

Taz 2

N06°90’33.7”

W006°43’78.9”

268

Swampy area; buildings; rice field; vegetable cultivation;

many constructions of houses underway

Zai 1

N06°94’97.8”

W006°67’35.7”

223

Swamps within a semi deciduous forest; stream; ponds;

grassy vegetation; clearing; rice field; cocoa plantation

Zai 2

N06°94’33.6”

W006°68’89.0”

222

Very large rice field; high grasses; edge of coffee and

cocoa plantations; forest patch

Zai 3

N06°94’65.9”

W006°66’58.7”

209

Rice field crossed by a stream: coffee plantation; tracks;

forest patch

Amphib. Reptile Conserv.

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Kouame et al.

Appendix 2. Amphibian survey periods in the urban and non-urban areas of Daloa (compare Appendix 1).

Surveyed periods

Site

10-21 Aug.

27 Aug.-l

27 Sep.-l

16-25 Aug.

18-27 Oct.

18-27 Jun.

16-24 Aug.

2011

Sep. 2011

Nov. 2012

2013

2014

Urban area

6 days

2 days

1 day

6 days

2 days

6 days

3 days

Bal

6

2

1

6

2

6

3

Eve

6

2

1

6

2

6

3

Gbo

6

2

1

6

2

6

3

Taz

6

2

1

6

2

6

3

Non-urban area

6 days

4 days

8 days

4 days

6 days

Sap

3

2

4

2

3

Zai

3

2

4

2

3

Total days

12

6

5

10

9

Appendix 3. List of amphibian voucher specimens from the district of Daloa and surroundings. Given are field and collection num-

bers (NG), collection site (compare Appendix 1) and collection date.

Arthroleptidae: Arthroleptis spp.: NG001 (Taz, 08 Nov. 2011); NG002 (Sap, 16 Oct. 2013); NG003 (Zai, 23 Oct. 2013); Leptopelis

spiritusnoctis : NG004 (Zai, 23 Oct. 2013); L. viridis : NG005 (Taz, 24 Oct. 2011); NG006 (Taz, 30 Oct. 2013); Bufonidae: Ami-

etophrynus maculatus: NG007 (Zai, 25 Oct. 2013); A. regularis : NG008 (Taz, 19 Oct. 2011); Dicroglossidae: Hoplobatrachus oc-

cipitalis'. NG009 (Taz, 25 Oct. 2011); Hemisotidae: Hemisus marmoratus: NG010 (Taz, 08 Sep. 2010); NG011 (Sap, 25 Oct. 2013);

Hyperoliidae: Afrixalus dorsalis : NG012 (Taz, 18 Aug. 2011); Hyperolius concolor concolor. NG013 (Taz, 18 Aug. 2011); NG014

(Sap, 18 Oct. 2013); H. fusciventris fusciventris'. NG015 (Sap, 19 Oct. 2013); H. guttulatus : NG016 (Zai, 23 Oct. 2013); NG017

(Zai, 25 Oct. 2013); H. nitidilus: NG018 (Taz, 25 Aug. 2011); NG019 (Sap, 16 Oct. 2013); H. picturatus: NG020 (Taz, 01 Sep.

2011); NG021 (Sap, 18 Oct. 2013); NG022 (Taz, 23 Oct. 2013); H. sp.: NG023 (Taz, 15 Sep. 2013); Kassina schioetzi: NG024-027

(Taz, 08 Sep. 2013); K. senegalensis : NG028 (Taz, 17 Aug. 2011); NG029 (Taz, 30 Aug. 2013); Microhylidae: Phrynomantis

microps'. NG030 (Taz, 08 Sep. 2013); Phrynobatrachidae: Phrynobatrachus calcaratus : NG03 1-036 (Sap, 19-20 Oct. 2013); P.

franciscv. NG037-038 (Taz, 01 Sep. 2011); NG039 (Taz, 17 Aug. 2013); P. gutturosus : NG040 (Taz, 19 Aug. 2013); NG041 (Sap,

20 Oct. 2013); P latifrons : NG042 (Taz, 20 Aug. 2011); NG043 (Taz, 25 Aug. 2011); NG044 (Sap, 19 Oct. 2010); NG045 (Zai, 23

Oct. 2013); Ptychadenidae: Ptychadena bibroni : NG046 (Taz, 09 Sep. 2012); NG047 (Zai, 24 Oct. 2013); P. tellinii : NG048 (Taz,

01 Sep. 2013); P. mascareniensis: NG049 (Taz, 16 Sep. 2013); NG050 (Zai, 24 Oct. 2013); P oxyrhynchus: NG051-052 (Taz, 06

Sep. 2012); NG053 (Zai, 24 Oct. 2013); P pumilio : NG054 (Taz, 16 Sep. 2013); NG055 (Sap, 19 Oct. 2013); P tournierv. NG056

(Taz, 10 Sep. 2012); Ranidae: Amnirana albolabris'. NG057 (Sap, 19 Oct. 2013); A. galamensis : NG058 (Taz, 24 Aug. 2014).

Amphib. Reptile Conserv.

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November 2015 I Volume 9 | Number 2 | el 06

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [Special Section]: 15-38 (el 08).

The reptiles of the summits of Mont Oku and the Bamenda

Highlands, Cameroon*

Mvan Ineich, 2 Matthew LeBreton, 3 Nathaly Lhermitte-Vallarino, and 4 Laurent Chirio

1 Museum national d’Histoire naturelle, Sorbonne Universites, ISyEB (Institut de Systematique, Evolution et Bio diver site), UMR 7205 (CNRS,

MNHN, UPMC, EPHE), CP 30, 25 rue Cuvier, F-75005 Paris, FRANCE 2 Mosaic (Environment, Health, Data, Tech), PO Box 35322, Yaounde,

CAMEROON 3 Museum national d’Histoire naturelle, Departement de Regulation, Developpement et Diversite moleculaire, UMR CNRS-MNHN

7245 (Molecule de Communication et Adaptation des Microorganismes) , CP 52, 61 rue Buffon, F-75005 Paris, FRANCE 4 14 rue des Roses, 06130

Grasse, FRANCE

Abstract . — The list of the non-avian reptiles occupying the summits above 1,400 m elevation of

Mount Oku and the Bamenda Highlands in Cameroon comprise 50 species (one tortoise, 18 lizards,

and 31 snakes) belonging to 12 families and 29 genera. This assemblage has a high biogeographic

interest because it harbors species with a large altitudinal spectrum and several high elevation

endemic forms (submontane). Those species are currently severely threatened by human expansion

in the area. Human impacts include direct collections of several endemic species with a high

commercial value for the international pet trade, but most importantly deforestation and the growing

encroachment of people, cattle, and agriculture. Efficient actions are urgently needed to preserve

this unique heritage for future generations.

Key words. Biogeography, conservation, biodiversity, afro-montane herpetofauna

Citation: Ineich I, LeBreton M, Lhermitte-Vallarino N, Chirio L. 201 5. The reptiles of the summits of Mont Oku and the Bamenda Highlands, Cameroon.

Amphibian & Reptile Conservation 9(2): 15-38 (el 08).

NoDerivatives 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 April 2015; Accepted: 13 November 2015; Published: 1'

Introduction

African mountain formations clearly show an island-like

distribution pattern, which explains their high biogeo-

graphical disposition and the importance of those moun-

tain ranges for the conservation of their distinctive fauna

(Gartshore 1986; Fjeldsa and Lovett 1997). The Camer-

oon Volcanic Dorsal extends in its southern part for 800

km, and is represented by a succession of insular-like re-

liefs (true or continental islands). It begins with the is-

land of Annobon (elevation 655 m; Equatorial Guinea),

located more than 360 km from the African mainland,

and extends through the islands of Sao Tome (2,024 m),

Principe (948 m) and Bioko (formerly called Fernando

Poo; 3,106 m). It continues on to the mainland, includ-

ing the highest volcanic summit of Western Africa, Mount

(Mt.) Cameroon, which rises to 4,085 m. North of Mt.

Cameroon, emerge Mt. Nlonako (1,822 m), the impor-

tant volcanic range of Manengouba (2,411 m), and the

Correspondence. Email: ineich@mnlm.fr (Corresponding auth<

*This paper was written in homage to our late colleague and friend

!■ December 2015

reliefs of the Bakossi Highlands. North of those first re-

liefs stands an imposing orographic set which includes

most of the Highlands generally called the Bamenda

Highlands (BH). Towards the south it starts with a large

and elevated volcanic edifice, the Bamboutos Mountains

(2,740 m). Through the Santa Range (Mt. Lefo or Peak

of Santa, 2,550 m elevation), the Bamboutos Mountains

connect to the main peak, Mt. Oku, at 3,011 m. Elevations

then decrease relatively quickly before joining the north-

ern part of the Cameroon Volcanic Dorsal that ends with

the Tchabal Mbabo (2,460 m). The relief then undergoes

an eastern shift in their orientation, to fit the septentine

border of the Adamaoua, with the smaller peaks of Mt.

Alantika (1,885 m) and Mt. Mandara (1,442 m) separated

by the depression of the Benoue valley, which does not

exceed 150 m elevation.

The central axis of the Cameroon Volcanic Dorsal has

lateral extensions including more or less important bas-

tions, including on the western flank, Mts. Rumpi (1,764

>r)

Dr. Odile Bain (CNRS, MNHN).

Amphib. Reptile Conserv.

15

December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

m) and in Nigeria, Sonkwala (also called Obudu Plateau)

and Gotel (2,418 m). In the east stand Mts. Bana (2,097

m), Mbapit (1,989 m), Nkogam (2,263 m), and Mbam

(2,335 m). The majority of these mountains truly function

like islands for orophilous species because their elevation

is substantially higher than surrounding territories of low

elevation (at most 100 m), thus usually prohibiting the

faunal exchange of climatically demanding, orophilous

species between neighboring mountain ranges.

To the northeast of the Cameroon Volcanic Dorsal rests

the largely tabular area of the Adamaoua, a vast middle

mountain barrier extending from east to west. Mean ele-

vation of that central Cameroon relief stays relatively low

(about 1,100 m), but is contiguous with the high west-

ern ranges and functions as a faunistic exchange corridor,

creating a zone of biogeographical interest. It is indeed

increasingly recognized that under the colder climate of

the Plio-Pleistocene climatic oscillations, the Adamaoua

represent a refugia, an efficient “hyphen” between the

Cameroon Volcanic Dorsal and the mountains scattered

across the eastern edge of the Congo Basin in Eastern Af-

rica (Wagner et al. 2008; Barej et al. 2011). Some impos-

ing volcanic relief is strewed on the Adamaoua Plateau,

especially towards its septentine rim. The most impor-

tant, about 40 km east of the city of Ngaoundere, is the

Hossere Nganha, which reaches 1 ,923 m elevation and is

a location where some endemic species of reptile and am-

phibians are encountered (Amiet 1971; Ineich and Chirio

2004).

In Cameroon, the highest peaks (above 2,000 m) are

located at Mt. Cameroon, Mts. Bamboutos, Mt. Oku, and

at Tchabal Mbabo. With the exception of Mt. Cameroon,

those formations have been significantly degraded by

man and most often comprise only forest remnants within

montane grasslands grazed by the abundant cattle of the

Fulani herdsmen (Fig. 1).

Mt. Oku (rarely called Mt. Kilum: 6.12°N and 10.28°E,

elevation 3,011 m) is located in the most septentine part

of the BH, not far from the transition zone between moun-

tain forest and savanna. Summits above 2,800 m are cov-

ered with an afro-alpine grassy lawn (Fig. 1), devoid of

Fig. 1. The beautiful cattle of the Fulani herdsmen observed

on pastures high in the region of Mt. Oku are fat and healthy.

6.2 1°N and 10.44°E. Picture: I. Ineich, May 8, 2007.

trees, in which there is even a bog. The north side is home

to one of the best-preserved mountain forest fragments in

the region (Figs. 2, 3). An associated vegetation is also

found there including wet mountain forests, which are

well developed around Fake Oku (6°12’N and 10°27’E),

and a crater lake located about 2,300 m above sea level.

Another lake, Fake Bambili (5°56’N and 10°15’E), is

present in the region. Cattle herds are common around

the massif and even into the montane forest protected ar-

eas. These forests are important elements in the economy

and local culture as they allow the production of a wide

range of forest products essential to the survival of lo-

cal populations (wood, honey, and medicinal plants, e.g.,

Prunus africanus used in the treatment of prostate cancer

and subjected to strict control by the Washington Conven-

tion on International Trade of Endangered Species) (Figs.

4, 5). Scared and felled trees are visible even in the forest

reserves, and caused by the overflow of human activity

along the many forest paths and trails that allow easy ac-

cess (Macleod 1987) (Fig. 2).

This report provides a critical inventory of the reptile

species recorded from the summit area (above 1,400 m

elevation) of the BH, demarcated by the valley that sepa-

rates it from Manengouba/Mt. Cameroon (less than 700

m) and the Tikar Plain that separates it from Tchabal Mba-

bo (Fig. 6). We also discuss the biogeographic affinities

of the study region. Many of the reptiles found there are

Fig. 2. The path leading from Oku Elak village to the summit of

Mt. Oku is very popular and easy to access. Picture: I. Ineich,

May 6, 2007.

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

Fig. 3. Just before the summit of Mt. Oku, the vegetation is cov-

ered with dense epiphytic altitude plants. Picture: I. Ineich, May

6, 2007.

Fig. 4. Villagers apply strong pressure on the fauna and flora of

Mt. Oku forests. It is common to find traps to catch bush meat,

here a brush-tailed porcupine (a forest porcupine species). Pic-

ture: I. Ineich, May 6, 2007.

Fig. 5. Hives, placed high in the trees almost to the top of Mt.

Oku (here), produce a thick, white honey of excellent quality,

highly sought after. Picture: I. Ineich, May 7, 2007.

endemic mountain species whose distribution is restricted

and unfortunately now highly fragmented. Our knowl-

edge of this herpetofauna has been greatly improved by

field work undertaken under the CamHerp project which

ultimately resulted in the publication of a complete Atlas

of the reptiles of the country (Chirio and LeBreton 2007).

The BH, as defined above, hosts 50 non avian reptile

species among which 16 are endemic to our study area.

Fig. 6. Map of Cameroon with the geographical area of the

mountain range (circled in black) retained as part of this study.

Country boundaries are shown in red.

Others overflow very locally into neighboring Nigeria

(Obudu plateau and Mts. Gotel), the Central African Re-

public (eastern borders of the Adamaoua), or Equatorial

Guinea (Bioko Island). Only one snake species, Dipsa-

doboa unicolor , is also present in Eastern Africa. The

mountain biodiversity is thus relatively low but the level

of endemicity is quite high. All of the area undoubtedly

represents significant challenges to preserve the richness

and originality of the endemic afro-montane herpeto-

fauna. The regions studied here represent a much drier

area than the larger mountains such as Mt. Cameroon lo-

cated southwest on the Cameroon Volcanic Dorsal. The

herpetofauna includes common species, in addition to

taxa more restricted to these formations and their climate.

Other studies on the altitude mountains of the Cameroon

Volcanic Dorsal ridge have produced impressive species

lists, but unlike our study (only 50 species found above

1,400 m), they also include herpetofauna from the base of

the mountain ranges (Herrmann et al. 2005, 2006).

Inventory of Taxa Present in the Study Area

Below we discuss all species of non-avian reptiles re-

ported from the BH above 1,400 m (Fig. 6) from our col-

lection (total 374 specimens), including our observations

and/or data available in the literature. We include chelo-

nians (one specimen available), lizards (266 specimens),

and snakes (107 specimens). In each of these groups

Amphib. Reptile Conserv. 17 December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

we deal with the species alphabetically, by family, and

by taxa in families, in alphabetical order. We indicate

the list of available specimen(s) in the MNHN-RA col-

lections (Museum national d’Histoire naturelle, Reptiles

and Amphibians collections, Paris), that originate mainly

from collections made during the CamHerp project. Note,

however, that only a subset of CamHerp lizards have been

entered into MNHN collections, while most snakes have

not been accessioned in to the MNHN-RA yet and only

field tag details (CamHerp xxx) are given.

CHELONIANS

Testudinidae Gray, 1 825

Kinixys homeana Bell, 1827 (One specimen)

Material: CamHerp 121 (Mboh village, 6.327°N and

10.348°E, elev. 1,800 m, coll. CamHerp M. LeBreton,

July 8, 2002).

This tortoise prefers relatively humid spots, where it

seems to feed on mushrooms. It is found in all the western

part of Cameroon, from the plain to 1,800 m at Mboh in

the BH. The main threat is its frequent use as bushmeat by

local people as well as collection for sale in the country’s

major markets (Lawson 2001; Luiselli and Diagne 2014).

LIZARDS

Agamidae Spix, 1 825

Agama agama (Linnaeus, 1758) (Two specimens)

Material: CamHerp 44831 (Boyui village, 6.242°N and

10.3 11°E, elev. 1400 m, coll. CamHerp M. LeBreton

and L. Chirio, April 19, 2000) - CamHerp 168 (Mbiame,

6.190°N and 10.849°E, elev. 1955 m, coll. CamHerp, De-

cember 14, 2002).

This agama is undoubtedly the most anthropophilic

species in its group; it occupies almost all the villages in

its range but is also common in savanna outcrops and de-

graded forests. It is present in Bafoussam (elev. 1,500 m),

and found from sea level to over 2,000 m on Mts. Bana.

Agama sp. 2 (in: Chirio and LeBreton 2007) (four speci-

mens)

Material: CamHerp 3576X-3579X (4 specimens, Dz-

indong waterfall, 5.622°N and 10.106°E, elev. 2,350 m,

coll. CamHerp M. LeBreton and L. Chirio, May 5, 2001).

This rare and endemic species of Cameroon has not

been described yet. It occurs from the Bamboutos and

Mbapit Mountains in the BH to Tchabal Mbabo in Ad-

amaoua at altitudes located between 1,900 and 2,350 m at

Dzindong waterfall (Chirio and LeBreton 2007).

Agama sp. 4 (in: Chirio and LeBreton 2007) (Nine speci-

mens)

Amphib. Reptile Conserv.

Material: MNHN-RA 1998.0277-0285 (Nine specimens,

Mt. Oku, five km north of Oku village, on rocky outcrops,

elev. 2,200 m, coll. L. Chirio, June 25, 1998).

This endemic species of Cameroon, identified by

Chirio and LeBreton (2007: 172-173), is still not de-

scribed; it is only known from two mountain stations. It

is a large agama living mainly on the rocky outcrops of

altitude savannas. It occurs only between 1,900 and 2,000

m above sea level like at the localities of Lungoi and Ta-

benken.

Chamaeleonidae Gray, 1 825

It is only recently that the molecular work of Tilbury and

Tolley (2009) demonstrated that the two subgenera of

the genus Chamaeleo auct., Chamaeleo Laurenti, 1768

sensu stricto, and Trioceros Swainson, 1839 should be

considered as two valid genera. Other studies have sub-

sequently confirmed this (Tolley et al. 2013). Cameroon

has great species richness of chameleons (14 species)

compared to its neighboring countries. This diversity is

mainly located in mountainous areas and is characterized

by a high level of endemism. The family is represented

by three genera: Chamaeleo (five species), Rhampholeon

Gunther, 1874 (at least one species), and Trioceros (eight

species and three subspecies; Barej et al. 2010). Within

Trioceros , the most common to occur at elevation in-

clude Trioceros oweni, the most basal taxon of the genus

in Cameroon, T. camerunensis, T. cristatus, T. montium,

T. perreti, T. wiedersheimi, T. serratus, T. quadricornis

eisentrauti, T. q. quadricornis , and T. q. gracilior. The

genus Rhampholeon occurs over 1,700 m in Mt. Cam-

eroun but it is curiously absent in the BH. Six species

are very clear mountain endemics occupying restricted

areas in the Cameroon Volcanic Dorsal mountain ridge.

Half of Cameroon chamaeleons are mountain endemics

with restricted ranges. A molecular phylogeny of the ge-

nus Trioceros in Cameroon was established by Pook and

Wild (1997) and completed by Barej et al. (2010). Three

altitudinal groups in Cameroon can be recognized within

the genus Trioceros : a plains group ( Trioceros oweni), a

plains and submontane group ( Trioceros camerunensis, T.

cristatus, and T. montium), and a submontane and moun-

tain group ( Trioceros pfefferi, T. perreti, T. serratus, T.

wiedersheimi, and T. quadricornis). Only species of the

last group are present in our study area.

Chamaeleo gracilis Hallowell, 1844 (three specimens)

Material: MNHN-RA 2005.3191-3192 (two specimens,

Bamessing, 6.004°N and 10.352°E, elev. 1,200 m, coll.

CamHerp L. Chirio, October 26, 2000) - MNHN-RA

2005.3590 (Balengu, 5.114°N and 10.450°E, elev. 1,480

m, coll. CamHerp L. Chirio, April 6, 2000).

This species is found in the Ethiopian Rift Valley from

200-1,900 m (Largen and Spawls 2010), whereas in

Cameroon it is reported between 5 and 1,775 m above sea

December 2015 | Volume 9 | Number 2 | el 08

18

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

level (Chirio and LeBreton 2007). It was observed but not

collected at Bafoussam by one of us (LC, June 22, 2000).

Chamaeleo laevigatus Gray, 1863 (seven specimens)

Material: MNHN-RA 2005.2721 (Fundong, 6.249°N and

10.3 15°E, elev. 1,500 m, coll. CamHerp, July 8, 2002) -

MNHN-RA 2005.3301-3305 (five specimens, Jakiri vil-

lage along the road from Bamenda to Nkambe, 6.055 °N

and 10.658°E, elev. 1,550 m, coll. CamHerp M. LeBret-

on, July 8, 2002) - MNHN-RA 2005.3398 (Awing village

(Benjom), 5.867°N and 10.266°E, elev. 1,747 m, coll.

CamHerp, M. LeBreton, July 8, 2002).

This species, presently known to occur in Cameroon,

was initially mistaken with Chamaeleo senegalensis

Daudin, 1802 by Chirio and LeBreton (2007), a taxon

whose distribution is more western. In East Africa, C.

laevigatus occurs in moist savanna between 1,000-1,500

m but can fall to 300 m elsewhere (Spawls et al. 2002;

Largen and Spawls 2010). It is reported from 350-1,550

m altitude in Cameroon (Chirio and LeBreton 2007).

Trioceros pfefferi (Tornier, 1900) (three specimens)

Material: MNHN-RA 2005.3396 (Mboh village, 6.327°N

and 10.348°E, elev. 1,900 m, coll. CamHerp M. LeBre-

ton and L. Chirio, July 8, 2002) - MNHN-RA 2007.1499

(male; Mt. Oku, Afua, Ijim Forest, western side of Mt.

Oku, 6.15°N and 10.40°E, elev. 2,000 m, coll. Cam-

Herp L. Chirio, June 1st, 2000) - MNHN-RA 2007.1500

(female; Bali Ngemba Forest Reserve, 5.825°N and

10.087°E, elev. 1,400 m, coll. CamHerp L. Chirio, June

6 , 2000 ).

This endemic species of the Cameroon Volcanic Dor-

sal is a typical inhabitant of the wet stations of the western

sub-montane forest in the country. It is rare throughout

its distribution and was only known from its original de-

scription from Nyassosso at Mt. Kupe for nearly a century

(Wild 1993). It is found at Mt. Manengouba, and in the

BH and Mt. Oku where it reaches almost 2,000 m above

sea level. Densities seem higher in populations at Mt.

Kupe (Hofer et al. 2003). Altitudinal distribution of the

species ranges from 1,200-1,500 m (Schuetze 1998) and

1,100-1,900 m according to Tilbury (2010); the species is

reported between 1,100 and 1,900 m from Mt. Kupe by

Anderson and Van Heygen (2013). Captive females lay

between six and nine eggs (Schuetze 1998).

The species is also present at Mt. Nlonako, very close

to Mt. Manengouba. T. pfefferi has horns (males only),

but its phylogenetic affinities are closer to the hornless

species of the T. wiedersheimi group than to other Cam-

eroon species (T. montium and T. quadricornis), indicat-

ing that the presence of horns has evolved several times

within the genus Trioceros.

Its distribution is comparable to that of the T. perreti

/ T. serratus / T. wiedersheimi group and the T. quadri-

cornis group (T. q. quadricornis, T. q. gracilior, and T.

Amphib. Reptile Conserv.

q. eisentrauti). These two groups of related taxa each

have an endemic taxon in the Manengouba area, another

in the BH and a third endemic in a peripheral region (to

the north and west respectively). The populations of T.

pfefferi recently discovered at Mbulu Hills and Ediango

to the north (Gonwouo et al. 2006) should therefore be

carefully compared with the more southern populations

to assess their taxonomic status. Like other submontane

and montane species from Cameroon, T. pfefferi occupies

only medium and high mountain areas with wet, mainly

pristine evergreen forests, often near streams (Jakubow-

icz and Van Tiggel 1998). It perches at heights between

l. 6 m and 2.1 m (Herrmann et al. 2005), 7 m at Mt. Kupe,

and 3.5 to 5.0 m at Manengouba (Anderson and Van Hey-

gen 2013). The species is threatened on Mt. Manengouba

by both logging and collecting for the pet trade.

Trioceros quadricornis gracilior Bohme and Klaver,

1981 (17 specimens)

Material: MNHN-RA 1998.0434-0435 (two specimens,

Mt. Oku, above Oku village, elev. 2,200 m, coll. Cam-

Herp L. Chirio, June 25, 1998) - MNHN-RA 2005.2715-

2720 (six specimens, Mt. Oku, Elak Oku village, 6.202°N

and 10.505°E, elev. 2,000 m, coll. CamHerp M. LeBre-

ton and L. Chirio, July 8, 2002) - MNHN-RA 2005.2722

(Oku Manchok, 6.241°N and 10.524°E, elev. 2,130 m,

coll. CamHerp M. LeBreton and L. Chirio, December

14, 2002) - MNHN-RA 2005.2723 (Mt. Oku, Lake Oku,

6.20°N and 10.45°E, elev. 2,250 m, coll. CamHerp M.

LeBreton and L. Chirio, April 19, 2000) - MNHN-RA

2005.2724, 2005.2726-2727 (three specimens, Mt. Oku,

Oku village, 6.202°N and 10.505°E, elev. 2,000 m, coll.

CamHerp M. LeBreton and L. Chirio, April 19, 2000)

- MNHN-RA 2005.2725 (Mt. Oku, Simonkuh village,

6.234°N and 10.572°E, elev. 2,109 m, coll. CamHerp M.

LeBreton, July 8, 2002) - MNHN-RA 2007.1423 (male;

Mt. Oku, Oku village, 6.202°N and 10.505°E, elev. 2,000

m, coll. I. Ineich and N. Lhermitte-Vallarino, May 8,

2007) - MNHN-RA 2007.1424 (male; Mt. Oku, Oku vil-

lage, 6.202°N and 10.505 °E, elev. 2,000 m, coll. I. Ineich

and N. Lhermitte-Vallarino, May 8, 2007) - MNHN-RA

2007.1426 (male; Mt. Oku, Oku village, 6.202°N and

10.505°E, elev. 2,000 m, coll. I. Ineich and N. Lhermitte-

Vallarino, May 7, 2007).

Barej et al. (2010) revised the T. quadricornis com-

plex with additional materials and molecular data. The

morphological differences between the populations of the

south (Mt. Kupe and Mt. Manengouba) and north (BH to

Obudu Plateau in Nigeria) were supported by genetics,

thus confirming the subspecific status of T. q. quadricor-

nis (Tornier, 1899) and T. q. gracilior Bohme and Klaver,

1981. T. q. gracilior is present at Mts. Bamboutos, Mbulu

Hills (Gonwouo et al. 2006), Mt. Lefo, Mt. Oku and onto

the Obudu Plateau in Nigeria, while T. q. quadricornis oc-

cupies the forests of Mt. Manengouba and Mt. Kupe. This

study also relegated Chamaeleo eisentrauti, once consid-

December 2015 | Volume 9 | Number 2 | el 08

19

Ineich et al.

ered a valid species, to subspecific status as T. q. eisen-

trauti (Mertens, 1968). This form is endemic to Rumpi

Hills in western Cameroon. All these taxa occupy primary

mountain forests, and T. q. gracilior occurs up to 2,700

m in altitude. Tilbury (2010) reported the taxon between

l, 600-2,500 m. The separation between these three sub-

specific taxa, attested by their low genetic divergence, is

thus probably recent and associated with the altitudinal

shifting of cool forests to the mountain peaks after the end

of Pleistocene glacial periods.

Trioceros q. gracilior (Fig. 7) is an endemic subspe-

cies of Cameroon and neighboring Nigeria (Plateau

Obudu). This is an arboreal montane forest lizard (mostly

met at the interface forest/grassland) that is still relative-

ly abundant locally, such as around the village of Elak

Oku (6.244°N, 10.508°E, elev. 1,970 m). Its altitudinal

distribution reaches 2,400 m above sea level at Mt. Oku

(Ijim Ridge; Wild 1994) and 2,700 m at Mt. Mekua in the

Bamboutos (Gonwouo et al. 2006; Barej et al. 2010). Its

perch height is much greater than that of T. serratus (see

below) and averages around 1 .9 m at Mt. Oku (Gonwouo

et al. 2006). Wild (1994) found the chameleon from one

m above the ground to the top of the canopy at Mt. Oku,

with a preference for branches near streams. The mini-

mum night temperature recorded in its habitat at 2,400 m

is 4.7 °C in December 1993 (Wild 1994). The female lays

from 6 to 24 eggs that are partially incubated before being

laid (Abate 1994).

This species is particularly threatened by trade in exot-

ic pets, and especially by rampant habitat destruction (de-

forestation, cultures, bush fires, grazing). Eucalyptus, an

alien tree widely introduced in the region creates unfavor-

able habitat. However, the species seems able to persist in

fragmented forest remnants and transitional habitats (Fig.

8). Its densities are estimated at four times higher at Mt.

Oku compared to populations in Mbulu Hills (Gonwouo

et al. 2006), and almost twice as high as at Mt. Manen-

gouba (T. q. quadricornis ). The conservation status of the

species remains nevertheless very fragile and sensitive

to environmental degradation. The threat of commercial

harvesting is now better regulated by effective measures

implemented mostly via European Union CITES regula-

tion.

Trioceros serratus (Mertens, 1922) (101 specimens)

(Figs. 9-14)

Material: MNHN-RA 1997.3642 (male; Mt. Oku, Oku

village, coll. CamHerp L. Chirio, May 1997) - MNHN-

RA 1998.0415 (female; Mt. Oku, Lake Oku, elev. 2,200

m, coll. CamHerp L. Chirio, July 6, 1998) - MNHN-

RA 1998.0416-0430 (15 specimens, Mt. Oku, elev.

2,000-2,500 m, coll. CamHerp L. Chirio, June 25, 1998)

- MNHN-RA 2005.2728-2732 (five specimens, Mt.

Oku area, Anyajua village, above Bello, 6.236°N and

10.394°E, elev. 2,100 m, coll. CamHerp M. LeBreton and

L. Chirio, April 19, 2000) - MNHN-RA 2005.2733-2734,

Amphib. Reptile Conserv.

Fig. 7. Despite its specific name, individuals of T. q. gracili-

or may have two to six horns. Adult male, Elak Oku village,

Mt. Oku. Note the presence of concentric rings on the horns, a

characteristic feature (synapomorphy) of the genus Trioceros.

MNHN-RA 2007. 1424. Picture: I. Ineich, May 13, 2007.

Fig. 8. Associated crops (beans, coffee, bananas, corn) en-

countered near villages (here around Elak Oku village) are not

completely adverse to chameleons when a large plant and shrub

cover is maintained. Picture: I. Ineich, May 8, 2007.

2005.2736 (three specimens, Awing village (Benjom),

5.867°N and 10.266°E, elev. 1,747 m, coll. CamHerp M.

LeBreton and L. Chirio, December 14, 2002) - MNHN-

RA 2005.2735 (Awing village (Benjom), 5.867°N and

10.266°E, elev. 1,747 m, coll. CamHerp M. LeBreton and

L. Chirio, July 8, 2002 - MNHN-RA 2005.2737 (Baba II

village, 5.857°N and 10.102°E, elev. 1,772 m, coll. Cam-

Herp M. LeBreton and L. Chirio, December 14, 2002)

- MNHN-RA 2005.2738-2744 (seven specimens, Baba II

village, 5.857°N and 10.102°E, elev. 1,772 m, coll. Cam-

Herp M. LeBreton and L. Chirio, July 8, 2002 [MNHN-

RA 2005.2739, .2741 and .2743: December 14, 2002]) -

MNHN-RA 2005.2745 (Bamboutos, Mt. Mekua, 5.688°N

and 1 0.095 °E, elev. 2,700 m, coll. CamHerp L. Chirio,

March 30, 2000) - MNHN-RA 2005.2748 (Bingo village,

6.166°N and 10.290°E, elev. 1,435 m, coll. CamHerp M.

LeBreton and L. Chirio, December 14, 2002) - MNHN-

RA 2005.2749-2752 (four specimens, Mt. Oku, Elak Oku

village, 6.202°N and 10.505°E, elev. 2,000 m, coll. Cam-

Herp M. LeBreton and L. Chirio, July 8, 2002) - MNHN-

RA 2005.2755-2759 (five specimens, Mbiame, 6.190°N

and 10.849°E, elev. 1,955 m, coll. CamHerp M. LeBreton

and L. Chirio, July 8, 2002 [MNHN-RA 2005.2758-2759:

December 14, 2002]) - MNHN-RA 2005.2760-2761 (two

specimens, Mbockghas, elev. 2,092 m, coll. CamHerp M.

December 2015 | Volume 9 | Number 2 | el 08

20

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

LeBreton and L. Chirio, December 14, 2002) - MNHN-

RA 2005.2762-2771 (10 specimens, Mboh village,

6.327°N and 10.348°E, elev. 1,900 m, coll. CamHerp

M. LeBreton and L. Chirio, July 8, 2002) - MNHN-RA

2005.2774-2775, MNHN-RA 2005.2777, MNHN-RA

2005.3381 (four specimens, Mufe village, 6.30°N and

10.35°E, coll. CamHerp M. LeBreton and L. Chirio, July

8, 2002) - MNHN-RA 2005.2776, 2005.2778-2780 (four

specimens, Njinkfuin, 6.187°N and 10.375°E, elev. 1,500

m, coll. CamHerp M. LeBreton and L. Chirio, April 19,

2000) - MNHN-RA 2005.2781-2787, 2005.2900 (five

males and three females; Mt. Oku, Simonkuh, 6.234°N

and 10.572°E, elev. 2,109 m, coll. CamHerp M. LeBre-

ton and L. Chirio, July 8, 2002) - MNHN-RA 2005.2788

(male; Mt. Oku, Oku village, 1 0.505 °E and 6.202°N,

elev. 2,000 m, coll. CamHerp M. LeBreton and L. Chirio,

April 19, 2000) - MNHN-RA 2005.2812-2815 (four

specimens, Tefo village, 6.30°N and 10.37°E, coll. Cam-

Herp M. LeBreton and L. Chirio, July 8, 2002) - MNHN-

RA 2005.2816-2824 (nine specimens, Veko village,

6.139°N and 10.578°E, elev. 2,044 m, coll. CamHerp M.

LeBreton and L. Chirio, December 14, 2002 [MNHN-RA

2005.2817, .2819-2824: coll. July 8, 2002]) - MNHN-RA

2005.2900 (Mt. Oku, Simonkuh, 6.234°N and 10.572°E,

elev. 2,109 m, coll. CamHerp M. LeBreton and L. Chirio,

July 8, 2002) - MNHN-RA 2005.3382 (Babadjou,

5.699°N and 10.187°E, elev. 1,580 m, coll. CamHerp

L. Chirio, no date) - MNHN-RA 2005.3383 (Mbiame,

6.190°N and 10.849°E, elev. 1,955 m, coll. CamHerp

M. LeBreton and L. Chirio, July 8, 2002) - MNHN-RA

2007.0461-0464 (two males and two females; Mt. Oku

Fig. 9. Trioceros serratus male observed near a house in the vil-

lage of Elak Oku at Mt. Oku. MNHN-RA 2007. 1463. Picture: I.

Ineich, May 8, 2007.

Fig. 10. The neotype of Trioceros serratus, MNHN-RA 2007.

1494, photographed several days after his capture (see also other

photographs below). Picture: I. Ineich, May 13, 2007.

area, around Elak Oku village, 6.244°N and 10.507°E,

elev. 1,973 m, coll. I. Ineich and N. Lhermitte-Vallarino,

May 6, 2007) - MNHN-RA 1 2007.1461 (Mt. Oku, Oku

village, elev. 2,000 m, coll. I. Ineich and N. Lhermitte-Val-

larino, May 7, 2007) - MNHN-RA 2007.1462 (Mt. Oku,

Oku village, elev. 2,000 m, coll. I. Ineich and N. Lher-

mitte-Vallarino, May 8, 2007) - MNHN-RA 2007.1463-

1464, 2007.1472 (three specimens, Mt. Oku, Oku village,

elev. 2,000 m, coll. I. Ineich and N. Lhermitte-Vallarino,

May 8, 2007) - MNHN-RA 2007.1465 (male; Mt. Oku

area, Lake Oku, 6.202°N and 10.461°E, elev. 2,272 m,

coll. I. Ineich and N. Lhermitte-Vallarino, May 8, 2007) -

MNHN-RA 2007.1494 (male, neotype of T. serratus ; Mt.

Oku, on the side along the road from Anyajua to Belo, not

far from Belo, coll. I. Ineich, May 9, 2007).

Klaver and Bohme (1992) described the subspecies T.

wiedersheimi perreti from Mt. Manengouba. Later mo-

lecular studies of Barej et al. (2010) highlighted the pos-

sible specific status of this taxon. This same study showed

that the nominal subspecies T. w. wiedersheimi comprises

two distinct genetic clades, separated geographically.

Previously T. w. wiedersheimi was considered to occupy

savanna and altitude grasslands from 1,400 to 2,450 m in

Mts. Bamboutos, Mbulu Hills, Mt. Lefo, Mt. Mbam, Mt.

Oku, and Mt. Tchabal Mbabo, and westwards into Nigeria

at Mts. Gotel and Mambilla and the Obudu Plateaus. The

original description of T. wiedersheimi was based on two

syntypes, a female from Tchabal Mbabo and a subadult

male from the BH. The female was designated as the lec-

totype of T. w. wiedersheimi by Klaver and Bohme (1992),

thus restricting the type locality to Tchabal Mbabo. This

restricted its distribution to the northern part of that previ-

ously accepted (Tchabal Mbabo and Tchabal Gangdaba).

The southern populations (BH, Mt. Mbam and Mt. Oku)

represent a distinct taxon that may also include the popu-

lations of the Koano, Mt. Lefo and Mbulu Hills, and Pla-

teau of southern Nigeria, but this has to be verified. An

available name, Chamaeleo serratus Mertens, 1922, was

revalidated to accommodate these southern populations

as Trioceros serratus (Mertens, 1922), although its name-

bearing type was unfortunately destroyed during the

Second World War. A neotype was designated by Barej

et al. (2010) in recent MNHN collections (MNHN-RA

2007.1494, Pigs. 10, 11). Its type locality is thus well at-

tached to the area just above the city of Belo on the west-

ern flank of Mt. Oku.

Trioceros serratus occupies high savannas of the BH,

Mt. Mbam and Obudu Plateau (Nigeria). Note, however,

that the reports of Gotel Mountains in Nigeria should be

attributed to T. wiedersheimi. In the BH region, the spe-

cies is cited from Bafoussam (Bangwa), Big Babanki (=

Kedjom Keku), the Bamileke region of Dschang, Kis-

hong, Mezam (Bafout), and Tsch’a Bekom (Barej et al.

‘Note that specimens MNHN-RA 2007.461-464 reported

by Barej et al. (2010) refers to MNHN-RA 2007.1461-

1464.

Amphib. Reptile Conserv.

21

December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

Fig. 11. Neotype of T. serratus (MNHN-RA 2007.1494) in situ

before collection at the edge of the road down from the summit

of Mt. Oku (Anyajua village), just a little over Belo (6.175°N

and 10.352°E). The chameleon was perched nearly 3 m up in a

palm tree. Picture: I. Ineich, May 9, 2007.

Fig. 12. Trioceros serratus widely used the herb layer where it

was comfortable. Here an individual seeking to hide on a blade

of grass by stiffening its tail to make it look like an herbaceous

branching. Not collected. Picture: I. Ineich, May 8, 2007.

2010). Our study allows addition of the following loca-

tions in the BH: Awing (Benjom), Baba II, Bali Ngemba,

Bingo, Mbiame, Mbockghas, Mboh, Mufe, Njinkfuin,

Tefo, and Veko. It was reported from Bafut (elev. 1,200

m, 6.08°N and 10.10°E) by Joger (1982) as Chamaeleo

wiedersheimi.

Fig. 13. Individuals assigned to T. serratus altitude populations

(top, Elak Oku village; MNHN-RA 2007.1463) differ from

those from lower altitudes like here (bottom) the neotype of T.

serratus (MNHN-RA 2007.1494) by some important scalation

and coloring characters. Picture: I. Ineich, May 2007.

Gonwouo et al. (2006) consider the taxon (named T.

w. wiedersheimi) to occur from 1,500 m to 2,450 m al-

titude, often in sympatry with T. quadricornis gracilior

on Bamboutos Mts. at Foto, Dschang, and Mts. Lefo, Mt.

Oku, and Obudu Plateau in Nigeria, and 2,700 m in Mt.

Mekua. Wild (1994) reported the species between 2,200

m and 2,500 m at Mt. Oku (Ijim Ridge). Tilbury (2010)

cited the species from 2,600 m above sea level at Mt.

Oku. Perch height average is 90 cm at Mt. Oku, the low-

est value found for different stations of its range (over two

m at Tchabal Mbabo). Wild (1994) reported a mean perch

height of 53 cm at Mt. Oku and a maximum height of 157

cm. However, we collected the neotype of the species in

a palm tree at three m height near the edge of a main road

(Fig. 11)! The low perch height observed in altitude at Mt.

Oku could be attributed to the scarcity of livestock and

predators that cause little disruption for chameleons, or

to a still unknown interaction between climate and veg-

etation (Fig. 12). The species tolerates some degree of

habitat degradation and does not hesitate to venture into

cultivated areas retaining some original vegetation. Yet it

is a sensitive species, recently threatened by the exotic pet

trade and especially the destruction of its habitat (culture,

fires, deforestation). The population at Mt. Oku, however,

Amphib. Reptile Conserv.

22

December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

is still abundant. The species is common around the vil-

lage of Elak Oku, including gardens and plantations. This

is the most abundant Cameroon mountain chameleon.

The species occupies relatively open habitats but does not

hesitate to venture into closed canopy forest. A min imum

night temperature of 2.9 °C was recorded in its habitat at

2,500 m altitude in December 1993 (Wild 1994). Trioc-

eros serratus mostly occupies herbaceous and shrub layer

below two m, while T. q. gracilior occupies bushy and

shrub layers above one meter, which generates a syntopy

area in the stratum located between one and two m (Wild

1994). Habitat separation in syntopy should be possible

through the important size differences between both taxa,

probably preventing dietary overlap.

The systematics of this species complex is not sat-

isfactory, despite the revision of Barej et al. (2010). In

fact, besides the obvious differences in size and color-

ation, the lowland form (larger) collected near Belo also

differs from the altitude form (smaller) of the summit of

Mt. Oku by the number of small scales around the large

granules on the flanks (Figs. 13, 14). Also one of us (II)

collected the neotype of T. serratus up in a palm tree and

this form seems much more arboreal than the altitude Mt.

Oku form. It is very unlikely that these two morphotypes

belong to the same taxon and further studies are required.

Gekkonidae Gray, 1825

Hemidactylus angulatus Hallowell, 1852 (nine speci-

mens)

Material: MNHN-RA 2005.1602-1603 (two speci-

mens, Mt. Oku, Anyajua village above Bello, 6.236°N

and 10.394°E, elev. 2,100 m, coll. CamHerp, respec-

tively April 14, 2001, and April 19, 2001) - MNHN-RA

2005.1616 (Bingo village, between Ijim and Bamenda,

6.162°N and 10.319°E, elev. 1,600 m, coll. CamHerp,

April 19, 2000) - MNHN-RA 2005.1692-1693 (two spec-

imens, Bingo village, 6.166°N and 10.290°E, elev. 1,435

m, coll. CamHerp M. LeBreton, respectively December

14, 2002, and July 8, 2002) - MNHN-RA 2005.1761

(Idjim, Birdlife Project, 6.226°N and 10.433°E, elev.

1,600 m, coll. CamHerp L. Chirio, April 19, 2000) -

MNHN-RA 2005.1927-1928 (two specimens, Njinkfuin,

6.187°N and 10.375°E, elev. 1,500 m, coll. CamHerp L.

Chirio, April 19, 2000) - MNHN-RA 2005.2496 (Boyui

village, 6.242°N and 10.311°E, elev. 1,400 m, coll. Cam-

Herp L. Chirio, April 19, 2000).

This house gecko is probably one of the most anthro-

pophilous species in the country, where it has a wide dis-

tribution throughout the northern region. The species is

abundant in homes but does not hesitate to shelter also in

rocks and trees in remote areas. It is found from sea level

to above 2,000 m at Tabenken and Nkambe.

Hemidactylus kamdemtohami Bauer and Pauwels, 2002

(one specimen)

Amphib. Reptile Conserv.

Fig. 14. Individuals assigned to altitude T. serratus populations

(on top, Elak Oku village; MNHN-RA 2007.1463) are very dif-

ferent from those from lower altitudes like here (on botom) the

neotype of T. serratus (MNHN- RA 2007.1494) by the confor-

mation of the large granules arranged on the flanks and also by

the number and arrangement of small scales placed around these

large granules. Pictures: I. Ineich, May 2007.

Material: MNHN-RA 2002.0739 (Balengou, elev. 1,480

m, 5.114°N and 10.450°E, coll. CamHerp, June 29, 2001).

Balengou remains the only known Cameroon location

for this gecko, which elsewhere is known from Equato-

rial Guinea (Mt. Allen) and Gabon (Mt. Iboundji). H.

kamdemtohami is without any doubt a submontane spe-

cies. Its occurrence at lower elevations in Gabon may be

because Mt. Iboundji, covered with evergreen forests, is

wetter than the BH and thus the altitudinal limit of the

species is reduced.

Scincidae Gray, 1825

Lacertaspis chriswildi (Bohme and Schmitz, 1996) (sev-

en specimens)

Material: MNHN-RA 1997.3649 (Mt. Oku, in a garden

of Oku village, elev. 2,000 m, coll. CamHerp L. Chirio,

June 8, 1997) - MNHN-RA 1997.3650 (Mt. Oku, in the

forest, elev. 2,350 m, coll. CamHerp L. Chirio, March 22,

1997) - MNHN-RA 1998.0286-0288 (three specimens,

Mt. Oku forest, elev. 2,200 m, coll. CamHerp L. Chirio,

June 25, 1998) -MNHN-RA 2005.2600-2601 (two speci-

mens, Mt. Oku, Oku forest, 6.250°N and 10.507°E, elev.

2,350 m, coll. CamHerp M. LeBreton and L. Chirio, re-

spectively May 5, 2000, and May 11, 2000).

This little lizard is endemic to the montane forests of

West Cameroon (Schmitz 2004; Schmitz et al. 2005; Her-

rmann et al. 2006). It is found at Mt. Kupe in the Taka-

manda forest, Mt. Oku, and the Tchabal Mbabo Massif.

It occurs up to 2,800 m altitude at Mt. Oku but does not

seem to fall below 1,000 m.

December 2015 | Volume 9 | Number 2 | el 08

23

Ineich et al.

Lacertaspis lepesmei (Angel, 1940) (35 specimens)

Material: MNHN-RA 1998.0295-0300, 1999.0401-0404,

1999.8418-8436 (29 specimens, without any precise loca-

tion, coll. CamHerp) - MNHN-RA 2004.0061 (Bambou-

tos, Fulbe house, 5.637°N and 10.106°E, elev. 2,450 m,

coll. CamHerp, May 5, 2001) - MNHN-RA 2005.2597-

2599 (Bamboutos, Mt. Mekua, 5.688°N and 10.095°E,

elev. 2,700 m, coll. CamHerp, respectively May 8,

2000, April 18, 2000, and April 19, 2000) - MNHN-RA

2005.2602-2603 (two specimens, without precise loca-

tion, coll. CamHerp).

This small, submontane endemic skink is only known

from the rocky slopes of Bamboutos Mountains, between

2,350 and 2,700 m altitude (Fig. 15). It is not present in

the Mt. Oku region. Its classification in the IUCN Red

List and the measures to undertake for the conservation

of its habitat should be a priority.

Lepidothyris fernandi (Burton, 1836) [formerly Mochlus

fernandi] (one specimen)

Material: MNHN-RA 2005.1265 (Tefo village, 6.30°N

and 10.37°E, elev. 1,700 m, coll. CamHerp M. LeB reton

and L. Chirio, July 8, 2002).

The genus was recently revised (Wagner et al. 2009).

In Eastern Africa, the species occurs between 600 and

2,100 m (Spawls et al. 2002) whereas in Cameroon it is

only reported from sea level to 1,200 m at Bafut. This

skink was also observed on the eastern sides of the BH

at Kenshi, at an elevation of 1,080 m on April 17, 2004

(6.107°N and 9.713°E).

Leptosiaphos ianthinoxantha (Bohme, 1975) (25 speci-

mens)

Material: MNHN-RA 2002.0798, 2002.0800, 2002.0928-

0930, 2002.0934 (six specimens, Mbockghas, 6.222°N

and 10.582°E, elev. 2,092 m, coll. CamHerp M. LeB-

reton, December 14, 2002) - MNHN-RA 2002.0942,

2005.2617-2620 (five specimens, Mbockghas, 6.222°N

and 10.582°E, elev. 2,092 m, coll. CamHerp M. Le-

Breton and L. Chirio, December 14, 2002) - MNHN-

RA 2005.2607 (Bamboutos, Fulbe house, 5.637°N and

10.106°E, elev. 2,450 m, coll. CamHerp, May 5, 2001)

- MNHN-RA 2005.2613-2616 (four specimens, Bam-

boutos, Mt. Mekua, 5.688°N and 10.095°E, elev. 2,700

m, coll. CamHerp, March 30, 2000, May 5, 2000 [.2615],

and May 8, 2000 [.2616]) - MNHN-RA 2005.2621-

2627, 2005.2629 (eight specimens, Mt. Oku, Simonkuh,

6.234°N and 10.572°E, elev. 2,109 m, coll. CamHerp

M. LeB reton, July 8, 2002, December 14, 2002 [.2625],

and January 16, 2003 [.2622, .2626]) - MNHN-RA

2005.2628 (Bamboutos, slopes of Mt. Mekua, 5.698°N

and 10.101°E, elev. 2,300 m, coll. CamHerp, March 19,

2002 ).

Amphib. Reptile Conserv.

Fig. 16. Leptosiaphos ianthinoxantha. Cameroon, Mt. Oku,

Oku Simonkou village. Picture: M. LeBreton, November 2002.

This small skink is endemic to montane grasslands of

the Western Highlands of Cameroon (Schmitz et al. 2005)

(Fig. 16). It is found at Mt. Lefo (Forest Reserve of Ba-

fut-Ngemba) and in the Bamboutos Mountains. Its occur-

rence at Mt. Oku had been suspected by Wild in 1994. It

is a semi-burrowing species living in open montane grass-

lands, and is oviparous. The species occurs up to 2,700 m

altitude at Mt. Mekua in the Bamboutos where its popula-

tions are highly localized but occur in high densities.

Leptosiaphos pauliani (Angel, 1940) (one specimen)

Material: MNHN-RA 1939.0082 (holotype; Bamboutos,

coll. J.-L. Perret).

This small endemic lizard was recorded by Perret

(1973) from Nyassosso on the slopes of Mt. Kupe at

1,100 m above sea level (holotype of Riopa erythropleu-

ron Mertens, 1968) and from Mts. Bamboutos at 2,300 m

above sea level (holotype of Lygosoma ( Liolepisma ) pau-

liani Angel, 1940). It was not found during the CamHerp

project work; its presence in the BH is questionable. This

strictly submontane species may be limited to the area of

submontane forests located between 1,100 and 2,000 m

above sea level in the Mts. Kupe and Bamboutos.

Leptosiaphos vigintiserierum (Sjostedt, 1897) (two speci-

mens)

December 2015 | Volume 9 | Number 2 | el 08

24

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

Material: MNHN-RA 1998.0294 (Mt. Oku, elev. 2,000

m, coll. CamHerp L. Chirio, September 1st, 1997) -

MNHN-RA 2004.0062 (Bamboutos, waterfall and sacred

forest, 5.622°N and 10.106°E, elev. 2,350 m, coll. Cam-

Herp, May 5, 2001) - Bamboutos, slopes of Mt. Mekua,

5.698°N and 10.101°E, elev. 2,300 m, coll. CamHerp,

March 19, 2002).

This species is endemic to the Cameroon Volcanic

Dorsal (Schmitz et al. 2005) (Fig. 17). It is found from

Bioko Island (Equatorial Guinea), Mt. Cameroon, and

Mt. Oku (Mt. Nkolodou, Mt. Kala, Mt. Kupe, and Mt.

Nlonako). It mainly occurs in the high meadows of the

peaks above the evergreen forest areas. It reaches 2,450 m

at Mts. Bamboutos and can be relatively abundant locally.

Trachylepis maculilabris (Gray, 1845) (33 specimens)

Material: MNHN-RA 1997.3643 (pass on the Bafoussam

road, elev. 1,850 m, coll. CamHerp L. Chirio, April 1997)

- MNHN-RA 1998.0289-0293 (five specimens, Mt. Oku,

five km north of Oku village, elev. 2,000 m, coll. CamHerp

L. Chirio, June 25, 1998) - MNHN-RA 2005.1610-1611

(two specimens, Baba II village, 5.857°N and 10.102°E,

elev. 1,772 m, coll. CamHerp M. LeBreton and L. Chirio,

respectively July 8, 2002, and December 14, 2002) -

MNHN-RA 2005.1616 (Bingo village, between Ijim and

Bamenda, 6.162°N and 10.3 19°E, elev. 1,600 m, coll.

CamHerp M. LeBreton and L. Chirio, April 19, 2000) -

MNHN-RA 2005.1617 (Bali Ngemba village, 5.833°N

and 10.077°E, elev. 1,398 m, coll. CamHerp M. LeBre-

ton and L. Chirio, July 8, 2002) - MNHN-RA 2005.1623

(Bamboutos, waterfall and sacred forest, 5.622°N and

10.106°E, elev. 2,350 m, coll. CamHerp, May 5, 2001) -

MNHN-RA2005. 1692-1693 (Bingo village, 6.166°N and

10.290°E, elev. 1,435 m, coll. CamHerp M. LeBreton and

L. Chirio, December 14, 2002) - MNHN-RA 2005.1761

(Idjim village, Birdlife Project, 6.226°N and 10.433°E,

elev. 1,600 m, coll. CamHerp M. LeBreton and L. Chirio,

April 19, 2000) - MNHN-RA 2005.1762 (Jakiri village,

road from Bamenda to Nkambe, 6.055°N and 10.658°E,

elev. 1,550 m, coll. CamHerp M. LeBreton and L. Chirio,

December 14, 2002) - MNHN-RA 2005.1847-1848 (two

specimens, Mbiame village, 6.190°N and 10.849°E, elev.

1,955 m, coll. CamHerp M. LeBreton and L. Chirio, De-

cember 14, 2002) - MNHN-RA 2005.1852 (Mbockghas,

6.222°N and 10.582°E, elev. 2,092 m, coll. CamHerp M.

LeBreton and L. Chirio, December 14, 2002) - MNHN-

RA 2005.1853-1858 (six specimens, Mboh village,

6.327°N and 10.348°E, elev. 1,900 m, coll. CamHerp

M. LeBreton and L. Chirio, December 14, 2002 [.1853],

and July 8, 2002 [.1854-1858]) - MNHN-RA 2005.1897

(Mufe village, 6.30°N and 10.35°E, coll. CamHerp M.

LeBreton and L. Chirio, July 8, 2002) - MNHN-RA

2005.1935-1938 (four specimens, Mt. Oku, Simonkuh

village, 6.234°N and 10.572°E, elev. 2,109 m, coll. Cam-

Herp M. LeBreton and L. Chirio, July 8, 2002, and De-

Amphib. Reptile Conserv.

Fig. 17. Leptosiaphos vigintiserierum. Cameroon, Mt. Mekua,

Bamboutos. Specimen CamHerp 36431. Picture: M. LeBreton,

March 12, 2002.

cember 14, 2002 [.1938]) - MNHN-RA 2005.1944 (Veko

village, 6.139°N and 10.578°E, elev. 2,044 m, coll. Cam-

Herp M. LeBreton and L. Chirio, December 14, 2002)

- MNHN-RA 2005.1958-1959 (two specimens, Sarkong

Hill, west of Jakiri, 6.054°N and 10.598°E, elev. 1,600 m,

coll. CamHerp, March 19, 2002) - MNHN-RA 2005.2484

(Tefo village, 6.30°N and 10.37°E, elev. 1,700 m, coll.

CamHerp M. LeBreton and L. Chirio, July 8, 2002).

This skink has a wide distribution in Africa and in

Cameroon it is found in a variety of habitats from lowland

forests to altitude grasslands. The species is also anthro-

pophilic and can be abundant in gardens and villages in

the southern half of the country. This lizard occurs from

sea level to above 2,550 m at Mt. Lefo or on the top of

Mt. Nlonako around 1,825 m (Herrmann et al. 2005). In

East Africa T. maculilabris is reported from the seaside to

above 2,300 m (Spawls et al. 2002; Largen and Spawls

2010). Note, however, that its taxonomy is not clearly es-

tablished (Mausfeld et al. 2004) and that it currently rep-

resents a species complex containing several cryptic taxa.

Trachylepis mekuana (Chirio and Ineich, 2000) (six spec-

imens)

Material: MNHN-RA 2001.0109 (Bamboutos, Mt. Me-

kua, 5.688°N and 10.095°E, elev. 2,700 m, coll. Cam-

Herp, April 19, 2000) - MNHN-RA 2002.0922 (Bali

Ngemba village, on rocks above the valley, 5.830°N

and 10.066°E, elev. 1,640 m, coll. CamHerp M. LeBre-

ton, July 8, 2002) - MNHN-RA 2005.1289-1291 (three

specimens, Bamboutos, slopes of Mt. Mekua, 5.698°N

and 10.086°E, elev. 2,600 m, coll. CamHerp, March 19,

2002) - MNHN-RA 2005.2606 (Bamboutos, 5.637°N

and 10.106°E, elev. 2,450 m, coll. CamHerp L. Chirio,

March 30, 2000).

This endemic mountain lizard of the BH in Cameroon

occupies only the top of Bamboutos Mountains (Mt. Me-

kua) and the Massif of Bali-Ngemba at elevations located

between 2,400 and 2,700 m (Fig. 18). The increasing use

of its habitat for grazing and planting food crops seriously

threatens the survival of this species. Its classification on

December 2015 | Volume 9 | Number 2 | el 08

25

Ineich et al.

Fig. 18. Trachylepis mekuana. Mt. Mekua, Bamboutos. March

18, 2002. Picture: L. Chirio.

Fig. 19. Atractaspis i. irregularis - Cameroon, Yaounde. Pic-

ture: M. LeBreton, January 4, 2011.

the IUCN Red List and habitat conservation measures

should be a priority.

SNAKES

Atractaspididae Gunther, 1858

Atractaspis irregularis irregularis (Reinhardt, 1843) (six

specimens)

Material: CamHerp 0627C, 0423C (two specimens,

Abu village, NE of Fundong, 6.297°N and 10.33 1°E,

elev. 1,750 m, coll. CamHerp M. LeBreton, December

14, 2002) - CamHerp 35011 (Awing village (Benjom),

5.867°N and 10.266°E, elev. 1,747 m, coll. CamHerp M.

LeBreton, December 14, 2002) - CamHerp 1269C, 14951

(two specimens, Baba II village, 5.857°N and 10.102°E,

elev. 1,772 m, coll. CamHerp M. LeBreton, December

14, 2002) - CamHerp 0158C (Mbiame, 6.190°N and

10.849°E, elev. 1,955 m, coll. CamHerp, December 14,

2002 ).

This burrowing and venomous snake (Barriere et al.

2006) exhibits an extensive African distribution (Fig. 19).

It occupies dense evergreen forests and degraded semi-

deciduous forests, forest-savanna mosaics (moist savan-

na), the Western Highlands, and the extreme south of the

Adamaoua. It is found in altitude from 500 m to 2,000 m

at Tabenken. This snake was mentioned in Wum (elev.

l, 023 m) by Bohme (1975). In East Africa, the species is

reported from 600 m to 2,000 m above sea level (Spawls

et al. 2002; Largen and Spawls 2010).

Polemon collaris (W. Peters, 1881) (four specimens)

Material: CamHerp 34681, 37071 (two specimens, Bin-

go village, 6.166°N and 10.290°E, elev. 1,435 m, coll.

CamHerp M. LeBreton, December 14, 2002) - CamHerp

37381 (Mbiame, 6.190°N and 10.849°E, elev. 1,955 m,

coll. CamHerp M. LeBreton, July 8, 2002) - CamHerp

36641 (Baba II village, 5.857°N and 10.102°E, elev. 1,772

m, coll. CamHerp M. LeBreton, December 14, 2002).

This small forest burrowing snake is found at altitudes

between 5 and 1,955 m in Cameroon. Joger (1982) men-

tions the species from Wum (elev. 1,023 m).

Colubridae Oppel, 1811

Crotaphopeltis hotamboeia (Laurenti, 1768) (four speci-

mens)

Material: CamHerp 0141, 24881 (two specimens, Jakiri

village on the road of Nkambe to Bamenda, 6.055 °N and

10.658°E, elev. 1,550 m, coll. CamHerp M. LeBreton,

July 8, 2002, and December 14, 2002) - CamHerp 24831

(Veko village, 6.139°N and 10.578°E, elev. 2,044 m, coll.

CamHerp M. LeBreton, July 8, 2002) - CamHerp 0159C

(Baba II village, 5.857°N and 10.102°E, elev. 1,772 m,

coll. CamHerp M. LeBreton, December 14, 2002).

This widely distributed snake occurs at elevations from

400-2,500 m in East Africa (Largen and Spawls 2010). In

Cameroon, it is found at altitudes between 160 and 2,044

m. Mountain populations in Cameroon show a particular

coloration, with a typical dark spotted belly; they could

belong to a distinct taxon (see below). The relationship

of individuals from Veko and Baba II villages to the sub-

montane species listed below should be reviewed.

Crotaphopeltis sp. (three specimens)

Material: CamHerp 4469, 4470 (two specimens, Mt.

Oku, Bello village, 6.170°N and 10.344°E, elev. 1,450 m,

coll. CamHerp, April 19, 2000) - CamHerp 03491 (City

of Bamenda, 5.958°N and 10.165°E, elev. 1,300 m, coll.

CamHerp, March 20, 2001).

This “species” has not been described yet but its va-

lidity, which remains to be confirmed, was indicated by

Chirio and LeBreton (2007: 400-401). It is considered

endemic to the mountains of Cameroon and occurs be-

tween 1,050 m and 1,500 m.

Amphib. Reptile Conserv.

26

December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

Dasypeltis confusa Trape and Mane, 2006 (three speci-

mens)

Material: CamHerp 24361 (Veko village, 6.139°N and

10.578°E, elev. 2,044 m, coll. CamHerp M. LeBreton,

July 8, 2002) - CamHerp 0097 (Awing village (Benjom),

5.867°N and 10.266°E, elev. 1,747 m, coll. CamHerp M.

LeBreton, July 8, 2002) - CamHerp 1367C (Bali Ngemba

village, 5.833°N and 10.077°E, elev. 1,398 m, coll. Cam-

Herp M. LeBreton, July 8, 2002).

This snake is a typical inhabitant of the humid savanna

of Cameroon where it occurs at altitudes between 510 m

and 2,044 m.

Dasypeltis fasciata A. Smith, 1849 (three specimens)

Material: CamHerp 0218C (Jakiri village on the road

from Bamenda to Nkambe, 6.055°N and 10.658°E, elev.

l, 550 m, coll. CamHerp M. LeBreton, December 14,

2002) - CamHerp 22721 (Baba II village, 5.857°N and

10.102°E, elev. 1,772 m, coll. CamHerp M. LeBreton,

December 14, 2002) - CamHerp 24361 (Veko village,

6.139°N and 10.578°E, elev. 2,044 m, coll. CamHerp M.

LeBreton and L. Chirio, July 8, 2002).

This semi-arboreal snake is found at altitudes between

4 and 1,380 m. It is reported from Bafut (elev. 1,200 m,

6.08°N and 10.10°E) by Joger (1982).

Dipsadoboa unicolor Gunther, 1858 (two specimens)

Material: MNHN-RA 1998.0438-0439 (two specimens,

Mt. Oku, Oku village, elev. 2,000 m, coll. CamHerp L.

Chirio, end 1997).

This nocturnal and semi-arboreal snake has a wide Af-

rican distribution from Guinea (Conakry) to Burundi. In

Cameroon, it occupies not only the altitude forest of the

west of the country but also evergreen degraded forests.

It occurs from around sea level up to 2,000 m at Mt. Oku

and up to 2,044 m in Veko, a village in the southeast of

Mt. Oku. At Mt. Nlonako, the species does not reach the

higher elevations of the massif (Herrmann et al. 2005).

This snake is still present over 1 ,600 m at Mt. Nimba

(Ineich 2003), but can occur elsewhere up to 3,000 m and

also can withstand low temperatures while remaining ac-

tive at night and hunting amphibians on which it feeds. In

East Africa, it is only reported between 1,500 m and 3,000

m elevation. The conspecificity of West African popula-

tions (Mt. Nimba, Cameroon Volcanic Dorsal) with those

of the East African mountains has not been confirmed.

Dipsadoboa weileri (Lindholm, 1905) (seven specimens)

Material: CamHerp 0835, 0101M, 0043C (three speci-

mens, Mboh village, 6.327°N and 10.348°E, elev. 1,900

m, coll. CamHerp M. LeBreton, July 8, 2002 (two speci-

mens), and December 14, 2002 (one specimen)) - Cam-

Herp 0606C (Fundong, 6.249°N and 10.3 15°E, elev. 1,500

Amphib. Reptile Conserv.

m, coll. CamHerp M. LeBreton, July 8, 2002) - CamHerp

0248C (Veko village, 6.139°N and 10.578°E, elev. 2,044

m, coll. CamHerp M. LeBreton, July 8, 2002) - Cam-

Herp 1437C (Mbiame village, 6.190°N and 10.849°E,

elev. 1,955 m, coll. CamHerp M. LeBreton, December 14,

2002) - CamHerp 119 (Awing village (Benjom), 5.867°N

and 10.266°E, elev. 1,747 m, coll. CamHerp M. LeBre-

ton, December 14, 2002).

This nocturnal forest semi-arboreal snake occurs in

Cameroon at altitudes from 10 m to above 2,000 m. The

species is more likely a central African species which was

erroneously reported from West Africa (Trape and Balde

2014).

Dispholidus typus (A. Smith, 1828) (one specimen)

Material: CamHerp 31971 (Baba II village, elev. 1,772

m, 5.857°N and 10.102°E, coll. CamHerp M. LeBreton,

December 14, 2002).

This diurnal semi-arboreal snake has a wide pan- Afri-

can distribution in the savannas. The subspecies Dispho-

lidus typus occidental is Perret, 1961 described from

Cameroon remains doubtful but requires a thorough revi-

sion before its validity can be evaluated (Broadley and

Wallach 2002). Perret (1961: 138) recognized D. t. oc-

cidental™ based on its color with green males, strongly

streaked with black, red and brown females, as well as

the presence of two elliptical black spots, slightly oblique,

situated laterally on each side of the neck in both sexes.

The species occupies forest-savanna mosaic, the western

Highlands and the high savannas. Its altitude record on its

whole range is 2,400 m (Spawls et al. 2002; Wagner and

Bohme 2007; Largen and Spawls 2010).

Grayia tholloni Mocquard, 1 897 (one specimen)

Material: CamHerp 2050C (Jakiri village, on the road

from Bamenda to Nkambe, 6.055°N and 10.658°E, elev.

1,550 m, coll. CamHerp M. LeBreton, July 8, 2002).

This water snake is found up to 1,400 m above sea

level in East Africa (Largen and Spawls 2010) and be-

tween 510 and 1,550 m in Cameroon.

Philothamnus angolensis Bocage, 1882 (two specimens)

Material: MNHN-RA 1998.0410 (Mt. Oku, above the

village, elev. 2,200 m - tail broken - formerly identified

as Philothamnus bequaerti, coll. CamHerp L. Chirio,

June 25, 1998) - CamHerp 37491 (Mbiame, 6.190°N and

10.849°E, elev. 1,955 m, coll. CamHerp M. LeBreton and

L. Chirio, July 8, 2002).

This arboreal snake of wet savanna occupies degraded

forests, forest- savanna mosaics, the western Highlands,

and altitude savannas like the Sudan savanna in the plains.

Herrmann et al. (2006) reported the species up to 2,450 m

at Mt. Meletan in the Bamboutos, as well as at Tchabal

Mbabo Range. A snake reported from the area as Philo-

December 2015 | Volume 9 | Number 2 | el 08

27

Ineich et al.

thamnus irregularis by Joger (1982) refers to this species

(Hughes 1985: 518; Bohme and Schneider 1987). In East

Africa, it occupies various habitats from the sea border

up to 2,000 m elevation (Spawls et al. 2002). This species

from Central and Eastern Africa only extends very little

west beyond the Cameroon border.

The Mt. Oku specimen deposited in the collections

(MNHN-RA 1998.0410) is a female formerly identi-

fied as Philothamnus bequaerti but here conservatively

considered to correspond to P. angolensis. It measures

565 mm SVL and stubby tail measurement is 201+ mm.

There are 15 dorsal scale rows in the middle of the body,

1+164 unkeeled ventral plates, and 79+ subcaudals, also

unkeeled. Anal plate is divided. The supralabials (right/

left) are 9 (4-6 touching the eye)/9 (4-6), infralabials

9/9, temporals 1 + 1/1 + 1, preoculars 1/1 and postoculars

2/2. The inside of the mouth is white. Its assignment to

P. angolensis is not entirely compatible with the species’

description, however.

Philothamnus hughesi Trape and Roux-Esteve, 1990 (one

specimen)

Material: CamHerp 880 (Veko village, 6.139°N and

10.578°E, elev. 2,044 m, coll. CamHerp M. LeBreton,

December 14, 2002).

This tree snake of wet savannas occurs at an altitudinal

range between 740 and 2,100 m.

Thrasops flavigularis (Hallo well, 1852) (one specimen)

Material: MNHN-RA 1998.0436 (skin, head and neck

only; Mt. Oku, Oku village, elev. 2,050 m, coll. CamHerp

L. Chirio, November 8, 1997).

This snake is a typical inhabitant of the dense forests of

Central Africa, from Cameroon to the Democratic Repub-

lic of Congo. It is common to find in the villages and plan-

tations. Thrasops flavigularis occupies the Highlands up

to 2,000 m at Mt. Oku. Gonwouo et al. (2007) recognize

it as an inhabitant of submontane forests in Cameroon.

This snake, once considered non-venomous, is capable of

inflicting serious envenomations (Ineich et al. 2006) and

should be handled with caution.

Our specimen, MNHN-RA 1998.0436, only consists

of the head, neck [in good condition], and the skin of an

individual eaten by the local population. It has 15 dorsal

scale rows in the middle of the body, which seems rare ac-

cording to Chippaux (2006), because there are more often

only 13 - however 15 dorsal scales seems more typical of

grass field populations (Stucki-Stirn 1979). Preoculars are

2/2 and the upper is the largest (>2 times the size of the

lower). The upper preoculars are widely separated from

the frontal. The first post-ocular prevents contact of the

supralabial 6 with the eye. Postoculars 3/3 and the lower

is much larger and elongated (>4 times) than the other

two substantially equal in size. The lower postocular con-

tacts two supralabials (5-6). There are only 7(4-5 )/7(4-5)

Amphib. Reptile Conserv.

supralabials and 10/11 infralabials. Temporals 1 +1/1+1.

This specimen slightly differs from the diagnosis given

by Chippaux (2006: 108-109) and Stucki-Stirn (1979:

320-328) for the species.

According to Chippaux (2006), our specimen differs

from Thrasops jacksoni because it has 2 preoculars (ver-

sus 3), its much larger lower postocular (vs. sup. and inf.

larger) and 7 supralabials (vs. 10-12) and from Thrasops

occidental is because the large postocular is in contact

with 2 supralabials (vs. postocular in contact with 3 su-

pralabials). We refer that damaged specimen to Thrasops

flavigularis and consider some of the characters indicated

in the diagnosis of the species given by Chippaux (2006)

as incomplete.

Elapidae Boie, 1827

Dendroaspis jamesoni jamesoni (Traill, 1843) (eight

specimens)

Material: MNHN-RA 2000.4360, 2000.4376, 2002.0385-

0389 (seven specimens, Bamenda, gift Latoxan, coll. Oc-

tober 30, 2000) - CamHerp 34281 (Jakiri village on the

road from Nkambe to Bamenda, 6.055°N and 10.658°E,

elev. 1,550 m, coll. CamHerp M. LeBreton, December 14,

2002 ).

This venomous tree snake has a wide distribution range

extending from Togo in West Africa to Angola in southern

Africa. It occupies dense evergreen and semi-deciduous

forests, forest-savanna mosaics, the Western Highlands,

and high savannas of Adamaoua (681 m at Tchabal Mba-

bo; Herrmann et al. 2006). It often frequents plantations

and gardens but is unaggressive. It occurs in altitude up to

2,000 m at Mts. Bana. Gonwouo et al. (2007) considered

the species as an inhabitant of mountain forests located

above 1,800 m. It seems to live up to 2,200 m elsewhere

on its range. In East Africa this green mamba is reported

from 600 m to 2,200 m above sea level (Spawls et al.

2002 ).

Naja melanoleuca Hallowell, 1857 (22 specimens)

Material: CamHerp 14881, 31841 (two specimens, Abu

village, northeast of Fundong, 6.297°N and 10.33 1°E,

elev. 1,750 m, coll. CamHerp M. LeBreton, Decem-

ber 14, 2002) - CamHerp 1222C, 3175C, 3736C (three

specimens, Baba II village, 5.857°N and 10.102°E, elev.

1,772 m, coll. CamHerp M. LeBreton, July 8, 2002) -

CamHerp 31401, 33941 (two specimens, Bali Ngemba

village, 5.833°N and 10.077°E, elev. 1,398 m, coll. Cam-

Herp M. LeBreton, December 14, 2002) - CamHerp

0880C, 32951 (two specimens, Bingo village, 6.166°N

and 10.290°E, elev. 1,435 m, coll. CamHerp M. LeBre-

ton, December 14, 2002) - CamHerp 4496 (Fundong,

6.249°N and 10.3 15°E, elev. 1,500 m, coll. CamHerp

L. Chirio, April 19, 2000) - CamHerp 31341 (Jakiri vil-

lage along the road from Bamenda to Nkambe, 6.055 °N

December 2015 | Volume 9 | Number 2 | el 08

28

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

and 10.658°E, elev. 1,550 m, coll. CamHerp M. LeBre-

ton, December 14, 2002) - CamHerp 12341, 0557C (two

specimens, Mbiame village, 6.190°N and 10.849°E, elev.

l, 955 m, coll. CamHerp M. LeBreton, December 14,

2002) - CamHerp 0856, 0014C, 0133C (three specimens,

Mbockghas village, 6.222°N and 10.582°E, elev. 2,092

m, coll. CamHerp M. LeBreton, December 14, 2002) -

CamHerp 0392C, 1086C, 23561, 33921 (four specimens,

Mboh village, 6.327°N and 10.348°E, elev. 1,900 m, coll.

CamHerp, July 8, 2002) - CamHerp 32911 (Sarkong Hill,

west of Jakiri village, 6.054°N and 10.598°E, elev. 1,600

m, coll. CamHerp, March 19, 2002) - CamHerp 1452C

(Veko village, 6.139°N and 10.578°E, elev. 2,044 m, coll.

CamHerp M. LeBreton, July 8, 2002).

This species (Fig. 20) has a wide distribution and the

systematics of the species complex remains problemat-

ic. The name N. melanoleuca has only to be applied to

central African populations. It occupies dense evergreen

and semi-deciduous forests, forest-savanna mosaics, and

the Western Highlands. It is found from sea level up to

2,700 m at Mt. Meletan in the Bamboutos. Gonwouo et

al. (2007) consider that this snake can occur in mountain

forests between 1,800 m and 3,000 m above sea level in

Cameroon. The cobra is quoted from Bafut (elev. 1,200

m, 6.08°N, 10.10°E) by Joger (1982). The species, as cur-

rently recognized (sensu lato), is reported up to 2,500 m

altitude in Kenya (Spawls et al. 2002; Wagner and Bohme

2007; Largen and Spawls 2010).

Naja nigricollis Reinhardt, 1843 (one specimen)

Material: CamHerp 1500C (Jakiri village along the road

from Bamenda to Nkambe, 6.055°N and 10.658°E, elev.

1,550 m, coll. CamHerp M. LeBreton, July 8, 2002).

This spitting cobra species seems not to exceed 1 ,000 m

elevation in East Africa where another related species,

Naja ashei Wtister and Broadley, 2007, can occur above

1,750 m (Largen and Spawls 2010). Naja nigricollis is

found between 20 and 1,800 m elevation in Cameroon.

Lamprophiidae Fitzinger, 1 843

The validity of this family was recently demonstrated

by Kelly et al. (2011). This work showed that the genus

Lamprophis was polyphyletic. A new genus was created

and other species previously included in the genus Lam-

prophis were divided into three groups: (1) virgatus and

fuliginosus, together with lineatus and olivaceus were

transferred to the revalidated genus Boaedon A.M.C. Du-

meril, Bibron, and A.H.A. Dumeril, 1854; (2) Lycodono-

morphus was nestled within Lamprophis sensu lato and

a sister taxon of Lamprophis inornatus-thQ latter species

was therefore transferred to the genus Lycodonomorphus;

(3) Lamprophis sensu stricto was restricted to a small

clade of four species endemic to South Africa, with Lam-

prophis aurora as type species. We follow this revised

taxonomy here.

Amphib. Reptile Conserv.

Fig. 20. Naja melanoleuca. Cameroon, Bamessing, October 31,

2003. Picture: M. LeBreton.

Boaedon fuliginosus (Boie, 1827) [formerly Lamprophis

fuliginosus ] (two specimens)

Material: CamHerp 0992C (Baba II village, 5.857°N and

10.102°E, elev. 1,772 m, coll. CamHerp M. LeBreton,

July 8, 2002) - CamHerp 1365C (Veko village, 6.139°N

and 10.578°E, elev. 2,044 m, coll. CamHerp M. LeBre-

ton, December 14, 2002).

Boaedon fuliginosus is a snake often encountered in

and around houses. Nocturnal and terrestrial, it has a

very wide African distribution, although populations in

southern and eastern Africa were referred to B. capensis

(Hughes 1997). It occupies a variety of habitats ranging

from dense evergreen and semi-deciduous degraded for-

ests, to forest-savanna mosaics through the Adamaoua

high savannas and Sudanian savannas. It occurs up to

2,044 m at Veko village in the BH and up to 2,400 m

in East Africa (Spawls et al. 2002; Largen and Spawls

2010 ).

Boaedon virgatus (Hallowell, 1854) (one specimen)

Material: CamHerp 37471 (Baba II village, 5.857°N and

10.102°E, elev. 1,772 m, coll. CamHerp M. LeBreton,

December 14, 2002).

This terrestrial forest species is present between 10 m

and 1,770 m elevation in Cameroon.

Bothrolycus ater Gunther, 1 874 (five specimens)

Material: CamHerp 0487C, 34031, 0174, 0306 (four

specimens, Mboh village, 6.327°N and 10.348°E, elev.

1,900 m, coll. CamHerp M. LeBreton, July 8, 2002 (two

specimens) and December 14, 2002 (two specimens) -

CamHerp 32381 (Baba II village, 5.857°N and 10.102°E,

elev. 1,772 m, coll. CamHerp M. LeBreton, December 14,

2002 ).

This terrestrial forest snake is present at elevations be-

tween 10 m and 1,500 m in Cameroon.

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December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

Gonionotophis stenophthalmus (Mocquard, 1887) (one

specimen)

Material: CamHerp 0897 (Jakiri village along the road

from Bamenda to Nkambe, 6.055°N and 10.658°E, elev.

1,550 m, coll. CamHerp M. LeBreton, July 8, 2002).

This semi-arboreal, ophiophagous forest snake is pres-

ent between 50 m and 1,500 m elevation in Cameroon.

Lycophidion multimaculatum Boettger, 1888 (two speci-

mens)

Material: MNHN-RA 2002.0943 (Awing village (Ben-

jom), 5°3’28”N and 10°1’4”E, elev. 1,747 m, coll. Cam-

Herp M. LeBreton, December 14, 2002) - CamHerp -

(Bamboutos, Fulbe house, elev. 2,450 m, coll. CamHerp

P. Makolowode, June 12, 1999).

The specimen MNHN-RA 2002.0943 is identified as

Lycophidion multimaculatum. It measures 250 mm SVL

and its tail is 28 mm. It has 17 dorsal rows at midbody. Its

non-keeled ventrals are 2+186 and unkeeled subcaudals

30. Anal plate is entire. Supralabials (right/left) 8 (3-5 in

contact with the eye)/8 (3-5), infralabials 8/8 (1-4 in con-

tact with the first pair of gular), temporals 1 +2+3/1 +2+3,

preocular 1/1, postoculars 2/2. An apical pit distinguished

on dorsal scales and anterior gulars are of the same size

as the posterior. That specimen is uniform grey bluish

dorsally and ventrally, only slightly lighter ventrally; no

marks, rings, or spots can be seen. Its diagnosis is not en-

tirely consistent with that of the species to which we refer

to tentatively. The species is found between 510 m and

2,450 m elevation (Mt. Meletan, Bamboutos) in Camer-

oon. So it is a partially submontane species in Cameroon

(i.e., but not strictly submontane, much like Dipsadoboa

unicolor).

Psammophiidae Boie, 1827

Psammophis cf. phillipsii (Hallowell, 1844) (three speci-

mens)

Material: CamHerp 180, 60 1C, 844C (three specimens,

Mbiame, 6.190°N and 10.849°E, elev. 1,955 m, coll.

CamHerp M. LeBreton and L. Chirio, July 8, 2002).

This terrestrial snake is common in Cameroon and

Central African Republic. It occupies a variety of habitats

ranging from degraded forests to high savannas. It does

not hesitate to frequent the villages and even large cit-

ies like Yaounde. The species is abundant in the whole

southern half of the country, except in undisturbed forest

areas, and is found up to 2,000 m at Tabenken. Species

status was granted to this taxon by Kelly et al. (2008) as

Psammophis occidentalis Wemer, 1919, but that name

does not apply to those populations of the P. phillipsii

complex (entire anal plate). They are however distinct

from P. phillipsii sensu stricto and their status is under

revision (Trape, pers. comm, to LC). Those populations

Amphib. Reptile Conserv.

were previously recognized as P. phillipsii by Chirio and

Ineich (2006) and Chirio and LeBreton (2007). They be-

long to a central African species whose distribution does

not occur west of the Cameroon border. This snake (as

Psammophis sibilans) was also reported from Bafut (elev.

1,200 m, 6.08°N, 10.10°E) by Bohme (1975).

Psammophis sp. 1 (in: Chirio and LeBreton 2007: 540-

541) (one specimen)

Material: CamHerp 0645 C (Oku Simokuh village,

6.234°N and 10.572°E, elev. 2,109 m, coll. CamHerp,

July 8, 2002).

This undescribed terrestrial species is an inhabitant

of the Cameroon mountains, and seems to share external

morphological affinities with an Ethiopian specimen from

MNHN-RA collections. It occupies the Western High-

lands, but also the Adamaoua high savannas. Currently

its distribution is limited to a few peaks of the Cameroon

Volcanic Dorsal, where it ascends to 2,109 m altitude at

Mt. Oku.

Typhlopidae Jan, 1 863

Afrotyphlops cf. punctatus (Leach, 1819) (11 specimens;

see below)

Material: CamHerp 0087C, 32371 (two specimens, Tefo

village, 6.30°N and 10.37°E, coll. CamHerp M. LeBre-

ton, and L. Chirio, July 8, 2002) - CamHerp 14121 (Mufe

village, 6.30°N and 10.35°E, coll. CamHerp M. LeBreton

and L. Chirio, July 8, 2002) - CamHerp 1018C 1208C

(two specimens, Mboh village, 6.327°N and 10.348°E,

elev. 1,900 m, coll. CamHerp L. Chirio, July 8, 2002);

CamHerp 1253C, 31351, (two specimens, Mboh village,

6.327°N and 10.348°E, elev. 1,900 m, coll. CamHerp L.

Chirio, December 14, 2002); CamHerp 0176C, 102 1C

(two specimens, Abuh village, NE of Fundong, 6.297°N

and 10.33 1°E, elev. 1,750 m, coll. CamHerp M. LeB-

reton, December 14, 2002) - CamHerp 0396C, 0180M

(two specimens, Baba II village, 5.857°N and 10.102°E,

elev. 1,772 m, coll. CamHerp M. LeBreton, December 14,

2002 ).

Specimens are only provisionally attributed to this

species pending further study and occur in marbled and

unmarbled forms. This burrowing snake is found at alti-

tudes between 5 m and 1,800 m in Cameroon from Mboh

village (1,800 m), Baba II village (1,770 m) and Idjim

village (1,600 m)). Afrotyphlops cf. punctatus is found

between 10 m and 1,800 m above sea level in Cameroon,

and has been reported from Wum (elev. 1,023 m, 6.39°N

and 10.07°E) by Bohme (1975).

Viperidae Oppel, 1811

Athens broadleyi Lawson, 1999 (one specimen)

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December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

Material: CamHerp 0974C (Bali Ngemba village,

5.833°N and 10.077°E, elev. 1,398 m, coll. CamHerp M.

LeBreton, July 8, 2002).

This small arboreal forest viper (Fig. 21) is found at al-

titudes between 332 m and 1,398 m in Cameroon (Chirio

and LeBreton 2007). The species is also present in the

Central African Republic. The geographic distribution of

this small tree viper is still unclear (Phelps 2010), but it

occurs with certainty in Cameroon and the Central Afri-

can Republic (Chirio and Ineich 2006).

Athens squamigera (Hallowell, 1854) (two specimens)

Material: CamHerp 0336, 1205C (two specimens, For-

est Reserve of Bali Ngemba, 5.825°N and 10.087°E, elev.

1,400 m, coll. CamHerp, March 19, 2002).

This semi-arboreal viper is an inhabitant of the dense

forests that occur from sea level up to 1 ,900 m (Broadley

1998). This is exceeded by Athens nitschei, an East Afri-

can species that occurs up to 2,700 m (Phelps 2010).

Bitis arietans arietans (Merrem, 1820) (two specimens)

Material: CamHerp 0694C (Veko village, 6.139°N and

10.578°E, elev. 2,044 m, coll. CamHerp M. LeBreton,

July 8, 2002) - CamHerp 35231 (Jakiri village along the

road from Bamenda to Nkambe, 6.055°N and 10.658°E,

elev. 1,550 m, coll. CamHerp M. LeBreton, December 14,

2002 ).

This big and massive snake has a pan- African distribu-

tion, and is also found on the Arabian Peninsula. It fre-

quents forest-savanna mosaics, the Western Highlands,

and all types of savannas (high, Sudanese, and Sahelian).

It lives at ground level and bites are frequent, making it a

feared snake. It occupies elevation areas up to 2,044 m in

the village of Veko in the BH. Its wide distribution in Af-

rica was largely influenced by the occupation of climatic

refuges during periods of glaciation (Barlow et al. 2013).

Other altitude populations exist such as those of the East

African Mountain Arc or of the Drakensberg mountains

in South Africa (Phelps 2010; Barlow et al. 2013). The

altitudinal record for the species is around 2,200 m but

the species seems able to occur even higher, up to 2,400

m (Spawls et al. 2002; Largen and Spawls 2010).

Bitis gabonica (A.M.C. Dumeril, Bibron and A.H.A. Du-

meril, 1 854) (one observed specimen)

Material: One specimen was observed but not collected

near Bangangte at 1,480 m elevation.

This big forest viper was reported from Bafut (elev.

1,200 m, 6.08°N and 10.10°E) by Stucky-Stirn (1979)

and found at 1,500 m in the western extension of the BH,

and also at Mende in the Takamanda. It was observed

by one of us (LC) at almost 1,500 m near Bangangte. In

Cameroon it is found at altitudes between 5 m and only

Fig. 21. Athens broadleyi. Megangme, 4.598°N and 12.225°E,

elev. 610 m, September 8, 2012. Picture: M. LeBreton.

1,500 m, but occurs over 2,300 m in East Africa (Kucha-

rzewski 2011).

Bitis nasicornis (Shaw, 1802) (no available specimen)

This bulky viper, characterized by its horn-shaped scales

at the snout tip, shows a vast African distribution. It oc-

cupies dense evergreen and semi-deciduous forests, the

Western Highlands, and the forest-savanna mosaics in

well-preserved forest pockets. It prefers moist valley bot-

toms in the dense forests, and is considered a dangerous

venomous snake. It occurs up to 2,000 m altitude at Lake

Awing in the BH in Cameroon (specimen observed but

not collected), and up to 2,400 m in East Africa (Spawls

et al. 2002; Kucharzewski 2011). It was reported from

Mbengwi, northwest of Bamenda (elev. 1,200 m) by

Stucky-Stirn (1979).

Causus maculatus (Hallowell, 1 842) (three specimens)

Material: CamHerp 1350C (Baba II village, 5.857°N and

10.102°E, elev. 1,772 m, coll. CamHerp M. LeBreton,

July 8, 2002) - CamHerp 0147C (Bali Ngemba village,

5.833°N and 10.077°E, elev. 1,398 m, coll. CamHerp

M. LeBreton, July 8, 2002) - CamHerp 08181 (Mbiame,

6.190°N and 10.849°E, elev. 1,955 m, coll. CamHerp M.

LeBreton and L. Chirio, July 8, 2002).

This small nocturnal viper is very common in wet sa-

vanna and degraded forests areas. It does not hesitate to

venture into the villages at night but its venom is only

slightly harmful. Its distribution is broad and includes

much of the African continent, from Mauritania to Ugan-

da and Angola. It can be present up to 1,950 m altitude

at Mbiame in the BH in Cameroon, which seems to be

its altitude record all over its range (Kucharzewski 2011).

Its presence in East Africa seems questionable and should

probably refer to an undescribed high-elevation species

close to the endemic species reported below. In Ethiopia

it is only known from a few specimens collected between

Amphib. Reptile Conserv.

31

December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

500 and 1,000 m above sea level (Largen and Spawls

2010 ).

Causus sp. (in: Chirio and LeBreton 2007: 612-613) (four

specimens)

Material: CamHerp 0964C (Mboh village, 6.327°N and

10.348°E, elev. 1,900 m, coll. CamHerp L. Chirio, July

9, 2002) - CamHerp 0196, 0695C, 0998C (three speci-

mens, Mbiame, 6.190°N and 10.849°E, elev. 1,955 m,

coll. CamHerp M. LeBreton and L. Chirio, July 8, 2002,

and December 14, 2002 [0998C]).

This scarce montane species occupies both Camer-

oon and the Central African Republic (far west). It is not

described yet but has numerous morphological affinities

with the forms of the Causus rhombeatus (Lichtenstein,

1823) group from East and South Africa. In Cameroon,

it occupies the Adamaoua high savannas and the Western

Highlands where it looks for wet lowlands and the banks

of mountain creeks. It is only found at altitude, from 700

m at Ngaouyanga (Adamaoua) up to 1,950 m at Mbiame

(BH).

Biogeographic Affinities of the Reptiles of Mt.

Oku and the Bamenda Highlands

The 50 reptile species in the study area are classified al-

phabetically below within each biogeographic region rec-

ognized.

Ubiquitous species (1): Agama agama.

Forest species from Western and Central Africa (9): Bitis

nasicornis - Boaedon virgatus - Dasypeltis fasciata -

Dendroaspis j. jamesoni - Dipsadoboa unicolor - Dipsa-

doboa weileri - Goniocephalus stenophthalmus - Kinixys

homeana - Trachylepis maculilabris .

Central African forest species (8): Athens broadleyi -

Athens squamigera - Bitis gabonica - Bothrolycus ater

- Lepidothyris fernandi - Naja melanoleuca - Polemon

collaris - Thrasops flavigularis.

African savanna species (12): Afrotyphlops cf. punctatus

- Atractaspis i. irregularis - Bitis a. arietans - Boaedon

fuliginosus - Causus maculatus - Chamaeleo graci-

lis - Crotaphopeltis hotamboeia - Dasypeltis confusa -

Dispholidus typus - Grayia tholloni - Hemidactylus an-

gulatus - Naja nigricollis.

Savanna species with eastern affinities (5): Chamaeleo

laevigatus - Lycophidion multimaculatum - Philotham-

nus angolensis - Philothamnus hughesi - Psammophis cf.

phillipsii.

Endemic Cameroon mountain species (13): Agama sp. 2

- Agama sp. 4 - Causus sp. - Crotaphopeltis sp. - Lacer-

taspis chriswildi - Lacertaspis lepesmei - Leptosiaphos

ianthinoxantha - Leptosiaphos pauliani - Leptosiaphos

vigintiserierum - Trachylepis mekuana - Trioceros pfef-

feri - Trioceros quadricornis gracilior - Trioceros ser-

ratus.

Montane species (2): Hemidactylus kamdemtohami -

Psammophis sp. 1 .

The species composition of our study area located on the

Cameroon Volcanic Dorsal is characterized by the pres-

ence of a similar number of species in the three dominant

elements: savanna forms, forest forms, and endemic mon-

tane forms.

Among the 50 reptile species in our study zone there are:

(1) two very anthropophilous species that rise high in el-

evation in the villages of the region: Agama agama and

Table 1. List of the 50 reptile species present in our study area at Mt. Oku and the Bamenda Highlands. For each species we indicate

if it is a low elevation or montane species (submontane) (in bold characters) and its altitudinal limits known in Cameroon. For each

family we indicate between brackets the number of species in our study area.

Altitudinal limits in Cameroon

(elevation indicated in meters)

Families

Species

Low elevation

species

Submontane

species

Testudinidae (1)

Kinixys homeana

0-1800

Agamidae (3)

Agama agama

Agama sp. 2

Agama sp. 4

0-2000

1900-2350

1900-2200

Chamaeleonidae (5)

Chamaeleo gracilis

Chamaeleo laevigatus

Trioceros pfefferi

Trioceros quadricornis gracilior

Trioceros serratus

0-1500

350-1550

1100-2000

1800-2700

1040-2700

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

Table 1 (continued). List of the 50 reptile species present in our study area at Mt. Oku and the Bamenda Highlands. For each spe-

cies we indicate if it is a low elevation or montane species (submontane) (in bold characters) and its altitudinal limits known in

Cameroon. For each family we indicate between brackets the number of species in our study area.

Altitudinal limits in Cameroon

(elevation indicated in meters)

Families

Species

Low elevation

species

Submontane

species

Gekkonidae (2)

Hemidactylus angulatus

Hemidactylus kamdemtohami

0-2000

1450-1500

Scincidae (8)

Lacertaspis chriswildi

1000-2800

Lacertaspis lepesmei

2350-2700

Lepidothyris fernandi

Leptosiaphos ianthinoxantha

0-1700

1300-2700

Leptosiaphos pauliani

1300-2000

Leptosiaphos vigintiserierum

1000-2450

Trachylepis maculilabris

Trachylepis mekuana

0-2100

2400-2700

Atractaspididae (2)

Atractaspis i. irregularis

500-2000

Polemon collaris

0-1950

Colubridae (11)

Crotaphopeltis hotamboeia

Crotaphopeltis sp.

160-2044

1050-1500

Dasypeltis confusa

500-1550

Dasypeltis fas data

Dipsadoboa unicolor

0-2050

80-2050

Dipsadoboa weileri

0-2050

Dispholidus typus

350-2150

Grayia tholloni

510-1550

Philothamnus angolensis

50-2450

Philothamnus hughesi

700-2100

Thrasops flavigularis

0-2050

Elapidae (3)

Dendroaspis j. jamesoni

0-2000

Naja melanoleuca

0-2700

Naja nigricollis

0-1800

Lamprophiidae (5)

Boaedon fuliginosus

250-2050

Boaedon virgatus

0-1800

Bothrolycus ater

0-1800

Gon ionotoph is stenophthalmus

50-1500

Lycophidion multimaculatum

500-2450

Psammophiidae (2)

Psammophis cf. phillipsii

Psammophis sp. 1

0-2000

1450-2100

Typhlopidae (1)

Afrotyphlops cf. pundatus

0-1800

Viperidae (7)

Athens broadleyi

300-1400

Athens squamigera

0-1500

Bids a. arietans

250-2000

Bids gabonica

0-1500

Bids nasicornis

0-2000

Causus maculatus

0-1950

Causus sp.

700-1950

Amphib. Reptile Conserv.

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December 2015 I Volume 9 I Number 2 I el 08

Ineich et al.

Trachylepis maculilabris . They both occur as well in

West Africa, Central and Eastern Africa. However, note

that T. maculilabris is an anthropophilic species that re-

quires more moisture than A. agama, which only enters in

the forest degraded by man;

(2) a mixed group of forest and savanna species that are

ecologically tolerant; they are also found in the plains but

they often reach 2,000 m in the BH and on the slopes of

Mt. Oku. Most of them are also found in West Africa, ex-

cept Bothrolycus ater , Chamaeleo laevigatus, Dendroas-

pis j. jamesoni, Lycophidion multimaculatum, Naja mela-

noleuca (sensu stricto), Philothamnus hughesi, Polemon

collaris, and Thrasops flavigularis , which are limited to

the large Central African forest block (and its surround-

ing areas);

(3) a group of mountain species, endemic or not to the

study area: Agama sp. 2, Agama sp. 4, Causus sp., Crota-

phopeltis sp., Dipsadoboa unicolor , Hemidactylus kam-

demtohami, Lacertaspis lepesmei, Lacertaspis chriswildi,

Leptosiaphos ianthinoxantha , Leptosiaphos pauliani,

Leptosiaphos vigintiserierum, Psammophis sp. 1, Trachy-

lepis mekuana, Trioceros pfefferi, Trioceros quadricornis

gracilior, and Trioceros serratus.

Altitudinal Distribution

Among the mountain endemic species of the Cameroon

Volcanic Dorsal, T. quadricornis gracilior , T. serratus,

and L. chriswildi reach the highest elevations on Mt. Oku,

although none occur beyond the treeline where subalpine

meadows appear around 2,600 m above sea level (Fig.

22, Table 1 ). So far, no reptile species has been identified

on the summit of Mt. Oku grasslands. However, as with

Mt. Cameroon, specific searches for them have not been

made, and amphibians are relatively well abundant as po-

tential prey for batrachophagous snakes. It is however a

harsh climate for reptiles, with cold nights and frequent

frosts.

A clear nomenclature describing the altitudinal distri-

bution patterns observed in Cameroon is difficult as dif-

ferences between zoological groups are important. Amiet

(1971) adopted the biogeographic classification of altitu-

dinal distributions in Cameroon proposed by Letouzey

(1968):

1,000 m/1,200 m = low and medium altitude rain

forest strata;

1,000 m-1,200 m/1,600-1,800 m = submontane

strata;

1,600 m-1,800 m/2,200-2,500 m = montane strata;

2,200 m-2,500 m/3,200-3,600 m = afro- subalpine

strata;

above 3,200-3,600 m = afro-alpine strata.

Fig. 22. The altitude grassland of the summit at Mt. Oku no

longer harbors any reptile. Chameleons can still be found in the

forest on the edge of the meadows, up almost 2,600 m above sea

level. However, one can observe there a tiny endemic viviparous

toad under the stones on the ground. Picture: I. Ineich, May 7,

2007.

Later he (Amiet 1975) defined a “oro-cameroon faunis-

tic element” of species with distributions above 1,000

m elevation. The term “orobiontes” (here replaced with

submontane species) was used for these high altitude spe-

cies. Montane species also present in lower areas around

mountains were also distinguished as “monticolous spe-

cies.” Amiet (1987) estimated that the average annual

temperature had lowered from 3.5 to 4.5 °C during the

last glaciation in Cameroon, and showed that the altitudi-

nal limit of 900 m to 1 ,000 m is an important ecological

boundary, marking the exclusion of many lowland spe-

cies and the appearance of true submontane species. This

boundary in the BH, however was not at 1,000 m eleva-

tion but increased to 1,400 m, before a distinctive “sub-

montane” herp assemblage occurs. The further mountain

ranges are located from the sea the altitudinal limit for a

species appears to increase. The separation of vicariant

Cameroon “submontane” reptile assemblage is relatively

recent and seems to mainly date from 25,000 to 15,000

BP (Amiet 1987).

Herrmann et al. (2005) presented a detailed study of

herpetofauna of Mt. Nlonako, and identified only four

of 89 species whose range exceeding 1,700 m altitude:

Trioceros pfefferi (Chamaeleonidae), Leptosiaphos vigin-

Amphib. Reptile Conserv.

34

December 2015 | Volume 9 | Number 2 | el 08

Reptiles of Mont Oku and the Bamenda Highlands, Cameroon

tiserierum, Trachylepis maculilabris (Scincidae), and

Chamaelycus fasciatus (Lamprophiidae). They noted that

within the Cameroon Volcanic Dorsal a mountain range

must exceed a certain altitude to allow the development

of an endemic herpetofauna, otherwise faunal exchanges

between ranges resulted in the presence of a shared sub-

montane Cameroon biota.

Supraspecific Diversity

Although many reptile fa mil ies in Cameroon, as in East

African mountains, have endemic montane species, e.g.,

Agamidae, Chamaeleonidae, Scincidae, Psammophiidae,

and Viperidae, there is a curious absence of montane La-

certidae in Cameroon. In Kenya, Adolfus alleni (Barbour,

1914) and Adolfus masavaensis (Wagner et al. 2014) oc-

cur in the summit grasslands of the Aberdares and Mt.

Kenya, respectively, with ranges from 2,700 to 4,500 m

(Spawls et al. 2002). In contrast, there are a number of

skinks, particularly small, semifossorial members of the

genera Lacertaspis and Leptosiaphos, that occur above

2,000 m, with Trachylepis mekuana and Lacertaspis lep-

esmei being high-altitude endemics.

Mountain dwelling taxa do not necessarily come from

the same genera: inside Viperidae, the genera Athens and

Bids often possess endemic montane forms sometimes

encountered over 3,000 m in East Africa, with the mono-

typic Montatheris hindii being also endemic to montane

heathlands. Only the genus Causus shows an endemic

submontane species in Cameroon which does not even

reach 2,000 m elevation. Note, however, that the genus

Athens holds endemic species in Cameroon or at least in

the Cameroon region ( Athens broadleyi, A. subocularis),

but curiously none of them are limited to the highlands,

contrary to what can be observed in East Africa. The

strongest affinities between East Africa and Cameroon

seem to mainly concern two particularly diverse lizard

families on the African continent, Chamaeleonidae and

Scincidae (Ineich and Chirio 2004).

Endemism

Endemism at the Cameroon Volcanic Dorsal has a gen-

eral pattern but with several exceptions. Speciation by

vicariance clearly dominates with close but distinct taxa

(except for Trioceros pfefferf see our comments above)

between separate massifs (e.g., Manengouba and BH-Mt.

Oku). The highest peak of the Cameroon Volcanic Dorsal,

Mt. Cameroon, an active volcano, is newer than the other

summits located further north in the Dorsal. It has no en-

demic mountain reptiles, however, and this is certainly re-

lated to its geological age. However, no detailed study has

been undertaken to estimate genetic divergences among

disjunct populations of Trioceros montium which reaches

1,100-1,200 m at Mt Kupe, but 1,500 m at Manengouba

(Anderson and Van Hey gen 2013). This species is cur-

rently assigned to a single taxon, without subspecific dis-

Amphib. Reptile Conserv.

tinction, but may well follow a similar evolutionary path-

way like other mountain chameleons of Cameroon.

Threats and Conservation

The threats to this submontane herpetofauna are numer-

ous (Euskirchen et al. 2000). The conservation status of

all the endemic species is fragile, and their limited ranges

are being rapidly degraded. However, they are character-

ized by locally high densities, which unfortunately also

makes them all the more easy to collect. In fact, species

of mountain chameleon from Cameroon are highly sought

after for the international exotic pet trade. However, the

most serious threat to their existence is the rapid human

population growth in the region of Mt. Oku and the West-

ern Highlands. It makes species preservation difficult be-

cause human pressure on land for agriculture and live-

stock, and consequent deforestation, is destructive and

growing with little regard to the conservation of endan-

gered species that are increasing in number.

Conclusions

Like most other highland areas, the highest reliefs of Mt.

Oku and the BH have only a limited herpetofauna. How-

ever the species assemblage is original in its composition.

First, it contains a ubiquitous fauna, able to occupy a wide

range of habitats from sea level to almost 2,000 m eleva-

tion. It also includes typical mountain species unable to

survive below 1,000 m, and climbing up to 2,800 m. The

vast majority of these latter species, highly specialized at

least climatically, are endemic to the Cameroon Volcanic

Dorsal and often to a single mountain range. The only

study on the herpetofauna of Mt. Oku mentioned only

two lizards and seven amphibians, including a scoleco-

morphid caecilian (Wild 1994). Our work considerably

increases this list but unfortunately five potential new

species first signaled by Chirio and LeBreton (2007) have

still to be described.

The unique herpetofauna of this region is seriously

threatened by exponential human growth and its associ-

ated impacts. The fertile volcanic soil in the region has al-

ways attracted humans, whose expanding population and

utilization of natural resources, inevitably encroaches on

the fragile habitats of reptiles. Survival and preservation

of these populations for future generations must be met

with prompt protective actions that are both robust and

effective. In addition it must gain the support of the local

human population if these endemic species are not to face

extinction in the near future.

Acknowledgments. — II and NLV wish to thank Sam-

uel Wanji (Research Foundation for Tropical Diseases

and the Environment, Buea, Cameroon) for his logistic

support during field work. Field research of II and NLV

in Cameroon was undertaken under the French ANR pro-

gramme (ANR Biodiversite - IFORA). Also thanks to

35

December 2015 | Volume 9 | Number 2 | el 08

Ineich et al.

the Ministry of Forestry and Wildlife and the Ministry of

Scientific Research and Innovation who provided autho-

rizations and facilitated this work in Cameroon. Funding

and support from the Bamenda Highlands Forest Project

and support from the Cameroon Biodiversity Conserva-

tion Society made also this work possible. Authors also

wish to thank Dan Portik and particularly Bill Branch for

useful comments on this paper.

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Ivan Ineich was the Reptile Curator for Squamates at Museum National d’Histoire Naturelle, Paris, France from

1988 to 2014. He has studied Pacific islands reptiles for more than 30 years, describing several new taxa and

contributing to a better knowledge of many areas in Oceania. He rediscovered Bocourt’s Terrific Skink ( Phobos -

cincus bocourti ) in 2003, a giant island skink that was considered extinct for nearly 150 years. He contributed

also to systematic and biodiversity studies in Africa and Asia, and participated in the description of numerous

new taxa of lizards and snakes. His research interests include taxonomy, biogeography, faunal turnovers and

conservation of lizards and snakes all over the world, particularly on islands and island-like continental areas.

Matthew LeBreton has worked at the intersection of the fields of health, environment, conservation, and wild-

life ecology for the past 25 years. His work has involved engagement and inclusion of government and com-

munity in research, program development, and implementation. He has coauthored around 80 scientific papers

related to health and environment and a book on the reptiles of Cameroon in collaboration with the National

Museum of Natural History in Paris. In central Africa, Matthew has worked on programs funded by various

governments universities and foundations. He is the founder and director of Mosaic which is based in Cameroon,

and provides technical advice, program assistance, and project implementation support to governments, NGOs,

and companies throughout the region.

Nathaly Lhermitte-Vallarino earned her Masters and Ph.D. at the Museum National d’Histoire Naturelle, Paris,

France, under the mentorship of Professor Odile Bain and Dr. Ivan Ineich. She has a strong interest in Sauropsids

and Lissamphibians, studying their host-parasite interactions. She has mainly studied the comparative distribu-

tion of nematode species from the East African mountains and Madagascar. Her research on parasitic nematodes

is focused on their morphology, biology, and co-evolutionary relationships with their hosts. Since 2002, she

has characterized and/or described numerous taxa of parasitic nematodes belonging to different Rhabdiasidae

§ Laurent Chirio completed his M.S. in Biology at Orleans University, France in 1980, and his Ph.D. in 1995

at Montpellier University, France, under the direction of Professor Charles Blanc. His research was completed

on the taxonomy and biogeography of the reptiles of the Aures mountains in Algeria. He has also been a Sci-

ences Teacher with Agregation since 1981. Beginning in 1986 he has worked in various African countries and

has reported undescribed reptile taxa from Niger, CAR, Cameroon, and Guinea. His research interests include

taxonomy and biogeography of African reptiles, amphibians, and fishes of the family Poecilidae.

Amphib. Reptile Conserv.

38

December 2015 | Volume 9 | Number 2 | el 08

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [Special Section]: 39-55 (el 10).

The snakes of Niger

Mean-Frangois Trape and Youssouph Mane

l Institut de Recherche pour le Developpement (IRD), UMR MIVEGEC, Lahoratoire de Paludologie el de Zoologie Medicate, B.P. 1386, Dakar,

SENEGAL

Abstract. — We present here the results of a study of 1,714 snakes from the Republic of Niger,

West Africa, collected from 2004 to 2008 at 28 localities within the country. Based on this data,

supplemented with additional museum specimens (23 selected specimens belonging to 10 species)

and reliable literature reports, we present an annotated checklist of the 51 snake species known

from Niger. Psammophis sudanensis is added to the snake fauna of Niger. Known localities for all

species are presented and, where necessary, taxonomic and biogeographic issues discussed.

Key words. Reptilia; Squamata; Ophidia; taxonomy; biogeography; species richness; venomous snakes; Niger Re-

public; West Africa

Citation: Trape J-F and Mane Y. 2015. The snakes of Niger. Amphibian & Reptile Conservation 9(2) [Special Section]: 39-55 (el 10).

Copyright: © 201 5 Trape and Mane. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommercial-

NoDerivatives 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: 1 1 July 201 5; Accepted: 25 November 2015; Published: 29 December 201 5

Introduction

Few studies have been dedicated to the snake fauna of

the Republic of Niger, the largest country of West Africa

with 1,267,000 km 2 between latitudes 11° and 24°N, and

longitudes 0° and 16°E (Fig. 1). The northern part of the

country is Saharan (Fig. 2), the central and southeastern

parts Sahelian (Fig. 3-4), and the southcentral and

southwestern parts Soudanian (Fig. 5). Elevation is low

in most parts of the country, ranging from 200 m to 700

m, the highest point reaching 2,022 m in Air Mountains,

an area of special biogeographical interest in the Sahara

desert (Fig. 6). Several snake specimens collected during

various Saharan expeditions were reported by Pellegrin

(1909), Angel (1932, 1936), Angel and Lhote (1938),

Villiers (1950a, 1950b) and Joger (1981). The snake

fauna of Air Mountains was investigated by Villiers

(1950a) and Kriska (2001). Important snake collections

were made in southwestern Niger by Roman (1974,

1984), and in W National Park by Chirio (2009). Snakes

observed in the Termit Massif were reported by Ineich

et al. (2014). These specimens and/or additional material

from Niger were included in several revisions or regional

studies, in particular by Papenfuss (1969), Leviton and

Anderson (1970), Roman (1972, 1974, 1977, 1984),

Roux-Esteve (1974), Hughes (1976, 1983, 1998), Hahn

and Roux-Esteve (1979), Broadley (1984), Chirio and

Ineich (1991), Hahn and Wallach (1998), Trape (2002),

Broadley and Hughes (2000), Wtister and Broadley

(2003), Trape and Mane (2006a, 2006b), Trape et al.

(2006, 2009, 2012), Crochet et al. (2008), Chirio et al.

(2011), and Sindaco et al. (2013).

Materials and Methods

In January 2004 and February-March 2005, we deposited

cans or buckets half filled with formaldehyde or ethanol in

22 villages in Niger. Cans or buckets — one per village —

were housed by the chief of the village. We asked the

villagers to deposit in these containers the snakes they

killed when they were occasionnaly encountered in

the vicinity of their village. A modest award (300

CFA, i.e., approximately 0.6 US $) was given for each

preserved specimen. In most parts of Niger — as in most

parts of Africa — all species of snakes are feared and

systematically killed when they are encountered. Thus,

the objective of the award was to acknowledge the effort

of carrying killed snakes from surrounding fields to the

village, this without encouraging snake search and killing.

Visits to the villages were organized in February-March

Correspondence. Email: ' jecin-francois.trape@ird.fr 2 y oussouph.mane@ird.fr

Amphib. Reptile Conserv.

39

December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

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Fig. 1. Map of Niger with location of collection localities. See Table 1 for locality numbers. Colors for vegetation areas: Sudanian /

Sahelo-Sudanian: green; Sahelian: light green; Saharan: yellow for sandy areas, white for stony areas, grey for rocky and mountain-

ous areas.

2005, September-October 2005, and January 2008 to

retrieve the specimens. During travels we also collected

snakes at six additional localities. The 28 collecting

localities (Table 1 and Fig. 1) were distributed either in

the southern part of the country (ll 0 52’N-14 0 52’N: 21

localities), where average annual rainfall ranges from

800 to 300 mm with a South-North gradient, or in the

northern arid part of the country (15°06’N-19°07 , N: 7

localities), including Air Mountains, where rains range

from 250 to less than 50 mm (Mahe et al. 2012).

Most specimens were deposited at the Institut de

Recherche pour le Developpement (Dakar, Senegal;

acronym: IRD), but some specimens — including those of

Rhagerhis moilensis used for comparison with the type

series of Rhamphiophis maradiensis — were donated

to the Museum national d’Histoire naturelle (Paris,

France; acronym: MNHN). We also examinated selected

specimens from Niger from the Institut Fondamental

d’Afrique Noire in Dakar (acronym: IFAN), the

Laboratoire de Bioecologie des Vertebres in Montpellier

(acronym: BEV), MNHN and Laurent Chirio private

collection.

Specimens were identified to species according to

classical identification keys for West African snakes

(Trape and Mane 2006b, Chippaux 2006), recent

revisions of several genera (Trape et al. 2009, Trape et

al. 2012) and further taxonomic analysis (Trape et al.,

unpublished). For recent changes in snake generic names,

we usually follow those adopted in the reptile database of

Uetz and Hosek (http://www.reptile-database.org/).

Results

We collected a total of 1,714 specimens and examined

23 selected additional specimens from IFAN (two speci-

mens), MNHN (17 specimens), BEV (one specimens) or

Chirio’s private collection (three specimens). They be-

longed to 43 species. Eight additional species are known

with certainty from Niger but were not represented

among the specimens we examined.

Family Typhlopidae Gray, 1845

Afrotyphlops lineolatus (Jan, 1864)

Material: One specimen.

Locality: Tela (1).

Literature records: Gaya (Chirio 2009, in error).

Remark: Our Tela specimen, the first known from Niger,

was quoted in error from Gaya by Chirio (2009).

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 10

The snakes of Niger

Table 1 . Collection localities of snakes in Niger (this study). *A: January 2004 - February 2005; B: March 2005 - October 2005;

C: November 2005 - January 2008; D: occasional encounters during travels.

N°

Locality

Latitude

Longitude

Elevation

Region

No of

specimens

No of

species

Sampling

period*

1

Aborah

15°53’N

06°53’E

510 m

Central

3

2

B

2

Ahole

13°33’N

04°01’E

225 m

South Central

150

9

A, B, C

3

Baboul

13°42’N

08°35’E

454 m

South Central

62

8

A, B

4

Chetimari

13°12’N

12°25’E

314 m

South East

60

6

A, B, C

5

Cissia

13°52’N

10°25’E

390 m

South East

80

13

A, B, C

6

Gaya

11°52’N

03°26’E

170 m

South West

1

D

7

Goudoumaria

13°42’N

lLll’E

348 m

South East

10

3

B

8

Gougaram

18°27’N

07’48’E

503 m

Air

1

A

9

Iferouane

19°03’N

08°25’E

660 m

Air

1

A

10

Karosofoua

13°37’N

06°37’E

316 m

South Central

91

10

A, B, C

11

Kelle

14°16’N

10°06’E

456 m

South East

9

B, C

12

Korri Solomi

17°37’N

07°40’E

467 m

Air

2

A

13

Kusa

13°42’N

09°34’E

406 m

South Central

19

8

A, B

14

Malbaza

13°57’N

05°30’E

324 m

South Central

51

5

B, C

15

Maradi

13°47’N

07°26’N

411 m

South Central

1

D

16

Niamey (airport)

13°28’N

02°10’E

226 m

South West

1

D

17

Piliki

13°08’N

01°57’E

210 m

South West

159

15

B, C

18

Saboulayi

13°30’N

07°50’E

440 m

South Central

70

8

A, B, C

19

Saouna

15°07’N

05°42’E

401 m

Central

1

B

20

Simiri (vicinity)

14°02’N

02°05’E

244 m

South West

1

D

21

Taghmert (6 km N)

19°06’N

09°02’E

794 m

Air

1

D

22

Tahoua

14°52’N

05°16’E

387 m

South Central

2

1

D

23

Tarka Dakouara

14°12’N

08°49’E

465 m

South Central

315

10

A, B, C

24

Tchintoulous

18°34’N

08°47’E

826 m

Air

1

A

25

Tekhe

14°01’N

06°01’E

323 m

South Central

209

11

B, C

26

Tela

12°08’N

03°28’E

193 m

South Central

170

21

A, B, C

27

Toundi Farkia

14°02’N

01°32’E

208 m

South West

20

5

B, C

28

Tounga Yacouba

13°55’N

05°26’E

306 m

South Central

223

10

A, B, C

Afrotyphlops punctatus (Leach, 1819)

Material: One specimen.

Locality: Bimi N’Konni (1, coll. MNHN).

Literature records: Birni N’Konni (Pellegrin 1909, Pa-

penfuss 1969, Roux-Esteve 1974); SW Niger (Roman

1974: One specimen).

Family Leptotyphlopidae Stejneger, 1892

Myriopholis adleri (Hahn and Wallach, 1998)

Material: Two specimens.

Locality: Gaya (2, coll. Chirio).

Literature records: Gaya (Chirio 2009).

Remarks: Despite the rarity of records, this species now

appears to occupy the whole sudano-sahelian belt from

Senegal to Chad but avoids the more sahelian areas

contrary to Myriopholis houeti (Trape 2006b, Trape, in

preparation).

Amphib. Reptile Conserv. 41

Myriopholis algeriensis (Jacquet, 1895)

Material: One specimen.

Locality: Agadez (1, coll. MNHN).

Literature records: Agadez (Angel 1932, as Leptotyph-

lops macrorhynchus), Agadez (Angel and Lhote 1938,

Villiers 1950a, as Leptotyphlops macrorhynchus ); Air

(Kriska 2001, as Leptotyphlops macrorhynchus ); Agadez

(Trape 2002, as Leptotyphlops algeriensis).

Myriopholis houeti (Chabanaud, 1917)

Material: Two specimens.

Locality: Kelle (1), Gaya (1, coll. Chirio).

Literature records: Gaya (Chirio 2009).

Myriopholis cairi (Dumeril and Bibron, 1844)

Material: Eight specimens.

Locality: Bilma (8, coll. MNHN).

December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

Fig. 2. The Tenere desert near Adrar Chiriet (19°17’N, 09°14’E).

Literature records: Bilma (Angel 1936, Angel and Lhote

1938, as Leptotyphlops macrorhynchus bilmaensis; Hahn

and Roux-Esteve 1979, Hahn and Wallach 1998, Trape

2002, as Leptotyphlops cairi)\ Teouar (Villiers 1950a,

1950b, as Leptotyphlops macrorhynchus bilmaensis).

Remarks: IFAN 47-4-38 from Teouar (Air Mountains)

is apparently lost: we have been unable to find it in Da-

kar or Paris. However, data on this specimen provided by

Villiers (1950b) exclude Myriopholis algeriensis, Myrio-

pholis boueti, Myriopholis adleri, and Myriopholis lan-

zai, and fit well with Myriopholis cairi.

Tricheilo stoma bicolor (Jan, 1 860)

Material: One specimen.

Locality: Niamey Airport (1).

Literature records: Niamey, Tapoa (Hahn and Roux-

Esteve 1979, Hahn and Wallach 1998, as Leptotyphlops

bicolor)\ Gaya, Campement Nigercar (Chirio 2009).

Family Boidae Gray, 1825

Eryx colubrinus (Linnaeus, 1758)

Material: Three specimens.

Localities: Cissia (1), Tarka Dakouara (2).

Literature records: Agadez, Tabello (Villiers 1950a,

1950b, Papenfuss 1969); Air (Kriska 2001).

Remarks: In Niger this species was known from Air

Mountains and Tamesna, i.e., 300 km north of Tarka

Dakouara and Cissia, but not from the southern part of

the country. Since Cissia is only 60 km from northeast-

ern Nigeria and shares similar sahelian vegetation, our

data suggest that this species may also reach this country

where it has never been mentioned.

Eryx muelleri Boulenger, 1892

Material: 104 specimens.

Localities: Aborach (1), Ahole (17), Baboul (2), Cheti-

mari (4), Cissia (2), Karosofoua (2), Kelle (1), Kusa (1),

Maradi (1), Saboulayi (8), Tarka Dakouara (30), Tekhe

(9), Tela (17), Toundi Farkia (2), Tounga Yacouba (7).

Literature records: SW Niger (Roman 1974: 27 speci-

mens); Air (Kriska 2001); Alambare, Gaya, Gourgou,

Koure (Chirio 2009); Termit (Ineich et al. 2014).

Family Pythonidae Fitzinger, 1826

Python regius (Shaw, 1802)

Material: No specimen collected.

Literature records: SW Niger (Roman 1974: Two speci-

mens); Alambare (Chirio 2009).

Python sebae (Gmelin, 1788)

Material: No specimen collected.

Literature records: SW Niger (Roman 1974: Four speci-

mens); 11 km NW of Niamey (Broadley 1984); Gaya,

Mekrou-Direct (Chirio 2009).

Remarks: In Sahelo and Sahelo-Soudanian areas, this

species is associated with perennial rivers, lakes, and

marshlands. None of our study villages was located near

the Niger River (Fig. 7), Lake Chad or other perennial

waters.

Family Lamprophiidae Fitzinger, 1843

Subfamily Atractaspidinae Bourgeois, 1968

Atractaspis micropholis Giinther, 1872

Material: 11 specimens.

Amphib. Reptile Conserv.

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December 2015 I Volume 9 I Number 2 I el 10

The snakes of Niger

Fig. 3. Atypical view of the Sahel north of Niamey (14°05’N, 01°42’E).

Localities: Kusa (1), Maradi (1, coll. MNHN), Saboulayi

(9).

Literature records: Kusa, Saboulayi, Maradi (Trape et al.

2006); Gaya (Chirio 2009).

Atractaspis watsoni Boulenger, 1908

Material: 33 specimens.

Localities: Bimi N’Konni (1, coll. MNHN), Chetimari

(2), Cissia (1), Karosofoua (5), Malbaza (1), Piliki (6),

Saboulayi (1), Tekhe (16).

Literature records: Birni N’Konni (Pellegrin 1909, as

Atractaspis nigra (holotype), see Trape et al. 2006); Birni

N’Konni (Laurent 1950, Papenfuss 1969, as Atractaspis

microlepidota micropholis ); SW Niger (Roman 1974,

as Atractaspis microlepidota micropholis ); Karosofoua,

Ader de Tahoua (Trape et al. 2006); Gourgou (Chirio

2009).

Subfamily Lamprophiinae Fitzinger, 1843

Boaedon fuliginosus (Boie, 1827)

Material: 16 specimens.

Localities: Chetimari (1), Cissia (2), Karosofoua (1), Pi-

liki (2), Tekhe (8), Tela (2).

Literature records: SW Niger (Roman 1974: Nine speci-

mens); Alambare, Dagaraga, Tapoa (Chirio 2009).

Boaedon lineatus Dumeril, Bibron and Dumeril, 1854

Material: Three specimens.

Locality: Tela (3).

Literature records: SW Niger (Roman 1974: Four speci-

mens); Gaya (Chirio 2009).

Gonionotophis grand (Gunther, 1863)

Material: No specimen exa min ed.

Literature records: Gourgou (Chirio 2009).

Lycophidion semicinctum (Dumeril, Bibron and Du-

meril, 1854)

Material: One specimen.

Locality: Tela (1).

Literature records: Gayia (Chirio 2009).

Mehelya crossi (Boulenger, 1895)

Material: 11 specimens.

Locality: Tela (11).

Literature records: Gayia (Chirio 2009).

Remarks: The Tela records were plotted on the grid map

in Trape and Mane (2006b). Recently, Kelly et al. (2011)

dumped several file snakes into the genus Gonionoto-

phis. However, on the basis of dentition and osteology

there appear to be several genera involved (D.G. Broad-

ley, in litt.) and thus we prefer to provisionally keep all

the West African file snakes in the genus Mehelya.

Subfamily Prosymininae Kelly, Barker, Villet and

Broadley, 2009

Prosymna greigerti collaris (Sternfeld, 1908)

Material: Five specimens.

Localities: Piliki (2), Tela (2), Tounga Yacouba (1).

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

Fig. 4. Field in the Sahel near Chetimari in southwestern Niger during the dry season (13°15’N, 12°28’E).

Literature records: SW Niger (Roman 1974, as Prosymna

meleagris : Two specimens); Alambare, Gaya, La Tapoa

(Chirio 2009); Alambare, Koure, La Tapoa, Malbaza (in

error), Piliki, Tounga Yacouba, Tela (Chirio et al. 2011).

Subfamily Psammophiinae Dowling, 1967

Hemirhagerrhis nototaenia (Gunther, 1864)

Material: One specimen.

Locality: Maradi (1, coll. MNHN).

Literature records: Maradi (Chirio and Ineich 1993,

Broadley and Hughes 2000; picture of the Maradi speci-

men in Trape and Mane 2006b).

Psammophis aegyptius Marx, 1958

Material: Three specimens.

Localities: Korri Solomi (1), Adrar Bous (1, BEV coll.),

Oued Er Roui (1, MNHN coll.).

Literature records: Agadez (Villiers 1950a, 1950b, Pa-

penfuss 1969, as Psammophis schokari ); cliff of Tiguidit

(Dragesco-Joffe 1993, as Psammophis schokari), Termit

(Ineich et al. 2014).

Remarks: It is unclear if P. schokari also occurs in Ni-

ger (see Dragesco-Joffe 1993), but all specimens we ex-

amined had the high number of ventrals of P. aegyptius

(Trape and Mane 2006b).

Psammophis elegans (Shaw, 1802)

Psammophis elegans univittatus Perret, 1961

Material: 32 specimens, including four univittatus.

Localities: Baboul (3 + 1 univittatus), Cissia (3), Gou-

doumaria (6), Kelle (1), Kusa (1), Piliki (6 + 3 univit-

tatus), Tela (8).

Literature records: SW Niger (Roman 1974); Gaya, La

Tapoa (Chirio 2009).

Remarks: The satus of univittatus initially described

from northern Cameroon is unclear. Hughes (circa 1998,

unpublished document) reports specimens from Mali,

Niger (La Tapoa, Garin, Maradi, Soku), Nigeria, Camer-

oon, and Central African Republic). This taxon is char-

acterized by a single vertebral brown line, and lacking

those usually present on the flanks in elegans. It appears

sympatric with elegans in Niger and is also distributed

in Burkina Faso where five specimens from Bam area

(13°20’N, 01°30’W) of Roman’s collection are attribut-

able to univittatus (J.-L. Trape, unpublished). Molecular

studies are needed to clarify whether univittatus deserves

taxonomic recognition or is simply intraspecific varia-

tion.

Psammophis lineatus (Dumeril, Bibron, and Dumeril,

1854)

Material: No specimen exa min ed.

Literature records: SW Niger (Roman 1974, as Dromo-

phis lineatus: 23 specimens); Point triple (Chirio 2009).

Psammophis praeornatus (Schlegel, 1837)

Material: Ten specimens.

Localities: Cissia (5), Kelle (1), Malbaza (1), Piliki (1),

Tekhe(l), Tela (1).

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 10

The snakes of Niger

Fig. 5. View of the Sudan savanna in W National Park in southwestern Niger during the dry season (12°25’N, 02°30’E).

Literature records: SW Niger (Roman 1974, as Dromo-

phis praeornatus : 13 specimens); Gaya, La Tapoa (Chirio

2009).

Psammophis sibilans (Linnaeus, 1758)

Material: 622 specimens.

Localities: Ahole (52), Baboul (22), Chetimari (42),

Cissia (50), Goudoumaria (3), Karosofoua (64), Kelle

(1), Kusa (6), Malbaza (30), Piliki (28), Saboulayi (30),

Saouna (1), Tarka Dakouara (100), Tekhe (80), Tela (20),

Toundi Farkia (4), Tounga Yacouba (89).

Literature records: Azzel (Villiers 1950a, 1950b, Papen-

fuss 1969); SW Niger (Roman 1974: 101 specimens);

Alambare, Dagaraga, Gaya, Gourgou, Koure, La Tapoa,

Moli Haoussa, campement Nigercar (Chirio 2009).

Remarks: We attribute these specimens to P. sibilans

(type locality: Egypt) pending a comprehensive molecu-

lar study that incorporates specimens from the full range

of the P sibilans complex. Such specimens are character-

ized by five infralabials in contact with the first pair of

mentals, a divided anal, and a more-or-less striped dorsal

pattern, with at least a black and white chain on the scales

of the vertebral line (this chain is occasionally absent in

the Sahel, but always present in Sudan and Guinea sa-

vanna areas).

Psammophis sudanensis Werner, 1919

Material: One specimen.

Locality: Tarka Dakouara (1).

Remarks: First record for Niger. This species is charac-

terized by four infralabials in contact with the first pair of

mentals and a typical head pattern, with a median yellow

line starting from the back of the rostral and reaching the

front of the parietals, i.e., crossing the median part of the

frontal contrary to P. sibilans.

Rhagerhis moilensis (Reuss, 1834)

Material: 18 specimens.

Localities: Ahole (4), Baboul (1), Chetimari (1), Cissia

(6), Gari’n Bakwai (3, MNHN coll.), Kelle (1), Kusa (1),

Tarka Dakouara (3), Tounga Yacouba (1).

Literature records: Between Air and Adrar (Angel and

Lhote 1938); Gari’n Bakwai (Chirio and Ineich 1991,

as Rhamphiophis maradiensis ); Termit (Dragesco-Joffe

1993); Air, Tamesna (Kriska 2001); Termit (Ineich et al.

2014).

Remarks: Chirio and Ineich (1991), when describing Rh-

amphiophis maradiensis on the basis of three specimens

from Gari’n Bakwai near Maradi (Niger), unfortunately

omitted to compare their new species with Rhagerhis

moilensis. We have examined the types of Rhamphiophis

maradiensis that are preserved in MNHN. We consider

the two species to be synonymous as they have the same

head shape, body color pattern, and meristic data. Ventral

counts ranged from 166 to 172 in males and from 165

to 1 82 in females for our material from Niger. To facili-

tate further comparisons, our material is now deposited

in MNHN.

Rhamphiophis oxyrhynchus (Reinhardt, 1843)

Material: 26 specimens.

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

Fig. 6. View of Air Mountains in northern Niger (19°06’N, 08°54’E).

Localities: Ahole (1), Karosofoua (1), Simiri (1), Tekhe

(5), Tela (3), Tounga Yacouba (15).

Literature records: SW Niger (Roman 1974: three speci-

mens); Dogondoutchi, Maradi, Sakabal, Gari’n Bakwai

(Chirio and Ineich 1991).

Family Colubridae Oppel, 1811

Subfamily Colubrinae Oppel, 1811

Crotaphopeltis hotamboeia (Laurenti, 1768)

Material: 14 specimens.

Localities: Ahole (4), Piliki (1), Tarka Dakouara (1), Tela

(5), Tounga Yacouba (5).

Literature records: Bebeye, Bimi N’Konni (Pellegrin

1909, as Leptodira hotamboeia ); Birni N’Konni (Pa-

penfuss 1969); SW Niger (Roman 1974: 34 specimens);

Alambare, La Tapoa, Mekrou-Direct, Point triple (Chirio

2009).

Dasypeltis gansi Trape and Mane, 2006

Material: Three specimens.

Localities: Cissia (1), Piliki (1), Tela (1).

Literature records: Cissia, Piliki, Tela (Trape and Mane

2006a); Alambare, Gaya, La Tapoa, Point triple (Chirio

2009).

Dasypeltis sahelensis Trape and Mane, 2006

Material: 70 specimens.

Localities: Ahole (2), Baboul (2), Cissia (3), Karosofoua

(4), Piliki (15), Korri Solomi (1), Saboulayi (1), Tarka

Dakouara (31), Tekhe (1), Tela (5), Tounga Yacouba (5).

Literature records: Ahole, Baboul, Karosofoua, Piliki,

Korri Solomi, Saboulayi, Tarka Dakouara (Trape and

Mane 2006a); Gaya (Chirio 2009).

Lytorhynchus diadema (Dumeril, Bibron, and Dumeril,

1854)

Material: No specimen examined.

Literature records: 39 miles N of Tanout (Leviton and

Anderson 1970).

Meizodon coronatus (Schlegel, 1837)

Material: Two specimens.

Localities: Karosofoua (1), Tela (1).

Literature records: Gaya (Chirio 2009).

Remark: The Tela specimen, the first known from Ni-

ger, appeared in the distribution map of Trape and Mane

(2006b).

Philothamnus irregularis (Leach, 1819)

Material: Nine specimens.

Locality: Tela (9).

Literature records: SW Niger (Roman 1974: seven speci-

mens); Gaya, Gourgou (Chirio 2009).

Philothamnus semivariegatus smithi Bocage, 1882

Material: Four specimens.

Locality: Tela (4).

Remarks: Trape and Mane (2006b) attributed West Afri-

can populations of P. semivariegatus to a distinct subspe-

cies “R semivariegatus ssp ” — differing from the nomi-

nal subspecies by its dorsal coloration: almost uniformly

green in West Africa, versus green with black crossbars

Amphib. Reptile Conserv.

46

December 2015 I Volume 9 I Number 2 I el 10

The snakes of Niger

Fig. 7. The Niger River near Ayorou in eastern Niger (14°42’N, 00°55’E).

in southern, eastern, and central Africa. Trape and Balde

(2014) revived smithi Bocage, 1882, for this subspecies.

Literature records: Gourgou (Chirio 2009).

Remark: The Tela specimens, the first known from Ni-

ger, appeared in the distribution map of Trape and Mane

(2006b).

Spalerosophis diadema cliff ordi (Schlegel, 1837)

Material: 86 specimens.

Localities: Ahole (18), Tchintoulous (1), Baboul (8), Cis-

sia (1), Karosofoua (3), Kelle (1), Kusa (3), Saboulayi

(7), Tarka Dakouara (33), Tekhe (5), Tounga Yacouba

(6).

Literature records: Vicinity of Agadez (Angel and Lhote

1938, as Coluber diadema)', Agadez, Tabello (Villiers

1950a, 1950b, as Coluber diadema ); Agadez, Tabello

(Papenfuss 1969); SW Niger (Roman 1974: 18 speci-

mens); Air (Kriska 2001).

Telescopus tripolitanus (Werner, 1909)

Material: 73 specimens.

Localities: Ahole (22), Baboul (1), Karosofoua (2), Kelle

(1), Malbaza (5), Piliki (7), Saboulayi (1), Tarka Dak-

ouara (5), Tekhe (18), Tela (2), Toundi Farkia (2), Toun-

ga Yacouba (7).

Literature records: Tahoua (Angel and Lhote 1938, Pa-

penfuss 1969, as Taborphis variegatus)', Agadez, Tabello

(Villiers 1950a, 1950b, Papenfuss 1969, as Taborphis ob-

tusus ); Niamey (Villiers 1951, Papenfuss 1969, as Tabor-

phis variegatus)', Agadez (Papenfuss 1969); SW Niger

(Roman 1974, as Telescopus obtusus : 19 specimens); SW

Niger (Roman 1977: six mapped localities); Air (Kriska

2001, as Telescopus obtusus)', Agadez, Tabelot, Maradi,

Piliki, Tela, Ahole, Tounga Yacouba, Malbaza, Tekhe,

Karosofoua, Saboulayi, Baboul, Kelle, Tondi Farkia

(Crochet et al. 2008); Gaya, Koure (Chirio 2009).

Subfamily Grayiinae Kelly, Barker and Villet,

2003

Grayia smithi (Leach, 1818)

Material: One specimen.

Localities: Gaya (1).

Literature records: SW Niger (Roman 1974: 24 speci-

mens).

Remarks: No specimen was collected by Chirio (2009) in

W National Park, but Roman’s collection comprised 24

specimens from southwestern Niger, most of them prob-

ably collected along the Niger River or its perennial and

semi-perennial tributaries.

Family Natricidae Boie, 1827

Natriciteres olivacea (Peters, 1854)

Material: No specimen exa min ed.

Literature records: southwestern Niger, without locality

(Roman 1984).

Remarks: Roman (1984) also reported Natriciteres full g-

inoides (Gunther, 1858) from Niger, but it was probably

a misidentified N. olivacea since he confused the two

species in Burkina Faso (see Trape 2005). The rare, con-

firmed records of N. fuliginoides in West Africa are all

located close to rainforest areas (Trape, in preparation).

Family Elapidae Boie, 1827

Elapsoidea semiannulata moebiusi (Werner, 1897)

Material: One specimen.

Amphib. Reptile Conserv. 47 December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

Locality: Tela (1).

Literature records: SW Niger (Roman 1974: one speci-

men); Gayia, La Tapoa (Chirio 2009).

Naja haje (Linnaeus, 1758)

Material: Eight specimens.

Localities: Cissia (3), Tahoua (2), Tekhe (3).

Literature records: Agadez (Villiers 1950a, Papenfuss

1969); SW Niger (Roman 1974: one specimen probably

attributable to Naja senegalensis)\ Air, Tamesna (Kriska

2001); Cissia, Tekhe, Tahoua, Zinder (Trape et al. 2009);

Gayia (Chirio 2009).

Naja melanoleuca Hallowell, 1857

Material: No specimen exa min ed.

Literature records: SW Niger (Roman 1974: four speci-

mens).

Naja nigricollis Reinhardt, 1843

Material: 66 specimens.

Localities: Goudoumaria (1), Kusa (4), Piliki (14), Tela

(39), Toundi Farkia (8).

Literature records: SW Niger (Roman 1974: 19 speci-

mens); Dagaraga, Gayia, La Tapoa, Moli Haoussa, Point

triple (Chirio 2009).

Naja nubiae Wtister & Broadley, 2003

Material: Two specimens.

Locality: Irabellaben (2, coll. IFAN).

Fiterature records: Irabellaben (Villiers 1950a, 1950b,

Papenfuss 1969, as Naja nigricollis , Wtister and Broad-

ley 2003, Trape and Mane 2006b); Air (Kriska 2001, as

Naja nigricollis).

Naja senegalensis Trape, Chirio, and Wtister, 2009

Material: Three specimens.

Focalities: Karosofoua (2), Tela (1).

Fiterature records: Karosofoua, Tela (Trape et al. 2009);

campement Nigercar (Chirio 2009).

Family Viperidae Oppel, 1811

Bitis arietans (Merrem, 1 820)

Material: Four specimens.

Focalities: Cissia (2), Kusa (2).

Fiterature records: Kimbouloua (Pellegrin 1909); Aga-

dez, Azzel, Dabaga, Tassesset (Villiers 1950a, as Bitis la-

chesis); Tassenet (Villiers 1950b, as Bitis lachesis); Tas-

sesset (Papenfuss 1969), SW Niger (Roman 1974: four

specimens); Air (Kriska 2001); Gaya, Mekrou-Direct

(Chirio 2009).

Causus maculatus (Hallowell, 1 842)

Material: One specimen.

Focadty: Piliki (1).

Amphib. Reptile Conserv. 48

Fiterature records: SW Niger (Roman 1974: six speci-

mens); Dagaraga, Gaya, Fa Tapoa, Mod Haoussa (Chirio

2009).

Cerastes cerastes (Finnaeus, 1758)

Material: Three specimens.

Focadty: Aborah (2), Iferouane (1).

Fiterature records: Dungas, Nguigmi (Pedegrin 1909, as

Cerastes cornutus)\ Agadez, Kaouar, Chirfa, Djado (An-

gel and Fhote 1938); Agadez, Dabaga, Oued In Kakane

near In Gad, Kori Tessouba (Villiers 1950a, Papenfuss

1969); Agadez (Villiers 1950b); 120 km SE of Arlit (Jo-

ger 1981); Air, Tamesna (Kriska 2001); Termit (Ineich et

al. 2014). See also Trape and Mane (2006b) and Sindaco

et al. (2013).

Remark: The Iferouane specimen had no “horns.”

Cerastes vipera (Finnaeus, 1758)

Material: One specimen.

Focality: Six km N of Taghmert (1).

Literature records: Erg of Bilma, erg of Tenere, cliff

of Tiguidit, Termit (Dragesco-Joffe 1993); Air (Kriska

2001); Termit (Ineich et al. 2014). See also Trape and

Mane (2006b) and Sindaco et al. (2013).

Echis leucogaster Roman, 1972

Material: 446 specimens.

Focalities: Ahole (29), Baboul (22), Chetimari (10),

Karosofoua (6), Kelle (1), Malbaza (14), Piliki (62), Sab-

oulayi (13), Tarka Dakouara (110), Tekhe (64), Tela (19),

Toundi Farkia (4), Tounga Yacouba (92).

Fiterature records: Tabello (Villiers 1950a, 1950b, as

Echis carinatus)', route de Dosso, Oualam, Boubon, Nia-

mey, five km W of Niamey, 10 km N of Niamey, 15 km

NW of Niamey, 27 km S of Niamey, Tondikouare, Kou-

tere, Hamdallaye, Koure, Sarandobeni, Tagabati, Saguia,

Tiourridi, Sargadji, Doulgou, Malgorou, Kolo, Sokorbe

(Roman 1972); SW Niger (Roman 1974: 82 specimens);

Boubon, Fido (Roman 1976); Agadez, Tabello, Boubon,

Doulgou, Koure, Malgorou, Niamey, Sargadji, Tin Akof,

Tiourdi (Hughes 1976); 10 km N of Dabnou, Dogon-

Doutchi (Joger 1981); Gaya, Koure (Chirio 2009); Ter-

mit (Ineich et al. 2014). See also Trape and Mane (2006b)

and Sindaco et al. (2013, as Echis pyramidum).

Echis ocellatus Stemmier, 1970

Material: 25 specimens.

Focalities: Piliki (9), Tela (17).

Fiterature records: Bebeye (Pedegrin 1909, as Echis car-

inatus ); Boubon, Gaya, Tiouridi (Roman 1972); SW Ni-

ger (Roman 1974: seven specimens); Boubon, Fido (Ro-

man 1976); Bebeye, Boubon, Gaya, Tiouridi (Hughes

1976); Alambare, Gaya (Chirio 2009).

December 2015 I Volume 9 | Number 2 | el 10

The snakes of Niger

Table 2. Checklist of snake species of Niger.

Species

First documented report

Ecological zone in Niger

Afrotyphlops lineolatus

Trape and Mane 2015

Sudan savanna

Afrotyphlops punctatus

Pellegrin 1909

Sudan savanna

Atractaspis micropholis

Trape et al. 2006

Sudan savanna / Sahel

Atractaspis watsoni

Trape et al. 2006

Sudan savanna / Sahel

Bids arietans

Pellegrin 1909

Sudan savanna / Sahel / Air

Boaedon fuliginosus

Roman 1974

Sudan savanna / Sahel

Boaedon lineatus

Roman 1974

Sudan savanna

Causus maculatus

Roman 1974

Sudan savanna / Sahel

Cerastes cerastes

Pellegrin 1909

Sahara / Air

Cerastes vipera

Dragesco-Joffe 1993

Sahara / Air

Crotaphopeltis hotamboeia

Pellegrin 1909

Sudan savanna / Sahel

Dasypeltis gansi

Trape and Mane 2006a

Sudan savanna

Dasypeltis sahelensis

Trape and Mane 2006a

Sudan savanna / Sahel / Air

Echis leucogaster

Roman 1972

Sudan savanna / Sahel / Sahara / Air

Echis ocellatus

Pellegrin 1909

Sudan savanna

Elapsoidea semiannulata

Roman 1974

Sudan savanna

Eryx colubrinus

Villiers 1950

Sahel / Air

Eryx muelleri

Roman 1974

Soudan savanna / Sahel / Air

Gonionotophis grand

Chirio 2009

Sudan savanna

Grayia smithi

Roman 1974

Sudan savanna

Hemirhagerrhis nototaenia

Chirio and Ineich 1993

Sudan savanna

Lycopl lid ion semicinctum

Chirio 2009

Sudan savanna

Lytorhynchus diadema

Leviton and Anderson 1970

Sahara

Mehelya crossi

Trape and Mane 2006b

Sudan savanna

Meizodon coronatus

Trape and Mane 2006b

Sudan savanna

Myriopholis algeriensis

Trape 2002

Sahara / Air

Myriopholis adleri

Chirio 2009

Sudan savanna

Myriopholis boned

Chirio 2009

Sudan savanna / Sahel

Myriopholis cairi

Hahn and Roux-Esteve 1979

Sahara / Air

Naja haje

Villiers 1950a

Sahel / Air

Naja melanoleuca

Roman 1974

Sudan savanna

Naja nigricollis

Roman 1974

Sudan savanna / Sahel

Naja nubiae

Wiister and Broadley 2003

Air

Naja senegalensis

Trape et al. 2009

Sudan savanna

Natriciteres olivacea

Roman 1984

Sudan savanna

Philothamnus irregularis

Roman 1974

Sudan savanna

Philothamnus semivariegatus

Trape and Mane 2006b

Sudan savanna

Prosymna greigerd

Roman 1974

Sudan savanna

Psammophis aegypdus

Trape and Mane 2006b

Sahara / Air

Psammophis elegans

Roman 1974

Sudan savanna / Sahel

Psammophis lineatus

Roman 1974

Sudan savanna

Psammophis praeornatus

Roman 1974

Sudan savanna / Sahel

Psammophis sibilans

Villiers 1950a

Sudan savanna / Sahel / Air

Psammophis sudanensis

Trape and Mane 2015

Sudan savanna

Python regius

Roman 1974

Sudan savanna

Python sebae

Roman 1974

Sudan savanna, Sahel

Rhagerhis moilensis

Angel and Lhote 1938

Sahara / Sahel / Air

Rhamphiophis oxyrhynchus

Roman 1974

Sudan savanna

Spalerosophis diadema

Villiers 1950a

Sudan savanna / Sahel / Air

Telescopus tripolitanus

Roman 1977

Sudan savanna / Sahel / Air

Tricheilostoma bicolor

Hahn and Roux-Esteve 1979

Sudan savanna

Amphib. Reptile Conserv.

49

December 2015 | Volume 9 | Number 2 | el 10

Trape and Mane

Discussion

Our collection of Nigerian snakes comprises 1,714 speci-

mens belonging to 38 species. With additional museum

material that we examined and accepting reliable litera-

ture reports the snake fauna of Niger comprises 5 1 species

(Table 2), i.e., 19 species more than the previous check-

list established by Roman (1984). The first checklist for

Niger (Papenfuss 1969) comprised only 15 species. It is

unclear whether P schokari also occurs in Niger, or if

only P. aegyptius is present. Data points probably in er-

ror for Naja katiensis and Atractaspis dahomeyensis in

maps by Chippaux (2006) are not retained here, but these

two species may still occur in southwestern Niger since

close records exist for Burkina Faso (Naja katiensis ) and

Benin (Atractaspis dahomeyensis ). As previously men-

tioned in Trape and Mane (2006b), Rhamphiophis mara-

diensis is a junior synonym of Rhagheris moilensis. The

occurrence of Psammophis sudanensis in Niger, a rare

species in West Africa (Trape and Mane 2006b, Trape

and Balde 2014), has not previously been noted.

North of 15°N, in the most arid part of the country

(rains < 250 mm), the snake fauna comprises at least 17

species; with six typical Saharan species: Myriopholis

algeriensis, Myriopholis cairi, Lytorhynchus diadema,

Psammophis aegyptius, Cerastes cerastes, and Ceras-

tes vipera\ eight Sahelo-Saharan species: Eryx colubri-

nus, Eryx muelleri, Dasypeltis sahelensis, Spalerosophis

diadema cliffordi, Telescopus tripolitanus, Rhagerhis

moilensis, Naja nubiae, and Echis leucogaster, one

Sahelo-Sudanian species: Naja haje; and two species

widely distributed in West African savannas including

the northern Sahel: Psammophis sibilans and Bit is ari-

etans. In these areas, only nine specimens were collected

during our study. Even if the duration of sampling was

much lower than south of 15°N for most sites, this may

reflect a lower density of snakes. However, it may also

reflect more limited participation in the study by nomads

contrary to settled agricultural workers. Some specific

beliefs may also have played a role, e.g., for some north-

ern populations killing a Psammophis is taboo. Our inter-

views of local populations suggested that at least Ceras-

tes cerastes and Psammophis aegyptius are common in

many areas of northern Niger.

Maximum diversity was observed in the southern

part of the country, between 12°00’N and 14°00’N,

where the snake fauna comprises at least 43 species, in-

cluding either: Sahelo-Saharan: Eryx colubrinus, Eryx

muelleri, Dasypeltis sahelensis, Spalerosophis diadema

cliffordi, Telescopus tripolitanus, Rhagerhis moilensis,

and Echis leucogaster, Sudanian and Sahelian: Myrio-

pholis adleri, Myriopholis boueti, Meizodon coronatus,

Prosymna greigerti collaris, Psammophis praeornatus,

Psammophis sudanensis, Rhamphiophis oxyrhynchus,

Elapsoidea semiamnnulata moebiusi, Naja haje, and

Naja senegalensis', or species widely distributed in West

African savannas: Afrotyphlops lineolatus, Afrotyph-

Amphib. Reptile Conserv. 50

lops punctatus, Tricheilo stoma bicolor. Python regius.

Python sebae, Boaedon fuliginosus, Boaedon lineatus,

Crotaphopeltis hotamboeia, Dasypeltis gansi, Goniono-

tophis grand, Grayia smithi, Hemirhagerrhis nototaenia,

Lycophidion semicinctum, Mehelya crossi, Natriciteres

olivacea, Philothamnus irregularis, Philothamnus semi-

variegatus smithi, Psammophis elegans, Psammophis

lineatus, Psammophis sibilans, Naja nigricollis, Naja

melanoleuca, Bids arietans, Causus maculatus, and Ech-

is ocellatus.

Despite the relatively high number of species record-

ed south of 14°N, many species were rarely collected

and diversity was low in most areas. Two species rep-

resented together almost two-third of the 1,705 snakes

that were collected south of 15°N: Psammophis sibilans

(621 specimens, 36.4 %), and Echis leucogaster (446

specimens, 26.2 %). Five additional species represented

at least 2% of the snakes that were collected: Eryx muel-

leri (104 specimens, 6.1 %), Spalerosophis diadema clif-

fordi (86 specimens, 5.0 %), Telescopus tripolitanus (72

specimens, 4.2%), Dasypeltis sahelensis (69 specimens,

4.0 %), and Naja nigricollis (66 specimens, 3.9 %). Two

species were close to 2%: Atractaspis watsoni (33 speci-

mens, 1.9%), and Psammophis elegans (32 specimens,

1.9%). In fact, except south of 13°N, snake diversity was

low in almost all sampling sites, e.g., only 10 different

species in Tarka Dakouara (14°12’N, 08°49’E) despite

315 specimens collected, but 21 species for 170 speci-

mens collected in Tela (12°08’N, 03°28’E), our south-

ernmost study area.

Regarding snakebite management, our data highlight

the danger represented by Echis leucogaster and Naja

nigricollis. These two highly venomous species are both

abundant and widely distributed in the most populated

areas of Niger, particularly Echis leucogaster which

probably occurs throughout the whole country. Among

the other dangerous species, Cerastes cerastes. Ceras-

tes vipera, Naja nubiae, and Naja haje are essentially

distributed in the most arid regions of the country, and

Echis ocellatus, Naja senegalensis, Naja melanoleuca,

Atractaspis watsoni, and Atractaspis micropholis in Su-

dan savanna areas.

The extensive collections made by Roman (1974,

1984) and Chirio (2009) in southwestern Niger, where

rains, permanent surface waters, and biodiversity are the

highest, combined with Air mountains records by Villiers

(1950) have provided a relatively comprehensive over-

view of the snake fauna of Niger. However, among the

species of our collection, five were new for Niger when

collected (i.e., Afrotyphlops lineolatus, Myriopholis bou-

eti, Meizodon coronatus, Philothamnus semivariegatus

smithi, and Psammophis sudanensis), three belonged

to new species that we described elsewhere (Dasypeltis

gansi, D. sahelensis (Trape and Mane 2006a) and Naja

senegalensis (Trape et al. 2009), and two belonged to

species that we have revived from the synonymy of

December 2015 | Volume 9 | Number 2 | el 10

The snakes of Niger

Atractaspis microlepidota (i.e., A. watsoni and A. micro-

pholis).

Acknowledgments. — We thank G. Diatta for assis-

tance during field work and G. Chauvancy for assistance

during preparation of the map and appendix. L. Chirio

contributed to snake collection in Air Moutains. L. Chirio

and I. Ineich provided useful complementary data for our

checklist of the snake fauna of Niger. L. Luiselli and an

anonymous reviewer provided useful comments on the

manuscript.

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Lamprophis Fitzinger 1843 and Mehelya Csiki 1903.

Molecular Phylogenetics and Evolution 58: 415^-26.

Kriska MA. 2001. Contribution a V inventaire

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Mahe G, Rouche N, Dieulin C, Boyer JF, Ibrahim B,

Cres A, Servat E, Valton C, Paturel JE. 2012. Carte

des pluies annuelles en Afrique / Annual rainfall map

of Africa. Bondy, IRD.

Papenfuss TJ. 1969. Preliminary analysis of the reptiles

of arid central West Africa. Wasman Journal of. Biol-

ogy 27: 249-325.

Pellegrin J. 1909. Reptiles du Soudan recoltes par la mis-

sion Tilho-Gaillard. Description d’une espece nou-

velle. Bulletin du Museum National d’Histoire Na-

turelle de Paris 15: 4 1 3 — 4 1 5 .

Roman B. 1972. Deux sous especes de la vipere Echis

carinatus (Schneider) dans les territoires de la Haute -

Volta et du Niger : Echis carinatus ocellatus Stem-

mier, Echis carinatus leucogaster n. ssp. Notes et

Documents voltai'ques 5: 1-15.

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glyphes) Wagler, 1830 en Haute-Volta et dans l’Ouest

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Trape and Mane

du Niger. Notes et Documents voltaiques 10: 92-98.

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1-313.

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Western Palearctic. 2. Annotated checklist and distri-

butional atlas of the snakes of Europe, North Africa,

Middle East and Central Asia, with an update of Vol.

1. Edizioni Belvedere, Latina, Italy. 543 p.

Trape JF. 2002. Note sur la repartition et le statut de

quelques Leptotyphlopides (Serpentes, Scolecophid-

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Bulletin de la Societe Herpetologique de France 102:

49-62.

Trape JF. 2005. Note sur quelques serpents meconnus

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39^19.

Trape JF, Balde C. 2014. A checklist of the snake fauna

of Guinea, with taxonomic changes in the genera Phi-

lothanmus and Dipsadoboa (Colubridae) and a com-

parison with the snake fauna of some other West Afri-

can countries. Zootaxa 3900: 301-338.

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(Serpentes: Colubridae) en Afrique de l’Ouest: de-

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velles. Bulletin de la Societe Herpetologique de

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Occidentale. Savane et Desert. IRD editions, Paris,

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logeography and systematic revision of the Egyptian

cobra (Serpentes: Elapidae: Naja haje) species com-

plex, with the description of a new species from West

Africa. Zootaxa 2236: 1-25.

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microlepidota, A. micropholis et A. watsoni en Af-

rique occidentale et centrale. Bulletin de la Societe

Herpetologique de France 119: 5-16.

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lour patterns reflect phylogeography - New spe-

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488-501.

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kar, Senegal, Africa. 155 p.

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(acquisitions 1950). Bulletin de l ’Institut Frangais

d’Afrique Noire 13: 813—836.

Wiister W, Broadley DG. 2007. Get an eyeful of this: a

new species of giant spitting cobra from eastern and

northeastern Africa (Squamata: Serpentes: Elapidae:

Naja). Zootaxa 1532: 51-68.

Jean-Frangois Trape is a French medical doctor, biologist, and herpetologist with lengthy ex-

perience in Africa, where he was born (1949). Since 1980 he has worked continuously in Africa

for the Institut de Recherche pour le Developpement (IRD, formerly ORSTOM), a French public

institution for research in Southern countries. Presently he is Emeritus Research Director at the

IRD Laboratory of Malarialogy and Medical Zoology located in Dakar, Senegal. Jean-Frangois has

authored or co-authored over 280 peer-reviewed papers and books on tropical medicine and her-

petology, including the books “Guide des serpents d’Afrique occidentale. Savane et desert” (2006)

and “Lezards, crocodiles et tortues d’Afrique occidentale et du Sahara” (2012). During his career

he has authored or co-authored the descriptions of 23 reptile and five tick species. He is also a ma-

laria expert for the World Health Organization, where he has served in several steering committees.

In 2010 he received the first IRD prize for research, and in 2013 the Lucien Tartois prize from the

French Foundation for Medical Research.

Youssouph Mane is a Senegalese biologist and herpetologist born in 1961 in the Casamance Prov-

ince of southern Senegal. His master dissertation at the University Cheikh Anta Diop of Dakar in

1992 investigated the snake fauna in the vicinity of Dielmo, a well preserved savanna area near the

Sine-Saloum National Park in central Senegal. In 1997 Youssouph‘s doctorate thesis was on the

ecology of bees in Casamance. After his thesis, he entered the Institut de Recherche pour le Devel-

oppement at Dakar, participated in many herpetological field surveys in West Africa, and served as

the curator of the IRD reptile collection. Youssouph has authored or co-authored 22 peer-reviewed

papers and the book entitled “Guide des serpents d’Afrique occidentale. Savane et desert” (2006,

with J-FT). Over his career to date, he has authored or co-authored the description of seven snake

and two amphisbaenian species.

Amphib. Reptile Conserv.

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The snakes of Niger

APPENDIX: list of specimens examined (IRD collection, Dakar).

Afrotyphlops lineolatus. Tela: TR.4448.

Atractaspis micropholis. Kusa: 5.N; Saboulayi: 34.N, 358.N, 375.N, 376.N, 377.N, 378.N, 379.N, 918.N, 930 N.

Atractaspis watsoni. Chetimari: 845.N, 861.N; Cissia: 1069.N; Karosofoua: 297. N, 298.N, 299.N, 903.N; Malbaza: 464.N; Pi-

liki: 301. N, 302.N, 352.N, 1407.N, 1444.N, 1450.N; Saboulayi: 357.N; Tekhe: 686.N, 757.N, 758.N, 762.N, 769.N, 775.N, 781.N,

787.N, 808.N, 815.N, 1274.N, 1282.N, 1290.N, 1303.N, 1336.N, 1353.N.

Bids arietans. Cissia: 1052.N, 1087.N; Kusa: 216.N, 227.N.

Boaedon fuliginosus. Chetimari: 863.N; Cissia: 1065. N, 1066.N; Karosofoua: 827.N; Piliki: 1412.N, 1457.N; Tekhe: 1275.N,

1335.N, 1346.N, 1347.N, 1362.N, 1379.N, 1386.N, 698.N; Tela: 272.N, 720.N.

Boaedon lineatus. Tela: 264.N, 711.N, 1564.N.

Causus maculatus. Piliki: 349.N.

Cerastes cerastes . Aborah: 356.N, TR.1513.

Cerastes vipera. Taghmert: TR.1548.

Crotaphopeltis hotamboeia. Ahole: 90.N, 91.N, 93.N, 580.N; Piliki: 345.N; Tarka Dakouara:125.N; Tela: 277.N, 286.N, 713.N,

714.N, 717.N; Tounga Yacouba: 33.N, 564.N, 1660.N.

Dasypeltis gansi. Cissia: 252.N; Piliki: 331.N; Tela: 273. N.

Dasypeltis sahelensis. Ahole: 1022.N, 587.N; Baboul: 394.N, 59.N; Cissia: 1051.N, 1071.N, 1083.N; Karosofoua: 820.N, 831.N,

899.N, 908.N; Korri Solomi: TR.1545; Piliki: 1405.N, 1418.N, 1434.N, 1437.N, 1445.N, 1452.N, 1466.N, 1467.N, 1468.N,

1470.N, 1473.N, 1491.N, 305.N, 309.N, 315.N; Saboulayi: 189.N; Tarka Dakouara: 10.N, 106.N, 115.N, 120.N, 130.N, 133.N,

163.N, 399.N, 400.N, 401.N, 402.N, 423.N, 431.N, 432.N, 433.N, 435.N, 444.N, 1106.N, 1112.N, 1149.N, 1195.N, 1206.N, 1240.N,

1262.N, 1269.N, 1273.N, 1703.N, 1704.N, 1705.N, 1706.N, 1707.N; Tekhe: 1363.N; Tela: 1543.N, 1552.N, 1561.N, 1569.N,

1579.N; Tounga Yacouba: 1662.N, 1686.N.

Echis leucogaster. Ahole: 95.N, 570.N, 572.N, 574.N, 593.N, 603.N, 609.N, 611.N, 615.N, 617.N, 623.N, 627.N, 628.N, 972.N,

974.N, 980.N, 990.N, 992.N, 1002.N, 1004.N, 1005.N, 1016.N, 1023.N, 1024.N, 1025.N, 1026.N, 1027.N, 1028.N, 1034.N;

Baboul: 45.N, 46.N, 52.N, 54.N, 62.N, 63.N, 65.N, 68.N, 69.N, 75.N, 76.N, 80.N, 81.N, 82.N, 83.N, 86.N, 386.N, 390.N, 391.N,

393.N, 395.N, 396.N; Chetimari: 233.N, 234.N, 235.N, 236.N, 240.N, 242.N, 243.N, 244.N, 656.N, 847.N; Karosofoua: 210.N,

212.N, 214.N, 215.N, 872.N, 878.N; Kelle: 934.N; Malbaza: 469.N, 471.N, 472.N, 937.N, 939.N, 944.N, 945.N, 946.N, 949.N,

952.N, 956.N, 958.N, 964.N, 966.N; Piliki: 310.N, 311.N, 312.N, 313.N, 314.N, 317.N, 320.N, 321.N, 329.N, 333.N, 334.N,

335.N, 336.N, 337.N, 339.N, 341.N, 342.N, 343.N, 346.N, 700.N, 701.N, 702.N, 703.N, 705.N, 706.N, 707.N, 709.N, 825.N,

1398.N, 1399.N, 1400.N, 1402.N, 1411.N, 1415.N, 1416.N, 1417.N, 1420.N, 1421.N, 1423.N, 1425.N, 1428.N, 1429.N, 1433.N,

1435.N, 1441. N, 1442.N, 1448.N, 1449.N, 1453.N, 1455.N, 1458.N, 1459.N, 1464.N, 1465.N, 1469.N, 1474.N, 1476.N, 1478.N,

1482.N, 1485.N, 1487.N, 1489.N; Saboulayi: 179.N, 363.N, 364.N, 366.N, 916.N, 917.N, 922.N, 923.N, 924.N, 925.N, 927.N,

931.N, 932.N; Tarka Dakouara: 105.N, 109.N, 114.N, 119.N, 122.N, 132.N, 141.N, 149.N, 152.N, 153.N, 156.N, 161.N, 162.N,

403. N, 404. N, 405.N, 406.N, 415.N, 416.N, 417.N, 418.N, 419.N, 421.N, 424.N, 426.N, 428.N, 436.N, 437.N, 440.N, 441.N,

445. N, 446. N, 451.N, 452.N, 1107.N, 1108.N, 1110.N, llll.N, 1113.N, 1114.N, 1115.N, 1116.N, 1118.N, 1121.N, 1123.N, 1125.N,

1127.N, 1133.N, 1136.N, 1137.N, 1141.N, 1142.N, 1143.N, 1146.N, 1147.N, 1148.N, 1150.N, 1152.N, 1153.N, 1154.N, 1155.N,

1156.N, 1157.N, 1164.N, 1165.N, 1167.N, 1171.N, 1173.N, 1178.N, 1180.N, 1181.N, 1182.N, 1183.N, 1185.N, 1187.N, 1199.N,

1201.N, 1203.N, 1204.N, 1207.N, 1209.N, 1210.N, 1211.N, 1212.N, 1215.N, 1217.N, 1220.N, 1223.N, 1226.N, 1228.N, 1229.N,

1233.N, 1234.N, 1236.N, 1237.N, 1239.N, 1244.N, 1248.N, 1251.N, 1252.N, 1253.N, 1254.N, 1258.N, 1259.N, 1261.N, 1263.N,

1265.N, 1266.N, 1267.N, 1268.N; Tekhe: 685.N, 759.N, 763.N, 764.N, 765.N, 766.N, 767.N, 772.N, 774.N, 782.N, 784.N, 785.N,

790.N, 792.N, 796.N, 799.N, 801.N, 802.N, 803.N, 813.N 1276.N, 1279.N, 1280.N, 1283.N, 1284.N, 1294.N, 1295.N, 1296.N,

1297.N, 1298.N, 1299.N, 1306.N, 1316.N, 1317.N, 1324.N, 1325.N, 1332.N, 1337.N, 1338.N, 1341.N, 1344.N, 1345.N, 1349.N,

1350.N, 1356.N, 1358.N, 1366.N, 1369.N, 1370.N, 1372.N, 1374.N, 1375.N, 1376.N, 1380.N, 1382.N, 1383.N, 1388.N, 1389.N,

1390.N, 1391.N, 1392.N, 1394.N, 1396.N, 1397.N; Tela: 4.N, 276.N, 287.N, 288.N, 292.N, 727.N, 734.N, 735.N, 740.N 1526.N,

1538.N, 1547.N, 1548.N, 1551.N, 1554.N, 1555.N, 1562.N, 1563.N, 1568.N; Toundi Farkia: 1044.N, 1045.N, 1047.N, 1049.N;

Tounga Yacouba: 42.N, 43.N, 44.N, 473.N, 486.N, 487.N, 496.N, 497.N, 502.N, 503.N, 504.N, 505.N, 507.N, 508.N, 509.N,

510.N, 513.N, 520.N, 522.N, 524.N, 525.N, 526.N, 530.N, 531.N, 537.N, 539.N, 541.N, 546.N, 548.N, 549.N, 551.N, 552.N,

554.N, 555.N, 556.N, 557.N, 558.N, 559.N, 560.N, 561.N, 562.N, 563.N, 565.N, 566.N, 1583.N, 1586.N, 1587.N, 1589.N, 1590.N,

1592.N, 1594.N, 1595.N, 1599.N, 1600.N, 1602.N, 1606.N, 1608.N, 1614.N, 1616.N, 1618.N, 1620.N, 1624.N, 1625.N, 1632.N,

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1634.N, 1635.N, 1637.N, 1641.N, 1648.N, 1650.N, 1653.N, 1654.N, 1661.N, 1665.N, 1666.N, 1667.N, 1670.N, 1671.N, 1673.N,

1675.N, 1676.N, 1677.N, 1679.N, 1680.N, 1683.N, 1685.N, 1690.N, 1691.N, 1692.N, 1694.N, 1699.N.

Echis ocellatus. Piliki: 323.N, 324.N, 145 l.N, 1475.N, 1479.N, 1480.N, 148 l.N, 1483.N, 1484.N; Tela: 716.N, 729.N, 733.N,

744.N, 750.N 1502.N, 1516.N, 1525.N, 1529.N, 1539.N, 1541.N, 1542.N, 1553.N, 1558.N, 1577.N, 1580.N.

Elapsoidea semiannulata moebiusi. Tela: 747.N.

Eryx colubrinus. Cissia: 1089.N; Tarka Dakouara: 1190.N, 1224.N.

Eryx muelleri. Aborah: 355.N; Ahole: 87.N, 94.N, 577.N, 578.N, 588.N, 590.N, 595.N, 610.N, 612.N, 616.N, 977.N, 998.N,

999.N, 1000.N, 1700.N, 1701. N, 1702.N; Baboul: 77.N, 392.N; Chetimari: 230.N, 23 l.N, 834.N, 854.N; Cissia: 108 l.N, 1086.N;

Karosofoua: 873.N, 890.N; Kelle; 642.N; Kusa: 226.N; Maradi: TR.4450; Saboulayi: 174.N, 362.N, 368.N, 369.N, 381.N,

914.N, 919.N, 920.N; Tarka Dakouara: 128.N, 129.N, 147.N, 407.N, 411.N, 414.N, 425.N, 427.N, 438.N, 448.N, 449.N, 453.N,

1124.N, 1126.N, 1132.N, 1144.N, 1168.N, 1179.N, 1191. N, 1200.N, 1205.N, 1208.N, 1214.N, 1216.N, 1218.N, 1242.N, 1250.N,

1255.N, 1257.N, 1272.N; Tekhe: 770.N, 783.N, 798.N, 1285.N, 1288.N, 1320.N, 1354.N, 1364.N, 1393.N; Tela: 294.N, 295.N,

715.N, 723.N, 724.N, 725.N, 726.N, 732.N, 736.N, 1505.N, 1506.N, 1513.N, 1528.N, 1550.N, 1557.N, 1560.N, 1575.N; Toundi

Farkia: 372.N, 1050.N; Tounga Yacouba: 495.N, 499.N, 533.N, 1597.N, 1631.N, 1663.N, 1698.N.

Lycophidion semicinctum. Tela: 1532.N.

Mehelya crossi. Tela: 271.N, 282.N, 285.N, 293.N, 710.N, 730.N, 1495.N, 1500.N, 1507.N, 1511.N, 1535.N.

Meizodon coronatus. Karosofoua: 882.N; Tela: 722.N.

Myriopholis boueti. Kelle: 936. N.

Naja haje. Cissia; 246.N, 248.N, 672.N; Tahoua: TR.4442, 832.N; Tekhe: 60.N, 690.N, 1395.N.

Naja nigricollis. Goudoumaria: 661.N; Kusa: 218.N, 220.N, 221.N, 223. N; Piliki: 303.N, 306.N, 307.N, 326.N, 338.N, 340.N,

348.N, 351.N, 1408.N, 1409.N, 1431.N, 1439.N, 1446.N, 1447.N; Tela: 265.N, 266.N, 267.N, 268.N, 269.N, 270.N, 278.N, 289.N,

29 l.N, 737.N, 738.N, 739.N, 741.N, 742.N, 743.N, 745.N, 746.N, 751.N, 752.N, 753.N, 754.N 1496.N, 1497.N, 1498.N, 1499.N,

1501.N, 1503.N, 1508.N, 1519.N, 1520.N, 1521.N, 1524.N, 1534.N, 1549.N, 1571.N, 1574.N, 1576.N, 1578.N, 1581.N; Toundi

Farkia: 1035.N, 1036.N1037.N, 1038.N, 1040.N, 1041. N, 1042.N, 1046.N.

Naja senegalensis. Karosofoua: 20 l.N, 910.N; Tela: 1504.N.

Philothamnus irregularis . Tela: 274.N, 275. N, 279.N, 280.N, 283. N, 290.N, 296.N, 712.N 1523.N.

Philothamnus semivariegatus smithi. Tela: 755.N, 1527.N, 1537.N.

Prosymna greigerti collaris. Piliki: 347.N, 1472.N; Tela: 1531.N, 1545.N; Tounga Yacouba: 536.N.

Psammophis aegyptius. Korri Solomi: TR.4449.

Psammophis elegans. Baboul: 73.N, 78.N, 85. N; Cissia: 262.N, 263.N, 674.N; Goudoumaria: 662.N, 663.N, 664.N, 665.N,

666. N, 669. N; Kelle: 935.N; Kusa: 648.N, Piliki: 308.N, 316.N, 318.N, 1422.N, 1443.N, 1454.N; Tela: 719.N, 1514.N, 1518.N,

1522.N, 1546.N, 1556.N, 1570.N, 1573.N.

Psammophis elegans univittatus. Baboul: 388.N; Piliki: 1432.N, 1436.N, 147 l.N.

Psammophis praeornatus. Cissia: 253.N, 257.N, 260.N, 261.N, 675.N; Kelle: 641.N, Malbaza: 467.N; Piliki: 1490.N; Tekhe:

1378.N; Tela: 1572.N.

Psammophis sibilans. Ahole. 88.N, 89.N, 92.N, 567.N, 568.N, 569.N, 571.N, 573.N, 576.N, 579.N, 581.N, 582.N, 583.N, 584.N,

586.N, 594.N, 596.N, 598.N, 599.N, 601.N, 602.N, 608.N, 613.N, 618.N, 619.N, 620.N, 622.N, 624.N, 630.N, 631.N, 632.N,

970.N, 973.N, 976.N, 981.N, 983.N, 984.N, 985.N, 986.N, 987.N, 988.N, 996.N, 997.N, 1001.N, 1008.N, 1014.N, 1017.N, 1018.N,

1020.N, 1029.N, 103 l.N, 1032.N; Baboul: 47.N, 48.N, 49.N, 53.N, 56.N, 57.N, 58.N, 60.N, 61.N, 64.N, 66.N, 67.N, 70.N, 71.N,

72.N, 74.N, 79. N, 84.N, 385.N, 387.N, 397.N, 398.N; Chetimari: 229.N, 232.N, 237.N, 238.N, 239.N, 241.N, 649.N, 650.N,

65 l.N, 652.N, 653.N, 654.N, 655.N, 657.N, 658.N, 659.N, 660.N, 833.N, 835.N, 836.N, 837.N, 838.N, 839.N, 840.N, 841.N,

842.N, 843. N, 844.N, 846.N, 848.N, 849.N, 850.N, 85 l.N, 852.N, 853.N, 855.N, 856.N, 858.N, 859.N, 860.N, 862.N, 864.N; Cis-

sia: 245. N, 247. N, 249.N, 250.N, 25 l.N, 254.N, 255.N, 256.N, 258.N, 259.N, 67 l.N, 673.N, 1053.N, 1054.N, 1055.N, 1056.N,

1057.N, 1058.N, 1059.N, 1060.N, 1061.N, 1062.N, 1063.N, 1064.N, 1067.N, 1068.N, 1072.N, 1073.N, 1074.N, 1075.N, 1076.N,

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The snakes of Niger

1077.N, 1078.N, 1082.N, 1084.N, 1085.N, 1088.N, 1090.N, 1091.N, 1092.N, 1095.N, 1096.N, 1097.N, 1098.N, 1099.N, 1100.N,

1101.N, 1102.N, 1103.N, 1104.N,; Goudoumaria: 667.N, 668.N, 670.N; Karosofoua: 190.N, 191.N, 192.N, 193.N, 194.N, 195.N,

196.N, 197.N, 198.N, 199.N, 200.N, 202.N, 203.N, 204.N, 205.N, 206.N, 207.N, 208.N, 211.N, 213.N, 817.N, 818.N, 819.N,

821.N, 822. N, 823.N, 824.N, 826.N, 828.N, 829.N, 830.N, 65.N, 866.N, 867.N, 868.N, 869.N, 870.N, 874.N, 875.N, 876.N,

877.N, 879.N, 880.N, 881.N, 883.N, 884.N, 885.N, 886.N, 888.N, 891.N, 892.N, 894.N, 895.N, 896.N, 897.N, 898.N, 901.N,

904.N, 905. N, 906.N, 907.N, 909.N, 911.N, 912.N; Kelle: 640.N; Kusa: 217.N, 222.N, 224.N, 225.N, 644.N, 646.N; Malbaza:

454.N, 455.N, 456.N, 457.N, 458.N, 460.N, 461.N, 462.N, 463.N, 465.N, 466.N, 470.N, 938.N, 940.N, 941.N, 942.N, 943.N,

947. N, 948.N, 950.N, 954.N, 955.N, 957.N, 960.N, 961.N, 962.N, 963.N, 967.N, 968.N, 969.N; Piliki: 300.N, 304.N, 319.N,

322.N, 325. N, 327.N, 328.N, 330.N, 353.N, 704.N,1403.N, 1404.N, 1406.N, 1410.N, 1413.N, 1414.N, 1419.N, 1424.N, 1426.N,

1427.N, 1430.N, 1438.N, 1440.N, 1462.N, 1463.N, 1477.N, 1486.N, 1488.N; Saboulayi: 165.N, 166.N, 167.N, 169.N, 170.N,

171.N, 172.N, 173.N, 175.N, 176.N, 177.N, 178.N, 180.N, 182.N, 184.N, 185.N, 186.N, 187.N, 188.N, 359.N, 360.N, 365.N,

380.N, 384.N, 913.N, 915.N, 921.N, 928.N, 929.N, 933.N; Saouna: 354.N; Tarka Dakouara: 96.N, 97.N, 98.N, 99.N, 100.N,

101.N, 102.N, 103.N, 104.N, 107.N, 108.N, 110.N, lll.N, 112.N, 113.N, 116.N, 117.N, 118.N, 121.N, 124.N, 126.N, 127.N,

131.N, 134.N, 135.N, 136.N, 137.N, 138.N, 139.N, 140.N, 142.N, 143.N, 144.N, 145.N, 146.N, 148.N, 150.N, 151.N, 154.N,

155.N, 157.N, 158.N, 160.N, 164.N, 408.N, 410.N, 412.N, 434.N, 439.N, 442.N, 443.N, 447.N, 450.N, 1105.N, 1109.N, 1120.N,

1128.N, 1129.N, 1130.N, 1131.N, 1135.N, 1138.N, 1145.N, U51.N, 1159.N, 1160.N, 1161.N, 1162.N, 1166.N, 1169.N, 1170.N,

1172.N, 1175.N, 1176.N, 1177.N, 1189.N, 1193.N, 1194.N, 1196.N, 1197.N, 1198.N, 1202.N, 1213.N, 1222.N, 1225.N, 1227.N,

123.N, 1230.N, 1232.N, 1235.N, 1238.N, 1241.N, 1243.N, 1245.N, 1247.N, 1249.N, 1256.N, 1260.N, 1270.N, 1271.N; Tekhe:

676.N, 677.N, 678.N, 679.N, 680.N, 681.N, 682.N, 683.N, 684.N, 687.N, 688.N, 689.N, 691.N, 692.N, 693.N, 694.N, 695.N,

696.N, 697. N, 699.N, 756.N, 761.N, 768.N, 771.N, 773.N, 776.N, 777.N, 780.N, 793.N, 797.N, 800.N, 809.N, 814.N, 816.N

1277.N, 1281.N, 1286.N, 1287.N, 1291.N, 1300.N, 1301.N, 1302.N, 1304.N, 1305.N, 1307.N, 1308.N, 1309.N, 1310.N, 1312.N,

1313.N, 1314.N, 1319. N, 1322.N, 1323.N, 1327.N, 1328.N, 1329.N, 1330.N, 1331.N, 1333.N, 1334.N, 1339.N, 1340.N, 1342.N,

1348.N, 1351.N, 1352.N, 1355.N, 1357.N, 1360.N, 1361.N, 1365.N, 1367.N, 1368.N, 1371.N, 1373.N, 1381.N, 1384.N, 1387.N;

Tela: 718.N, 721.N, 728.N, 748.N, 1492.N, 1493.N, 1494.N, 1509.N, 1510.N, 1512.N, 1515.N, 1517.N, 1530.N, 1533.N, 1540.N,

1544.N, 1559.N, 1565.N, 1566.N, 1567.N; Toundi Farkia: 370.N, 373.N, 1039.N, 1043.N; Tounga Yacouba: 36.N, 37.N, 38.N,

39. N, 41. N, 374.N, 474.N, 475.N, 476.N, 477.N, 478.N, 479.N, 480.N, 481.N, 482.N, 485.N, 489.N, 490.N, 492.N, 500.N, 501.N,

506.N, 511.N, 512.N, 514.N, 515.N, 516.N, 517.N, 518.N, 519.N, 523.N, 527.N, 528.N, 529.N, 532.N, 534.N, 535.N, 542.N,

543. N, 544.N, 550.N, 553.N, 642.N, 1585.N, 1591.N, 1593.N, 1596.N, 1598.N, 1603.N, 1604.N, 1605.N, 1607.N, 1609.N, 1610.N,

1612.N, 1613.N, 1615. N, 1617.N, 1619.N, 1621.N, 1623.N, 1626.N, 1628.N, 1629.N, 1630.N, 1633.N, 1638.N, 1639.N, 1643.N,

1644.N, 1645.N, 1646.N, 1649.N, 1651.N, 1652.N, 1655.N, 1656.N, 1657.N, 1658.N, 1659.N, 1668.N, 1672.N, 1674.N, 1681.N,

1684.N, 1687.N, 1689.N, 1693.N, 1695.N.

Psammophis sudanensis. Tarka Dakouara: 17.N.

Rhagerhis moilensis. Ahole: 636.N, 1012.N, 1019.N, 1033.N; Baboul: 389.N; Chetimari: 857.N; Cissia: l.N, 2.N, 3.N, 1080.N,

1093.N, 1094.N; Kelle: 639.N; Kusa: 645.N, Tarka Dakouara: 409.N, 1158.N, 1174.N; Tounga Yacouba: 1588.N.

Rhamphiophis oxyrhynchus. Ahole: 1013. N; Karosofoua: 209.N; Simiri: TR.270: Tekhe: 811.N 1315.N, 1318.N, 1321.N,

1343.N; Tela: 281.N, 284.N, 731.N; Tounga Yacouba: 40.N, 484.N, 488.N, 491.N, 498.N, 1582.N, 1584.N, 1601.N, 1622.N,

1627.N, 1640.N, 1647.N, 1664.N, 1669.N, 1678.N.

Spalerosophis diadema cliffordi. Ahole: 6.N, 7.N, 575.N, 591.N, 597.N, 600.N, 605.N, 606.N, 607.N, 621.N, 625.N, 635.N, 637.N,

979.N, 989.N, 994.N, 995.N, 1011.N; Baboul: 26.N, 27.N, 28.N, 29.N, 30.N, 50.N, 51.N, 55.N; Cissia: 1070.N; Karosofoua:

87 l.N, 889.N, 900.N; Kelle: 638.N; Kusa: 219.N, 228.N, 647.N; Saboulayi: 168.N, 181.N, 183.N, 361.N, 382.N, 383.N, 926.N;

Tarka Dakouara: 8.N, 9.N, ll.N, 12.N, 13.N, 14.N, 15.N, 16.N, 18.N, 19.N, 20.N, 21.N, 22.N, 23.N, 24.N, 25.N, 159.N, 413.N,

420.N, 422.N, 429.N, 430.N, 1119.N, 1134.N, 1139.N, 1140.N, 1163.N, 1184.N, 1186.N, 1188.N, 1192.N, 1219.N, 1221.N; Tchin-

toulous: TR.4453; Tekhe: 786.N, 789.N, 795.N, 807.N, 1359.N; Tounga Yacouba: 483.N, 494.N, 545.N, 547.N, 1688.N, 1697.N.

Telescopus tripolitanus. Ahole: 1003.N, 1006.N, 1007.N, 1009.N, 1010.N, 1015.N, 1021.N, 1030.N, 585.N, 589.N, 592.N, 604.N,

614.N, 626.N, 629.N, 633.N, 634.N, 971.N, 975.N, 978.N, 982.N, 991.N, 993.N; Baboul: 31.N; Gayia: TR.2351; Karosofoua:

35.N, 902.N; Kelle: 643.N; Malbaza: 459.N, 468.N, 951.N, 953.N, 965.N; Piliki: 332.N, 350.N, 708.N, 1401.N, 1456.N, 1460.N,

1461.N; Saboulayi: 367.N; Tarka Dakouara: 1117.N, 1122.N, 1231.N, 1246.N, 1264.N, Tekhe: 778.N, 779.N, 788.N, 791.N,

794.N, 804.N, 805.N, 806.N, 810.N, 812.N, 1278.N, 1289.N, 1292.N, 1293.N, 1311.N, 1326.N, 1377.N, 1385.N; Tela: 749.N;

Toundi Farkia: 371.N, 1048.N; Tounga Yacouba: 32.N, 493.N, 521.N, 538.N, 540.N 1636.N, 1682.N.

Tricheilostoma bicolor . Niamey (airport): TR.4451.

Amphib. Reptile Conserv.

55

December 2015 | Volume 9 | Number 2 | el 10

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [Special Section]: 56-84 (el 11).

The tadpoles of eight West and Central African Leptopelis

species (Amphibia: Anura: Arthroleptidae)

1 *Michael F. Barej, ^ilo Pfalzgraff, ^areike Hirschfeld, 23 H. Christoph Liedtke,

Johannes Penner, 4 Nono L. Gonwouo, Matthias Dahmen, Tranziska Grozinger,

5 Andreas Schmitz, and ^ark-Oliver Rodel

1 Museum fiir Naturkunde, Leibniz Institute for Evolution and Biodiversity Science, Invalidenstr. 43, 10115 Berlin, GERMANY department of

Environmental Science (Biogeography), University of Basel, Klingelbergstrasse 27, 4056 Basel, SWITZERLAND 3 Ecology \ Evolution and

Developmental Group, Department of Wetland Ecology, Estacion Biologica de Dohana (CS1C), 41092 Sevilla, SPAIN 4 Cameroon Herpetology-

Conservation Biology Foundation (CAMHERP-CBF), PO Box 8218, Yaounde, CAMEROON 5 Natural History Museum of Geneva, Department of

Herpetology and Ichthyology, C.P. 6434, 1211 Geneva 6, SWITZERLAND

Abstract. — The tadpoles of more than half of the African tree frog species, genus Leptopelis, are

unknown. We provide morphological descriptions of tadpoles of eight species from Central and West

Africa. We present the first descriptions for the tadpoles of Leptopelis boulengeri and L. millsoni. In

addition the tadpoles of L. aubryioides, L. calcaratus, L. modestus, L. rufus, L. spiritusnoctis, and

L. viridis are herein reinvestigated and their descriptions complemented, e.g., with additional tooth

row formulae or new measurements based on larger series of available tadpoles.

Key words. Anuran larvae, external morphology, diversity, mitochondrial DNA, DNA barcoding, lentic waters, lotic

waters

Citation: Barej MF, Pfalzgraff T, Hirschfeld M, Liedtke HC, Penner J, Gonwouo NL, Dahmen M, Grozinger F, Schmitz A, Rodel M-0. 2015. The

tadpoles of eight West and Central African Leptopelis species (Amphibia: Anura: Arthroleptidae). Amphibian & Reptile Conservation 9(2) [Special

Section]: 56-84 (el 11).

NoDerivatives 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: 02 October 201 5; Accepted: 14 November 2015; Published: 31 December 201 5

Introduction

Given that sequences of correctly determined species

are available, the application of DNA-barcoding has

facilitated species-assignment of tadpoles. Thus, tad-

pole morphology is more and more frequently included

in species descriptions (e.g., Blackburn 2008a; Das and

Haas 2010; Rodel et al. 2012; Lima et al. 2014; Portillo

and Greenbaum 2014b; Vassilieva et al. 2014) and nu-

merous publications even focus exclusively on tadpole

descriptions. Insights from larval morphology have been

important for recognizing, or hinting at, cryptic species

(e.g., Randrianiaina et al. 2012; Pfalzgraff et al. 2015),

have contributed to systematics (Haas 2003; Muller et

al. 2005) or indicated the presence of range-restricted

taxa and the appropriateness of a habitat for elusive, i.e.,

semi-fossorial species (e.g., Cardioglossa : Hirschfeld et

al. 2012; Leptodactylodon : Cruz et al. 2013; Mapouyat

et al. 2014).

Moreover, detection of tadpoles can be informative for

habitat preferences of species and even more important-

ly, provides direct evidence of successful reproduction

of recorded species even in the absence of adult vouch-

ers (e.g., Hirschfeld et al. 2012). Thus, determination of

tadpoles is beneficial for conservation assessments and

long-term management strategies. However, due to the

bi-phasic life-cycle of anurans, tadpoles and adults are

exposed to different threats in their habitat or during mi-

gration, and conservation efforts should be considered

accordingly (e.g., Becker et al. 2007; Wells 2007).

While four herpetological journals provided insight

on tadpoles of more than 80 species in the last two years

(20 1 4-20 1 5 those dealing with or describing African tad-

poles were relatively few; e.g., Herpetologica : 0/4; Zoo-

taxa : 11/70; Salamandra: 4/9; The Herpetological Jour-

nal. 1/1; accessed 30 September 2015) our knowledge is

still far from complete (Channing et al. 2012).

Correspondence. Email: hnichael@barej.de (Corresponding author)

Amphib. Reptile Conserv.

56

December 201 5 | Volume 9 | Number 2 | el 1 1

Barej et al.

This likewise applies to the genus Leptopelis Gunther,

1859 which is endemic to sub-Saharan Africa and cur-

rently comprises 53 species (Frost 2015). New species

are continuously being added to this list (e.g., Lotters et

al. 2005; Kohler et al. 2006; Rodel 2007; Portillo and

Greenbaum 2014a,b; Gvozdik et al. 2014) and further

species complexes are already known (Portillo et al.

2015; Barej and Rodel, unpubl. data). These medium

to large-sized frogs inhabit a wide variety of vegetation

types, from tropical and subtropical forests to open grass-

lands (Rodel 2000; Channing 2001; Minter et al. 2004;

Channing and Howell 2006; Amiet 2012). The common

name “treefrogs” is not applicable to the entire genus, as

some species are adapted to burrowing and a terrestrial

lifestyle (e.g., Poynton and Broadley 1987; Rodel 2000

and references therein).

Generally, knowledge of the biology and natural his-

tory of Leptopelis is rather incomplete although adver-

tisement calls of more than half of the species are known

(e.g., Amiet and Schiptz 1974; Schiptz 1999; Grafe et

al. 2000; Rodel 2000; Kohler et al. 2006; Greenbaum et

al. 2012; Portillo and Greenbaum 2014b) and anecdotal

observations on predation events by spiders (Barej et

al. 2009), death-feigning reflexes (de Witte 1941; Per-

ret 1966; Kofron and Schmitt 1992; Schmitz et al. 1999;

Rodel et al. 2000), cocoon building (Grafe 2000), and

malacophagy (Perret 1966; Amiet 2012) have been docu-

mented. Furthermore, Leptopelis are featured as magi-

cal creatures used in traditional wars and modern sports

(Pauwels et al. 2003).

Details on the reproduction of Leptopelis species are

generally scarce. As far as known egg deposition occurs

outside water in or on top of moist soil, the development

is slow, and hatching starts when the eggs in their nest

are inundated during the beginning of the rainy season.

Hatched tadpoles then move towards the water where

they develop and metamorphosis takes place (Schiptz

1963, 1975; Oldham 1977; Wager 1986; Rodel 2007). It

is presumed that tadpoles are exotrophic (developmental

energy derived from ingested food as a free-living tad-

pole) and live in the thin muddy layer in the benthos of

lentic waters (Altig and McDiarmid 1999a; Channing

et al. 2012). However, Amiet (2012) also reports on re-

production in lotic waters. Direct development has been

speculated forL. brevirostris (Schiptz 1999).

A simplified morphological description of the de-

scribed Leptopelis tadpoles comprises: an elongated and

eel-like shape, in particular a very long tail with low fins,

and a predominantly dark coloration of body and tail

(Perret 1966; Channing et al. 2012).

Recently, Channing et al. (2012) compiled avail-

able data on African tadpoles including 22 Leptopelis

tadpoles, nine being described for the first time. Since

then, two more Leptopelis tadpoles have been described

(Portillo and Greenbaum 2014b; Penske et al. 2015).

However, several tadpole descriptions in Channing et al.

(2012) were often based on single specimens and require

a through comparison with larger series of specimens as

it is well known that tadpole morphology can be very

variable due to genetic and environmental factors as well

as during development (e.g., Duellman and Trueb 1994;

Laurila and Kujasalo 1999; Relyea 2001; Kraft et al.

2006; Wells 2007).

We herein use larger voucher series to re-describe the

tadpoles of four Central African (L. aubryioides n = 20,

L. calcar atus n= 16, L. modestus n = 3, L. rufus n = 18),

and two West African Leptopelis species (L. spiritusnoc-

tis n = 20, L. viridis n = 2). In addition, we provide the

first descriptions of two other Central African species: L.

boulengeri (n = 16) and L. millsoni ( n = 1).

Materials and Methods

Sampling. Field surveys were carried out in Liberia

and Guinea by M.F. Barej and J. Penner (June 2011);

in Cameroon on Mt. Manengouba, Littoral and South-

West Province by M. Hirschfeld and F. Grozinger (No-

vember 2010 to October 2011), in the Abo Forest, North

West Province by T.M. Doherty-Bone (August 2012),

in the Ebo forest, Littoral Province by M.-O. Rodel, M.

Dahmen, F. Grozinger, and M. Hirschfeld (September

2010 to October 2011), on Mt. Nlonako, Littoral Prov-

ince by M.F. Barej, H.C. Liedtke, N.L. Gonwouo, and

M. Hirschfeld (October 2011), and around Kribi, South

Province and Etome, South-West Province by M.F. Barej,

H.C. Liedtke, and N.L. Gonwouo (October to November

2011). Detailed locality data of investigated tadpoles are

provided in Appendix Table Al. Tadpoles were caught

either by hand or with dip nets. They were anaesthetized

in a tricaine methane sulphonate (MS222, Thomson &

Joseph Ltd), chlorobutanol, or benzocaine solution. For

molecular analyses a piece of tail muscle was removed

and preserved in ethanol (99%) from at least one individ-

ual for each set of morphologically distinct tadpoles for

every locality. All tadpoles were then fixed in formalin

(8%) and later transferred into ethanol (75%).

Determination. Species identity of the tadpoles was

verified by DNA-barcoding, comparing 16S ribosomal

RNA sequences from tadpoles to those of adult vouchers

and/or available GenBank sequences. For comparison

of the partial 16S rRNA a total of 37 sequences (474-

554 bp) has been generated and deposited in GenBank

(KT967076-KT967 112; Appendix Table Al). For details

of extraction, primers, and PCR protocols, and sequenc-

ing see Barej et al. (2014). Sequences were aligned using

ClustalX (Thompson et al. 1997; default parameters) and

manually checked using the original chromatograph data

in the program BioEdit (Hall 1999). Uncorrected p-dis-

tances for the partial 16S rRNA gene between included

Leptopelis species were calculated with PAUP* 4.0b 10

(Swofford 2002).

All tadpoles could be unambiguously assigned to a

valid Leptopelis species. Intraspecific genetic divergenc-

December 2015 I Volume 9 | Number 2 | el 1 1

Amphib. Reptile Conserv.

57

The tadpoles of eight West and Central African Leptopelis species

es ranged from 0.0-0. 8% (Table 1), except in L. rufus

where a 1.5% difference indicated two distinct lineages

herein referred to as L. rufus _1 and L. rufus _2. Voucher

IDs and GenBank numbers of adults and tadpoles are

provided in Appendix Table Al. For further synonyms

and chresonyms used in older publications on Leptopelis

tadpoles see Frost (2015).

Character assessment. Measurements were taken with

a dissecting microscope or digital calliper by one per-

son (TP). Summaries for several individuals are given

as mean values. The following measurements were taken

(for details see Appendix Figure Al): EL (entire length),

BL (body length), TL (tail length), BH (body height at

the point of the spiracle insertion), BW (maximum body

width, in dorsal view), AW (width of the tail muscle

[axis], at the tail base), AH (maximum tail muscle (axis)

height), VF (maximum height of ventral fin), DF (maxi-

mum height of dorsal fin), TTH (total tail height), ED

(horizontal eye diameter), IOD (interocular distance),

IND (internostril distance), SND (snout-nostril distance),

SED (snout-eye distance), ODW (oral disc width), SL

(spiracle length), and SSD (snout- spiracle distance). Dis-

tances including eyes and/or nostrils were taken from

respective centers (e.g., SED: centre of the eye to snout

tip). Measurements of all examined specimens are pro-

vided in Appendix Table A2. The following ratios were

calculated: BL/TL, BH/BL, BW/BL, SND/SED, ED/BL,

IOD/IND, TL/EL, DF/VF, AH/DF, TTH/BH, AW/BW,

AH/BH, SL/BL, ODW/BW, and SSD/BL. Ratios of all

examined specimens are provided in Appendix Table A3;

mean ratios for each species are provided in Appendix

Table A4. The relation of body length to total length was

mostly not measurable in genotyped vouchers, as tail tips

have been removed for tissue sampling. Specimens were

staged according to Gosner (1960) and labial tooth row

formulae are based on Rodel (2000).

Illustrations of genotyped representatives in the best

condition of each taxon were prepared with the help of a

camera lucida on a dissecting microscope. Missing parts

resulting from tissue sampling are drawn as outlines

based on non-genotyped vouchers. Schematic sketches

were made of the oral discs of genotyped tadpoles.

Comparative morphometries. Morphological features

like fin height, body shape or tail length point to adapta-

tions to particular habitat types (e.g., Altig and McDi-

armid 1999b). To assess morphological adaptations in

Leptopelis tadpoles to particular habitats all 1 8 measure-

ments were log 1() transformed and subjected to a rigid ro-

tation via a Principal Component Analysis. Only individ-

uals with full sets of measurements were included, and

so L. viridis and L. rufus _1 were not represented in the

final dataset and L. millsoni and L. modestus were only

represented by one and two individuals, respectively. The

preomp function was used in R v3.2 (R core team 2013),

data was scaled and centered and the ordispider function

Amphib. Reptile Conserv.

Table 1 . Intraspecific genetic distances (uncorrected p) in the

mitochondrial 16S ribosomal RNA between Leptopelis species,

compared to adult individuals (for GenBank# see Appendix

Table Al); SD = standard deviation, n = number of pairwise

comparisons, alignment: 558 bp. Note that the maximum value

in L. rufus results from two lineages in this species; if indepen-

dently analysed both lineages show p-distance values within

the range of remaining taxa: rufus _ 1 (n = 1): 0.43%; rufus _ 2

(n = 10): 0%.

Species

min

max

mean

SD

n

aubryioides

0

0.75

0.37

0.24

36

boulengeri

0

0.19

0.08

0.1

10

calcaratus

0.18

0.6

0.39

0.21

3

millsoni

—

0

—

1

modestus

0

0.2

0.13

0.11

3

rufus

0

1.5

0.66

0.68

21

spiritusnoctis

0

0.83

0.21

0.27

28

viridis

0

0.21

0.11

0.12

6

interspecies

1.92

13.03

8.8

2.2

712

in the vegan package (Oksanen et al. 2013) was used to

add a cluster dendrogram to species groupings.

Results and Discussion

The tadpoles of eight Leptopelis species are described

herein: Leptopelis aubryioides (Andersson, 1907), L.

boulengeri (Werner, 1898), L. calcaratus (Boulenger,

1906), L. millsoni (Boulenger, 1895), L. modestus Wer-

ner, 1898, L. rufus Reichenow, 1874 from Central Africa,

and L. spiritusnoctis Rodel, 2007, and L. viridis (Gun-

ther, 1869) from West Africa. The morphology of the

analyzed tadpoles is generally consistent with the simpli-

fied tadpole diagnosis of the genus Leptopelis provided

by Altig and McDiarmid (1999a): oval/depressed body

shape; generally uniformly dark colored; dorsal eyes;

small nares, nearer snout than eye; labial tooth row for-

mula 3-5/3, usually 2-n rows on upper labium broken

medially and one row on lower labium may be broken;

typical, anteroventral oral apparatus; wide dorsal gap

on marginal distribution; uniserial dorsally and biserial

ventrally; submarginal papillae absent; wide upper jaw

sheath with medial indentation; wide, V-shaped lower

jaw sheath; dextral vent tube; sinistral spiracle; low dor-

sal fin which originates near dorsal tail body junction

ends in a pointed tip.

Leptopelis aubryioides (Andersson, 1907)

The description of L. aubryioides tadpoles is based on

twenty tadpoles: ZMB 79604 (two tadpoles, at Gosner

stages 30 and 36, near Etome, Cameroon, 4.83 17°N;

9.9253°E, 476 m a.s.l., 23 October 2011, the tadpoles

were found in a small muddy puddle along a stream

bank; stream characterised by lots of little rapids), ZMB

79605 (one tadpole at Gosner stage 25) and ZMB 79606

December 2015 | Volume 9 | Number 2 | el 1 1

58

Barej et al.

Fig. 1 . Lateral (A) and dorsal (B) view of Leptopelis aubryioides (ZMB 79605) at Gosner stage

25; coloration of tadpole (ZMB 79604) in life (C); adult L. aubryioides (ZMB 83029) (D); oral

disc opened in life (F); sketch of the oral disc (E); scale bars: 1 mm.

(nine tadpoles at Gosner stages 25 to 40, near Ekom-

tolo, at the foot of Mt. Nlonako, Cameroon, 4.8329°N;

9.9259°E, 477 m a.s.l., 24 October 2011, the tadpoles

were found in a slow flowing forest stream), ZMB 79607

(three tadpoles, at Gosner stages 36 and 39, Njuma, Ebo

Forest, Cameroon, 4.3483°N; 10.2329°E, 238 m a.s.l.,

08 August 2011), ZMB 79608 (one tadpole, at Gosner

stage 40, Njuma, Ebo Forest, Cameroon, 4.3483°N;

10.2329°E, 238 m a.s.l., 19 August 2011), ZMB 79609

(one tadpole, at Gosner stage 31, Njuma, Ebo Forest,

Cameroon, 4.3394°N; 10.2458°E, 320 m a.s.l., 20 Au-

gust 2011), ZMB 79610 (one tadpole, at Gosner stage 36,

Njuma, Ebo Forest, Cameroon, 4.3483°N; 10.2329°E,

238 m a.s.l., 07 October 2011), ZMB 79611 (one tad-

pole, at Gosner stage 41, Njuma, Ebo Forest, Cameroon,

4.3483°N; 10.2329°E, 238 m a.s.l., 08 October 2011)

and ZMB 79612 (one tadpole, at Gosner stage 34, Camp

Njuma, Ebo Forest, Cameroon, 4.3480°N; 10.2323°E,

Amphib. Reptile Conserv.

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December 201 5 I Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

Fig. 2. Lateral (A) and dorsal (B) view of Leptopelis boulengeri (ZMB 79616) at Gosner stage 38; sketch of the oral disc (C); adult

L. boulengeri (ZFMK 87857) (D); scale bars: 1 mm.

315 m a.s.l., 23 September 2011, the locality was situ-

ated in primary rainforest). Proportions including total or

tail length were only available for non-genotyped indi-

viduals.

Description. Body oval with nearly rounded snout in

dorsal view (Fig. IB); ovoid to slightly compressed in

lateral view (Fig. 1A); tail length-body length ratio 2.38

(TL/BL); body height 0.44 of body length (BH/BL);

body width 0.58 of body length (BW/BL); maximum

body width slightly behind the spiracle’s posterior end;

nostrils situated dorsally, slightly closer to snout tip than

eyes (SND/SED = 0.42), distance snout-nostrils 0.20 of

body length (SND/BL); eyes positioned laterally; eye di-

ameter 0. 1 1 of body length (ED/BL); interocular distance

exceeds internostril distance by a factor of 1.93 (IOD/

IND); tail length 0.70 of entire length (TL/EL), with

moderately pronounced fins and narrow fin tip; dorsal

fin originates at dorsal tail-body junction, barely rising

at the first quarter of the tail; dorsal fin slightly curved

with maximum height at three-quarters of the tail length;

ventral fin originates on the ventral terminus of the body;

ventral fin narrower than tail axis with maximum height

at three-quarters of the tail length; maximum fin height in

dorsal fin higher (DF/VF = 1 .29); fin tip pointed; maxi-

mum tail height including fins lower than body height

(TTH/BH = 0.90); tail axis width (in dorsal view) 0.42 of

body width (AW/BW); maximum height of tail axis (at

base) 0.56 of body height (AH/BH); tail axis height (at

base) distinctly higher than maximum height of dorsal fin

(AH/DF = 2.07); dextral vent tube, positioned basicau-

dally; spiracle sinistral, visible in dorsal view, originat-

ing anterior to mid-body (SSD/BL = 0.45); spiracle tube

length 0.14 of body length (SL/BL); mouth opens antero-

ventrally; oral disc width less than quarter of body width

(ODW/BW = 0.24); one row of papillae (with rounded

tips) laterally at anterior lip with huge rostral gap, these

connected to papillae in labial angles and posterior lip;

second row of papillae caudal at posterior lip (Fig. IF);

labial tooth row formula 1/3+3//3 (Fig. IE) or 1/2+2//3;

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December 201 5 I Volume 9 | Number 2 | el 1 1

Barej et al.

jaw sheaths black, of equal width and serrated; upper jaw

widely V-shaped; lower jaw U-shaped.

Coloration in preservation. Dorsolateral part of the

body, tail axis and dorsal fin mostly speckled dark brown

on light brown ground at the body and yellowish ground

at the tail; areas without brown spots shine through as

yellow blots; ventral part of the body light brown with

some dark brown spots at the anterior third of the body;

vent tube translucent; spiracle translucent or pigmented;

ventral fin predominantly translucent with few brown

spots composed of dense melanophores towards tail tip.

Coloration in life (Fig. 1C). Pale brown with shiny

golden speckles at dorsolateral part of the body, tail axis

and dorsal fin; ventral fin translucent with few speckles;

ventral part of the body translucent.

Remarks. Leptopelis aubryioides occurs from eastern

Nigeria through Cameroon to Gabon and the Republic

of the Congo (e.g., Schiptz 1967, 1999, 2007; Fretey and

Blanc 2001; Blanc and Fretey 2004; Amiet 2012). Ami-

et and Schiptz (1974) and Amiet (2006, 2012) reported

on habitat use and the call activity of the species. The

tadpole of L. aubryioides has already been described by

Channing et al. (2012) based on a single specimen, which

belongs to a larger series of tadpoles examined herein

(MH198 = ZMB 79612). Shape of body and tail, as well

as tail shape and overall pigmentation are congruent

with the available description. In addition to the labial

Fig. 3. Lateral (A) and dorsal (B) view of Leptopelis calcaratus (ZMB 79618) at Gosner stage 28; coloration in life of tadpole (ZMB

79618) in lateral (top) and dorsal (below) view (C); adult L. calcaratus (ZFMK 75590) (D); sketch of the oral disc (E); scale bars:

1 mm. Note that the greenish coloration at the tail tip results from a leaf used as the background.

Amphib. Reptile Conserv.

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December 201 5 I Volume 9 I Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

tooth row formula presented by Charming et al. (2012:

1/2+2//3) a second labial tooth formula has been recog-

nized 1/3+3//3 (Fig. IE). While Channing et al. (2012)

refer to a tail length-body length ratio of 2.2, the mean

value of our measures was slightly higher (2.4) in the

present series. Regarding the coloration, pale blotches

are present in our material on the tail as well as the lateral

part of the body (Fig. 1A, C). The spiracle was translu-

cent, lacking any pigmentation.

Leptopelis boulengeri (Werner, 1 898)

The description of L. boulengeri tadpoles is based on

sixteen tadpoles: ZMB 79613 (one tadpole, at Gosner

stage 37, Bekob, Ebo Forest, Cameroon, 4.3578°N;

10.4170°E, 921 m a.s.l., 27 August 2011), ZMB 79614

(four tadpoles, at Gosner stage 36 to 40, Bekob, Ebo

Forest, Cameroon, 4.3578°N; 10.4170°E, 921 m a.s.l.,

28 August 2011), ZMB 79615 (three tadpoles, at Gos-

ner stage 36, Bekob, Ebo Forest, Cameroon, 4.3575°N;

10.4168°E, 903 m a.s.l., 29 August 2011), ZMB 79616

(one tadpole, at Gosner stage 38) and ZMB 79617 (seven

tadpoles, at Gosner stages 36 to 40), Bekob, Ebo Forest,

Cameroon, 4.3578°N; 10.4170°E, 921 m a.s.l., 08 Sep-

tember 2011. Proportions including total or tail length

were only available for non-genotyped individuals.

Description. Body oval with subovoid snout in dorsal

view (Fig. 2B); ovoid to slightly compressed in lateral

view (Fig. 2A); tail length-body length ratio 2.44 (TL /

BL); body height 0.43 of body length (BH/BL); body

width 0.53 of body length (BW/BL); maximum body

width on the level of the spiracle’s posterior end; nostrils

situated dorsally, closer to snout tip than eyes (SND/SED

= 0.41), distance snout-nostrils 0. 14 of body length (SND/

BL); eyes positioned laterally; eye diameter 0.08 of body

length (ED/BL); interocular distance exceeds internostril

distance by a factor of 2.35 (IOD/IND); tail length 0.71

of entire length (TL/EL), with moderately pronounced

fins with narrow fin tip; dorsal fin originates at dorsal

tail-body junction with maximum height at half of the

tail length; dorsal fin and ventral fin particularly curved;

ventral fin originates on the ventral terminus of the body;

ventral fin narrower than tail axis with maximum height

at half of the tail length; maximum fin height in dorsal fin

slightly higher (DF/VF = 1 . 12); fin tip pointed; maximum

tail height including fins exceeds body height (TTH/BH

= 1.20); the tail axis width (in dorsal view) 0.34 of body

width (AW/BW); maximum height of tail axis (at base)

0.55 of body height (AH/BH); tail axis height (at base)

higher than maximum height of dorsal fin (AH/DF =

1.63); dextral vent tube, positioned basicaudally; spira-

cle sinistral, visible in dorsal view, originating anterior to

mid-body (SSD/BL = 0.43); spiracle tube length 0.18 of

body length (SL/BL); mouth opens anteroventrally; oral

disc width wider than a third of body width (ODW/B W =

0.36); one row of papillae (with rounded tips) laterally at

Amphib. Reptile Conserv.

anterior lip with huge rostral gap, these connected to pa-

pillae in labial angles and posterior lip; second and third

row of papillae at posterior lip; labial tooth row formula

1/3+3//3 (Fig. 2C); jaw sheaths black, of equal width and

serrated; upper jaw very widely U-shaped; lower jaw U-

shaped.

Coloration in preservation. Dorsolateral part of the

body mostly speckled dark brown on yellowish ground,

tail axis and dorsal fin speckled with lighter brown spots

on yellowish ground; areas without brown spots shine

through as yellow blots; ventral part of the body yellow

without any spots; spiracle and vent tube yellowish; ven-

tral fin translucent without any brown spots.

Remarks. Leptopelis boulengeri is known from Nige-

ria to Gabon, the Republic of the Congo in the south

and the Democratic Republic of the Congo to the east

(e.g., de la Riva 1994; Schiptz 1967, 1999; Amiet 2012).

Similar to L. aubryioides the species inhabits dense for-

ests with small rivulets and ponds (Schiptz 1967; Amiet

2012). The call and call activity have been reported by

Amiet and Schiptz (1974) and Amiet (2006). The tad-

pole is herein described for the first time. The tadpole of

L. boulengeri exhibits the generic diagnostic characters:

elongated and slender body with a long thin tail (TL/BL

= 2.4) and acute tip (Fig. 2A). The coloration is similar

to other Leptopelis tadpoles with brown spots on yellow-

ish ground, the spots however, being brighter than usual.

The chromatophores on the dorsal part of the body and

the tail are less dense in L. boulengeri than in the remain-

ing examined species, the fin has dorsally only very few

chromatophores and is translucent ventrally (Fig. 2A).

Likewise, the labial tooth row formula 1/3+3//3 is com-

mon in the genus but the keratodonts are relatively long.

Further typical characters of L. boulengeri tadpoles are

small eyes (ED/BL= 0.08), a very high tail (including

fins) in comparison to its congeners despite a narrow tail

axis, and the presence of three rows of caudal papillae on

the lower lip (Fig. 2C), the latter character being unique

in the genus (compare Channing et al. 2012; Penske et al.

2015; Portillo and Greenbaum 2014b).

Leptopelis calcaratus (Boulenger, 1906)

The description of L. calcaratus tadpoles is based on

eleven tadpoles: ZMB 79618 (one tadpole at Gosner

stage 28) and ZMB 79619 (nine tadpoles at Gosner stag-

es 25 to 40), all on Mt. Nlonako, Cameroon, 4.9250°N;

9.98 17°E, 1,035 m a.s.l., 25 October 2011, the tadpoles

were found in a stream near a village) and ZMB 79620

(one tadpole at Gosner stage 41, near Manengouba vil-

lage, Mt. Manengouba, Cameroon, 4.9502°N; 9.8639°E,

1,116 m a.s.l., 23 November 2011, the tadpoles were

found in a stream near the village). Proportions including

total or tail length were only available for non-genotyped

individuals and ZMB 79620.

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December 2015 | Volume 9 | Number 2 | el 1 1

Barej et al.

Description. Body oval with nearly rounded snout in

dorsal view (Fig. 3B); ovoid to slightly compressed in

lateral view (Fig. 3A); tail length-body length ratio 2.27

(TL/BL); body height 0.43 of body length (BH/BL);

body width 0.54 of body length (BW/BL); maximum

body width on the level of the spiracle’s posterior end;

nostrils situated dorsally, closer to snout tip than eyes

(SND/SED = 0.38), distance snout-nostrils 0.16 of body

length (SND/BL); eyes positioned dorsolaterally; eye di-

ameter 0. 10 of body length (ED/BL); interocular distance

exceeds internostril distance by a factor of 2.56 (IOD/

IND); tail length 0.69 of entire length (TL/EL), with

moderately pronounced fins with narrow fin tip; dorsal

fin originates posterior to the dorsal tail-body junction

with maximum height at three-quarters of the tail length;

dorsal fin slightly curved; ventral fin originates on the

ventral terminus of the body; ventral fin narrower than

tail axis with maximum height at three-quarters of the

tail length; maximum fin height in dorsal fin higher (DF/

VF = 1 . 1 8); fin tip pointed; maximum tail height includ-

ing fins equals body height (TTH/BH = 1 .00); tail axis

width (in dorsal view) 0.50 of body width (AW/BW);

maximum height of tail axis (at base) 0.53 of body height

(AH/BH); tail axis height (at base) higher than maxi-

mum height of dorsal fin (AH/DF = 2.15); dextral vent

tube, positioned basicaudally; spiracle sinistral, visible

in dorsal view, originating anterior to mid-body (SSD/

BL = 0.43); spiracle tube length 0.17 of body length (SL/

BL); mouth opens anteroventrally; oral disc width less

than fifth of body width (ODW/BW = 0.19); one row of

papillae (with rounded tips) laterally at anterior lip with

huge rostral gap, these connected to papillae in labial

angles and posterior lip; second row of papillae caudal

at posterior lip with slightly pointed tips; labial tooth row

Fig. 4. Lateral (A) and dorsal (B) view of Leptopelis millsoni (ZMB 79621) at Gosner stage 39; coloration in life of tadpole (ZMB

79621) in lateral (top) and dorsal (below) view (C); sketch of the oral disc (D); adult L. millsoni (ZFMK 87708) (E); scale bars: 1

mm. Note that the greenish coloration on the lower fin results from a leaf used as the background.

Amphib. Reptile Conserv. 63 December 2015 | Volume 9 | Number 2 | el 11

The tadpoles of eight West and Central African Leptopelis species

formula 1/3+3//3 (Fig. 3E); jaw sheaths black, of equal

width and serrated; upper jaw very widely U-shaped with

median concavity; lower jaw widely V-shaped.

Coloration in preservation. Dorsolateral part of the

body, tail axis and dorsal fin mostly mottled brown on

yellowish ground; areas without brown spots shine

through as yellow blots; ventral part of the body pale yel-

low with some homogeneously distributed brown spots;

spiracle and vent tube translucent; ventral fin translucent

without any brown spots.

Coloration in life (Fig. 3C). Dark brown with shiny

golden speckles at dorsolateral part of the body, tail axis

and dorsal fin; ventral fin predominantly translucent with

few spots towards tail tip; ventral part of the body with-

out golden speckles.

Remarks. Leptopelis calcaratus is known from Nigeria

to Gabon and the Republic of the Congo in the south and

the Central African Republic and the Democratic Repub-

lic of the Congo to the east (e.g., de la Riva 1994; Schiptz

1963, 1999; Fretey and Blanc 2001; Fretey et al. 2006;

Jackson and Blackburn 2007; Amiet 2012). Reproduc-

tion takes place in more or less swampy forests that are

crossed by small rivers (Amiet 2012). Notes on habitat

use and call activity of this species were documented

by Schiptz (1967, 1999), Amiet and Schiptz (1974)

and Amiet (2006, 2012). The tadpole of L. calcaratus

has been described by Lamotte and Perret (1961) and

Channing et al. (2012). Shape of body and tail, as well

as overall pigmentation are congruent with the available

tadpole descriptions. In addition to the above recorded

labial tooth row formula Lamotte and Perret (1961) men-

tion 1/2+2//3. The eyes are positioned dorsolaterally in

our material, as described by Channing et al. (2012); in

contrast, Lamotte and Perret (1961) refer to a dorsal po-

sition; however, it cannot be excluded that their series

comprised material of different species (their descrip-

tions were usually based on morphological series and not

on tadpoles from known parents). The tail length-body

length ratio of 2.3 was higher in comparison to both for-

S' Lmuni II > M-J I il :

l*PJ|j,,n||!IL , ^ | >

Fig. 5. Lateral (A) and dorsal (B) view of Leptopelis modestus (ZMB 79622) at Gosner stage 34; adult L. modestus (MCZ A138023,

photo courtesy David C. Blackburn ) (C); sketch of the oral disc (D); habitat of L. modestus on Mt. Manengouba (E and F); scale

bars: 1 mm.

Amphib. Reptile Conserv.

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December 201 5 | Volume 9 | Number 2 | el 1 1

Barej et al.

mer descriptions (1.9). Examined tadpoles also differed

in coloration to the voucher examined by Channing et al.

(2012). While these authors note black pigments on tail

and fins, pigmentation in our material was mottled brown

on a pale ground or forming large, almost uniform brown

blotches, with a small translucent spiracle and vent tube

as described by Lamotte and Perret (1961). Pigmentation

tended to decrease from body to tail.

Leptopelis millsoni (Boulenger, 1895)

The description of L. millsoni tadpoles is based on one

tadpole: ZMB 79621 (at Gosner stage 39, the tadpole

was found in Kribi, near Miangasio Lendi, Cameroon,

2.8930°N; 9.9542°E, 31m a.s.l., 04 November 2011, in a

slow flowing, sandy bottom forest stream).

Description. Body oval with semi-circular snout in dor-

sal view (Fig. 4B); ovoid to slightly compressed in lateral

view (Fig. 4A); tail length-body length ratio 1.82 (TL /

BL); body height 0.38 of body length (BH/BL); body

width 0.58 of body length (BW/BL); maximum body

width on the level of the spiracle’s anterior end; nostrils

situated dorsally, slightly closer to snout tip than eyes

(SND/SED = 0.43), distance snout-nostrils 0.16 of body

length (SND/BL); eyes positioned laterally; eye diameter

0.12 of body length (ED/BL); interocular distance ex-

ceeds intemostril distance by a factor of 2.33 (IOD/IND);

tail length 0.65 of entire length (TL/EL), with moderately

pronounced fins with narrow fin tip; dorsal fin originates

at dorsal tail-body junction; dorsal fin moderately curved

with maximum height at two-thirds of the tail length;

ventral fin originates on the ventral terminus of the body;

ventral fin narrower than tail axis with maximum height

at half of the tail length; maximum fin height of dorsal

fin higher (DF/VF = 1 .25); fin tip pointed; maximum tail

height including fins exceeds body height (TTH/BH =

1.06); tail axis width (in dorsal view) 0.52 of body width

(AW/BW); maximum height of tail (axis at base) 0.69 of

body height (AH/BH); tail axis height at its base higher

than maximum height of dorsal fin (AH/DF = 2.50); dex-

tral vent tube, positioned basicaudally; spiracle sinistral,

visible in dorsal view, originating slightly anterior to

mid-body (SSD/BF = 0.47); spiracle tube length 0.11 of

body length (SF/BF); mouth opens anteroventrally; oral

disc width more than a third of body width (ODW/BW

= 0.36); one row of short papillae (with slightly pointed

tips) laterally at anterior lip with huge rostral gap, these

connected to papillae in labial angles and posterior lip;

second row of papillae at posterior lip; labial tooth row

formula 1/3+3//3 (Fig. 4D); jaw sheaths black, of equal

width and serrated; upper jaw widely U-shaped with me-

dian concavity; lower jaw widely V-shaped.

Coloration in preservation. Body, tail axis, dorsal fin

and ventral fin mostly speckled dark brown on yellowish

ground, areas without brown spots shine through as yel-

Amphib. Reptile Conserv.

low blots, ventral part of the body yellow with some light

brown spots; spiracle and vent tube in the same color as

body and tail.

Coloration in life (Fig. 4C). Dark brown with shiny

golden speckles at dorsolateral part of the body, tail axis,

dorsal fin and ventral fin; speckles very dense at dorsal

part of the body; dorsoventral part of the body with few

speckles.

Remarks. Leptopelis millsoni is known from Nigeria to

Gabon and the eastern Democratic Republic of the Con-

go (e.g., Schiptz 1967, 1999; Fotters et al. 2001; Blanc

and Fretey 2004; Rodel et al. 2014). As in the other spe-

cies male calling sites are found close to streams in the

breeding season but reproduction most probably occurs

in stagnant water (Amiet 2012). The call has been record-

ed by Amiet and Schiptz (1974) and call activity is de-

tailed in Amiet (2006). The tail with low fins is long (TF /

BE =1.8), but not as long as observed in other Leptopelis

species. Because we had only one tadpole available we

cannot check if this is a peculiarity of our specimen or a

general trend in this Gosner stage. What distinguishes L.

millsoni from the other studied tadpoles is the shape of

the papillae. While all other Leptopelis species showed

papillae with rounded tips, the papillae of L. millsoni had

fairly pointed tips (Fig. 4D). The eyes of our voucher

were relatively big compared to the other species (ED/

BE = 0.12); only L. viridis had similar sized eyes in rela-

tion to body length. We cannot evaluate whether the TF/

BE value reflects a species specific state, an individual

character state or the advanced Gosner stage.

Leptopelis modestus (Werner, 1 898)

The description of L. modestus tadpoles is based on three

tadpoles: ZMB 79622 (one tadpole, at Gosner stage 34),

ZMB 79623 (one tadpole, at Gosner stage 31), near sum-

mit of Mt. Manengouba, Cameroon, 5.0098°N; 9.8568°E,

2,135 m a.s.l., 27 September 2011, the tadpoles were

found in a medium sized river in a gallery forest) and

ZMB 79624 (one tadpole, at Gosner stage 36, Northwest

Province Abo Forest, Cameroon, 24 August 2012). Pro-

portions including total or tail length were only available

for the non-genotyped individual and ZMB 79624.

Description. Body oval with nearly rounded snout in

dorsal view (Fig. 5B); ovoid to slightly compressed in

lateral view (Fig. 5A); tail length-body length ratio 2.27

(TF/BF); body height 0.49 of body length (BH/BF);

body width 0.57 of body length (BW/BF); maximum

body width slightly behind the level of the spiracle’s

posterior end; nostrils situated dorsally, closer to snout

tip than eyes (SND/SED = 0.39), distance snout-nostrils

0.19 of body length (SND/BF); eyes positioned laterally;

eye diameter 0.09 of body length (ED/BF); interocu-

lar distance exceeds internostril distance by a factor of

65

December 2015 | Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

1.94 (IOD/IND); tail length 0.70 of entire length (TL /

EL), with moderately pronounced fins with narrow fin

tip; dorsal fin originates at dorsal tail-body junction ris-

ing barely at the first eighth of the tail length; dorsal fin

slightly curved with maximum height at half of the tail

length; ventral fin originates on the ventral terminus of

the body; ventral fin narrower than tail axis with maxi-

mum height around half of the tail length; maximum fin

height in dorsal fin higher (DF/VF = 1 .25); fin tip pointed;

maximum tail height including fins equals body height

(TTH/BH = 1.00); tail axis width (in dorsal view) 0.36 of

body width (AW/BW); maximum height of tail axis (at

base) 0.46 of body height (AH/BH); tail axis height at its

base higher than maximum height of dorsal fin ( AH/DF

= 1.55); dextral vent tube, positioned basicaudally; spira-

cle sinistral, visible in dorsal view, originating anterior to

mid-body (SSD/BL = 0.53); spiracle tube length 0.07 of

body length (SL/BL); mouth opens anteroventrally; oral

disc width more than a third of body width (ODW/BW

= 0.34); one row of papillae (with rounded tips) later-

ally at anterior lip with huge rostral gap, these connected

to papillae in labial angles and posterior lip; second row

of papillae at posterior lip, also with rounded tips; labial

tooth row formula 1/3+3//3 or 1/4+4//3 (Fig. 5D); jaw

sheaths black, of equal width and serrated; upper jaw and

lower jaw widely Fl-shaped.

Coloration in preservation. Dorsolateral part of the

body, tail axis and dorsal fin mostly speckled dark brown

on brownish ground on the body and yellowish ground

on the tail; areas without brown spots shine through as

yellow blots; ventral part of the body yellowish with

some homogeneously distributed dark brown spots at the

anterior third of the body; spiracle and vent tube translu-

cent; ventral fin at the anterior part translucent with some

brown spots towards tail tip.

Remarks. Since a record of Leptopelis modestus from

eastern Congo (Laurent 1972) and subsequent recogni-

tion as a distinct sub-species (Laurent 1973), L. modestus

has been regarded as a species with a disjunct distribu-

tion with known occurrences in Nigeria, Cameroon, and

Bioko - Equatorial Guinea (Schiptz 1967, 1999; Amiet

2012; Fretey et al. 2012) and the eastern Democratic Re-

public of the Congo and Kenya (Kohler et al. 2006; Por-

tillo and Greenbaum 2014b). However, the latter popula-

tions have been recently recognized as several distinct

species (Schiptz 1975: L.fiziensis from South Kivu Prov-

ince, DRC; Kohler et al. 2006: L. mackayi from the West-

ern Province, Kenya; Portillo and Greenbaum 2014b: L.

mtoewaate from South Kivu Province, DRC). Although

males congregate close to streams and torrents during the

Fig. 6. Lateral (A) and dorsal (B) view of Leptopelis rufus (ZMB 79627) at Gosner stage 36; adult L. rufus (female: ZMB 78398

and male: ZMB 78399) (E); sketch of the oral disc (D), scale bars: 1 mm.

Amphib. Reptile Conserv. 66 December 2015 | Volume 9 | Number 2 | el 11

Barej et al.

breeding season, reproduction takes place in slow run-

ning and stagnant water bodies (Amiet 2012). Further

notes on call activity and the advertisement call are pro-

vided by Schiptz (1999) and Amiet (2006). Based on two

vouchers the tadpole has been described by Channing et

al. (2012). Our observations are in agreement with their

description. Minor differences refer to coloration and the

interocular distance-internostril distance ratio and an ad-

ditional labial tooth row formula (1/4+4//3; Fig. 5D). The

IOD/IND was marginally lower (1.94) in comparison to

the value of 2 recorded by Channing et al. (2012). Note-

worthy, the tail length-body length ratio differed between

different Gosner stages (stage 31 : TL/BL= 2.1; stage 36:

TL/BL= 2.5). Concerning the coloration, the anterior

half of the ventral fin lacked speckles in Gosner stages

31 and 34 (Fig. 5 A) while it was pigmented in the more

developed tadpole (Gosner stage 36).

Taxonomic remark. Amiet (2012) discussed the possi-

bility of cryptic speciation based on a modestus- like fe-

male from Mwandong, West Cameroon, which differed

in coloration of skin and iris, size of tympanum, and

snout-vent length from remaining populations and co-

occurred with congeneric species (L. brevirostris, L. cal-

caratus, and L. modestus ). The herein investigated tad-

poles have been collected on Mt. Manengouba, in close

proximity to Mwandong, and in the Abo Forest. The bar-

coded sequences included a specimen (MCZ A138023;

Fig. 5C) collected near Nsoung on Mt. Manengouba.

MCZ A138023 exhibits characters that assign the speci-

men to the “true” L. modestus. Although the two geno-

typed tadpoles originate from high elevation localities on

Mt. Manengouba and Mt. Oku (both app. 2,150 m a.s.l.),

with a distance of more than 150 km between them, they

show no difference in the analysed 16S fragment and

point to the occurrence of the same taxon on both moun-

tain ranges.

Leptopelis rufus Reichenow, 1874

The description of L. rufus tadpoles is based on eighteen

tadpoles (remark: two different molecular lineages have

been recognized in L. rufus in the course of the present

analyses, thus we herein refer to L. rufus _1 and L. rufus _2

in order to assure differentiation of the examined mate-

rial): ZMB 79625 (L. rufus_ 1, three tadpoles, at Gosner

stages 26 and 29, Camp Bekop, Ebo Forest, Cameroon,

4.35 19°N; 10.4244°E, 845 m a.s.l., 07 January 2011, the

tadpoles were found in secondary forest), ZMB 79626

(L. rufus_ 2; two tadpoles, at Gosner stages 28 and 29,

Mt. Nlonako, Cameroon, 4.8309°N; 9.9255°E, 459 m

a.s.l., 23 October 2011, the tadpoles were found in a

small rock pool of approximately 50 cm diameter), ZMB

79627 (L. rufus _2; one tadpole, at Gosner stage 36, Nju-

ma, Ebo Forest, Cameroon, 4.3394°N; 10.2458°E, 320 m

a.s.l., 20 August 2011, the tadpole was found in primary

rainforest), ZMB 79628 ( L . rufus_ 2; one tadpole, at Gos-

ner stage 29, Ndogbanguengue, Ebo Forest, Cameroon,

4.4069°N; 10.1653°E, 96 m a.s.l., 19 September 2010,

the tadpole was found in farmbush) and ZMB 79629

(L. rufus_ 2; seven tadpoles, at Gosner stages 28 to 36,

Ekom Khan, Mt. Manengouba, Cameroon, 5.0633°N;

10.0163°E, 587 m a.s.l., 29 December 2010, the tadpoles

were found in a medium sized river in a forest fragment).

Proportions including total or tail length were only avail-

able for non-genotyped individuals.

Description. Body oval with nearly rounded snout in

dorsal view (Fig. 6B); ovoid to slightly compressed

in lateral view (Fig. 6A); tail length-body length ratio

2.04 (TL/BL); body height 0.37 of body length (BH/

BL); body width 0.53 of body length (BW/BL); maxi-

mum body width between the level of the eyes and the

spiracle’s anterior end; nostrils situated dorsally, closer

to snout tip than eyes (SND/SED = 0.40), distance snout-

nostrils 0.20 of body length (SND/BL); eyes positioned

laterally; eye diameter 0.10 of body length (ED/BL); in-

terocular distance exceeds internostril distance by a fac-

tor of 1.78 (IOD/IND); tail length 0.67 of entire length

(TL/EL), with moderately pronounced fins with narrow

fin tip; dorsal fin originates at dorsal tail-body junction,

but very low, not visible in lateral view; rising behind

anterior sixth of tail length; dorsal fin moderately curved

with maximum height at three-quarters of the tail length;

ventral fin originates on the ventral terminus of the body;

ventral fin narrower than tail axis with maximum height

at three-quarters of the tail length; maximum tin height

higher in dorsal tin (DF/VF = 1 . 1 8); fin tip pointed; maxi-

mum tail height including tins nearly equals body height

(TTH/BH= 0.98); tail axis width (in dorsal view) 0.36 of

body width (AW/BW); maximum height of tail axis (at

base) 0.65 of body height (AH/BH); tail axis height at

its base higher than maximum height of dorsal fin (AH/

DF = 1.75); dextral vent tube, positioned basicaudally;

spiracle sinistral, visible in dorsal view, originating at

mid-body (SSD/BL = 0.50); spiracle tube length 0.13 of

body length (SL/BL); mouth opens anteroventrally; oral

disc width more than a third of body width (ODW/BW

= 0.36); one row of papillae (with rounded tips) laterally

at anterior lip with huge rostral gap, these connected to

papillae in labial angles and posterior lip; second row of

longer papillae caudal at posterior lip, also with rounded

tips; labial tooth row formula 1/3+3//3 or 1/4+4//3 (Fig.

6D); jaw sheaths black and serrated, upper jaw sheath

thicker; upper jaw widely U-shaped with median concav-

ity; lower jaw widely V-shaped.

Coloration in preservation. Dorsolateral part of the

body, tail axis and dorsal fin mostly speckled dark brown

on brownish ground at the body and yellowish ground

at the tail; areas without brown spots shine through as

small yellow blots; ventral part of the body yellowish

with many homogeneously distributed dark brown spots

at the anterior third of the body and fewer spots at the

December 2015 I Volume 9 | Number 2 | el 1 1

Amphib. Reptile Conserv.

67

The tadpoles of eight West and Central African Leptopelis species

Fig. 7. Lateral (A) and dorsal (B) view of Leptopelis spiritusnoctis (ZMB 79634) at Gosner stage 34; adult L. spiritusnoctis (ZMB

79578) (C); sketch of the oral disc (D), scale bars: 1 mm.

posterior two-thirds of the body; spiracle and vent tube

translucent; ventral fin at the anterior part translucent

with some brown spots towards tail tip.

Remarks. Leptopelis rufus is known from Nigeria to

northern Angola (de la Riva 1994; Schiptz 1963, 1999;

Amiet 2012). Adults are common on branches and lia-

nas in proximity to streams during the breeding season

(Amiet 1975). The call has been reported by Amiet and

Schiptz (1974). The tadpole of has been described by

Channing et al. (2012) based on a single tadpole belong-

ing to a larger series examined herein (MH399 = ZMB

79629; herein assigned to L. rufus _2). Generally our ob-

servations of the larger series coincide with the former

description. However, while early tadpole stages of L.

rufus exhibit the labial tooth row formula 1/3+3//3, also

reported in Channing et al. (2012), we observed an in-

crease of tooth rows on the upper lip in more developed

tadpoles (Gosner stage 29: 1/4+4//3; Fig. 6D). Further

differences refer to a lower tail length-body length ratio

(TL/BL= 2.0) than in Channing et al. (2012; TL/BL =

2.6).

Taxonomic remark. A comparison of 16S sequences of

adults and tadpoles revealed two molecular lineages in L.

rufus , diverging by app. 1.5% in the mitochondrial 16S

gene (Tab. 2). Each lineage could be assigned to adult

specimens that have morphologically been assigned to L.

rufus. While no obvious differences have been assessed,

neither in tadpoles nor adults, we herein refer to L. ru-

fus _1 and L. rufus fl in order to highlight this molecular

divergence beyond intraspecific variance in remaining

species analysed herein. A similar genetic divergence

(0. 9-1.1% in 16S) has recently been uncovered between

two species in the eastern Democratic Republic of the

Congo (Portillo and Greenbaum 2014a) warranting fur-

ther morphological and bio-acoustical analyses to exam-

ine the status of lineages of L. rufus in western Central

Africa.

Leptopelis spiritusnoctis Rodel, 2007

The description of L. spiritusnoctis tadpoles is based

on twenty tadpoles: ZMB 79630 (five tadpoles, at Gos-

ner stages 25 to 40, 7.2347°N; 9.3096°E, 398 m a.s.l.),

ZMB 79631 (one tadpole, at Gosner stage 40, 7.2347°N;

9.3096°E, 398 m a.s.l.), ZMB 79632 (one tadpole, at

Gosner stage 31, 7.2316°N; 9.3118°E, 382 m a.s.l.),

ZMB 79633 (eight tadpoles, at Gosner stages 25 to 36,

7.2308°N; 9.3023°E, 387 m a.s.l.), ZMB 79634 (one

Amphib. Reptile Conserv.

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December 201 5 | Volume 9 | Number 2 | el 1 1

Barej et al.

tadpole, at Gosner stage 36, 7.2308°N; 9.3023°E, 387

m a.s.l.), ZMB 79635 (three tadpoles, at Gosner stages

25 and 27, 7.2376°N; 9.3117°E, 417 m a.s.l.), and ZMB

79636 (one tadpole, at Gosner stage 25, 7.2376°N;

9.3117°E, 417 m a.s.l.). All L. spiritusnoctis tadpoles

were caught near Gbanju, Liberia, 08 June 2011. Propor-

tions including total or tail length were only available for

non-genotyped individuals, ZMB 79630, 79632, 79634,

and 79636.

Description. Body oval with subovoid snout in dorsal

view (Fig. 7B); ovoid to slightly compressed in lateral

view (Fig. 7A); tail length-body length ratio 2.33 (TL /

BL); body height 0.49 of body length (BH/BL); body

width 0.60 of body length (BW/BL); maximum body

width on the level of the spiracle’s anterior end; nostrils

situated dorsally, closer to snout tip than eyes (SND/SED

= 0.37), distance snout-nostrils 0.21 of body length (SND/

BL); eyes positioned laterally; eye diameter 0.09 of body

length (ED/BL); interocular distance exceeds internostril

distance by a factor of 1.76 (IOD/IND); tail length 0.70

of entire length (TL/EL), with moderately pronounced

fins with narrow fin tip; dorsal fin originates at dorsal tail-

body junction; dorsal fin moderately curved with maxi-

mum height at three-quarters of the tail length; ventral

fin originates on the ventral terminus of the body; ventral

fin narrower than tail axis with maximum height at three-

quarters of the tail length; maximum fin height in dor-

sal fin higher (DF/VF = 1 .28); fin tip pointed; maximum

tail height including fins slightly exceeds body height

(TTH/BH = 1.08); tail axis width (in dorsal view) 0.41

of body width (AW/BW); maximum height of tail axis

(at base) 0.56 of body height (AH/BH); tail axis height at

its base higher than maximum height of dorsal fin (AH/

DF = 1.93); dextral vent tube, positioned basicaudally;

spiracle sinistral, visible in dorsal view, originating at

mid-body (SSD/BL = 0.50); spiracle tube length 0.12 of

body length (SL/BL); mouth opens anteroventrally; oral

disc width more than a quarter of body width (ODW/B W

= 0.30); one row of papillae (with rounded tips) laterally

at anterior lip with huge rostral gap, these connected to

papillae in labial angles and posterior lip; second row of

papillae (also with rounded tips) at posterior lip; labial

tooth row formula 1/4+4//3 (Fig. 7D); jaw sheaths black,

of equal width and serrated; upper jaw widely U-shaped;

lower jaw U-shaped.

Coloration in preservation. Dorsolateral part of the

body mostly speckled dark brown on yellowish ground,

tail axis, dorsal fin and spiracle speckled with less brown

spots on yellowish ground; ventral part of the body yel-

low with some homogeneously distributed brown spots

Fig. 8. Lateral (A) and dorsal (B) view of Leptopelis viridis (ZMB 79638) at Gosner stage 40; adult L. viridis (ZMB 83028) (C);

sketch of the oral disc (ZMB 79637) at Gosner stage 30 (D), scale bars: 1 mm.

Amphib. Reptile Conserv.

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December 201 5 | Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

at the anterior third of the body; vent tube translucent;

ventral fin translucent at anterior part with some brown

spots towards tail tip.

Remarks. Leptopelis spiritusnoctis is known from the

entire West African forest belt ranging from Guinea,

through Sierra Leone, Liberia, Cote d’Ivoire, Ghana,

Togo, Benin to western Nigeria (e.g., Schiptz 1963, 1967;

Rodel et al. 2000, 2004; Hillers and Rodel 2007; Rodel

2007; Segniagbeto et al. 2007). Male calling sites have

been reported from close to various water bodies, from

fast flowing creeks with rocky bed to tiniest puddles on

the forest floor (Rodel 2007). Females deposit up to 140

eggs below the soil surface (Schiptz 1963; Rodel 2007).

After three weeks tadpoles hatch and wriggle up to 50

cm towards the water (Schiptz 1963; Oldham 1971). The

tadpole was described by Lamotte and Perret (1961),

Schiptz (1963, 1967), Rodel (2007), and Channing et al.

(2012). Prior to the description of L. spiritusnoctis by

Rodel (2007) records of the species, including tadpole

descriptions, have been named L. hyloides. Generally

the observations of our larger series agree with former

descriptions. However, an additional labial tooth row

formula has been encountered 1/4+4//3 (Fig. 7D). La-

motte and Perret (1961) reported a change of the number

of tooth rows during tadpole growth. The observed tail

length-body length ratio was marginally higher (TL/BL=

2.3) than in the previous descriptions of Channing et al.

(2012: TL/BL= 2.2) and Lamotte and Perret (1961: TL /

BL= 2). The position of nostrils was closer to the snout

tip than to the eyes while they are closer to the eye ac-

cording to Channing et al. (2012).

Taxonomic remark. Amiet (2012) assumed the West

African L. spiritusnoctis and the Central African L. au-

bryi to be conspecific. However, based on genetics and

bioacoustics Rodel et al. (2014) recently confirmed their

specific distinctness. This is herein further supported by

tadpole morphology, as tadpoles of the two species dif-

fered in their size (tadpoles of L. aubryi growing larg-

er 53 mm; Schiptz 1963), tail length-body length ratio

(higher in in L. aubryi ; TL/BL = 3.4x) and labial tooth

row formulae 1/3+3//3 in L. aubryi ; diverse in L. spiri-

tusnoctis ; see above).

Leptopelis viridis (Gunther, 1869)

The description of L. viridis tadpoles is based on two

tadpoles: ZMB 79637 (one tadpole, at Gosner stage 30)

and ZMB 79638 at (one tadpole, at Gosner stage 40).

Both tadpoles were caught near Banambala, Guinea,

7.9899°N; 9.1312°E, 449 m a.s.l., 01 June 2011. Propor-

tions including total or tail length for this species were

not available, because there were only two individuals to

examine, both with incomplete tail as tail tips were used

for DNA analysis.

Description. Body oval with subelliptical snout in dorsal

view (Fig. 8B); ovoid to slightly compressed in lateral

view (Fig. 8A); body height 0.50 of body length (BH/

BF); body width 0.58 of body length (BW/BF); maxi-

mum body width on the level of the spiracle’s posterior

end; nostrils situated dorsally, closer to snout tip than

eyes (SND/SED = 0.35), distance snout-nostrils 0.20 of

body length (SND/BF); eyes positioned laterally; eye di-

ameter 0.12 of body length (ED/BF); interocular distance

exceeds intemostril distance by a factor of 1.92 (IOD/

IND); tail with moderately pronounced fins; dorsal fin

originates at dorsal tail-body junction; dorsal fin nearly

parallel; ventral fin originates on the ventral terminus of

the body; ventral fin narrower than tail axis and parallel

to it; maximum fin height in dorsal fin higher (DF/VF =

1.60); maximum tail height including fins equals body

height (TTH/BH= 1.00) at the level, where the tail was

cut; tail axis width (in dorsal view) 0.49 of body width

(AW/BW); maximum height of tail axis (at base) 0.61 of

body height (AH/BH); tail axis height at its base higher

than maximum height of dorsal fin (AH/DF = 2. 19); dex-

tral vent tube, positioned basicaudally; spiracle sinistral,

visible in dorsal view, originating at mid-body (SSD/BF

= 0.53); spiracle tube length 0.10 of body length (SF/

BF); mouth opens anteroventrally; oral disc width about

a quarter of body width (ODW/BW = 0.24); one row of

papillae (with rounded tips) laterally at anterior lip with

huge rostral gap, these connected to papillae in labial an-

gles and posterior lip; second row of papillae (also with

rounded tips) at posterior lip; labial tooth row formula

1/2+2//1+1/2 or 1/2+2//2+2/1 (Fig. 8D); jaw sheaths

black and serrated, upper jaw sheath broader than lower

jaw sheath; upper jaw widely U-shaped; lower jaw wide-

ly U-shaped as well.

Coloration in preservation. Dorsolateral part of the

body mostly speckled dark brown on brownish ground;

tail axis with less brown spots on yellow ground; ventral

part of the body yellowish with some brown spots at the

anterior third of the body; spiracle and vent tube trans-

lucent; ventral fin predominantly translucent with few

brown spots composed of dense melanophores, dorsal fin

brownish with some dark brown spots particularly at the

anterior part of the tail.

Remarks. Leptopelis viridis covers a wide geographic

range from Senegal to Nigeria and the north-eastern

Democratic Republic of the Congo (e.g., Perret 1966;

Schiptz 1963, 1967, 1999; Rodel 2000; Amiet 2012).

Females produce up to 220 eggs of 3. 1-4.7 mm that are

rich in yolk and of yellowish-white color (Barbault 1984;

Rodel 2000). Rodel (2000) assumed egg deposition in

rock-pools or transport of tadpoles by adults as the el-

evated surrounding was rocky and did not make digging

of burrows possible. The tadpole of Leptopelis viridis has

already been described in the past (Famotte and Perret

Amphib. Reptile Conserv.

70

December 2015 | Volume 9 | Number 2 | el 1 1

Barej et al.

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Fig. 9. Biplot of the second and third components of a Principal Component Analyses of morphological measures of Leptopelis

tadpoles (A). Illustrations are of genotyped representatives (not necessarily included in the PC A) roughly to scale. Boxplots show

morphometric ratios of variables contributing most to these components (B-F).

Amphib. Reptile Conserv.

71

December 201 5 I Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

1961; Schiptz 1963, 1967; Rodel 2000; Channing et al.

2012). Both of our tadpole vouchers had a cut tail, thus

we can only refer to formerly reported tail length-body

length ratios (Lamotte and Perret 1961: TL/BL = 2.1;

Rodel 2000: TL/BL = 2.5; Channing et al. (2012): TL/BL

= 2). In comparison to other Leptopelis species, the tooth

rows are very variable. Rodel (2000) mentioned the la-

bial tooth row formula of 1//2 for a tadpole two days after

hatching and various formulae are known in more devel-

oped tadpoles: 1/2+2//3 (Lamotte and Perret 1961; Rodel

2000; Channing et al. 2012), 1/2+2// 1+1/2 (Lamotte and

Perret 1961; Channing et al. 2012; this study, in Gosner

stage 40 in ZMB 79638), 1/3+3//3 and 1/3+3//1+1/2

(both Lamotte and Perret 1961) and 1/2+2//2+2/1 (this

study, in Gosner stage 30 in ZMB 79637; Fig. 8D). A

dark pigmentation of dorsal parts of the body has already

been reported in the past and is more conspicuous than in

other known Leptopelis tadpoles. While the spiracle was

translucent in ZMB 79637, a condition also reported by

Lamotte and Perret (1961) and Channing et al. (2012),

it contained some chromatophores in ZMB 79638 (Fig.

8 A). Lamotte and Perret (1961) reported the presence

of pigmentation on the fins, ventral body parts and ab-

sence of chromatophores at the intestinal region, which

could be confirmed herein. Likewise the presence of low

and nearly parallel fins of similar height (Channing et

al. 2012) and large eyes (Lamotte and Perret 1961) is in

agreement with our observations (ED/BL = 0.12).

Comparative Morphometries and Habitat

Morphometric patterns in Leptopelis tadpoles were com-

pared to investigate whether species occupy different ar-

eas of morpho-space. This was achieved by subjecting

log ]Q -transformed body measurements to a rigid rotation

(Principal Component Anlaysis; PC A) and by compar-

ing morphometric ratios based on measurements that are

contributing most to PCI and PC2. The first component

of the PCA was largely dominated by overall size dif-

ferences (likely also influenced by differences in Gos-

ner stages), but the second and third components could

clearly separate species into distinct morphological clus-

ters (Fig. 9A). PC2 is loaded negatively by AW, ED, and

IOD, and positively by ODW, VF, and DF. This means

that species clusters with negative PC2 values (L. calca-

ratus, L. aubryioides ) have wider, more muscular tails,

bigger eyes and wider interocular distances, compared

to clusters with positive PC2 values (L. boulengeri, L.

rufus, and L. modestus ), which have wider oral discs and

deeper tail fins. Leptopelis millsoni and L. spiritusnoctis

are intermediate for these traits (PC2 values close to 0).

Leptopelis rufus, L. modestus, and L. boulengeri show

strongly overlapping values for these traits, but L. bou-

lengeri is distinct from the other two, by having a nar-

rower internarial distance (similar to L. calcaratus), the

main loading of PC3. The relevant ratios (AH/DF, AW/

BW, OWD/BW, IOD/IND, ED/BL; Fig. 9B-F) reiterate

these patterns and in addition, show that L. rufus_\ tad-

poles have similar body proportions to L. rufus _2 and

that L. viridis is most similar to L. aubryioides in mor-

phology, with possibly a wider tail muscle, more similar

to L. calcaratus.

It should be noted however, that tadpole morphol-

ogy, especially tail shape, can be plastic in response to

extrinsic conditions (Duellman and Trueb 1994; Lau-

rila and Kujasalo 1999; Relyea 2001; Kraft et al. 2006;

Wells 2007) and due to limited sampling, morphologi-

cal variation due to differences in Gosner stage could not

be investigated. Nonetheless, the eight tadpoles includ-

ed in the analyses occur in differing microhabitats that

can roughly be grouped into temporary ponds, marshes

or slow running to stationary parts of streams (L. viri-

dis, L. spiritusnoctis, L. modestus, and L. aubryioides),

versus faster flowing running streams (L. calcaratus, L.

millsoni, L. rufus, and L. boulengeri). Differences in fea-

tures, such as the hydrodynamics of the tail shape, may

thus be experiencing diverging selective pressures across

these differing habitats (Altig and McDiarmid 1999b).

Greater sampling and more empirical data on microhabi-

tat of these tadpoles is needed however, to thoroughly

test whether such morphological differences are indeed

correlated to environmental parameters or a result of phe-

notypic plasticity or development.

Concluding Summary of Morphological

Characters

On a continental scale, and taking into account the lat-

est taxonomic decisions (Gvozdik et al. 2014; Portillo

and Greenbaum 2014a), tadpoles of only 25 of the 53

recognized Leptopelis species have been described. This

is astonishing as most species are abundant during the

breeding season.

Generally, tadpoles in the genus Leptopelis are mor-

phologically conservative and can be unambiguously as-

signed to that genus directly in the field. They possess

either the labial tooth row formula 1/3+3//3 or 1/2+2//3.

Only L. gramineus has strongly divergent formulae

(LTRF: 1/4+4//4, 1/4+4//1+1/2), and the first anterior

tooth row may sometimes be interrupted (Channing et al.

2012). Future studies on these tadpoles should consider

a potential ontogenetic change as increase of tooth rows

has been reported in L. aubryioides (this study), L. cal-

caratus (Lamotte and Perret 1961), and L. viridis (Rodel

2000 ).

West and western Central African regions experi-

enced an increase in herpetological surveys and subse-

quent taxonomic works in the last decades. But despite

this positive development and the present descriptions of

eight Leptopelis tadpoles, detailed accounts of the larval

morphology for ten western African congeners are still

missing: West Africa: Leptopelis bufonides, L. macrotis.

Amphib. Reptile Conserv. 72 December 2015 | Volume 9 | Number 2 | el 1 1

Barej et al.

L. occidentalis; western Central Africa: Leptopelis boca-

gii, L. brevirostris, L. bufonides, L. christyi, L. crystal-

linoron, L. palmatus, and L. zebra.

Among the eight herein described tadpoles, a super-

ficial similarity is conspicuous. However, preliminary

analyses not only reveal their morphological distinctness

but tentatively indicate morphological adaptations to the

respective habitat (lentic or lotic). Two species were un-

derrepresented (L. millsoni, L. modestus ) or even missing

completely (L. viridis ) in the analysis.

Acknowledgments. — We thank all respective authori-

ties from Cameroon, Guinea, and Liberia for research,

access, collection, and export permits as well as our many

guides and field assistants for their courageous help.

Thomas M. Doherty-Bone (Leeds) kindly provided com-

parative material from Abo Forest, Mt. Oku. Fieldwork

of MH was supported by scholarships from the Federal

State of Berlin (Elsa-Neumann-Stipendium) and the Ger-

man Academic Exchange Service (DAAD). MD’s field

work was supported by the German Herpetological Soci-

ety (DGHT, Wilhelm-Peters-Fonds). Simon Loader sup-

ported, and secured funding for, the field- and lab-work

of HCL (Swiss National Science Foundation: 3 1003 A-

133067). Barcoding of tadpoles was financially support-

ed by the Forderverein des Museums fur Naturkunde,

Berlin and the Swiss National Science Foundation.

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Michael F. Barej is interested in the diversity of African amphibians and reptiles, their biology, biogeographic

7" patterns, as well as resulting puzzles in taxonomy and systematics. He received his Ph.D. from the Humboldt

University Berlin (Germany) for unravelling patterns and systematics of African toiTent frogs.

Amphib. Reptile Conserv.

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December 2015 | Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

Tilo Pfalzgraff graduated with a B.S. in Biology from the Freie Universitat Berlin, Germany. He worked for the

Museum fur Naturkunde Berlin studying Central African tadpoles, and is currently a M.S. student of aquacul-

ture and sea ranching in the Faculty of Agricultural and Environmental Sciences at the University of Rostock,

Germany. He conducts surveys for alternative fish nutrition and is interested in ichthyology, herpetology, bio-

diversity, and sustainable development.

Mareike Hirschfeld studied Biological Sciences at the Universities of Marburg and Wurzburg, Germany and

received her Diplom (M.Sc. equivalent) in 2009. Currently she is a scientist and Ph.D. candidate at the Museum

fur Naturkunde in Berlin. She is interested in amphibian ecology and the taxonomy of West and Central African

anurans.

H. Christoph Liedtke is a zoologist currently based at the Estacion Biologica de Donana (Seville, Spain). His

main interests are comparative phylogenetics and life history evolution and his Ph.D. thesis, undertaken at the

University of Basel (Switzerland), was centered on understanding the life history and species diversity of Afri-

can amphibians, particularly the evolution of terrestrial modes of reproduction in anurans.

Johannes Penner received his Diploma from the University of Wurzburg and Ph.D. from the Humboldt Uni-

versity Berlin (both in Germany). Engaged in various herpetological projects for the last 14 years, he is cur-

rently located at the Museum fur Naturkunde Berlin (Germany) and focuses on African amphibians and reptiles

with a geographic emphasis on West Africa. In addition he works for a large European Biodiversity project

(EU BON) and as a freelance consultant in West Africa. He is also engaged in various IUCN specialist groups,

mainly the IUCN Viper Specialist Group.

Nono L. Gonwouo, as a conservation biologist, is interested in the responses of biodiversity to environmen-

tal change. His research to date has focused on the effects of anthropogenic activities on the herpetofauna of

Cameroon, with particular interest in the distributions and dynamics of endemic species near their geographic

range margins. His interest as well involves systematics and taxonomy of reptiles and amphibians, community

ecology, ethology, biology of vertebrates, and environmental impact assessment studies.

Matthias Dahmen studied Applied Biogeography at Trier University. He undertook research on degradation

tolerances of Central African rainforest anurans and their life-cycle strategies. Thereby he focused on species

of lower altitude range and conducted his fieldwork in the Ebo Forest in Cameroon. Meanwhile, he is work-

ing as a landscape planner and thereby, specialized on the detection, legal consideration, and conservation of

amphibians and reptiles.

Franziska Grozinger studied biology at the Universities of Heidelberg and Wurzburg, Germany. She com-

pleted her Ph.D. thesis at the Museum fur Naturkunde in 2014, where she focused on the phenotypic plasticity

of the European Common Frog (Rana temporaria). She is an experienced field biologist, interested in the inter-

action between organisms and their environment.

Andreas Schmitz received both his Diploma and his Ph.D. at the University of Bonn in Germany. He has

worked in herpetology for over 20 years, first at the Zoological Research Museum in Bonn (ZFMK), Germany

and since 2003 he is the leading Research Officer of Herpetology at the Natural History Museum of Geneva

(MHNG), Geneva, Switzerland. He works on the systematics, taxonomy, phylogenetics, and conservation of a

multitude of amphibian and reptile groups mostly in Africa and Southeast Asia.

Mark-Oliver Rodel is the Curator of Herpetology and head of the department of “Diversity Dynamics” at the

Museum fur Naturkunde, Berlin. Since his teens he has studied amphibians and reptiles, mostly those from

Europe and Africa. With his team he investigates the taxonomy, systematics, ecology, and biogeography of

amphibians and reptiles, but in particular uses amphibians as model organisms in order to understand the effect

of environmental change on species and ecosystems.

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Barej et al.

Figure Appendix 1.

A

Fig. Al. Schematic tadpole in dorsal (A), lateral (B) view and sketch of the mouth part in ventral view (C) showing assessed dis-

tances and mouth parts. Abbreviations: G - dorsal gap; A1-A3 - anterior papillae; L - lateral papillae; P1-P3 - posterior papillae;

LJS - lower jaw sheath; UJS - upper jaw sheath; for abbreviations of measurements see material and methods.

Amphib. Reptile Conserv.

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December 201 5 I Volume 9 I Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

Table Appendix 1.

Table Al. Collection numbers (Museum flir Naturkunde, Berlin, ZMB; Zoologisches Forschungsmuseum Alexander Koenig, Bonn,

ZFMK), localities of Leptopelis tadpoles studied herein, and GenBank data analysed in our 16S DNA-barcoding analysis; n = num-

ber of tadpoles (a single one genotyped, see Appendix Tables A2 and A3).

Species

Collection

number

Stage

n

Country

Region

Site

Latitude

Longi-

tude

Elevation

[m a.s.l.]

GenBank

number

Reference

aubryioides

ZFMK

81604

adult

—

Cameroon

foot of Mt.

Nlonako

near

Ekomtolo

4.8397°N

9.9303°E

470

KT967076

this study

aubryioides

ZMB

79604

tadpole

2

Cameroon

Etome

near

Etome

4.8317°N

9.9253°E

476

KT967077

this study

aubryioides

ZMB

79605

tadpole

1

Cameroon

foot of Mt.

Nlonako

near

Ekomtolo

4.8329°N

9.9259°E

477

KT967078

this study

aubryioides

ZMB

79606

tadpole

9

Cameroon

foot of Mt

Nlonako

near

Ekomtolo

4.8329°N

9.9259°E

477

—

aubryioides

ZMB

79607

tadpole

3

Cameroon

Ebo Forest

Njuma

4.3483°N

10.2329°E

238

KT967079

this study

aubryioides

ZMB

79608

tadpole

1

Cameroon

Ebo Forest

Njuma

4.3483°N

10.2329°E

238

KT967080

this study

aubryioides

ZMB

79609

tadpole

1

Cameroon

Ebo Forest

Njuma

4.3394°N

10.2458°E

320

KT967081

this study

aubryioides

ZMB

79610

tadpole

1

Cameroon

Ebo Forest

Njuma

4.3483°N

10.2329°E

238

KT967082

this study

aubryioides

ZMB

79611

tadpole

1

Cameroon

Ebo Forest

Njuma

4.3483°N

10.2329°E

238

KT967083

this study

aubryioides

ZMB

79612

tadpole

1

Cameroon

Ebo Forest

Camp

Njuma

4.3480°N

10.2323°E

315

KT967084

this study

boulengeri

ZFMK

87860

adult

—

Cameroon

—

Amebishu

6.1239°N

9.6875°E

165

KT967085

this study

boulengeri

ZMB

79613

tadpole

1

Cameroon

Ebo Forest

Bekob

4.3578°N

10.4170°E

921

KT967086

this study

boulengeri

ZMB

79614

tadpole

4

Cameroon

Ebo Forest

Bekob

4.3578°N

10.4170°E

921

KT967087

this study

boulengeri

ZMB

79615

tadpole

3

Cameroon

Ebo Forest

Bekob

4.3575°N

10.4168°E

903

KT967088

this study

boulengeri

ZMB

79616

tadpole

1

Cameroon

Ebo Forest

Bekob

4.3578°N

10.4170°E

921

KT967089

this study

boulengeri

ZMB

79617

tadpole

7

Cameroon

Ebo Forest

Bekob

4.3578°N

10.4170°E

921

—

calcaratus

ZFMK

75509

adult

—

Cameroon

Mt

Nlonako

Nguengue

4.9172°N

9.9892°E

1140

KT967090

this study

calcaratus

ZMB

79618

tadpole

1

Cameroon

Mt

Nlonako

—

4.9250°N

9.9817°E

1035

KT967091

this study

calcaratus

ZMB

79619

tadpole

9

Cameroon

Mt

Nlonako

—

4.9250°N

9.9817°E

1035

—

calcaratus

ZMB

79620

tadpole

1

Cameroon

Mt

Manen-

gouba

Manen-

gouba

village

4.9502°N

9.8639°E

1116

KT967092

this study

millsoni

ZFMK

87708

adult

—

Cameroon

—

near

Nkoelon

2.3972°N

10.0352°E

75

KF888342

Rodel et al.

(2014)

millsoni

ZMB

79621

tadpole

1

Cameroon

Kribi

near

Miangasio

Lendi

2.8930°N

9.9542°E

31

KT967093

this study

modestus

MCZ

A138023

adult

—

Cameroon

Mt.

Manen-

gouba

Nsoung

4.98 14°N

9.8133°E

1346

JQ715683

Blackburn

(2008b)

modestus

ZMB

79622

tadpole

1

Cameroon

Mt

Manen-

gouba

near

summit

5.0098°N

9.8568°E

2135

KT967094

this study

modestus

ZMB

79623

tadpole

1

Cameroon

Mt

Manen-

gouba

near

summit

5.0098°N

9.8568°E

2135

—

modestus

ZMB

79624

tadpole

1

Cameroon

North

West

Province

Abo Forest

6.2857°N

10.3580°E

2162

KT967095

this study

Amphib. Reptile Conserv.

78

December 201 5 I Volume 9 | Number 2 | el 1 1

Barej et al.

Table A1 (continued). Collection numbers (Museum flir Naturkunde, Berlin, ZMB; Zoologisches Forschungsmuseum Alexander

Koenig, Bonn, ZFMK), localities of Leptopelis tadpoles studied herein, and GenBank data analysed in our 16S DNA-barcoding

analysis; n = number of tadpoles (a single one genotyped, see Appendix Tables A2 and A3).

Species

Collection

number

Stage

n

Country

Region

Site

Latitude

Longi-

tude

Elevation

[m a.s.l.]

GenBank

number

Reference

rufus_ 1

ZFMK

87897

adult

—

Cameroon

—

near

Nkoelon

2.3972°N

10.0352°E

75

KT967096

this study

rufus_\

ZMB

79625

tadpole

3

Cameroon

Camp

Bekop

Ebo Forest

4.3519°N

10.4244°E

845

KT967097

this study

rufusjl

ZFMK

67382

adult

—

Cameroon

Bakossi

Mts.

Kodmin

4.9833°N

9.7000°E

1065

KT967098

this study

rufusjl

ZMB

79626

tadpole

2

Cameroon

Mt

Nlonako

—

4.8309°N

9.9255°E

459

KT967099

this study

rufusjl

ZMB

79627

tadpole

1

Cameroon

Ebo Forest

Njuma

4.3394°N

10.2458°E

320

KT967100

this study

rufusjl

ZMB

79628

tadpole

5

Cameroon

Ebo Forest

Ndog-

banguen-

gue

4.4069°N

10.1653°E

96

KT967101

this study

rufusjl

ZMB

79629

tadpole

7

Cameroon

Mt

Manen-

gouba

Ekom

Khan

5.0633°N

10.0163°E

587

KT967102

this study

spiritusnoctis

ZMB

79582

adult

—

Liberia

—

near

Jarwodee

5.4938°N

8.3636°W

220

KF888336

Rodel et al.

(2014)

spiritusnoctis

ZMB

79630

tadpole

5

Liberia

—

near

Gbanju

7.2347°N

9.3096°W

398

KT967103

this study

spiritusnoctis

ZMB

79631

tadpole

1

Liberia

—

near

Gbanju

7.2347°N

9.3096°W

398

KT967104

this study

spiritusnoctis

ZMB

79632

tadpole

1

Liberia

—

near

Gbanju

7.23 16°N

9.3118°W

382

KT967105

this study

spiritusnoctis

ZMB

79633

tadpole

8

Liberia

—

near

Gbanju

7.2308°N

9.3023°W

387

KT967106

this study

spiritusnoctis

ZMB

79634

tadpole

1

Liberia

—

near

Gbanju

7.2308°N

9.3023°W

387

KT967107

this study

spiritusnoctis

ZMB

79635

tadpole

3

Liberia

—

near

Gbanju

7.2376°N

9.3117°W

417

KT967108

this study

spiritusnoctis

ZMB

79636

tadpole

1

Liberia

—

near

Gbanju

7.2376°N

9.3117°W

417

KT967109

this study

viridis

ZMB

83027

adult

—

Liberia

—

near

Gbanju

7.3242°N

9.3035°W

380

KT967110

this study

viridis

ZMB

79637

tadpole

1

Guinea

—

near

Banam-

bala

7.9899°N

9.1312°W

449

KT967 111

this study

viridis

ZMB

79638

tadpole

1

Guinea

—

near

Banam-

bala

7.9899°N

9.1312°W

449

KT967112

this study

Amphib. Reptile Conserv.

79

December 201 5 I Volume 9 I Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

Table A2. Morphometries of Leptopelis tadpoles; G = Gosner stage; measurements in mm; genotyped specimens are marked with an

asterisk genotyped and drawn specimens are marked with two asterisks for abbreviations see Materials and Methods.

species

ZMB#

G

BL

TL

EL

BW

BH

AH

VF

DF

TTH

AW

IOD

IND

SND

SED

ED

SSD

ODW

SL

aubryioides

79604*

30

9

-

5.3

3.8

2

0.5

-

2.2

3.5

1.8

1

2.4

0.9

5

1.3

0.7

aubryioides

79604

36

10.4

-

6.5

5.1

2.5

1

1.3

4.3

2.5

3.8

2

1

2.5

1

4.5

1.1

0.9

aubryioides

79605**

25

5.5

-

3.2

2.4

1.5

0.4

0.6

1.9

1.3

2.1

1.3

0.7

1.5

0.5

2.2

0.8

0.9

aubryioides

79606

40

10.3

23.8

34.1

6.3

5

2.3

1

1.4

4.2

2.6

3.8

2

1.1

2.6

1

4.6

1.1

0.9

aubryioides

79606

27

7.4

18.7

26.1

4.1

3.4

2

0.6

0.7

2.9

1.8

3.1

1.6

0.8

2.2

1

3.3

1.2

1

aubryioides

79606

25

7.1

16.9

24

4.2

3.3

1.9

0.7

0.8

3

1.9

3

1.5

0.8

2.3

1.1

3.6

1.1

1.2

aubryioides

79606

27

7.9

18.9

26.8

4.6

3.5

2

0.6

0.9

3.1

1.7

2.9

1.6

0.8

2.1

1

3.4

1.1

1

aubryioides

79606

27

7.7

-

4.8

3.8

2.1

0.7

1

3.4

1.8

2.8

1.5

0.8

2.2

1.1

3.5

1.2

1.3

aubryioides

79606

28

8.2

19.5

27.7

5

4.1

2

0.8

1.1

3.5

1.9

3

1.5

0.8

2.2

1.1

3.8

1.1

1.5

aubryioides

79606

37

10.4

23.6

34

5.5

4.5

2.4

1

1.3

4.2

2.4

3.9

2

1.1

2.6

1.2

4.7

1.4

1.7

aubryioides

79606

37

10.3

23.8

34.1

5.8

4.6

2.5

1.1

1.4

4.5

2.6

4.1

2.2

1.1

2.5

1.1

4.6

1.5

aubryioides

79606

37

10.5

24.4

34.9

6.2

4.5

2.7

1

1.4

4.5

2.5

3.9

2

1.1

2.6

1.2

4.7

1.4

1.8

aubryioides

79607*

36

10.7

-

6.1

5

2.7

1.1

1.3

3.8

2.6

3.8

2.1

1.4

2.9

1

4.5

1.6

1.7

aubryioides

79607

39

11.6

26.3

37.9

6.6

4.8

2.9

1.2

1.6

5.1

2.9

4.3

2.3

1.2

2.7

1.2

4.9

1.5

1.8

aubryioides

79607

36

10

24.8

34.8

6

4.4

2.6

1.1

1.3

4.5

2.5

4

2.1

1.1

2.5

1.1

4.3

1.4

1.7

aubryioides

79608*

40

10.6

-

5.5

4.5

2.4

1

1.3

2.9

2.4

3.9

2

1.1

2.6

1.2

4.7

1.4

1.7

aubryioides

79609*

31

8.8

-

5.2

3.2

2.2

0.9

1

2.9

2.3

3.3

1.7

1

2.3

0.8

4.2

1.3

1.1

aubryioides

79610*

36

9.8

-

5.5

4.4

2.5

1

1.3

3

2.3

3.8

1.8

1

2.4

1

4.7

1.2

aubryioides

79611*

41

10

-

5.4

3.9

2.3

0.9

1

3

2.3

4.1

1.9

1.2

2.6

1.3

4.4

1.3

0.7

aubryioides

79612*

34

9.4

-

5.2

3.9

2.3

0.9

1.1

2.8

2.1

3.3

1.6

0.9

2

0.9

4.1

1.3

boulengeri

79613*

37

10.9

-

5.5

3.5

2.2

1

1.2

4.4

1.8

3.5

1.5

1.1

2.9

0.9

5

1.9

1.5

boulengeri

79614*

37

11.5

-

7

5.6

2.9

1.5

1.7

6.1

2.3

4

1.6

1.1

2.8

1

5.5

2.1

1.7

boulengeri

79614

40

12.3

-

6.3

5

2.9

2

2.3

7.2

2.2

4.3

1.9

1.5

3.3

1

4.9

2.3

2.2

boulengeri

79614

40

12.2

34.7

46.9

5.9

4.9

2.8

1.8

2

6.6

2.4

4.1

1.9

1.4

3.2

1

5

2.4

boulengeri

79614

36

11.7

28.9

40.6

5.8

4.7

2.6

1.7

1.9

6.2

2.1

4

1.8

1.3

3.1

1

4.5

2.1

boulengeri

79615*

36

10.3

-

5.7

4.5

2.4

1

1.2

4.6

2

3.5

1.5

1.2

3

1

4.6

2

boulengeri

79615

36

9.3

20.9

30.2

5.1

4.3

2.3

1

1.1

4.4

1.7

3

1.3

1

2.4

0.7

4.2

1.8

1.7

boulengeri

79615

36

9.9

23.8

33.7

5.2

4.4

2.2

1.2

1.4

4.8

1.6

3.1

1.4

1

2.3

0.7

4.3

1.9

1.8

boulengeri

79616**

38

11.6

-

7.1

5.7

2.8

1.6

1.7

6.1

2.4

4

1.6

1.1

2.8

1

5.1

2.2

2

boulengeri

79617

40

10.8

24.7

35.5

5.7

4.6

2.5

1.3

1.5

5.3

1.9

3.7

1.5

1.2

2.7

0.8

4.4

2.2

2

boulengeri

79617

40

11.5

30.1

41.6

5.9

4.8

2.6

1.5

1.6

5.7

2

3.8

1.7

1.3

3.1

1

4.8

2

2.1

boulengeri

79617

36

10

24.2

34.2

5.3

4.5

2.4

1.5

1.7

5.6

1.8

3.5

1.6

1.2

2.8

0.8

4.7

2.3

2

boulengeri

79617

40

11.7

28.8

40.5

6

4.9

2.7

1.6

1.7

6

1.9

3.8

1.5

1

2.9

1.1

4.3

2.1

2.2

boulengeri

79617

38

10.8

25.8

36.6

5.7

4.5

2.4

1

1.2

4.6

2

3.7

1.5

1.2

3

1

4.6

2

boulengeri

79617

40

11.2

27.1

38.3

5.8

4.7

2.8

1.7

1.8

6.3

2.1

3.8

1.6

1.3

3.1

1

4.8

2.1

2.3

boulengeri

79617

40

11.8

30.6

42.4

6.2

4.8

2.5

1.8

1.9

6.2

2

4

1.6

1.2

3

1

4.5

2.1

2.3

calcaratus

79618**

28

8.8

-

4.5

3.3

1.8

0.3

-

2.2

1.7

3.2

1.3

0.7

2.3

0.8

4.9

1.4

0.7

calcaratus

79619

27

9.7

18.7

28.4

5.3

4

2.2

0.9

1.2

4.3

2.4

4

1.5

1

2.7

1

4.5

0.9

1.9

calcaratus

79619

29

9.2

22.4

31.6

5.2

4.1

2.3

0.9

1

4.2

2.3

3.8

1.4

0.9

2.6

1

4.3

0.9

2

calcaratus

79619

25

6.5

15.5

22

4

3.2

1.5

0.7

0.8

3

1.6

3

1.2

0.6

2

0.7

2.9

0.7

1.2

calcaratus

79619

40

11.8

26.8

38.6

5.8

4.5

2.5

1

1.1

4.6

3.2

4.6

1.8

1.1

2.7

1.1

4.6

0.9

2.1

calcaratus

79619

40

12

28.2

40.2

6

4.8

2.6

1

1.2

4.8

3.3

4.7

1.9

1.2

2.8

1.2

4.8

1

2.3

calcaratus

79619

40

11.6

26.4

38

5.9

4.7

2.4

1.1

1.2

4.7

3.2

4.6

1.7

1.2

2.9

1.1

4.3

0.9

1.7

calcaratus

79619

25

8

18.7

26.7

4.6

4.2

2

0.8

1

3.8

2.2

3.6

1.4

0.8

2.2

0.9

4.4

0.8

1.5

calcaratus

79619

36

11.4

24.9

36.3

5.6

5

2.6

1.1

1.3

5

3.5

4.5

1.8

1.3

3

1

4.2

0.9

2

calcaratus

79619

38

10.8

24

34.8

5.7

4.5

2.4

1

1.2

4.6

3.1

4.2

1.5

1.2

2.8

1.1

4.1

0.9

1.8

calcaratus

79620*

41

13.2

30.2

43.4

8

6

3.7

1

1.2

5.9

3.9

5.5

2.5

1.3

3.3

1.4

5

2.1

NA

millsoni

79621**

39

9.5

17.3

26.8

5

3.6

2.5

0.8

1

3.8

2.6

3.5

1.5

1

2.3

1.1

4.5

1.8

1

modestus

79622**

34

11.3

-

6.5

6

2.5

1.5

1.8

5.8

2.3

4.3

2.2

1.2

2.8

1

6.2

2.5

0.7

Amphib. Reptile Conserv.

80

December 201 5 | Volume 9 | Number 2 | el 1 1

Barej et al.

Table A2 (continued). Morphometries of Leptopelis tadpoles; G = Gosner stage; measurements in mm; genotyped specimens are marked

with an asterisk genotyped and drawn specimens are marked with two asterisks for abbreviations see Materials and Methods.

species

ZMB#

G

BL

TL

EL

BW

BH

AH

VF

DF

TTH

AW

IOD

IND

SND

SED

ED

SSD

ODW

SL

modestus

79623

31

7.9

16.8

24.7

4.3

3.3

1.4

0.9

1.1

3.4

1.3

2.8

1.5

0.9

2.5

0.6

4.7

1.5

0.5

modestus

79624 *

36

14.1

35.2

49.3

8.3

7.5

4

1.5

2

7.5

3.5

5

2.5

1.1

2.8

1.3

6.5

2.4

1

rufus_l

79625 *

26

6.4

-

3.7

2.9

1.7

0.8

0.9

2.9

1.5

2.5

1.4

0.7

2.1

0.6

3.3

1.3

0.8

rufus_\

79625

29

7

-

3.7

2.3

1.6

0.9

1

2.2

1.2

2.7

1.6

0.8

2

0.7

3.7

1.5

0.9

rufus_\

79625

29

7.4

-

3.6

2.2

1.7

1

1.1

2.1

1.1

2.6

1.5

0.9

2.1

0.8

3.5

1.4

0.9

rufus_2

79626 *

29

7.2

-

3.7

2.3

1.5

0.3

-

1.2

2.8

1.7

0.7

2

0.7

3.7

1.5

0.9

rufus_2

79626

28

5.7

-

2.7

2

1.2

0.6

0.8

2

1.1

2.1

1.3

0.8

1.8

0.5

3.1

1.1

0.7

rufus_2

79627 **

36

10.2

-

5.6

4

2.4

1.1

1.3

3.9

2.3

3.5

1.5

1

2.6

1

5.3

1.8

1.6

rufus_2

79628 *

29

8.8

-

4.7

3.5

2

0.9

1

3.4

1.9

3

1.8

0.9

2.4

0.8

4.3

1.6

1.2

rufus_2

79628

36

8.6

16.4

25

4.9

3.1

1.9

0.9

1.1

3

1.4

2.9

1.6

0.9

2

0.8

3.9

1.6

1

rufus_2

79628

32

7.6

15.5

23.1

3.7

2.4

1.9

0.9

1.2

2.4

1.2

2.7

1.6

1

2.1

0.8

3.6

1.4

0.9

rufus_2

79628

31

7.1

14.4

21.5

3.7

2.3

1.5

0.7

0.9

2.2

1.2

2.8

1.7

0.7

2

0.7

3.7

1.5

0.9

rufus_2

79628

40

8.8

14.7

23.5

4.7

3.5

2

0.9

1

3.5

1.9

3

1.8

0.9

2.4

0.8

4.3

1.6

1.2

rufus_2

79629 *

36

12.6

-

8

5.8

3.3

1.7

2

5.6

3.4

4.5

2.4

1.2

3.3

1.1

6.4

2.5

1.5

rufus_2

79629

28

6.7

14.2

20.9

3.7

2.9

1.7

0.8

0.9

2.8

1.5

2.5

1.4

0.7

2.1

0.6

3.3

1.3

0.8

rufus_2

79629

29

7.7

15.1

22.8

3.8

2.4

1.8

1

1.2

2.4

1.2

2.7

1.6

1

2.1

0.8

3.6

1.4

0.9

rufus_2

79629

29

7.5

15.6

23.1

3.6

2.2

1.8

1

1.1

2.1

1.1

2.6

1.5

0.9

2.1

0.8

3.5

1.4

0.9

rufus_2

79629

35

10.2

22.7

32.9

5.6

4

2.4

1.1

1.3

4

2.3

3.5

1.9

1

2.6

1

5.3

1.8

1.6

rufus_2

79629

31

8.5

20.5

29

4.8

3

2

0.9

1

2.9

1.5

3

1.6

0.9

2

0.8

3.9

1.6

1

rufus_2

79629

34

9.9

22

31.9

5.5

3.9

2.3

1

1.2

3.9

2.2

3.4

1.8

1

2.5

1

5.1

1.8

1.5

spiritusnoctis

79630 *

-

9.5

18.8

28.3

4.8

4

-

1

1.3

-

2.5

3.1

1.9

1

2.6

0.8

4.3

1.6

1.2

spiritusnoctis

79630

25

4.4

11.1

15.5

3.4

2.7

1.2

0.4

0.6

2.2

1

1.8

1

0.5

1.3

0.4

2.4

0.8

0.5

spiritusnoctis

79630

29

7.8

19.8

27.6

4.5

3.4

1.9

1

3.9

1.6

2.9

1.3

0.8

2.1

0.7

3.9

1.3

1

spiritusnoctis

79630

36

10.7

25.8

36.5

5.5

4.3

3

1

1.6

5.6

3

3.6

2

1

3

1

4.8

1.9

1.5

spiritusnoctis

79630

40

10.5

26.1

36.6

5.4

4.1

2.9

1

1.5

5.4

2.9

3.4

1.9

1

2.9

1

4.6

1.8

1.4

spiritusnoctis

79631 *

40

13.3

-

7.4

6.3

4

1.4

1.7

7.1

3.8

5

2.7

1.2

2.7

1.4

6.4

2.1

1.5

spiritusnoctis

79632 *

31

12.5

26.8

39.3

7.2

5.2

3.7

1.4

1.9

7

4

2.4

1.5

3.9

1.2

6.5

2.1

1.5

spiritusnoctis

79633 *

25

6.6

-

4

3.3

1.7

0.8

1

3.5

1.4

2.3

1.5

0.7

2

0.6

3.6

1.2

0.7

spiritusnoctis

79633

25

5

12.1

17.1

2.8

2.7

1.3

0.5

0.7

2.5

1.1

1.9

1.1

0.5

1.4

0.4

2.6

0.9

0.6

spiritusnoctis

79633

25

4.8

11.6

16.4

3

2.9

1.1

0.5

0.7

2.3

1.1

1.9

1.1

0.5

1.4

0.4

2.6

0.9

0.6

spiritusnoctis

79633

36

8.7

22.8

31.5

4.7

3.6

2.1

1.2

4.5

1.8

3.1

1.6

1

2.3

0.9

4.2

1.3

spiritusnoctis

79633

27

7.2

17.7

24.9

4.1

3.5

1.9

0.7

0.8

3.4

1.8

2.4

1.5

0.7

2

0.6

3.2

1.2

0.7

spiritusnoctis

79633

26

5.9

-

4

3.3

1.7

0.8

1

3.5

1.4

2.3

1.5

0.7

2

0.6

3.6

1.2

0.7

spiritusnoctis

79633

26

6.2

14.2

20.4

4.1

3.2

1.8

0.8

1

3.6

1.5

2.5

1.4

0.7

2

0.6

3.8

1.3

0.8

spiritusnoctis

79633

30

7.6

17.9

25.5

4.4

3.3

1.9

0.9

1

3.8

1.6

2.8

1.3

0.7

2.1

0.7

3.9

1.3

1

spiritusnoctis

79634 **

34

11.5

26.1

37.6

6.3

4.5

3.2

1.1

1.7

6

3.1

3.8

2.2

1.1

3.1

1.1

5.3

2

1.6

spiritusnoctis

79635 *

27

6.3

-

3.8

3.2

1.5

0.7

0.8

3

1.3

2.2

1.3

0.6

1.7

0.5

2.9

1

0.7

spiritusnoctis

79635

25

5.5

12.2

17.7

3.6

3

1.4

0.6

0.7

2.7

1.2

2

1.2

0.6

1.5

0.5

2.7

1

0.6

spiritusnoctis

79635

25

4.9

10.7

15.6

3.5

2.8

1.3

0.5

0.7

2.5

1.1

1.9

1.1

0.5

1.4

0.4

2.6

0.9

0.6

spiritusnoctis

79636 *

25

6.8

17.1

23.9

3.8

3.5

1.9

0.7

0.8

3.4

1.8

2.4

1.5

0.7

2

0.6

3.2

1.2

0.7

viridis

79637 *

30

9.8

-

5.8

5.3

3.4

-

4.1

2.4

3.5

2.1

1

2.7

1.1

5.4

1.4

0.9

viridis

79638 **

40

13.5

-

7.7

6.1

3.5

1

1.6

6

4.4

5.2

2.4

1.1

3.3

1.7

7

1.8

1.4

Amphib. Reptile Conserv.

81

December 201 5 I Volume 9 I Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

Table A3. Ratios of Leptopelis tadpoles; G = Gosner stage; measurements in mm; genotyped specimens are marked with an asterisk

genotyped and drawn specimens are marked with two asterisks for abbreviations see Materials and Methods.

Species

ZMB#

BL/

BH /

BW/

SND /

ED/

IOD/

TL/

DF/

AH/

TTH/

AW/

AH/

SL/

ODW/

SSD /

TL

BL

SED

BL

IND

EL

VF

DF

BH

BW

BH

BL

BW

BL

aubryioides

79604 *

-

0.42

0.59

0.42

0.10

1.94

-

0.42

0.53

0.08

0.25

0.56

aubryioides

79604

-

0.49

0.63

0.40

0.10

1.90

-

1.30

1.92

0.84

0.38

0.49

0.09

0.17

0.43

aubryioides

79605 **

-

0.44

0.58

0.47

0.09

1.62

-

1.50

2.50

0.79

0.41

0.63

0.16

0.25

0.40

aubryioides

79606

0.43

0.49

0.61

0.42

0.10

1.90

0.70

1.40

1.64

0.84

0.41

0.46

0.09

0.17

0.45

aubryioides

79606

0.40

0.46

0.55

0.36

0.14

1.94

0.72

1.17

2.86

0.85

0.44

0.59

0.14

0.29

0.45

aubryioides

79606

0.42

0.46

0.59

0.35

0.15

2.00

0.70

1.14

2.38

0.91

0.45

0.58

0.17

0.26

0.51

aubryioides

79606

0.42

0.44

0.58

0.38

0.13

1.81

0.71

1.50

2.22

0.89

0.37

0.57

0.13

0.24

0.43

aubryioides

79606

-

0.49

0.62

0.36

0.14

1.87

-

1.43

2.10

0.89

0.38

0.55

0.17

0.25

0.45

aubryioides

79606

0.42

0.50

0.61

0.36

0.13

2.00

0.70

1.38

1.82

0.85

0.38

0.49

0.18

0.22

0.46

aubryioides

79606

0.44

0.43

0.53

0.42

0.12

1.95

0.69

1.30

1.85

0.93

0.44

0.53

0.16

0.25

0.45

aubryioides

79606

0.43

0.45

0.56

0.44

0.11

1.86

0.70

1.27

1.79

0.98

0.45

0.54

0.15

0.26

0.45

aubryioides

79606

0.43

0.59

0.42

0.11

1.95

0.70

1.40

1.93

1.00

0.40

0.60

0.17

0.23

0.45

aubryioides

79607 *

-

0.47

0.57

0.48

0.09

1.81

-

1.18

2.08

0.76

0.43

0.54

0.16

0.26

0.42

aubryioides

79607

0.44

0.41

0.57

0.44

0.10

1.87

0.69

1.33

1.81

1.06

0.44

0.60

0.16

0.23

0.42

aubryioides

79607

0.40

0.44

0.60

0.44

0.11

1.90

0.71

1.18

2.00

1.02

0.42

0.59

0.17

0.23

0.43

aubryioides

79608 *

-

0.42

0.52

0.42

0.11

1.95

-

1.30

1.85

0.64

0.44

0.53

0.16

0.25

0.44

aubryioides

79609 *

-

0.36

0.59

0.43

0.09

1.94

-

1.11

2.20

0.91

0.44

0.69

0.13

0.25

0.48

aubryioides

79610 *

-

0.45

0.56

0.42

0.10

2.11

-

1.30

1.92

0.68

0.42

0.57

0.12

0.22

0.48

aubryioides

79611 *

-

0.39

0.54

0.46

0.13

2.16

-

1.11

2.30

0.77

0.43

0.59

0.07

0.24

0.44

aubryioides

79612 *

-

0.41

0.55

0.45

0.10

2.06

-

1.22

2.09

0.72

0.40

0.59

0.14

0.25

0.44

boulengeri

79613 *

-

0.32

0.50

0.38

0.08

2.33

-

1.20

1.83

1.26

0.33

0.63

0.14

0.35

0.46

boulengeri

79614 *

-

0.49

0.61

0.39

0.09

2.50

-

1.13

1.71

1.09

0.33

0.52

0.15

0.30

0.48

boulengeri

79614

-

0.41

0.51

0.45

0.08

2.26

-

1.15

1.26

1.44

0.35

0.58

0.18

0.37

0.40

boulengeri

79614

0.35

0.40

0.48

0.44

0.08

2.16

0.74

1.11

1.40

1.35

0.41

0.57

0.20

0.41

boulengeri

79614

0.40

0.50

0.42

0.09

2.22

0.71

1.12

1.37

1.32

0.36

0.55

0.18

0.36

0.38

boulengeri

79615 *

-

0.44

0.55

0.40

0.10

2.33

-

1.20

2.00

1.02

0.35

0.53

0.19

0.35

0.45

boulengeri

79615

0.44

0.46

0.55

0.42

0.08

2.31

0.69

1.10

2.09

1.02

0.33

0.53

0.18

0.35

0.45

boulengeri

79615

0.42

0.44

0.53

0.43

0.07

2.21

0.71

1.17

1.57

1.09

0.31

0.50

0.18

0.37

0.43

boulengeri

79616 **

-

0.49

0.61

0.39

0.09

2.50

-

1.06

1.65

1.07

0.34

0.49

0.17

0.31

0.44

boulengeri

79617

0.44

0.43

0.53

0.44

0.07

2.47

0.70

1.15

1.67

1.15

0.33

0.54

0.19

0.39

0.41

boulengeri

79617

0.38

0.42

0.51

0.42

0.09

2.24

0.72

1.07

1.63

1.19

0.34

0.54

0.18

0.34

0.42

boulengeri

79617

0.41

0.45

0.53

0.43

0.08

2.19

0.71

1.13

1.41

1.24

0.34

0.53

0.20

0.43

0.47

boulengeri

79617

0.41

0.42

0.51

0.34

0.09

2.53

0.71

1.06

1.59

1.22

0.32

0.55

0.19

0.35

0.37

boulengeri

79617

0.42

0.53

0.40

0.09

2.47

0.70

1.20

2.00

1.02

0.35

0.53

0.19

0.35

0.43

boulengeri

79617

0.41

0.42

0.52

0.42

0.09

2.38

0.71

1.06

1.56

1.34

0.36

0.60

0.21

0.36

0.43

boulengeri

79617

0.39

0.41

0.53

0.40

0.08

2.50

0.72

1.06

1.32

1.29

0.32

0.52

0.19

0.34

0.38

calcaratus

79618 **

-

0.38

0.51

0.30

0.09

2.46

-

0.67

0.38

0.55

0.08

0.31

0.56

calcaratus

79619

0.52

0.41

0.55

0.37

0.10

2.67

0.66

1.33

1.83

1.08

0.45

0.55

0.20

0.17

0.46

calcaratus

79619

0.41

0.45

0.57

0.35

0.11

2.71

0.71

1.11

2.30

1.02

0.44

0.56

0.22

0.17

0.47

calcaratus

79619

0.42

0.49

0.62

0.30

0.11

2.50

1.29

1.14

1.88

0.94

0.40

0.47

0.18

0.45

calcaratus

79619

0.44

0.38

0.49

0.41

0.09

2.56

0.69

1.10

2.27

1.02

0.55

0.56

0.18

0.16

0.39

calcaratus

79619

0.43

0.40

0.50

0.43

0.10

2.47

0.70

1.20

2.17

1.00

0.55

0.54

0.19

0.17

0.40

calcaratus

79619

0.44

0.41

0.51

0.41

0.09

2.71

0.69

1.09

2.00

1.00

0.54

0.51

0.15

0.37

calcaratus

79619

0.43

0.53

0.58

0.36

0.11

2.57

0.70

1.25

2.00

0.90

0.48

0.19

0.17

0.55

calcaratus

79619

0.46

0.44

0.49

0.43

0.09

2.50

0.69

1.18

2.00

1.00

0.63

0.52

0.18

0.16

0.37

calcaratus

79619

0.45

0.42

0.53

0.43

0.10

2.80

0.69

1.20

2.00

1.02

0.54

0.53

0.17

0.16

0.38

calcaratus

79620 *

0.44

0.45

0.61

0.39

0.11

2.20

0.70

1.20

3.08

0.98

0.49

0.62

-

0.26

0.38

millsoni

79621 **

0.55

0.38

0.53

0.43

0.12

2.33

0.65

1.25

2.50

1.06

0.52

0.69

0.11

0.36

0.47

modestus

79622 **

-

0.53

0.58

0.43

0.09

1.95

-

1.20

1.39

0.97

0.35

0.42

0.06

0.38

0.55

Amphib. Reptile Conserv.

82

December 201 5 | Volume 9 | Number 2 | el 1 1

Barej et al.

Table A3 (continued). Ratios of Leptopelis tadpoles; G = Gosner stage; measurements in mm; genotyped specimens are marked with an

asterisk genotyped and drawn specimens are marked with two asterisks for abbreviations see Materials and Methods.

Species

ZMB#

BL/

BH/

BW/

SND /

ED/

IOD/

TL/

DF/

AH/

TTH/

AW/

AH/

SL/

ODW/

SSD /

TL

BL

SED

BL

IND

EL

VF

DF

BH

BW

BH

BL

BW

BL

modestus

79623

0.47

0.42

0.54

0.36

0.08

1.87

0.68

1.22

1.27

1.03

0.30

0.42

0.06

0.35

0.59

modestus

79624 *

0.40

0.53

0.59

0.39

0.09

2.00

0.71

1.33

2.00

1.00

0.42

0.53

0.07

0.29

0.46

rufus_ 1

79625 *

-

0.45

0.58

0.33

0.09

1.79

-

1.13

1.89

1.00

0.41

0.59

0.13

0.35

0.52

rufus_ 1

79625

-

0.33

0.53

0.40

0.10

1.69

-

1.11

1.60

0.96

0.32

0.70

0.13

0.41

0.53

rujus_ 1

79625

-

0.30

0.49

0.43

0.11

1.73

-

1.10

1.55

0.95

0.31

0.77

0.12

0.39

0.47

rufus_2

79626 *

-

0.32

0.51

0.35

0.10

1.65

-

0.32

0.65

0.13

0.41

0.51

rufus_2

79626

-

0.35

0.47

0.44

0.09

1.62

-

1.33

1.50

1.00

0.41

0.60

0.12

0.41

0.54

rufus_2

79627 **

-

0.39

0.55

0.38

0.10

2.33

-

1.18

1.85

0.98

0.41

0.60

0.16

0.32

0.52

rufus_2

79628 *

-

0.40

0.53

0.38

0.09

1.67

-

1.11

2.00

0.97

0.40

0.57

0.14

0.34

0.49

rufus_2

79628

0.52

0.36

0.57

0.45

0.09

1.81

0.66

1.22

1.73

0.97

0.29

0.61

0.12

0.33

0.45

rufus_2

79628

0.49

0.32

0.49

0.48

0.11

1.69

0.67

1.33

1.58

1.00

0.32

0.79

0.12

0.38

0.47

rufus_2

79628

0.49

0.32

0.52

0.35

0.10

1.65

0.67

1.29

1.67

0.96

0.32

0.65

0.13

0.41

0.52

rufus_2

79628

0.60

0.40

0.53

0.38

0.09

1.67

0.63

1.11

2.00

1.00

0.40

0.57

0.14

0.34

0.49

rufus_2

79629 *

-

0.46

0.63

0.36

0.09

1.88

-

1.18

1.65

0.97

0.43

0.57

0.12

0.31

0.51

rufus_2

79629

0.47

0.43

0.55

0.33

0.09

1.79

0.68

1.13

1.89

0.97

0.41

0.59

0.12

0.35

0.49

rufus_2

79629

0.51

0.31

0.49

0.48

0.10

1.69

0.66

1.20

1.50

1.00

0.32

0.75

0.12

0.37

0.47

rufus_2

79629

0.48

0.29

0.48

0.43

0.11

1.73

0.68

1.10

1.64

0.95

0.31

0.82

0.12

0.39

0.47

rufus_2

79629

0.45

0.39

0.55

0.38

0.10

1.84

0.69

1.18

1.85

1.00

0.41

0.60

0.16

0.32

0.52

rufus_2

79629

0.41

0.35

0.56

0.45

0.09

1.88

0.71

1.11

2.00

0.97

0.31

0.67

0.12

0.33

0.46

rufus_2

79629

0.45

0.39

0.56

0.40

0.10

1.89

0.69

1.20

1.92

1.00

0.40

0.59

0.15

0.33

0.52

spiritusnoctis

79630 *

0.51

0.42

0.51

0.38

0.08

1.63

0.66

1.30

-

0.52

-

0.13

0.33

0.45

spiritusnoctis

79630

0.40

0.61

0.77

0.38

0.09

1.80

0.72

1.50

2.00

0.81

0.29

0.44

0.11

0.24

0.55

spiritusnoctis

79630

0.39

0.44

0.58

0.38

0.09

2.23

0.72

1.00

1.90

1.15

0.36

0.56

0.13

0.29

0.50

spiritusnoctis

79630

0.41

0.40

0.51

0.33

0.09

1.80

0.71

1.60

1.88

1.30

0.55

0.70

0.14

0.35

0.45

spiritusnoctis

79630

0.40

0.39

0.51

0.34

0.10

1.79

0.71

1.50

1.93

1.32

0.54

0.71

0.13

0.33

0.44

spiritusnoctis

79631 *

-

0.47

0.56

0.44

0.11

1.85

-

1.21

2.35

1.13

0.51

0.63

0.11

0.28

0.48

spiritusnoctis

79632 *

0.47

0.42

0.58

0.38

0.10

1.67

0.68

1.36

1.95

1.35

0.56

0.71

0.12

0.29

0.52

spiritusnoctis

79633 *

-

0.50

0.61

0.35

0.09

1.53

-

1.25

1.70

1.06

0.35

0.52

0.11

0.30

0.55

spiritusnoctis

79633

0.41

0.54

0.56

0.36

0.08

1.73

0.71

1.40

1.86

0.93

0.39

0.48

0.12

0.32

0.52

spiritusnoctis

79633

0.41

0.60

0.63

0.36

0.08

1.73

0.71

1.40

1.57

0.79

0.37

0.38

0.13

0.30

0.54

spiritusnoctis

79633

0.38

0.41

0.54

0.43

0.10

1.94

0.72

1.00

1.75

1.25

0.38

0.58

0.15

0.28

0.48

spiritusnoctis

79633

0.41

0.49

0.57

0.35

0.08

1.60

0.71

1.14

2.38

0.97

0.44

0.54

0.10

0.29

0.44

spiritusnoctis

79633

-

0.56

0.68

0.35

0.10

1.53

-

1.25

1.70

1.06

0.35

0.52

0.12

0.30

0.61

spiritusnoctis

79633

0.44

0.52

0.66

0.35

0.10

1.79

0.70

1.25

1.80

1.13

0.37

0.56

0.13

0.32

0.61

spiritusnoctis

79633

0.42

0.43

0.58

0.33

0.09

2.15

0.70

1.11

1.90

1.15

0.36

0.58

0.13

0.30

0.51

spiritusnoctis

79634 **

0.44

0.39

0.55

0.35

0.10

1.73

0.69

1.55

1.88

1.33

0.49

0.71

0.14

0.32

0.46

spiritusnoctis

79635 *

-

0.51

0.60

0.35

0.08

1.69

-

1.14

1.88

0.94

0.34

0.47

0.11

0.26

0.46

spiritusnoctis

79635

0.45

0.55

0.65

0.40

0.09

1.67

0.69

1.17

2.00

0.90

0.33

0.47

0.11

0.28

0.49

spiritusnoctis

79635

0.46

0.57

0.71

0.36

0.08

1.73

0.69

1.40

1.86

0.89

0.31

0.46

0.12

0.26

0.53

spiritusnoctis

79636 *

0.40

0.51

0.56

0.35

0.09

1.60

0.72

1.14

2.38

0.97

0.47

0.54

0.10

0.32

0.47

viridis

79637 *

-

0.54

0.59

0.37

0.11

1.67

-

0.77

0.41

0.64

0.09

0.24

0.55

viridis

79638 **

-

0.45

0.57

0.33

0.13

2.17

-

1.60

2.19

0.98

0.57

0.10

0.23

0.52

Amphib. Reptile Conserv.

83

December 201 5 I Volume 9 | Number 2 | el 1 1

The tadpoles of eight West and Central African Leptopelis species

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0.45

[ 0 . 40 - 0 . 56 ]

0.43

[ 0 . 37 - 0 . 48 ]

0.43

[ 0 . 37 - 0 . 56 ]

0.47

0.53

[ 0 . 46 - 0 . 59 ]

0.51

[ 0 . 47 - 0 . 53 ]

0.50

[ 0 . 45 - 0 . 54 ]

0.50

[ 0 . 45 - 0 . 54 ]

0.50

[ 0 . 44 - 0 . 61 ]

0.53

[ 0 . 52 - 0 . 55 ]

ODW/BW

0.24

[ 0 . 17 - 0 . 29 ]

0.36

[ 0 . 30 - 0 . 43 ]

0.19

[ 0 . 15 - 0 . 31 ]

0.36

0.34

[ 0 . 29 - 0 . 38 ]

0.38

[ 0 . 35 - 0 . 41 ]

0.36

[ 0 . 31 - 0 . 41 ]

0.36

[ 0 . 31 - 0 . 41 ]

0.30

[ 0 . 24 - 0 . 35 ]

0.24

[ 0 . 23 - 0 . 24 ]

SL/BL

0.14

[ 0 . 07 - 0 . 18 ]

0.18

[ 0 . 14 - 0 . 21 ]

0.17

[ 0 . 08 - 0 . 22 ]

ll'O

0.07

[ 0 . 06 - 0 . 07 ]

0.13

[ 0 . 12 - 0 . 13 ]

0.13

[ 0 . 12 - 0 . 16 ]

0.13

[ 0 . 12 - 0 . 16 ]

0.12

[ 0 . 10 - 0 . 15 ]

[ oro - 60 ' o ]

oro

AH/BH

0.56

[ 0 . 46 - 0 . 69 ]

0.55

[ 0 . 49 - 0 . 63 ]

0.53

[ 0 . 47 - 0 . 62 ]

690

0.46

[ 0 . 42 - 0 . 53 ]

0.68

[ 0 . 59 - 0 . 77 ]

0.64

[ 0 . 57 - 0 . 82 ]

0.65

[ 0 . 57 - 0 . 82 ]

0.56

[ 0 . 38 - 0 . 71 ]

0.61

[ 0 . 57 - 0 . 64 ]

AW/BW

0.42

[ 0 . 37 - 0 . 45 ]

0.34

[ 0 . 31 - 0 . 41 ]

0.50

[ 0 . 38 - 0 . 63 ]

0.52

0.36

[ 0 . 30 - 0 . 42 ]

0.35

[ 0 . 31 - 0 . 41 ]

0.36

[ 0 . 29 - 0 . 43 ]

0.36

[ 0 . 29 - 0 . 43 ]

0.41

[ 0 . 29 - 0 . 56 ]

0.49

[ 0 . 41 - 0 . 57 ]

TTH/BH

0.86

[ 0 . 64 - 1 . 06 ]

1.20

[ 1 . 02 - 1 . 44 ]

0.97

[ 0 . 67 - 1 . 08 ]

1.06

1.00

[ 0 . 97 - 1 . 03 ]

0.97

[ 0 . 95 - 1 . 00 ]

0.98

[ 0 . 95 - 1 . 00 ]

0.98

[ 0 . 95 - 1 . 00 ]

1.08

[ 0 . 79 - 1 . 35 ]

0.88

[ 0 . 77 - 0 . 98 ]

AH/DF

2.07

[ 1 . 64 - 2 . 86 ]

1.63

[ 1 . 26 - 2 . 09 ]

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2.50

1.55

[ 1 . 27 - 2 . 00 ]

1.68

[ 1 . 55 - 1 . 89 ]

1.77

[ 1 . 50 - 2 . 00 ]

1.75

[ 1 . 50 - 2 . 00 ]

1.93

[ 1 . 57 - 2 . 38 ]

61‘Z

DF/VF

1.29

[ 1 . 11 - 1 . 50 ]

1.12

[ 1 . 06 - 1 . 20 ]

1.18

[ 1 . 09 - 1 . 33 ]

1.25

[ 1 . 20 - 1 . 33 ]

[ eri - ori ]

111

1.19

[ 1 . 10 - 1 . 33 ]

1.18

[ 1 . 10 - 1 . 33 ]

1.28

[ 1 . 00 - 1 . 60 ]

1.60

TL/EL

0.70

[ 0 . 69 - 0 . 72 ]

0.71

[ 0 . 69 - 0 . 74 ]

0.75

[ 0 . 66 - 1 . 29 ]

0.65

0.70

[ 0 . 68 - 0 . 71 ]

1

0.67

[ 0 . 63 - 0 . 71 ]

0.67

[ 0 . 63 - 0 . 71 ]

0.70

[ 0 . 66 - 0 . 72 ]

1

I0D/IND

1.93

[ 1 . 62 - 2 . 16 ]

2.35

[ 2 . 16 - 2 . 53 ]

2.56

[ 2 . 20 - 2 . 80 ]

2.33

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1.74

[ 1 . 69 - 1 . 79 ]

1.78

[ 1 . 62 - 2 . 33 ]

1.78

[ 1 . 62 - 2 . 33 ]

1.76

[ 1 . 53 - 2 . 23 ]

1.92

[ 1 . 67 - 2 . 17 ]

ED/BL

0.11

[ 0 . 09 - 0 . 15 ]

[010-Z.00]

80'0

0.10

[ 0 . 09 - 0 . 11 ]

0.12

[ 60 ' 0 _ 80 ' 0 ]

600

0.10

[ 0 . 09 - 0 . 11 ]

0.10

[ 0 . 09 - 0 . 11 ]

0.10

[ 0 . 09 - 0 . 11 ]

0.09

[ 0 . 08 - 0 . 11 ]

0.12

[ 0 . 11 - 0 . 13 ]

SND/SED

0.42

[ 0 . 35 - 0 . 48 ]

0.41

[ 0 . 34 - 0 . 45 ]

0.38

[ 0 . 30 - 0 . 43 ]

0.43

0.39

[ 0 . 36 - 0 . 43 ]

0.39

[ 0 . 33 - 0 . 43 ]

0.40

[ 0 . 33 - 0 . 48 ]

0.40

[ 0 . 33 - 0 . 48 ]

0.37

[ 0 . 33 - 0 . 44 ]

0.35

[ 0 . 33 - 0 . 37 ]

BW/BL

0.58

[ 0 . 52 - 0 . 63 ]

0.53

[ 0 . 48 - 0 . 61 ]

0.54

[ 0 . 49 - 0 . 62 ]

0.53

0.57

[ 0 . 54 - 0 . 59 ]

0.53

[ 0 . 49 - 0 . 48 ]

0.53

[ 0 . 47 - 0 . 63 ]

0.53

[ 0 . 47 - 0 . 63 ]

0.60

[ 0 . 51 - 0 . 77 ]

0.58

[ 0 . 57 - 0 . 59 ]

BH/BL

0.44

[ 0 . 36 - 0 . 50 ]

0.43

[ 0 . 32 - 0 . 49 ]

0.43

[ 0 . 38 - 0 . 53 ]

0.38

0.49

[ 0 . 42 - 0 . 53 ]

0.36

[ 0 . 30 - 0 . 45 ]

0.37

[ 0 . 29 - 0 . 46 ]

0.37

[ 0 . 29 - 0 . 46 ]

0.49

[ 0 . 39 - 0 . 61 ]

0.50

[ 0 . 45 - 0 . 54 ]

BL/TL

0.42

[ 0 . 40 - 0 . 44 ]

0.41

[ 0 . 35 - 0 . 44 ]

0.44

[ 0 . 41 - 0 . 52 ]

0.55

0.44

[ 0 . 40 - 0 . 47 ]

1

0.49

[ 0 . 41 - 0 . 60 ]

0.49

[ 0 . 41 - 0 . 60 ]

0.43

[ 0 . 38 - 0 . 51 ]

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Amphib. Reptile Conserv.

84

December 201 5 I Volume 9 I Number 2 | el 1 1

Ecnomiohyla rabborum. Rabb’s Fringe-limbed Treefrog is one of the most significantly threatened amphibians in Central America.

This species is one of the most unusual anurans in the region because of its highly specialized reproductive mode, in which the eggs

are laid in water-containing tree cavities and are attached to the interior of the cavity just above the water line. Females depart the

tree cavity after oviposition, leaving the males to brood the eggs and the developing tadpoles, and parental care apparently extends

to feeding the tadpoles flecks of skin from the male’s body (AmphibiaWeb site: accessed 24 July 2014). Mendelson et al. (2008)

described this tree canopy treefrog from “montane cloudforest in the immediate vicinity of the town of El Valle de Anton” (Am-

phibiaWeb site: accessed 24 July 2014) in central Panama, at elevations from 900 to 1,150 m. This mode of reproduction is typical

of the members of the genus Ecnomiohyla, which now comprises 14 species (Batista et al. 2014) with a collective distribution

extending from southern Mexico to northwestern South America (Colombia and Ecuador). This treefrog appears to be one of the

many casualties of a sweep-through of Panama by the fungal pathogen Batrachochytrium dendrobatidis in 2006. The arrival of this

pathogen was anticipated by a team of amphibian biologists, who observed the disastrous effects of B. dendrobatidis on the popula-

tions of anurans in the El Valle de Anton region. Individuals of E. rabbororum were taken into captivity and housed at Zoo Atlanta,

but only a single male remains alive. We determined its EVS as 20, placing it at the upper end of the high vulnerability category,

and its IUCN status is Critically Endangered. Since the species is known to survive only in captivity, its IUCN status should be

considered as Extinct in the Wild. Additionally, since the animal now is known from a single male, its IUCN status should change

to Extinct once it dies. This individual is from the type locality. Photo by Brad Wilson.

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

1

Bothriechis guifarroi. This green palm-pitviper is known only from the type locality in the Refugio de Vida Silvestre Texiguat in

north-central Honduras, where it occurs in Premontane Wet Forest at elevations of 1,015 to 1,450 m. We calculated its EVS as 19,

placing it in the upper portion of the high vulnerability category, but its IUCN status has not been determined. Its EVS is the highest

for any snake in Central America. Molecular analysis of this species indicates that it is part of a clade containing the Lower Central

American taxa B. lateralis and B. nigroviridis . Two pattern phases are seen in juveniles, of which one resembles the juveniles of its

apparent closest relative, B. lateralis, which is distributed in the chain of mountains in the central portions of Costa Rica and western

Panama. This snake was named in honor of the Honduran environmental leader Mario Guifarro, who was slain by unknown assail-

ants while heading grassroots attempts to stop illegal logging in the indigenous Tawahka territory in the Mosquitia of eastern Hon-

duras. Don Mario was the guide on several herpetological expeditions undertaken in the Mosquitia by Wilson and co-researchers

during the last decade. This individual is from the type locality. Photo by Josiah H. Townsend.

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

2

DEDICATION

We are pleased to dedicate this contribution to our friend and colleague Louis W. Porras, for the many ways he has

supported our efforts to conserve the rich herpetodiversity of Mesoamerica. As editor, publisher, and contributor to

Conservation of Mesoamerican Amphibians and Reptiles (2010), he remained solidly behind this multi-year project. In

addition, his amazing skills as a copy-editor and knowledge of graphic design were extremely important in the produc-

tion of th q Amphibian & Reptile Conservation Special Mexico Issue, published in 2013. Most recently, he has become

the force behind the journal Mesoamerican Herpetology in which a number of our contributions have appeared. In

general, we continually find it worthwhile to seek his counsel on a broad range of matters relating to herpetology and

conservation. Most importantly, however, we consider it an honor to call him friend.

Porthidium porrasi. The White -tailed Hog-

nosed Pitviper is endemic to the region of

the Osa Peninsula of southwestern Costa

Rica, where it occurs in Lowland Moist

Forest at elevations from near sea level to

200 m. We assessed its EVS as 18, placing

it in the upper portion of the high vulnera-

bility category, and its IUCN status is Least

Concern. This individual is from Rincon,

province of Puntarenas. Photo by Alejan-

dro Solorzano.

Louis W. Porras photographed on 19 April

2014 with a pair of Mormon Racers ( Colu-

ber mormon) in the Lake Shore Mountains

in Utah County, Utah. Louis said the fol-

lowing: “Eve been monitoring a den in

these mountains for about 25 years. In the

spring I often hike up there with my grand-

son and other family members. This was

an unusually productive day, because we

found 25 snakes of four species.” Photo by

Robbie Eagleston.

Amphib. Reptile Conserv.

3

August 2015 | Volume 9 | Number 2 | el 00

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [General Section]: 1-94 (el 00).

A conservation reassessment of the Central American

herpetofauna based on the EVS measure

1 Jerry D. Johnson, 2 Vicente Mata-Silva, and 3 Larry David Wilson

1,2 Department of Biological Sciences, The University of Texas at El Paso, El Paso, Texas 79968-0500, USA "Centro Zamorano de Biodiversidad,

Escuela Agricola Panamericana Zamorano, Departamento de Francisco Morazan, HONDURAS

Abstract. — Mesoamerica, the area composed of Mexico and Central America, is the third largest

of the world’s biodiversity hotspots. The Central American herpetofauna currently consists of 493

species of amphibians and 559 species of crocodylians, squamates, and turtles. In this paper, we

use a revised EVS measure to reexamine the conservation status of the native herpetofauna of this

region, utilize the General Lineage Concept of Species to recognize species-level taxa, and employ

phylogenetic concepts to determine evolutionary relationships among the taxa. Since the publication

of Conservation of Mesoamerican Amphibians and Reptiles, in 2010, 92 species of amphibians and

squamates have been described, resurrected, or elevated from subspecies to species level, and

one species of anuran has been synonymized. The herpetofaunal diversity of Central America is

comparable to that of Mexico, an especially significant finding because the land area of Mexico is

3.75 times larger. The number of amphibian species is 1.3 times greater in Central America, whereas

the number of species of turtles, crocodylians, and squamates is 1.5 times greater in Mexico.

Endemicity also is significant in Central America (65.6% among amphibians, 46.5% among turtles,

crocodylians, and squamates), with a combined average of 55.6%. We regard the IUCN system as

expensive, time-consuming, tending to fall behind systematic advances, and over-dependent on

the Data Deficient and Least Concern categories. Conversely, the EVS measure is economical, can

be applied when species are described, is predictive, simple to calculate, and does not “penalize”

poorly known species. Our EVS analysis of amphibians demonstrates that on average salamanders

are more susceptible to environmental deterioration, followed by caecilians, and anurans. Among

the remainder of the herpetofauna, crocodylians are the most susceptible and snakes the least, with

turtles and lizards in between. We compared the EVS results for the Central American herpetofauna

with those reported for Mexico; the results from those regions show an increase in numbers and

percentages from low through medium to high. Arguably, attempting to conserve biodiversity is one

of the most important and intransigent issues facing humanity, a situation partially due to humanity’s

lack of appreciation for its most serious concerns, and brought about by its anthropocentric focus.

Key words. EVS, anurans, salamanders, caecilians, crocodylians, turtles, lizards, snakes, IUCN categorizations, sur-

vival prospects

Resumen. — Mesoamerica, el area comprendida por Mexico y Centroamerica, es el centro de

biodiversidad mas grande del planeta. La herpetofauna de Centroamerica actualmente consiste de

493 especies de anfibios y 559 especies de crocodilidos, esquamados, y tortugas. En este articulo,

usamos la medida de EVS revisada para reexaminar el estado de conservacion de la herpetofauna

nativa de esta region, usamos el Concepto del Linaje General de Especie para reconocer taxones al

nivel de especie, y empleamos conceptos filogeneticos para determinar relaciones evolutivas entre

taxones. Desde la publicacion del libro Conservation of Mesoamerican Amphibians and Reptiles,

en 2010, 92 especies de anfibios y esquamados han sido descritas, resucitadas, o elevadas

de subespecie al nivel de especie y una especie de anuro ha sido sinonimizada. La diversidad

herpetofaunistica en Centroamerica es comparable a la de Mexico, un resultado especialmente

significativo dado que la superficie de Mexico es 3.75 veces mas grande. El numero de especies

de anfibios es 1.3 veces mayor en Centroamerica, mientras que el numero de especies de tortugas,

cocodrilidos y esquamados es 1.5 veces mayor en Mexico. El endemismo es tambien significativo

en Centroamerica (65.6% entre anfibios, 46.5% entre tortugas, cocodrilidos y esquamados), con un

Correspondence. Emails: fjohnson@utep.edu ; 2 vmata @ utep.edw, 3 bufodoc@ aol.com

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

4

Johnson et al.

promedio combinado de 55.6%. Consideramos el sistema de UICN como costoso, consume mucho

tiempo, con una tendencia a quedarse rebasado por los avances sistematicos, y sobre dependiente

de las categorias de Datos Deficientes y de Preocupacion Menor. Inversamente, la medida de EVS es

economica, puede ser aplicada cuando una especie es descrita, es predictiva, es facil de calcular y no

“penaliza” especies por ser pobremente conocidas. Nuestro analisis del EVS en anfibios demuestra

que en promedio las salamandras son las mas susceptibles al deterioro ambiental, seguidas por las

cecilias y los anuros. Entre el resto de la herpetofauna, los cocodrilidos son los mas susceptibles

y las serpientes las menos susceptibles, con las tortugas y las lagartijas en medio. Comparamos

los resultados del EVS de la herpetofauna de Centroamerica con la herpetofauna de Mexico; los

resultados para ambas regiones muestran un incremento en los numeros y porcentajes de baja

a mediana, a alta vulnerabilidad. Posiblemente, intentar conservar la biodiversidad es uno de los

problemas mas importantes y arduos que enfrenta la humanidad, una situacion parcialmente debida

a la falta de apreciacion de las preocupaciones mas serias por parte de la humanidad, y exacerbada

por su enfoque antropocentrico.

Palabras claves. EVS, anuros, salamandras, cecilias, cocodrilidos, tortugas, lagartijas, culebras, categorias de UICN,

perspectivas de supervivencia

Citation: Johnson JD, Mata-Silva V, Wilson LD. 2015. A conservation reassessment of the Central American herpetofauna based on the EVS mea-

sure. Amphibian & Reptile Conservation 9(2) [General Section]: 1-94 (el 00).

NoDerivatives 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: 11 March 2015; Accepted: 09 July 2015; Published: 14 August 2015

Currently, the global extinction rate far exceeds the rate

of speciation, and consequently, loss of species is the pri-

mary driver of changes in global biodiversity .. .Since the

advent of the Anthropocene, humans have increased the

rate of species extinction by 100-1,000 times the back-

ground rates that were typical over Earth’s history ...

Until recently, most extinctions (since 1500) occurred on

oceanic islands. In the last 20 years, however, about half

of the recorded extinctions have occurred on continents,

primarily due to land-use change, species introductions,

and increasingly climate change, indicating that biodi-

versity is now broadly at risk throughout the planet.

Rockstrom et al. 2009: 14

Introduction

The most significant problem facing humanity is biodi-

versity decline. Our attempts to estimate the total number

of species and our knowledge and appreciation of envi-

ronmental relationships within and among the large plan-

etary spheres are woefully inadequate. Strangely enough,

given the immense diversity of life on our planet and the

endless intellectual fulfillment its study can foster, hu-

mans have become increasingly focused on their own

activities and become increasingly removed from the rest

of the living world. In spite of this loss of perspective,

we are beginning to learn that our existence as a species

depends on our understanding of how life on this planet

operates, and the role we play in this process.

Amphib. Reptile Conserv. 5

In a Special Mexico Issue of the journal Amphibian

& Reptile Conservation , we conducted a conservation

reassessment of the reptiles (Wilson et al. 2013a) and

amphibians (Wilson et al. 2013b) of Mexico based on

the use of the Environmental Vulnerability Score (EVS).

These works allowed us to examine the results obtained

by the International Union for Conservation of Nature

(IUCN) and published in the Red List website (www.

iucnredlist.org), and compare them to our EVS results.

In total, we assayed 1,227 species (378 amphibians, 849

reptiles) of the Mexican herpetofauna. Our conclusions

from those studies were that, “both groups are highly im-

periled, especially the salamanders, lizards, and turtles”

(Wilson et al. 2013b: 98). Because the term “reptile” has

been demonstrated increasingly to have a paraphyletic

standing in phylogenetic systematics (www.iflscience.

com/plants-and-animals/there-s-no-such-thing-reptiles-

any-more-and-here-s-why), instead we use the names

“crocodylians, squamates, and turtles” when referring to

these groups.

The purpose of this paper is to reexamine the con-

servation status of the herpetofauna of Central America,

updating and broadening the treatments that appeared in

Conservation of Mesoamerican Amphibians and Rep-

tiles (CMAR; Wilson et al. 2010). A substantial amount

of systematic work has been published since the cutoff

point of 31 December 2008 used by Wilson and Johnson

(2010); our cutoff date for the present paper was 1 March

2015. In the interim, 92 species-level taxa have been

described, resurrected, or elevated, and one species was

August 2015 | Volume 9 | Number 2 | el 00

Conservation reassessment of Central American herpetofauna

synonymized. In addition, 30 species have undergone

status changes (usually placement in another genus). In

this study, therefore, we treat 1,052 species (493 am-

phibians; 559 crocodylians, squamates and turtles) and

use a revised EVS designed to encompass all of Central

America.

Our Taxonomic Positions

Transitions in systematics. — For herpetologists work-

ing in Mesoamerica, these are interesting times. We live

in a period of transition, from conditions characterizing

the past to those we envision will come in the future. The

element of transition is evident in much of what we pres-

ent in this paper and in our taxonomic positions, which

we deliberate below.

In trying to understand the biological aspects of the

Mesoamerican herpetofauna, we must be interested in

systematics, the study of the pattern of relationships

among living taxa (www.ucmp.berkeley.edu/clad/clad4.

html). Most systematists today practice phylogenetic

systematics, defined as “the way that biologists recon-

struct the pattern of events that has led to the distribu-

tion and diversity of life” (www.ucmp.berkeley.edu/clad/

clad4.html). The word “phylogenetic” refers to a system

based on evolutionary relationships, in this case among

members of biotic groups that commonly are depicted

as segments on a phylogeny (an evolutionary tree). As

with any reasoned system that has developed over time,

today’s phylogenetic systematics represents our current

understanding of the way life has diversified and changed

over time (www.ucmp.berkeley.edu/clad/clad4.html).

The degree that phylogenetic systematics has influenced

our present understanding of Mesoamerican herpetofau-

nal diversity has depended on the group of amphibians,

crocodylians, squamates, or turtles studied, and by the

level of acceptance of modem philosophical ideas and

techniques by taxonomists. Thus, our understanding of

phylogenetic systematics is in a state of transition, as we

keep moving from the ideas and techniques of the past

into those of the present and future.

Our predecessors attempted to catalogue all life, and

from Carolus Linnaeus we received a system of binomial

nomenclature that provided a means for biologists to com-

municate. Under the Linnaean system of nomenclature,

first and second names (generic and specific epithets) are

provided for living organisms. A system for the place-

ment of organisms into a set of hierarchically positioned

taxonomic categories followed. Another idea that near-

ly all biologists embrace is that life changes over time.

Charles Darwin delivered his theories of biotic evolution,

of which some still constitute fundamental themes of

modern-day biology. Presently, we combine the ideas of

Linnaeus and Darwin and recognize the species category

as the fundamental starting point of taxonomic inquiry.

Anything systematically linked to populations, below the

species level, is consigned to the ecologically regulated

expression of individual and geographic variation within

a species’ genotypic and related phenotypic characters;

geographic variation is how individual variation within

a species fluctuates in space. Genera and all other higher

taxonomic categories are not applicable until species are

recognized. Once recognized, species are named, and in

doing so must be placed within an existing genus or a

new one erected to incorporate the newly named spe-

cies. According to the rules of zoological nomenclature,

named taxa also are placed into a specified set of higher

taxonomic categories; major ones are genera, families,

orders, classes, phyla, kingdoms, and domains.

Species concepts and their evolution. — Biologists

also have inherited the part of systematics that deals with

understanding how species come to exist and how they

can be defined, and throughout history have provided

a suite of species concepts. Within the context of these

conceptions, the Biological Species Concept (BSC) pro-

posed ideas of definitive reproductive isolation and the

use of subspecies as a formal taxonomic category. The

BSC gained primacy as a means of objectively defining

and recognizing a species during the early to mid 20th

century. In those days, the modern synthesis of evolu-

tionary thought established genetic background as the

source for evolutionary processes, through the early

works of groundbreaking geneticists like Thomas H.

Morgan and Wilhelm Johannsen, and later by the sys-

tematists Theodosius Dobzhansky and especially Ernst

Mayr, whose book Systematics and the Origin of Spe-

cies from the Viewpoint of a Zoologist (1942) served as a

turning point for views about what constitutes a species.

Together with like-minded biologists, such as the herpe-

tologist and anatomist Hobart M. Smith, Mayr viewed a

species as a group of populations of organisms that are

capable of reproducing with each other and are reproduc-

tively isolated from other species. This species concept

enjoyed great popularity among biologists who worked

with sexually reproducing organisms, such as Mayr, who

was an ornithologist. Nonetheless, the BSC never ap-

pealed much to biologists who focused on asexually re-

producing organisms, because these creatures do not en-

joy sexual reproductive compatibility. Although the BSC

still holds sway in some comers of the biological world,

it has gradually been replaced by species concepts that

purport to work for all organisms, irrespective of their

means of reproduction, and which are part of an over-

arching view of how life has changed over time.

These efforts gained remarkable focus and became

part of the modern theory and practice of phylogenetic

systematics, which rests on a foundation of cladistic the-

ory pioneered by Willi Hennig in the 1930’s. Cladistic

analysis provided a means of erecting testable hypoth-

eses about evolutionary initiated connections among or-

ganisms, and currently is considered by many as the best

means for phylogenetic analysis (www.ucmp.berkeley.

edu/clad/cladl .html), which now we recognize predomi-

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Amphib. Reptile Conserv.

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Johnson et al.

Abronia vasconcelosii. This arboreal alligator lizard is endemic to the Guatemalan Plateau in the south-central portion of the coun-

try, where occurs in Lower Montane Wet Forest at elevations from 2,000 to 2,100 m. We assessed its EVS as 16, placing it in the

middle portion of the high vulnerability category, and its IUCN status is Vulnerable. This individual is from Cerro Alux, department

of Sacatepequez, Guatemala. Photo by Gunther Kohler.

Andinobates claudiae. This poison dart frog is endemic to islands on the Atlantic side of Panama, where occurs in Lowland Moist

Forest at elevations from 5 to 140 m. We gauged its EVS as 18, placing it in the upper portion of the high vulnerability category,

and its IUCN status is Data Deficient. This individual is from Isla Colon, province of Bocas del Toro. Photo by Brian Freiermuth.

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

7

Conservation reassessment of Central American herpetofauna

nantly among groups at higher taxonomic categories

(see below). Most importantly, cladistic analysis gives

biologists a way to use scientific methodology to study

how organisms are related to one another on an accepted

ancestor-descendent evolutionary basis. Cladistic proto-

cols recognize synapomorphies, namely shared derived

homologous characteristics, which uniquely distinguish

the related groups in which they are present from all oth-

er such groups, most specifically by sharing the derived

traits that originated during evolutionary modification of

the direct ancestor to the descendants comprising phylo-

genetic segments of an evolutionary lineage. Therefore,

cladistic systematics does not use reproductive capacity

as a universal character to identify sister species on a

phylogeny.

Reproduction is a characteristic of life, and sexual

reproduction is common to a large portion of living spe-

cies. Today, however, speciation in bisexual organisms

is properly recognized to arise by cladogenesis, which

is the splitting of a single lineage into two new geneti-

cally separate lineage segments. This idea, in part, dates

to at least Darwin and his supporters, and was expanded

upon by more modern phylogenetically-based species

concepts, like the Evolutionary and Phylogenetic Species

Concepts of George G. Simpson and Edward O. Wiley

for the former, and Niles Eldridge and Joel Cray craft for

the latter, among others. It was Kevin de Queiroz, in a se-

ries of papers dating from the late 1990s (e.g., de Queiroz

2005, 2007) that proposed a General Lineage Concept of

Species (GLCS) that reiterated species to be genetically

separated lineages, but uniquely embraced both clonal

(asexual) and bisexual reproductive systems. We inter-

prete the GLCS and its inclusive phylogenetically based

principles to falsify some traditionally used doctrines

that are deemed unusable in a modem phylogenetically

assembled taxonomic system; below we identify the ma-

jor ones associated with bisexual species.

As a consequence of modern phylogenetic theory, the

BSC as a universal definition for bisexual species essen-

tially was relegated to the systematics of the past, because

reproductive capability is not a synapomorphic state but

rather a plesiomorphic one, which is the ancestral state

before the feature evolved into the derived condition in

groups making up separate lineage segments found on a

phylogeny. Plesiomorphic characters cannot be used to

show sister relationships among other members of a phy-

logeny, because they can remain in that primitive condi-

tion in some or all taxa making up lineage segments of

the phylogeny. The demise of the BSC to depict phyloge-

netic relationships among related taxa because of its reli-

ance on an unusable trait (reproductive isolation) to show

sister relationships also led to the finale for the short-

lived impact of numerical taxonomy (Sneath and Sokal

1973). Numerical taxonomy used the overall similarity

of many unweighted phenetic traits to cluster sister taxa

together on a supposed phylogeny (actually a similarity

phenogram). The high number of plesiomorphic traits

Amphib. Reptile Conserv. 8

shared among closely related species, however, makes

the phenograms untenable for depicting phylogenetic

sister relationships because such primitive traits cannot

reflect evolutionary sister relationships among them.

Taxonomic processes at the species-lineage level usu-

ally do not follow cladistic principles, because specia-

tion regularly does not rely on shared synapomorphies

to specify sister relationships. In many of those cases

evolutionary relationships were established when new

evolutionary lineage segments were formed during allo-

patric speciation (geographic separation of ancestor into

two separate evolutionary lineage segments), so features

identifying separate sister species at that level can be

an assortment of traits that were present in the ancestor

to the two new lineage segments. Allopatric speciation

typically is not due to genetic changes, but rather to spa-

tial separation that eliminates gene flow. In other words,

newly defined sister species can be very similar (if not

identical) in genotypic structure and phenotypic expres-

sion during early periods of their lineage diversification.

Still, a synapomorphy could define sister species if its

attainment in the ancestral lineage is the reason for spe-

ciation, which generally would be expected in a sympat-

ric situation. The primary function of taxonomists work-

ing at the species level is to determine if gene flow has

ceased or not, and then decide what suite of taxonomic

characters will define the taxon as a new separate evolu-

tionary lineage segment.

A lineage is “any series of organisms connected by re-

production by parent of offspring” (www.ucmp.berkeley.

edu/glossary/glosslphylo.html). Thus, in bisexual organ-

isms, speciation occurs as soon as an ancestral gene pool

splits into two genetically isolated gene pools (lineage

segments), as depicted at the nodes of a phylogeny. Con-

sequently, there are no “stages of speciation,” other than

the initial complete separation of an ancestral lineage

into two new sister lineages, which can be rapid or pro-

longed depending on the source of separation. Evolution-

ary character divergences are not stages of speciation, but

rather changes within a single lineage’s gene pool during

its evolutionary lifespan. Some people consider stages

of speciation alongside some speculative rule when they

report that their sampled population has not changed ad-

equately in genetic distance or morphological divergence

to be considered a full species, as though some indefin-

able amount of evolutionary change is necessary to be

considered a different species. Frost and Hillis (1990)

correctly pointed out that “invoking a particular arbitrary

level of genetic distance or morphological divergence

as a species criterion is neither appropriate nor fruitful.”

This means that a species currently is defined only as a

separate evolutionary lineage and not by some subjec-

tive amount of evolutionary change. Because a single

lineage, say a species, does not develop into a new taxon

without a genetic split, the idea of anagenesis (develop-

ment of a new taxon without a genetic split) is negated,

along with the related idea that stages of speciation occur

August 2015 | Volume 9 | Number 2 | el 00

Johnson et al.

Agkistrodon howardgloydi. The Southern Cantil is distributed from southern Honduras to northwestern Costa Rica, where it occurs

in Lowland Arid and Dry forests at elevations from near sea level to 470 m. We determined its EVS as 17, placing it in the middle

portion of the high vulnerability category, but its IUCN status has not been determined. This individual is from Volcan Masaya,

Nicaragua. Photo by Javier Sunyer.

within a single lineage. Anagenesis has been considered

a valid concept in the past, but in order to form a new

species gene pool separation must exist.

The demise of the subspecies category. — Even though

the subspecies category has been associated with taxo-

nomically recognized geographic variants within a spe-

cies that are connected by gene flow (intergradation),

some tend to consider a subspecies as a stage of specia-

tion, even in light of clinal intergradation (gene-flow be-

tween members of the same species along a geographic

cline). Thus, the subspecies category no longer is useful

in systematics as a formal taxon, because by definition it

does not constitute a separate evolutionary lineage, nor

is it a stage of speciation. Disposing with this catego-

ry also eliminates the conundrum created with another

definition of a subspecies, as an entity consisting of or-

ganisms capable of interbreeding and producing fertile

offspring with other subspecies of the same species, but

cannot do so in nature because of geographic isolation

or other factors. This “they can but they don’t” paradox

remains because it is not a testable hypothesis through

scientific methodology. This definition also is misleading

because allopatric populations, by definition, are separate

evolutionary lineages due to genetic isolation and should

be considered full species, not subspecies. If supposedly

allopatric populations do not exhibit distinct genetic or

morphological differences at a particular point in time,

the only reasonable conclusion is that their gene pools

are not incontrovertibly separated, so those populations

should continue to be considered the same species until

empirical data reverse that conclusion. So again, a fore-

most issue for taxonomists is to determine if gene pool

separation has transpired or not.

Our understanding of the lowest-level phylogenetic

relationships is that only species are separate evolution-

ary lineages and, thus, only species can be depicted ap-

propriately on phylogenetic trees as lineage segments di-

verging from the nodes. Inserting subspecies as a lineage

segment branching at nodes might seem to give subspe-

cies a legitimate position as a formal taxonomic category,

but it does not because a population that is not a separate

evolutionaly lineage legitimately cannot be placed onto a

resolved phylogeny.

In summary, our position is that subspecies, as for-

merly defined, are not separate evolutionary lineages and

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Amphib. Reptile Conserv.

9

Conservation reassessment of Central American herpetofauna

cannot be placed into a phylogeny. Furthermore, subspe-

cies do not conform to an anagenetic stage of speciation

because those stages do not exist. The focus of phylo-

genetic systematics, therefore, including its association

with conservation biology, requires species to be the

fundamental unit of diversification as identified by their

binomial scientific name.

In addition, hybridization between two species in a

contact zone should not be a factor in determining the

presence of one or two species, because the ability to suc-

cessfully reproduce is a plesiomorphic characer that can-

not be used to identify phylogenetic sister relationships

among the species being investigated. Hybridization in

contact zones often is observed in natural situations; hy-

brids have no taxonomic status unless they lead to a sepa-

rate lineage segment.

Persistent issues in publication of systematic results.

— Another aspect of this discussion is our need to com-

ment on the GLCS theory and its practice in modem sys-

tematics, because of its scientific relevance in officially

published and unpublished literature. With the focus of

modern phylogenetic systematics being centered on evo-

lutionary divergence at the species level, our recognition

of amphibians, crocodylians, squamates, and turtles in

Central America is based on our interpretation of the in-

formation available in peer-reviewed scientific literature.

With continued advances in communication, especially

through the Internet, recognition of taxa should not be

founded on what one might find on a Facebook page, in

a blog, from someone’s tweet, or in a private non-peer-

reviewed journal (see Kaiser et al. 2013), no matter what

attempts are made to masquerade them as legitimate sci-

entific contributions. Thus, in documenting the makeup

of the Cental American herpetofauna, we cite our sources

as in Wilson et al. (2013a, b). Unfortunately, problems in

scientific publication still persist, which are identified to

clarify our position, as follows: (a) a lack of appropriate

taxon representation; (b) a lack of appropriate taxonomic

follow-through; and (c) taxon recognition based on non-

phylogenetic grounds. We discuss some of these prob-

lems below and in the section entitled “Controversial

Taxonomic Issues.”

An example of lack of appropriate taxon representa-

tion is evident in the manner in which recognition of the

genus Masticophis has been treated in recent literature.

We believe efforts to synonymize Masticophis with Col-

uber have been hampered by a serious lack of appropri-

ate taxon representation by previous investigators (e.g.,

Utiger et al. 2005; Pyron et al. 2013; and others). As

traditionally recognized (e.g., Wallach et al. 2014), Mas-

ticophis contains at least 11 species, and no taxonomic

analysis to date has included more than a small sample of

those. In addition, little effort has been made to examine

the phylogenetic relationships of the 1 1 species to more

than a handful of the other genera and their constituent

species that likely are close relatives of Masticophis and

Coluber (for elaboration, see section on Controversial

Taxonomic Issues).

Another example of a lack of appropriate taxon rep-

resentation regarding racers in the Burbrink et al. (2008)

study is the absence of samples of C. constrictor from

Mexico, Belize, or Guatemala, where the “subspecies”

C. c. oaxaca has been recognized (Kohler 2008). Lack of

appropriate taxon representation is a common inconsis-

tency in taxonomic studies of the herpetofauna that oc-

cur in the United States and neighboring Latin America,

where taxon sampling often stops at or near the United

States and Mexico border.

The single species recognized in the genus Coluber

(C. constrictor) is what used to be recognized as the

generotype of a much larger constellation of species that

mostly occur in the Old World, which now have been

segregated into seven genera (including the six listed

in Wallach et al. 2014, and another genus, Argyrogena,

resurrected by Wilson 1967, to contain the species A.

fasciolatus). Wallach et al. (2014) noted that Burbrink

et al. (2008) studied C. constrictor from a phylogenetic

perspective and recognized “six unnamed clades .” The

clades or lineages they recognized are reminiscent of the

“subspecies” arrangement held prior to the publication

of their study (e.g., Conant and Collins 1998; Stebbins

2003). Burbrink et al. (2008) concluded that, “according

to the general lineage concept of species, the racer may

not be a single taxon, particularly since several lineages

are well-defined geographically and are of very ancient

origin.” So, our questions to these authors are: (1) what

happened to the taxonomic follow-through; (2) what is

the taxonomic status of the six recognized but unnamed

lineages; and (3) given that the lineages are noticeably

not named in Burbrink et al. (2008), is there somewhere

else where they are, or will be named? The logical place

to find this information would be at the Center for North

American Herpetology website, but the standard comple-

ment of 1 1 subspecies is listed there (accessed 1 March

2015). Nonetheless, it would be simple to figure out the

names of the six lineages recognized in Burbrink et al.

(2008), but it is not our responsibility to second-guess

the authors and apply the designations to their recognized

lineages. We believe, however, that the authors of this

study and others like it are responsible for providing the

necessary taxonomic follow-through and place some bi-

nomial on the lineages in question, at least until someone

else reports different conclusions.

The last issue is taxon recognition based on non-phy-

logenetic grounds. Recognition of taxa must be founded

on conclusions reached in phylogenetic studies using ev-

idence-based data published in peer-reviewed scientific

outlets. Once published, the information can be applied

to resolve a variety of problems, such as determining

conservation status. Importantly, such resolutions must

be founded entirely on solid phylogenetic grounds. We

cite a perplexing recent example to the contrary. Sand-

ers et al. (2013) studied the phylogeny of the viviparous

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Amphib. Reptile Conserv.

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Johnson et al.

Bolitoglossci cerroensis. This web-footed salamander is dis-

tributed in the Cordillera de Talamanca in central Costa Rica,

where it occurs Lower Montane and Montane Wet forests at

elevations from 2,530 to 2,990 m. We determined its EVS as

16, placing it in the middle portion of the high vulnerability

category, and its IUCN status is Least Concern. This individual

is from near Cerro de la Muerte. Photo by Tobias Eisenberg.

Bolitoglossa centenorum. This web-footed salamander is only

known from the type locality on Cerro Bobic in west-central

Guatemala, in the Sierra de Cuchumatanes, department of

Huehuetenango, where it occurs in Montane Wet Forest at en

elevation of 3,250 m. We gauged its EVS as 18, placing it in

the upper portion of the high vulnerability category, but its

IUCN status has not been determined. This individual is from

near San Mateo Ixtatan. Photo by Todd Pierson.

seasnakes using both mitochondrial and nuclear markers

from 39 of 62 species and 15 of 16 genera. We found

one of their conclusions of particular interest because

they allocated the long-recognized genus Pelamis, with

its single species, to the genus Hydrophis (the name thus

became Hydrophis platurus). This view later was sup-

ported by the broader study of Pyron et al. (2013), so

we accept it based on the suggestions presented in both

studies. We take issue, however, with the last sentence

in Sanders et al. (2013), which reads: “The taxon Hy-

drophis is well known as comprising dangerously ven-

omous sea snakes; hence, retaining this name (instead of

adding multiple new genera) will create less confusion

for conservationists, medical professionals, and fishing

Bolitoglossa diaphora. This Cusuco web-footed salamander is

known only from Parque Nacional Cusuco, Sierra de Omoa,

in northwestern Honduras, where it occurs in Lower Montane

Wet Forest at elevations from 1,450 to 2,200 m. We calculated

its EVS as 18, placing it in the upper portion of the high vulner-

ability category, and its IUCN status is Critically Endangered.

This individual is from the vicinity of the type locality. Photo

by Todd Pierson.

Bolitoglossa aureogularis. The Yellow-throated Web-footed

Salamander is known only from two localities in Costa Rica, of

which one is the vicinity of the type locality on the Atlantic ver-

sant of the Cordillera de Talamanca; it occurs in Lower Mon-

tane Wet forest (cloud forest) at elevations from 1,680 to 2,100

m. We estimated its EVS as 18, placing it in the upper portion

of the high vulnerability category, but its IUCN status has not

been determined. This individual is from the headwaters of the

Rio Coen, province of Limon. Photo by Roney Santiago and

Eduardo Boza Oviedo.

industries/communities as well as herpetologists.” The

level of confusion agonized over by the types of people

indicated, including those compiling taxonomic lists

(taxonomic inflation - Isaac et al. 2004; Will et al. 2005)

is not a valid reason for reaching taxonomic conclusions,

in this case whether one genus {Hydrophis) should be

recognized or multiple genera (including, according to

the authors, five new genera). Making life easier for per-

sons not evolutionarily driven is not a valid motive for

disregarding phylogenetic conclusions.

We also contend that recognizing subspecies as a for-

mal taxonomic category, or placing them as separate evo-

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lutionary lineage segments on a phylogeny, are examples

of taxon recognition based on non-phylogenetic grounds.

Unfortunately, many studies continue this practice and

sometimes unnessessarily confound taxonomic issues.

In particular, the wrongful use of subspecies as a formal

taxonomic category can obscure the issue when geo-

graphic pattern classes (Grismer 2002) of the same spe-

cies are acknowledged with official taxonomic names.

Such variation can be erratic when it expresses vastly

inconsistent spatial features in ecological conditions and

in the size of intergrade zones, and is a non-phylogenetic

and speculative concept without basis in authenticity. In

a similar context, Uetz et al. (2014) unfairly criticized

Wallach et al. (2014) for not recognizing subspecies in

their Snakes of the World, which in our opinion was the

correct thing for the latter authors to do because of the

invalid status of subspecies in modem phylogenetically

based taxononmy. Identifying subspecies today only has

relevance in historical perspectives.

Controversial Taxonomic Issues

Our work deals with over one thousand species of am-

phibians, crocodylians, squamates, and turtles occurring

in Central America. Thus, differences in taxonomic opin-

ion are expected between our position and those held by

other systematic herpetologists. We discussed some of

these differences above in Our Taxonomic Positions sec-

tion, and discuss others below.

Trachemys in Central America. — In recent years,

the taxonomy of the turtle genus Trachemys in Meso-

america has been examined numerous times with in-

consistent results. Seidel and Smith (1986) transferred

the taxon Pseudemys scripta and its subspecies into the

genus Trachemys. Legler (1990) continued recognizing

Pseudemys as the genus containing T. scripta and ac-

knowledged the Central American forms as P. s. venusta

(Caribbean versant of southern Mexico and the Yucatan

Peninsula), P s. grayi (Pacific side from the Isthmus of

Tehuantepec to western Guatemala), and P. s. emolli (Ni-

caraguan lakes and Costa Rica). Ernst (1990) accepted

the genus Trachemys and similarly recognized the sub-

species T. s. venusta and T. s. grayi, but considered T. s.

ornata as occurring from Honduras to Panama. Seidel

(2002) later elevated two of the Central American forms

to T. emolli and T. venusta. Bonin et al. (2006) considered

T. ornata to be a Mexican Pacific versant endemic, T. ve-

nusta as occurring on the Atlantic slopes from Veracruz,

Mexico, to Panama and on the Pacific side from south-

eastern Oaxaca, Mexico, to Guatemala, and T. emolli as

restricted to Nicaragua and adjacent Costa Rica. Kohler

(2008) reviewed the most recent literature on this species

complex, but preferred to take a “conservative approach”

and relegated all Central American populations to inde-

terminate status as part of the wide-ranging Trachemys

scripta, but commented that he expected the taxonomy

to be revised.

Fritz et al. (2011) examined the molecular phylog-

eny of the slider turtles of Mexico, Central America, and

South America and determined previous allocations to

be incorrect, therein identifying two species in Central

America: T. grayi and T. ornata. Their evidence indicated

that T. grayi occurred intermittently on the Pacific low-

lands of Oaxaca, Mexico, through Panama and included

species or subspecies of taxa previously considered as

T. venusta panamensis, T. v. grayi, and T. emolli. Their

information also specified that T. ornata ranged sporadi-

cally on the Pacific versant from Sinaloa, Mexico (type

locality, Mazatlan), to a depicted allopatric population in

the vicinity of Acapulco, Guerrero, the only locality in

western Mexico from which they had samples. Trache-

mys ornata also was reported to occur from Tamaulipas,

Mexico, on the Atlantic versant into South America.

Populations of T. ornata from that area previously were

listed as comprised of T. venusta cataspila, T. v. venusta,

T. v. uhrigi, and two subspecies of T. callirostris in South

America. In a paper associated primarily with Antillean

Trachemys, Parham et al. (2013) continued to recognize

T. venusta for Atlantic versant turtles without analyzing

any T. ornata from western Mexico (except from the

supposed isolated population around Acapulco), and T.

emolli on the Pacific vesant of middle Central America

because of its supposed allopatric distribution. McCra-

nie et al. (2013), in reporting the taxon T. g. emolli in

southern Honduras, added new data that corroborated the

taxonomy of Fritz et al. (2011), although they cited the

publication date of that paper as 2012. The main problem

with both Fritz et al. (2011) and McCranie et al. (2013),

as with most recent sources, is that these authors contin-

ued to utilize subspecies as a formal taxonomic category.

The question arises as to what these recent studies

demonstrate regarding which species-level taxa of slider

turtles should be recognized in Central America. In our

effort to arrive at a decision, we examined the latest ver-

sion of the world turtle checklist published by the IUCN/

SSC Tortoise and Freshwater Turtle Specialist Group

(van Dijk et al. 2014). These authors adopted a position

that allows users of the checklist to arrive at their own

conclusion on what taxa at what level can or should be

recognized, which leads to a curious situation. They rec-

ognized three taxa of slider turtles in Central America.

One was T. venusta, which supposedly was distributed

principally along the Atlantic versant from Tamaulipas,

Mexico, to extreme northwestern Colombia, but also

on the Pacific versant in Panama, van Dijk et al. (2014),

however, suggested that this taxon also could be called,

in addition to T. venusta, T. ornata venusta, or T. venus-

ta venusta. They also listed T. grayi (Pacific versant of

Oaxaca, Mexico, to eastern El Salvador), but indicated

that it could also be called T. venusta grayi. Finally, they

included T. emolli (Pacific versant from eastern El Salva-

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Johnson et al.

Bolitoglossci indio. This web-footed salamander is known from the lowlands of the Rio San Juan area in extreme southeastern Nica-

ragua and north-central Costa Rica, where it occurs in Lowland Moist Forest at elevations from 25 to 68 m. We evaluated its EVS as

17, placing it in the middle portion of the high vulnerability category, and its IUCN status is Data Deficient. This individual is from

the type locality, Dos Bocas de Rio Indio, department of Rio San Juan, Nicaragua. Photo by Javier Sunyer.

dor to northwestern Costa Rica), but also listed the taxon

as T. grayi emolli. So, the reader could make a choice

among three species and/or subspecies (grayi, ornata,

and venusta ) into which to place the three Central Ameri-

can populations of slider turtles under a total of seven

preferred names.

We then examined Legler and Vogt’s (2013) book on

Mexican turtles to see how they handled the issue, and

quickly realized that their taxonomic arrangements were

permeated with subspecies, and that they continued to

treat all populations as subspecies of T. scripta. When

we ignored the trinomials and just concentrated on the

subspecific names as potential species names, Legler and

Vogt’s (2013) scheme would recognize the following: T.

ornata as occurring on the Pacific side of Mexico from

Sinaloa southwestward to the area around Acapulco,

Guerrero, the latter location depicted as a broadly al-

lopatric population (also illustrated that way by Legler

1990, and Seidel 2002); T. venusta as ranging on the At-

lantic versant from southeastern Veracruz through Cen-

tral America into Colombia, and on the Pacific side in

western Panama and adjacent Costa Rica; and T. grayi as

occurring on the Pacific versant from south-central Oax-

aca into El Salvador. They did not recognize the taxon

emolli that had been considered a subspecies of P. scripta

by Legler (1990) and T. scripta by Iverson (1992), as a

full species by Seidel (2002) and Jackson et al. (2008),

and as T. grayi by Fritz et al. (2011).

McCranie et al. (2013) also produced a subspecies in-

fused phylogeny, so again if their trinomials are ignored,

their taxa as based on distributional information found on

their phylogeny, included the following potential Central

American fonns: T. ornata ranging from Sinaloa, Mexi-

co, on the Pacific versant to Acapulco, Guerrero, and on

the Atlantic slope from Tamaulipas, Mexico, southward

and eastward through Central America to Venezuela; and

T. grayi occurring on the Pacific slope from southeastern

Oaxaca, Mexico, to Panama. A major difference of Fritz

et al. (2011) and McCranie et al. (2013), when compared

to the other papers, was that of all the species of Trache-

mys in Central America, only T. ornata occurred on both

Atlantic and Pacific versants of Mexico above the Isth-

mus of Tehuantepec and on the Atlantic slope of Central

America. Below the Isthmus, however, only T. grayi was

present on the Pacific side, from southwestern Mexico to

Panama. The pattern of species distributed on the Pacific

and Atlantic sides connected near the Isthmus of Tehuan-

tepec, as in T. ornata, also is found among crocodylians,

squamates, and other turtles (see maps in Kohler 2008).

Seidel (2002) and Legler and Vogt (2013) regarded

the population of Trachemys located on the Pacific side

of Panama and Costa Rica as T. venusta, a species that

almost everywhere else in Mesoamerica was an Atlantic

versant form. Parham et al. (2013) thought that T. venusta

and T. emolli probably intergraded in southern Nicaragua

and northern Costa Rica, although they apparently had no

access to the information in McCranie et al. (2013). Fritz

et al. (2011) and McCranie et al. (2013) both reported

that T. grayi was the species present from Pacific Costa

Rica and Panama, which was conspecific with other pop-

ulations to the northwest on the Pacific slopes, and not to

those on the Atlantic side. The question of what species

name to use for the Atlantic versant population occurring

from Tamaulipas into South America tentatively is an-

swered by recognizing the conclusions of the published

positions of Fritz et al. (2011) and McCranie (2013) that

T. ornata is the valid name, because it has publication

date priority over T. venusta. The decision by Parham et

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Conservation reassessment of Central American herpetofauna

Bothrops punctatus. This semiarboreal pitviper is distributed

from extreme eastern Panama to northwestern Ecuador, where

it occurs in Lowland Wet, Premontane Wet, and Lower Mon-

tane Wet forests at elevations from near sea level to 2,300 m.

We evaluated its EVS as 16, placing it in the middle portion

of the high vulnerability category, but its IUCN status has not

been determined. This individual is from the Serrania de Pirre,

province of Darien, Panama. Photo by Abel Batista.

Bothriechis marchi. The Honduran Emerald Tree Viper is en-

demic to northwestern and north-central Honduras, where it

occurs in Premontane Wet and Lower Montane Wet forests at

elevations from 500 to 1,840 m. We calculated its EVS as 16,

placing it in the middle portion of the high vulnerability cat-

egory, and its IUCN status is Endangered. This individual is

from Parque Nacional Cusuco, Sierra de Omoa, department of

Cortes. Photo by Silvia Petrovan.

Bradytriton silus. This salamander, the sole member of its ge-

nus, is endemic to the Sierra de Cuchumatanes in northwestern

Guatemala, where it is known only from two localities in Pre-

montane and Lower Montane Wet forests at elevations of 1,3 10

and 1,640 m. We established its EVS as 18, placing it in the

upper portion of the high vulnerability category, and its IUCN

status is Critically Endangered. This individual is from San

Jose Maxbal, department of Huehuetenango. Photo by Sean

Michael Rovito.

al. (2013) to revert to calling the Atlantic versant turtles

T. venusta is curious. These authors admitted that Atlan-

tic and Pacific Mexico populations probably were con-

specific and that the valid name would be T. ornata. Still,

they decided to maintain the name T. venusta because

they had no data from Mexican west coast T. ornata other

than samples from the supposed allopatric population in

the vicinity of Acapulco, which they thought might have

been introductions, and speculated that genetic introgres-

sion was the reason for their alliance with T. ornata-, to

us, this indicates that wild T. ornata probably were pres-

ent in the area. We also question the allopatric nature of

the Acapulco population because another Guerrero local -

Bolitoglossa insularis. This web-footed salamander is endemic

to Volcan Maderas on Ometepe Island in southwestern Nicara-

gua, where it occurs in Premontane Moist Forest at elevations

from 800 to 1,050 m. We assessed its EVS as 18, and its IUCN

status is Vulnerable. This individual is from Volcan Maderas,

Isla de Ometepe, department of Rivas. Photo by Javier Sunyer.

ity for T. ornata was reported by Mertz et al. (2015) from

200 km NW of Acapulco, which bridges a portion of the

distributional gap between Cabo Corrientes, Jalisco, and

Acapulco (Legler and Vogt 2013).

For our purposes in this paper and to try to reduce the

confusion created in the van Dijk et al. (2014) checklist

and other papers, we consider that the equivalent data in

Fritz et al. (2011) and McCranie (2013) best explain the

present knowledge of the taxonomic status of Trachemys

in Central America, so we recognize two species-level

taxa of slider turtles: T. grayi on the Pacific lowlands

and T. ornata on the Atlantic side, with their ranges as

indicated above. Nonetheless, we reject all reference to

subspecies due to taxonomic recognition based on non-

phylogenetic grounds.

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Johnson et al.

Taxonomy of Chelonia my das. — The Green Turtle,

Chelonia mydas, is a cosmopolitan species of marine tur-

tle that occurs in all the tropical to temperate oceans, and

has been regarded as showing considerable individual

and geographic variation in morphological and genetic

characters (see discussion in Ernst and Lovich 2009).

Chelonia agassizii, a supposedly Pacific Ocean form,

was named by Bocourt (1868) for an individual from the

Pacific coast of Guatemala, which some authorities have

determined to be a local variant of C. mydas (Karl and

Bowen 1999), others have considered it a subspecies of

C. mydas (Kamezaki and Matsui 1995), and still others

as a full species (Iverson 1992; Pritchard 1999; Savage

2002; Bonin et al. 2006). In a morphological study of C.

mydas from coastal waters around Japan, Okamoto and

Kamezaki (2014) found differences between two sam-

ples of turtles that appeared to validate C. mydas and C.

agassizii as separate species (at least around Japan), and

they commented on other studies in the Pacific Ocean

that agreed with their findings (e.g., Parker et al. 2011).

We consider that the possibility of the two species ar-

rangement eventually might stand or even expand. We

also feel, however, that accepting the two species scenar-

io is premature because of a serious lack of appropriate

taxon representation, especially in the Atlantic and In-

dian Oceans, as well as the need for using more relevant

phylogenetic criteria to decipher species-level taxonomic

status within the composite of populations associated

with C. mydas.

Status of Cryptochelys. — Taxon delimitation among the

turtles historically placed in the family Kinosternidae has

been challenging at all taxonomic levels, and this con-

troversy continues to the present. Two recent studies are

relevant to the status of members of this group in Central

America. As noted in van Dijk et al. (2014), “Iverson et al.

(2013) sequenced three mtDNA and three nuclear mark-

ers for every recognized species and most subspecies of

kinosternids. Their analyses revealed three well-resolved

clades within the Kinosternidae, corresponding to Ster-

notherus, a previously unnamed clade that they described

as the new genus Cryptochelys , and Kinosternon sensu

stricto. Their molecular data support for Cryptoche-

lys was strong, but data support for non-monophyly of

Kinosternon with respect to Sternotherus was weak. The

identified groups are broadly consistent with morpho-

logical and biogeographical features. Their new genus

Cryptochelys was diagnosed based on an extensive set

of morphological and molecular characters, and contains

the designated type species leucostoma, as well as acuta,

angustipons, creaseri, dunni, and herrerai .” van Dijk

et al (2014) referenced “a parallel study of kinosternid

phylogenetics . . . that reaches different taxonomic con-

clusions.” The title of this paper by Spinks et al. (2014),

“Multilocus phylogeny of the New-World mud turtles

(Kinosternidae) supports the traditional classification of

the group,” indicated the principal conclusion of this pa-

Amphib. Reptile Conserv. 15

per, i.e., a rebuttal of the Iverson et al. (2013) classifica-

tion, as well as an argument for maintaining stability in

organismic classifications. Their abstract provides a good

statement of their position, as follows: “A goal of modern

taxonomy is to develop classifications that reflect current

phylogenetic relationships and are as stable as possible

given the inherent uncertainties in much of the tree of

life. Here, we provide an in-depth phylogenetic analysis,

based on 14 nuclear loci comprising 10,305 base pairs

of aligned sequence data from all but two species of the

turtle family Kinosternidae, to determine whether recent

proposed changes to the group’s classification are jus-

tified and necessary. We conclude that those proposed

changes were based on (1) mtDNA gene tree anomalies,

(2) preliminary analyses that do not fully capture the

breadth of geographic variation necessary to motivate

taxonomic changes, and (3) changes in rank that are not

motivated by non-monophyletic groups. Our recommen-

dation, for this and other similar cases, is that taxonomic

changes be made only when phylogenetic results that are

statistically well-supported and corroborated by multiple

independent lines of genetic evidence indicate that non-

phylogenetic groups are currently recognized and need to

be corrected. We hope that other members of the phylo-

genetics community will join us in proposing taxonomic

changes only when the strongest phylogenetic data de-

mand such changes, and in so doing that we can move

toward stable, phylogenetically infonned classifications

of lasting value.” Operating on this basis, Spinks et al.

(2014) rejected the Iverson et al. (2013) genus Crypto-

chelys, moved the six above-mentioned species back into

the genus Kinosternon, and maintained recognition of

the genus Sternotherus. The Spinks et al. (2014) arrange-

ment appears to rest on a more secure basis, does not

support recognition of Cryptochelys, and is the approach

we tentatively adopted. Nonetheless, we wish to caution

those same “members of the phylogenetics community”

that attempting stability of organismic classification is

only desirable if it does not limit scientific discourse.

Given that humans will always be dealing with the inher-

ent uncertainties in much of the tree of life, and that their

scientific toolbox can only hope to recover phylogenies

of organisms about which we are aware, systematic bi-

ologists must have the freedom to attempt such recovery

in a spirit of cooperative enlightenment. After all, we are

guided in this effort by the conventions of peer review

in scientific publications and the principles of zoological

nomenclature. Even with these conventions, it will never

be possible for systematists to locate a comfortable arm-

chair from which to reflect on stable, phylogenetically

informed classifications of lasting value.

Staurotypinae vs. Staurotypidae. — Divergent ap-

proaches to the family-level classification of the genera

Claudius and Staurotypus were taken in the Iverson et al.

(2013) and Spinks et al. (2014) papers discussed above,

with the former arguing for the placement of these genera

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Conservation reassessment of Central American herpetofauna

Craugastor laevissimus. This species is distributed from west-

ern and east-central Honduras to northern and southwestern

Nicaragua, where it occurs in Lowland Moist, Lowland Dry,

Premontane Wet, Premontane Moist, Premontane Dry, and

Lower Montane Moist forests at elevations from near sea level

to 2,000 m. We assessed its EVS as 12, placing it in the up-

per portion of the medium vulnerability category, and its IUCN

status is Endangered. This individual is from Cerro Kilambe,

department of Jinotega, Nicaragua. Photo by Javier Sunyer.

Craugastor nefrens. The distribution of this ranita de hojarasca

(little litter frog) is restricted to a narrow elevational band (800-

1,000 m) of Premontane Wet Forest in the Sierra de Caral of

eastern Guatemala, near the border with Honduras. We estab-

lished its EVS as 18, placing it in the upper portion of the high

vulnerability category, and its IUCN status is Data Deficient.

This individual is from Finca la Firmeza, Morales, department

of Izabal. Photo by Sean Michael Rovito.

Craugastor chingopetaca. This rainfrog is known only from

the type locality along the Rio San Juan in extreme southeast-

ern Nicaragua, department of Rio San Juan, where it occurs in

Lowland Wet Forest at an elevation of 40 m. We evaluated its

EVS as 18, placing it in the upper portion of the high vulner-

ability category, and its IUCN status is Data Deficient. This in-

dividual is from Reserva de Vida Silvestre Rio San Juan. Photo

by Javier Sunyer.

Ctenosaura palearis. This Guatemalan spiny-tailed iguana is

endemic to the Motagua Valley in eastern Guatemala, where it

occurs in Lowland Arid and Premontane Dry forests at eleva-

tions from 150 to 700 m. We calculated its EVS as 19, placing

it in the upper portion of the high vulnerability category, and

its IUCN status is Endangered. This individual is from Zacapa,

Motagua River Valley, department of Zacapa. Photo by Antonia

Pachmann.

in the family Staurotypidae and the latter in the subfami-

ly Staurotypinae. Iverson et al. (2013) followed Bickham

and Carr (1983) in recognizing two clades, one consisting

of Claudius and Staurotypus and another of Kinosternon

and Sternotherus , as separate families, based on the esti-

mated age of the clades and their unambiguously distinct

morphologies and sex-determining mechanisms (genetic

sex determination in the former clade and temperature-

dependent sex determination in the latter), as well as the

concatenated sequences of three nuclear and three mi-

tochondrial genes. Spinks et al. (2014: 258), however,

argued that, “in the interest of maintaining taxonomic

stability ... we suggest that the community maintain the

historical treatment of Staurotypinae as a subfamily as

has been done for decades.” We briefly explained our po-

sition on this matter above, and in this case follow the

recommendations of Iverson et al. (2013) and recognize

the genera Claudius and Staurotypus in the family Stau-

rotypidae, distinct from the family Kinosternidae that in-

cludes the genera Kinosternon and Sternotherus .

Single-genus vs. multiple-genera approaches to anole

classification. — A sizeable number of herpetologists

are interested in anoles and their classification. Over the

years, many herpetologists have tried to make sense of

a group of lizards that presently contains 395 species

Amphib. Reptile Conserv.

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Johnson et al.

(Reptile Database website; accessed 28 February 2015),

with more added each year (e.g., see our listing of pres-

ently added taxa to the Central American herpetofauna,

in which we document the recognition of 15 additional

species-level taxa since the publication of Wilson et al.,

2010). Gunther Kohler and his colleagues undertook

most of this work and with one exception {Dactyloa gi-

naelisae) described or resurrected the remainder under

the genus Anolis. In our present work, we list 95 species

of anoles in Central America, and Wilson et al. (2013a)

recorded 50 species from Mexico; presently 129 species

comprise the anole fauna of Mesoamerica (16 species oc-

cupy both regions; www.mesoamericanherpetology.com;

accessed 28 February 2015). In Wilson et al. (2013a), we

listed all 50 Mexican species under the genus Anolis.

We took that position because a controversy was brew-

ing over the classification proposed by Nicholson et al.

(2012), especially with the harsh rebuttal of this paper

by Poe (2013), and we were uncertain where the contro-

versy would go. Since that time, however, Nicholson et

al. (2014) provided a detailed response addressing Poe’s

concerns. Most anyone with an interest in anole system-

atics knows the backstory, beginning with Guyer and

Savage’s (1986) revolutionary cladistic analysis of the

anoles. The effect of that study was to segment the huge

and unwieldy genus Anolis into a series of eight genera.

Subsequently, Williams (1989) authored a scathing cri-

tique of the Guyer- Savage approach, asking if the data

were available to reclassify the anoles; herpetologists

varied in their opinions. During the ensuing years, stu-

dents of tropical American herpetology basically fell into

two camps, those who supported or opposed the Guyer-

Savage scheme. In recent years, we sided with the latter

camp (Wilson and Johnson 2010; Johnson et al. 2010;

Wilson et al. 2013a), but did not undertake an exhaustive

study of the matter. Nonetheless, after the publication of

Poe’s (2013) critique of the Nicholson et al. (2012) paper

and the Nicholson et al. (2014) rebuttal, we decided to

take a fresh look at this issue. Principally, the controversy

that developed over the last two years results from two

approaches to the classification of anoles. The Nichol-

son et al. (2012, 2014) approach was to recognize eight

genera of these lizards. Poe’s (2013) tactic was to jetti-

son entirely the Nicholson et al. (2012) approach and to

recognize a single genus that contained 391 species, the

largest genus of squamates. Fundamentally, Poe’s criti-

cism of the eight-genus approach was two-fold, i.e., that

“some of the proposed genera are not monophyletic” and

that Nicholson et al. (2012) did not study enough taxa

or enough characters. Nicholson et al. (2014) presented

their rebuttal “to explain how Poe erred in characteriz-

ing our work, and missed the opportunity to present an

alternative comprehensive taxonomy to replace the one

against which he argues so strenuously. In this contribu-

tion we explain, and correct, Poe’s errors and misrepre-

sentations, and argue that our taxonomy is likely to be

adopted because it (1) eliminates the obvious problem

that will arise if the family Dactyloidae contains only a

single large genus (i.e., that a single genus obscures the

evolution and diversity within the group and misrepre-

sents or cloaks it), (2) it conforms with the long historical

trend of dissecting large, cumbersome groups into small-

er sub-units, (3) is consistent with all recent phylogenetic

studies for anoles in membership within clades we rec-

ognize as genera, and (4) aids in associating these lizards

with the ancient land masses that shaped their history.”

We consider that Nicholson and her coauthors adequate-

ly responded to Poe’s criticisms and we are confident in

adopting the portion of their scheme relevant to the situ-

ation in Central America, and Mesoamerica as a whole.

So, what impact does the Nicholson et al. approach have

on the taxonomy of anoles in Mesoamerica? As it turns

out, only three of the eight genera Nicholson et al. (2012,

2014) recognized contain Mesoamerican species as fol-

lows: Anolis (one species), Dactyloa (10 species), and

Norops (118 species). The distribution of the genus Ano-

lis is stated by Nicholson et al. (2012) to be in “the Ba-

hamas, Cuba, and adjacent islands, Navassa Island, Little

Cayman [Ijsland, Hispaniola, and the southeastern Unit-

ed States west to Oklahoma and Texas.” They further in-

dicated that, “one Cuban species (A. allisoni ) occurs on

Isla Cozumel, Mexico and Islas de la Bahia, Honduras,

and on coastal islands off Belize.” Distribution of the ge-

nus Dactyloa is indicated by Nicholson et al. (2012) to

be on the “Atlantic and Pacific slopes of Costa Rica and

Panama, then south through the Choco region of Colom-

bia and Ecuador, including Malpelo Island; highlands

of Colombia, Ecuador, Peru, and Venezuela; Caribbean

slope of Colombia and Venezuela; Bonaire and Blanquil-

la Islands and the southern Lesser Antilles; south on the

Atlantic versant through the Guayanas to Espiritu Santo

State in eastern Brazil, and throughout the Orinoco and

Amazon Basins in Colombia, Ecuador, Peru, Venezu-

ela, Bolivia, and Brazil.” The remainder of the anoles in

Central America (as well as all of the species in Mexico

except for Anolis allisoni) are placed in the genus No-

rops, which Nicholson et al. (2012) reported to occur in

“Cuba, Jamaica, Bahamas, Grand and Little Cayman,

Cayman Brae, Mexico, Central America, and many ad-

jacent islands, including Cozumel, the Bay Islands, the

Corn Islands, Swan Island, San Andres and Providencia

(Caribbean) and Isla del Coco (Pacific); south to west-

ern Ecuador, northern South America (Colombia and

Venezuela), including Isla Gorgona (Pacific), the islands

of Aruba, Curasao, and Margarita (Caribbean), Trinidad

and Tobago; then south through the Guyanas to south-

eastern and southern Brazil, and Paraguay, and through-

out the Orinoco and Amazon Basins (Colombia, Ven-

ezuela, Ecuador, Peru, Brazil, and Bolivia).” We agree

that Nicholson and her coauthors provided a perceptive

set of reasons why their classification will be accepted

in time, just as with other classifications that sought to

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Amphib. Reptile Conserv.

17

Conservation reassessment of Central American herpetofauna

Cryptotriton nasalis. This small salamander is endemic to the Sierra de Omoa in northwestern Honduras, where it occurs in Premon-

tane and Lower Montane Wet forests at elevations from 1,220 to 2,200 m. We estimated its EVS as 18, placing it in the upper portion

of the high vulnerability category, and its IUCN status is Endangered. This individual is from the Sierra de Omoa, department of

Cortes. Photo by Sean Michael Rovito.

make sense of formerly unmanageable genera, such as

Eleutherodactylus, which now not only is segmented into

a number of genera, but also a number of families.

Coluber versus Mastic op his. — We base most of this

commentary on information discussed in Wilson and

Johnson (2010), along with a fresh look at the available

data and on our reliance on the proper use of phyloge-

netic systematics to produce accurate conclusions. The

major issue is: should the genus Masticophis be synony-

mized with the genus Coluber based on the information

available today? This question has been contentious for

many years, and the disagreement stems from a number

of factors, including overall molecular, morphological,

and ontogenetic similarities between the two genera; a

prodigious lack of appropriate taxon representation in

seminal papers of recent vintage (see Our Taxonomic Po-

sition section), especially those that reflected molecular

comparisons; and the overt continuation of recognizing

groups at the subspecies level.

Nagy et al. (2004), in a molecular study using mito-

chondrial and nuclear genes, agreed with Schatti’s (1987)

morphogical investigation that the genus Coluber {sensu

stricto ) should be restricted to the New World; both de-

clined to synonymize Masticophis with Coluber based

on their own data. Utiger et al. (2005), with low support,

found Masticophis flagellum to be nested within Colu-

ber constrictor , with M. taeniatus as the sister to the C.

constrictor-M. flagellum clade, which made Mastico-

phis paraphyletic, therein recommending the placement

of Masticophis into Coluber (the older generic name).

Burbrink et al. (2008) examined C. constrictor from

throughout upper North America and concluded the spe-

cies to be monophyletic and composed of six unnamed

lineages; they also considered M. flagellum the sister

species to C. constrictor , thus negating Utiger et al.’s.

(2005) verdict that a population of C. flagellum was more

closely related to C. constrictor than to other populations

of C. flagellum. The Burbrink et al. (2008) treatment also

is afflicted with a lack of taxonomic follow-through, in-

asmuch as the separate lineages within the C. constrictor

complex they disclosed are not named. In addition, they

did not indicate the species to which M. flagellum is the

sister taxon. Collins and Taggart (2008) correctly noted

that because of incomplete taxon sampling by Utiger et

al. (2005), the generic status of certain taxa could not be

addressed adequately. Wilson and Johnson (2010) also

presented summary information on this debate, and com-

mented that Utiger et al. (2005) did not provide adequate

samples from throughout the range of the respective taxa

(e.g., at least nine other species of Masticophis were not

included in their study). Both Collins and Taggart (2008)

and Wilson and Johnson (2010) recommended the con-

tinued recognition of both genera as separate taxa, al-

though some publications have continued to use Coluber

for all the species of Masticoph is , most notedly C. flagel-

lum and C. taeniatus , species occurring sympatrically in

the southwestern United States.

Importantly, no comparison has been made between

M. flagellum and the wide-ranging M. mentovarius, as

presently envisioned, which long were thought to be

sister species (e.g., Wilson 1970; Johnson 1977). Also,

only a small amount of genetic material has been avail-

able to examine and compare the relationships of Colu-

ber and Masticophis to other genera of North American

racer-like colubrids (e.g., Dendrophidion, Drymobius,

Leptodrymus, Leptophis, Mastigodryas, Salvadora), of

which most do not occur northward outside of Mexico.

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

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Johnson et al.

Craugastor polyptychus. This frog is distributed along the

lowlands of the Atlantic versant from extreme southeastern Ni-

caragua to extreme northwestern Panama, where it occurs in

Lowland Moist Forest at elevations from near sea level to 260

m. We estimated its EVS as 17, placing it in the middle portion

of the high vulnerability category, and its IUCN status is Least

Concern. This individual is from the Refugio Nacional de Vida

Silvestre Gandoca-Manzanillo, province of Limon, Costa Rica.

Photo by Maciej Pabijan.

Dactyloa ibanezi. This anole is distributed on the Caribbean

versant from southeastern Costa Rica to western Panama,

where it occurs in Lowland Moist and Premontane Wet forests

at elevations from 400 to 1,070 m. We established its EVS as

15, placing it in the lower portion of the high vulnerability cat-

egory, but its IUCN status has not been determined. This indi-

vidual is from Donoso, province of Colon, Panama. Photo by

Abel Batista.

In an extensive review of squamates, Pyron et al. (2013)

also showed C. constrictor and C. flagellum as sister spe-

cies and C. taeniatus as the sister to that clade, but didn’t

mention the overt lack of appropriate taxon representa-

tion when producing their phylogeny. Pyron et al. (2013)

included some samples of other racer-like genera in their

phylogeny, but still maintained a lack of sufficient taxon

sampling in those genera, of which most have not un-

dergone recent phylogenetic analyses. After all, if the

phylogenetic interpretation is not accurate or based on

inadequate taxonomic representation, it could lead to er-

roneous conclusions.

Crocodylus acutus. The American Crocodile is broadly distrib-

uted in the Caribbean Basin from southern Florida and the Yu-

catan Peninsula south to Colombia and Venezuela, and on the

Pacific coast of Latin America from Sinaloa in Mexico to Peru

in South America. We evaluated its EVS as 14, at the lower

end of the high vulnerability category, and its IUCN status is

Vulnerable. This individual is from Isla Juan Venado, a barrier

island constituting a nature reserve, department of Leon, Nica-

ragua. Photo by Javier Sunyer.

Dactyloa kunayalae. This anole is distributed in western and

central Panama, where it occurs in Lowland Moist and Pre-

montane Wet forests at elevations from 320 to 1,050 m. We

estimated its EVS as 15, placing it in the lower portion of the

high vulnerability category, but its IUCN status has not been

determined. This individual is from the Rio Tuquesa, province

of Darien, Panama. Photo by Abel Batista.

Another germane question about the generic status of

Masticophis could be resolved by determining the phy-

logenetic position of M. taeniatus compared with that of

the above-mentioned genera of racer-like species. After

an all-encompasing phylogenetic comparison, the pos-

sibility exists that a monophyletic M. taeniatus group

(eight species), could be assigned to a genus other than

Masticophis or Coluber ; which would remove the para-

phyletic status of Masticophis, and make its inclusion

into Coluber inconsequential.

In conclusion, because pertinent phylogenetic studies

on the issue of Coluber versus Masticophis have not in-

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Amphib. Reptile Conserv.

19

Conservation reassessment of Central American herpetofauna

eluded appropriate taxonomic representation of members

of the genera Masticophis and Coluber or genera of other

Western Hemisphere racer-like colubrids, we accept the

recommendations of Collins and Taggart (2008) and Wil-

son and Johnson (2010) and use the name Masticophis

for the 1 1 species traditionally included in this genus, in-

cluding M. mentovarius in Central America.

A Revised Environmental Vulnerability Measure

Wilson et al. (2013a, b) adapted the Environmental Vul-

nerability Score developed by Wilson and McCranie

(2004) for use in Mexico. The Mexican EVS only dif-

fered from that used for Honduras by Wilson and Mc-

Cranie (2004) in the design of the geographic component

(considering, however, that the third component of the

measure differed between amphibians and the remainder

of the herpetofauna). Herein, we revised the same com-

ponent for use with the Central American herpetofauna,

as follows:

1 = distribution broadly represented both inside and

outside of Central America (large portions of

the range are both inside and outside of Central

America)

2 = distribution prevalent inside of Central America,

but limited outside of Central America (most of

the range is inside of Central America)

3 = distribution limited inside of Central America, but

prevalent outside of Central America (most of the

range is outside of Central America)

4 = distribution limited both inside and outside of

Central America (most of range is marginal to

areas near the border of Central America and

Mexico or South America, respectively)

5 = distribution only within Central America, but not

restricted to the vicinity of the type locality

6 = distribution limited to Central America in the

vicinity of the type locality

The second component of the EVS measure, for eco-

logical distribution based on occurrence in different veg-

etaion formations, is the same for Central America as for

Mexico, as follows:

1 = occurs in eight or more formations

2 = occurs in seven formations

3 = occurs in six formations

4 = occurs in five formations

5 = occurs in four formations

6 = occurs in three formations

7 = occurs in two formations

8 = occurs in one formation

The third component, for amphibians, deals with the

type of reproductive mode, as follows:

Amphib. Reptile Conserv.

1 = both eggs and tadpoles are found in large to small

bodies of lentic or lotic water

2 = eggs are deposited in foam nests, and tadpoles are

found in small bodies of lentic or lotic water

3 = tadpoles are found in small bodies of lentic or

lotic water, and eggs elsewhere

4 = eggs are laid in moist situations on land or in

moist arboreal situations, and tadpoles (larvae)

are carried (or move) to water or undergo direct

development

5 = eggs and/or tadpoles are carried in the dorsal

pouch of the female or are imbedded in the dor-

sum of female, larval or direct development, or

viviparous

6 = eggs and tadpoles are found in water-retaining

arboreal bromeliads or in water-filled tree cavities

The third component, for crocodylians, squamates,

and turtles, deals with the degree of human persecution,

as follows:

1 = fossorial, usually escape human notice

2 = semifossorial, or nocturnal arboreal or aquatic,

nonvenomous and usually non-mimicking, some-

times escape human notice

3 = terrestrial and/or arboreal or aquatic, generally

ignored by humans

4 = terrestrial and/or arboreal or aquatic, thought to be

harmful, might be killed on sight

5 = venomous species or mimics thereof, killed on

sight

6 = commercially or non-commercially exploited for

hides, meat, eggs and/or the pet trade

Once these three components are added, the EVS

can range from 3 to 20 in both groups. Wilson and Mc-

Cranie (2004) placed the range of scores for Honduran

amphibians into three categories of vulnerability to en-

vironmental degradation, as follows: low (3-9); medium

(10-13); and high (14-19). The categories for the rest

of the herpetofauna were similar, with the high category

encompassing values of 14-20. Herein, we employ the

same categorizations: low (3-9); medium (10-13); and

high (14-20). In Appendices 1 and 2, these categories

are signified by the abbreviations L (low), M (medium),

and H (high).

Recent Changes to the Central American

Herpetofauna

Due to ongoing fieldwork in Central America by a num-

ber of herpetologists from around the globe, and the

systematic research emanating from their fieldwork, the

composition of the region’s herpetofauna constantly is

being updated. In most cases, the number of recognized

taxa increases. These changes add or subtract from the

taxonomic lists that appeared in Wilson et al. (2010);

August 2015 | Volume 9 | Number 2 | el 00

20

Johnson et al.

Dactyloa latifrons. This anole is distributed from eastern Panama to northwestern Ecuador, where it occurs in Premontane Wet

Forest at elevations from 665 to 780 m. We gauged its EVS as 13, placing it at the upper end of the medium vulnerability category,

but its IUCN status has not been determined. This individual is from the Serrania de Pirre, province of Darien, Panama. Photo by

Abel Batista.

since that work appeared, the following 92 species have

been described, resurrected, or elevated to species level:

Anomaloglossus astralogaster. Myers et al. 2012.

American Museum Novitates 3,763: 1-19. New

species.

Anomaloglossus isthminus : Myers et al. 2012. Ameri-

can Museum Novitates 3763: 1-19. New species.

Atelopus chirripoensis: Savage and Bolanos. 2009.

Revista Biologia Tropical 57: 381-386. New spe-

cies.

Incilius aurarius : Mendelson et al. 2012. Journal of

Herpetology 46: 473^179. New species.

Incilius karenlipsae: Mendelson and Mulcahy. 2010.

Zootaxa 2396: 61-68. New species.

Craugastor evanesco : Ryan et al. 2010b. Copeia

2010: 405^109. New species.

Andinobates geminisae: Batista et al. 2014b. Zootaxa

3866: 333-352. New species.

Diasp orus citrinobapheus: Hertz et al. 2012. ZooKeys

196: 23-46. New species.

Diasp orus igneus: Batista et al. 2012. Zootaxa 3410:

51-60. New species.

Ecnomiohyla bailarina : Batista et al. 2014c. Zootaxa

3826: 449^174. New species.

Ecnomiohyla sukia : Savage and Kubicki. 2010. Zoo-

taxa 2719: 21-34. New species.

Ecnomiohyla veraguensis: Batista et al. 2014c. Zoo-

taxa 3826: 449^174. New species.

Pristimantis adnus : Crawford et al. 2010. Herpeto-

logica 66: 171-185. New species.

Bolitoglossa aureogularis: Boza-Oviedo et al. 2012.

Zootaxa 3309: 36-61. New species.

Bolitoglossa centenorum : Campbell et al. 2010. Mis-

cellaneous Publications, Museum of Zoology, Uni-

versity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa chucantiensis: Batista et al. 2014d. Me-

soamerican Herpetology 1: 96-121. New species.

Bolitoglossa daryorum: Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa eremia : Campbell et al. 2010. Miscella-

neous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa huehuetenanguensis: Campbell et al.

2010. Miscellaneous Publications, Museum of Zo-

ology, University of Michigan (200): i-iv, 1-60.

New species.

Bolitoglossa jugivagans: Hertz et al. 2013. Zootaxa

3636: 463^175. New species.

Amphib. Reptile Conserv.

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August 2015 | Volume 9 | Number 2 | el 00

Conservation reassessment of Central American herpetofauna

Bolitoglossa kamuk: Boza-Oviedo et at. 2012. Zoo-

taxa 3309: 36-61. New species.

Bolitoglossa kaqchikelorum: Campbell et al. 2010.

Miscellaneous Publications, Museum of Zoology,

University of Michigan (200): i-iv, 1-60. New spe-

cies.

Bolitoglossa la: Campbell et al. 2010. Miscellaneous

Publications, Museum of Zoology, University of

Michigan (200): i-iv, 1-60. New species.

Bolitoglossa ninadormida: Campbell et al. 2010. Mis-

cellaneous Publications, Museum of Zoology, Uni-

versity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa nussbaumi: Campbell et al. 2010. Mis-

cellaneous Publications, Museum of Zoology, Uni-

versity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa nympha : Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa omniums anctorum: Campbell et al.

2010. Miscellaneous Publications, Museum of Zo-

ology, University of Michigan (200): i-iv, 1-60.

Resurrection from synonymy.

Bolitoglossa pacaya: Campbell et al. 2010. Miscella-

neous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa psephena: Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa pygmaea: Bolanos and Wake. 2009. Zoo-

taxa 1981: 57-68. New species.

Bolitoglossa robinsoni: Bolanos and Wake. 2009.

Zootaxa 1981: 57-68. New species.

Bolitoglossa splendida: Boza-Oviedo et al. 2012.

Zootaxa 3309: 36-61. New species.

Bolitoglossa suchitanensis: Campbell et al. 2010.

Miscellaneous Publications, Museum of Zoology,

University of Michigan (200): i-iv, 1-60. New spe-

cies.

Bolitoglossa tenebrosa: Vasquez-Almazan and Ro-

vito. 2014. Journal of Herpetology 48: 518—524.

New species.

Bolitoglossa tzultacaj: Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa xibalba: Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Bolitoglossa zacapensis: Rovito et al. 2010. Journal

of Herpetology 44: 516-525. New species.

Cryptotriton necopinus: McCranie and Rovito. 2014.

Zootaxa 3795: 61-70. New species.

Cryptotriton sierraminensis: Vasquez-Almazan et al.

2009. Copeia 2009: 313-319. New species.

Dendrotriton chujorum: Campbell et al. 2010. Miscel-

laneous Publications, Museum of Zoology, Univer-

sity of Michigan (200): i-iv, 1-60. New species.

Dendrotriton kekchiorum: Campbell et al. 2010. Mis-

cellaneous Publications, Museum of Zoology, Uni-

versity of Michigan (200): i-iv, 1-60. New species.

Nototriton matama: Boza-Oviedo et al. 2012. Zootaxa

3309: 36-61. New species.

Nototriton mime: Townsend et al. 2013c. Zootaxa

3666: 359-368. New species.

Nototriton picucha: Townsend et al. 2011. Systemat-

ics and Biodiversity 9: 275-287. New species.

Oedipina chortiorum: Brodie et al. 2012. Journal of

Herpetology 46: 233-240. New species.

Oedipina koehleri: Sunyer et al. 2011. Breviora 526:

1-16. New species.

Oedipina motaguae: Brodie et al. 2012. Journal of

Herpetology 46: 233-240. New species.

Oedipina nica: Sunyer et al. 2010. Zootaxa 2613:

29-39. New species.

Oedipina nimaso: Boza-Oviedo et al. 2012. Zootaxa

3309: 36-61. New species.

Oedipina petiola: McCranie and Townsend. 2011.

Zootaxa 2990: 59-68. New species.

Oedipina tzutujilorum: Brodie et al. 2012. Journal of

Herpetology 46: 233-240. New species.

Dactyloa ginaelisae: Lotzkat et al. 2013. Zootaxa

3626: 1-54. New species.

Dactyloa ibanezi: Poe et al. 2009. Phyllomedusa 8:

81-87. New species.

Norops alocomyos: Kohler et al. 2014. Zootaxa 3915:

1 1 1-122. New species.

Norops beckeri: Kohler. 2010. Zootaxa 2354: 1—8.

Resurrection from the synonymy of A. pentaprion.

Norops benedikti: Lotzkat et al. 2011. Zootaxa 3125:

1-21. New species.

Norops charlesmyersi: Kohler. 2010. Zootaxa 2354:

1-8. New species.

Norops gaigei: Kohler et al. 2012. Zootaxa 3348:

1-23. Resurrection of A. gaigei from the synony-

my of A. tropidogaster.

Norops leditzigorum: Kohler et al. 2014. Zootaxa

3915: 111-122. New species.

Norops marsupialis: Kohler et al. 2015. Zootaxa 3915:

111-122. Resurrection of A. marsupialis from the

synonymy of A. humilis. Previously recognized at

the species level without comment by Bolanos et

al. (2011).

Norops monteverde : Kohler. 2009. Journal of Herpe-

tology 43: 11-20. New species.

Norops osa: Kohler et al. 2010a. Zootaxa 2718: 23-

38. New species.

Norops tenorioensis: Kohler 2011. Zootaxa 3120:

29^12. New species.

Norops triumphalis: Nicholson and Kohler. 2014.

Zootaxa 3895: 225-237 . New species.

Norops unilobatus: Kohler and Vesely. 2010. Herpe-

tologica 66: 186-207. New species.

Amphib. Reptile Conserv.

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Johnson et al.

Dendrotriton chujorum. This salamander is endemic to the

northern portion of the Sierra de Cuchumatanes in northwest-

ern Guatemala, where occurs in the lower extent of Montane

Wet Forest at elevations from 2,697 to 2,792 m in. We gauged

its EVS as 18, placing it in the upper portion of the high vulner-

ability category, and its IUCN status is Critically Endangered.

This individual is from near San Mateo Ixtatan, Sierra de los

Cuchumatanes, Guatemala. Photo by Todd Pierson.

Diploglossus bilobatus. This anguid lizard is distributed along

the Atlantic lowlands and premontane slopes of Costa Rica and

northwestern Panama, where it occurs in Lowland Moist and

Wet forests, Premontane Wet Forest and Premontane Rainfor-

est at elevations from near sea level to 1,360 m. We determined

its EVS as 16, placing in the middle of the high vulnerability

category, and its IUCN status is Least Concern. This individual

is from Isla Bopa, province of Bocas del Toro, Panama. Photo

by Abel Batista.

Norops wellbornae: Kohler and Vesely. 2010. Herpe-

tologica 66: 186-207. Resurrection from the syn-

onymy of A. sericeus.

Ctenosaura praeocularis: Hasbun and Kohler. 2009.

Journal of Herpetology 43: 192-204. New species.

Marisora magnacornae: Hedges and Conn. 2012.

Zootaxa 3288: 1-244. New species.

Marisora roatanae: Hedges and Conn. 2012. Zootaxa

3288: 1-244. New species.

Phyllodactylus paralepis: McCranie and Hedges.

2013b. Zootaxa 3694: 51-58. New species.

Sphaerodactylus alphas: McCranie and Hedges.

2013a. Zootaxa 3694: 40-50. New species.

Diasporus igneus. The Fiery Rainfrog is known only from the

eastern and western slopes of Cerro Santiago in the Serrania de

Tabasara in central Panama, where it occurs in Lower Montane

Wet Forest at elevations from 1,699 to 1,815 m. We determined

its EVS as 1 8, placing it in the upper portion of the high vulner-

ability category, but its IUCN status has not been determined.

This individual is from Llano Tugri, in the Comarca Ngobe Bu-

gle. Photo by Abel Batista.

Dipsas articulata. This slug-eating snake is found along the At-

lantic versant from southeastern Nicaragua to western Panama,

where it occurs in Lowland Moist and Wet forests at elevations

from near sea level to 500 m. We assessed its EVS as 15, plac-

ing it in the lower portion of the high vulnerability category,

and its IUCN status is Least Concern. This individual is from

Greytown, department of Rio San Juan, Nicaragua. Photo by

Javier Sunyer.

Sphaerodactylus continentalis: McCranie and Hedg-

es. 2012. Zootaxa 3492: 65-76. Resurrection from

synonymy.

Sphaerodactylus guanajae: McCranie and Hedges.

2012. Zootaxa 3492: 65-76. New species.

Sphaerodactylus leonardovaldesi: McCranie and

Hedges. 2012. Zootaxa 3,492: 65-76. New species.

Sphaerodactylus poindexteri: McCranie and Hedges.

2013. Zootaxa 3694: 40-50. New species.

Ameiva praesignis : Ugueto and Harvey. 2011. Herpe-

tological Monographs 25: 113-170. Elevation to

species level from within A. ameiva.

Cnemidophorus duellmani: McCranie and Hedges.

2013c. Zootaxa 3722: 301-316. New species.

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Amphib. Reptile Conserv.

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Conservation reassessment of Central American herpetofauna

Dermophis occidentalis. This caecilian is endemic to the southern Pacific versant of Costa Rica, where it occurs in Lowland Moist

and Premontane Wet forests at elevations from 50 to 970 m. We determined its EVS as 17, placing it in the middle portion of the

high vulnerability category, and its IUCN status is Data Deficient. This individual is from Rio Piro, province of Puntarenas. Photo

by Eduardo Boza Oviedo.

Cnemidophorus ruatanus: McCranie and Hedges.

2013c. Zootaxa 3722: 301-316. Resurrection from

the synonymy of C. lemniscatus.

Boa imperator: Hynkova et al. 2009. Zoological Sci-

ence 26: 623-631. Elevation to species level from

within B. constrictor.

Dendrophidion apharocybe: Cadle 2012. Bulletin of

the Museum of Comparative Zoology 160: 183—

240. New species.

Dendrophidion crybelum : Cadle 2012. Bulletin of the

Museum of Comparative Zoology 160: 183-240.

New species.

Dendrophidion rnfiterminonmr. Cadle and Savage.

2012. Zootaxa 3513: 1-50. New species.

Tantilla olympia : Townsend et al. 2013a. Journal of

Herpetology 47: 191-200. New species.

Tantilla psittaca : McCranie 2011b. Zootaxa 3037:

37—44. New species.

Omoadiphas cannula: McCranie and Cruz Diaz.

2010. Zootaxa 2690: 53-58. New species.

Sibon merendonensis: Rovito et al. 2012. Zootaxa

3266: 62-68. New species.

Sibon noalamina: Lotzkat et al. 2012. Zootaxa 3485:

26^10. New species.

Sibon perissostichon: Kohler et al. 2010b. Herpeto-

logica 66: 80-85. New species.

Epictia magnamaculata: Adalsteinnsson et al. 2009.

Zootaxa 2244: 1-50. Transfer from genus Eepto-

typhlops and resurrection from the synonymy of E.

goudotii.

Bothriechis guifarroi: Townsend et al. 2013b. Zoo-

Keys 298: 77-105. New species.

Cerrophidion sasai: Jadin et al. 2012. Zoological

Scripta doi: 10.1111/j.l463-6409.2012.00547.x.

New species.

Cerrophidion wilsoni: Jadin et al. 2012. Zoological

Scripta doi: 10.1111/j.l463-6409.2012.00547.x.

New species.

These 92 species represent an increase of 9.7% over

the 952 species listed for Central America by Wilson and

Johnson (2010: Appendix 1).

The following species has undergone synonymization:

Pristimantis educatoris: Ryan et al. 2010a. Journal of

Herpetology 44: 193-200. Synonymized with P.

caryophyllaceus (Batista et al. 2014).

The following 29 species have undergone status changes:

Incilius chompipe: Mendelson et al. 2011. Zootaxa

3138: 1-34. Transfer from genus Crepidophryne.

Incilius epioticus: Mendelson et al. 2011. Zootaxa

3138: 1-34. Transfer from genus Crepidophryne.

Incilius guanacaste: Mendelson et al. 2011. Zootaxa

3138: 1-34. Transfer from genus Crepidophryne.

Andinobates claudiae: Brown et al. 2011. Zootaxa

3083: 1-120. Transfer from genus Ranitomeya.

Andinobates fulguritus: Brown et al. 2011. Zootaxa

3083: 1-120. Transfer from genus Ranitomeya.

Andinobates minutus: Brown et al. 2011. Zootaxa

3083: 1-120. Transfer from genus Ranitomeya.

Agalychnis lemur: Faivovich et al. 2010. Cladistics

26: 227-261. Transfer from genus Hylomantis.

Trachycephalus typhonius: Lavilla et al. 2010. Zoo-

taxa 2671 : 17-30. New name for T. venulosus.

Leptodactylus insularum: Heyer and de Sa. 2011.

Smithsonian Contributions to Zoology 635: i-vii,

1-58. Name E. insularum applied to populations

in Costa Rica and Panama, as well as Colombia,

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Amphib. Reptile Conserv.

24

Johnson et al.

Venezuela, and Trinidad. Called L. bolivianus in

Wilson et al. (2010).

Ctenophryne aterrima : de Sa et al. 2012. BMC Evolu-

tionary Biology 12: 241 (21 pp.). Formerly placed

in the genus Nelsonophryne, now placed in syn-

onymy of Ctenophryne.

Elachistocleis panamensis: de Sa et al. 2012. BMC

Evolutionary Biology 12: 241 (21 pp.). Transfer

from genus Chiasmocleis .

Elachistocleis pearsei: de Sa et al. 2012. BMC Evo-

lutionary Biology 12: 241 (21 pp.). Return to ge-

nus Elachistocleis from Relictivomer. This species

was not considered in Wilson et al. (2010), but was

shown to occur in Panama by Kohler (2011b).

Hypopachus pictiventris: Streicher et al. 2012. Mo-

lecular Phylogenetics and Evolution 64: 645-653.

Tentative transfer from genus Gastrophryne.

Transfer supported by de Sa et al. 2012. BMC Evo-

lutionary Biology 12: 241 (21 pp.).

Hypopachus ustus: Streicher et al. 2012. Molecular

Phylogenetics and Evolution 64: 645-653. Tenta-

tive transfer from genus Gastrophryne. Spelling of

specific epithet corrected by Frost (2013). Transfer

supported by de Sa et al. 2012. BMC Evolutionary

Biology 12: 241 (21 pp.).

Marisora alliacea : Hedges and Conn. 2012. Zootaxa

3288: 1-244. Transfer from the genus Mabuya.

Marisora brachypoda: Hedges and Conn. 2012.

Zootaxa 3288: 1-244. Transfer from the genus

Mabuya.

Marisora unimarginata: Hedges and Conn. 2012.

Zootaxa 3288: 1-244. Transfer from the genus

Mabuya.

Holcosus chaitzami : Harvey et al. 2012. Zootaxa

3459: 1-156. Transfer from the genus Aineiva.

Holcosus festivus: Harvey et al. 2012. Zootaxa 3459:

1-156. Transfer from the genus Ameiva.

Holcosus leptophrys: Harvey et al. 2012. Zootaxa

3459: 1-156. Transfer from the genus Ameiva.

Holcosus quadrilineatus: Harvey et al. 2012. Zootaxa

3459: 1-156. Transfer from the genus Ameiva.

Holcosus undulatus: Harvey et al. 2012. Zootaxa

3459: 1-156. Transfer from the genus Ameiva.

Epictia magnamaculata: Adalsteinnsson et al. 2009.

Zootaxa 2244: 1-50. Resurrection from the syn-

onymy of E. goudotii.

Trichello stoma mac role pis'. Adalsteinnsson et al.

2009. Zootaxa 2244: 1-50. Transfer from the ge-

nus Leptotyphlops. Later established as the type

species of a new leptotyphlopid genus Trilepida by

Hedges (2011).

Amerotyphlops costaricensis: Hedges et al. 2014. Ca-

ribbean Herpetology 49: 1-61. Transfer from the

genus Typhlops.

Amerotyphlops microstomus: Hedges et al. 2014. Ca-

ribbean Herpetology 49: 1-61. Transfer from the

genus Typhlops.

Amerotyphlops stadelmani: Hedges et al. 2014. Ca-

ribbean Herpetology 49: 1-61. Transfer from the

genus Typhlops.

Amerotyphlops tenuis: Hedges et al. 2014. Caribbean

Herpetology 49: 1-61. Transfer from the genus

Typhlops.

Amerotyphlops tycherus: Hedges et al. 2014. Carib-

bean Herpetology 49: 1-61. Transfer from the ge-

nus Typhlops.

Streicher et al. (2014) examined evolutionary relation-

ships among some members of the Craugastor rhodopis

species group and recognized four major clades, includ-

ing one identified as C. occidental^, which required its

movement from the C. mexicanus species series to the C.

rhodopis species group. A clade in eastern Mexico corre-

sponds to C. rhodopis and one on both the Pacific and At-

lantic versants of southeastern Mexico, Guatemala, and

El Salvador to C. loki. Further, they identified a haplo-

type from Volcan San Martin in southern Veracruz, Mex-

ico, which might correspond to a separate evolutionary

lineage. The authors also indicated that, “a small group

of specimens was reported from the northern department

of Cortes in Honduras [that report appeared in McCranie

and Wilson, 2002], but the actual occurrence of C. loki

in Honduras is questionable given the abundance of the

morphologically similar C. time, C. gollmeri, and C. la-

tic eps, in this region ...” The authors left the identity of

the Honduran material and the status of other populations

in the rhodopis species group to future work.

In a broad-scale paper on blindsnake taxonomy,

Hedges et al. (2014) transferred five Central American

typhlopid species from Typhlops to a new genus, Amero-

typhlops. This study, based on morphological and molec-

ular data, supported the recognition of four subfamilies,

of which three were described anew, and contains essen-

tially geographically cohesive groups of genera and spe-

cies. Recognition of the three new subfamilies restricts

the remaining subfamily, the Typhlopinae, to genera and

species in the New World. The authors recognized four

genera, of which Amerotyphlops , Antillotyphlops , and

Cubatyphlops were described as new. Interestingly, the

first of these genera is composed of 14 species distrib-

uted “primarily on the mainland, ranging from eastern

Mexico (Veracruz) to southern South America (Bolivia

and Argentina), and includes a West Indian species, A.

tasymicris in Grenada and the Grenadines” (Hedges et

al. 2014: 44). Five of the 14 species are distributed in

Central America (Appendix 2).

Torres et al. (2013) reported Abronia lythrochila,

formerly a Mexican endemic, from northwestern Guate-

mala, thus adding this species to the Central American

herpetofauna.

Griffin and Powell (2014) reported Tropidodipsas

fasciata, formerly a Mexican endemic, from Guatemala,

thus adding this species to the Central American herpe-

tofauna.

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Amphib. Reptile Conserv.

25

Conservation reassessment of Central American herpetofauna

Ecnomiohyla bailarina. The Golden-eyed Fringe-limbed Tree-

frog is known only from the type locality in extreme southwest-

ern Panama near the border with Colombia (but, see Adden-

dum), where it occurs in Premontane Wet Forest at an elevation

of 750 m. We calculated its EVS as 20, placing it at the upper

end of the high vulnerability category, but its IUCN status has

not been determined. This individual is from the northern slope

of the Jingurudo mountain range in the Comarca Embera-Wou-

naan, in the Darien region. Photo by Abel Batista.

Olson and David (2014) changed the spelling of the

specific name of the single species of Chelonoidis oc-

curring in Central America to carbonarius , given the

ICZN requirement (ICZN 1999; Article 30,2.4) to treat

the generic name as masculine since the original author

(Fitzinger 1835) did not state it explicitly to be feminine.

Some other qualifications concerning the taxonomic

status of certain species to which we adhere are found in

the above Controversial Taxonomic Issues section.

Diversity and Endemism in the Central

American Herpetofauna

Mesoamerica is one of the world’s most important bio-

diversity reservoirs, and Central America contains a sub-

stantial component of that region’s herpetofauna (Wilson

and Johnson 2010). The Central American herpetofauna

presently consists of 1,052 species (319 anurans, 159

salamanders, 15 caecilians, 3 crocodylians, 532 squa-

mates, and 24 turtles; Table 1). Compared to the her-

petofauna of Mexico, which currently consists of 1,252

species (239 anurans, 141 salamanders, 3 caecilians, 3

crocodylians, 818 squamates, and 48 turtles; J. D. John-

son, unpublished data), the number of species in Cen-

tral America is significant given that the area of Mexico

is about 3.75 times larger than that of Central America

(www.cia.gov; accessed 14 December 2013). Compared

to Mexico, Central America also is a haven for anurans,

salamanders, and caecilians, as it contains 1 .3 times more

species. In contrast, however, Mexico contains 1.6 times

more crocodylians, squamates, and turtles than Central

America. Evidently, these differences are related to the

environmental requirements for these two groups of ver-

tebrates, and the variety of ecosystems in the two regions.

Helodenna charlesbogerti. The Motagua Valley Beaded Liz-

ard is restricted to the Motagua Valley in eastern Guatemala,

where it occurs in Lowland Arid and Premontane Dry forests at

elevations from 300 to 900 m. We assessed its EVS as 18, plac-

ing it in the upper portion of the high vulnerability category,

but its IUCN status has not been determined. This individual is

from the Motagua River Valley in Guatemala. Photo by Antonia

Pachmann.

The 493 species amphibians in Central America are

classified in 16 families and 69 genera (Table 1). The

Hylidae contains the most genera (21); the remaining

15 families contain 1-8 genera. The Dendrobatidae and

Plethodontidae contain eight genera each; the remaining

anuran and caecilian families five or fewer (Table 1). The

number of species per family ranges from one (Pipidae

and Rhinophrynidae) to 159 (Plethodontidae). Three

families (Craugastoridae, Hylidae, and Plethodontidae)

contain close to or considerably more than 100 species

each, and collectively total 358 (72.6%) of all the am-

phibian species. The remaining 13 families contain 1-39

species (the latter number is for the Bufonidae). In total,

there are 13 families and 57 genera of anurans, one fam-

ily and eight genera of salamanders, and two families and

four genera of caecilians.

The 559 species of crocodylians, squamates, and tur-

tles in Central America are classified in 42 families and

145 genera (Table 1). The Colubridae and Dipsadidae are

the largest, with 24 and 35 genera, respectively; the re-

maining families contain 1-8 genera. Two families con-

tain eight genera (Gymnophthalmidae and Viperidae),

and the others contain five or fewer (Table 1). The num-

ber of species per family ranges from one (seven fami-

lies) to 144 (Dipsadidae). Two families (Dactyloidae and

Dipsadidae) contain close to or considerably more than

100 species (Table 1), collectively 239 (42.8%) of all

the squamate species. The remaining 40 families contain

1-32 species (the latter number is for Viperidae). In total,

there are two families and two genera of crocodylians,

nine families and 14 genera of turtles, and 31 families

and 129 genera of squamates.

The herpetofauna of Central America also is charac-

terized by a high degree of endemism (Table 1). Of the

493 species of anurans, salamanders, and caecilians in

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Amphib. Reptile Conserv.

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Johnson et al.

Table 1 . Family composition and endemicity of the Central American herpetofauna.

Families

Genera

Species

Endemic Species

Percentage of Endemicity

Aromobatidae

2

3

2

66.7

Bufonidae

4

39

23

59.0

Centrolenidae

5

14

3

21.4

Craugastoridae

3

101

77

76.2

Dendrobatidae

8

19

12

63.2

Eleutherodactylidae

2

11

6

54.5

Hemiphractidae

2

3

0

Hylidae

21

98

53

54.1

Leptodactylidae

3

9

1

11.1

Microhylidae

4

9

1

11.1

Pipidae

1

100

Ranidae

1

11

5

45.5

Rhinophrynidae

1

0

Total Anurans

57

319

184

57.5

Plethodontidae

8

159

133

83.6

Total Salamanders

8

159

133

83.6

Caeciliidae

2

7

3

42.9

Dermophiidae

2

8

4

50.0

Total Caecilians

4

15

7

46.7

Total Amphibians

69

493

324

65.7

Alligatoridae

1

0

Crocodylidae

1

2

0

Total Crocodylians

2

3

0

Amphisbaenidae

1

2

0

Anguidae

5

28

22

78.6

Corytophanidae

3

9

0

Dactyloidae

3

95

67

70.5

Eublepharidae

1

2

0

Gymnophthalmidae

8

14

5

35.7

Helodermatidae

1

2

1

50.0

Hoplocercidae

2

0

Iguanidae

2

11

7

63.6

Mabuyidae

1

5

4

80.0

Phrynosomatidae

2

17

2

11.8

Phyllodactylidae

2

5

3

60.0

Polychrotidae

1

0

Scincidae

2

3

0

Sphaerodactylidae

4

19

10

52.6

Sphenomorphidae

1

4

1

25.0

Teiidae

4

12

4

33.3

Xantusiidae

1

4

1

25.0

Xenosauridae

1

0

Anomalepididae

3

1

33.3

Boidae

3

4

0

Charinidae

1

2

0

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Conservation reassessment of Central American herpetofauna

Table 1 (continued). Family composition and endemicity of the Central American herpetofauna.

Families

Genera

Species

Endemic Species

Percentage of Endemicity

Colubridae

24

74

26

35.1

Dipsadidae

35

144

78

54.2

Elapidae

2

18

8

44.4

Leptotyphlopidae

2

5

1

20.0

Loxocemidae

1

0

Natricidae

2

5

0

Sibynophiidae

1

2

0

Tropidophiidae

1

0

Typhlopidae

1

5

3

60.0

Viperidae

8

32

15

46.9

Total Squamates

129

532

259

48.7

Chelonndae

4

5

0

Chelydridae

1

2

0

Dermatemydidae

1

0

Dermochelyidae

1

0

Emydidae

1

2

0

Geoemydidae

1

5

1

20.0

Kinosternidae

1

4

1

25.0

Staurotypidae

2

3

0

Testudinidae

1

0

Total Turtles

14

24

2

8.3

Total “Reptiles”

145

559

261

46.7

Total Herpetofauna

214

1,052

585

55.6

this region, 324 (65.7%) are endemic, and of the 559 spe-

cies of crocodylians, squamates, and turtles, 261 (46.7%)

are endemic. The percentage of endemicity for the en-

tire herpetofauna is 55.6%. These figures are somewhat

comparable to those for the Mexican herpetofauna (J.D.

Johnson, unpublished data). Of the 383 Mexican amphib-

ian species, 258 (67.4%) are endemic, and of the 869 spe-

cies of crocodylians, squamates, and turtles, 499 (57.4%)

are endemic. The percentage of endemicity for the entire

herpetofauna is 60.5% (J.D. Johnson, unpublished data).

Among the Central American amphibians, the per-

centage of endemicity at the family level ranges from

zero (Hemiphractidae and Rhinophrynidae) to 100 (Pipi-

dae). Interestingly, each of these anuran families contains

1-3 species in Central America. The largest number of

endemic species is in the family Plethodontidae (133);

its percentage of endemicity is 83.6. The Bufonidae (23),

Craugastoridae (77), and Hylidae (53) also contain siz-

able numbers of endemic species. Collectively, these

four families contain 286 (88.3%) of all the amphibian

endemic species.

Among the crocodylians, squamates, and turtles, the

percentage of endemicity at the family level ranges from

zero (in 21 families) to 80.0% (Mabuyidae). As with am-

phibians, the 21 families with no endemics contain rela-

tively few species (nine or fewer). The largest number

of endemic squamates is in the family Dipsadidae (78),

with the next largest being the Dactyloidae (67). The next

most sizable numbers of endemic species are in the fami-

lies Colubridae (26) and Viperidae (15). Collectively,

these four families contain 186 (71.8%) of all the squa-

mate endemic species.

In summary, four amphibian and four squamate fami-

lies contain the largest numbers of endemic species in

Central America (472; 81.1%) of the 585 endemic spe-

cies known from this region (Table 1). In total, these

eight families contain 742 species, of which 63.6% are

endemic to Central America (Table 1). With additional

exploration and systematic research, the number and

proportion of endemic species in Central America should

continue to rise.

IUCN Red List Assessment of the Central

American Herpetofauna

In response to the emerging picture of global amphib-

ian population decline, the IUCN began a conservation

assessment of the world’s amphibians (see Stuart et al.

2004). Consequently, in 2002, a workshop to assess the

Mesoamerican amphibians was held at the La Selva

Biological Station in Costa Rica, followed by one in

Jalisco, Mexico, to assess the crocodylians, squamates,

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Johnson et al.

Table 2. IUCN Red List categorizations for Central American herpetofaunal families.

Families

Number

of

Species

IUCN Red List categorizations

Extinct

Critically

Endan-

gered

Endan-

gered

Vulner-

able

Near

Threat-

ened

Least

Concern

Data

Deficient

Not

Evaluated

Aromobatidae

3

—

1

—

2

Bufonidae

39

1

9

7

3

1

12

4

2

Centrolenidae

14

—

1

12

1

—

Craugastoridae

101

2

23

16

10

8

26

14

2

Dendrobatidae

19

—

3

1

8

5

1

Eleutherodactylidae

11

—

3

—

5

1

2

Hemiphractidae

3

—

1

—

1

—

Hylidae

98

—

33

14

5

35

3

Leptodactylidae

9

—

1

—

8

—

Microhylidae

9

—

1

—

8

—

Pipidae

1

—

1

—

Ranidae

11

—

3

1

6

—

1

Rhinophrynidae

1

—

1

—

Total Anurans

319

3

66

42

26

18

123

28

13

Plethodontidae

159

—

25

33

17

8

19

18

39

Total Salamanders

159

—

25

33

17

8

19

18

39

Caeciliidae

7

—

3

4

—

Dermophiidae

8

—

1

—

2

5

—

Total Caecilians

15

—

1

—

5

9

—

Total Amphibians

493

3

91

75

44

26

147

55

52

Alligatoridae

1

—

1

—

Crocodylidae

2

—

1

—

1

—

Total Crocodylians

3

—

1

—

2

—

Amphisbaenidae

2

—

1

—

1

Anguidae

28

—

2

8

2

6

5

3

Corytophanidae

9

—

5

—

4

Dactyloidae

95

—

3

1

—

4

3

84

Eublepharidae

2

—

2

—

Gymnophthalmidae

14

—

1

4

—

9

Helodermatidae

2

—

2

Hoplocercidae

2

—

2

Iguanidae

11

—

1

4

—

1

3

Mabuyidae

5

—

1

—

3

1

—

Phrynosomatidae

17

—

17

—

Phyllodactylidae

5

—

1

—

1

—

3

Polychrotidae

1

—

1

Scincidae

3

—

3

—

Sphaerodactylidae

19

—

10

—

9

Sphenomorphidae

4

—

2

1

Teiidae

12

—

6

1

5

Xantusiidae

4

—

1

3

—

Xenosauridae

1

—

1

—

Anomalepididae

3

—

2

1

Boidae

4

—

4

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Amphib. Reptile Conserv.

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Conservation reassessment of Central American herpetofauna

Table 2 (continued). iUCN Red List categorizations for Central American herpetofaunal families.

Families

Number

of

Species

IUCN Red List categorizations

Extinct

Critically

Endan-

gered

Endan-

gered

Vulner-

able

Near

Threat-

ened

Least

Concern

Data

Deficient

Not

Evaluated

Charinidae

2

—

2

Colubridae

74

—

1

2

3

—

37

5

26

Dipsadidae

144

—

7

11

5

8

66

15

32

Elapidae

18

—

1

—

12

—

5

Leptotyphlopidae

5

—

1

—

4

Loxocemidae

1

—

1

—

Natricidae

5

—

5

—

Sibynophiidae

2

—

1

—

1

Tropidophiidae

1

—

1

Typhlopidae

5

—

1

—

3

—

1

Viperidae

32

—

2

1

12

1

15

Total Squamates

532

—

13

30

15

14

206

35

219

Cheloniidae

5

—

2

1

—

Chelydridae

2

—

1

—

1

Dermatemydidae

1

—

1

—

Dermochelyidae

1

—

1

—

Emydidae

2

—

2

Geoemydidae

5

—

3

—

2

Kinosternidae

4

—

1

—

2

Staurotypidae

3

—

3

—

Testudinidae

1

—

1

Total Turtles

24

—

4

2

3

7

—

8

Total “Reptiles”

559

—

17

32

19

21

208

35

227

Total Herpetofauna

1,052

3

108

107

63

47

355

90

279

and turtles of that country. Several years later, in 2012,

a workshop to assess the squamates of Central America

was held at Parque Nacional Palo Verde in Costa Rica.

The results of the first two workshops appeared on the

IUCN Red List website, but to date those for the third

remain incomplete. Wilson et al. (2013a, b) presented an

overview and conclusions of these assessments for the

Mexican herpetofauna.

We accessed the IUCN website (www.iucnredlist.org)

to summarize the present situation for Central American

amphibians (Table 2). The data in this table are some-

what more complete than for crocodylians, squamates,

and turtles, given that the Global Reptile Assessment

still is underway. Nonetheless, of 493 species of Central

American amphibians, 52 species (10.5%) have not been

evaluated as of this writing, so we placed them in the

NE (Not Evaluated) category. The remaining categories

are: Extinct (EX, 3 [0.6%]); Critically Endangered (CR,

91 [18.5%]); Endangered (EN, 75 [15.2%]); Vulnerable

(VU, 44 [8.9%]); Near Threatened (NT, 26 [5.3%]); Least

Concern (LC, 147 [29.8%]); and Data Deficient (DD, 55

[11.2%]). A total of 210 species (42.6%), therefore, have

been assessed in one of the three threat categories (CR,

EN, or VU), which is slightly more than 10% higher than

what was reported for these categories on a global scale

(32.3%) by Stuart et al. (2010). If the EX and DD spe-

cies are added to those in the threat categories, then 268

(54.4%) species are extinct, threatened with extinction,

or too poorly known to allow for an assessment; these

results are similar to those reported for the global situa-

tion (EX+CR+EN+VU+DD = 3,181 [55.4%]; Stuart et

al. 2010). This percentage, however, is about 10 points

lower than that reported for the Mexican amphibians

(Wilson et al. 2013b).

The families Craugastoridae (49 of 101 species;

48.5%), Hylidae (52 of 98 species; 53.1%), and Plethod-

ontidae (75 of 159 species; 47.2%) contain the greatest

number and proportion of threatened species. For the

salamanders, if the numbers of DD and NE species are

added to those considered threatened (18+39+75 = 132),

then 83.0% of the 159 Central American species are

threatened, poorly known, or have not been evaluated.

Collectively, the 358 species in the three largest families

comprise 72.6% of the amphibian taxa in Central Ameri-

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Amphib. Reptile Conserv.

30

Johnson et al.

ca, and the 176 threatened species in these families make

up 83.8% of the 210 total. A similar proportion is seen

among the Mexican amphibians (88.6% of 211 totals).

As startling as the statistics for amphibian population

decline are on a global scale (Stuart et al. 2010), on a re-

gional scale for Central America they are more alarming.

According to the IUCN criteria, about four out of every

10 species of amphibians are judged as threatened, and

more than one-half of those are threatened or too poorly

known to allow for an assessment, which is the case for

the most speciose families in the region. Two factors are

expected to keep increasing the severity of this decline,

even without considering the rate of accelerating envi-

ronmental deterioration. The first is that new species

will continue to be described, as indicated above in the

discussion on taxonomic changes since the publication

of Wilson et al. (2010). The other factor is that advanc-

ing molecular studies, especially on broadly distributed

taxa, will continue to reveal the presence of new species.

Both of these factors will increase the number of threat-

ened taxa. As an example, Ruane et al. (2014) studied

the molecular systematics of Lampropeltis triangulum , a

species that for many decades was considered one of the

world’s most broadly distributed terrestrial snakes (Wil-

liams 1988). These authors recognized seven species in

what previously was considered a single species-level

taxon, and noted that additional species in this complex

likely will be recognized in the future.

Critique of the IUCN Assessment

In conservation reassessments for the Mexican herpeto-

fauna, we criticized the IUCN system of categorization

and provided distinctions between this system and the

EVS (Wilson et al. 2013a, b). Alvarado-Diaz et al. (2013)

also criticized this system. The principal criticisms levied

by these authors are as follows:

1. Using the IUCN system of conservation assessment

is expensive and time-consuming. Stuart et al. (2010)

provided a figure of $534.12 for the average cost of cre-

ating an IUCN threat assessment for a single species.

If this figure were applied to the 1,052 species making

up the Central American herpetofauna, the total ex-

penditure would be $561,894.24. In comparison, costs

for our EVS assessments were negligible because they

were accomplished using the resources of the Inter-

net and our own volunteered time. Creating the IUCN

Global Amphibian Assessment, of which the results

appeared in 2004 (Stuart et al. 2004) involved a num-

ber of years. For example, one of us (LDW) attended

the Mesoamerican Amphibian Workshop undertaken

at the La Selva Biological Station in Costa Rica in No-

vember 2002, so a period of close to two years elapsed

before the global results were published (Stuart et al.

2004). Another example is that the complete results

of the Central American Reptile Workshop, attended

by two of us (JDJ and LDW), have not appeared two

years and two months since this workshop was con-

ducted at Palo Verde National Park in Costa Rica in

May of 2012 (as of 1 March 2015). The delay pri-

marily has been caused because evaluations for most

of the anoles have not been completed, and because

evaluations for a sizable number of species that oc-

cur in both Central America and South America will

not be available until all of the relevant workshops

for the latter region are completed. In contrast, we be-

gan working on the present paper in early October,

2013. We completed most of our EVS assessment of

the Central American herpetofauna by the early por-

tion of January, 2014, but the publication of this pa-

per was delayed because we needed to wait until the

entire results of the Palo Verde Workshop appeared at

the IUCN Red List website (but see above). Accord-

ingly, we consider it pertinent to quote the “important

note” or proviso indicated on the Overview paper at

the amphibians.org website, as follows: “Given the

current quality control requirements needed for con-

servation assessments to be published on the IUCN

Red List, and our very limited human resources, we

are unable to process large numbers of assessments

at this time. Country-level global reassessments may

be possible if requests come with the funding and re-

sources necessary to conduct such reassessments, or

if the herpetological community of the country or re-

gion in question is willing to take over stewardship of

its global assessments through its respective regional/

national working group.” Thus, the expense for such

IUCN assessments has overwhelmed the ability of

this organization to continue undertaking this work.

2. New herpetofaunal taxa are described more rapidly

than the IUCN procedures can provide a conservation

assessment. As noted in the previous section, 52 spe-

cies of amphibians (13 anurans and 39 salamanders)

remain unevaluated by the IUCN, which is 10.5% of

the 493 species known from Central America as of

this writing. Comparable figures are not available for

the remainder of the herpetofauna, since the Global

Reptile Assessment is ongoing, but we can state that

32 species of squamates (lizards and snakes) have

been described since the publication of Wilson and

Johnson (2010). This figure represents 6.0% of the

532 species of squamates now known from Central

America. The data in Table 2 indicate that 227 species

of crocodylians, squamates, and turtles (40.6% of the

total of 559 species) have not been evaluated. Given

the provisos indicated in the above paragraph and the

consequences indicated, a much more rapid and cost-

effective mode of conservation assessment is needed,

not only for keeping up with the advances of system-

atic knowledge, but more importantly because of the

increasing rate of environmental deterioration.

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Conservation reassessment of Central American herpetofauna

3. Once new herpetofaunal species are incorporated into

the IUCN Red List, often they are placed in the Data

Deficient category due to an expected lack of initial

information on their population status. In particular,

this situation occurs with taxa described from a sin-

gle specimen and/or a single locality. Species in this

category were termed “threat species in disguise” by

Wilson et al. (2013b), because of the likelihood that

such species, once evaluated, would fall into one of

the three threat categories. One of our recommenda-

tions deals with this issue.

4. Typically, large numbers of taxa are assigned to the

Least Concern (LC) category, described by Wilson

et al. (2013b) as a “dumping ground” for species that

might require “a more discerning look that would

demonstrate that many of these species should be par-

titioned into IUCN categories other than LC,” such

as the three threat categories and the Near Threatened

one. This opinion was expressed after the authors ex-

amined the relationship between the IUCN catego-

rizations and the EVS assessments for Mexican am-

phibians, and is corroborated here by the assessment

Table 3. Environmental Vulnerability Scores for Central American herpetofaunal species, arranged by family. Shaded area to the left encompasses low vulner-

ability scores, and to the right high vulnerability scores.

Families

Number

of

species

Environmental Vulnerability Scores

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

Aromabatidae

3

—

1

—

1

—

1

—

Bufonidae

39

1

—

1

2

3

1

3

8

9

4

2

1

—

Centrolenidae

14

—

1

2

4

1

—

1

2

—

Craugastoridae

101

—

1

—

1

6

2

6

15

35

17

18

—

Dendrobatidae

19

1

6

3

—

Eleutherodactylidae

11

1

—

3

1

2

3

—

Hemiphractidae

3

1

2

—

Hylidae

98

1

2

—

3

4

2

6

9

11

18

8

1

2

4

3

6

Leptodactylidae

9

—

1

—

1

—

3

1

—

Microhylidae

9

—

1

—

2

—

1

3

—

1

—

Pipidae

1

Ranidae

11

1

—

1

—

1

2

1

—

3

—

1

—

Rhinophrynidae

1

Total Anurans

319

3

2

5

11

10

11

18

34

30

37

39

50

26

29

3

6

Total Anuran %

—

0.9

0.6

1.6

3.4

3.1

3.4

5.6

10.7

9.4

11.6

12.2

15.7

8.2

9.1

0.9

1.9

Plethodontidae

159

—

1

—

2

6

4

17

34

35

57

—

Total Salamanders

159

—

1

—

2

6

4

17

34

35

57

—

Total Salamander %

—

0.6

—

1.3

3.8

2.5

10.7

21.4

22.0

35.8

—

Caeciliidae

7

2

1

—

2

—

Dermophiidae

8

—

1

—

2

1

—

1

2

—

Total Caecilians

15

—

1

—

2

1

2

3

2

—

Total Caecilian %

—

6.7

—

13.3

6.7

13.3

20.0

13.3

—

Total Amphibians

493

3

2

6

12

11

20

36

38

42

58

87

63

88

5

6

Total Amphibian %

—

0.6

0.4

1.2

2.4

2.2

4.1

7.3

7.7

8.5

11.8

17.6

12.8

17.8

1.0

1.2

Alligatoridae

1

Crocodylidae

2

1

—

Total Crocodylians

3

1

—

1

—

Total Crocodylian %

33.3

—

33.3

—

Amphisbaenidae

2

1

Anguidae

28

—

1

—

1

2

7

9

3

5

—

Corytophanidae

9

—

1

—

2

1

2

—

2

—

1

—

Dactyloidae

95

—

2

4

1

4

12

11

23

13

22

—

Eublepharidae

2

—

1

—

1

—

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Johnson et al.

Table 3 (continued). Environmental Vulnerability Scores for Central American herpetofaunal species, arranged by family. Shaded area to the left encom-

passes low vulnerability scores, and to the right high vulnerability scores.

Families

Number

of

species

Environmental Vulnerability Scores

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

Gymnophthalmidae

14

—

1

—

1

5

3

2

—

Helodermatidae

2

—

1

—

1

—

Hoplocercidae

2

—

1

—

1

—

Iguanidae

11

—

1

—

1

—

1

—

1

—

3

4

—

Mabuyidae

5

1

2

1

—

Phrynosomatidae

17

—

1

—

3

1

5

2

—

1

—

Phyllodactylidae

5

—

2

—

1

2

—

Polychrotidae

1

Scincidae

3

1

—

Sphaerodactylidae

19

—

1

—

1

2

4

2

4

—

Sphenomorphidae

4

—

1

—

1

—

1

—

Teiidae

12

—

1

—

1

—

1

—

1

2

3

—

Xantusiidae

4

—

2

—

2

—

Xenosauridae

1

Total Lizards

236

—

1

2

5

9

11

7

6

15

26

29

45

33

34

9

4

—

Anomalepididae

3

—

1

—

1

Boidae

4

—

2

—

1

—

1

—

Charinidae

2

—

1

—

1

—

Colubridae

74

—

1

6

5

2

5

3

5

10

8

13

2

12

2

—

Dipsadidae

144

—

2

1

3

7

10

5

17

24

23

27

2

—

Elapidae

17

—

2

1

—

2

—

4

2

4

2

—

Leptotyphlopidae

5

—

2

—

1

—

2

—

Loxocemidae

1

Natricidae

5

—

1

2

1

—

1

—

Sibynophiidae

2

1

Tropidophiidae

1

Typhlopidae

5

2

—

1

—

Viperidae

32

—

1

2

1

3

6

3

1

—

Total Snakes

295

—

2

4

9

8

19

18

35

31

41

35

47

14

5

1

—

Total Squamates

531

—

2

5

11

13

17

30

25

24

50

57

70

80

48

14

5

—

Total Squamate %

—

0.4

0.9

2.1

2.4

3.2

5.6

4.7

4.5

9.4

10.7

13.2

15.1

9.0

2.6

0.9

—

Chelydridae

2

—

1

—

1

—

Dermatemydidae

1

Emydidae

2

1

—

Geoemydidae

5

—

1

—

1

—

Kinosternidae

4

—

2

—

1

—

1

—

Staurotypidae

3

1

2

—

Testudinidae

1

Total Turtles

18

—

3

—

2

1

2

3

1

2

3

—

1

—

Total Turtle %

—

16.7

—

11.1

5.5

11.1

16.7

5.5

11.1

16.7

—

5.5

—

Total “Reptiles”

552

—

2

5

11

13

20

30

25

26

51

60

74

81

83

51

14

6

—

Total “Reptile” %

—

0.4

0.9

2.0

2.4

3.6

5.4

4.5

4.7

9.2

10.9

13.4

14.7

15.0

9.2

2.5

1.1

—

Total Herpetofauna

1,045

3

5

7

13

19

32

41

36

46

87

98

116

139

170

114

102

11

6

Total Herpetofauna %

—

0.3

0.5

0.7

1.2

1.8

3.1

3.9

3.4

4.4

8.3

9.4

11.1

13.3

16.3

10.9

9.8

1.1

0.6

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Conservation reassessment of Central American herpetofauna

Imantodes phantasma. This blunt-headed treesnake is distrib-

uted in the Darien region of eastern Panama, where it occurs in

Premontane Wet Forest at elevations from 1,000 to 1,100 m. We

established its EVS as 16, placing in the middle portion of the

high vulnerability category, and its IUCN status is Data Defi-

cient. This individual is from the Serrania de Pirre, province of

Darien, Panama. Photo by Abel Batista.

Isthmohyla picadoi. This treefrog occurs in the Cordillera

Central and Cordillera de Talamanca of Costa Rica and west-

ern Panama, where it occurs in Lower Montane and Montane

rainforests at elevations from 1,920 to 2,770 m. We assessed

its EVS as 19, placing it in the upper portion of the high vulner-

ability category, and its IUCN status is Near Threatened. This

individual is from near Tres Colinas, Parque Internacional La

Amistad, Cordillera de Talamanca, province of Puntarenas,

Costa Rica. Photo by Sean Michael Rovito.

for Central American species we provide in a follow-

ing section.

Given our opinion about the nature of the IUCN sys-

tem of conservation assessment, as in our assessments

for the Mexican herpetofauna, we employ the EVS mea-

sure to conduct our own assessment of the conservation

status of the Central American herpetofauna.

EVS for the Central American Herpetofauna

In our prior conservation reassessments of the mem-

bers of the Mexican herpetofauna (Wilson et al. 2013a,

b), we specified a number of advantages for using the

Isthmohyla zeteki. This treefrog is distributed from central

Costa Rica to western Panama, where it occurs in Premontane

Wet Forest and Rainforest on into Lower Montane Rainforest at

elevations from 1,200 to 1,804 m. We evaluated its EVS as 13,

placing it at the upper end of the middle vulnerability category,

and its IUCN status is Critically Endangered. This individual

is from the Cordillera de Tilaran, province of Alajuela, Costa

Rica. Photo by Brian Kubicki.

Kinosternon angustipons. The Narrow-bridged Mud Turtle is

distributed along the Atlantic versant from southeastern Nicara-

gua to northwestern Panama, where it occurs in Tropical Moist

Forest at elevations from near sea level to 260 m. We estimated

its EVS as 16 in the middle portion of the high vulnerability

category, and its IUCN status is Vulnerable. This individual is

from the Rio Papaturro, Los Guatuzos, department of Rio San

Juan, Nicaragua. Photo by Javier Sunyer.

EVS system. Based on the information in Wilson et al.

(2013b: 107), we summarize these advantages below.

1 . “This measure can be applied as soon as a species is

described, because the information necessary for its

application generally is known at that point.” If the

information is not entirely known (e.g., about amphib-

ian reproductive mode), it can be estimated based on

the phylogenetic relationships of the newly described

species.

2. “The calculation of the EVS is an economical under-

taking and does not require expensive, grant-sup-

ported workshops, such as those held in connection

with the Global Reptile Assessment sponsored by the

Amphib. Reptile Conserv.

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Johnson et al.

IUCN.” Given that any conservation assessment is

only an estimate of an organism’s status in nature, it

will always remain subject to modification based on

additions to our knowledge. As an extreme example,

conservation biologists might reach a point where

they feel justified to report that a certain species is ex-

tinct (i.e., the creature no longer is in existence). This

theoretical definition of the term can be problematical,

however, because it can be difficult or impossible to

determine if any individuals of the species remain in

nature. Thus, a practical definition of the term extinc-

tion can be adopted to mean that no individuals of the

species have been found by anyone qualified to make

such a determination. Interestingly, on 30 December

2013 the AmphibiaWeb site indicated that an anuran

from Sri Lanka ( Pseudophilautus hypomelas ) that had

been declared extinct was rediscovered (Wickramas-

inghe et al. 2013). The authors suggested that its status

be changed to Critically Endangered “under the IUCN

Red List Categories and Criteria because of the extent

of occurrence (EOO) is less than 1 00 km 2 , the area of

occupancy (AOO) is less than 10 km 2 , and is recorded

from a single location. The habitat is under severe

anthropogenic activities such as overexploitation of

natural resources for tea cultivation, forest fragmenta-

tion, use of agrochemicals, soil erosion, inadequately

planned constructions and illegal constructions, mini-

hydro power plants, forest die back, and discharge of

pollutants to the environment.” This familiar litany of

reasons for organismic endangerment also applies to

environments in Central America, where similar re-

ports have been published (Abarca et al. 2010).

3. “The EVS is predictive, because it provides a measure

of succeptibility to anthropogenic pressure, and can

pinpoint taxa in need of immediate attention and con-

tinuing scrutiny.” We provide an example of two re-

cently described species of the hylid genus Ecnomio-

hyla. Batista et al. (2014) described E. bailarina and

E. veraguensis from southeastern and west-central

Panama, respectively. The authors noted that, “the se-

cretive habits of Ecnomiohyla bailarina , as with other

Ecnomiohyla species, make it difficult to obtain an as-

sessment of its population size. Considering that the

status of the E. bailarina population is unknown, the

data deficient (DD) criterion, according to the IUCN

. . . seems appropriate for this species, until data on its

population trend become available. Moreover, due to

the fact that E. bailarina and E. thysanota occur in a

region affected by social problems and political con-

flicts along the border between Panama and Colombia,

it is unlikely that there will be sufficient opportunity

to visit the region to assess population sizes.” With

respect to their other new species, Batista et al. (2014)

indicated that, “since Ecnomiohyla veraguensis oc-

curs along with relatively widely distributed species,

it is not suspected to be endemic to the Cerro Negro

surroundings [the vicinity of the type locality]. How-

ever, all species in the genus are known to be very sus-

ceptible to habitat degradation and thus most are listed

under a threatened category ... So it is very likely that

E. veraguensis also will qualify for a threatened cat-

egory as soon as additional data are available.” These

authors implied that this species should be given a

Data Deficient status based on the same sort of rea-

soning used for E. bailarina. We reviewed the Batista

et al. (2014) paper and determined the EVS for the

two species based on the information provided in the

original descriptions. As a result, the EVS for both

species are the highest possible (Appendix 1), i.e., 20

(6+8+6), based on being known only from the type

locality in a single vegetation formation and having

presumably a reproductive mode like other species of

Ecnomiohyla (i.e., with eggs and tadpoles in water-

filled tree cavities). This EVS is the same as that for

the recently described and famously endangered Ec-

nomiohyla rabborum (Appendix 1; Mendelson 2011).

As a consequence of our ability to calculate EVS for

both of the newest Ecnomiohyla species, we can bring

attention to their plight and their conservation status

to the point that they can be used as flagship species,

along with E. rabborum , to publicize the issues sur-

rounding the conservation of the Panamanian herpeto-

fauna as a whole (also see Jaramillo et al. 2010).

4. “Finally, this measure is simple to calculate and does

not ‘penalize’ species that are poorly known.” In our

opinion, this penalizing comes when a species is des-

ignated as Data Deficient, because it then enters into

a conservation status limbo until and unless informa-

tion is available that will allow for the application of

another IUCN category to be applied (most likely

one of the three threat categories). For this reason, as

previously discussed, we consider the DD species as

“threat species in disguise.” Given the pace at which

organismic endangerment proceeds and the survival

chances for many species, obviously they cannot af-

ford such delays.

We calculated the EVS scores for each of the 1,045

species of amphibians, crocodylians, squamates, and

turtles in Central America to which it can be applied (see

Appendix 1). We placed these data alongside those for

the IUCN categorizations we obtained from the IUCN

Red List website (www.iucnredlist.org) and used the des-

ignation NE for those species presently not evaluated by

the IUCN.

Theoretically, the EVS scores can range from 3 to 20

for amphibians, crocodylians, squamates, and turtles. A

score of 3 would be assigned to broadly distributed spe-

cies both inside and outside of Central America, which

occurs in eight or more forest formations, and, if an am-

phibian has both its eggs and tadpoles in large to small

bodies of lentic or lotic water or, if a squamate, if a spe-

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Conservation reassessment of Central American herpetofauna

Lepidophyma reticulatum. This night lizard is distributed on the

Pacific versant of Costa Rica and western Panama, where it oc-

curs in Lowland Moist and Wet and Premontane Moist and Wet

forests at elevations from 10 to 1,250 m. We estimated its EVS

as 13, placing it at the upper end of the medium vulnerability

category, and its 1UCN status is Least Concern. This individual

is from Porton, province of Chiriqui, Panama. Photo by Abel

Batista.

cies is fossorial and thus usually escapes human notice.

The amphibian species receiving an EVS score of 3 are

the ranid Lithobates forreri, the bufonid Rhinella marina ,

and the hylid Smilisca baudinii (Appendix 1). We did not

assign this score to any crocodylian, squamate, or turtle.

At the other extreme, an EVS score of 20 would be as-

signed to a species known only from the vicinity of its

type locality, is restricted to a single forest fonnation,

and, if an amphibian, has both its eggs and tadpoles in

water-retaining arboreal bromeliads or water-filled tree

cavities, or, if a crocodylian, squamate, or turtle is com-

mercially or non-commercially exploited for hides, meat,

eggs and/or the pet trade. We assigned an EVS score of

20 to six species of hylid anurans, including four in the

genus Ecnomiohyla, one in Isthmohyla, and one in Pty-

chohyla (Appendix 1). As with the lowest possible score,

no crocodylian, squamate, or turtle received the highest

possible score. The remaining EVS scores ranged from 4

to 19. We provide a summary of the EVS scores for the

Central American herpetofaunal species in Table 3. The

EVS range falls into the following three categories: low

(3-9), medium (10-13), and high (14-19).

The range and mean EVS scores for the major herpe-

tofaunal groups are as follows: anurans = 3-20 (13.8);

caecilians = 7-19 (15.4); and salamanders = 8-18 (16.5);

crocodylians = 13-16 (14.3); lizards = 5-19 (14.0);

snakes = 4-19 (12.8); and turtles = 8-19 (13.5). We

found that on average among amphibians, salamanders

are more susceptible to environmental deterioration, and

anurans are less susceptible than caecilians; among the

remainder of the herpetofauna, crocodylians are the most

susceptible and snakes the least susceptible, with turtles

and lizards falling in between. The average scores fell

into the upper portion of the medium category (anurans,

snakes, and turtles), and the lower portion of the high cat-

egory (caecilians, salamanders, and lizards). We found

the average EVS scores for all amphibian species as 14.7,

a value near the lower end of the high range of vulnera-

Lithobates miadis. This leopard frog is endemic to Little Corn

Island off the Caribbean coast of Nicaragua, whose area con-

sists of only 3 km2; it occurs in Lowland Moist Lorest and

breeds in permanent ponds. We established its EVS as 15, plac-

ing it in the lower portion of the high vulnerability category,

and its IUCN status is Vulnerable. This individual is from the

Region Autonoma del Atlantico Sur. Photo by Javier Sunyer.

bility, and that for crocodylians, squamates, and turtles as

13.3, a value slightly above the upper end of the medium

range of vulnerability. Based on these average EVS val-

ues, amphibians are somewhat more vulnerable to envi-

ronmental degradation than the rest of the herpetofauna.

Our results show an EVS score of 16, near the middle

portion of the high vulnerability category, in the highest

percentage (15.6) of anuran species, and an EVS score

of 1 8, near the upper end of the high vulnerability cat-

egory, in the highest percentage (35.8) of the salamander

species. For caecilians, we found the same percentage of

species (13.3) with EVS values ranging from 13 to 19.

When organized by EVS category, the lowest number

of species of amphibians (39 [7.9%]) fell into the low

category, an intermediate number (105 [21.3%]) into the

medium category, and the highest number (349 [70.8%])

into the high category. These figures are more alarming

than those reported for the Mexican amphibian fauna;

Wilson et al. (2013b) noted that of the 378 total taxa,

50 (13.2%) fell into the low vulnerability category, 106

(28.0%) into the medium category, and 222 (58.7%) into

the high category.

We discovered that the EVS scores for crocodylians

are too few and too scattered to confirm a pattern. With

squamates, however, we found EVS scores of 15 and 16,

in the lower portion of the high vulnerability category,

in the highest percentage (14.9%) of the species. Over-

all, the frequency of EVS values for all crocodylians,

squamates, and turtles increased to peak at the value of

16, and decreased steeply thereafter. When organized

by EVS category, as with amphibians we found that the

lowest number of species (81 [14.7%]) fell into the low

category, an intermediate number (162 [29.3%]) into the

medium category, and the highest number (309 [56.0%])

into the high category. These statistics differ only slight-

ly from those reported for Mexican crocodylians, squa-

mates, and turtles by Wilson et al. (2013a), who indicated

that of the 841 total taxa that could be scored, 99 (11.8%)

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Johnson et al.

fell into the low vulnerability category, 272 (32.3%) into

the medium category, and 470 (55.9%) into the high cat-

egory.

For the total Central American herpetofauna, our re-

sults show 120 species (11.5%) with EVS scores in the

low category of vulnerability, 267 (25.6%) in the medi-

um category, and 658 (63.0%) in the high category. For

Mexico, the comparable figures are 149 (12.2%), 378

(31.0%), and 692 (56.8%). Amazingly, we found more

than six of every 10 species in Central America in the

high category of vulnerability to environmental deterio-

ration. This figure is more elevated than that for the Mex-

ican herpetofauna, although in both regions more than

one-half of the herpetofauna has been judged to have the

highest level of vulnerability to environmental damage.

This statistic has astounding implications for efforts to

conserve this hugely significant herpetofauna.

Given that our EVS results show such a high percent-

age of the Central American herpetofauna in the high

vulnerability category, this situation needs to be exam-

ined more closely. Thus, we indicate the EVS categoriza-

tions of low, medium, and high in the last column in Ap-

pendices 1 and 2. We summarized these categorizations

and compared them to the scores for each of the three

components that contribute to the total EVS, as well as

the total EVS itself, and organized these data according

to the herpetofaunal families and orders (Table 4).

As noted above, amphibians generally are more en-

vironmentally vulnerable than the remainder of the her-

petofauna (percentage of high EVS 70.8% vs. 55.8%).

The relatively high figure for amphibians primarily is

due to the extremely high number of salamander spe-

cies placed in the high vulnerability category (92.4%)

compared to the situation among anurans (59.6%). All

salamanders in Central America are categorized in the

family Plethodontidae. According to the accounting at

the AmphibiaWeb site (accessed 9 December 2014),

this family consists of 444 species; thus, the 159 Central

American species comprise 35.8% of the total. This fig-

ure also represents 60.9% of the 261 species known from

Mesoamerica (www.mesoamericanherpetology.com; ac-

cessed 9 December 2014). The elevated vulnerability of

Central American salamanders largely is due to the small

geographic ranges and limited vegetational occurrence

of most species (respective average component scores

of 5.1 of 6 and 7.3 of 8; Table 4). All Central Ameri-

can plethodontids are direct developers, so the score for

reproductive mode always is 4. The average total EVS

score is 16.5, which is in the middle of the range of high

vulnerability scores (14-20).

The next most vulnerable group of amphibians con-

tains the caecilians, of which only 15 species occur in

Central America. Typically, these amphibians are more

broadly distributed, both geographically and vegetation-

ally (average component scores of 3.9 and 6.8, respec-

tively). Although their reproductive biology remains

poorly understood, all species likely are direct develop-

ers or viviparous, and thus are allocated reproductive

mode scores of 4 or 5.

Anurans generally are less environmentally vulner-

able than caecilians or salamanders (average EVS of

13.7). This situation principally is due to the relatively

fewer species with high scores for reproductive mode

(average score 2.8 of 6). Otherwise, the other compo-

nent scores for anurans are similar to those for caecilians

(4.4 vs. 3.9 for geographic distribution and 6.7 vs. 6.8

for ecological distribution). Nonetheless, our assessment

showed 59.6% of the 319 anuran species with high EVS

scores.

Of the 319 anuran species, 238 (74.6%) are catego-

rized in three families, the Bufonidae (39 species), Crau-

gastoridae (101), and Hylidae (98). Generally, members

of these families are more geographically widespread

than the typical salamander (respective average geo-

graphic component scores of 4.4, 4.8, and 4.4 compared

to that of 5.1 for salamanders). This situation also is the

case with vegetational occurrence (6.6, 7.0, and 6.7 vs.

7.3) . Typical bufonid and hylid anuran species lay eggs

in standing or flowing water, whereas craugastorid spe-

cies have direct development. Thus, the component for

reproductive mode is lower for bufonids and hylids

(1.3 and 2.0, respectively) than for craugastorids (4.0).

Nonetheless, we found the species with the highest EVS

scores, including the highest possible score, among the

hylid anurans. We calculated a total score of 20 for six

hylids (Appendix 1 ), four in the genus Ecnomiohyla ( E .

bailarina, E. rabborum, E. tliysanota, and E. veraguen-

sis ), one in the genus Isthmohyla (/. melacaena ), and one

in the genus Ptychohyla (P. dendrophasma). Given a to-

tal score of 20, each of these species is known only from

their respective type localities, from a single vegetation

zone, and has a reproductive mode of either laying eggs

in tree holes or in bromeliads (Appendix 1).

The reason why we assessed fewer crocodylians,

squamates, and turtles in the high EVS category than

amphibians primarily is due to their greater breadth in

geographic and ecological distribution (respective aver-

age values of 4.0 vs. 4.6 and 6.1 vs. 6.9). Nevertheless,

a slightly higher average score for degree of persecution

is present in these creatures (3.6) than for reproductive

mode in amphibians (3.2).

We found turtles and squamates slightly less vulner-

able than crocodylians (13.5 and 13.3, respectively, vs.

14.3) . Obviously, the patterns of vulnerability are skewed

toward the squamates, since 96.2% of the Central Ameri-

can crocodylians, squamates, and turtles are squamates.

Most squamates are classified in the families Dacty-

loidae (95 species), Sphaerodactylidae (19), Colubridae

(74), Dipsadidae (144), and Viperidae (32). Their total

number (364) represents 68.4% of the 532 species for

which an EVS can be calculated. We found the average

EVS scores for these families, respectively, as follows:

14.4, 14.4, 11.9, 13.0, and 15.1. Only the values for the

colubrids and dipsadids fell outside of the high value

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Conservation reassessment of Central American herpetofauna

Nothopsis rugosus. This unusual snake is distributed from

northeastern Honduras to northwestern Colombia on the Atlan-

tic versant, and on the Pacific versant from southwestern Costa

Rica to northwestern Ecuador, where it occurs in Lowland

Moist and Premontane Wet forests at elevations from near sea

level to 900 m. We estimated its EVS as 10, placing it at the

lower end of the medium vulnerability category, and its IUCN

status is Least Concern. This individual is from the Serrania de

San Bias, in Panama. Photo by Abel Batista.

Nototriton matama. The Matama Moss Salamander is known

only from the type locality at the southeastern end of the Fila

Matama, a ridge on the Atlantic slope of Cerro Chirripo in

southeastern Costa Rica, where it occurs in Premontane Wet

Forest at an elevation of 1,300 m. We calculated its EVS as 18,

placing it in the upper portion of the high vulnerability catego-

ry, but its IUCN status has not been determined. This individual

is the holotype of the species. Photo by Eduardo Boza Oviedo.

range (14-20). The proportion of high EVS species fell

below 50% only in colubrids (39.2%). In the other fami-

lies, the percentage values were, respectively, 72.7, 73.7,

52.8, and 78.1. We did not assign a total EVS score of

20 to any crocodylian, squamate, or turtle, although we

accorded a score of 19 to five species (four iguanids and

one viperid; Appendix 2). The four iguanids all are mem-

bers of the genus Ctenosaura (C. bakeri, C. oedirhina, C.

palearis, and C. quinquecarinata). The single viperid is

the recently described Bothriechis guifarroi.

In the case of amphibians and the remainder of the

herpetofauna, the typical member is a species allocated

to either the lower portion of the high vulnerability range

(14.7) or slightly above the upper portion of the medium

vulnerability category (13.3). Consequently, manage-

ment plans for the general protection of the herpetofauna,

Micrurus steward. This coralsnake is distributed in central Pan-

ama, where it occurs in Lowland Moist and Premontane Wet

forests at elevations from 500 to 1,200 m. We gauged its EVS

as 17, placing it in the middle portion of the high vulnerability

category, and its IUCN status is Least Concern. This individual

is from Donoso, province of Colon, Panama. Photo by Abel Ba-

tista.

Mastigodryas dorsalis. This racer is distributed from western

Guatemala to north-central Nicaragua, where it occurs in Pre-

montane Wet, Lower Montane Wet, and Lower Montane Moist

forests at elevations from 635 to 2,200 m. We determined its

EVS as 14, placing it at the lower end of the high vulnerability

category, and its IUCN status is Least Concern. This individual

is from Cerro Kilambe, department of Jinotega, Nicaragua.

Photo by Javier Sunyer.

and particularly the high vulnerability species, require

development in all regions of Central America.

Comparison of IUCN Categorizations and

EVS Values

Wilson et al. (2013a) stated that, “since the IUCN cat-

egorizations and EVS values both measure the degree of

environmental threat impinging on a given species, a cer-

tain degree of correlation between the results of these two

measures is expected.” They also noted that, “Townsend

and Wilson (2010) demonstrated this relationship with

reference to the Honduran herpetofauna, by comparing

the IUCN and EVS values for 362 species of amphibians

and terrestrial reptiles in their table 4.”

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Johnson et al.

Table 4. Summary of Environmental Vulnerability Scores by component, total score, and category, arranged by family. The num-

bers in the Environmental Vulnerability Scores columns represent ranges followed by means in parentheses. Values for EVS cat-

egories are percentages. L = low vulnerability; M = medium vulnerability; H = high vulnerability.

Environmental Vulnerability Scores

Families

Geographic

Distribution

(range 1-6)

Ecological

Distribution

(range 1-8)

Reproductive

Mode/Degree

of Persecution

(range 1-6)

Total Score

(range 3-20)

EVS Categories

Aromobatidae (3)

1-6 (4.0)

6-8 (7.0)

4 (4.0)

11-18(15.0)

L = 0.0, M = 33.3,

H = 66.7

Bufonidae (39)

1-6 (4.4)

1-8 (6.6)

1-4 (1.3)

3-17(12.2)

L = 20.5, M =

41.0, H = 38.5

Centrolenidae (14)

1-5 (2.3)

4-8 (6.6)

3 (3.0)

8-16(11.1)

L = 14.3, M =

64.3, H = 21.4

Craugastoridae (101)

2-6 (4.8)

3-8 (7.0)

4 (4.0)

9-18(15.8)

L = 1.0, M = 7.9, H

= 91.1

Dendrobatidae (19)

4-6 (4.8 )

6-8 (7.3)

4 (4.0)

14-18(15.3)

L = 0.0, M = 0.0, H

= 100

Eleutherodactylidae (11)

2-6 (4.5)

5-8 (7.2)

4 (4.0)

11-18(15.6)

L = 0.0, M= 18.2,

H = 81.8

Hemiphractidae (3)

3^1 (3.7)

7 (7.0)

5 (5.0)

15-16(15.7)

L = 0.0, M = 0.0, H

= 100

Hylidae (98)

1-6 (4.4)

1-8 (6.7)

1-6 (2.0)

3-20(13.1)

L = 12.2, M =

44.9, H = 42.9

Leptodactylidae (9)

1-5 (2.7)

2-8 (5.2)

2 (2.0)

5-14(10.0)

L = 44.4, M =

44.4, H= 11.2

Microhylidae (9)

2-5 (3.3)

1-8 (5.8)

1(1.0)

4-14(10.1)

L = 33.3, M =

55.6, H= 11.1

Pipidae (1)

4 (4.0)

8 (8.0)

5 (5.0)

17(17.0)

L = 0.0, M = 0.0, H

= 100

Ranidae (11)

1-6 (4 .1)

1-8 (4.8)

1(1.0)

3-15 (9.9)

L = 45.4, M =

36.4, 18.2

Rhinophrynidae (1)

2 (2.0)

5 (5.0)

1(1.0)

8 (8.0)

L = 100, M = 0.0,

H = 0.0

Total Anurans (319)

1-6 (4.4)

1-8 (6.7)

1-6 (2.8)

3-20 (13.7)

L = 11.3, M =

29.2, H = 59.6

Plethodontidae (159)

1-6 (5.1)

3-8 (7.3)

4 (4.0)

8-18(16.5)

L = 1.3, M = 6.3, H

= 92.4

Total Salamanders (159)

1-6 (5.1)

3-8 (7.3)

4 (4.0)

8-18 (16.5)

L = 1.3, M = 6.3,

H = 92.4

Caeciliidae (7)

3-6 (4.4)

7-8 (7.9)

4-5 (4.4)

15-19(16.7)

L = 0.0, M = 0.0, H

= 100

Dermophiidae (8)

1-5 (3.4)

1-8 (5.9)

5 (5.0)

7-18(14.3)

L = 12.5, M =

25.0, H = 62.5

Total Caecilians (15)

1-6 (3.9)

1-8 (6.8)

4-5 (4.7)

7-19 (15.4)

L = 6.7, M = 13.3,

H = 80.0

Total Amphibians (493)

1-6 (4.6)

1-8 (6.9)

1-6 (3.2)

3-20 (14.7)

L = 7.9, M = 21.3,

H = 70.8

Alligatoridae (1)

3 (3.0)

7 (7.0)

6 (6.0)

16(16.0)

L = 0.0, M = 0.0, H

= 100

Crocodylidae (2)

2-3 (2.5)

5 (5.0)

6 (6.0)

13-14 (13.5)

L = 0.0, M = 50.0,

H = 50.0

Total Crocodylians (3)

2-3 (2.7)

5-7 (5.7)

6 (6.0)

13-16 (14.3)

L = 0.0, M = 33.3,

H = 66.7

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Conservation reassessment of Central American herpetofauna

Table 4 (continued). Summary of Environmental Vulnerability Scores by component, total score, and category, arranged by family.

The numbers in the Environmental Vulnerability Scores columns represent ranges followed by means in parentheses. Values for

EVS categories are percentages. L = low vulnerability; M = medium vulnerability; H = high vulnerability.

Environmental Vulnerability Scores

Families

Geographic

Distribution

(range 1-6)

Ecological

Distribution

(range 1-8)

Reproductive

Mode/Degree

of Persecution

(range 1-6)

Total Score

(range 3-20)

EVS Categories

Amphisbaenidae (2)

3 (3.0)

7-8 (7.5)

1(1.0)

11-12(11.5)

L = 0.0, M = 100,

H = 0.0

Anguidae (28)

2-6 (4.9)

3-8 (7.2)

3^1 (3.6)

8-18(15.7)

L = 3.6, M = 3.6, H

= 92.8

Corytophanidae (9)

1-5 (3.0)

3-7 (4.9)

3 (3.0)

7-15(10.9)

L = 33.3, M =

55.6, H= 11.1

Dactyloidae (95)

1-6 (4.7)

2-8 (6.7)

3 (3.0)

7-17(14.4)

L = 8.4, M= 18.9,

H = 72.7

Eublepharidae (2)

3-5 (4.0)

4 (4.0)

10-14(12.0)

L = 0.0, M = 50.0,

H = 50.0

Gymnophthalmidae (14)

2-5 (4.0)

3-8 (7.0)

2-3 (2.7)

9-16(13.7)

L = 7.1, M = 21.5,

H = 71.4

Helodermatidae (2)

3-5 (4.0)

6-8 (7.0)

5 (5.0)

14-18(16.0)

L = 0.0, M = 0.0, H

= 100

Hoplocercidae (2)

3^1 (3.5)

7-8 (7.5)

3 (3.0)

13-15 (14.0)

L = 0.0, M = 50.0,

H = 50.0

Iguanidae (11)

1-5 (4.0)

3-8 (6.5)

3-6 (5.7)

10-19(16.3)

L = 0.0, M = 27.3,

H = 72.7

Mabuyidae (5)

1-6 (4.4)

2-8 (6.4)

3 (3.0)

6-17 ( 13.8)

L = 20.0, M = 0.0,

H = 80.0

Phrynosomatidae (17)

1-5 (3.6)

1-8 (5.0)

3 (3.0)

5-15(11.6)

L = 11.8m M =

64.7, H = 23.5

Phyllodactylidae (5)

1-6 (3.8)

4-8 (6.4)

3 (3.0)

8-17 (13.2)

L = 40.0, M = 0.0,

H = 60.0

Polychrotidae (1)

1 (1.0)

8 (8.0)

3 (3.0)

12(12.0)

L = 0.0,M= 100,

H = 0.0

Scincidae (3)

4-5 (4.3)

5-6 (5.7)

3 (3.0)

12-14(13.0)

L = 0.0, M = 66.7,

H = 33.3

Sphaerodactylidae (19)

1-6 (4.3)

3-8 (7.1)

3 (3.0)

8-17(14.4)

L = 10.5, M =

15.8, H = 73.7

Sphenomorphidae (4)

2-6 (4.0)

2-8 (4.8)

3 (3.0)

7-17(11.8)

L = 50.0, M = 0.0,

H = 50.0

Teiidae (12)

1-5 (3.6)

2-8 (6.3)

3 (3.0)

6-16(12.9)

L = 16.7, M =

25.0, H = 58.3

Xantusiidae (4)

2-5 (3.5)

4-7 (5.5)

2 (2.0)

9-13 (11.0)

L = 50.0, M =

50.0, H = 0.0

Xenosauridae (1)

3 (3.0)

1(1.0)

3 (3.0)

7 (7.0)

L = 100, M = 0.0,

H = 0.0

Anomalepididae (3)

2-5 (3.3)

5-8 (6.3)

1 (1.0

9-12 (10.7)

L = 33.3, M =

66.7, H = 0.0

Boidae (4)

1-3 (1.5 )

1-8 (5.5)

2-6 (3.0)

8-13 (10.0)

L = 50.0, M =

50.0, H = 0.0

Charinidae (2)

2^1 (3.0)

5-6 (5.5)

2 (2.0)

9-12 (10.5)

L = 50.0, M =

50.0, H = 0.0

Colubridae (74)

1-6 (3.6)

1-8 (5.1)

2-5 (3.2)

5-17(11.9)

L = 25.7m M =

35.1, H = 39.2

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Johnson et al.

Table 4 (continued). Summary of Environmental Vulnerability Scores by component, total score, and category, arranged by family.

The numbers in the Environmental Vulnerability Scores columns represent ranges followed by means in parentheses. Values for

EVS categories are percentages. L = low vulnerability; M = medium vulnerability; H = high vulnerability.

Environmental Vulnerability Scores

Families

Geographic

Distribution

(range 1-6)

Ecological

Distribution

(range 1-8)

Reproductive

Mode/Degree

of Persecution

(range 1-6)

Total Score

(range 3-20)

EVS Categories

Dipsadidae (144)

1-6 (4.1)

1-8 (6.2)

2-5 (4.0)

4-17(13.0)

L = 13.9, M =

33.3, H = 52.8

Elapidae (17)

2-5 (4.0)

1-8 (5.7)

5 (5.0)

9-18(14.7)

L = 11.8, M = 17.6,

H = 70.6

Leptotyphlopidae (5)

3-5 (3.6)

1-8 (4.2)

1(1.0)

5-12 (8.8)

L = 40.0, M =

60.0, H = 0.0

Loxocemidae (1)

1(1.0)

5 (5.0)

4 (4.0)

11 (11.0)

L = 0.0,M = 100,

H = 0.0

Natricidae (5)

1-4 (2.8)

1-5(34)

2-4 (3.6)

8-13 (9.8)

L = 60.0, M =

40.0, H = 0.0

Sibynophiidae (2)

1(1.0)

5-7 (6.0)

5 (5.0)

11-13 (12.0)

L = 0.0,M = 100,

H = 0.0

Tropidophiidae (1)

3 (3.0)

5 (5.0)

3 (3.0)

11 (11.0)

L = 0.0, M = 100,

H = 0.0

Typhlopidae (5)

4-5 (4.6)

5-8 (6.4)

1(1.0)

11-14(12.0)

L = 0.0, M = 80.0,

H = 20.0

Viperidae (32)

1-6 (4.0)

2-8 (6.0)

5 (5.0)

9-19(15.1)

L = 3.1,M= 18.8,

H = 78.1

Total Squamates (531)

1-6 (4.1)

1-8 (6.1)

1-6 (3.5)

4-19 (13.3)

L = 14.7, M =

29.6, H = 55.7

Chelydridae (2)

1-4 (2.5)

4-7 (5.5)

6 (6.0)

11-17(14.0)

L = 0.0, M = 50.0,

H = 50.0

Dermatemydidae (1)

4 (4.0)

7 (7.0)

6 (6.0)

17(17.0)

L = 0.0, M = 0.0, H

= 100

Emydidae (2)

1-5 (3.0)

4-8 (6.0)

6 (6.0)

11-19(15.0)

L = 0.0, M = 50.0,

H = 50.0

Geoemydidae (5)

1-5 (3.2)

4-8 (6.6)

3 (3.0)

8-16(12.8)

L = 20.0, M =

40.0, H = 40.0

Kinosternidae (4)

1-5 (2.8)

4-8 (5.8)

3 (3.0)

8-16(11.5)

L = 50.0, M = 0.0,

H = 50.0

Staurotypidae (3)

4 (4.0)

4-8 (6.7)

3 (3.0)

13-14(13.7)

L = 0.0, M = 33.3,

H = 66.7

Testudinidae (1)

3 (3.0)

8 (8.0)

6 (6.0)

17 (17.0)

E = 0.0, M = 0.0, H

= 100

Total Turtles (18)

1-5 (3.2)

4-8 (6.3)

3-6 (4.0)

8-19 (13.5)

L = 16.7, M =

27.8, H = 55.5

Total “Reptiles” (552)

1-6 (4.0)

1-8 (6.1)

1-6 (3.6)

4-19 (13.3)

L = 14.7, M =

29.5, H = 55.8

Total Herpetofauna

(1045)

1-6 (4.3)

1-8 (6.5)

1-6 (3.4)

3-20 (14.0)

L = 11.5, M =

25.6, H = 62.9

As Wilson et al. (2013a, b) developed for the Mexi-

can herpetofauna, we constructed a pair of tables (Tables

5 and 6) to judge whether such a correspondence exists

between these two measures of conservation status for

the Central American herpetofauna. The results for the

Mexican and Central American amphibian faunas are

Amphib. Reptile Conserv. 41

comparable to a point, but not otherwise. With respect to

the IUCN categorizaitons, the absolute numbers for the

three threat categories and the NT category are similar

to one another (Central American values indicated first;

CR = 91 vs. 88, EN = 75 vs. 79, VU = 44 vs. 44, NT

= 26 vs. 21), even though 114 more amphibian species

August 2015 | Volume 9 | Number 2 | el 00

Conservation reassessment of Central American herpetofauna

occur in Central America than in Mexico (493 vs. 379).

Interestingly, the values for the LC, DD, and NE catego-

ries are not similar between the two regions, especially

with respect to the latter two (LC = 147 vs. 91, DD =

55 vs. 38, NE = 52 vs. 17). Apparently, a correlation ex-

ists between the greater number of amphibian species in

Central America and those in Mexico, and the number of

species relegated to the LC, DD, and NE categories in

the two regions. Of the 493 Central American amphib-

ian species, 236 (47.9%) have been categorized as CR,

EN, VU, and NT. In Mexico, 232 (61.4%) of these spe-

cies have been assessed in these categories. In Central

America, however, a significantly larger percentage of

the species have been assessed in the LC, DD, and NE

categories (254 [51.5%]) than in Mexico (146 [38.6%]).

Note that the two percentage figures for Central America

do not add up to 100, because three species in this region

have been judged as extinct (Appendix 1). Why such a

relatively large percentage of DD + NE species (21.7%)

is present in Central American amphibians compared to

those in Mexico (14.6%) is not evident, but it means that

more than one in every five species in Central America

has not been evaluated or is too poorly known to allow

for an evaluation. This situation provided us with a spe-

cial impetus to conduct an EVS analysis on these crea-

tures.

Like Wilson et al. (2013b) did for the Mexican am-

phibians, we detennined the mean EVS for each of the

IUCN columns in Table 5, including the NE species and

the total species. The results are as follows: CR (91 spp.)

= 15.4 (range 7-20); EN (75 spp.) = 15.3 (9-18); VU

(44 spp.) = 14.8 (7-18); NT (26 spp.) = 14.9 (8-20); LC

(147 spp.) = 8.0 (3-17); DD (55 spp.) = 16.8 (13-20);

NE (52 spp.) = 17.3 (8-20); and total (493 spp.) = 14.7

(3-20). Some interesting resemblances are evident be-

tween these data and those for the Mexican amphibians

(Wilson et al. 2013b). As with the Mexican species, the

mean EVS value decreases steadily (though not as dra-

matically) from the CR category (15.4) through the EN

(15.3), and VU (14.8) categories, with the value for the

NT species (14.9) almost the same as that for the VU

species. A precipitous drop also is evident from the VU

and NT values to those for the LC species (8.0), more so

than for the Mexican amphibians. Although this decrease

was expected, as for the Mexican amphibians we did not

anticipate the size of the mean value for the DD species

in Central America (16.8), which is almost the highest

mean value for these categories. Thus, this value is sub-

stantially higher than that for any of the threat species.

Even more surprising is that the mean value for the NE

species is even higher (17.3) than that for the DD spe-

cies. The value for the DD species supports our stated

opinion about these species; apparently the NE group

also is comprised of such species. As expected, the EVS

values for almost all the DD (54 of 55 [98.2%]) and the

NE species (51 of 52 [98.1%]) fell into the high vulner-

ability category, including the average total value (14.7).

These additional reasons provide a compelling argument

for conducting a reassessment of the Central American

herpetofauna based on the EVS measure.

A revealing statistic is that the average EVS value

for each IUCN category, except for the LC, fell into the

high vulnerability category. With the LC category, of

the 38 amphibian species with EVS values in the low

vulnerability category, 30 (78.9%) have been placed in

this IUCN category; however, 51 (34.7%) of the LC spe-

cies fell into the medium category, with the remaining 66

(44.9%) species into the high category. Thus, as with our

work on the Mexican herpetofauna, these data support

our opinion that the LC category is applied too broadly in

IUCN assessments to be of significant value in conserva-

tion planning.

As with Table 5, the data in Table 6 illustrate the rela-

tionship between the IUCN ratings and EVS values for

the 552 Central American crocodylians, squamates, and

turtles. These data can be compared to those for these

creatures in Mexico (see Wilson et al. 2013a). With refer-

ence to the IUCN categorizations, the absolute numbers

for the three threat categories and the NT category for

the two regions are not as similar to one another for the

crocodylians, squamates, and turtles as they are for the

amphibians (Central American values listed first; CR =

14 vs. 6, EN = 30 vs. 36, VU = 18 vs. 44, NT = 21 vs.

26). The figures for Central America total 83, compared

to 112 for Mexico. The total figures for the two regions,

however, comprise reasonably close percentages of the

respective total non-amphibian herpetofaunas (i.e., 83 of

552 [15.0%] vs. 112 of 841 [13.3%]). We believe, how-

ever, that once the IUCN categorizations are available

for Central American anoles that the ranks of the three

threat categories and the NT category will be augmented,

similar to when the categorizations are published for the

species in Central and South America. With respect to

the remainder of the IUCN categorizations, however, the

total relative figures are comparable for Central America

and Mexico. The comparable absolute figures for the two

regions, respectively, are as follows: LC = 208 vs. 422,

DD = 35 vs. 1 1 8, NE = 226 vs. 1 89. For Central America,

the three absolute values total 469 species (85.0% of the

total of 552); for Mexico, the comparable figures are 729

and 86.7%.

Equivalent to the approach in Wilson et al. (2013a)

for Mexican crocodylians, squamates, and turtles, we

ascertained the mean EVS scores for each of the IUCN

columns in Table 6, including the NE species and the to-

tal species. The results are as follows: CR (14 spp.) =

16.6 (15-19); EN (30 spp.) = 15.9 (13-19); VU (18 spp.)

= 15.0 (7-18); NT (21 spp.) = 14.3 (12-16); LC (208

spp.) = 12.3 (4-18); DD (35 spp.) = 15.6 (11-18); NE

(226 spp.) =13.2 (4-19). In common with Mexican cro-

codylians, squamates, and turtles (Wilson et al. 2013a),

a corresponding increase in average EVS scores is evi-

dent with ascending degrees of threat, from LC through

CR. Similar to the situation with Mexican crocodylians,

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

42

Johnson et al.

Table 5. Comparison of Environmental Vulnerability Scores (EVS) and IUCN categorizations for Central American amphibians.

Shaded area at the top encompasses low vulnerability scores, and that at the bottom high vulnerability scores.

IUCN categories

EVS

Extinct

Critically

Endangered

Endan-

gered

Vulner-

able

Near

Threatened

Least

Concern

Data

Deficient

Not

Evaluated

Totals

3

4

5

6

7

—

1

—

1

—

3

2

4

—

3

2

6

8

—

1

9

—

1

12

9

—

1

—

8

—

11

10

—

2

1

—

6

—

11

—

2

—

4

—

14

—

20

12

—

5

6

3

5

17

—

36

13

—

14

4

2

14

1

38

14

—

9

8

3

2

17

4

—

43

15

2

11

8

6

4

19

7

1

58

16

—

7

24

7

25

10

5

85

17

—

12

18

5

1

4

11

12

63

18

1

23

4

10

2

—

19

30

89

19

—

2

—

1

—

2

—

5

20

—

2

—

1

—

1

2

6

Totals

3

91

75

44

26

147

55

52

493

Table 6. Comparison of Environmental Vulnerability Scores (EVS) and IUCN categorizations for Central American crocodylians,

squamates, and turtles. Shaded area at the top encompasses low vulnerability scores, and that at the bottom high vulnerability scores.

IUCN categories

EVS

Critically

Endangered

Vulnerable

Near

Threatened

Least

Concern

Data

Deficient

Not

Evaluated

Totals

3

4

—

1

—

1

2

5

—

1

—

4

5

6

—

6

—

5

11

7

—

1

—

7

—

6

14

8

—

9

—

10

19

9

—

17

—

12

29

10

—

15

—

11

26

11

—

13

2

12

27

12

—

1

33

1

17

52

13

—

3

—

4

28

1

24

60

14

—

4

6

27

1

30

74

15

l

5

2

7

25

6

34

80

16

8

7

5

3

18

17

26

84

17

1

5

3

—

6

4

30

49

18

3

4

1

—

2

3

1

14

19

20

1

2

—

3

6

Totals

14

30

18

21

208

35

226

552

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Amphib. Reptile Conserv.

43

Conservation reassessment of Central American herpetofauna

squamates, and turtles, the average EVS scores for the

DD species (15.6) is closest to that for the VU species

(15.5), which also suggests that if and when these species

are better known, they likely will be judged as VU, EN,

or CR. The amount of decrease in average EVS scores

for Central American crocodylians, squamates, and tur-

tles from the NT to the LC categories (14.3 to 12.3) is

comparable to the same groups in Mexico (12.9 to 10.5),

although those for Central America are higher. The NE

species constitute the largest component of the Central

American crocodylian, squamate, and turtle fauna (226

species [40.9%] of the total). The average EVS score for

these species is closest to that for the LC species (13.2 vs.

12.3), the second largest group (208 species). The ranges

in their EVS values are similar to one another (4-19 for

NE species, 4-18 for LC species). Eventually, the large

number of NE species likely will join the large number

of LC species when the IUCN categorizations have been

determined for the former group. If so, then the combined

226 NE and 208 LC species would comprise 434 species

(78.6% of the Central American crocodylian, squamate,

and turtle fauna). If this eventually occurs, it would con-

stitute a travesty of conservation effort, allowing for a

serious degradation of the significance of this fauna.

Similar to the situation with Central American am-

phibians, most of the average EVS values we assessed

for the crocodylians, squamates, and turtles, except for

those in the LC and NE categories, fell into the high vul-

nerability category. The LC and NE averages, however,

fell into the upper portion of the medium vulnerability

category (12.3 and 13.2, respectively). Interestingly, the

ranges of EVS values for the LC and NE species are sim-

ilar (4-18 and 4-19, respectively). Both of these ranges

are close to the total possible EVS range of 3-20. The

EVS values, however, were not evenly distributed among

the three vulnerability categories in either case. With the

LC category, 41 of the 208 species (19.7%) fell into the

low vulnerability grouping, 89 (42.8%) into the medium

category, and 78 (37.5%) into the high category. With

regard to the NE category, the comparable values are

38 (16.8%), 64 (28.3%), and 122 (54.9%). As with the

Central American amphibians, the LC category appears

too broadly applied to a large a group of species to be

of meaningful conservation value for decision-making.

Given the large number of species that remain unevalu-

ated, many of these species likely will be allocated to

the LC category, thus inflating the lack of utility of this

category to reasonably reflect the conservation status of

the species involved.

We harbor no illusions that the EVS measure will

come to replace the IUCN system of categorization of

conservation status and do not necessarily desire for this

change to happen, but we maintain that the IUCN system

has serious disadvantages when compared with the EVS

measure. For the purposes of this analysis, if we divide

the IUCN categories into three groups so they can be

compared with the three EVS categories, and determine

the absolute and relative numbers of species occupying

each, the results are germane to our conclusions.

The three groupings of the IUCN categories are as

follows: EX+CR+EN+VU; NT+LC; and DD+NE. Sum-

ming the numbers of species from tables 5 and 6 in each

of these groupings for amphibians and the remainder of

the herpetofauna provides a set of absolute values for the

entire herpetofauna, in respective order as follows: 275

species (26.3%); 402 (38.5%); and 368 (35.2%). For the

three EVS groupings, from high through medium to low,

the results are as follows: 656 (62.8%); 270 (25.8%); and

119 (11.4%). The three IUCN groups and the three EVS

groupings are not entirely comparable; nonetheless, the

first IUCN grouping (EX+CR+EN+VU), i.e., the threat

categories plus the extinct one, can be compared to the

high vulnerability EVS grouping. Only 275 species

(26.3%) of the total are allocated to the IUCN group-

ing, whereas 656 species (62.8%) are placed in the EVS

grouping. The second IUCN grouping (NT+LC) is gross-

ly comparable to the low vulnerability EVS grouping;

the respective values are: 402 (38.5%) and 119 (11.4%).

The third IUCN grouping is not comparable to any of the

EVS groupings, since all the species can be evaluated

using the latter, whereas a substantial proportion (367

species [35.2%]) of the former remain unevaluated. Even

with the discrepancies between the IUCN and EVS sys-

tems, the use of the latter identifies a substantially larger

absolute and relative number of species in need of seri-

ous conservation attention (275 [26.3%] vs. 656 [62.8%],

respectively) and a substantially smaller absolute and

relative number of species least needing this attention

(402 [38.5%] vs. 119 [11.4%], respectively). These high-

ly divergent results have profound consequences in ef-

forts to conserve the highly significant Central American

herpetofauna. The IUCN evaluation implies that this is

a much simpler task to accomplish than the EVS evalu-

ation. Such a conservation effort presently is a huge un-

dertaking, which will grow increasingly in extent into the

forseeable future.

Comparison of EVS Results for Central

America and Mexico

We demonstrated that a large proportion of the Central

American herpetofauna is highly vulnerable to envi-

ronmental deterioration, more so than for the Mexican

herpetofauna. To examine this situation in more detail,

we constructed Table 7, in which the absolute and rela-

tive distribution of EVS values is indicated for the major

herpetofaunal groups. For ease of understanding, we col-

lapsed these data (Table 8) into the three categories of

vulnerability generally recognized for the EVS measure,

i.e., low, medium, and high.

Perusal of the data in Table 8 indicates that the general

pattern for amphibians is for the numbers and percent-

ages to increase from the low through the medium to the

high categories. This pattern is evident in both regions

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

44

Johnson et al.

Table 7. Comparison of Environmental Vulnerability Scores and Percentages for the Central American and Mexican herpetofauna, arranged by major

groups. Shaded area to the left encompasses low vulnerability scores, and to the right high vulnerability scores. CA = Central American. Data for Central

American taxa are from Table 3, and for Mexican taxa from Wilson et al. (2013a, b).

Number Environmental Vulnerability Scores

Major groups

of

species

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

C A Anurans

319

3

2

5

11

10

11

18

34

30

37

39

50

26

29

3

6

Percentages

—

0.9

0.6

1.6

3.4

3.1

3.4

5.6

10.7

9.4

11.6

12.2

15.7

8.2

9.1

0.9

1.9

Mexican Anurans

237

4

3

4

9

12

14

13

20

25

29

36

30

8

14

12

1

—

Percentages

—

1.7

1.3

1.7

3.8

5.1

5.9

5.4

8.4

10.5

12.2

15.2

12.7

3.4

5.9

5.1

8.4

—

CA Salamanders

159

—

1

—

2

6

4

17

34

35

57

—

Percentages

—

0.6

—

1.3

3.8

2.5

10.7

21.4

22.0

35.8

—

Mexican Salamanders

139

—

1

2

6

7

13

23

13

36

—

Percentages

—

0.7

1.4

4.3

5.0

9.4

16.6

9.4

25.9

—

CA Caecilians

15

—

1

—

2

—

Percentages

—

6.7

—

13.3

—

Mexican Caecilians

3

1

—

1

—

Percentages

33.3

—

33.3

—

C A Amphibians

493

3

2

6

12

11

20

36

38

42

58

87

63

88

5

6

Percentages

—

0.6

0.4

1.2

2.4

2.2

4.1

7.3

7.7

8.5

11.8

17.6

12.8

17.8

1.0

1.2

Mexican Amphibians

379

4

3

4

9

12

15

23

32

36

49

53

22

50

48

1

—

Percentages

—

1.1

0.8

1.1

2.4

3.2

4.0

6.1

8.4

9.5

12.9

14.0

5.8

13.2

12.7

0.3

—

CA Crocodylians

3

1

—

1

—

Percentages

33.3

—

33.3

—

Mexican Crocodylians

3

1

—

1

—

Percentages

33.3

—

33.3

—

C A Lizards

236

—

1

2

5

9

11

7

6

15

26

29

45

33

34

9

4

—

Percentages

—

0.4

0.8

2.1

3.8

4.7

3.0

2.5

6.4

11.0

12.3

19.1

14.0

14.4

3.8

1.7

Mexican Lizards

413

—

1

3

6

11

13

14

28

39

49

54

67

78

38

10

2

—

Percentages

—

0.2

0.7

1.5

2.7

3.1

3.4

6.8

9.4

11.9

13.1

16.2

18.9

9.2

2.4

0.5

—

CA Snakes

295

—

2

4

9

8

19

18

35

31

41

35

47

14

5

1

—

Percentages

—

0.7

1.4

3.1

2.7

6.4

6.1

11.9

10.5

13.9

11.9

15.9

4.7

1.7

0.3

—

Mexican Snakes

383

1

7

10

9

19

17

30

25

31

47

52

50

44

24

9

7

—

Percentages

—

0.3

1.8

2.6

2.3

5.0

4.4

7.8

6.5

8.1

12.3

13.6

13.1

11.5

6.3

2.3

1.8

—

C A Turtles

18

—

3

—

2

1

2

3

1

2

3

—

1

—

Percentages

—

16.7

—

11.1

5.5

11.1

16.7

5.5

11.1

16.7

—

5.5

—

Mexican Turtles

42

—

1

—

3

1

3

8

6

4

3

5

6

1

Percentages

—

2.4

—

7.1

2.4

7.1

19.0

14.3

9.5

7.1

11.9

14.3

2.4

CA “Reptiles”

552

—

2

5

11

13

20

30

25

26

51

60

74

81

83

51

14

6

—

Percentages

—

0.4

0.9

2.0

2.4

3.6

5.4

4.5

4.7

9.2

10.9

13.4

14.7

15.0

9.2

2.5

1.1

—

Mexican “Reptiles”

841

1

8

13

15

31

30

47

54

71

100

115

123

127

65

24

15

1

Percentages

—

0.1

1.0

1.5

1.8

3.7

3.6

5.6

6.4

8.4

11.9

13.7

14.6

15.1

7.7

2.9

1.8

0.1

(Central America and Mexico), and in each of the major

groups (anurans, caecilians, and salamanders). The rela-

tionship of the numbers and percentages changes, how-

ever, between the two regions and among the three major

groups. Among the anurans, proportionately more taxa

were assigned to the high category in Central America

(59.5%) than in Mexico (42.6%). Among the salaman-

ders, the same situation is evident (92.4% vs. 87.1%).

This relationship is not evident among the caecilians,

since there is only one of three Mexican caecilians, in-

cluding the recently reported Gymnopis syntrema with

an assessed score falling into the high category. Overall,

more taxa were assessed in the high category in Central

America than Mexico (70.8% vs. 58.8%, respectively).

In both Central America and Mexico, the group of am-

phibians exhibiting the greatest vulnerability to environ-

mental damage were the salamanders, with about nine

of every 10 species assessed in the high category. A ma-

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Amphib. Reptile Conserv.

45

Conservation reassessment of Central American herpetofauna

Table 8. Summary comparison of EVS category values and percentages from Table 6 for the Central American and Mexican her-

petofauna, arranged by major groups.

Major groups l

EVS Categories

jumDer ot species

Low

Medium

High

C A Anurans

319

36

93

190

Percentages

—

11.3

29.2

59.5

Mexican Anurans

237

49

87

101

Percentages

—

20.7

36.7

42.6

CA Salamanders

159

2

10

147

Percentages

—

1.3

6.3

92.4

Mexican Salamanders

139

1

17

121

Percentages

—

0.7

12.2

87.1

CA Caecilians

15

1

2

12

Percentages

—

6.7

13.3

80.0

Mexican Caecilians

3

—

2

1

Percentages

—

66.7

33.3

C A Amphibians

493

39

105

349

Percentages

—

7.9

21.3

70.8

Mexican Amphibians

379

50

106

223

Percentages

—

13.2

28.0

58.8

CA Crocodylians

3

—

1

2

Percentages

—

33.3

66.7

Mexican Crocodylians

3

—

1

2

Percentages

—

33.3

66.7

CA Lizards

236

28

54

154

Percentages

—

11.9

22.9

65.2

Mexican Lizards

413

34

130

249

Percentages

—

8.2

31.5

60.3

CA Snakes

295

50

102

143

Percentages

—

16.9

34.6

48.5

Mexican Snakes

383

64

133

186

Percentages

—

16.7

34.7

48.6

CA Turtles

18

3

5

10

Percentages

—

16.7

27.8

55.5

Mexican Turtles

42

1

8

33

Percentages

—

2.4

19.0

78.6

CA “Reptiles”

552

81

162

309

Percentages

—

14.7

29.3

56.0

Mexican “Reptiles”

841

99

272

470

Percentages

—

11.8

32.3

55.9

jor distinction is evident between the salamanders and

the anurans, given that about four of every 10 species

of anurans in Mexico and about six of every 10 species

in Central America were assessed in the high category.

In both Central America and Mexico (thus, all of Meso-

america) salamanders are of most crucial conservation

concern.

The same general pattern we found among the am-

phibians also is evident among the remainder of the her-

Amphib. Reptile Conserv.

petofauna, i.e., an increase in the numbers and percentag-

es from low through medium to high in both regions and

within each group. Again, as with the amphibians, some

distinctions can be made among the proportions of taxa

falling into the three categories of vulnerability. Among

the turtles, a greater proportion fell into the high category

in Mexico than in Central America (78.6% vs. 55.5%).

Among the lizards, however, the proportions falling into

the three categories are similar to one another in Cen-

46

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Johnson et al.

Nototriton lignicola. This salamander is endemic to mountains in north-central Honduras, where it occurs in Lower Montane Wet

Forest at elevations from 1,760 to 2,000 m. We determined its EVS as 18, placing it in the upper portion of the high vulnerability

category, and its 1UCN status is Critically Endangered. This individual is from Cataguana, Parque Nacional Montana de Yoro, de-

partment of Yoro. Photo by Josicih H. Townsend.

tral America and Mexico, respectively (low: 11.9 vs. 8.2;

medium: 22.9 vs. 31.5; high: 65.2 vs. 60.3). Interestingly,

among the snakes, the proportions were almost the same

in Central America and Mexico (low: 16.9 vs. 16.7; me-

dium: 34.6 vs. 34.7; high: 48.5 vs. 48.6). Considering

the two larger species groups, lizards and snakes, lizards

generally were more vulnerable to environmental dam-

age than snakes in both Central America and Mexico

(65.2% and 60.3% for lizards and 48.5% and 48.6% for

snakes, respectively). Nonetheless, just as with amphib-

ians, more than one-half of the combined Central Ameri-

can and Mexican crocodylians, squamates, and turtles

fell into the high category of vulnerability (56.0% and

55.9%, respectively), which is of major conservation

concern.

Discussion

Biodiversity conservation requires one of the greatest ef-

forts for crisis intervention ever undertaken by humanity.

As we stated in the Introduction, the fundamental sig-

nificance of this effort generally goes unappreciated by

humanity at large. Thus, the attempt to salvage planet

Earth as a haven for life essentially falls to the interest

of an extremely small number of professional conserva-

tion biologists and a somewhat larger group of commit-

ted non-professional environmentalists. In essence, this

tiny group of people is pitted against the remainder of

humanity, collectively termed the “planetary killer” by

Wilson (2002), which, knowingly or unknowingly, has

cooperated to create the sixth mass extinction episode in

Earth’s history (Wake and Vredenburg 2008).

No matter what the actual number of people devoted

to conserving biodiversity is, it pales in significance when

compared to the number of humans who collectively rep-

resent the reason why biodiversity decline exists. At the

time of this writing (4:20 PM on 10 December 2014), the

global human population was estimated as 7,210,491,630

(www.census.gov). This constantly increasing figure is

the most important statistic in attempting to determine

the impact of humanity on the natural world. According

to the Population Reference Bureau World (PRB) 2013

Population Data Sheet (available at www.prb.org), the

current rate of natural increase is 1.2 (i.e., crude birth

rate - crude death rate / 10). Thus, the current doubling

time of the global population is 58.3 years. In other

words, the current world population indicated above will

double to 14,420,098,326 by early April, 2073, assum-

ing that the growth rate remains constant. Nonetheless,

the growth rate has been declining since peaking in the

period from 1962 to 1963 and is projected to fall to zero

in about 2080; thus, the total human population might

peak at about 10.3 billion (Population growth, Wikipe-

dia, en.wikipedia.org; accessed 9 January 2014). The ac-

tual pattern of growth will depend on the extent of family

planning on the growth rate. The 2013 Population Data

Sheet projects that the mid-2050 global population will

be 9.727 billion, and that the greatest amount of growth

(1.3 billion) will come in Sub-Saharan Africa. This fig-

ure exceeds the growth expected in Asia, the population

giant. By the year 2050, Nigeria will surpass the United

States to become the world’s third most populous nation,

after India and China (which will switch positions to be-

come the largest and second most populous nations, re-

spectively). In contrast, by 2050 the populations of North

America and Europe are projected to remain at their cur-

rent levels (at 0.4 and 0.7 billion, respectively).

Given this projected pattern of growth, what conse-

quences can we expect? With respect to human impact on

planetary biodiversity, we can expect that “nearly all fu-

ture population growth will be in the world’s less devel-

oped countries” (PRB 2012 Population Data Sheet: 5).

The current population level in the less developed coun-

tries is 4.7 times greater than that of the more developed

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Conservation reassessment of Central American herpetofauna

Nototriton mime. This moss salamander is known only from the

type locality, Cerro de Ulloa on the border of the departments of

Colon and Olancho in north-central Honduras, where it occurs

in Lower Montane Wet Forest at an elevation of 1,705 m. We

evaluated its EVS as 18, placing it in the upper portion of the

high vulnerability category, but its IUCN status has not been

determined. This individual is from the type locality. Photo by

Josiah H. Townsend.

Oedipina nica. This worm salamander is known only from

three isolated localities in north-central Nicaragua, where it oc-

curs in Lower Montane Wet Forest at elevations from 1,360 to

1,660 m. We gauged its EVS as 17, placing it in the middle por-

tion of the high vulnerability category, but its IUCN status has

not been determined. This individual is from Finca Monimbo,

department of Matagalpa. Photo by Javier Sunyer.

Nototriton stuarti. Stuart’s Moss Salamander is known only

from the type locality, Montanas del Mico in extreme eastern

Guatemala, where it occurs in Premontane Wet Forest at an el-

evation of 744 m. We assessed its EVS as 18, placing it in the

upper portion of the high vulnerability category, and its IUCN

status is Data Deficient. Photo by Sean Michael Rovito.

Oedipina carablanca. This worm salamander is known only

from the vicinity of the type locality in east-central Costa Rica,

where it occurs in Lowland Moist Forest at elevations from 60

to 260 m. We established its EVS as 18, placing it in the upper

portion of the high vulnerability category, and its IUCN sta-

tus is Endangered. This individual is from Pocora, 15 km NW

Siquirres, province of Limon. Photo by Brian Kubicki.

ones (PRB 2012 Population Data Sheet). Based on the

population projection in this same data sheet, the level

will rise to 6.2 in the year 2050. This increasing disparity

is expected to continue into the foreseeable future, again

assuming that growth rates remain constant. The more

developed countries are all of those in Europe and North

America (i.e., Canada and the United States), as well as

Australia, Japan, and New Zealand. The less developed

ones comprise the world’s remaining countries. The re-

markable disparity in growth patterns between the more

and less developed countries also is evident by compar-

ing the rates of natural increase between the two regions.

For the more developed countries, the figure is 0.1%, and

for the less developed ones 1 .4%. Thus, the growth rate

for the less developed region is 14 times greater than that

for the more developed area.

Because this paper deals with the Central American

herpetofauna, we will examine the population growth

trends in this region. The mid-2013 population for the

seven Central American nations is 45.2 million (PRB

2013 Population Data Sheet: 8). The rate of natural in-

crease ranges from a low of 1 .2 in Costa Rica and El Sal-

vador to 2.6 in Guatemala; the latter figure is 2.2 times

greater than that of the former. Thus, the doubling time

in Costa Rica and El Salvador is 58.3 years, the same

as for the entire globe. That for Guatemala, however, is

26.9 years, which is slightly more than for Nigeria (25.0

years); as noted above, Nigeria is projected to become the

world’s third most populous nation by 2050. The average

rate of natural increase for all of Central America is 1.8,

which provides a doubling time of 38.9 years. Assuming

no change in the average growth rate, the population of

Central America would double to 90.4 million by about

2052. The growth rate for the region is predicted to de-

crease, however, so the PRB 2013 Population Data Sheet

provides an estimate of 74 million by 2050. Nonetheless,

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Amphib. Reptile Conserv.

48

Johnson et al.

this figure is about 29 million more than the mid-2013

figure, a 64.2% size increase.

The decrease in growth rate is relatively good news,

but largely will depend on the rate of increase in the use

of contraceptives and the consequential frequency of

decrease of the total fertility rate. The current average

percentage rate for the use of all types of contraceptives

by married women ages 1 5—49 is 65.9% (PRB Popula-

tion Data Sheet 2013). This statistic, however, does not

consider the use of contraceptives by unmarried women

or women outside of the usual reproductive age range,

or the failure rate of contraceptive use by women or the

use of contraceptives by men. Assuming that the rate of

natural increase will decrease in the next 36 years to al-

low for a population of 73.5 million by the year 2050,

this growth pattern will measurably increase the impact

of population pressure on the remaining natural areas in

Central America. The current average density of human

population in the region is 103. 1 people per km 2 , and this

figure should grow to 167.7 in the intervening 36 years.

The rate of deforestation can be expected to be roughly

comparable to that of the addition of people to the popu-

lation. Deforestation, therefore, can be predicted to con-

tinue, especially given the income disparity in the region.

The average percentage comprising the poorest one-fifth

of the population is 3.7, and the wealthiest one-fifth 55.3.

These figures exceed those for the entire world, which

are 6.7 and 45.8, respectively (PRB Population Data

Sheet 2013).

Our examination of the average figures for popula-

tion growth and related factors tell only a portion of the

story. The average figures hide rather sizable disparities

in these statistics on a country level. When we examined

these statistics on a country basis, it became evident that

Guatemala is faced with the most serious problems and

Belize the least. Of the 45.2 million people currently in-

habiting Central America, 15.4 million (34.1%) live in

Guatemala; the next most populous country is Honduras,

with 8.6 million (19.0%); and the least populated country

is Belize with 0.3 million (0.7%). Any reduction in the

human population growth rate in Central America would

be highly desirable in terms of biodiversity conservation,

but will the projected decrease in growth rate be suffi-

cient to allow for the continued protection of this biodi-

versity?

The most significant reason for biodiversity decline

generally is conceded as habitat destruction, fragmenta-

tion, and degradation (Raven and Berg 2004; Vitt and

Caldwell 2009). This premise is easy to understand,

because the word habitat is defined as “the local envi-

ronment in which an organism, population, or species

lives” (Raven and Berg 2004). Living organisms derive

the resources to support their lives and their efforts at

reproduction from their habitats. The relationship be-

tween an organism and its habitat has evolved over time,

and thus is an outcome of the evolutionary process. An-

thropogenic damage to habitats reduces the capability of

the resident organisms to survive and reproduce in their

natural homes. The extent of such damage is evident

in the following statement in Vitt and Caldwell (2009):

“Humans have modified the environment everywhere.”

They further noted that “such a comment may seem to be

an exaggeration, but it is not an overstatement . . . Glob-

ally, our activities have resulted in a rising average an-

nual temperature and in a rise in ultraviolet radiation at

the earth’s surface. These climatic effects are only one

facet of our environmental alteration, which ranges from

global climatic change to the local loss of a marsh or a

patch of forest.”

Habitat alteration proceeds at a rate commensurate

with the following three principal factors: 1) an increase

in the number of people inhabiting the Earth; 2) an in-

crease in standards of living; and 3) the level of techno-

logical advantage enjoyed by these people. These three

factors have a combined environmental effect that is de-

scribed by the formula I = PAT, in which I stands for

“human impact,” P for “population,” A for “affluence,”

and T for “technology” (Chertow 2000). This formula

describes how our growing population, affluence, and

technology contribute to our increasing environmental

impact. It also predicts that the increase in any one of

these factors, or in any combination, can increase the

amount of environmental impact felt not only by us, but

also by the biosphere at large. This formula also predicts

that environmental impact can increase as a consequence

of rising affluence and technological capability, most

evident in the more developed countries, just as it does

with increasing population numbers, most evident in the

less developed countries. Thus, environmental impact

arises from all outcomes of the human experiment on our

planet. Nonetheless, not all technological advances are

undesirable (Chertow 2000). What is undesirable is hu-

manity’s willingness to augment the undesirable aspects

of such technology, i.e., planned obsolescence, lack of

recycling of resources, accumulation of pollutants, and

so forth.

The human experiment has been an effort, ostensibly

successful, to move away from being under the control

of the environmental limiting factors that impinge on all

organisms. In human terms, this has meant attempting

to improve the standards of living of human beings. No

matter how desirable this effort might be, however, it

has resulted in the creation of an unsustainable society,

of which the defects and the consequences are becom-

ing increasingly apparent over time. Perusal of the data

on income distribution in the PRB 2013 Population Data

Sheet is informative in this regard. Improvements to stan-

dards of living have been more beneficial to the wealthy

than the poor, both at the global and individual levels.

Currently, the distinction in the gross national income in

purchasing power parity (GNI PPP) between the more

developed and less developed sectors is startling; in the

former it is $35,800 and in the latter $6,600, a disparity

of 5.4 times between the two. The PRB data also indicate

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Amphib. Reptile Conserv.

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Conservation reassessment of Central American herpetofauna

Oedipina koehleri. This worm salamander is limited in distri-

bution to three isolated montane regions in northern Nicaragua,

where it occurs in Premontane Moist and Premontane Wet for-

ests at elevations from about 600 to 945 m. We estimated its

EVS as 16, placing it in the middle portion of the high vulner-

ability category, but its IUCN status has not been assessed. This

individual is from the Reserva Natural Cerro Musun, depart-

ment of Matagalpa. Photo by Javier Sunyer.

Rhinobothryum bovallii. This arboreal false coralsnake occurs

from southeastern Honduras to northwestern Venezuela and

northwestern Ecuador, where it occurs in Lowland Moist and

Wet forests at elevations from near sea level to 550 m. We cal-

culated its EVS as 16, placing it in the middle portion of the

high vulnerability category, and its IUCN status is Least Con-

cern. This individual is from Guayacan, Costa Rica. Photo by

Tobias Eisenberg.

Rhinoclemmys funerea. The Black River Turtle is distributed

from the Rio Coco on the border between Honduras and Nica-

ragua southward to central Panama, where it occurs in Lowland

Moist Forest at elevations from near sea level to 600 m. We

established its EVS as 16, placing it in the middle portion of the

high vulnerability category, and its IUCN status is Near Threat-

ened. This individual is from the Rio Puerto Viejo, Sarapiqui,

province of Alajuela, Costa Rica. Photo by Alejandro Solor-

zano.

Oscaecilia osae. This caecilian is endemic to the Golfo Dulce

region of southwestern Costa Rica, where it occurs in Lowland

Moist Forest at elevations from near sea level to 40 m. We cal-

culated its EVS as 19, placing it in the upper portion of the high

vulnerability category, and its IUCN status is Data Deficient.

This individual is from La Gamba, province of Puntarenas.

Photo by Peter Weish.

that the percentage growth of the gross domestic product

(GDP) has been decreasing both in the more developed

and less developed portions of the world. In the more

developed nations, the percentage dropped from 6.3 dur-

ing the period of 2000-2006 to 1 .9 during 2007-2011 . In

the less developed nations, the drop was from 10.2 to 7.8.

As unnerving as these statistics are, living in the more

developed portion of the world does not confer insula-

tion from economic disparity. The percent share of in-

come between the poorest one-fifth and the richest one-

fifth in the less developed and more developed regions

of the world essentially is the same (6.7 and 46.3 in the

former, 6.7 and 43.4 in the latter). This economic reality

is relevant in the United States, where the PRB report

(p. 4) concludes that “the rich get richer and the poor

get poorer,” a common way to characterize this dispar-

ity. Moreover, “despite having one of the world’s highest

standards of living, the gap between the income share

of the wealthiest and poorest households in the United

States is one of the widest among industrialized coun-

tries” and has increased over time. In 1967, the richest

one-fifth controlled 43.6 percent of household income,

compared to 4.0 percent for the poorest one-fifth. In

2011, the poorest one-fifth of households received only

3.2 percent of total national household income, while the

wealthiest one-fifth received 51.1 percent. This inequal -

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Amphib. Reptile Conserv.

50

Johnson et al.

ity is expected to continue to grow and the economy of

the United Sates will continue to suffer, even though this

country’s economy is discussed widely on a daily basis.

So, the affluence factor in the I = PAT formula looks a

bit shaky.

Since Earth presents a finite quantity of area for the

human population to occupy, the density of this popu-

lation will continue to increase with time. Actually, the

amount of habitable land will continue to decrease with

time, as a predicted consequence of global warming (In-

tergovernmental Panel on Climate Change [IPCC] Ap-

proved Climate Change Summary for Policymakers: 27

September 2013). Currently, the average density of the

population in the less developed world is 71 people per

square kilometer, which is 2.6 times greater than that

in the more developed world (27/km 2 ). The increasing

movement of people from rural to urban areas across the

globe worsens this overall pattern. As noted in the PRB

report, “in 1950, 117 million people lived in the top 30

[metropolitan areas] but that number rose to 426 million

by 2011. In 1950, 19 of the top 30 [“mega-cities”] were

in industrialized countries. By 2011, that number had

shrunk to eight. In 1950, Delhi was not even in the top

30 but it is now second behind only Tokyo. Such phe-

nomenal growth usually is due to rural-urban migration,

as migrants seek a better life in cities. Unfortunately, the

better life being sought often proves illusory, inasmuch

as rural areas are those that provide the resources neces-

sary to support life in both rural and urban areas. The

economic investment necessary to support people in ur-

ban settings increases the impact on the resource base in

rural regions. As these unsustainable practices continue

environmental degradation mounts, and the impact on

the remaining natural areas increases commensurately.

These features of human social evolution portend

disaster for the maintenance of biodiversity. Economic

primacy, especially in the more developed world, and

uncontrolled population growth, especially in the less

developed world, combines to create an unsustainable

society for humanity (Raven and Berg 2004). Unsustain-

ability increases the environmental pressure on organ-

ismic populations. Increasing environmental pressure

promotes increasing endangerment of the other members

of the living world. Thus, the job for conservation biolo-

gists grows more diffcult with the passing of time. Con-

sequently, the time lost to inaction becomes increasingly

important.

Perhaps the most unfortunate aspect of attempts at

conserving biodiversity is that the most biodiverse ar-

eas overlap those that support the most rapidly growing

human populations. As posted at the Conservational In-

ternational website (www.conservation.org/hotspots),

“the world’s most remarkable places are also the most

threatened.” The most biodiverse areas of the planet have

been termed “biodiversity hotspots.” Thirty-four such

areas are recognized (www.conservation.org/hotspots).

Four of these areas, as recognized by Conservation In-

ternational, lie in North and Central America. Almost all

of two of these areas, however, lie in what we define as

Mesoamerica, i.e., Mexico and Central America (Wilson

and Johnson, 2010). These two are tenned the Madrean

Pine-Oak Woodlands and Mesoamerica; the latter name a

different usage of the term than that of Wilson and John-

son (2010). The former encompasses the main mountain

chains in Mexico and isolated islands in Baja California,

and the southern United States (actually the southwest-

ern United States in southeastern Arizona, southwestern

New Mexico, and southwestern Texas). Apart from the

northernmost portions lying in the southwestern United

States, the remainder of this hotspot lies in Mexico. The

other hotspot includes the lowland and premontane areas

from northern Sinaloa on the Pacific versant and the Gulf

coastal plain as far north as Tampico, Tamaulipas, on the

Atlantic versant south to eastern Panama. This hotspot

encompasses essentially all of Central America. Al-

though the Mesoamerican forests, as defined by Conser-

vation International, constitute the third largest hotspot

in the world, the original extent of 1,130,019 km 2 has

been reduced to 226,004 km 2 (to 20.0% of the original).

Of the original extent, only 142,103 km 2 (12.6%) are

protected, with only 63,902 km 2 (5.7%) afforded higher

levels of protection. We presume that the relative figures

for the entire hotspot also apply to its portion in Central

America.

Ultimately, answering all the questions about biodi-

versity conservation will depend on finding fundamental

answers to the questions about why biodiversity decline

occurs. Until we uncover why humans represent such a

great threat to the rest of the planet’s organisms, i.e., why

they have assembled themselves into unsustainable soci-

eties of one sort or another, we will have no hope of de-

vising lasting solutions to this problem. Even though we

do not intend to explore this subject in depth, at least we

can offer what we consider some important comments

indicating the seriousness of biodiversity decline.

1 . If, as Wake and Vredenburg (2008) reported “we

are entering or in the midst of the sixth great mass

extinction,” and that “intense human pressure,

both direct and indirect, is having profound ef-

fects on natural environments,” then our species is

predicted to be responsible for a mass extinction

episode that will be equivalent in impact to those

that have preceded it. Scientists have documented

that “in each of the five events” generally thought

to have occurred during Earth’s history, “there was

a profound loss of biodiversity during a relatively

short period” (Wake and Vredenburg 2008). “The

most recent mass extinction was at the Cretaceous-

Tertiary boundary (~65 Mya); 16% of the families,

47% of the genera of marine organisms, and 1 8%

of the vertebrate families were lost. Most notable

was the disappearance of nonavian dinosaurs;

causes continue to be debated (Wake and Vreden-

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Conservation reassessment of Central American herpetofauna

Sphaerodactylus homolepis. This gecko is distributed from extreme southeastern Nicaragua to north-central Panama, where it oc-

curs in Lowland Moist and Wet forests at elevations from near sea level to 600 m. We established its EVS as 16, placing in the

middle portion of the high vulnerability category, and its 1UCN status is Least Concern. This individual is from the province of

Bocas del Toro, Panama. Photo by Adam G. Clause.

burg 2008). Given that whole genera and families

of organisms, including vertebrates, disappeared

during this most recent event, then a central ques-

tion for humanity is whether the progenitor of the

sixth mass extinction episode will survive its own

malevolent creation.

2. Organisms persist on our planet because sufficient

quantities of resources exist over time to support

their populations. These resources arise from the

atmosphere, hydrosphere, and lithosphere, as well

as from the sphere of life. The three abiotic spheres

interact among themselves and with the biosphere,

and these interactions allow life to exist and persist

on our planet. These statements are very simple

and can be confirmed by a cursory examination of

any ecology or environmental science textbook;

however, humanity proceeds as though its collec-

tive actions are exempt from these fundamental

rules of survival.

3. Natural science is one of the principal intellectual

undertakings of the human species (Wilson 1998).

What we know about the natural world is the re-

sult of the application of scientific methodology to

the endless questions that arise from our bound-

less curiosity. The design of science and its use is

the result of the way in which rationality operates.

We generally consider that humans are the best

exemplars of the rational being. Only a few other

creatures (e.g., cetaceans) are thought to have the

mental ability to compare favorably with our ratio-

nal capacity. No other organism, however, has the

benefit of our brain design coupled with bipedal

posture and an opposable thumb on a five-fingered

hand. Interestingly, finding an operational defini-

tion of rationality is elusive; the effort commonly

results in the construction of circular definitions

(i.e., definitions that do not actually define, but

eventually lead back to the word one is attempting

to define). Irrespectively, rationality is a function

of our nervous system that allows for the connec-

tion of cause to effect from the past through the

present to the future. It allows us to understand the

consequences of our actions. Strangely, rationality

also allows us to “ignore” the consequences of our

actions. Thus, the use of scientific methodology,

which is one outcome of rationality, can allow us to

ask and answer questions about the natural world,

within limits, but whether the answers lead to ap-

Amphib. Reptile Conserv.

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Johnson et al.

propriate actions depends on a number of other

factors, such as can be understood from the view-

ing of any day’s events in the human world.

4. Scientific advance depends on the use of scientific

methodology to generate tangible and sometimes

reproducible evidence to falsify hypotheses in or-

der to support philosophies. In turn, assembling

such evidence depends upon the functioning of our

sensory structures, as assisted by scientific instru-

mentation. Other sorts of systems exist, however,

that do not depend on structuring ideas based on

evidence. Many people use these types of be-

lief systems in the conduct of their lives (Ehrlich

and Ehrlich 1996). To illustrate our meaning, we

will use the example of evolution. As any biolo-

gist knows, the theory of evolution is the central

concept of modem-day biology; examination of

any modern-day university-level biology text-

book will confirm this statement (e.g., Reese et al.

2013). Broad-based disciplines such as environ-

mental science and conservation biology have the

same conceptual foundation. Among the general

public, however, the reality of the evolutionary

process often is thought to be a matter of opin-

ion. The word “opinion” is defined as “a belief or

conclusion held with confidence but not substanti-

ated by positive knowledge or proof’ (American

Heritage Dictionary, 3 rd edition). A recent report

(30 December 2013) of the Pew Research Center

(www.pewresearch.org; accessed 2 January 2014)

entitled “Public’s Views on Human Evolution” is

based on telephone interviews conducted from 2 1

March to 8 April 2013 among a national sample

of 1,983 adults (age 18 and older) living in all 50

U.S. states. The question asked of the respondents

was whether “humans and other living things

have existed in their present form since the begin-

ning of time, or humans and other living things

have evolved over time.” Thirty-three percent of

the respondents agreed with the former statement

and 60 percent with the latter. The greatest diver-

gence from the results for all adults was evidenced

among white evangelical Protestants (64 vs. 27%),

which also was the largest group to think that evo-

lution has been guided by a supreme being (36 vs.

36%), and Republicans (43 vs. 48%). Since the

Pew Research Center’s survey questions ask for

yes or no responses, the basis for the variation in

the responses was not explored, although it seems

unlikely that it has to do with the scientific exami-

nation of the evidence for the theory of evolution

through natural selection.

5. Climate change is another issue subject to the vaga-

ries of public opinion. This term refers to the phe-

nomenon of the anthropogenic alteration of global

Amphib. Reptile Conserv. 53

climatic patterns. In the sense of this definition,

climate change is an environmental superproblem,

in the sense of Bright (2000). Wilson and McCra-

nie (2004) reflected that Bright “uses this term to

describe environmental synergisms resulting from

the interaction of two or more environmental prob-

lems, so that their combined effect is greater than

the sum of their individual effects. These problems

represent an environmental worst-case scenario —

the point when environmental problems become

so serious that they produce unanticipated results,

the successful resolution of which threatens to slip

forever from the grasp of humanity.” This global

superproblem has been studied by the Intergovern-

mental Panel of Climate Change (IPCC), which

released its latest report in September of 2013. The

panel produced an “approved summary for poli-

cymakers,” which includes several conclusions of

great importance. The most significant conclusion

is as follows (p. 3): “Warming of the climate sys-

tem is unequivocal, and since the 1950s, many of

the observed changes are unprecedented over de-

cades to millenia. The atmosphere and ocean have

warmed, the amounts of snow and ice have dimin-

ished, sea level has risen, and the concentrations

of greenhouse gases have increased.” With respect

to the atmosphere, the report concluded that, “each

of the last three decades has been successively

warmer at the Earth’s surface than any preceding

decade since 1850. In the Northern Hemisphere,

1983-2012 was likely the warmest 30-year peri-

od of the last 1,400 years (medium confidence).”

Concerning the ocean, the report concluded that,

“ocean warming dominates the increase in energy

stored in the climate system, accounting for more

than 90% of the energy accumulated between

1971 and 2010 (high confidence). It is virtually

certain that the upper ocean (0-700 m) warmed

from 1971 to 2010.” The IPCC report summary

also indicated that with regard to the cryosphere

“over the last two decades, the Greenland and ant-

arctic ice sheets have been losing mass, glaciers

have continued to shrink almost worldwide, and

Arctic sea ice and Northern Hemisphere spring

snow cover have continued to decrease in extent

(high confidence).” As a consequence of this dimi-

nution of ice and snow at the polar regions, “the

rate of sea level rise since the mid- 19th century

has been larger than the mean rate during previous

two millennia (high confidence). Over the period

of 1901-2010, global mean sea level rose by 0.19

[0.17 to 0.21] m .” Finally, the report indicated that,

“the atmospheric concentrations of carbon dioxide

(CO.,), methane, and nitrous oxide have increased

to levels unprecedented in at least the last 800,000

years. CO., concentrations have increased by 40%

since pre-industrial times, primarily from fossil

August 2015 | Volume 9 | Number 2 | el 00

Conservation reassessment of Central American herpetofauna

Tantilla vermiformis. This centipede snake is distributed from El Salvador to northwestern Costa Rica, where it occurs in Lowland

Dry Forest at elevations from near sea level to 520 m. We evaluated its EVS as 14, placing it at the lower end of the high vulner-

ability category, and its 1UCN status is Least Concern. This juvenile individual is from Volcan Masaya, Nicaragua. Photo by Jose

Gabriel Martinez-F onseca.

fuel emissions and secondarily from net land use

change emissions. The ocean has absorbed about

30% of the emitted anthropogenic carbon dioxide,

causing ocean acidification.” Taken in their en-

tirety, these conclusions about the anthropogenic

impact on the global climate system are extreme-

ly frightening and portend future environmental

changes that will have worldwide effects of hugely

significant consequence. These conclusions also

point very clearly to the way in which the litho-

sphere, the home of humanity, interacts with the

atmosphere and how the atmosphere interacts with

the hydrosphere and, in turn, the lithosphere. Thus,

climate change is a best-case example of how an

environmental superproblem evolves. In light of

the general high confidence levels for the summary

statements in the IPCC report, we examined the

results of a Pew Research Center report published

5 November 2013 (available at www.pewresearch.

org) and entitled “Climate Change: Key Data

Points from Pew Research,” which concluded that

“the American public routinely ranks dealing with

global warming low on its list of priorities for the

president and Congress. This year, it ranked at the

bottom of the 21 tested.” Of the people surveyed

in January of 2013, just 28% indicated that dealing

with global warming is a top priority. This statis-

tic contrasts most markedly with strengthening the

economy, which was identified as a top priority by

86% of the survey respondents. Even dealing with

“moral breakdown” at 40% beat out global warm-

ing as a top priority. Interestingly, people in the

United States, who collectively are major contribu-

tors to global climate change, fell behind people in

most other countries in recognizing global climate

change as a major threat. Beyond all this opinion,

some people opine that global warming is “just not

happening.” Another view of the significance of

global climate change is provided in the report of

the World Economic Forum entitled “Outlook on

the Global Agenda 2014” (2013). One portion of

this report identifies the Top Trends of 2014. In-

terestingly, “inaction on climate change” is on the

list, but only at spot number five and after “rising

societal tensions in the Middle East and North Af-

rica,” “widening income disparities,” “persistent

structural unemployment,” and “intensifying cyber

threats ” Addressing issues of biodiversity decline,

however, does not appear on the list. Given the

glacial pace at which scientific research results are

transformed into governmental policy and, beyond

that, into sufficiently comprehensive plans of ac-

tion that are put into effect, the question obviously

arises as to whether humanity, even with its vaunt-

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Johnson et al.

ed rational capacity, has the wherewithal to deal

with the gargantuan problems of its own creation,

especially since those problems increase in sever-

ity at a rate commensurate with the exponential

growth of human population. As always, however,

time will tell.

6. Given that all of humanity is faced with environ-

mental superproblems, exemplified by global cli-

mate change, and that these problems originate

in planetary spheres remote from human control,

the question arises as to what effect these super-

problems will have on efforts to conserve organ-

ismic populations in particular, and the structure

and function of the biosphere in general. The bio-

sphere, the entire compendium of life on Earth,

exists at the interface of the three abiotic spheres

based on the retrieval of resources from them. In-

asmuch as the three abiotic spheres and their inter-

relationships evolve over time, the biosphere gen-

erally persists over time by also evolving to adapt

to these environmental changes. The adaptability

of organisms depends on the process of evolution

according to natural selection, which obviously is

a powerful enough force to allow life on Earth to

survive several mass extinction episodes that date

back to as far as ~439 My a (Wake and Vredenburg

2008). All of these past episodes have been geo-

logical in nature. As noted by Wake and Vreden-

burg (2008), “many scientists think that we are just

now entering a profound spasm of extinction and

that one of its main causes is global climate change

... Furthermore, both global climate change and

many other factors (e.g., habitat destruction and

modification) responsible for extinction events are

directly related to activities of humans” Thus, per-

haps the major question facing humanity now and

in the future is what portion of the biosphere will

disappear into the extinction void, and if ultimately

humans will join these other unfortunate creatures.

7. Presently we do not know the answers to these

fundamental questions, but we are beginning to

understand the extent of the impact on selected

groups of organisms, especially the best known.

Most zoologists work on vertebrate animals and

we three are among them. As herpetologists work-

ing in one of Earth’s most significant biodiversity

hotspots (Mesoamerica), and attempting to assess

the conservation status of the herpetofaunal species

resident in this hotspot, we offer some ideas about

how the sixth mass extinction episode will impact

these creatures. We bring to this subject some 102

person-years of experience, as judged by the date

of publication of the first scientific paper for each

of us. All three of us were involved in the produc-

tion of the 2010 volume entitled Conservation

Amphib. Reptile Conserv. 55

of Mesoamerican Amphibians and Reptiles , and

last year we coauthored two papers in the Special

Mexico Issue of the journal Amphibian & Reptile

Conservation entitled “A conservation reassess-

ment of the reptiles of Mexico based on the EVS

measure” and “A conservation reassessment of

the amphibians of Mexico based on the EVS mea-

sure.” Other herpetologists also have weighed in

on these questions, most importantly Gibbons et al.

(2000), Wake and Vredenburg (2008), Stuart et al.

(2010), and Bohm et al. (2013). The Gibbons et al.

(2000) study was written in part to document that

crocodylians, squamates, and turtles are undergo-

ing population declines similar in scope on a global

scale “to those experienced by amphibians in terms

of taxonomic breadth, geographic scope, and se-

verity.” Bohm et al. (2013) presented “the first-

ever global analysis of extinction risk in reptiles,

based on a random representative sample of 1,500

species (16% of all currently known species)” and

concluded that, “nearly one in five reptilian species

[is] threatened with extinction, with another one in

five species classed as Data Deficient.” They fur-

ther concluded that, “conservation actions specifi-

cally need to mitigate the effects of human- induced

habitat loss and harvesting, which are the predomi-

nant threats to reptiles.” The Stuart et al. (2010)

paper reiterated the Global Amphibian Assessment

analysis presented in the Stuart et al. (2004) study

and concluded that “a plethora of threats impact

amphibian species globally, with habitat loss and

degradation being the principal threat followed by

pollution. Disease is a less significant threat on a

global scale, but can bring about rapid population

declines leading to extinction. Deforestation is a

significant threat to amphibian population stabil-

ity, inasmuch as the vast majority of species de-

pend on forest for their survival. A sizable number

also depends on flowing and still freshwater habi-

tats, largely due to their biphasic lifestyle. If the

observed declines are not quickly understood and

reversed, hundreds of species of amphibians will

face extinction within the next few decades.” Fi-

nally, Wake and Vredenburg (2008) attempted to

answer the question “Are we in the midst of the

sixth mass extinction?” using amphibians as a test

group. These authors concluded in the most sweep-

ing way of any of these four papers that “multiple

factors acting synergistically are contributing to the

loss of amphibians. But we can be sure that behind

all of these activities is one weedy species, Homo

sapiens, which has unwittingly achieved the ability

to directly affect its own fate and that of most of the

other species on this planet. It is an intelligent spe-

cies that potentially has the capability of exercis-

ing necessary controls on the direction, speed, and

intensity of factors related to the extinction crisis.

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Conservation reassessment of Central American herpetofauna

Education and changes of political direction take

time that we do not have, and political leadership

to date has been ineffective largely because of so

many competing, short-term demands. A primary

message from the amphibians, other organisms,

and environments, such as the oceans, is that little

time remains to stave off mass extinction, if it is

possible at all ” (emphasis ours). Using the conclu-

sions of Wake and Vredenburg (2008) as a starting

point, we provide our conclusions and recommen-

dations on the conservation status of the Central

American herpetofauna.

Conclusions and Recommendations

One or more of us previously have provided sets of con-

clusions and recommendations for addressing the issues

of conservation of the Mesoamerican herpetofauna (Wil-

son and Townsend 2010; Wilson et al. 2013a, b). We used

this information as a partial framework and starting point

for our conclusions and recommendations concerning the

conservation of the Central American herpetofauna.

1. Biodiversity decline is an environmental problem

of global dimensions, comparable to the more

commonly publicized problem of climate change.

Both of these environmental superproblems exist

because of human action and inaction, exacerbated

by humanity’s anthropocentric focus.

2. Our work deals with the scientific study of the her-

petofauna, of which all groups are prominent com-

ponents of terrestrial ecosystems in temperate and

tropical regions across the globe. Only crocodyl-

ians, squamates, and turtles have made relatively

limited inroads into marine habitats. Some of our

earlier work dealt with the conservation status of

the herpetofauna of Mexico; in this study, we are

concerned with the herpetofauna of Central Amer-

ica.

3. Central America is a major component of Meso-

america, the other component consisting of Mex-

ico. Together, these two regions contribute to and

extend beyond the limits of the third largest of the

34 biodiversity hotspots identified by Conserva-

tion International. The herpetofauna of Central

America is of major significance and presently

consists of 493 amphibians and 559 crocodylians,

squamates, and turtles, for a total of 1,052 species.

Our knowledge of the dimensions of this herpeto-

fauna will continue to augment with time. In the in-

terim between 31 December 2008 and the present,

92 species have been added to this herpetofauna,

an increase of 9.7% percent over the number con-

sidered in Wilson and Johnson (2010). Presently,

there are more amphibians in Central America than

in Mexico (493 vs. 383), and more crocodylians,

squamates, and turtles collectively in Mexico than

in Central America (869 vs. 559). Although more

amphibians, crocodylians, squamates, and turtles

occur in Mexico than in Central America (1,252

vs. 1,052), Mexico is about three and three-quar-

ters the size of Central America, indicating signifi-

cantly greater herpetofaunal numbers per unit area

in Central America than in Mexico.

4. Herpetofaunal endemism also is significant in Cen-

tral America. Of the 493 amphibians known from

the region, 324 (65.7%) are endemic. Of the 559

reptiles found there, 261 (46.7%) are endemic. The

entire herpetofauna is characterized by an endemic -

ity of 55.6%. These figures are fairly comparable

to those for Mexico. Amphibian endemism is only

slightly higher in Mexico than in Central America

(67.4 vs. 65.7%). Endemism of the remainder of

the herpetofauna is about 11 percentage points

higher in Mexico than in Central America (57.4

vs. 46.7%). Endemism for the total herpetofauna

is only a few percentage points higher in Mexico

than in Central America (60.4% vs. 55.6%). Thus,

more than one-half of the Central American herpe-

tofauna is endemic to the region, compared to six

of every 10 species in Mexico.

5. The IUCN employs the most commonly used

means of conservation status assessment. The

implementation of this system, however, is expen-

sive, time-consuming, slow to respond to system-

atic advances, and likely to resort to the Data Defi-

cient category when assessing taxa described from

single specimens and/or single localities, and to the

Least Concern category as a kind of conservation

“dumping ground” for species that deserve a more

careful examination.

6. Given the problems we see with the use of the

IUCN system of categorizaitons, we employed a

revised Environmental Vulnerability Score (EVS)

measure that allowed us to address the deficiencies

of the IUCN system and to provide a conservation

assessment for all of the species now known to

comprise the Central America herpetofauna. The

EVS values can range from 3-20 and are placed in

three categories: low (3-9); medium (10-13); and

high (14-20). Our calculations indicate that the

EVS values for amphibians are categorized as fol-

lows: low (39 species of 493 [7.9%]); medium (105

[21.3%]); and high (349 [70.8%]). For the croco-

dylians, squamates, and turtles, the values are: low

(81 of 552 [14.7%]); medium (162 [29.3%]); and

high (309 [56.0%]). For the entire herpetofauna,

the values are: low (119 of 1,045 [11.4%]); medi-

um (267 [25.6%]); and high (658 [63.0%]). Thus,

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Johnson et al.

Ungaliophis panamensis. This small arboreal boa is found on the Atlantic versant from southeastern Nicaragua to northwestern

Colombia, and on the Pacific versant from northwestern Costa Rica to western Panama, where it occurs in Lowland Moist and Wet,

Premontane Wet, and Lower Montane Wet forests at elevations from near sea level to 2,100 m. We gauged its EVS as 12, placing

it in the upper portion of the medium vulnerability category, but its IUCN status has not been determined. This individual is from

the Rio Indio Lodge located in the Indio Maiz Biological Reserve, department of Rio San Juan, in southeastern Nicaragua. Photo

by Javier Sunyer.

our analysis indicates that more than six of every

10 herpetofaunal species are highly vulnerable to

environmental damage from anthropogenic causes.

7. In 2013, we conducted a similar study of the Mexi-

can herpetofauna. When comparing our results for

Central America and Mexico, a greater proportion

of amphibians in Central America fell into the high

vulnerability category than in Mexico (70.8% vs.

58.8%). In both regions, salamanders are the most

vulnerable when compared to anurans and caeci-

lians. Among the rest of the herpetofauna, howev-

er, we found about the same proportion in the high

vulnerability category in Central America (56.0%)

as in Mexico (55.9%). Considering the two highest

species groups (lizards and snakes), in both Mex-

ico and Central America lizards are more vulner-

able to environmental damage than snakes.

8. Given the length of time it takes for an IUCN as-

sessment to appear at the Red List website after a

new species is described and the expense involved

to produce such an assessment, we recommend that

the original describers provide at least an estimate

of the conservation status of the taxon in ques-

tion in the original description. In addition, since

this task might be difficult to undertake, given the

deficiencies of the IUCN system we have identi-

fied here and elsewhere, we also recommend that

the original describers calculate an Environmental

Vulnerability Score to provide an additional as-

sessment of the conservation status for the species

being described.

9. Assessments of the conservation status of any

group of organisms essentially remain academic

exercises, unless sufficient attention is provided

to the imperatives underlying the threats to biodi-

versity created by humanity. Humanity lives un-

sustainably on planet Earth. The pressure placed

on limited resources by an exponentially growing

human population creates this reality. Humans are

cosmopolitan animals that become more so with

the passage of time. The approach is the same

wherever one finds humans, as essentially it is a

unidirectional track from point A (what humans

want) to point B (what humans obtain), with the

minimal amount of possible diversion between

the two points. Unidirectionality, however, is not

a feature of the structure and function of Earth.

Amphib. Reptile Conserv.

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Conservation reassessment of Central American herpetofauna

Rather, this planet, especially the portion of most

concern to humanity, consists of four primary

spheres that intertwine among themselves to cre-

ate an environment in which humanity can exist.

All of these spheres, the atomsphere, hydrosphere,

lithosphere, and biosphere provide resources to our

species, without which its survival is impossible.

One way of looking at this matter is that human-

ity, in return for life support and from its perch on

Earth’s surface, favors these spheres with a pletho-

ra of environmental problems that retrace the same

pathways as exist in the natural world to make the

resources for life support available to humans. As

an example, burning forests and fossil fuels pumps

CO., into the atmosphere and this pollutant causes

its temperature to rise and creates global warming,

which in turn produces climate change that impacts

the planet’s solid and liquid surfaces. Burning for-

ests to make way for agriculture also degrades hab-

itats for the world’s creatures, especially those that

live on land, creating biodiversity decline.

10. More than two decades ago on 1 8 November 1992,

the Union of Concerned Scientists issued the

World Scientists’ Warning to Humanity (www.uc-

susa.org). To date, this statement has been signed

by “some 1,700 of the world’s leading scientists,

including the majority of Nobel laureates in the

sciences” (www.ucsusa.org/about/1992-world-sci-

entists.html; accessed 2 February 2014). The one-

paragraph introduction to the statement is cogently

powerful. “Human beings and the natural world

are on a collision course. Human activities inflict

harsh and often irreversible damage on the envi-

ronment and on critical resources. If not checked,

many of our current practices put at serious risk the

future that we wish for human society and the plant

and animal kingdoms, and may so alter the living

world that it will be unable to sustain life in the

manner that we know [emphasis ours]. Fundamen-

tal changes are urgent if we are to avoid the colli-

sion our present course will bring about.” For all

intents and purposes, we have lost the intervening

two decades to inaction and further encroachment.

1 1 . The warning to humanity contained a simple and

elegant statement of “what we must do.” This state-

ment consists of “five inextricably linked areas that

must be addressed simultaneously,” as follows:

“We must bring environmentally damaging ac-

tivities under control to restore and protect the

integrity of the earth’s systems we depend on.

We must, for example, move away from fossil fu-

els to more benign, inexhaustible energy sources

to cut greenhouse gas emissions and the pollution

of our air and water. Priority must be given to the

development of energy sources matched to Third

World needs — small-scale and relatively easy to

implement.”

“We must manage resources crucial to human

welfare more effectively. We must give high pri-

ority to efficient use of energy, water, and other

materials, including expansion of conservation and

recycling.”

“We must stabilize population. This wifi be pos-

sible only if all nations recognize that it requires

improved social and economic conditions, and the

adoption of effective, voluntary family planning.”

“We must reduce and eliminate poverty.”

“We must ensure sexual equality, and guaran-

tee women control over their own reproductive

decisions.”

12. Only within the context of simultaneously address-

ing the above-indicated “inextricably linked” so-

cial imperatives can we sensibly discuss “what

we must do” to safeguard organismic populations,

including those of the herpetofauna of Central

America. So, our most significant recommendation

is to address these imperatives in the shortest time

possible.

Acknowledgments. — Although this contribution

is dedicated to Fouis W. Porras, beyond his help in the

past, we would like to thank him sincerely for the various

courtesies he extended to us during the writing of this

paper, including a complete copy editing. We are also

grateful to Gunther Kohler, who checked for accuracy

our list of taxonomic additions and changes to the Cen-

tral American herpetofauna. We also owe an indirect debt

to the many contributors to the chapters of Conservation

of Mesoamerican Amphibians and Reptiles (Wilson et al.

2010), who were responsible for presenting data in their

work that has been of great use to us in this paper. We

also are indebted hugely to those colleagues who sup-

plied us with the excellent photographs that grace the

pages of this publication, including: Abel Batista, Edu-

ardo Boza Oviedo, Adam G. Clause, Robbie Eagleston,

Tobias Eisenberg, Brian Freiermuth, Gunther Kohler,

Brian Kubicki, Jose G. Martinez-Fonseca, Maciej Pabi-

jan, Antonia Pachmann, Silviu Petrovan, Todd Pierson,

Sean Michael Rovito, Roney Santiago, Alejandro Solor-

zano, Javier Sunyer, Josiah H. Townsend, Peter Weish,

and Brad Wilson. Finally, we are very appreciative of the

splendid efforts of Manuel Acevedo, Abel Batista, and

Javier Sunyer as reviewers to improve the quality and

accuracy of our paper.

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Amphib. Reptile Conserv.

58

Johnson et al.

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Addendum (changes past conclusion of analyses)

We chose a cut-off date of 1 March 2015 for which to discontinue revising the hundreds to thousands of numbers and

calculations dealing with the 1,052 herpetofaunal species in this paper. After this date, we continued adding pertinent

taxa and publications in this addendum, as follows:

(1) Hyalinobatrachium dianae. Kubicki et al. (2015) described this new species of glassfrog from the lowland and

premontane forests of Caribbean Costa Rica, which is known from the provinces of Heredia and Limon at elevations

from 400 to 900 m. Its EVS can be calculated as 5+7+3=15.

(2) Gerrhonotus liocephalus. Morales et al. (2015) reported this alligator lizard, formerly limited in distribution to

Mexico and Texas in the United States, from Guatemala, thus adding this species to the Central American herpeto-

fauna. Its EVS remains as 2+l+3=6.

(3) Ecnomiohyla bailarina. Kubicki and Salazar (2015) reported this fringe-limbed treefrog, formally known only

from the type locality in Panama, from the Caribbean foothills of southeastern Costa Rica. As a consequence, its EVS

needs to be recalcuated as 5+7+6=18.

(4) Holcosus spp. Meza-Lazaro and Nieto Montes de Oca (2015) revised the species Holcosus undulatus and elevated

nine former subspecies to species level in Mesoamerica, including five taxa in Central America {H. hartwegi, H. mia-

dis, H. parvus , H. pule her, and H. thomasi ). As a consequence, the ranges of these elevated taxa naturally are smaller

and the resulting EVS will be higher than that calculated in Appendix 2 for the former H. undulatus.

(5) Brady triton silus. Since its description in 1983, this plethodontid salamander species, the single member of its ge-

nus, has been considered endemic to Guatemala and, therefore, to Central America. Recently, however, a specimen was

collected by a field crew associated with Sean Rovito at San Francisco Jimbal in northern Chiapas, which constitutes

the first record for this species in Mexico (Bouzid et al. 2015). Thus, B. silus no longer is a Central American endemic.

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Conservation reassessment of Central American herpetofauna

Jerry D. Johnson is Professor of Biological Sciences at The University of Texas at El Paso, and has exten-

sive experience investigating the herpetofauna of Mesoamerica, especially in Mexico. Presently, he is the

Director of the 40,000 acre “Indio Mountians Research Station” located in the Chihuahuan Desert near the

Mexican border. Jerry is a co-editor of the recently published Conservation of Mesoamerican Amphibians

and Reptiles, is Mesoamerica/Caribbean section editor for Geographic Distribution segment of Herpetolog-

ical Review, and is an Associate Editor and Co-chair of the Taxonomic Board for the Journal Mesoamerican

Herpetology. Johnson has authored or co-authored 100 peer-reviewed papers, including two 2010 articles,

“Geographic distribution and conservation of the herpetofauna of southeastern Mexico” and “Distributional

patterns of the herpetofauna of Mesoamerica, a biodiversity hotspot,” as well as two 2013 articles in the

Special Mexican Edition of Amphibian & Reptile Conservation entitled, “A conservation reassessment of

the reptiles of Mexico based on the EVS measure, and “A conservation reassessment of the amphibians of

Mexico based on the EVS measure.” He was also Co-editor for the books Middle American Herpetology:

A Bibliographic Checklist and Mesoamerican Herpetology: Systematics, Zoogeography, and Conservation.

Vicente Mata-Silva is a herpetologist interested in ecology, conservation, geographic distribution, and the

monitoring of amphibians and reptiles in Mexico and the southwestern United States. His bachelor’s thesis

at the Universidad Nacional Autonoma de Mexico (UNAM) compared herpetofaunal richness in Puebla,

Mexico, in habitats with different degrees of human-related disturbance. Vicente’s master thesis focused

primarily on the diet of two syntopic whiptail lizard species, one unisexual and the other bisexual, in the

Trans-Pecos region of the Chihuanhuan Desert. His dissertation was on the ecology of the rock rattlesnake,

Crotalus lepidus, in the northern Chihuahuan Desert. To date, Vicente has authored or co-authored over 60

peer-reviewed scientific publications. Currently, he is a research fellow and lecturer at the University of

Texas at El Paso, where his work focuses on the ecology of rattlesnake populations in a Chihuahuan Desert

habitat; he also is a Distribution Notes Section Editor for the journal Mesoamerican Herpetology.

Larry David Wilson is a herpetologist with lengthy experience in Mesoamerica, totaling six and one-half

collective years (combined over the past 49). Larry is the senior editor of Conservation of Mesoamerican

Amphibians and Reptiles (2010) and the co-author of seven of its chapters. He is retired from 35 years of

service as a professor of biology at Miami-Dade College in Miami, Florida. Larry is the author or co-author

of over 3 1 5 peer-reviewed papers and books on herpetology, including the 2004 Amphibian & Reptile

Conservation paper entitled “The conservation status of the herpetofauna of Honduras” and the two 2013

papers entitled “A conservation reassessment of the amphibians of Mexico based on the EVS measure” and

“A conservation reassessment of the reptiles of Mexico based on the EVS measure.” His other books in-

clude The Snakes of Honduras (1985), Middle American Herpetology (1988), The Amphibians of Honduras

(2002), Amphibians & Reptiles of the Bay Islands and Cayos Cochinos, Honduras (2005), The Amphib-

ians and Reptiles of the Honduran Mosquitia (2006), and Guide to the Amphibians & Reptiles of Cusuco

National Park, Honduras (2008). For 33 years he served as the Snake Section Editor for the Catalogue of

American Amphibians and Reptiles. Over his career to date, he has authored or co-authored the descriptions

of 70 currently recognized herpetofaunal species and six species have been named in his honor, including

the anuran Craugastor lauraster and the snakes Cerrophidion wilsoni, Myriopholis wilsoni, and Oxybelis

wilsoni.

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Appendix 1. Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environmental Vulner-

ability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems. EVS category

abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode estimated based

on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Order Anura (319 species)

Family Aromobatidae (3 species)

Allob cites talamancae

LC

1

6

4

11

M

Anomaloglossus astralogaster *

NE

6

8

4?

18

H

Anomaloglossus isthminus *

NE

5

7

4?

16

H

Family Bufonidae (39 species)

Atelopus certus*

EN

5

8

1

14

H

Atelopus chiriquiensis *

CR

5

8

1

14

H

Atelopus chirripoensis *

CR

6

8

1?

15

H

Atelopus glyphus

CR

4

8

1

13

M

Atelopus limosus*

EN

5

8

1

14

H

Atelopus senex*

CR

5

7

1

13

M

Atelopus varius*

CR

5

1

11

M

Atelopus zeteki*

CR

5

7

1

13

M

Incilius aucoinae*

LC

5

8

1

14

H

Incilius aurarius

NE

4

8

1

13

M

Incilius bocourti

LC

4

6

1

11

M

Incilius campbelli

NT

4

7

1

12

M

Incilius canaliferus

LC

4

3

1

8

L

Incilius chompipe*

VU

5

7

1?

13

M

Incilius coccifer

LC

3

5

1

9

L

Incilius coniferus

LC

1

6

1

8

L

Incilius epioticus*

LC

5

7

4?

16

H

Incilius fastidiosus *

CR

5

7

1

13

M

Incilius guanacaste*

DD

5

8

4?

17

M

Incilius holdridgei *

CR

5

8

1

14

H

Incilius ibarrai*

EN

5

7

1

13

M

Incilius karenlipsae*

NE

6

8

1?

15

H

Incilius leucomyos *

EN

5

6

1

12

M

Incilius luetkenii

LC

3

1

7

L

Incilius macrocristatus

VU

4

6

1

11

M

Incilius melanochlorus*

VU

5

6

1

12

M

Incilius periglenes*

EX

6

8

1

15

H

Incilius peripatetes*

CR

5

8

1?

14

H

Incilius porteri*

DD

5

8

1?

14

H

Incilius signifer*

LC

5

8

1?

14

H

Incilius tacanensis

EN

4

1

9

L

Incilius tutelarius

EN

4

5

1

10

M

Incilius valliceps

LC

3

2

1

6

L

Rhaebo haematiticus

LC

1

7

1

9

L

Rhinella acrolopha

DD

4

8

4?

16

H

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Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Rhinella alata

DD

4

7

4?

15

H

Rhinella centralis*

LC

5

8

1

14

H

Rhinella chrysophora *

EN

5

7

1

13

M

Rhinella marina

LC

1

3

L

Family Centrolenidae (14 species)

Cochranella euknemos

LC

1?

6

3

10

M

Cochranella granulosa *

LC

5

7

3

15

H

Espadarana prosoblepon

LC

1?

5

3

9

L

Hyalinobatrachium aureoguttatum

NT

3

7

3

13

M

Hyalinobatrachium chirripoi

LC

2

7

3

12

M

Hyalinobatrachium colymbiphyllum

LC

1?

6

3

10

M

Hyalmobatrachium fleischmanni

LC

1?

4

3

8

L

Hyalinobatrachium talamancae *

LC

5

8

3

16

H

Hyalinobatrachium valerioi

LC

1?

7

3

11

M

Hyalinobatrachium vireovittatum *

DD

5

8

3

16

H

Sachatamia albomaculata

LC

2

7

3

12

M

Sachatamia ilex

LC

2?

7

3?

12

M

Teratohyla pulverata

LC

2?

7

3

12

M

Teratohyla spinosa

LC

1?

7

3

11

M

Family Craugastoridae (101 species)

Craugastor adamastus*

DD

6

8

4

18

H

Craugastor alfredi

vu

2

5

4

11

M

Craugastor amniscola

DD

4

6

4

14

H

Craugastor anciano*

CR

5

7

4

16

H

Craugastor andi*

CR

5

8

4

17

H

Craugastor angelicas *

CR

5

6

4

15

H

Craugastor aphanus*

VU

5

8

4

17

H

Craugastor aurilegulus*

EN

5

6

4

15

H

Craugastor azueroensis*

EN

5

7

4

16

H

Craugastor bocourti *

VU

5

7

4

16

H

Craugastor bransfordii*

LC

5

4

13

M

Craugastor brocchi

VU

4

6

4

14

H

Craugastor campbelli*

DD

5?

7

4

16

H

Craugastor catalinae*

CR

5

8

4

17

H

Craugastor chac*

NT

5

7

4

16

H

Craugastor charadra*

EN

5

6

4

15

H

Craugastor chingopetaca*

DD

6

8

4

18

H

Craugastor chrysozetetes*

EX

6

8

4

18

H

Craugastor coffeus *

CR

6

8

4

18

H

Craugastor crassidigitus

LC

2

6

4

12

M

Craugastor cruzi *

CR

6

8

4

18

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

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Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Craugastor cuaquero*

DD

6

8

4

18

H

Craugastor cyanochthebius*

NT

6

8

4

18

H

Craugastor daryi*

EN

5

8

4

17

H

Craugastor emcelae*

CR

5

8

4

17

H

Craugastor emleni*

CR

5

6

4

15

H

Craugastor epochthidius *

CR

5

7

4

16

H

Craugastor escoces*

EX

5

6

4

15

H

Craugastor evanesco*

NE

5

8

4

17

H

Craugastor fecundus *

CR

5

7

4

16

H

Craugastor fitzingeri

LC

2

6

4

12

M

Craugastor fleischmanni *

CR

5

7

4

16

H

Craugastor gollmeri*

LC

5

7

4

16

H

Craugastor greggi

CR

4

7

4

15

H

Craugastor gulosus*

EN

5

8

4

17

H

Craugastor inachus *

EN

5

8

4

17

H

Craugastor jota*

DD

6

8

4

18

H

Craugastor laevissimus*

EN

5

3

4

12

M

Craugastor laticeps

NT

4

12

M

Craugastor lauraster*

EN

5

7

4

16

H

Craugastor lineatus

CR

4

7

4

15

H

Craugastor loki

LC

4

12

M

Craugastor longirostris

LC

3

7

4

14

H

Craugastor matudai

vu

4

7

4

15

H

Craugastor megacephalus*

LC

5

7

4

16

H

Craugastor melanostictus*

LC

5

7

4

16

H

Craugastor merendonensis *

CR

6

8

4

18

H

Craugastor milesi*

CR

5

7

4

16

H

Craugastor mimus*

LC

5

7

4

16

H

Craugastor monnichorum*

DD

5

7

4

16

H

Craugastor myllomyllon*

DD

6

8

4

18

H

Craugastor nefrens*

DD

6

8

4

18

H

Craugastor noblei*

LC

5

7

4

16

H

Craugastor obesus*

EN

5

8

4

17

H

Craugastor olanchano*

CR

6

8

4

18

H

Craugastor omoaensis*

CR

6

8

4

18

H

Craugastor opimus

LC

4

7

4

15

H

Craugastor palenque

DD

4

7

4

15

H

Craugastor pechorum*

EN

5

7

4

16

H

Craugastor persimilis*

VU

5

7

4

16

H

Craugastor phasma*

DD

6

8

4

18

H

Craugastor podiciferus*

NT

5

6

4

15

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Craugastor polyptychus*

LC

5

8

4

17

H

Craugastor psephosypharus *

vu

5

7

4

16

H

Craugastor punctariolus*

EN

5

7

4

16

H

Craugastor pygmaeus

VU

2

3

4

9

L

Craugastor raniformis

LC

4

7

4

15

H

Craugastor ranoides*

CR

5

6

4

15

H

Craugastor rayo*

DD

5

7

4

16

H

Craugastor rhyacobatrachus*

EN

5

7

4

16

H

Craugastor rivulus*

VU

5

8

4

17

H

Craugastor rostralis*

NT

5

7

4

16

H

Craugastor rugosus*

LC

5

7

4

16

H

Craugastor rupinius

LC

4

5

4

13

H

Craugastor sabrinus*

EN

5

7

4

16

H

Craugastor saltuarius*

CR

6

8

4

18

H

Craugastor sandersoni*

EN

5

7

4

16

H

Craugastor stadelmani*

CR

5

7

4

16

H

Craugastor stejnegerianus*

LC

5

4

14

H

Craugastor stuarti

EN

4

7

4

15

H

Craugastor tabasarae*

CR

5

8

4

17

H

Craugastor talamancae*

LC

5

8

4

17

H

Craugastor taurus *

CR

5

8

4

17

H

Craugastor trachydermus*

CR

6

8

4

18

H

Craugastor underwoodi*

LC

5

7

4

16

H

Craugastor xucanebi*

VU

5

7

4

16

H

Pristimantis achatinus

LC

3

7

4

14

H

Pristimantis adnus*

NE

6

8

4

18

H

Pristimantis altae*

NT

5

7

4

16

H

Pristimantis caryophyllaceus *

NT

5

6

4

15

H

Pristimantis cerasinus*

LC

5

7

4

16

H

Pristimantis cruentus

LC

4

6

4

14

H

Pristimantis gaigeae

LC

4

8

4

16

H

Pristimantis morn

LC

4

8

4

16

H

Pristimantis museosus*

EN

5

8

4

17

H

Pristimantis pardalis *

NT

5

8

4

17

H

Pristimantis pirrensis*

DD

6

8

4

18

H

Pristimantis ridens

LC

2

6

4

12

M

Pristimantis taeniatus

LC

4

8

4

16

H

Strabomantis bufoniformis

LC

4

8

4

16

H

Strabomantis laticorpus*

DD

5

8

4

17

H

Family Dendrobatidae (19 species)

Ameerega maculata*

DD

6

8

4?

18

H

August 2015 | Volume 9 | Number 2 | el 00

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Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Andinobates claudiae*

DD

6

8

4

18

H

Andinobcites fulguritus

LC

4

7

4

15

H

Andinobates geminisae*

NE

6

8

4

18

H

Andinobates minutus

LC

4

7

4

15

H

Colostetlms latinasus*

DD

5

6

4

15

H

Colostethus panamensis

LC

4

7

4

15

H

Colostetlms pratti

LC

4

7

4

15

H

Dendrobates auratus

LC

4

7

4

15

H

Hyloxalus chocoensis

DD

4

8

4

16

H

Oophaga arborea*

EN

5

7

4

16

H

Oophaga granulifera*

VU

5

8

4

17

H

Oophaga pumilio *

LC

5

7

4

16

H

Oophaga speciosa *

EN

5

7

4

16

H

Oophaga vicentei*

DD

5

7

4

16

H

Phyllobates lugubris*

LC

5

8

4

17

H

Phyllobates vittatus *

EN

5

8

4

17

H

Silver stoneia flotator*

LC

5

7

4

16

H

Silverstoneia nubicola

Family Eleutherodactylidae (11 species)

NT

4

6

4

14

H

Diasporus citrinobapheus*

NE

5

8

4

17

H

Diasporus diastema*

LC

5

6

4

15

H

Diasporus hylaeformis*

LC

5

8

4

17

H

Diasporus igneus*

NE

6

8

4

18

H

Diasporus quidditus

LC

4

8

4

16

H

Diasporus tigrillo*

DD

6

8

4

18

H

Diasporus ventrimaculatus*

VU

6

8

4

18

H

Diasporus vocator

LC

4

7

4

15

H

Eleutherodactylus leprus

VU

2

6

4

12

M

Eleutherodactylus pipilans

LC

2

5

4

11

M

Eleutherodactylus rubrimaculatus

Family Hemiphractidae (3 species)

VU

4

7

4

15

H

Gastrotheca cornuta

EN

4

7

5

16

H

Gastrotheca nicefori

LC

3

7

5

15

H

Hemiphractus fasciatus

Family Hylidae (98 species)

NT

4

7

5

16

H

Agalychnis annae*

EN

5

7

3

15

H

Agalychnis callidryas

LC

3

5

3

11

M

Agalychnis lemur

CR

2

7

3

12

M

Agalychnis litodryas

VU

4

8

3

15

H

Agalychnis moreletii

CR

1

3

7

L

Agalychnis saltator *

LC

5

6

3

14

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Agalychnis spurrelli

LC

4

7

3

14

H

Anotheca spinosa

LC

3

6

15

H

Bromeliohyla bromeliacia

EN

4

7

6

17

H

Cruziohyla calcarifer

LC

4

8

3

15

H

Dendropsophus ebraccatus

LC

3

6

3

12

M

Dendropsophus microcephalus

LC

3

1

7

L

Dendropsophus phlebodes

LC

3

7

1

11

M

Dend ropsophus robe rtme rtensi

LC

4

1

9

L

Dendropsophus subocularis

LC

4

8

1

13

M

Duellmanohyla lythrodes *

EN

5

8

1

14

H

Duellmanohyla rufioculis *

LC

5

8

1

14

H

Duellmanohyla salvavida*

CR

5

7

1

13

M

Duellmanohyla schmidtorum

VU

4

3

1

8

L

Duellmanohyla soralia*

CR

5

6

1

12

M

Duellmanohyla uranochroa *

EN

5

6

1

12

M

Ecnomiohyla bailarina*

NE

6

8

6?

20

H

Ecnomiohyla fimbrimembra

CR

6

7

6

19

H

Ecnomiohyla miliaria*

VU

5

7

6

18

H

Ecnomiohyla minera *

EN

5

7

6

18

H

Ecnomiohyla rabborum *

CR

6

8

6

20

H

Ecnomiohyla salvaje*

CR

5

8

6

19

H

Ecnomiohyla sukia*

NE

5

7

6

18

H

Ecnomiohyla thysanota*

DD

6

8

6?

20

H

Ecnomiohyla veraguensis*

NE

6

8

6?

20

H

Exerodonta catracha*

EN

5

8

1

14

H

Exerodonta perkinsi*

CR

6

8

1

15

H

Hyla bocourti*

CR

5

8

1

14

H

Hyla walkeri

VU

4

6

1

11

M

Hyloscirtus colymba

CR

4

8

1

13

M

Hyloscirtus palmeri

LC

4

8

1

13

M

Hypsiboas boans

LC

3

8

1

12

M

Hypsiboas crepitans

LC

3

8

1

12

M

Hypsiboas pugnax

LC

4

8

1

13

M

Hypsiboas rosenbergi

LC

4

8

1

13

M

Hypsiboas rufitelus*

LC

5

8

1

14

H

Isthmohyla angustilineata *

CR

5

7

1

13

M

Isthmohyla calypsa*

CR

5

8

3

16

H

Isthmohyla debilis*

CR

5

8

1

14

H

Isthmohyla graceae*

CR

5

7

1

13

M

Isthmohyla infucata*

DD

5

8

1

14

H

Isthmohyla insolita*

CR

6

8

3

17

H

August 2015 | Volume 9 | Number 2 | el 00

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Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Isthmohyla lancasteri*

LC

5

8

1

14

H

Isthmohyla melacaena*

NT

6

8

6

20

H

Isthmohyla picadoi *

NT

5

8

6

19

H

Isthmohyla pictipes*

EN

5

8

1

14

H

Isthmohyla pseudopuma *

LC

5

7

1

13

M

Isthmohyla rivularis*

CR

5

7

1

13

M

Isthmohyla tica*

CR

5

7

1

13

M

Isthmohyla xanthosticta *

DD

6

8

1

15

H

Isthmohyla zeteki*

NT

5

7

6

18

H

Phyllomedusa venusta

LC

4

8

1

13

M

Plectrohyla acanthodes

CR

4

7

1

12

M

Plectrohyla avia

CR

4

8

1

13

M

Plectrohyla chrysopleura*

CR

5

7

1

13

M

Plectrohyla dasypus*

CR

6

7

1

14

H

Plectrohyla exquisita *

CR

6

8

1

15

H

Plectrohyla glandulosa *

EN

5

6

1

12

M

Plectrohyla guatemalensis

CR

4

1

9

L

Plectrohyla hartwegi

CR

4

5

1

10

M

Plectrohyla ixil

CR

4

7

1

12

M

Plectrohyla matudai

VU

4

6

1

11

M

Plectrohyla pokomchi*

CR

5

7

1

13

M

Plectrohyla psiloderma*

EN

5

8

1

14

H

Plectrohyla quecchi*

CR

5

7

1

13

M

Plectrohyla sagorum

EN

4

5

1

10

M

Plectrohyla tecunumani*

CR

5

8

1

14

H

Plectrohyla teuchestes*

CR

6

8

1

15

H

Ptychohyla dendrophasma*

CR

6

8

6?

20

H

Ptychohyla euthysanota

NT

4

3

1

8

L

Ptychohyla hypomykter*

CR

5

4

1

10

M

Ptychohyla legleri*

EN

5

8

1

14

H

Ptychohyla macrotympanum

CR

4

6

1

11

M

Ptychohyla panchoi*

EN

5

7

1

13

M

Ptychohyla salvadorensis*

EN

5

6

1

12

M

Ptychohyla sanctaecrucis*

CR

6

7

1

14

H

Ptychohyla spinipollex *

EN

5

6

1

12

M

Scinax altae*

LC

5

8

1

14

H

Scinax boulengeri

LC

4

6

1

11

M

Scinax elaeochroa*

LC

5

7

1

13

M

Scinax rostrata

LC

3

7

1

11

M

Scinax rubra

LC

3

7

1

11

M

Scinax staufferi

LC

2

1

4

L

August 2015 | Volume 9 | Number 2 | el 00

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73

Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Smilisca baudinii

LC

1

3

L

Smilisca cyanosticta

NT

4

7

1

12

M

Smilisca phaeota

LC

4

6

1

11

M

Smilisca puma *

LC

5

8

1

14

H

Smilisca sila

LC

4

5

1

10

M

Smilisca sordida

LC

2

5

1

8

L

Tlalocohyla loquax

LC

3

1

7

L

Tlalocohyla picta

LC

2

5

1

8

L

Trachycephalus typhonius

LC

1

2

1

4

L

Triprion petasatus

LC

4

5

1

10

M

Family Leptodactylidae (9 species)

Engystomops pustulosus

LC

3

2

7

L

Leptodactylus fragilis

LC

1

2

5

L

Leptodactylus fuscus

LC

3

7

2

12

M

Leptodactylus insularum

LC

3

7

2

12

M

Leptodactylus melanonotus

LC

1

3

2

6

L

Leptodactylus poecilochilus

LC

4

6

2

12

M

Leptodactylus savagei

LC

2

5

2

9

L

Leptodactylus silvanimbus*

CR

5

7

2

14

H

Pleurodema brachyops

LC

3

8

2

13

M

Family Microhylidae (9 species)

Ctenophryne aterrima

LC

4

7

1

12

M

Elachistocleis ovalis

LC

3

7

1

11

M

Elachistocleis panamensis

LC

4

7

1

12

M

Elachistocleis pearsei

LC

3

8

1

12

M

Gastrophryne elegans

LC

2

5

1

8

L

Hypopachus barberi

vu

4

5

1

10

M

Hypopachus pictiventris*

LC

5

8

1

14

H

Hypopachus ustus

LC

3

4

1

8

L

Hypopachus variolosus

LC

2

1

4

L

Family Pipidae (1 species)

Pip a myersi *

EN

4

8

5

17

H

Family Ranidae (11 species)

Lithobates brownorum

NE

4

3

1

8

L

Lithobates forreri

LC

1

3

L

Lithobates juliani*

NT

5

6

1

12

M

Lithobates macroglossa

VU

4

7

1

12

M

Lithobates maculatus

LC

3

1

5

L

Lithobates miadis*

VU

6

8

1

15

H

Lithobates pipiens complex

LC

4

1

9

L

Lithobates taylori*

LC

5

6

1

12

M

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

74

Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Lithobates vaillanti

LC

3

5

1

9

L

Lithobates vibicarius*

vu

5

8

1

14

H

Lithobates warszewitschi i *

Family Rhinophrynidae (1 species)

LC

5

4

1

10

M

Rhinophrynus dorsalis

Order Caudata (159 species)

Family Plethodontidae (159 species)

LC

2

5

1

8

L

Bolitoglossa alvaradoi*

EN

5

7

4

16

H

Bolitoglossa anthracina*

DD

6

8

4

18

H

Bolitoglossa aureogularis*

NE

6

8

4

18

H

Bolitoglossa biseriata

LC

1

8

4

13

M

Bolitoglossa bramei*

DD

5

8

4

17

H

Bolitoglossa carri*

CR

6

8

4

18

H

Bolitoglossa cataguana*

NE

6

8

4

18

H

Bolitoglossa celaque*

EN

5

8

4

17

H

Bolitoglossa centenorum *

NE

6

8

4

18

H

Bolitoglossa cerroensis*

LC

5

7

4

16

H

Bolitoglossa chucantiensis

NE

6

8

4

18

H

Bolitoglossa colonnea*

LC

5

7

4

16

H

Bolitoglossa compacta*

EN

5

8

4

17

H

Bolitoglossa conanti*

EN

5

7

4

16

H

Bolitoglossa copia*

DD

6

8

4

18

H

Bolitoglossa cuchumatana*

NT

5

4

14

H

Bolitoglossa cuna*

DD

5

8

4

17

H

Bolitoglossa dairy orurn *

NE

5

8

4

17

H

Bolitoglossa decora *

CR

6

8

4

18

H

Bolitoglossa diaphora*

CR

6

8

4

18

H

Bolitoglossa diminuta*

VU

6

8

4

18

H

Bolitoglossa dofleim*

NT

5

6

4

15

H

Bolitoglossa dunni*

EN

5

7

4

16

H

Bolitoglossa engelhardti

EN

4

7

4

15

H

Bolitoglossa epimela*

DD

5

8

4

17

H

Bolitoglossa eremia*

NE

6

8

4

18

H

Bolitoglossa flavimembris

EN

4

7

4

15

H

Bolitoglossa flaviventris

EN

4

5

4

13

M

Bolitoglossa franklini

EN

4

6

4

14

H

Bolitoglossa gomezi*

DD

5

7

4

16

H

Bolitoglossa gracilis *

VU

6

8

4

18

H

Bolitoglossa hartwegi

NT

4

12

M

Bolitoglossa heiroreias*

EN

5

8

4

17

H

Bolitoglossa helmrichi *

NT

5

7

4

16

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Bolitoglossa huehuetenanguensis *

NE

6

8

4

18

H

Bolitoglossa indio*

DD

5

8

4

17

H

Bolitoglossa insularis*

VU

6

8

4

18

H

Bolitoglossa jacksoni*

DD

6

8

4

18

H

Bolitoglossa jugivagans*

NE

6

8

4

18

H

Bolitoglossa kamuk*

NE

6

8

4

18

H

Bolitoglossa kaqchikelorum*

NE

5

8

4

17

H

Bolitoglossa la *

NE

5

8

4

17

H

Bolitoglossa lignicolor *

VU

5

7

4

16

H

Bolitoglossa lincolni

NT

4

5

4

13

M

Bolitoglossa longissima*

CR

6

8

4

18

H

Bolitoglossa magnifica*

EN

5

7

4

16

H

Bolitoglossa marmorea*

EN

5

8

4

17

H

Bolitoglossa medemi

VU

4

7

4

15

H

Bolitoglossa me liana *

EN

5

7

4

16

H

Bolitoglossa mexicana

LC

1

3

4

8

L

Bolitoglossa minutula*

EN

5

8

4

17

H

Bolitoglossa mombachoensis *

VU

5

8

4

17

H

Bolitoglossa morio*

LC

5

4

13

M

Bolitoglossa mulleri

VU

2

7

4

13

M

Bolitoglossa nigrescens*

EN

5

7

4

16

H

Bolitoglossa ninadormida*

NE

6

8

4

18

H

Bolitoglossa nussbaumi*

NE

6

8

4

18

H

Bolitoglossa nympha*

NE

5

7

4

16

H

Bolitoglossa obscura*

VU

6

8

4

18

H

Bolitoglossa occidentalis

LC

4

3

4

11

M

Bolitoglossa odonnelli*

EN

5

7

4

16

H

Bolitoglossa omniumsanctorum *

NE

5

7

4

16

H

Bolitoglossa oresbia*

CR

5

8

4

17

H

Bolitoglossa pacaya*

NE

5

8

4

17

H

Bolitoglossa pesrubra*

VU

5

6

4

15

H

Bolitoglossa phalarosoma

DD

4

8

4

16

H

Bolitoglossa porrasorum*

EN

5

7

4

16

H

Bolitoglossa psephena*

NE

6

8

4

18

H

Bolitoglossa pygmaea*

NE

5

8

4

17

H

Bolitoglossa robinsoni*

NE

5

7

4

16

H

Bolitoglossa robusta *

LC

5

7

4

16

H

Bolitoglossa rostrata

VU

4

6

4

14

H

Bolitoglossa rufescens

LC

1

4

9

L

Bolitoglossa salvinii*

EN

5

7

4

16

H

Bolitoglossa schizodactyla *

LC

5

6

4

15

H

August 2015 | Volume 9 | Number 2 | el 00

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Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Bolitoglossa sombra *

vu

5

7

4

16

H

Bolitoglossa sooyorum *

EN

5

7

4

16

H

Bolitoglossa splendida *

NE

6

8

4

18

H

Bolitoglossa striatula*

LC

5

7

4

16

H

Bolitoglossa stuarti

DD

4

7

4

15

H

Bolitoglossa subpalmata*

EN

5

6

4

15

H

Bolitoglossa suchitanensis *

DD

6

8

4

18

H

Bolitoglossa synoria*

CR

5

8

4

17

H

Bolitoglossa taylori*

DD

5

8

4

17

H

Bolitoglossa tenebrosa*

NE

5

8

4

17

H

Bolitoglossa tica *

EN

5

8

4

17

H

Bolitoglossa tzultacaj*

NE

6

8

4

18

H

Bolitoglossa xibalba*

NE

5

8

4

17

H

Bolitoglossa yucatana

LC

4

7

4

15

H

Bolitoglossa zacapensis*

NE

6

8

4

18

H

Bradytriton silus*

CR

6

8

4

18

H

Cryptotriton monzoni *

CR

6

8

4

18

H

Cryptotriton nasalis*

EN

6

8

4

18

H

Cryptotriton necopinus

NE

6

8

4

18

H

Cryptotriton sierraminensis *

DD

5

8

4

17

H

Cryptotriton veraepacis*

CR

5

8

4

17

H

Dendrotriton bromeliacius*

CR

5

8

4

17

H

Dendrotriton chujorum*

CR

6

8

4

18

H

Dendrotriton cuchumatanus *

CR

6

8

4

18

H

Dendrotriton kekchiorum *

EN

6

8

4

18

H

Dendrotriton rabbi *

CR

5

8

4

17

H

Dendrotriton sanctibarbarus*

VU

6

8

4

18

H

Nototriton abscondens*

LC

5

7

4

16

H

Nototriton barbouri*

EN

5

7

4

16

H

Nototriton brodiei*

CR

5

8

4

17

H

Nototriton gamezi *

VU

6

8

4

18

H

Nototriton guanacaste*

VU

5

8

4

17

H

Nototriton lignicola*

CR

6

8

4

18

H

Nototriton limnospectator*

EN

5

8

4

17

H

Nototriton major *

CR

6

8

4

18

H

Nototriton matama*

NE

6

8

4

18

H

Nototriton mime *

NE

6

8

4

18

H

Nototriton picadoi*

NT

5

7

4

16

H

Nototriton picucha *

NE

6

8

4

18

H

Nototriton richardi*

NT

5

7

4

16

H

Nototriton saslaya*

VU

6

8

4

18

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

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Conservation reassessment of Central American herpetofauna

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

Species

IUCN

rating

Environmental Vulnerability Score

c\/c

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

t Vo

Category

Nototriton stuarti*

DD

6

8

4

18

H

Nototriton tapanti*

VU

6

8

4

18

H

Nototriton tomcimorum*

NE

6

8

4

18

H

Nyctanolis pernix

EN

4

7

4

15

H

Oedipina alfaroi*

VU

5

7

4

16

H

Oedipina alleni*

LC

5

7

4

16

H

Oedipina altura*

CR

6

8

4

18

H

Oedipina carablanca*

EN

6

8

4

18

H

Oedipina chortiorum*

NE

6

8

4

18

H

Oedipina collaris*

DD

5

8

4

17

H

Oedipina complex

LC

1

6

4

11

M

Oedipina cyclocauda*

LC

5

6

4

15

H

Oedipina elongata

LC

2

7

4

13

M

Oedipina fortunensis *

NE

6

8

4

18

H

Oedipina gephyra*

EN

5

8

4

17

H

Oedipina gracilis *

EN

5

7

4

16

H

Oedipina grandis*

EN

5

8

4

17

H

Oedipina ignea*

DD

5

6

4

15

H

Oedipina kasios*

NE

5

7

4

16

H

Oedipina koehleri*

NE

5

7

4

16

H

Oedipina leptopoda*

NE

5

8

4

17

H

Oedipina maritima*

CR

6

8

4

18

H

Oedipina motaguae*

NE

6

8

4

18

H

Oedipina nica *

NE

5

8

4

17

H

Oedipina nimaso *

NE

6

8

4

18

H

Oedipina pacificensis*

LC

5

7

4

16

H

Oedipina parvipes

LC

4

7

4

15

H

Oedipina paucidentata*

CR

6

8

4

18

H

Oedipina petiola*

NE

6

8

4

18

H

Oedipina poelzi*

EN

5

7

4

16

H

Oedipina pseudouniformis *

EN

5

7

4

16

H

Oedipina quadra*

NE

5

8

4

17

H

Oedipina savagei *

DD

6

8

4

18

H

Oedipina stenopodia*

EN

5

8

4

17

H

Oedipina stuarti *

DD

5

6

4

15

H

Oedipina taylori*

LC

5

4

14

H

Oedipina tomasi*

CR

6

8

4

18

H

Oedipina tzutujilorum*

NE

6

8

4

18

H

Oedipina uniformis*

NT

5

6

4

15

H

Pseudoeurycea brunnata

CR

4

7

4

15

H

Pseudoeurycea exspectata*

CR

6

8

4

18

H

Pseudoeurycea goebeli

CR

4

7

4

15

H

Pseudoeurycea rex

CR

4

12

M

Amphib. Reptile Conserv.

78

August 2015 | Volume 9 |

Number 2 | el 00

Johnson et al.

Appendix 1 (continued). Comparison of the IUCN Ratings from the Red List website (updated to 10 August 2014) and Environ-

mental Vulnerability Scores for 493 Central American amphibians. See text for explanations of the IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America. ? = reproductive mode

estimated based on phylogenetic relationships.

IUCN

rating

Environmental Vulnerability Score

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Reproductive

Mode

Total

Score

Order Gymnophiona (15 species)

Family Caeciliiidae (7 species)

Caecilia isthmica

DD

4

8

4?

16

H

Caecilia leucocephala

LC

3

8

4?

15

H

Caecilia nigricans

LC

3

8

4?

15

H

Caecilia volcani*

DD

5

8

4?

17

H

Oscaecilia elongata*

DD

6

8

5

19

H

Oscaecilia ochrocephala

LC

4

7

5

16

H

Oscaecilia osae*

DD

6

8

5?

19

H

Family Dermophiidae (8 species)

Dermophis costaricensis*

DD

5

8

5

18

H

Dermophis glandulosus

DD

2

6

5?

13

M

Dermophis gracilior *

DD

5

8

5

18

H

Dermophis mexicanus

VU

1

5

7

L

Dermophis occidentalism

DD

5

7

5

17

H

Dermophis parviceps

LC

2

6

5?

13

M

Gymnopis multiplicata*

LC

5

4

5

14

H

Gymnopis syntrema

DD

4

7

5

16

H

Appendix 2. Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental Vulnerability

Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rating systems.

EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Environmental Vulnerability Scores

Species

IUOIN

Ratings

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

tVd

Category

Order Crocodylia (3 species)

Family Alligatoridae (1 species)

Caiman crocodilus

LC

3

7

6

16

H

Family Crocodylidae (2 species)

Crocodylus acutus

VU

3

5

6

14

H

Crocodylus moreletii

LC

2

5

6

13

M

Order Squamata (532 species)

Family Amphisbaenidae (2 species)

Amphisbaena fuliginosa

LC

3

7

1

11

M

Amphisbaena spurrelli

NE

3

8

1

12

M

Family Anguidae (28 species)

Abronia anzuetoi*

VU

6

8

4

18

H

Abronia aurita*

EN

5

7

4

16

H

Abronia campbelli*

CR

6

8

4

18

H

Abronia fimbriata*

NE

5

7

4

16

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E Vo

Category

Abronia frosti*

CR

6

8

4

18

H

Abronia gaiophantasma*

EN

5

7

4

16

H

Abronia lythrochila

LC

4

7

4

15

H

Abronia matudai

EN

4

7

4

15

H

Abronia meledona*

EN

6

8

4

18

H

Abronia montecristoi*

EN

5

8

4

17

H

Abronia ochoterenai

DD

4

8

4

16

H

Abronia salvadorensis *

EN

5

8

4

17

H

Abronia vasconcelosii *

VU

5

7

4

16

H

Celestus adercus *

DD

6

8

3

17

H

Celestus atitlanensis *

NE

5

7

3

15

H

Celestus bivittatus *

EN

5

7

3

15

H

Celestus cyanochloris *

LC

5

6

3

14

H

Celestus hylaius*

NT

5

8

3

16

H

Celestus montanus*

EN

5

7

3

15

H

Celestus orobius*

DD

5

8

3

16

H

Celestus rozellae

LC

4

6

3

13

M

Celestus scansorius*

NT

5

7

3

15

H

Coloptychon rhomb if er*

DD

5

8

3

16

H

Diploglossus bilobatus *

LC

5

7

4

16

H

Diploglossus monotropis

NE

4

7

4

15

H

Diploglossus montisilvestris *

DD

6

8

4

18

H

Mesaspis monticola*

LC

5

6

3

14

H

Mesaspis moreletii

Family Corytophanidae (9 species)

LC

2

3

8

L

Basiliscus basiliscus

NE

4

3

11

M

Basiliscus galeritus

NE

3

7

3

13

M

Basiliscus plumifrons

LC

5

7

3

15

H

Basiliscus vittatus

NE

1

3

7

L

Corytophanes cristatus

NE

2

5

3

10

M

Corytophanes hernandesii

LC

4

6

3

13

M

Corytophanes percarinatus

LC

4

3

11

M

Laemanctus longipes

LC

1

5

3

9

L

Laemanctus serratus

Family Dactyloidae (95 species)

LC

3

9

L

Anolis allisoni

NE

3

7

3

13

M

Dactyloa casildae*

NE

5

8

3

16

H

Dactyloa chloris

NE

3

8

3

14

H

Dactyloa chocorum

NE

4

8

3

15

H

Dactyloa frenata

NE

4

7

3

14

H

Dactyloa ginaelisae*

NE

5

4

3

12

M

August 2015 | Volume 9 | Number 2 | el 00

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80

Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Species

II lOKI

Environmental Vulnerability Scores

EVS

Category

lUUIN

Ratings

1

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

Dactyloa ibanezi*

NE

5

7

3

15

H

Dactyloa insignis*

NE

5

6

3

14

H

Dactyloa kunayalae*

NE

5

7

3

15

H

Dactyloa latifrons

NE

3

7

3

13

M

Dactyloa microtus *

NE

5

7

3

15

H

Norops alocomyos

NE

5

8

3

16

H

Norops altae*

LC

5

7

3

15

H

Norops amplisquamosus *

EN

6

8

3

17

H

Norops apletophallus*

NE

5

7

3

15

H

Norops aquaticus*

NE

5

7

3

15

H

Norops auratus

NE

3

7

3

13

M

Norops beckeri

NE

3

6

3

12

M

Norops benedikti*

NE

5

8

3

16

H

Norops bicaorum*

NE

6

8

3

17

H

Norops biporcatus

NE

2

4

3

9

L

Norops campbelli*

NE

6

8

3

17

H

Norops capito

NE

2

6

3

11

M

Norops carpenteri*

LC

5

8

3

16

H

Norops charlesmyersi*

NE

5

8

3

16

H

Norops cobanensis*

NE

5

3

13

M

Norops crassulus

NE

2

4

3

9

L

Norops cristifer

DD

4

6

3

13

M

Norops cryptolimifrons *

NE

5

8

3

16

H

Norops cup reus*

NE

5

3

13

M

Norops cusuco*

EN

6

8

3

17

H

Norops datzorum*

NE

5

7

3

15

H

Norops dollfusianus

NE

4

6

3

13

M

Norops fortunensis *

DD

6

8

3

17

H

Norops fungosus *

NE

5

7

3

15

H

Norops fuscoauratus

NE

3

7

3

13

M

Norops gaigei

NE

4

7

3

14

H

Norops gruuo*

NE

6

8

3

17

H

Norops haguei*

NE

6

8

3

17

H

Norops heteropholidotus*

NE

5

8

3

16

H

Norops humilis*

NE

5

6

3

14

H

Norops intermedius*

NE

5

6

3

14

H

Norops johnmeyeri*

NE

5

8

3

16

H

Norops kemptoni*

NE

5

7

3

15

H

Norops kreutzi *

NE

6

8

3

17

H

Norops laeviventris

NE

2

3

8

L

Norops leditzigorum

NE

5

7

3

15

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Species

IUCN -

Ratings

1

Environmental Vulnerability Scores

Geographic Ecological Degree of Total

Distribution Distribution Human Score

Persecution

EVS

Category

Norops lemurinus

NE

2

3

7

L

Norops limifrons *

NE

5

7

3

15

H

Norops lionotus *

LC

5

6

3

14

H

Norops loveridgei *

EN

5

6

3

14

H

Norops macrophallus *

NE

5

7

3

15

H

Norops magnaphallus *

NE

6

8

3

17

H

Norops marsupialis *

NE

5

8

3

16

H

Norops matudai

NE

4

6

3

13

M

Norops monteverde*

NE

6

8

3

17

H

Norops morazani*

NE

6

8

3

17

H

Norops muralla*

VU

6

8

3

17

H

Norops ocelloscapularis*

NE

5

7

3

15

H

Norops osa*

NE

5

8

3

16

H

Norops pachypus*

LC

5

7

3

15

H

Norops pentaprion*

NE

5

4

3

12

M

Norops petersii

NE

2

4

3

9

L

Norops pijolensis*

NE

6

7

3

16

H

Norops poecilopus

NE

4

7

3

14

H

Norops polylepis*

NE

5

7

3

15

H

Norops pseudokemptoni*

NE

6

8

3

17

H

Norops pseudopachypus*

NE

6

8

3

17

H

Norops purpurgularis *

NE

5

8

3

16

H

Norops quaggulus*

NE

5

7

3

15

H

Norops roatanensis*

NE

6

8

3

17

H

Norops rodriguezii

NE

4

3

10

M

Norops rubribarbaris*

NE

6

8

3

17

H

Norops sagrei

NE

3

7

3

13

M

Norops salvini*

NE

5

7

3

15

H

Norops sericeus

NE

2

3

8

L

Norops serranoi

NE

4

5

3

12

M

Norops sminthus*

DD

5

7

3

15

H

Norops tenorioensis*

NE

6

8

3

17

H

Norops townsendi*

NE

6

8

3

17

H

Norops triumphalis *

NE

6

8

3

17

H

Norops tropidogaster

NE

3

7

3

13

M

Norops tropidolepis*

NE

5

7

3

15

H

Norops tropidonotus

NE

4

2

3

9

L

Norops uniformis

NE

4

6

3

13

M

Norops unilobatus

NE

1

3

7

L

Norops utilensis*

NE

6

8

3

17

H

Norops villai*

NE

6

8

3

17

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

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Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E VO

Category

Norops vittigerus

NE

4

7

3

14

H

Norops wcimpuensis*

NE

6

8

3

17

H

Norops wellbornae*

NE

5

7

3

15

H

Norops wennuthi*

NE

5

8

3

16

H

Norops woodi*

NE

5

6

3

14

H

Norops yoroensis*

NE

5

7

3

15

H

Norops zeus *

NE

5

7

3

15

H

Family Eublepharidae (2 species)

Coleonyx elegans

LC

3

4

10

M

Coleonyx mitratus

LC

5

4

14

H

Family Gymnophthalmidae (14 species)

Anadia ocellcita*

NE

5

8

3

16

H

Anadia vittata

NE

4

7

3

14

H

Bachia blairi*

NT

5

8

2

15

H

Bachia pallidiceps

NE

4

8

2

14

H

Cercosaura vertebralis

NE

3

7

3

13

M

Echinosaura palmeri

NE

3

7

2

12

M

Echinosaura pcinamensis*

LC

5

7

2

14

H

Gymnophthalmus speciosus

NE

3

9

L

Leposoma rugiceps

LC

4

8

3

15

H

Leposoma southi

NE

4

7

3

14

H

Potamites apodemus*

LC

5

7

3

15

H

Ptychoglossus festae

NE

4

7

3

14

H

Ptychoglossus myersi *

LC

5

8

3

16

H

Ptychoglossus plicatus

NE

2

6

3

11

M

Family Helodermatidae (2 species)

Heloderma alvarezi

NE

3

6

5

14

H

Heloderma charlesbogerti*

NE

5

8

5

18

H

Family Hoplocercidae (2 species)

Enyalioides heterolepis

NE

3

7

3

13

M

Morunasaurus groi

NE

4

8

3

15

H

Family Iguanidae (11 species)

Ctenosaura acanthura

NE

3

4

6

13

M

Ctenosaura alfredschmidti

NT

4

8

3

15

H

Ctenosaura bakeri*

CR

5

8

6

19

H

Ctenosaura flavidorsalis*

EN

5

7

6

18

H

Ctenosaura melanosterna*

EN

5

7

6

18

H

Ctenosaura oedirhina*

EN

5

8

6

19

H

Ctenosaura palearis*

EN

5

8

6

19

H

Ctenosaura praeocularis*

DD

5

7

6

18

H

Ctenosaura quinquecarinata*

NE

5

8

6

19

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

83

Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E Vo

Category

Ctenosaura similis

LC

1

4

6

11

M

Iguana iguana

Family Mabuyidae (5 species)

NE

1

3

6

10

M

Marisora alliacea*

LC

5

7

3

15

H

Marisora brachypoda

LC

1

2

3

6

L

Marisora magnacornae*

DD

6

8

3

17

H

Marisora roatanae*

CR

5

8

3

16

H

Marisora unimarginata*

Family Phrynosomatidae (17 species)

LC

5

7

3

15

H

Phrynosoma asio

LC

3

6

3

12

M

Sceloporus acanthinus

LC

4

7

3

14

H

Sceloporus carinatus

LC

4

5

3

12

M

Sceloporus chrysostictus

LC

4

6

3

13

M

Sceloporus internasalis

LC

4

3

11

M

Sceloporus lunaei*

LC

5

7

3

15

H

Sceloporus lundelli

LC

4

7

3

14

H

Sceloporus malachiticus*

LC

5

2

3

10

M

Sceloporus melanorhinus

LC

3

4

3

10

M

Sceloporus prezygus

LC

4

8

3

15

H

Sceloporus serrifer

LC

3

1

3

7

L

Sceloporus siniferus

LC

3

6

3

12

M

Sceloporus smaragdinus

LC

4

5

3

12

M

Sceloporus squamosus

LC

2

5

3

10

M

Sceloporus taeniocnemis

LC

4

5

3

12

M

Sceloporus teapensis

LC

4

6

3

13

M

Sceloporus variabilis

Family Phyllodactylidae (5 species)

LC

1

3

5

L

Phyllodactylus insularis*

VU

6

8

3

17

H

Plryllodactylus palmeus*

NE

5

8

3

16

H

Phyllodactylus paralepis*

NE

6

8

3

17

H

Phyllodactylus tuberculosus

LC

1

4

3

8

L

Thecadactylus rapicauda

Family Polychrotidae (1 species)

NE

1

4

3

8

L

Polychrus gutturosus

Family Scincidae (3 species)

NE

1

8

3

12

M

Mesoscincus managuae

LC

5

6

3

14

H

Mesoscincus schwartzei

LC

4

6

3

13

M

Plestiodon sumichrasti

Family Sphaerodactylidae (19 species)

LC

4

5

3

12

M

Aristelliger georgeensis

NE

3

7

3

13

M

Aristelliger praesignis

NE

3

8

3

14

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

84

Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E Vo

Category

Gonatodes albogularis

NE

1

5

3

9

L

Lepidoblepharis sanctaemartae

LC

4

7

3

14

H

Lepidoblepharis xantho stigma

LC

4

6

3

13

M

Sphaerodactylus alphas*

NE

6

8

3

17

H

Sphaerodactylus continentalis

NE

2

3

8

L

Sphaerodactylus dunni *

LC

5

7

3

15

H

Sphaerodactylus glaucus

LC

4

5

3

12

M

Sphaerodactylus graptolaemus *

LC

5

8

3

16

H

Sphaerodactylus guanaje*

NE

6

8

3

17

H

Sphaerodactylus homolepis *

LC

5

8

3

16

H

Sphaerodactylus leonardovaldesi *

NE

5

8

3

16

H

Sphaerodactylus lineolatus

NE

4

7

3

14

H

Sphaerodactylus millepunctatus *

LC

5

7

3

15

H

Sphaerodactylus notatus

LC

3

8

3

14

H

Sphaerodactylus pacificus*

LC

6

8

3

17

H

Sphaerodactylus poindexteri*

NE

6

8

3

17

H

Sphaerodactylus rosaurae*

Family Sphenomorphidae (4 species)

LC

5

8

3

16

H

Scincella assatus

LC

3

2

3

8

L

Scincella cherriei

LC

2

3

7

L

Scincella incerta

NE

5

7

3

15

H

Scincella rara*

Family Teiidae (12 species)

DD

6

8

3

17

H

Ameiva praesignis

NE

3

8

3

14

H

Aspidoscelis angusticeps

LC

4

6

3

13

M

Aspidoscelis deppii

LC

1

4

3

8

L

Aspidoscelis maslini

LC

4

8

3

15

H

Aspidoscelis motaguae

LC

4

5

3

12

M

Cnemidophorus duellmani *

NE

5

8

3

16

H

Cnemidophorus ruatanus*

NE

5

7

3

15

H

Holcosus chaitzami

DD

4

7

3

14

H

Holcosus festivus

NE

2

5

3

10

M

Holcosus leptophrys *

NE

5

8

3

16

H

Holcosus quadrilineatus*

LC

5

8

3

16

H

Holcosus undulatus

Family Xantusiidae (4 species)

LC

1

2

3

6

L

Lepidophyma flavimaculatum

LC

2

5

2

9

L

Lepidophyma mayae

NT

4

7

2

13

M

Lepidophyma reticulatum*

LC

5

6

2

13

M

Lepidophyma smithii

LC

3

4

2

9

L

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

85

Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/c

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

t Vo

Category

Family Xenosauridae (1 species)

Xenosaurus grandis

vu

3

1

3

7

L

Family Anomalepididae (3 species)

Anomalepis mexicanus

DD

2

8

1

11

M

Helminthophis frontalis *

DD

5

6

1

12

M

Liotyphlops albirostris

NE

3

5

1

9

L

Family Boidae (4 species)

Boa imperator

NE

1

6

8

L

Corallus annulatus

NE

1

8

2

11

M

Corallus ruschenbergerii

NE

3

8

2

13

M

Epicrates maurus

NE

1

5

2

8

L

Family Charinidae (2 species)

Ungaliophis continentalis

NE

2

5

2

9

L

Ungaliophis panamensis

NE

4

6

2

12

M

Family Colubridae (74 species)

Chironius exoletus

NE

3

5

4

12

M

Chironius flavopictus

DD

4

7

4

15

H

Chironius grandisquamis

NE

1

6

4

11

M

Coluber constrictor

LC

3

6

3

12

M

Dendrophidion apharocybe

NE

5

7

4

16

H

Dendrophidion crybelum*

NE

5

8

4

17

H

Dendrophidion clarkii

NE

4

6

4

14

H

Dendrophidion paucicarinatum *

LC

5

7

4

16

H

Dendrophidion percarinatum

NE

1

6

4

11

M

Dendrophidion rufiterminorum *

NE

5

7

4

16

H

Dendrophidion vinitor

LC

3

7

3

13

M

Drymarchon melanurus

LC

1

4

6

L

Drymobius chloroticus

LC

1

3

4

8

L

Drymobius margaritiferus

NE

1

4

6

L

Drymobius melanotropis*

LC

5

7

4

16

H

Drymobius rhombifer

LC

3

7

4

14

H

Ficimia publia

LC

4

3

2

9

L

Lampropeltis abnorma

NE

1

3

5

9

L

Lampropeltis micropholis

NE

4

1

5

10

M

Leptodrymus pulcherrimus*

LC

5

4

13

M

Leptophis ahaetulla

NE

3

4

10

M

Leptophis depressirostris

NE

3

7

4

14

H

Leptophis mexicanus

LC

1

4

6

L

Leptophis modestus

VU

3

7

4

14

H

Leptophis nebulosus*

LC

5

4

14

H

Leptophis riveti

NE

3

7

4

14

H

Masticophis mentovarius

NE

1

4

6

L

Mastigodryas alternatus*

LC

5

3

4

12

M

Amphib. Reptile Conserv. 86 August 2015 | Volume 9 | Number 2 | el 00

Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Environmental Vulnerability Scores

Species

Ratings

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

cvo

Category

Mastigodryas dorsalis *

LC

5

4

14

H

Mastigodryas melanolomus

LC

3

4

11

M

Mastigodryas pleei

NE

3

7

4

14

H

Oxybelis aeneus

NE

1

3

5

L

Oxybelis brevirostris

NE

1

7

4

12

M

Oxybelis fulgidus

NE

1

2

4

7

L

Oxybelis wilsoni*

EN

5

8

4

17

H

Phrynonax poecilonotus

LC

1

3

7

L

Pituophis lineaticollis

LC

4

2

4

10

M

Pseudelaphe flavirufa

LC

4

12

M

Rhinobothryum bovallii

LC

3

8

5

16

H

Scolecophis atrocinctus*

LC

5

3

5

13

M

Senticolis triaspis

LC

3

1

3

7

L

Spilotes pullatus

NE

1

4

6

L

Stenorrhina degenhardtii

NE

3

9

L

Stenorrhina freminvillii

LC

1

2

4

7

L

Symphimus mayae

LC

4

7

3

14

H

Tantilla albiceps*

DD

6

8

2

16

H

Tantilla alticola

NE

4

5

2

11

M

Tantilla armillata*

LC

5

4

2

11

M

Tantilla bairdi*

DD

6

8

2

16

H

Tantilla brevicauda*

LC

5

6

2

13

M

Tantilla cuniculator

LC

4

7

2

13

M

Tanti llahende rson i *

DD

6

8

2

16

H

Tantilla impensa

LC

2

5

2

9

L

Tantilla jani*

VU

4

8

2

14

H

Tantilla lempira*

EN

5

7

2

14

H

Tantilla melanocephala

NE

3

7

2

12

M

Tantilla moesta

LC

4

7

2

13

M

Tantilla olympia*

NE

6

8

2

16

H

Tantilla psittaca*

VU

5

8

2

15

H

Tantilla reticulata

NE

4

7

2

13

M

Tantilla rubra

LC

3

1

2

6

L

Tantilla ruficeps*

LC

5

2

12

M

Tantilla schistosa

LC

2

3

2

7

L

Tantilla supracincta

NE

4

7

5

16

H

Tantilla taeniata*

LC

5

2

12

M

Tantilla tecta*

DD

6

8

2

16

H

Tantilla tritaeniata*

CR

6

8

2

16

H

Tantilla vermiformis*

LC

5

7

2

14

H

Tantilla vulcani*

LC

5

6

2

13

M

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

87

Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E Vo

Category

Tantillita brevissima

LC

4

3

2

9

L

Tantillita canula

LC

4

6

2

12

M

Tantillita lintoni

LC

4

6

2

12

M

Trimorphodon biscutatus

NE

3

1

4

8

L

Trimorphodon quadruplex*

Family Dipsadidae (144 species)

LC

5

4

14

H

Adelphicos daryi*

EN

6

8

2

16

H

Adelphicos ibarrorum*

EN

5

8

2

15

H

Adelphicos quadrivirgatum

LC

4

2

10

M

Adelphicos sargii

LC

4

6

2

12

M

Adelphicos veraepacis*

VU

5

7

2

14

H

Amastridium sapperi

LC

4

2

10

M

Amastridium veliferum

LC

4

7

2

13

M

Atractus clarki

NE

4

8

2

14

H

Atractus darienensis*

DD

6

8

2

16

H

Atractus depressiocellus*

DD

6

7

2

15

H

Atractus ho stilitr actus *

DD

6

8

2

16

H

Atractus imperfectus*

DD

6

8

2

16

H

Chapinophis xanthocheilus *

EN

5

8

3

16

H

Clelia clelia

NE

1

5

4

10

M

Clelia equatoriana

NE

4

6

4

14

H

Clelia scytalina

LC

3

5

4

12

M

Coniophanes bipunctatus

LC

2

5

3

10

M

Coniophanes fissidens

NE

1

3

7

L

Coniophanes imperialis

LC

3

9

L

Coniophanes joanae*

DD

5

7

3

15

H

Coniophanes piceivittis

LC

1

3

7

L

Coniophanes quinquevittatus

LC

4

6

3

13

M

Coniophanes schmidti

LC

4

6

3

13

M

Conophis lineatus

LC

4

3

4

11

M

Conophis vittatus

LC

3

5

4

12

M

Crisantophis nevermanni *

LC

5

7

4

16

H

Cubophis brooksi

NE

3

8

3

14

H

Diaphorolepis wagneri

NE

3

8

3

14

H

Dipsas articulata*

LC

5

8

2

15

H

Dipsas bicolor*

LC

5

7

5

17

H

Dipsas brevifacies

LC

4

7

4

15

H

Dipsas nicholsi*

LC

5

8

2

15

H

Dipsas temporalis

NE

3

8

2

13

M

Dipsas tenuis sima*

NT

5

7

2

14

H

Dipsas viguieri*

LC

4

7

2

13

M

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

88

Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Species

IUCN -

Ratings

1

Environmental Vulnerability Scores

Geographic Ecological Degree of Total

Distribution Distribution Human Score

Persecution

EVS

Category

Enuliophis sclateri

NE

4

7

2

13

M

Enulius bifoveatus*

CR

6

8

2

16

H

Enulius flavitorques

NE

1

2

4

L

Enulius roatanensis*

EN

6

8

2

16

H

Erythrolamprus bizona

LC

3

4

5

12

M

Erythrolamprus mimus

LC

4

6

5

15

H

Geophis bellus *

DD

6

8

2

16

H

Geophis brachycephalus*

LC

5

4

2

11

M

Geophis cancellatus

LC

4

6

2

12

M

Geophis carinosus

LC

3

4

2

9

L

Geophis championi*

DD

6

8

2

16

H

Geophis damiani*

CR

6

8

2

16

H

Geophis downsi*

DD

6

8

2

16

H

Geophis dunni*

DD

6

8

2

16

H

Geophis fulvoguttatus *

EN

5

7

2

14

H

Geophis godmani*

LC

5

7

2

14

H

Geophis hoffmanni*

NE

5

2

12

M

Geophis immaculatus

LC

4

8

2

14

H

Geophis nasalis

LC

4

3

2

9

L

Geophis nephodrymus*

VU

6

8

2

16

H

Geophis rhodogaster

LC

2

7

2

11

M

Geophis ruthveni*

LC

5

7

2

14

H

Geophis talamancae*

EN

5

8

2

15

H

Geophis tectus*

LC

5

6

2

13

M

Geophis zeledoni*

LC

5

8

2

15

H

Hydromorphus concolor*

LC

5

2

12

M

Hydromorphus dunni*

DD

6

8

2

16

H

Imantodes cenchoa

NE

1

3

2

6

L

Imantodes gemmistratus

NE

1

3

2

6

L

Imantodes inornatus

LC

4

6

2

12

M

Imantodes phantasma*

DD

6

8

2

16

H

Imantodes tenuis simus

LC

4

7

2

13

M

Leptode i ra frenata

LC

4

12

M

Leptodeira maculata

LC

3

1

4

8

L

Leptodeira nigrofasciata

LC

1

3

4

8

L

Leptodeira rhombifera*

LC

5

3

4

12

M

Leptodei ra rub ri cat a *

LC

5

8

4

17

H

Leptodeira septentrionalis

NE

1

2

4

7

L

Liophis epinephelus

NE

1

4

5

10

M

Liophis lineatus

NE

3

8

4

15

H

Ninia atrata

NE

3

8

2

13

M

Ninia celata*

NT

5

8

2

15

H

Amphib. Reptile Conserv. 89 August 2015 | Volume 9 | Number 2 | el 00

Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

Species

IUCN -

Ratings

1

Environmental Vulnerability Scores

Geographic Ecological Degree of Total

Distribution Distribution Human Score

Persecution

EVS

Category

Ninia diademata

LC

1

3

2

6

L

Ninia espinali *

NT

5

7

2

14

H

Ninia mciculata*

LC

5

2

12

M

Ninia pavimentata*

LC

5

8

2

15

H

Ninia psephota*

LC

5

6

2

13

M

Ninia sebae

LC

1

2

4

L

Nothopsis rugosus

LC

1

7

2

10

L

Omoadiphas aurula*

VU

6

8

2

16

H

Omoadiphas cannula

CR

6

8

2

16

H

Omoadiphas texiguatensis *

CR

6

8

2

16

H

Oxyrhopus petolarius

NE

1

6

5

12

M

Phimophis guianensis

NE

3

8

2

13

M

Pliocercus elapoides

LC

4

1

5

10

M

Pliocercus euryzonus

LC

1

6

5

12

M

Pseudoboa neuwiedii

NE

3

6

5

14

H

Rhadinaea calligaster*

LC

5

7

2

14

H

Rhadinaea decorata

NE

1

6

2

9

L

Rhadinaea pulveriventris*

NE

5

7

2

14

H

Rhadinaea sargenti *

LC

5

7

2

14

H

Rhadinaea stadelmani*

EN

5

6

2

13

M

Rhadinaea vermiculaticeps*

NT

5

8

2

15

H

Rhadinella anachoreta*

LC

5

7

2

14

H

Rhadinella godmani

LC

2

5

2

9

L

Rhadinella hannsteini

DD

4

5

2

11

M

Rhadinella hempsteadae*

EN

5

6

2

13

M

Rhadinella kinkelini*

LC

5

6

2

13

M

Rhadinella lachrymans

LC

4

2

8

L

Rhadinella montecristi*

VU

5

7

2

14

H

Rhadinella pegosalyta *

VU

6

8

2

16

H

Rhadinella pilonaorum*

NE

5

8

2

15

H

Rhadinella posadasi

EN

4

8

2

14

H

Rhadinella rogerromani*

NT

6

8

2

16

H

Rhadinella serperaster*

LC

5

6

2

13

M

Rhadinella tolpanorum*

CR

6

8

2

16

H

Sibon annulatus *

LC

5

7

2

14

H

Sibon anthracops*

LC

5

15

H

Sibon argus*

LC

5

7

4

16

H

Sibon carri*

NE

5

7

2

14

H

Sibon dimidiatus

LC

1

5

4

10

M

Sibon lamari*

EN

6

8

2

16

H

Sibon longifrenis *

LC

5

7

2

14

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

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Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E VO

Category

Sibon manzanaresi*

NT

5

8

2

15

H

Sibon merendonensis*

CR

6

8

2

16

H

Sibon miskitus*

NT

5

8

2

15

H

Sibon nebulatus

NE

1

2

5

L

Sibon noalamina*

NE

5

8

2

15

H

Sibon perissostichon *

DD

6

8

2

16

H

Sibon sanniolus

LC

4

6

2

12

M

Siphlophis cervinus

NE

3

8

5

16

H

Siphlophis compressus

LC

3

8

5

16

H

Tretanorhinus mocquardi*

NE

5

8

2

15

H

Tretanorhinus nigroluteus

NE

2

5

2

9

L

Trimetopon bcirbouri*

DD

5

8

2

15

H

Trimetopon grcicile*

LC

5

7

2

14

H

Trimetopon pliolepis *

LC

5

2

12

M

Trimetopon simile*

EN

5

6

2

13

M

Trimetopon slevini*

NT

5

7

2

14

H

Trimetopon viquezi *

CR

5

8

2

15

H

Tropidodipsas fas data

NE

4

12

M

Tropidodipsas fischeri

LC

4

3

2

9

L

Tropidodipsas sartorii

LC

3

2

5

10

M

Urotheca decipiens

NE

2

6

2

10

M

U rotheca fulviceps

NE

3

8

2

13

M

Urotheca guentheri *

LC

5

2

12

M

Urotheca myersi *

DD

5

8

2

15

H

Urotheca pachyura*

LC

5

7

2

14

H

Xenodon rabdocephalus

NE

1

5

11

M

Family Elapidae (18 species)

Hydrophis platurus

LC

_

Micrurus alleni*

LC

5

6

5

16

H

Micrurus ancoralis

NE

3

7

5

15

H

Micrurus browni

LC

3

1

5

9

L

Micrurus clarki*

NE

5

7

5

17

H

Micrurus diastema

LC

3

1

5

9

L

Micrurus dissoleucus

LC

3

7

5

15

H

Micrurus dumerilii

NE

3

8

5

16

H

Micrurus elegans

LC

4

5

13

M

Micrurus hippocrepis *

LC

5

8

5

18

H

Micrurus latifasciatus

LC

4

5

13

M

Micrurus mipartitus

NE

3

7

5

15

H

Micrurus mosquitensis *

LC

5

7

5

17

H

Micrurus multifasciatus *

LC

5

15

H

August 2015 | Volume 9 | Number 2 | el 00

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Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E Vo

Category

Micrurus nigrocinctus

NE

2

3

5

10

M

Micrurus ruatanus*

CR

5

8

5

18

H

Micrurus steward *

LC

5

7

5

17

H

Micrurus stuarti*

Family Leptotyphlopidae (5 species)

LC

5

7

5

17

H

Epictia ater*

LC

5

4

1

10

M

Epictia goudotii

NE

3

1

5

L

Epictia magnamaculata

NE

4

7

1

12

M

Epictia phenops

NE

3

1

5

L

Trilepida macrolepis

Family Loxocemidae (1 species)

NE

3

8

1

12

M

Loxocemus bicolor

Family Natricidae (5 species)

LC

1

5

4

10

M

Storeria dekayi

LC

3

4

2

9

L

Thamnophis cyrtopsis

LC

3

1

4

8

L

Thamnophis fulvus

LC

4

5

4

13

M

Thamnophis marcianus

LC

1

5

4

10

M

Thamnophis proximus

Family Sibynophiidae (2 species)

LC

3

2

4

9

L

Scaphiodontophis annulatus

LC

1

5

11

M

Scaphiodontophis venustissimus

Family Tropidophiidae (1 species)

NE

1

7

5

13

M

Trachyboa boulengeri

Family Typhlopidae (5 species)

NE

3

5

3

11

M

Amerotyphlops costaricensis *

LC

5

1

11

M

Amerotyphlops microstomus

LC

4

7

1

12

M

Amerotyphlops stadelmani*

NE

5

6

1

12

M

Amerotyphlops tenuis

LC

4

6

1

11

M

Amerotyphlops tycherus*

Family Viperidae (32 species)

vu

5

8

1

14

H

Agkistrodon bilineatus

NT

3

5

13

M

Agkistrodon howardgloydi*

NE

5

7

5

17

H

Agkistrodon russeolus

NE

4

6

5

15

H

Atropoides indomitus *

EN

5

8

5

18

H

Atropoides mexicanus

LC

2

4

5

11

M

Atropoides occiduus

LC

4

6

5

15

H

Atropoides olmec

LC

4

6

5

15

H

Atropoides picadoi*

LC

5

6

5

16

H

Bothriechis aurifer

VU

4

6

5

15

H

Bothriechis bicolor

LC

4

5

14

H

Bothriechis guifarroi

NE

6

8

5

19

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

92

Johnson et al.

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations: L = low; M = medium; H = high. * = species endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

c\/o

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

E VO

Category

Bothriechis lateralis*

LC

5

6

5

16

H

Bothriechis marchi*

EN

5

6

5

16

H

Bothriechis nigroviridis *

NE

5

7

5

17

H

Bothriechis schlegelii

NE

2

4

5

11

M

Bothriechis supraciliaris *

NE

5

7

5

17

H

Bothriechis thalassinus *

NE

5

7

5

17

H

Bothrops asper

NE

1

4

5

10

M

Bothrops punctatus

NE

3

8

5

16

H

Cerrophidion godmani

LC

4

3

5

12

M

Cerrophidion sasai*

NE

5

6

5

16

H

Cerrophidion wilsoni*

NE

5

15

H

Crotalus simus

LC

2

5

9

L

Crotalus tzabcan

LC

4

7

5

16

H

Lachesis acrochorda

NE

3

6

5

14

H

Lachesis melanocephala*

NE

5

7

5

17

H

Lachesis stenophrys*

NE

5

7

5

17

H

Porthidium lansbergii

NE

3

7

5

15

H

Porthidium nasutum

LC

1

6

5

12

M

Porthidium ophry omegas*

LC

5

4

5

14

H

Porthidium porrasi*

LC

5

8

5

18

H

Porthidium volcanicum*

Order Testudines (24 species)

Family Cheloniidae (5 species)

DD

5

8

5

18

H

Caretta caretta

EN

—

Chelonia mydas

EN

—

Eretmochelys imbricata

CR

—

Lepidochelys kempii

CR

—

Lepidochelys olivacea

Family Chelydridae (2 species)

VU

—

Chelydra acutirostris

NE

1

4

6

11

M

Chelydra rossignonii

Family Dermatemydidae (1 species)

VU

4

7

6

17

H

Dermatemys mawii

Family Dermochelyidae (1 species)

CR

4

7

6

17

H

Dermochelys coriacea

Family Emydidae (2 species)

CR

—

Trachemys grayi

NE

4

8

6

18

H

Trachemys ornata

Family Geoemydidae (5 species)

NE

1

4

6

11

M

Rhinoclemmys annulata

NT

2

7

3

12

M

Rhinoclemmys areolata

NT

4

6

3

13

M

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

93

Conservation reassessment of Central American herpetofauna

Appendix 2 (continued). Comparison of IUCN Ratings from the Red List website (updated to 16 July 2014) and Environmental

Vulnerability Scores for 559 Central American crocodilians, squamates, and turtles. See text for explanation of IUCN and EVS rat-

ing systems. EVS category abbreviations:

L = low; M

= medium; H =

high. * = species

endemic to Central America.

IUCN

Ratings

Environmental Vulnerability Scores

EVS

Category

Species

Geographic

Distribution

Ecological

Distribution

Degree of

Human

Persecution

Total

Score

Rhinoclemmys funerea *

NT

5

8

3

16

H

Rhinoclemmys melanosterna

NE

4

8

3

15

H

Rhinoclemmys pulcherrima

NE

1

4

3

8

L

Family Kinosternidae (4 species)

Kinosternon acutum

NT

4

7

3

14

H

Kinosternon angustipons *

VU

5

8

3

16

H

Kinosternon leucostomum

NE

1

4

3

8

L

Kinosternon scorpioides

NE

1

4

3

8

L

Family Staurotypidae (3 species)

Claudius august at us

NT

4

7

3

14

H

Staurotypus salvinii

NT

4

6

3

13

M

Staurotypus triporcatus

NT

4

7

3

14

H

Family Testudinidae (1 species)

Chelonoidis carbonarius

NE

3

8

6

17

H

August 2015 | Volume 9 | Number 2 | el 00

Amphib. Reptile Conserv.

94

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [General Section]: 95-99 (el 01).

SHORT COMMUNICATION

First report of the salamanders Bolitoglossa leandrae and

B. tamaense (Urodela, Plethodontidae) for Venezuela

^esar L. Barrio-Amoros, 2 Andres Chacon-Ortiz, 3 ’ 4 Fernando J.M. Rojas-Runjaic

l Fundacion AndtgenA, Apartado Postal 210, 5 101 -A Merida, VENEZUELA. Current address: Doc Frog Expeditions, Uvita, COSTA RICA 2 Centro

de Estudios de Vectores de Enfermedades, Decanato de Investigacion, Vicerrectorado Academico, Universidad Nacional Experimental del Tdchira,

San Cristobal, estado Tdchira, VENEZUELA Museo de Historia Natural La Salle, Fundacion La Scdle de Ciencias Naturales. Apartado Posted

1930, Caracas 1010-A, VENEZUELA 4 Laboratdrio de Sistemdtica de Vertebrados, Pontificia Universidade Catolica do Rio Grande do Sul (PUCRS).

Av. Ipiranga 6681, Porto Alegre, RS, 90619-900, BRAZIL

Key words. Caudata, biogeography, Amazon, Orinoquia, Andes, Colombia

Citation: Barrio-Amoros CL, Chacon-Ortiz A, Rojas-Runjaic FJM. 201 5. First report of the salamanders Bolitoglossa leandrae and B. tamaense (Uro-

dela, Plethodontidae) for Venezuela. Amphibian & Reptile Conservation 9(2) [General Section]: 95-99 (el 01).

Copyright: © 201 5 Barrio-Amoros et al. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCommer-

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-

reptile-conservation. org> .

Received: 11 May 2015; Accepted: 15 August 2015; Published: 5 September 2015

Salamanders of the Family Plethodontidae constitute a

major batrachological element in the Neotropic realm,

though descending in species richness from North to

South. Venezuela has an impoverished list of five spe-

cies of salamanders so far: Bolitoglossa altamazonica

(Cope 1874), B. borhurata Trapido 1942; B. guarama-

calensis Schargel, Garcfa-Perez, and Smith 2002; B. or-

estes Brame and Wake 1962, and the recently described

B. mucuyensis Garcfa-Gutierrez, Escalona, Mora, Draz

de Pascual, and Fermrn 2013. The best studied species,

both taxonomically and genetically, is B. orestes. An ap-

parently isolated population was described as B. spongai

by Barrio-Amoros and Fuentes (1999), and later some

ecological traits were published (Barrio-Amoros et al.

2010). Inconsistencies of the formal description and mo-

lecular data led Fermrn et al. (2012) to conclude that B.

spongai is a junior synonym of B. orestes , a position that

we follow here. We use the order name Urodela Dumeril

1805 instead of Caudata Fischer von Waldheim 1813,

following the Dubois and Raffaelli (2012) rationale.

Schargel and Rivas (2003) assigned tentatively the

juvenile specimen ULABG (Universidad de Los Andes,

Laboratorio de Biogeograffa, Merida, Venezuela) 3392

to Bolitoglossa altamazonica , but the evidence they of-

fered (a series of measurements) are hard to corrobo-

rate as clearly diagnostic for this species, especially be-

cause the only comprehensive description of the species

(Brame and Wake 1963) is old and needs verification and

comparison with topotypic specimens (D.B. Wake, pers.

com.; Brcko et al. 2013).

Recently, Acevedo et al. (2012) described two sala-

manders of the genus Bolitoglossa from the Colombian

side of the Tama Massif in the Cordillera Oriental de

Colombia. The southwestern half of this massif is Co-

lombian and the northeastern half is Venezuelan, but

geologically and ecologically it represents a continuum.

Bolitoglossa leandrae Acevedo, Wake, Marquez, Silva,

Franco, and Amezquita 2013 was diagnosed as the small-

est Bolitoglossa known from Colombia, with 30.3 mm

mean snout-vent length (SVL) for males and the only

female known of 39.2 mm SVL, 23-24 maxillary teeth

(MT), and 18-19 vomerine teeth (VT). It inhabits low-

land piedmont rainforest at around 600 m asl. On the

other hand, B. tamaense Acevedo, Wake, Marquez, Silva,

Franco, and Amezquita 2013 is a somewhat larger spe-

cies with males up to 40.3 mm and females up to 52.7

mm, 38-42 maxillary teeth, and 17-23 vomerine teeth

(including both males and females). Genetic data also

confirm the proper specific status of both species.

Here we report for the first time the presence on Ven-

ezuelan territory of two species of salamanders ( Boli-

toglossa leandrae and B. tamaense ). The citation of B.

altamazonica by Schargel and Rivas (2003) is probably a

misidentified B. leandrae , but we cannot be certain as we

Correspondence. Email: 1 cesarlba@yahoo.com (Corresponding author); 2 aecortiz@yahoo.com; 3 rojas _runjaic@yahoo.com

September 2015 | Volume 9 | Number 2 | el 01

Amphib. Reptile Conserv.

95

Barrio-Amoros et al.

were unable to access the specimen ULABG 3392 — the

specimen is a juvenile that makes its proper identification

more difficult.

Specimens CVUNET (Coleccion de Vertebrados,

Universidad Experimental del Tachira, San Cristobal,

Venezuela) 644 (female; Fig. IB), CVUNET 645 (male;

Fig. 1A), CVUNET 669 (male), and CVUNET 670

(male) from Quebrada La Espuma, Rio Frio, Tachira

state 7.3540 N, 72.1012 W, 650 m asl (Fig. 2), are herein

assigned to B. leandrae by having all the set of charac-

ters diagnostic for the species in Acevedo et al. (2013),

such as a very small size; actually the smallest species of

Bolitoglossa of Colombia and Venezuela (females up to

39.2 mm; in our sample, females up to 34.4 mm and our

sample of males expands the maximum size to 35 mm),

extensive webbed digits on hands and feet (see Fig 1A

and IB; Table 1 for measurements).

CVUNET 615 (female; Fig. 1C), CVUNET 626 (fe-

male; Fig. ID), CVUNET 703 (sex unknown) CVUNET

726 (sex unknown), from Matamula, between Bramon

and Delicias, Tachira state, 7.2833 N, 72.4333 W, 2,020

m asl (Fig. 2), and MHNLS 1268 (male) from Rio Chiq-

uito, Junfn, Tachira state, 7.32 N, 72.20 W, ca. 2,000 m

asl (Fig. 2), are assigned herein to B. tamaense following

the diagnostic characters given by Acevedo et al. (2013),

such as small size between the range given by the origi-

nal description, the webbed hands with broadly triangu-

lar and pointed finger tips (Fig 1C), coloration similar to

that in Fig 3E and G in Acevedo et al. (2012) (see our Fig

1 C, D); measurements presented in Table 1.

With the data at hand, the range of MT is slightly

wider in both species, ranging now from 21-24 for males

and 28-29 for females of B. leandrae ; and from 31-39

for males of B. tamaense. The same is valid for VT, with

males now ranging from 17-19 and females from 18-20

in B. leandrae ; and males ranging from 16-19, and fe-

males 17-23 in B. tamaense.

As explained, the area where both species occur in

Colombia and Venezuela, conform a continuum, only

separated by an artificial frontier line on maps. Bolito-

glossa leandrae inhabits primary (in Venezuela) and sec-

ondary (in Colombia) lowland rainforests up to 650 m asl

(600 m in Colombia). It is active at night on vegetation

up to 1.5 m (own observations). Bolitoglossa tamaense

occurs at cloud forests between 2,000 and 2,700 m asl

(2,000 to 2,020 m in Venezuela), also on low vegetation

and mossy rocks.

The Valle del Rio Doradas is an important area for

Orinoquian and Upper Amazonian herpetofauna (con-

trasting with the surrounding typical Llanos and Andes

elements), as demonstrated by Barrio-Amoros and col-

leagues for other amphibian species like Hypsiboas lan-

ciformis Cope 1871, H. boans (Linnaeus 1758), Scinax

wandae (Pybum and Fouquette 1971), Lithodytes linea-

Fig. 1 . Bolitoglossa leandrae : subadult male CVUNET 645 (A) and adult female CVUNET 644 (B), both from Quebrada La

Espuma, Rio Frio, Tachira state, Venezuela. Bolitoglossa tamaense-. adult females CVUNET 615 (C) and CVUNET 626 (D), both

from Matamula, between Bramon and Delicias, Tachira state, Venezuela. All photos by CBA except D by ACO.

September 2015 | Volume 9 | Number 2 | el 01

Amphib. Reptile Conserv.

96

Two new salamander species reported in Venezuela

71°W

Fig. 2. Known distribution of Bolitoglossa leandrae (open squares) and B. tamaense (open circles) in Colombia and Venezuela.

Colombian records are from Acevedo et al. (2013). 1: Matamula, between Bramon and Delicias. 2: Rio Chiquito. 3: Quebrada la

Espuma, Rio Frio.

tus (Schneider 1799), and Rhaebo glaberrimus (Gunther

1869), among others (respectively Barrio et al. 1999;

Barrio, 1999; 2001; Barrio-Amoros and Chacon, 2004;

Chacon et al., 2002) and therefore, we cannot rule out

that ULABG 3392 is indeed Bolitoglossa altamazonica,

though we retain it as B. aff. altamazonica. Thus, we do

not exclude this late species from the list of Venezuelan

amphibians, but caution about the proper identification

of further specimens from the same general area. Both,

morphological and genetic data would be desirable to

identify this species complex in the Upper Amazon of

Peru, Ecuador, and Colombia, continuing the study of

Brazilian material by Brcko et al. (2013).

Acknowledgments. — We thank Andres Orellana,

William Tovar, and Valeria Bellazzini for their company

in the field and to Carla Ochoa for helping to take mea-

surements at the UNET’s Lab. Sean Rovito and three

anonymous reviewers improved the original version of

the paper.

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the eastern Colombian Andes. Zootaxa 3609: 69-84.

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dae) de los Andes venezolanos, con comentarios sobre

el genero en Venezuela. Acta Biologica Venezuelica

September 2015 | Volume 9 | Number 2 | el 01

Amphib. Reptile Conserv.

97

Barrio-Amoros et al

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September 2015 | Volume 9 | Number 2 | el 01

Amphib. Reptile Conserv.

98

Two new salamander species reported in Venezuela

19: 9-19.

Barrio CL, Orellana A, Manrique R. 1999. Geographic

distribution: Anura: Hyla lanciformis. Herpetological

Review 30: 106-107.

Barrio-Amoros CL, Garcia J, Fuentes-Ramos O. 2010.

Preliminary data on natural history and intraespecific

variation of the endangered salamander Bolitoglossa

spongai in the Venezuelan Andes. Salamandra 46:

108-113.

Brame AH, Wake DB. 1963. The salamanders of South

America. Contributions in Science 69: 1-72.

Brcko IC, Hoogmoed MS, Neckel-Oliveira S. 2013.

Taxonomy and distribution of the salamander genus

Bolitoglossa Dumeril, Bibron & Dumeril, 1854 (Am-

phibia, Caudata, Plethodontidae) in Brazilian Amazo-

nia. Zootaxa 3686: 401-431.

Chacon A, Diaz de Pascual A, Barrio CL. 2002 “2000.”

Presencia de Bufo glaberrimus (Anura: Bufonidae) en

Venezuela. Acta Biologica Venezuelica 20: 65-69.

Dubois A, Raffaelli J. 2012. A new ergotaxonomy of the

order Urodela Dumeril, 1805 (Amphibia, Batrachia).

Alytes 28: 77-161.

Fer mfn G, Garcia J, Escalona M, Mora A, Diaz A. 2012.

Molecular taxonomic reassessment of the Cloud For-

est’s Bolitoglossa salamanders (Caudata: Plethodon-

tidae) from Cordillera de Merida (Merida state, Ven-

ezuela). Zootaxa 3356: 47-56.

Frost DR. 2013. Amphibian Species of the World: An on-

line reference. Version 6.0 (09 March 2015). Ameri-

can Museum of Natural History, New York, New

York, USA. Available: http://research.amnh.org/her-

petology/amphibia/index.html [Accessed: 09 March

2015]. ’

Schargel WE, Rivas G. 2003. Two new country records

of salamanders of the genus Bolitoglossa from Co-

lombia and Venezuela. Herpetozoa 16: 94-95.

Cesar L. Barrio-Amoros is an anthropologist who has worked with herpetofauna in Spain, Venezuela, and Costa

p i Rica. His research interests include the biogeography and systematics of amphibia and reptilia, with emphasis in

j| Dendrobatoidea and Terrarana from Venezuela, especially of the Guiana Shield. Now a resident of Costa Rica, he is

H II 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.

Andres Chacon-Ortiz is Associate Professor at the National Experimental University of Tachira (UNET) in Ven-

ezuela. He is the director of the Center for Studies for Vector Diseases at UNET and is interested in Venezuelan

Andean herpetology, especially in amphibian taxonomy and ecology.

Fernando J. M. Rojas-Runjaic is a researcher at the Museo de Historia Natural La Salle in Caracas, Venezuela, and

curator of the amphibian, reptile, and arachnid collections. Fernando is a graduate in biology from the Universidad

del Zulia (Venezuela) and has a master’s degree in biodiversity in tropical areas and its conservation from the Uni-

versidad Internacional Menendez Pelayo (Spain). Currently he is in a Ph.D. programe in zoology at the Pontificia

Universidade Catolica do Rio Grande do Sul (Brazil). His research is broad and includes amphibian, reptile, and

scoipion diversity, phylogenetic systematics, taxonomy, biogeography, and conservation.

Appendix 1. Specimens examined

Bolitoglossa adspersa. MBUCV (Museo de Biologia Uni-

versidad Central de Venezuela, Caracas) 418, from Paramo

de Cruz Verde, Cordillera Oriental, Cundinamarca, Colom-

bia.

Bolitoglossa borburata. EBRG (Museo de la Estacion Bi-

ologica Rancho Grande, Maracay) 3173, from Fila la Guer-

rillera, Sierra de Aroa, Yaracuy state, Venezuela. MBUCV

6563, Altos de Choronf, Aragua state, Venezuela. MBUCV

6664, Rancho Grande, Aragua state, Venezuela.

Bolitoglossa leandrae. CVUNET 644, CVUNET 645,

both from Quebrada La Espuma, Rio Frio, Tachira state,

Venezuela. 7.3540 N, 72.1012 W, 650 m asl, collected on

20 May 2012 by W. Tovar, A. Chacon, and C.L. Barrio-

Amoros. CVUNET 669, CVUNET 670 both from Quebra-

da La Espuma, Rio Frio, Tachira state, Venezuela. 7.3540

N, 72.1012 W, 650 m asl, collected on May 2013, by Wil-

liam Tovar, Lionel Fernandez, and Andres Chacon Ortiz.

Amphib. Reptile Conserv.

Bolitoglossa orestes. MBUCV 6570 (holotype of B. spon-

gai), from Hato La Carbonera, Fila la Cuchilla, Merida state,

Venezuela. MBUCV 6571-72, MCNC (Museo de Ciencias

Nacional de Caracas, Caracas) 8116-17, EBRG 3583-84,

all from the same last locality and referred as paratypes of

B. spongai. MCNC 6432, 6484, from San Eusebio, Andres

Bello District, Merida state, Venezuela.

Bolitoglossa tamaense. MHNLS 1268. Rio Chiquito, Junin

municipality, Tachira state, Venezuela (7.32 N, 72.20 W,

ca. 2,000 m asl), collected on February 1956, by Ramon

Urbano. CVUNET 615, CVUNET 626, both from Matam-

ula, between Bramon and Delicias, Tachira state, 7.2833 N,

72.4333 W, 2,020 m asl, collected on February 2012 by W.

Tovar, A. Chacon and C.L. Barrio-Amoros, CVUNET 703,

CVUNET 726 both from Matamula, between Bramon and

Delicias, Tachira state, 7.2833 N, 72.4333 W, 2,020 m asl,

collected on June and August 2013 respectively, by Marian

Chacon Jaimes, Andres Chacon Ortiz, and Carla Ochoa.

September 2015 | Volume 9 | Number 2 | el 01

99

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [General Section]: 100-110 (el 02).

Breeding and rearing the Critically Endangered Lake Oku

Clawed Frog ( Xenopus longipes Loumont and Kobel 1991)

1V1 Christopher J. Michaels, ^ Benjamin Tapley, ^uke Harding, ^oe Bryant, Sebastian Grant,

George Sunter, ^ri Gill, 2 Oscar Nyingchia, and 3 Thomas Doherty-Bone

1 Zoological Society of London, Regent’s Park, London, UNITED KINGDOM NW1 4RY "-Conservation Research and Action for Amphibians Unique

to Cameroon (CRAAUC), Royal Zoological Society of Scotland, Edinburgh Zoo, Corstorphine Road, Edinburgh, UNITED KINGDOM Needs

University, Woodhouse Lane Leeds UNITED KINGDOM LS2 9JT

Abstract . — The Lake Oku Clawed Frog Xenopus longipes is a Critically Endangered, dodecaploid

anuran endemic to Lake Oku in Cameroon. An ex situ population of this species was established

at Zoological Society of London (ZSL), London Zoo in 2008, as well as at several other institutions,

with the intention of providing data on the biology and husbandry of this species. We report the first

captive breeding of the species. Adult frogs maintained under environmental conditions designed

to mimic field data produced clutches of 7-300 eggs; eggs measured 1.23 mm in diameter, and

were laid singly after a period of 6.5 hours in axial amplexus. Spawning took place only during the

day. Tadpoles hatched in 2-3 days and development was very long compared to congeners, lasting

193-240+ days until metamorphosis. Tadpoles grew very large (maximum 79 mm total length),

particularly compared with the relatively small adult size (maximum 36 mm Snout to Vent Length

[SVL]). Tadpoles proved to be highly sensitive to total dissolved solids (TDS) in the water and only

thrived when low levels (20 mg/L) were used. Metamorphosis concluded with an SVL of 19-25 mm

and FI animals began first sexual activity at 5-6 months post metamorphosis. These data will

inform future husbandry in captivity as well as illuminating facets of biology previously unknown

and difficult to determine in the field.

Key words. Amphibian; ex situ ; captive husbandry; water quality; Cameroon; West Africa; field data

Citation: Michaels CJ, Tapley B, Harding L, Bryant Z, Grant S, Sunter G, Gill I, Nyingchia O, Doherty-Bone T. 2015. Breeding and rearing the Criti-

cally Endangered Lake Oku Clawed Frog ( Xenopus longipes Loumont and Kobel 1991). Amphibian & Reptile Conservation 9(2) [General Section]:

100-110 (el 02).

NoDerivatives 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 June 2015; Accepted: 20 August 201 5; Published: 1 8 September 201 5

The creation of ex situ populations for research and con-

servation breeding has become an important part of the

international conservation response to global amphibian

declines (Browne et al. 2011; Gascon 2007; Koute et al.

2012; Wilkinson et al. 2013), which represent one of the

greatest conservation challenges in history (Zippel et al.

2011). The requirements of amphibians in captivity are

poorly understood and many species are presently dif-

ficult to maintain and breed (Antwis et al. 2014; Antwis

and Browne 2009; Browne et al. 2006; Dugas et al. 2013;

King 2011; Ogilvy et al. 2012; Verschooren et al. 2011).

Ex situ programs have experienced difficulty in provid-

ing conditions under which animals survive (Norris 2007;

Gagliardo et al. 2008) or successfully breed (Birkett et al.

1999; Gratwick 2012). Moreover, information on how to

rear tadpoles is particularly lacking in peer reviewed lit-

erature (Pryor 2014).

The Lake Oku Clawed Frog Xenopus longipes Lou-

mont and Kobel 1991 (Fig. 1) is an entirely aquatic, do-

decaploid frog found only in Lake Oku, a high elevation

crater lake in the north west region of Cameroon. Xeno-

pus longipes is classified as Critically Endangered by

the IUCN (Stuart et al. 2008) due to its restricted range

and therefore vulnerability to stochastic factors. Between

2006 and 2010 recurring, enigmatic X. longipes morbidi-

ties and mortalities were observed, but the overall impact

of these events is unknown (Doherty-Bone et al. 2013).

A captive-breeding program was considered vital in case

of a catastrophic collapse of the population due to the

potential introduction of fish to the lake as well as habi-

Correspondence. Email: 1 christophenmichaels@zsl.org (Corresponding author). These authors contributed equally to the work.

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

100

Michaels et al.

tat degradation and disease threats (Tinsley and Measey

2004a). Xenopus longipes is ranked as the 35 th global pri-

ority for amphibian conservation on the basis of threat

and evolutionary history by the Zoological Society of

London’s Evolutionarily Distinct and Globally Endan-

gered (EDGE) program (Isaac et al. 2012).

Captive colonies of the Critically Endangered X. lon-

gipes were established in 2008 at Antwerp Zoo (later

moved to Cologne Zoo and one private breeder), Zoo-

logical Society of London (ZSL), London Zoo, and more

recently in 2013, at the Steinhart Aquarium in the USA,

for conservation research purposes (Browne et al. 2009;

T. Ziegler pers. comm.; P. Janzen pers. comm.; D. Black-

burn pers. comm.). The zoo colonies were intended to be

assurance populations for conservation breeding. How-

ever, due to concerns over biosecurity and suitability

of animals for release to the wild, the ZSL population

was assimilated into the main collection with the focus

now on conservation research aiming to document the

reproductive biology of the species, as little is currently

known. Such information is of importance for developing

in situ conservation management strategies. Despite re-

peated attempts in all these institutions, however, efforts

to breed and rear this species in captivity have failed,

even with the use of artificial reproductive techniques (P.

Janzen, pers. comm.; D. Blackburn pers. comm.).

A

Fig. 1 . Male (top) and female (bottom) adults of Xenopus lon-

gipes in the collection at ZSL London Zoo (ZIMS ID 7441).

Here we report the first captive breeding success of X.

longipes and the rearing of the tadpoles until metamor-

phosis.

Methods

In 2008, frogs were collected from Cameroon after con-

sultation with local communities (Permit No. 0742/CO/

MINFOF/SG/DFAP/SDVEF/SC and No. 0928/PRBS/

MINFOF/SG/DFAP/SDVEF/SC). Lake Oku is consid-

ered sacred by the Oku villages and permission had to be

granted before any contact with the lake could be made.

Thirty-nine founders were housed at Zoological Society

of London (ZSL), London Zoo.

Table 1 summarizes the initial and subsequent hus-

bandry used for these frogs between 2008 and 2014. In

2012 the husbandry of X. longipes was reviewed (Table

1) as breeding had not occurred and the temperature re-

gime and water parameters did not reflect conditions in

the field (Table 2). Captive management should be in-

formed by field data (Tapley and Acosta 2010; Michaels

and Preziosi 2013; Michaels et al. 2014) and replicating

field conditions has improved captive breeding success

of X. laevis (Godfrey and Sanders 2004). In 2012, all

32 (30.2) remaining founders were sexed; males be-

ing smaller, slimmer, and having keratinized nuptial

pads (Fig. 2A and C) and females possessing a trio of

cloacal papillae (Fig. 2B). These features became more

prominent around breeding events, but were noticeable

year round. All 30 female frogs were continuously heav-

ily gravid and amplexus was occasionally observed, but

without spawning. Additional founders including four

more males were imported from Cameroon in July 2012

and after completing their quarantine period were assimi-

lated into the existing X. longipes colony.

In June 2013 mixed sex groups varying from 1.6 to

3.3 were transferred to a custom-made system (Fig. 3;

Table 1). A new environmental regime based on longi-

tudinal field data collected monthly from Lake Oku by

Doherty Bone et al. (2013) was adopted (Tables 1 and 2;

Fig. 4). Lake Oku water temperature and pH were simu-

lated initially, and Total Dissolved Solids (TDS) was

subsequently added to the parameters being replicated in

2014 (Table 1). Total Dissolved Solids was measured us-

ing Micro 800 Optical DO meter (Palintest) and pH using

a Micro 600 pH meter (Palintest). The feeding regime

was also modified (Table 1) and a more diverse array of

food items were offered to compensate for potential di-

etary deficiencies as knowledge regarding the nutritional

requirements of amphibians is lacking (Densmore and

Green 2007).

Results

On 20.3.14, two pairs of X. longipes spawned naturally

and without hormonal induction, followed by a number

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

101

Breeding and rearing the Lake Oku Clawed Frog

Table 1 . Changes in enclosures, life support systems, environmental parameters and diets used for X. longipes between 2008 and

2014. Reproduction occurred in 2014.

Dates

Enclosure

Life support systems

Photo-

Water

Diet

type and size

and furnishings

period

parameters

Acrylic aquaria

pH c. 8.5

Blood worm ( Chironomus ), Nutrafin

2008-

(Exo Terra, Rolf

Air-stream sponge filter;

10:14

TDS: c. 350 mg/L

Max cichlid sinking capsules, Tetra

2012

C. Hagen) 20 L

Plastic plants

Temperature: 19-22 °C

prima granules and King British blood

enclosures

pH c. 8.5

worm (freeze dried)

Mixed invertebrates: blood worm ( Chi-

ronomus sp); glass worm ( Chaoborus

2012

Acrylic aquaria

Air-stream sponge filter;

10:14

TDS: c. 350 mg/L

crystallinus ); water fleas ( Daphnia sp);

20 L enclosures

Plastic plants

Temperature: 18-19 °C

hatchling crickets ( Gryllus bimaculatus

and G. assimilis ) and worms (. Eisenia

sp.)

48 L enclosures

linked to 100 L

TRIO Teco chiller/heater and

UV filter; External canister

pH c. 7.5

2013

filter (FX6 Fluval). Plastic

tubes, plastic and five plants

12:12

TDS: c. 150 mg/L

Temperature: 17-19 °C,

Mixed invertebrates (as above)

sump

(' Vallisneria spp., Echinodor-

us spp.)

with seasonal variation

48 L enclosures

linked to 100 L

sump

TRIO Teco chiller/heater and

UV filter; External canister

pH c. 7.5

TDS: 20 mg/L

Temperature: 17-19 °C

with seasonal variation

2014

filter (FX6 Fluval). Plastic

tubes, plastic and five plants

(' Vallisneria spp., Echinodor-

us spp.)

12:12

Mixed invertebrates (as above)

of other spawning events (Table 3). In the initial spawn-

ing event a single pair in each of two tanks containing

1.2 animals spawned. Audible vocalizations, consisting

of metallic clicks typical for Xenopus (Tinsley and Kobel

1996) were only heard very infrequently from spawning

and non-spawning males and were not closely associated

with spawning activity; being heard sporadically during

both spawning and non-spawning periods. Amplexus and

spawning behavior were only observed throughout the

day, with no evidence of spawning occurring over-night.

Amplexus was axial and the process of oviposition lasted

6.5 hours from initiation to termination of amplexus.

Eggs, numbering seven to 300 per clutch (Table 3), were

deposited singly over all available surfaces in aquaria.

Egg diameter was 1.23 mm one hour after laying. Oc-

casionally, multiple males attempted to amplex single

females, but were dislodged by vigorous kicking on the

part of the original male. Laying and non-laying females

were observed feeding on the eggs, even during amplex-

us, so non-amplectant animals were removed immediate-

ly. Mating pairs were removed as soon as spawning was

complete. Animals could not be individually identified,

so it is unclear how many clutches were produced by

individual animals. The initial spawning event occurred

after increasing the temperature from 17.5 to 19.1 °C.

This was done by adding warm tap water to the system,

resulting in a pH shift from 7.5 to 8.09 to replicate the

seasonal temperature and pH regime in Lake Oku, al-

though being done two months earlier than this shift oc-

curs in the field (Fig. 4). This shift occurred over a period

of less than one hour after warm water was added in a

single dose. However, as the breeding season is not docu-

mented in the field, there is no evidence that this seasonal

change accompanies the initiation of breeding in nature,

other than this relationship observed in congeneric spe-

cies (Kobel et al. 1996). Later spawning events in the

following months (see Table 3) were not associated with

manipulation of water parameters, but did follow heavy

feeds with earthworms ( Eisenia sp.). Lertility was highly

variable; some clutches were almost entirely infertile, but

in most cases fertility rates were close to 100%. Eggs de-

veloped and hatched in 2-4 days, with tadpoles initially

clinging to hatch sites via the cement gland. Eventually

eyes and pigmentation developed before becoming free

swimming after 2^1 days. Free-swimming tadpoles ini-

tially congregated in areas of slow current, swimming

against the water flow. Hatch rate varied between clutch-

es, with later clutches being more consistently successful

than earlier clutches.

A variety of combinations of conditions were used in

attempts to rear tadpoles (see Table 4). However, we only

had success by maintaining tadpoles in water with a very

low TDS of 20 mg/L (measured at roughly weekly inter-

vals) and without any live plants or accumulation of hu-

mic detritus, and only in aquaria isolated from the adult

system possibly as a result of secretions from adults or

toxins from PVC pipework used in the aquatic system.

Mortality of tadpoles remained high until the TDS of

the systems fell below 80 mg/L, with tadpoles becom-

ing weak, opaque, and finally sinking to the floor of the

aquaria before dying. Following gradual replacement of

high TDS water with low TDS reverse-osmosis water,

surviving tadpoles began to feed, swim normally, and to

develop. Doherty-Bone et al. (2013) report a TDS of <10

mg/L (See Table 2), but our value of 20 mg/L was the

lowest possible output from the RO system in use (Pen-

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

102

Michaels et al.

A

Fig. 2. Keratinized nuptial pads on the inside surfaces of the front limbs of male (A and C) and cloaca of a female X. longipes (B);

note the cloacal papillae, which are absent in male frogs.

tair PRF; Fileder) and appears to be adequate for larval

rearing.

Tadpole enclosures were glass aquaria measuring 50 x

36 x 30 cm (L x W x H) held in a temperature controlled

room with water temperature at 18-20 °C. Between three

and 15 tadpoles were housed per aquarium (maximum

density of one tadpole per 3.6 litres). Aquaria were fil-

tered with air-stream sponge filters set to the minimum

effective flow to reduce turbulence, which would have

disturbed the swimming and foraging behavior of tad-

poles. Tadpoles were fed 2-4 times throughout the day

on a suspension consisting initially of blanched and

blended spinach or nettle, commercial Xenopus tadpole

food, SERA Micron powdered food, and Spirulina alga,

which was strained prior to use to remove larger plant

fragments. After several weeks, the diet was changed to

only include commercial Xenopus tadpole food, SERA

Micron (SERA), and Spirulina (3:1:1 by mass, suspend-

ed in water before adding to aquaria) to avoid the high

oxalate content of spinach (Noonan and Savage 1999),

which may interfere with calcium metabolism (Rosol et

al. 1995). Food was added throughout the day depen-

dent on the rate at which food was consumed in a given

aquarium, with food density of 5.3 mg/L aquarium wa-

ter provided immediately after feeding; density reduced

gradually as food was consumed by tadpoles. Uneaten

suspended food accumulated on the bottom of aquaria,

where tadpoles were unable to consume it. Additionally,

the low carbonate content of the water reduced the capac-

ity for biological filtration. Consequently, nitrogenous

Table 2. Water temperature, pH, and TDS measured at the Lake Oku shoreline (modified from Doherty-Bone et al. 2013).

Parameter

Mean Value

+

Units

Water temperature

17.27

4.17

Celsius

pH

7.58

0.24

-

Total Dissolved Solids

8.72

2.27

Ppm

Amphib. Reptile Conserv.

103

September 2015 |

Volume 9 | Number 2 | el 02

Breeding and rearing the Lake Oku Clawed Frog

Fig. 3. Aquarium for X. longipes, set within a custom built, centrally filtered system (inset photograph) at ZSL London Zoo. Life

support system and sump not shown - see text for details.

waste (measured using Photometer 7100 [Palintest]) was

difficult to manage and tadpoles were briefly exposed to

high levels of ammonia (>1 mg/L) and, later, nitrite (up

to 2.4 mg/L) without mortality. A regime of 10% water

changes in the morning and 50% water changes in the

afternoon, both accompanied by removal of uneaten food

on the bottom of tanks by siphon and thorough cleaning

of sponge filters in aquarium water, helped to suppress

nitrogenous waste to more acceptable, but still detectable

levels (Ammonia: <0.1 mg/L; Nitrite: <0.5 mg/L) for

most of the tadpole rearing period.

The tadpoles of X. longipes are described separately

(Tapley et al. 2015). Development in the most rapidly

developing tadpole (Fig. 5) lasted 193 days between

hatching and metamorphosis. We report development

using Gosner (1960) stages, as it was impossible to ac-

curately apply the more detailed Nieuwkoop and Faber

(1994) stages for Xenopus laevis development to live

tadpoles without restraining them. This would likely

have proven fatal for these delicate and Critically En-

dangered tadpoles, though could be employed in future

offspring once captive population growth has been as-

sured. However, developmental rates were highly vari-

able and the more slowly developing tadpoles had not

yet metamorphosed at the time of writing. A maximum

total length of 68 mm was reached in the first tadpole to

metamorphose (Fig. 5), and the largest tadpole reached a

maximum total length of 79 mm. Metamorphs measured

19-25 mm SVL and captive-bred males began to exhibit

amplexus six months post metamorphosis, by which time

they had nearly reached adult size. Further details of tad-

pole development are provided by Tapley et al. (2015).

Once front limbs emerged from the operculum, tadpoles

were separated by placing them in identical systems with

Table 3. Spawning dates and clutch sizes for X. longipes.

Date

Clutch number

Clutch size

20.03.14

1

93

2

115

21.03.14

1

190

22.03.14

1

40-50

2

40-50

05.04.14

1

40

25.08.14

1

50

04.09.14

1

Not counted

16.09.14

1

20

17.09.14

1

120

18.09.14

1

80

20.09.14

1

Not counted

29.09.14

1

50

04.10.14

1

120

05.10.14

1

300

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

104

Michaels et al.

Month

Fig. 4. Monthly water temperatures (circles) and pH (triangles) recorded from the shoreline of Lake Oku between 2008/2009 and

2013. Error bars represent SEM.

Table 4. Combinations of conditions used to rearX. longipes tadpoles, and the outcome in terms of tadpole survival.

Water TDS

(mg/L)

Refugia

(live plants)

Detritus

Lighting

Tannins

Isolated from

adult system?

Tadpoles

survived?

-

+

-

+

-

+

-

+

-

+

-

+

-

20

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

+

-

150

-

+

-

+

-

+

-

+

-

+

-

+

-

+

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

105

Breeding and rearing the Lake Oku Clawed Frog

Qi

cuo

to

■M

1—

0 )

c

o

(5

46

44

42

40

38

36

34

32

30

28

26

24

22

20

18

20

40

60

80

100

120

140

160 180

Days after hatching

Fig. 5. Gosner stage progression of the most rapidly developing X. longipes tadpole. Hatching to metamorphosis took 193 days, but

smaller tadpoles had only reached stage 35 by this point.

sponge filters that had been matured in the system hous-

ing the adult frogs, but with a shallower water depth of

15 cm to facilitate access to the surface for breathing.

Metamorphosis from this point took around seven days

to complete. Froglets fed on a similar range of prey items

to adults.

Discussion

Although the husbandry of adult X. longipes is largely

similar to that established for other Xenopus species

(Green 2012), and adult frogs are able to survive a range

of water parameters, the tadpoles are more sensitive. The

dietary requirements of tadpoles are similar if not identi-

cal to those of X. laevis and X. tropicalis, but tadpoles

appear to be more sensitive to the mineral/solute content

of water. Tadpoles maintained in water with a TDS high-

er than around 80 ppm died rapidly and tadpoles devel-

oped well with a TDS of around 20 ppm. Total Dissolved

Solids represent the total amount of dissolved mobile

charged ions, including minerals, salts or metals and is

closely related to hardness, but includes a broader range

of dissolved substances. Typically, very low solute con-

tent of aquarium water can lead to osmotic imbalances in

amphibians, but species may adapt evolutionarily to rela-

tively pure water (Odum and Zippel 2008). Sensitivity to

hardness or TDS values in Xenopus is not without prece-

dent. The tadpoles of the softwater specialist X. gilli from

the Cape also appears to be intolerant of hard, alkaline

conditions (Rau 1978), while the reproductive success of

captive X. laevis is improved by matching the hardness

of their wild environment (Godfrey and Sanders 2004).

Other amphibians including the Hellbender Cryptobran-

chus alleganiensis have been shown to be reproductively

sensitive to TDS levels (Ettling et al. 2013). The closely

related X. amieti has been reproduced with hormonal

induction in captivity (Xenopus Express pers. comm.),

and the tadpoles of this species were maintained in hard

water with success. However, there are no field data for

water quality in its wild range and the larger distribu-

tion of X. amieti , which is not restricted to a single lake

(Tinsley and Measey 2004b), may have led to the evo-

lution of less specific environmental requirements. Our

combinations of environmental conditions, summarized

in Table 3, were not fully exhaustive and so the effects

of some parameters (particularly tannins) cannot be fully

elucidated based on these data. Tannins are thought to be

important in reducing the frequency of fungal infections

in the tadpoles of some anuran species (e.g., Theloderma

corticale ; Rauhaus et al. 2012), but there are no data

concerning the tannin levels in Lake Oku. The forested

shores of Lake Oku do produce inputs of leaf litter (so

far unquantified) suggesting some levels of tannins, but

this needs to be confirmed. Underwater photographs of

the lake suggest relatively clear water (T. Doherty-Bone

pers. obs.), which may mean that tannins are unimportant

or potentially harmful in this species.

Amplexus and egg-laying behavior is similar to other

Xenopus, although we did not observe calling in close

association to spawning. Indeed, calls were very rare

in general and we were unable to record them despite

repeated efforts. Amplexus and oviposition were ex-

clusively diurnal, in comparison to the often nocturnal

habit of X. laevis (Green 2012) and the apparent strict-

ly nocturnal amplexus, calling, and spawning reported

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

106

Michaels et al.

from hormone induced X. amieti (Xenopus Express pers.

comm.). Specific triggers involved in stimulating spawn-

ing activity remain unclear. In a species from a habitat

that is relatively stable year round (Fig. 4), and with no

periods of drought and pond drying, it is possible that

reproduction can take place year round and strong en-

vironmental stimuli are not required. Although initial

spawning was associated with a change in temperature

and pH, the breeding season is not documented in the

wild and there is no evidence that this seasonal change

accompanies the initiation of breeding in nature. Our ob-

servations suggest that heavy feeding may contribute to

spawning activity, and so breeding may be more linked

to a threshold in body condition than to external triggers.

Kobel et al (1996) have suggested that some Xenopus

species breed following first rains, when nutrients in the

water have increased and secondary productivity of in-

vertebrates is thus stimulated. Our observations indicated

the initial stimuli of changing temperature, but correlated

more strongly with increased availability of food. These

speculations merit further investigation.

Clutch size (7-300; Table 3) was smaller than that

produced by X. laevis (500-30,000 eggs Green, 2012)

or X. tropicalis (1,000-3,000 eggs; Green, 2012). This

may partly reflect the smaller body size of X. longipes,

but may also be a function of breeding in a more stable

lake system habitat, where there may be advantage in

producing a smaller number of larger eggs. The fact that

egg size is similar for X. laevis and X. longipes (1.3 mm

[Brown 2004] and 1.23 mm, respectively), as well as for

a number of other Xenopus species much larger than X.

longipes (Kobel et al. 1996) supports this hypothesis.

The pattern of small clutch size and relatively large eggs

is continued in the very large tadpoles of this species

(maximum total length 79 mm), particularly compared

with adult size (32-36 mm snout-to-vent length [SVL];

Loumont and Kobel 1991); see Tapley et al. (2015). The

closely related X. amieti , which has larger adults than X.

longipes, has a tadpole of only 40 mm total length (Chan-

ning and Rodel 2012), while the very large X. laevis has

tadpoles of 80 mm compared with adults of over 140 mm

SVL (Green 2012).The metamorphs of X. longipes are

correspondingly large relative to adult size, being similar

in size to the metamorphs of X. laevis despite a fivefold

difference in adult size between the two species (see Ta-

pley et al. (2015), for further discussion of larval size).

Larval development was slower in X. longipes than

congeners. Larval duration was 193 days at 17-19 °C for

the fastest developing larva, in comparison to the faster

development of X. laevis (42-56 days (Green 2012); 53

days at 18 °C; Gomez-Mestre et al. 2010) orX. tropica-

lis (21—42 days; Green 2012). Several healthy tadpoles

of X. longipes remained untransformed at 240 days post

hatching. This may, again, be linked to a relatively sta-

ble breeding habitat at higher altitude, where very low

seasonal variation in environmental parameters (Fig. 4),

lower temperatures, and no risk of the water body drying

out may select for a longer larval phase (Werner 1986).

In X. gilli, which is found in more temperate lowland

habitat in the extreme south of the African Cape, lower

temperatures comparable with those measured in Lake

Oku are also associated with the long developmental du-

ration of this species (120 days; Rau 1978), albeit still

shorter than for X. longipes.

The observations presented herein provide the first

insight into the behavior, development, and captive re-

quirements in X. longipes. This is of particular note as

to the best of our knowledge the tadpoles of this spe-

cies have never been observed alive in the field and so

nothing is known of their habits in nature. In particular,

the high sensitivity to mineral content and smaller clutch

size of this species than in commonly maintained Xeno-

pus may make X. longipes more susceptible to aquatic

pollution and less able to recover quickly from declines.

Moreover, this characteristic highlights the limitations of

the “analogue species” concept (Preece 1998; Michaels

et al. 2014), whereby common relatives of a threatened

species are used as models to develop husbandry strate-

gies before working with target, usually Critically En-

dangered, species. The relative ease of breeding and rear-

ing X. laevis in captivity does not entirely transfer to X.

longipes, particularly where water TDS for tadpoles is

concerned.

Our findings will hopefully improve success with this

species in other institutions, and contribute to the long-

term viability of captive colonies. This includes attain-

ing reproduction from the first generation of captive bred

X. longipes. Once reproduction is achieved regularly, a

studbook should be developed to ensure that a viable

population of this species is maintained in captivity long-

term, both for conservation breeding and for research

purposes. A studbook would require individual marking

techniques as X. longipes do not have distinctive skin

markings. Such marking techniques has not yet been tri-

alled in this species. Xenopus longipes is one of only two

vertebrates known to be dodecaploid (the other being X.

ruwenzoriensis ) and so there is considerable interest in

this species as a model laboratory organism. Inclusion

of X. longipes in research captive colonies may help to

secure the future of this species in captivity.

Although the current captive populations of X. lon-

gipes are not managed under strict enough biosecurity

controls to be suitable for reintroduction efforts (IUCN/

SSC 2014), laboratory techniques for other Xenopus

exist to generate “clean” animals (e.g., Kay and Peng

1991). There is therefore potential to use these tech-

niques to create biosecure cohorts that could safely be

used for reintroduction should it be required. Moreover,

husbandry protocols can also be distributed to Cameroo-

nian specialists so that conservation breeding facilities

can be developed in country if necessary; this option is

often preferable due to reduced risk of disease transmis-

sion and reduced cost. More work is required to fully un-

September 2015 | Volume 9 | Number 2 | el 02

Amphib. Reptile Conserv.

107

Breeding and rearing the Lake Oku Clawed Frog

derstand and control the reproduction of this species in

the laboratory as well as the field.

Conclusions

Although superficially similar to other Xenopus species

better established in captivity, the husbandry and cap-

tive breeding of X. longipes differs in several important

aspects. The breeding triggers are poorly defined and

less obvious than for many other species, which often

breed in response to large water changes with cool water.

Clutches are small and eggs are relatively large for the

adult body size compared with other Xenopus species.

The tadpoles are also very large and take a very long time

to develop in comparison with other species. Moreover,

they are highly sensitive to dissolved solids. These char-

acters may reflect adaptation to a single volcanic lake

with a stable environment.

Acknowledgments. — The captive colony of Xenopus

longipes was exported under permit from the Cameroon

Ministry of Forestry & Wildlife (0928/PRBS/MINFOF/

SG/DFAP/SDVEF/SC and 0 1 93/CO/MINFOF/SG/

DFAP/SDVEF/SC), following prior consultation with

the Oku community, who also sanctioned access to their

lake. We thank Gonwouo Nono LeGrand, Ndifon David,

Roland Ndifon, Henry Kolem, and Robert Browne for

logistical assistance within Cameroon. Field work was

supported by the Royal Zoological Society of Scotland,

an Erasmus Darwin Barlow grant from the ZSL, a Small

Project Ecological Grant from the British Ecological So-

ciety, an Amphibian Conservation Fund grant from the

European Association of Zoos and Aquaria, and a Mo-

hammed bin Zayed Conservation grant. We also extend

our thanks to Brian Zimmerman, Alex Cliffe, and Ra-

chel Jones (Aquarium, ZSL London Zoo) for their as-

sistance in managing aquatic systems. We would like to

thank Andrew Cunningham (Institute of Zoology, ZSL)

for his support in initiating the foundation of the captive

colony of X. longipes. Finally, we would like to thank the

three referees, including Matthias Goetz (Durrell Wild-

life Conservation Trust) who waived anonymity, for their

comments on the manuscript.

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Christopher Michaels is a conservation biologist and herpetoculturist working in the Herpetology Section at the

Zoological Society of London. Chris gained his doctorate in amphibian husbandry from the University of Manches-

ter and his interests include using captive husbandry to inform and complement conservation in the field, as well as

direct involvement in field conservation. His primary taxonomic interest is in tailed amphibians and he is associated

with several field conservation projects on this group in Mexico, but also works on projects involving anurans and

caecilian amphibians.

Benjamin Tapley is a conservation biologist at the Zoological Society of London. Benjamin’s primary interests

include the conservation breeding and captive management of amphibians and he is chair of the British and Irish

Association of Zoos and Aquariums, Reptile and Amphibian Working Group, and co-chair of the Amphibian Cap-

tive Breeding Working Group. Benjamin is currently involved in several amphibian conservation programs and is

currently working with Chinese Giant Salamanders, Mountain Chicken Frogs, and Megophryid frogs in Vietnam.

Luke Harding is a senior keeper in the Herpetology Section of the Zoological Society of London, London Zoo.

He has extensive experience in the training of animals from police dogs to crocodiles and is particularly interested

in using these techniques to manage reptiles in zoo settings. He has a long-standing involvement in the Mountain

Chicken Frog conservation program, having lived on Dominica to take part in fieldwork on the species, as well as

being involved in a number of other field monitoring and conservation projects for reptiles and amphibians.

Zoe Bryant is a qualified keeper in the Herpetology Section of Zoological Society of London, London Zoo. She is

interested in the husbandry of a wide variety of amphibians and reptiles and has particularly focused on developing

enrichment and training methods for captive reptiles.

Sebastian Grant is a senior keeper at the Zoological Society of London, London Zoo with a broad interest in captive

husbandry, particularly of amphibians and reptiles, as well as invertebrates and fish.

George Sunter is a lead keeper at the Zoological Society of London, London Zoo, having worked there for the past

17 years. He is an experienced herpetologist and Komodo Dragon enthusiast and currently his interests include the

management of nocturnal mammals and naked mole rats in captivity.

Iri Gill is deputy team leader of the Herpetology Section of the Zoological Society of London, London Zoo. He has

particular interests in the management of of crocodilians and venomous reptiles in captivity. He is a member of the

IUCN Crocodilian Specialist Group, EAZA Venomous Snake RCP Co-ordinator, and BIAZA Venomous Reptile

Focus Group Co-ordinator. Iri is currently involved in Gharial conservation in Nepal and India.

Oscar Nyingchia is a member of the Oku community and has been working as a field technician for the CRAAC

Project since 2008. He has principally been involved with surveying Lake Oku’s frogs and water chemistry, as

well as working on community education events. He has taken part in surveys of caecilians and other lakes across

Cameroon.

Thomas Doherty-Bone is the manager of the Conservation Research and Action for Amphibians of Cameroon

(CRAAC) Project, hosted by the Royal Zoological Society of Scotland, which has been running since 2008. Work

primarily revolves around surveying amphibians and their habitats, training other amphibian conservationists, as-

sessing threats, engaging with stakeholders, and deploying conservation action. Work is especially directed at high

priority amphibian species and habitats such as Xenopus longipes and Lake Oku. Other work has involved assessing

amphibian chytrid fungus in Cameroon, caecilian baseline surveys, and crater lake surveys. Thomas has a B.Sc.

(Hons) in zoology from the University of Aberdeen and a Masters in advanced methods in taxonomy & biodiversity

from Imperial College London. Thomas is currently completing a doctorate on the impacts of alien invasive species

on freshwater biodiversity and ecosystem functioning at the University of Leeds.

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Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [General Section]: 111-119 (el 03).

Notes on the natural history and morphology of the Ning-

shan Lined Snake ( Stichophanes ningshaanensis Yuen, 1983;

Ophidia: Colubridae) and its distribution in the Shennongjia

National Nature Reserve, China

1A3 Kevin R. Messenger and 2 Yong Wang

1 Nanjing Forestry University, Nanjing, Jiangsu, CHINA 2 Alabama A&M University, Department of Biological and Environmental Sciences, Normal,

Alabama 35762, USA

Abstract. — The present study reports on the natural history of the Ningshan Lined Snake

( Stichophanes ningshaanensis) in the Shennongjia National Nature Reserve (NNR) in western Hubei

Province, China. Prior to this work, little was known about the natural history of this species due to

a paucity of specimens since the original description in 1983. Since its discovery, only the original

three specimens were known to science, all of which are now lost or destroyed. Over the course

of five summers, we observed 24 specimens within the Shennongjia NNR. We report on its natural

history, including seasonal activity, habitat and environmental preferences, breeding behavior, sexual

dimorphism, and incubation data for the eggs. We reiterate the morphological differences between

Stichophanes, its former genus Oligodon, and members of Pareatidae. In China, Stichophanes is

not protected under law due to the species being classified as “Data Deficient.” The species exhibits

sexual dimorphism and dichromatism, i.e., males are smaller than females and the sexes differ in

color. The species has unique breeding habits in mid-summer, and copulation occurs immediately

after oviposition of the females. The number of eggs per clutch ranges from eight to nine, and takes

64 days to hatch.

Key words. Oligodon, P areas, Asia, slug eaters, reproduction

Citation: Messenger KR, Wang Y. 2015. Notes on the natural history and morphology of the Ningshan Lined Snake ( Stichophanes ningshaanensis

yuen, 1983; Ophidia: Colubridae) and its distribution in the Shennongjia National Nature Reserve, China. Amphibian & Reptile Conservation 9(2)

[General Section]: 111-119 (e103).

Copyright: © 201 5 Messenger and Wang. This is an open-access article distributed under the terms of the Creative Commons Attribution-NonCom-

mercialNoDerivatives 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: 01 May 2015; Accepted: 11 September 2015; Published: 30 September 2015

Little is known about the Ningshan Lined Snake, Sticho-

phanes ningshaanensis Yuen, 1983. The species was

discovered in Ningshan County, Shaanxi Province,

China, in the southern part of the Qinling Mountains

(Yuen 1983) and was described based on three speci-

mens, which presently are all lost or destroyed (Wang et

al. 2014). No additional specimens were found until in

2006 a survey revealed 17 new specimens in Shennon-

gjia NNR, western Hubei Province (Yang et al. 2009).

In the original description, the species was assigned to

family Colubridae and placed in the genus Oligodon.

But few of its morphological characters match up to the

genus, additionally, none of the characters fit easily into

any other Asia genera. It was for this reason the species

was recently assigned to the new genus Stichophanes

(Wang, Messenger, Zhao, and Zhu 2014). The specific

epithet ningshaanensis is named for Ningshan County in

Shaanxi Province (note the double “aa” in the specific

epithet, which is not a typo and distinguishes Shaanxi

Province from Shanxi Province. In this circumstance, it

is used to correctly pronounce the extended vowel sound

of Shaanxi compared to Shanxi in the Mandarin lan-

guage), where the type specimen was found. The generic

epithet Stichophanes breaks down into stichos- (Greek),

meaning “line or row,” and -phanes (Greek), meaning

“appearing, conspicuous,” in reference to the dorsal and

lateral lines of the body.

Due to its elusive behavior, and the paucity of speci-

mens, little was known about the natural history of the

species. Aside from the initial description, the only other

Correspondence. Email: 3 kevinrmessenger@ gmail.com (Corresponding author).

September 2015 | Volume 9 | Number 2 | el 03

Amphib. Reptile Conserv.

Ill

Messenger and Wang

work of the species was an examination of the micro-

structure of the skin by Li and Liang (2007), which re-

vealed a canaliculated type structure. Additionally, prior

to the 2006 field work, this snake was among the rarest

of China’s species, with only three specimens known to

science at the time. Even though the species is locally

common at select locations within Shennongjia NNR,

this species could very well be a species of conserva-

tion interest if no additional populations are found in its

range. For these reasons, it is important to understand the

natural history of the species.

This study reports on the natural history, breeding

ecology, and distribution of the species as observed in

the Shennongjia NNR, and additional commentary on

aspects of its unique morphology, with special regard

toward the genus Oligodon and members of the family

Pareatidae.

Materials and Methods

Fourteen field sites were surveyed within the Shennon-

gjia NNR in western Hubei Province, China (Fig. 1). Ap-

proximately one week was spent at each field station. At

field stations, the primary surveying technique was walk-

ing habitat day and night, flipping natural cover objects

such as rocks and logs, and actively searching using vi-

sual and auditory stimuli. The first survey was conducted

in 2006 from May to September. A second survey was

completed in July 2008. A final intense survey was car-

ried out during the summer of 2011. Beginning in 2012,

only one month every summer was surveyed opportunis-

tically.

If the reserve museum did not have a specimen, then

an animal was preserved as a voucher. Subsequent indi-

viduals were photographed and released unless they dif-

fered from the previous specimens in such attributes as

pattern, gender, or age. Specimens were deposited with

the museum officials in Shennongjia, headquartered in

the town of Muyu. Specimens were later relocated to the

research lab at Guanmenshan within the reserve. Foca-

tions of finds were marked with GPS coordinates (accu-

racy < 3 m). Environmental data such as ambient temper-

ature, substrate temperature, habitat, and elevation were

recorded as well as precipitation and time of day or night.

Upon capture, each animal was sexed via probing and

measured snout-to-vent (SVF) and total length (TF) to

the closest 0.25 cm using a tape measure. Measurements

of eggs were taken with digital slide calipers to the clos-

est 0.01 mm. Dorsal scale rows were counted one head

length posterior to the head, at mid-body, and one head-

length distance anterior to the vent.

Fig. 1 . Locations of field stations and of Stichophanes ningshaanensis (n = 22) within the Shennongjia NNR.

September 2015 | Volume 9 | Number 2 | el 03

Amphib. Reptile Conserv.

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Natural history and morphology of Stichophanes ningshaanensis Yuen

Gans

Hubei

Legend

Shennongjia

# Stichophanes ningshaanensis

0 55 110

220 Kilometers

I 1 1 1 1 1 1 1

Fig. 2. Current known range of S. ningshaanensis ; type locality located in Shaanxi province.

Results

From 24-26 June 2006, we found six females and three

males near Pingqian. Three of the females were dead (one

beat to death by villagers, two were road kill); all males

were alive. In July 2008, we found eight live specimens,

and one dead specimen, all adults except one sub-adult.

Three new locations within the reserve were recorded:

a high mountain road near the town of Xiangshui, an-

other record between the towns of Banqiao and Pingq-

ian, and several individuals (n = 3) on the outskirts of

Muyu. In July 2011, we found an additional three speci-

mens in Pingqian and another on the mountain road near

Xiangshui (N31. 531231° E110. 113914°). In the years

2012-2014, no additional new locations were discov-

ered. In 2012, we had a single observation, representing

the earliest known observation of the species since its

discovery in 1983. In 2012, surveys were conducted in

late May, early June, and early August. In 2013, major

construction and development at the core site, Pingqian,

began, and only a single specimen was found in July,

ironically crossing habitat just bulldozed. The year 2014

represented the first year that surveys failed to find an in-

dividual, despite surveying during the active time of year

for the species. The development started in 2013 was

much more extensive in 2014 and much of the habitat in

Pingqian, where individuals had been found previously,

was completely destroyed or urbanized.

The finds in Shennongjia NNR represented a range

extension of 280 km to the southeast of the type locality,

and the first major population found since the species’

discovery in 1983 (Yang et al. 2009; Fig. 2).

Description Based on Specimens from

Shennongjia NNR

Dorsal scales are smooth, with counts of 13, 13, and 12

anteriorly, mid-body, and posteriorly, respectively. The

anal scale is divided. Head scales consist of two post-

oculars, one pre-ocular, no loreal scale, six supralabials

(3 rd and 4 th in contact with eye), five infralabials, one an-

terior temporal, and two (sometimes one) posterior tem-

porals. The rostral scale is smooth and not upturned or

protruding as is characteristic of the genus Oligodon. The

subcaudal scales are paired.

Males are olive-brown or olive-green, and females

are yellow-brown in color. In both sexes, the venter is

a cream-colored version of the dorsal background color.

Both sexes have a single row of dots on the lateral edges

of each ventral scale. These spots tend to fade posteri-

orly. Anteriorly, there are five distinct black lines imme-

diately posterior to the head. One line is along the spine

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113

Messenger and Wang

Fig. 3. Close up of the head, showing detail of the scales, and

illustrating the indistinct neck of the species. Photo by Kevin

R. Messenger.

Fig. 5. Typical habitat of Stichophanes ningshaanensis in the

Pingqian area pre-2013. Photo by Kevin R. Messenger

but quickly fades from black to brown to indistinct and

blending with the background coloration toward the tail.

Two pairs of lines are situated dorso-laterally and run the

length of the body with consistent boldness, often the in-

ferior edge of the line is brown and the superior edge re-

mains black. The final two pairs of lines are located ven-

tro -laterally, between or along the 1 st and 2 nd scale rows.

The iris is golden-brown in females and golden-yellow

in males. The head is indistinct from the neck (Fig. 3).

The largest individual found was a female measur-

ing 730 mm total length (TL) and 578 mm Snout-to-vent

length (SVL). The largest male measured 654 mm TL

and 495 mm SVL. Hatchlings (n = 17) averaged 150 mm

TL (SD + 4 mm) and 119 mm SVL (SD + 2 mm).

Natural History Notes

Specimens were found during the day and in the evening,

as late as 80 minutes after sunset. Twenty four specimens

were found: locals beat one specimen to death, two were

found dead on the road, two specimens were under rocks,

and the rest were actively moving about. Species obser-

vations were primarily terrestrial, but lacking specimens

outside the breeding season, species are suspected to be

primary fossorial and only move above ground during

Fig. 4. Seasonal activity of Stichophanes ningshaanensis ob-

servations (n = 24).

the breeding season. No specimens were found in an ar-

boreal setting. During the breeding season, several speci-

mens ( n = 8) were also found dead on the road. The aver-

age elevation of provenances of specimens was 1628 (SD

+ 126 m) (range 1,550-2,200 m). May surveys failed to

find specimens. The earliest observation date was a fe-

male found on 07 June 2012, but in general, late June and

early July were best times finding species as this is the

breeding season and snakes were actively moving above

surface (Fig. 4). The latest documented date occurrence

was on 20 July 2008. Surveys in August and September

failed to detect any specimens. The species was usual-

ly in proximity to water, i.e., within -300 m of a water

source, and was often encountered actively moving after

rainstorms. The species was active on overcast and cool

days with temperatures ranging 20-24 °C.

With respect to habitat, between 1,500-2,600 m el-

evation, the habitat is classified as temperate deciduous

broadleaf coniferous forest including Farges’ Fir and For-

tune’s Rhododendron (Zhao et al. 2005). Average annual

temperature of locations where individuals were found

were 16.2 °C (range: 15.6-16.7). Average annual pre-

cipitation of locations where individuals were found was

222.85 mm (range: 209-235). Individuals were found

in ephemeral stream beds, in short grasses, commonly

found on the outskirts of agriculture land, and in habitat

adjacent to permanent streams (Fig. 5). Individuals were

never far from a source of water.

Stichophanes ningshaanensis shares its habitat with

the following species: Snakes: Achalinus spinalis, Azem-

iops feae, Dinodon [ =Lycodon ] rufozonatum, Elaphe

carinata, Lycodon cf. fasciatus, Orthriophis taeniurus,

Protobothrops jerdonii, Pseudoxenodon macrops, Rhab-

dophis nuchalis, Sibynophis chinensis ; Lizards: Ples-

tiodon capito, P elegans, Scincella modesta, Spheno-

morphus indicus, Takydromus septentrionalis', Anurans:

Amolops chunganensis, A. granulosus, Bufo cf. andrew-

si, Megophrys wushanensis, Odorrana margaratae, P aa

quadranus, Rana chensinensis; Salamanders: Lina shihi

and Ranodon tsinpaensis (pers. obs.).

When confronted, the species was reluctant to bite.

No amount of provocation elicited a defensive bite.

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Natural history and morphology of Stichophanes ningshaanensis Yuen

Fig. 6. Tail-curling defensive behavior characteristic of Oligodon : O. formosanus (left), O. ornatus (right), and enlarged rostral

scale. Photos by Kevin R. Messenger.

Fig. 7. Courtship behavior by the male, rubbing his chin along

the female, observed on 28 June 2006. Photo by Kevin R. Mes-

senger.

Fig. 8. Courtship behavior by the male observed on 28 June

2006, and illustration of dichromatic differences. Photo by

Kevin R. Messenger.

Many members of Oligodon effectively use their unique

teeth when restrained and harassed and will bite readily.

Stichophanes thrashes about and readily produces musk

but does not display the characteristic tail-coiling known

to some other species within the genus Oligodon (Ses-

hadri 2014; Fig. 6).

Notes on Reproduction

The species exhibits strong sexual dimorphism, not only

in size, but also in color (sexual dichromatism), an un-

common trait among snakes (Boulenger 1913; Jacob

and Altenbach 1977; Shine and Madsen 1992). There

are only a handful of other species that have been re-

ported to exhibit sexual dichromatism, such as Crotalus

lepidus klauberi, in which males have a greenish hue and

females have a purple hue (Jacob and Altenbach 1977).

Shine and Madsen (1992) noted dichromatism in the ge-

nus Vipera. In S. ningshaanensis, males are smaller than

females. Females are yellowish-brown, while males are

olive-brown or olive-green. Males also have a longer tail

than females. In males, the tail is 24-27% of the total

body length, whereas the value for females is 21% (Wang

etal. 2014).

Despite the fact snakes were found in mid-summer

(late June), males attempted to mate with gravid females.

Courtship behavior was observed on multiple occasions.

It consisted of a male rubbing his chin along the length

of a female and positioning his cloaca next to hers (Figs.

7, 8). No copulation was observed with these gravid fe-

males before oviposition. There is no documentation of

other colubrids trying to copulate with gravid females

nearly full term, although this is commonly observed in

crotalids (Duvall et al. 1992) in which mating and birth-

ing occur in the same season, typically fall. Stichophanes

ningshaanensis is similar. Immediately after females laid

eggs in late summer, males commence with copulation.

A clutch of eight and nine eggs was recorded from two

females on 29 and 30 June 2006 (Fig. 9). The time span

between successive eggs was 15 minutes, and each egg

took two minutes to exit the cloaca. In the first female,

after oviposition, a male immediately courted her and

successfully copulated (Fig. 10).

The eggs measured 26.98 mm long and 9.52 mm wide.

All 17 eggs were placed in a plastic container and cov-

ered with a damp paper towel. They were kept at room

temperature (generally 24 °C but reaching a maximum

of 29 °C). After 62 days, the first eggs started to pip. By

64 days all 17 eggs had pipped, and the young began to

emerge from the eggs (Fig. 11).

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Messenger and Wang

Fig. 9. Nine eggs from a female measuring 533 mm SVL and

673 mm TL on 30 June 2006. Photo by Kevin R. Messenger.

Fig. 11. Hatching and emergence after 64 days of incubation.

Photo by Kevin R. Messenger.

Fig. 13. Comparison of typical head scales and head shapes of

Oligodon (top); 8 supralabials, 4 and 5 in contact with eye, 2

pre-oculars, 2 post-oculars, 1+2 temporals, 1 loreal, enlarged,

upturned rostral scale, to the head scales of Stichophanes (bot-

tom); 6 supralabials, 3 and 4 in contact with eye, 1 pre-ocular, 2

post-oculars, 1+2 temporals, no loreal, blunt rostral scale. Pho-

tos by Kevin R. Messenger

Fig. 10. Copulation on 30 June 2006, post oviposition by the

female. Photo by Kevin R. Messenger

1 mm

Fig. 12. Comparison of right maxillae; Oligodon on the top,

with the characteristic kukri-shaped rear teeth which it uses to

saw into eggs, distinguished from the anterior teeth (from Cole-

man et al. 1993), Stichophanes on the bottom, anterior teeth

all the same, and a lack of rear- specialized teeth (from Wang

et al. 2014).

Fig. 14. Photograph of P areas formosensis (van Denburgh

1909) from Taiwan, illustrating the concave tongue notch

opening that is typical of Pareas members. Photo by Daniel

Rosenberg.

September 2015 | Volume 9 | Number 2 | el 03

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Natural history and morphology of Stichophanes ningshaanensis Yuen

Comparison with Species of Oligodon

The genus Oligodon Fitzinger 1826 is a very broadly

characterized genus. There are approximately 74 species

within the genus as of 2013 and as such bring a wide

variety of characteristics and diversity (Green 2010; Vas-

silieva et al. 2013). Five robust characters tend to apply

to most species (Green 2010). These are:

1) Presence of unique posterior maxillary teeth, ap-

pearing in shape to Ghurka kukri knives, for which the

genus gets its common name, “Kukri Snake.”

2) Large, slightly upturned rostral shield, protruding

when viewed from above.

3) Many species possess a distinct dark chevron mark

on the nape and a stripe across the anterior part of the

head and down over/through the eye.

4) Majority of species have blotched and/or reticulate

pattern, usually not prominently striped.

5) Most species possess a loreal scale.

Stichophanes ningshaanensis differs on several lev-

els and conflicts with each of these five robust charac-

ters: in addition to the defensive behavioral differences

mentioned previously (i.e., lack of tail curling, refusing

to bite defensively), S. ningshaanensis does not possess

the distinctive rear teeth for which Oligodon was named

(Fig. 12). Most Oligodon use these specialized teeth to

slice or “saw” into reptile eggs (Coleman et al. 1993).

They use their upturned snout to dig up eggs, similar to

species in the North American genera Cemophora and

Phyllorhynchus. Once an egg is opened, they insert their

head inside the egg to consume the contents. Sticho-

phanes ningshaanensis lacks this upturned rostral shield,

instead, having a very blunt and squared-off head (Fig.

13). Additionally, the species does not prey on eggs or

Fig. 15. Left: underside view of Pareas vindumi (from Vogel

2015), showing the lack of a mental groove due to asymmetri-

cal chin shields. Right: underside view of Stichophanes (from

Wang et al. 2014), showing symmetrical chin shields and the

presence of a mental groove.

any of the known prey ingested by other Oligodon spe-

cies but rather eats snails and slugs exclusively (Wang

et al. 2014). The species lacks chevron markings on the

nape and lacks a stripe across the anterior part of the head

or through the eye. The species is distinctly striped and

not blotched, and lastly, all specimens lack a loreal scale.

From an internal perspective, the hemipene morphology

does not conform to that of Oligodon. From a morpho-

logical and behavioral standpoint, these key differences

give credence to the species not belonging to the genus

Oligodon.

The next most likely genus for the species to be placed

in, from a morphological and dietary standpoint, is Par-

eas, the Asian snail eaters.

Comparison with Asian Snail and Slug Eat-

ing Species

Due to its shortened, square head, as well as its exclu-

sive diet of gastropods, it seems likely that Stichophanes

could be closely related to members of the Asian snail

and slug eaters: Pareatidae. Currently only three genera

are known in Pareatidae. These are:

Aplopeltura : a genus containing a single species, A.

boa, the Blunt-headed Slug Eating Snake. This genus

is arboreal. The head is very distinct from the neck.

This genus is located outside of China.

Asthenodipsas : a genus containing five species (Lore-

do et al. 2013). Members are characterized by a large

head, distinct neck, lacking a mental groove, very

large eyes, and an arboreal lifestyle. The mouth pos-

sesses a slotted opening that facilitates ingestion of

snails and slugs. All members of the genus are located

outside of China.

Pareas : a genus containing 13 species (You et al.

2015; Vogel 2015). Members are characterized by a

blunt snout, lacking a mental groove, distinct neck,

and no teeth on the anterior part of the maxilla (Guo

and Deng 2009). The tongue notch possesses a con-

cave opening to facilitate the ingestion of snails and

slugs (Fig. 14). The majority of the species are found

in China.

Stichophanes ningshaanensis does not fit into any of

these genera. The species is strictly terrestrial and fos-

sorial, there is little to no distinction between the head

and neck, it possesses teeth on the anterior part of the

maxilla (Fig. 12), it possesses a mental groove (Fig. 15),

and does not have a slotted notch on the mouth. From

a morphological, dietary, and behavioral standpoint, the

species does not fit into any known Asian genus and is

quite unique, not only in appearance but also in its ecol-

ogy. Genetic work by Wang et al. (2014) further support-

ed what the morphological data suggested and could not

September 2015 | Volume 9 | Number 2 | el 03

Amphib. Reptile Conserv.

117

Messenger and Wang

Fig. 16. Downtown Pingqian. Stichophanes ningshaanensis

was commonly found crossing this road and in the habitat ad-

jacent to the road. Picture taken June 2011. Photo by Kevin R.

Messenger.

place the species in any known genus, at which point, a

new genus was erected, Stichophanes.

Discussion

The purpose of this paper was to provide insight into

this rarely observed and studied species. The population

in the Shennongjia NNR has provided opportunities to

observe several aspects of the species’ natural history,

from activity periods, to seasonal differences, to court-

ship, breeding, and incubation of eggs. The species has

a unique reproductive strategy, which is not documented

among other species of colubrids, or is, at the very least,

quite uncommon.

Prior to 2013, the species was locally abundant in

Shennongjia NNR, and specifically in Pingqian, and was

among the more common and predictable species when

in its habitat. The changing habitat due to development of

the Pingquin village may be a turning point for the spe-

cies in the area, for the worse (Figs. 16, 17). Future inves-

tigations in this area will hopefully yield knowledge on

the urban tolerance (or intolerance) of the species. Inves-

tigations in 2014, despite being done during the height of

the breeding season, failed to turn up a single specimen.

Currently, the species is not under any special protection,

currently classified as “Data Deficient.” It is hoped this

paper will bring us closer to understanding the species

and its potential distribution, and this information will

reduce the deficiency of data for this species.

Aside from the natural history aspects, another goal

was to further illustrate the morphological distinction

of the species from closely aligned genera, such as its

original placement in Oligodon and its next most likely

genus, Pareas — these differences were briefly touched

on in Wang et al. (2014), and deserved greater scrutiny.

Acknowledgments. — We want to especially thank the

Shennongjia National Nature Reserve for all of the sup-

port they have provided us over the years, specifically

Fig. 17. View of Downtown Pingqian in June 2014, covering

same view (different angle) as Figure 16. Photo by Kevin R.

Messenger.

my friends Dong Xue, Ming Wong, Linsen Yang, and

Jianhuan Yang. We thank Shennongjia NNR, Alabama

A&M University, Nanjing Forestry University, and the

National Science Foundation for funding, either indirect-

ly or directly, over the years.

Literature Cited

Boulenger GA. 1913. On the geographic races of Vipera

ammodytes. Annuals and Magazine of Natural His-

tory 8(11): 283-287.

Coleman K, Rothfuss LA, Ota H, Kardong KV. 1993.

Kinematics of egg-eating by the specialized Taiwan

snake Oligodon formosanus (Colubridae). Journal of

Herpetology 27(3): 320-327 .

Duvall D, Arnold SJ, Schuett GW. 1992. Pitviper mat-

ing systems: Ecological potential sexual selection,

and microevolution. Pp. 321-336 In: Biology of the

Pitvipers. Editors, Campbell JA, Brodie ED Jr. Selva

Publishing, Arlington, Texas, USA. 467 p.

Green M. 2010. Molecular phylogeny of the snake genus

Oligodon (Serpentes: Colubridae), with an annotated

checklist and key. M.Sc. Thesis. University of Toron-

to, Toronto, Ontario, Canada. 169 p.

Guo K, Deng X. 2009. A new species of Pareas (Ser-

pentes: Colubridae: Pareatinae) from the Gaoligong

Mountains, southwestern China. Zootaxa (2009):

53-60.

Jacob JS, Altenbach JS. 1977. Sexual dimorphism in

Crotalus lepidus klauberi Gloyd (Reptilia, Serpentes,

Viperidae). Journal of Herpetology 11(1): 81-84.

Li L, Liang G. 2007. Microdermatoglyphis patterns of

Oligodon ningshaanensis. Journal of Xian University

of Arts and Science (Natural Science Edition) 10(3):

66-68. [In Chinese with English abstract].

Loredo AI, Wood PL Jr, Quah ESH, Anuar S, Greer LF,

Ahmad N, Grismer LL. 2013. Cryptic speciation

within Asthenodipsas vertebralis (Boulenger, 1900)

(Squamata: Pareatidae), the description of a new spe-

cies from Peninsular Malaysia, and the resurrection of

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A. tropidonotus (Lidth de Jude, 1923) from Sumatra:

An integrative taxonomic analysis. Zootaxa 3664(4):

505-524.

Seshadri KS. 2014. Display behaviour in response to a

perceived threat by Indian short-headed kukri snake

Oligodon brevicaudus (Gunther, 1862) (Squamata:

Colubridae). Herpetology Notes 7(2014): 447-450.

Shine R, Madsen T. 1992. Sexual dichromatism in snakes

of the genus Vipera: A review and a new evolutionary

hypothesis. Journal of Herpetology 28(1): 114-117.

Vassilieva AB, Geissler P, Galoyan EA., Poyarkov Jr

NA, Van Devender W, Bohme W. 2013. Anew species

of Kukri Snake ( Oligodon Fitzinger, 1826; Squamata:

Colubridae) from the Cat Tien National Park, south-

ern Vietnam. Zootaxa 3702(3): 233-246.

Vogel G. 2015. Anew montane species in the genus P ar-

eas Wagler, 1830 (Squamata: Pareatidae) from north-

ern Myanmar. Taprobanica 7(1): 1-7.

Wang X, Messenger K, Zhao E, Zhu C. 2014. Reclas-

sification of Oligodon ningshaanensis Yuan, 1983

(Ophidia: Colubridae) into a new genus, Stichophanes

gen. nov. with description on its malacophagous be-

havior. Asian Herpetological Research 5(3): 137-149.

Yang L, Messenger K, Liao MY. 2009. Reptile and am-

phibian diversity of Shennongjia National Nature

Preserve. Sichuan Journal of Zoology 2009 28(2):

286-291. [In Chinese with English abstract].

You CW, Poyarkov Jr NA, Lin SM. 2015. Diversity of

the snail-eating snakes Pareas (Serpentes, Pareatidae)

from Taiwan. Zoologica Scripta 44(4): 349-361.

Yuen H. 1983. Anew species of the genus Oligodon from

Shaanxi Province, China. Acta Herpetologica Sinica

2: 65-67. [In Chinese with English abstract].

Zhao CM, Chen WL, Tian ZQ, Xie ZQ. 2005. Altitudi-

nal pattern of plant species diversity in Shennongjia

Mountains, central China. Journal of Integrative

Plant Biology 47(12): 1,431-1,449.

Kevin Messenger is a graduate student pursuing dual Ph.D.’s, one from Alabama A&M University under

Professor Yong Wang in Wildlife Conservation, and another from Nanjing Forestry University in Nanjing,

China in Zoology. He received his B.S. (zoology) from North Carolina State University under Dr. Harold

Heatwole, followed by a M.S. (biology) from Marshall University under Dr. Thomas Pauley. Kevin’s primary

interest is in behavior, conservation, ecology, and natural history. He is especially interested in snakes, the

herpetofauna of the southeastern US, and Asian herpetology.

Yong Wang is a Professor of Wildlife Biology and Biometry at the Department of Biological and Environ-

mental Science, Alabama A&M University. He has a B.S degree from Shanghai Normal University of China

and a doctoral degree from University of Southern Mississippi. He has worked as a post-doctoral wildlife

biologist at the Rocky Mountain Research Station of the USDA Forest Service. His current research interests

include the relationships between forest management practices and wildlife communities including avian and

herpetofauna, stopover ecology of songbird migrants, and modeling spatial and temporal patterns of forest

and wildlife community using statistical, geographic information systems, and remote sensing technology.

September 2015 | Volume 9 | Number 2 | el 03

Amphib. Reptile Conserv.

119

Official journal website:

amphibian-reptile-conservation.org

Amphibian & Reptile Conservation

9(2) [General Section]: 120-125 (el 04).

Short Communication

On the distribution, taxonomy, and natural history of the

Indian Smooth Snake, Coronella brachyura (Gunther, 1866)

14 Harshil Patel, 25 Raju Vyas, and 3 ’ 6 Shantilal K. Tank

1 Department of Biosciences, Veer Narmad South Gujarat University, Surat-395007, Gujarat, INDIA 2 505 Krishnadeep Tower, Mission Road,

Fatehgunj, Vadodara, Gujarat, INDIA 3 Department of Biosciences, Veer Narmad South Gujarat University, Surat-395007, Gujarat, INDIA

Abstract . — The Indian Smooth Snake Coronella brachyura is one of the least studied endemic species

of snake from India with regard to distribution, taxonomy, and natural history. In the present study,

we verified literature, museum specimens and distributional records which enabled us to correct

erroneous reports and map the distribution of this species. Additionally, we provide information on

taxonomy, morphology, microhabitat, and behavior of the species based on three live specimens

and voucher specimens in the collection of the Bombay Natural History Society, Mumbai.

Key words. Colubridae, endemic, India, rare, morphology, scalation

Citation: Patel H, Vyas R, Tank SK. 201 5. On the distribution, taxonomy, and natural history of the Indian Smooth Snake, Coronella brachyura

(Gunther, 1866). Amphibian & Reptile Conservation 9(2) [General Section]: 120-125 (el 04).

NoDerivatives 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: 23 April 201 5; Accepted: 1 3 June 2015; Published: 03 October 201 5

Members of colubrid snake genus Coronella Laurenti,

1768 are among the least studied snakes across the world.

The genus is represented by three species namely C.

austriaca Laurenti, 1768, C. girondica (Daudin, 1803),

and C. brachyura (Gunther, 1866) (Wallach et al. 2014;

Uetz and Hosek 2015). The former two species are dis-

tributed in western Palaearctic (from southern Norway in

the north to northern Algeria in the south; Portugal in the

west to northern Iran in the east) and the latter, endemic

to India (Wallach et al. 2014; Uetz and Hosek 2015).

Gunther (1866) described this species from Poona (Pune)

in the Indian state of Maharashtra. Subsequently, the spe-

cies was reported from several localities based on which

the distribution range of the species was considered to

be restricted to three states in the western part of India

namely; Maharashtra, Gujarat, and Madhya Pradesh.

Reported localities from Maharashtra state are: “Wun,

S. E. Berar” (now Wani, Yavatmal district) referred by

Blanford (1870), Anderson (1871), Theobald (1876),

Boulenger (1890), Sclater (1891), and Wall (1923);

Chink Hill and Kurduwadi in Solapur district (Lindberg

1932); Visapur, Ahmednagar district (Gharpurey 1935);

Marole (Andheri) — Salsette Islands, Mumbai (Abdulali

1935); Nashik (Mistry 2005); Melghat, Amravati dis-

trict (Nande and Deshmukh 2007); Latur, Latur district

(Kamble 2010); Khed, Pune district (Ghadage et al.

2013), and Jalna (Z. Mirza, pers. comm.). Furthermore,

the species was reported from Gujarat state (Vyas and

Patel 2007) and Ujjain, Madhya Pradesh state (Ingle and

Sarsavan 2011). Sarasin (1910) referred to this species

but did not provide any specific localities. Whitaker and

Captain (2004) gave the range of this species as “few lo-

calities in Maharashtra.” According to Smith (1943) the

range of this species is “Northern India. Poona district

and Visapur, near Bombay; S. E. Berar,” however, it is

unclear why he included “Northern India” in its range.

In the recent past, we came across three live individuals

of C. brachyura from Surat, Gujarat. Based on museum

specimens, published literature, and additional data from

live individuals we provide additional morphological and

distributional data, as well as natural history observations

for this poorly known species.

Correspondence. Email: 4 har shilpatell21@gmail.com (Corresponding author) 5 razoovyas@hotmail.com 6 drtanksk@ gmail.com

Amphib. Reptile Conserv.

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Patel et al.

Material and Methods

Three live specimens (two females and one male: field

number assigned as: NCS 01-03) rescued by snake res-

cuers and brought to us (they were photographed, ex-

amined, and released at the same locality within a few

days), and seven specimens catalogued in the museum

of Bombay Natural History Society (BNHS), Mumbai as

Coronella brachyura from six localities were also exam-

ined. The pholidosis and morphometric data of museum

specimens and live specimens are given in Table 1 .

Ventral scales were counted following the method

proposed by Dowling (1951). Head measurements of

voucher specimens were measured with a digital calli-

per to the nearest 0.01 mm and other body measurements

were recorded with string and a ruler to the nearest mm.

Descriptions and mensural characters were compared

with available literature (Smith 1942; Mistry 2005; Vyas

and Patel 2007). The number of dorsal scale rows were

counted at approximately one head length behind the

head, midbody, and one head length before the vent, re-

spectively. Subcaudal counts reported here do not include

the terminal scute. The supralabials touching the eye are

given in brackets after the number of supralabials. Val-

ues for symmetric head characters are given in right/left

order. Abbreviations used to describe scalation and other

comparable characters are: V, ventrals; SC, subcaudals;

D, dorsal rows; SL, supralabials; L, loreal; PrO, preoc-

ular; PO, postocular; T, temporal; IL, infralabial; SVL,

snout-ventral length; TaL, tail length; TL, total length;

HL, head length; and HW, head width.

Results

Morphology and coloration : Head short, comprising

2.3% of total length; longer than wide (HL/HW ratio:

1.55); slightly distinct from neck; eyes circular with

round pupil; nostrils large; body circular. Dorsal color

of live individuals was olive brown, with indistinct light

variegation on head and forebody (Fig. 1); labials pale

olive; lateral scale rows dark brown, forming indistinct

lateral stripe on each side from nostril to tail, which is

prominent between nostril to eye; underside cream white.

Lepidosis : Dorsal scale rows (DSR) smooth, in most

specimens 23:23:19 (23:23:21 in BNHS 3407; 23:23:17

in NCS 2); with single apical pit on the posterior margin.

Ventrals 209-237 (maximum 224 fide. Smith 1943); anal

undivided; subcaudals 43-54 (46-53 fide. Smith 1943);

rostral wider than high, scarcely visible from above; 2 in-

ternasals, wider than long; 2 prefrontals, as long as wide,

longer than the internasals; frontal bell shaped, slightly

longer than wide; parietals longer than wide, slightly lon-

ger than frontal; 1 loreal, as long as high, rarely longer

than high; 1 preocular reaching top of head; 2 postocu-

lars; 2 anterior temporal scales; 2, rarely 1 posterior tem-

poral scale(s); 8, sometimes 9 (8 fide. Smith 1943) supra-

Fig. 1 . Dorsal aspect of Coronella brachyura in life, from Surat, Gujarat, India.

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Distribution, taxonomy, and natural history of Coronella brachyura

Fig. 2. Lateral aspect of Coronella brachyura (NCS 2); a, left

side showing 8 supralabials and 5 Ul supralabial partly divided;

b, right side showing 9 supralabials, 4— 6 lh touching eye.

labials, the 4 th and 5 th , sometimes 5 th and 6 th and rarely 4 th

to 6 th (4 th and 5 {h fide. Smith 1943) touch the eye (Fig. 2);

9-11 infralabials.

Distribution-. The present study and published records

(Gunther 1866; Blanford 1870; Anderson 1871; Theo-

bald 1876; Boulenger 1890; Scarlet 1891; Wall 1923;

Lindberg 1932; Gharpurey 1935; Smith 1943; Whitaker

and Captain 2004; Mistry 2005; Vyas and Patel 2007;

Fig. 3. BNHS 794, collected by Abdulali (1935) from Mumbai,

India.

Nande and Deshmukh 2007; Ingle and Sarsavan 2011;

Ghadage et al. 2013) shows that the species is narrowly

distributed in western India (Table 2).

Four museum specimens BNHS 793, 796, 798, and

3407 were examined. Two specimens BNHS 795 and

797 were damaged; therefore unable to examine for

pholidosis and morphometric data. The specimen BNHS

794 (Fig. 3) from Marol, Mumbai collected and reported

as C. brachyura by Abdulali (1935); was re-examined by

the senior author. It had 23 scale rows at mid body; 217

ventrals; anal scale damaged; 96+ subcaudals, divided;

8 supralabials; 1 presubocular; 2+3 temporals; and mea-

sured 285 mm total length. All these characters clearly

matched with Argyrogena fasciolata (Shaw, 1802). The

coloration of this specimen has faded likely due to long

Table 1. Scale counts, measurements (mm), and collection details for specimens of Coronella brachyura.

Specimen No

BNHS 793

BNHS 796

BNHS 798

BNHS 3407

NCS 1

NCS 2

NCS 3

Locality

Visapur,

Ahmednagar,

Maharashtra

Talegaon,

Pune, Maha-

rashtra

Bhopal, Mad-

hya Pradesh

Piplod, Surat,

Gujarat

Surat, Gujarat

Piplod, Surat,

Gujarat

Piplod, Surat,

Gujarat

Date

—

October 27,

1956

July 1945

March 2006

December 5,

2012

February 12,

2014

Feburary 17,

2014

TL

375

523

507

495

410

620

560

SVL

322

447

443

445

360

552

480

TaL

53

66

64

50

68

80

D

23:23:19

23:23:21

23:23:19

23:23:17

23:23:19

V

221

216

209

237

223

220

A

Undivided

SC

45

49

47

45

47

43

54

SL

9(5,6)/8(4,5)

8(4,5)/8(4,5)

9(5,6)/8(4,5)

9(4 to 6)/8(4,5)

8(4,5)/8(4,5)

L

1/1

IL

10/10

9/10

10/10

9/9

10/10

11/11

9/9

PreO

1/1

PO

2/2

T

2 + 212+2

2+ 1/2+1

2 + 212+2

Sex

ND=Not Deter-

mined

ND

Female

Male

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Patel et al.

Fig. 4. Map showing distribution range of Coronella brachyura (For all the localities: 1-13, reference Table 2).

25.00

23.00

21.00

19.00

term preservation. However, it shows remains of 27+ ves-

tigial whitish bands in the forebody which became paler

in posterior half and became indistinguishable — which is

found in juveniles of A. fasciolata. Based on our obser-

vations we here conclude that the specimen cited by Ab-

dulali (1935) is conspecific with A. fasciolata and is an

erroneous report from Mumbai, and should be removed

from the known distribution range of C. brachyura.

Vyas and Patel (2007) collected C. brachyura from

Surat, Gujarat and in the same publication they also pre-

sented two more localities (Ahmedabad and Bhavnagar)

from Gujarat based on photographs of a striped snake

which they attributed to C. brachyura. However, spec-

imens were not available to the authors and hence the

exact identity of specimens from these two localities re-

mains in question. Fresh specimens are needed to con-

firm the presence of C. brachyura from these localities.

Habit, habitat, and natural history. Coronella brachyura

are found in plains and hillocks; majority of known local-

ities are situated around 500 m a.s.l. The species appears

to occur in a wide range of habitats from arid scrub lands

to dry deciduous forests; they are also found in human

habitations. Two live individuals (NCS 02 and 03) were

found in a water body near a newly developing urban

area; NCS 01 was found near a water body. The speci-

mens were active during day time and did not show any

aggression when handled. Live individuals were kept for

a few days; juveniles of Hemidactylus sp. were offered

food but none accepted. However, some authors report-

ed that the species feeds on juvenile geckos in captivity

(Whitaker and Captain 2004; Ingle and Sarsavan 2011).

Discussion

Distribution: Our observations coupled with published

information of the species shows this endemic species is

widely distributed encompassing a geographical area of

2,80,000 sq. km across three Indian states, namely Ma-

harashtra, Madhya Pradesh (west), and Gujarat (south),

only (Fig. 4). This has a very si mil ar distribution range

recorded in another endemic colubrid snake, Psammo-

phis longifrons (Vyas and Patel 2013).

Conservation status: Coronella brachyura is legally pro-

tected as a Schedule IV species under the Indian Wildlife

Protection Act of 1972 and categorized as Least Concern

by the IUCN Red List of Threatened Species (Sriniva-

sulu et al. 2013). During the study no specific threats to

the species were observed, except the general threats to

the reptilian fauna as reported by Vyas (2007), includ-

ing expansion of urbanization, agricultural lands, habitat

loss, and habitat alteration, and large numbers of snakes

killed by laymen due to fear.

Taxonomy: The genus Coronella has shown to be para-

phyletic based on molecular data from western Palaeart-

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October 2015 I Volume 9 I Number 2 | el 04

Distribution, taxonomy, and natural history of Coronella brachyura

Table 2. List of localities for Coronella brachyura based on new collections or observations 1 , examined specimens 2 , literature or

database records, 3 and photographic records. 4

No

Locality

Coordinates

Elevation, m a.s.l.

District

State

1

Pune 2,3

18.31°N 73.51°E

561

Pune

Maharashtra

2

Talegaon 2 - 3

18.72°N 73.68°E

670

Pune

Maharashtra

3

Wani 3

20.03°N 78.57°E

228

Yavatmal

Maharashtra

4

Kurduwadi 23

18.08°N 75.43°E

502

Solapur

Maharashtra

5

Visapur 2 - 3

18.48°N 74.35°E

620

Ahmednagar

Maharashtra

6

Nashik 3

20.00°N 73.78°E

600

Nashik

Maharashtra

7

Melghat 3

21.26°N 77.11°E

575

Amravati

Maharashtra

8

Latur 3

18.23°N 76.36°E

620

Latur

Maharashtra

9

Khed 3

18.56°N 73.43°E

715

Pune

Maharashtra

10

Bhopal 2

23.15°N 77.25°E

527

Bhopal

Madhya Pradesh

11

Ujjain 3

23.10°N 75.47°E

511

Ujjain

Madhya Pradesh

12

Surat 1 ’ 23

21.18°N 72.83°E

13

Surat

Gujarat

13

Jalna 4

19.83°N 75.88°E

489

Jalna

Maharashtra

ic species by recent workers (Pyron et al. 2010, 2013;

Utiger et al. 2002). Recently, Hoser (2012) removed C.

brachyura from the genus Coronella and allocated it to

the genus Wallophis; it was earlier suggested by Werner

(1929). In doing so, Hoser (2012) did not provide any

valid taxonomic characters to support partitioning the

genus Coronella. Coronella brachyura differs from its

congeners by the higher number of scale rows at mid

body (23 vs. 21 in C. girondica and 19 in C. austriacaf

by the higher number of supralabials (8-9 vs. 7 in C. aus-

triaca and 8 in C. girondica). However, the status of In-

dian taxa remains unresolved as there is no comparative

study on the morphology or molecular data of Coronella

with other colubrid genera. We believe for now, the In-

dian species should be considered as a member of the

genus Coronella. Future studies involving detailed com-

parison of the genus Coronella , with the aid of molecular

techniques, will be essential for the correct allocation of

Indian species.

Acknowledgments. — We are thankful to Bhautik

Dudhatra and Bhavin Mistri for sharing information and

allowing us to examine the snake specimens. Rahul Khot

(BNHS) kindly facilitated examining material under his

care. Vithoba Hegde, Priya Warekar, Pinal Patel, and

Saunak Pal provided valuable assistance at the BNHS,

Mumbai. Viral Mistry and Frank Tillack provided some

important literature. Zeeshan Mirza and Deepak Veerap-

pan are thanked for valuable comments for which the

manuscript benefited. HP was supported by a INSPIRE

Fellowship (IF 130480) from the Department of Science

and Technology (DST), New Delhi, India.

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Harshil Patel is a young herpetologist, currently pursuing a Ph.D. in the Department of Biosciences, Veer

Narmad South Gujarat University, Surat, India. He is interested in the systematics and distribution of colubrid

snakes and geckos of genus Hemidactylus from India. His doctoral work and study is on the “Taxonomic

study of herpetofauna of Northern Western Ghats of Gujarat State.”

Raju Vyas is an enthusiastic herpetologist, presently working at the Sayaji Baug Zoo, Vadodara- Gujarat,

India as a Zoo Biologist. After his post graduate education in Zoology, he pursued a doctorate in philosophy,

his research dissertation titled “Snakes of Gujarat State,” from Bhavnagar University, Gujarat-India (1995).

For almost two decades, he has extensively explored the natural heritage of Gujarat state and its territo-

rial extensions contributing significantly toward the enrichment of base line data on amphibian and reptiles

of the state. Apart from his exposure to ex-situ conservation, he’s active in breeding many native reptilian

species. Presently, his activities include conservation of urban wildlife, especially Mugger Crocodiles, and

has an affiliation with the Vishwamitri River Project, Vadodara Municipal Corporation, Vadodara. Raju has

been monitoring the urban crocodile population in Gujarat and has published several reports about the same.

Lastly, Raju is optimistically promoting mitigation measures for man-animal conflicts locally and nationally.

Shantilal K. Tank is a Professor at the Department of Biosciences, Veer Narmad South Gujarat University,

Surat. For the past two decades Dr. Tank has worked in the fields of environmental toxicology, bioremedia-

tion, and ichthyology. Recently, he works in biodiversity documentation and conservation.

Amphib. Reptile Conserv.

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October 2015 | Volume 9 I Number 2 | el 04

CONTENTS

Special Section

N’Goran Germain Kouame, Caleb Ofori-Boateng, Gilbert Baase Adum, Germain Gourene, Mark-

Oliver Rodel — A The anuran fauna of a West African urban area 1

Ivan Ineich, Matthew Lebreton, Nathalie Lhermitte-Vallarino, Laurent Chirio — The reptiles of the

summits of Mont Oku and the Bamenda Highlands, Cameroon 15

Jean-Fran^ois Trape and Youssouph Mane — The snakes of Niger 39

Michael F. Barej, Tilo Pfalzgraff, Mareike Hirschfeld, H. Christoph Liedtke, Johannes Penner, Nono

L. Gonwouo, Matthias Dahmen, Franziska Grozinger, Andreas Schmitz, Mark-Oliver Rodel —

The tadpoles of eight West and Central African Leptopelis species (Amphibia: Anura: Arthroleptidae). ... 56

General Section

Jerry D. Johnson, Vicente Mata-Silva, Larry David Wilson — A conservation reassessment of the Cen-

tral American herpetofauna based on the EVS measure 1

Cesar L. Barrio-Amoros, Andres Chacon-Ortiz, Fernando J.M. Rojas-Runjaic — First report of the sala-

manders Bolitoglossa leandrae and B. tamaense (Urodela, Plethodontidae) for Venezuela 95

Christopher J. Michaels, Benjamin Tapley, Luke Harding, Zoe Bryant, Sebastian Grant, George

Sunter, Iri Gill, Oscar Nyingchia, Thomas Doherty-Bonec — Breeding and rearing the Critically En-

dangered Lake Oku Clawed Frog {Xenopus longipes Loumont and Kobel 1991) 100

Kevin R. Messenger and Yong Wang — Notes on the natural history and morphology of the Ningshan Lined

Snake (, Stichophanes ningshaanensis Yuen, 1983; Ophidia: Colubridae) and its distribution in the Shennon-

gjia National Nature Reserve, China Ill

Harshil Patel, Raju Vyas, Shantilal K. Tank — On the distribution, taxonomy, and natural history of the Indi-

an Smooth Snake, Coronella brachyura (Gunther, 1866) 120

Table of Contents Back cover

Cover: A juvenile Lined House Snake ( Bothrophthalmus lineatus) caught in a funnel trap in closed canopy lowland

evergreen forest during a herpetological survey of the West Nimba Nature Reserve, Nimba County, Libera (07°

29’33.0”N, 008° 4F59.5”W, 432 m a.s.l., 19 November 2011). Photograph: Bill Branch.

Instructions for Authors: Located at the Amphibian & Reptile Conservation website:

http://amphibian-reptile-conservation.org/submissions.html

volume 9

2015

NUMBER 2