Museum of Comparative Zoology Breviora 20

A
B

RE VI O

M

e HI iin o I v^oimpaFattive

HI s

us ISSN
CAMBRiDCiE, Mass.

2

R
ogy

0006-9698

Number

February 2009

515

NEW SQUEAKER FROG (ARTHROLEPTIDAE: ARTHROLEPTIS) FROM THE
CAMEROON VOLCANIC LINE WITH REDESCRIPTIONS OF ARTHROLEPTIS
ADOLFIFR/EDER/C/ NIEDEN, 1911 "1910" AND A. VARIABILIS MATSCHIE, 1893
A

David C. Blackburn,' Legrand N. Gonwouo,- Raffael Ernst,^ and Mark-Oliver Rodel^
Abstract. We describe a new species of squeaker frog {Arthrolcpiis) from Mt. Manengouba in southwestern
Cameroon. The new species is distinguished from other Cameroonian Artlirolcptis by moderately larger body size; a
darkened throat and posterior thigh, both with many white spots; and, in females, a fourth fmger longer than the first
and second fingers. This species corresponds to a Cameroonian taxon previously identified as Artlirolcptis
adolfijriederici but which has been long recognized as distinct. Multivariate morphometric analysis demonstrates that
the new taxon is distinct from Arthroleptis variabilis, which occurs in the surrounding lowlands. Because of the
general similarity of the new species to A. variabilis and its previous confusion with A. adolfifriederici, redescriptions
of the latter two species are provided. This refinement of taxonomic knowledge will facilitate future study of cryptic
or undescribed large Arthroleptis from western, central, and eastern Africa.

Key words:

Amphibia; Anura; new

species; biodiversity hotspot;

Cameroon

highlands; central Africa

Resume. Nous decrivons une nouvelle espece de grenouille (Arthroleptis) du Mont Manengouba au sud du
Cameroun. La nouvelle espece se distingue des autres Arthroleptis Camerounaises par sa moderement plus grande
taille corporelle, de nombreux points blancs trouves a la fois sur le cou sombre et sur les cuisses posterieures, et, chez
les femelles, un quatrieme doigt plus long que les premier et deuxieme doigts. Cette espece correspond a un taxon
Camerounais prealablement identifie en tant que A. adolfifriederici. mais qui est depuis longtemps reconnu comme
distinct de celui-ci. Des analyses morphometriques multivariees demontrent que ce nouveau taxon est different de A.
variabilis, qui est

trouve dans

les

espece et A. adolfifriederici, et du

'

Natural History

Museum and

basses plaines environnantes.
fait

de sa ressemblance a A.

Du

fait

variabilis,

de

la

confusion anterieure entre

la

nouvelle

nous decrivons a nouveau ces dernieres deux

Biodiversity Research Center, University of Kansas. Lawrence,

Kansas 66045,

U.S. A; e-mail: dblackb(ajfas. harvard.edu

"Cameroon

Biodiversity Conservation Society,

Yaounde, Cameroon;

'Department of Biodiversity Dynamics, Technische

Universitiit

e-mail:
Berlin,

lgonwouo(a yahoo.com
D-12165,

Berlin,

Germany;

e-mail:

raffael.ernst(a tu-berlin.de
*

Humboldt

University.

Museum

of Natural

History,

10115

Berlin.

Germany;

hu-berlin.de

© The

President and Fellows of Harvard College 2009.

e-mail:

mo. roedelf« museum.


BREVIORA
especes.

No. 515

Ce raffinement de notre connaissance taxonomique facilitera les etudes futures portant
ou non decrits, et en provenance d'Afrique de TOuest, de I'Est et Centrale.

sur

les

Artlvoleptis

larges, cryptiques

Squeaker frogs

{ArtJiroleptis sensii Frost,

Amphibian Assessment, 2006;

1986; Global

much

of

Herrmann

sub-Saharan Africa. This genus occurs

in

tions previously assigned to A. adolfifrieder-

2007) are distributed throughout

two biodiversity hotspots deemed conservation priorities (Myers et al., 2000) and hkely
contains high levels of cryptic species diver-

Poynton, 2003b; Rodel and Ban2004). The taxonomic history of

ici

2005). Tanzanian popula-

et al.,

correspond to several currently recognized

taxa, including Arthroleptis affiuis Ahl, 1939

"1938," Arthroleptis

fa77«^r/

Grandison, 1983,

sity (e.g.,

and possibly Arthroleptis

goura,

Bourgeois, 1961 (Grandison, 1983; Poynton,

Artlvoleptis

characterized by long-stand-

is

ing disagreements between researchers about
the status of both species

taxa

(e.g.,

and supraspecific

Laurent, 1940, 1954, 1973; Perret,

1991; Poynton,

1976, 2003a;

Poynton and

Skelton-

leleupi

2003b; Poynton and Loader, 2008), and
is

it

unlikely that A. adolfifriederici occurs in

eastern Tanzania

(e.g.,

Poynton and Loader,

2008).

Broadley, 1985; Schmidt and Inger,

1959).

During recent fieldwork on Mt. Manengouba in southwestern Cameroon, specimens

Typically, these disagreements focus

on the

of an undescribed Arthroleptis species were

smallest

species,

which

placed in Schoutedenella but

now

ed within Arthroleptis (Frost

et al.,

also Blackburn,

2008a).

were

previously

One

are includ-

2006; see

large species

taxonomic confusion is Arthroleptis adolfifriederici Nieden
1911 "1910," which was originally described
on the basis of two specimens from the
mountains of Rwanda. Shortly thereafter,
Nieden (1913) assigned specimens from
northeastern Tanzania to A. adolfifriederici,
and Noble (1924) stated that this species
occurs from "Cameroon eastward to the
Lake Region." It is unclear, however, on
what basis Noble's (1924) range was established. Over the past century, A. adolfifriederici has been claimed to occur in Cameroon at the western extent of its distribution
that has caused significant

(e.g.,

Noble,

1924;

Perret,

1966)

to

the

found. These specimens are distinct from
other described Cameroonian species.

between

similarity

ly

10

km

initiated

of Mt.

southeast

specimens as a new species.

MATERIALS AND METHODS
Specimens were collected during visual
encounter surveys on Mt. Manengouba
the Republic of

Cameroon

Specimens

in

Surveys

1).

chloretone

in

standard

following

Kenya at its eastern extent (e.g., Barbour
and Loveridge, 1928; Channing and Howell,
1957).

(Fig.

and near the village of
Nsoung, as well as near Moabi and Ebonemin and in forests near the summit. Animals
were euthanatized in an aqueous solution of

were conducted

samples were

1942.

Manengouba,

which describes these

study,

this

Eastern Arc Mountains of Tanzania and

Loveridge,

adolfifrie-

from near Nkongsamba, approximate-

derici

procedures (Simmons,

2006;

specimens

these

Perret's (1966) description of A.

all

The
and

2002).

preserved

in

collections

Liver

95%

tissue

ethanol;

remaining voucher specimens were preserved
overnight in

10%

neutral buffered formalin

70%

Type material

from Cameroon have long been recognized
as distinct from A. adolfifriederici; however,

before storage in

taxon has remained undescribed (e.g.,
Amiet, 1987; Frost, 1985, 2007; Gartshore,

those of Arthroleptis stenodactylus Pfeffer,

this

was examined of all

ethanol.

species discussed, except

1893, which were destroyed during

World


2009

NEW CAMEROONIAN

ARTIIROLEPTIS

War

measurements (±
and
a dissecting microscope. Limb measurements
were taken on the right side. Measurements
follow Blackburn (2005), which is a modification ofMatsui (1984). Images of preserved
specimens were taken with a JVC 3-CCD
digital camera mounted on a dissecting
microscope with AutoMontage Pro 5.0
0.1

(Frost, 2007). All

II

mm)

were taken with

Museum

(Synoptics).

Leviton

et

digital calipers

abbreviations follow

(1985), with the addition of

al.

Museums of Malawi, Blantyre (MMB).
To determine whether the new taxon
differs

morphologically from the superficialArthroleptis variabilis, a multivar-

ly similar

based on the type specimens was

iate analysis

used

Twenty-seven

ordination.

for

adult

specimens (including three syntypes) of A.

were measured (Appendix

variabilis

1).

Be-

cause few male specimens have been collectof

ed

were

new

the

species,

female A.

to

restricted

identified

by

this

analysis

variabilis.

large

body

is

Females
size,

the

presence of ova (visible either in dissection

through

or

the

skin),

the

lack

of male

secondary sexual characters typical of the
genus, or a combination of factors (Black-

burn, in press). All data were natural log-

transformed before analyses were conducted

R 2.7.1

by
for

for

Mac OS X

(The
Computing,

Statistical

A

R-project.org).

technique,

R

multivariate

principal

Foundation
http://www.
ordination

component

analysis

(PCA), was used to analyze patterns of
variation and covariation within the measurement data. The covariance matrix, rather
than a correlation matrix, was used, and all
component axes were scaled to be equal to
their eigenvalues. Those components ac-

Figure

1

.

Distribution

Mt. Manengouba
nates

type

in

locality

of Anlirolcpiis perreti on

southwest Cameroon. Star designear village

indicate paratype localities.

of Nsoung;

circles


BREVIORA

85% of the cumulative variance
were retrieved from the analysis. The relationship between the two species in morphocounting for

space was evaluated by
component (PC) scores.

plotting principal

No. 515

(705G), adult male (27.4 mm). Republic of

Cameroon,

NEW

Arthroleptis perreti^
Perret's

Arthroleptis

SPECIES

new

species

2, 3

adolfifriederici:

Perret

(1966):

Gonwouo; ZMB 71475 (0883N), adult
(34.7 mm). Republic of Cameroon,

N.

Arthroleptis sp.

female

7:

Gartshore (1986): 220.

Amiet (1987): 100.
Hohtype. MCZ A- 137978

J.-

L. Amiet;

DCB

34368),

Republic

female.

adult

number

(field

Southwest Province,

L.N. Gonwouo; ZMB 71474 (103G), aduU
male (30.6 mm). Republic of Cameroon,
Littoral Province, Mt. Manengouba, 05°00'N,
009°49'E, 2,045 m elevation, 6 August 2005, L.

396.

Following personal communication with

of Cameroon,

Mt. Manengouba, near Nsoung, 04°59'N,
009°48'E, 1,420 m elevation, 4 July 2004,

Squeaker Frog

Figures

of

Littoral Province, Mt.
summit, 05°00'N, 009°51

17

March

2006, L. N.

Manengouba,

'E,

Gonwouo;

Cameroon,

gouba,04°59'05.5"N,009°48'41.0"E, 1,400 m
elevation, 13 July 2006, D. C. Blackburn, K.

28 September 2006, L. N.

Blackburn, and M. T. Kouete.

MCZ

Paratypes.

A- 136931

A-1 36932

(DCB

juvenile,

respectively.

(DCB

Republic of Camer-

oon, Littoral Province, Mt.
type

locality

m

eleva-

of Cardioglossa nuinen-

gouba (Blackburn, 2008b), 27 September
2004, D. C. Blackburn, J. L. Diffo, and L.
N. Gonwouo; MCZ A- 137980 (DCB 34432),
adult female. Republic of Cameroon, Southwest
Province, Mt.
Manengouba, 05°0r48.9"N,
009°50'37.3"E,

2,110

m

elevation,

21

July

S. Blackburn, M.
A- 137981 (DCB 34434),
juvenile. Republic of Cameroon, Southwest
Province, Mt. Manengouba, 05°01'09.1"N.
009°5r04.0"E, ~ 2,180 m elevation, same
date and collectors as A- 137980; ZMB 71471
(86G), Republic of Cameroon, Littoral Province, Mt. Manengouba, near summit, adult
female (42.0 mm), 05°0rN, 009°52'E, 22

2006, D. C. Blackburn, K.

T.

Kouete;

March

MCZ

2006, L. N.

Gonwouo;

ZMB

71472

Littoral

Province,

Manen-

Mt.

gouba, 05°00'N, 009°51'E, 2,185

m

elevation,

Gonwouo.

Specimens collected
by Legrand N. Gonwouo on Mt. Manengouba, Republic of Cameroon; sex and
snout-vent length (SVL) are provided for
Additioiud Material.

each specimen.

ZMB 71484 (0263LG), juvenile (22.9

Manengouba,

05°00'38.9"N, 009°51'24.8"E, 2,160
tion,

34208),

34209), subadult female and

near

m elevation,
ZMB 71476

2,200

(0023LG), adult female (39.3 mm). Republic of

Cameroon, Southwest Province, near village
of Nsoung, southwest slope of Mt. Manen-

S.

Mt. Manen-

71473 (560G), aduU female (39.6 mm), Republic

DESCRIPTION OF

Littoral Province,

gouba, 05°00'N, 009°49'E, 2,045 m elevation, 6 August 2005, L. N. Gonwouo; ZMB

western fiank, no data available,

~

mm),

1,500

m

December 2006; ZMB 71478
(0899N), aduh female (41.5 mm), near Manengouba lakes. 05°0r49"N, 009°50'39"E,
2,155 m elevation, 20 March 2006; ZMB
71479 (282C), juvenile (22.6 mm), near
elevation, 20

Nsoung, 04°59'N, 009°48'E, 1,420
4 July 2005;

tion,

female

(37.9

009°50'E,

ZMB

m

eleva-

71485 (0393N), adult

mm), near Moabi, 05°10'N,

1,980

m

elevation,

12

August

ZMB

71486 (0396N). juvenile (23.6
mm), near Moabi, 05°10'N, 009°50'E, 1,980
2005;

m

elevation,

12

August 2005;

ZMB

71487

(0804N), juvenile (23.1 mm), near Ebonemin,

05°00'N, 009°44'E,

1,650

ZMB

m

elevation,

18

November

2005;

nile

mm), near Ebonemin, 05°01'N.
1.465 m elevation, 16 November

(25.1

()09°46'E,

71488 (0992N), juve-

NEW CAMHROONIAN

2009

Figure

2.

Arihrulepii.s pcrrcii in

137980)

in ventral

2005;

ZMB

mm), no data
13

life:

holotype

view (B). Scale bar, 10

August 2005;

ZMB

~

2.085

(MCZ A-137978)

view (A), and paratype

(MCZ

A-

(25.1

71492 (0865N). adult female (42.7 mm), near

m elevation.

summit. ()5°00'N. 0()9°5rE, 2,185 m elevation. 15 March 2006; ZMB 71494 (0884N),

71490 (0963N), adult

female (41.1 mm), no data available,

~

2,085

September 2005; ZMB 71491

(88G), near summit, adult female (44.7 mm),

m

in roslrokitcral

mm.

71489 (0945N), juvenile
available.

ARI II ROI.EI'TIS

elevation, 5

05°0rN. 009°52'E, 22 March 2006;

ZMB

mm), near summit, 05°00'N,
009°5rE. 2.200 m elevation, 17 March 2006;
ZMB 71495 (0893N), adult female (33.0 mm),
near summit, ()5°00'N. 009°5rE. 2,200 m

juvenile (21.7


BREVIORA

No. 515

(0006LG), adult female (38.3 mm), 04°59'N,
elevation, 25 September

m

009°50'E, 2,010

ZMB

2006;
(30.4

(0007LG),

71501

adult

mm), 04°59'N, 009°50'E, 2,010

vation,

ZMB

September 2006;

25

male

m

ele-

71502

(0020LG), adult female (37.2 mm), 04°59'N,
m elevation, 25 September

009°50'E, 2,010

ZMB

2006;
(37.6

71503 (0022LG), adult female

mm), 05°00'N, 009°5rE, 2,185

vation,

ZMB

September 2006;

28

m

ele-

71504

(0024LG), adult female (35.5 mm), 05°00'N,
009°51'E, 2,185

m

September

elevation, 28

ZMB

71505 (0027LG), juvenile (22.0
mm), 05°00'N, 009°51'E, 2,185 m elevation,
2006;

ZMB

28 September 2006;

m

71506 (0044LG),

mm), 05°02'N, 009°86'E, 2,066

juvenile (27.2

elevation, 28

September 2006;

ZMB

71507

(0045LG), adult female (36.8 mm), 05°02'N,
009°86'E, 2,066

ZMB

2006;

mm),

(43.8

m

September

elevation, 28

71508 (0048 LG), adult female

009M9'E,

05°00'N,

September 2006;

elevation, 30

m

2,184

ZMB

71509

(0053LG), adult female (27.4 mm), 05°00'N,

m elevation, 30 September
71510 (0182LG), adult female
(36.8 mm), 04°59'N, 009M7'E, 1,393 m ele-

009°49'E, 2,184
2006;

ZMB

November

vation, 19

A

Diagnosis.
that

is

2006.

medium-sized

Arthroleptis

very similar to A. variabilis but readily

distinguished from that species by lacking a

midline gular stripe (Fig. 2) and by having a
longer snout and, in females, a

relatively

fourth fmger that
Figure
137978)
10

3.

in

Artliroleptis perreti

holotype

(MCZ

A-

dorsal (A) and ventral (B) views. Scale bar,

mm.

2,045

m

71498

defined light mottling on a dark-brown or

August 2005;

6

female

adult

ZMB

2,045

m

71499

(40.2

-gray background.

mm),

perreti (44.7

elevation,

(843C),

elevation,

6 August

2005;

ZMB

3),

ZMB

6

adult

female (34.2 mm), 05°00'N, 0()9°49'E, 2,045

m

In

ZMB 71496 (31G),

2006;

elevation,

2005;

and

mm), 4°59'N, 009°48'E,

05°00'N, 009M9'E,

August

first

section).

dark gray with many small white
whereas in A. variabilis, the
posterior surface of the thigh exhibits poorly

March

(553G),

Variation

is

spots (Fig.
adult female (35.5

longer than the

addition, the posterior surface of the thigh of
A. perreti

elevation, 17

is

second fingers (see

71500

than

that

mm

(37.8

SVL,

of other

Cameroonian
iabilis

Maximum body

ZMB

size in A.

71491)

described

is

larger

species

of

Arthroleptis. including A. var-

mm

SVL; Blackburn, 2008a);

mean male SVL of A.

perreti

is

29.5

mm (n

=


.

NEW CAMEROONIAN

2(HW

±

3;

differs

medium

SVL >
less

=

25;

±

is

Anhroleptis perreti

from other

Arihrolcpiis

large

to

4.3).

following ways

the

in

SVL

whereas mean female

1.8).

mm (n

37.5

(maximum

mm): from A. culolfifricdcrici by
granular skin, more extensive pigmenta35

on

tion

throat,

metatarsal

and

tubercle;

globular

less

from

A.

inner

by

a/finis

lacking supernumerary tubercles on the

feet;

from Avthroleptis francei and A. steuodactyby tibiofibula length greater than 50%
SVL; from Arthroleptis krokosua by lighter
pigmentation on throat and vent, lacking
large well-defined black spots on the lateral
surface of the body, and by a fourth finger
that is longer than the first and second
fingers; from A. stenodactylus by an inner
his

metatarsal tubercle length

less

than

80%

eye-narial distance; eye diameter 2.1 times
distance from naris to rostral
region

of Holotype.

mm

SVL), moderately robust female
with slender limbs (Figs. 2A, 3; Online
Table 1 ); head somewhat broad, head length
79% head width; snout projecting approximately 1
beyond lower jaw; rostral tip
(37.4

mm

rounded
slightly

in

dorsal

blunt

view,

and only

curving posteroventrally in lateral

view; eyes barely project

beyond

eyelids in

interor-

ovoid,

distinctly

tympanic annulus poorly defined; supratympanic fold absent; tongue very broad,

eye;

covered

heart-shaped,
pustules;

many

with

attachment

anterior

small

tongue

of

narrow; posterior notch of tongue approximately one fourth anteroposterior length of
tongue; prominent, fiat, fiap-like median

on dorsal tongue surface near antechoana hidden by maxillary
shelf in ventral view; premaxillary and
papilla

rior attachment;

maxillary teeth present, but hidden by

lips;

vomerine teeth absent.
Skin of limbs, dorsal surface of head and

and

body,

smooth

head

ventral

tuberculate in Hfe; Fig.

2);

(more

skin of lateral

body glandular, becoming more indistinct on
median skin raphe promi-

ventral surface;

nent in

life

(Fig. 2), but very indistinct in

preservative; cloacal region weakly glandular.

Limbs

Medium-sized

tympanum

internarial

90%

height slightly less than half the diameter of

globose inner metatarsal tubercle.
Description

tip;

internarial distance

fiat;

distance;

bital

of

from Arthroleptis nikeae by
much smaller adult body size (> 50 mm SVL
in A. nikeae); from A. taiineri by a relatively
narrower head; from A. variabilis by lacking

a white stripe on the midline of throat; and
from both A. stenodactylus and A. variabilis
by having a relatively smaller and more
toe length;

first

ARI II ROLIIPTIS

and

slender;

digits

well

digits

of

developed

slender; relative length of fingers: III

>

II

>

>

pes

IV

fingertips rounded, slightly swollen,

I;

approximately same width as
finger with
single

but

manus and

both

subarticular

metacarpal

rest

of finger;

rounded, prominent, globular,

tubercles;

present

tubercles

palmar and
but weakly

developed and barely projecting from surface
of hand;

webbing between manual

93%

digits

of crus length;

dorsal view; eyes placed well medial of, and

absent;

not projecting laterally beyond, margins of

IV > III > V > II >
and slightly expanded to
approximately same width as interphalange-

head; eyes fiush with dorsal margin of head
in lateral view;

eye diameter approximately

thigh

relative length of toes:
I;

toe tips pointed

equal to interorbital distance; pupil round to

al

slightly horizontally elliptical in preservative;

elongate,

loreal

region weakly concave; naris small,

rounded, facing
in

dorsal

indistinct,

view;

laterally,

and barely

canthus

rostralis

rounded; eye diameter

1.5

visible

short,

times

length

regions;

toes

with

prominent,

single,

and slightly conical subarticular
tubercles; webbing between pedal digits
absent; prominent fiange-like inner metatarsal tubercle,

length

72%

of

first

toe length.

Measurements. See Online Table

I


BREMORA
Coloration of Holotype (in Alcohol). Dor-

ground color silvery brown with small,
weakly defined gray and well-defined darkbrown spots (Fig. 3); iris dark brown with
pale cream pupil; snout silvery gray to dark
brown near rostral tip; loreal and suborbital
regions nearly solid medium to dark brown

becoming lighter and slightly broken postesal

medium brown

rior to eye; interorbital bar

No. 515

um-brown

foot silvery

brown

tips

of

toes

slightly

continuous

with

dark

mask

of

phalangeal

in

and

unpigmented

I-III

ultimate

skin;

tending from posterodorsal margin of orbit,

general

dark brown; ultimate phalangeal joints

brown spots on posterodorsal

head; dark-brown supratympanic band ex-

melanocytes;

medial toes; dorsal surface of toe

lateral to

but broken and poorly delimited from other

of

and creamy with many small

well-defined

trend of lighter and less pigmentation from

lighter

surface

spot on lateral margin of foot at

base of metatarsal V; base color of dorsal

IV

V

than

surrounding

triangle with

apex directed

pigmentation

dark-brown

creamy;

of toes

joints

dorsally centered on cloaca.

Lateral margin of lower

jaw dark brown

with small white spots at regular intervals;

brown

suborbital region, extending posteroventrally

throat

and terminating anterior

(Fig. 3B); posterior throat, just anterior to

of arm;
flecks,

tympanum
and

to

and

just dorsal

translucent, with silver

darkened

dorsally

becoming

confluent with supratympanic band; dorsal
surface of arms silvery gray grading into

dark brown on forearm and

wrist;

single,

broken dark-brown band on forearm; dorsal

surface of hands mottled gray and brown
with large cream-colored spots on second
and first fingers; ultimate interphalangeal
joint of all fingers unpigmented and cream in
color; posterior to skull, two dark-brown,
highly broken chevrons with apices directed
anteriorly, separated by light gray-brown
region approximately 3.8 mm in anteroposterior length; highly broken, dark-brown
band at lateral margin of dorsum extending
from just above arm along lateral surface
toward inguinal region before becoming
indistinct; lateral surface of body with many
small medium- and dark-brown spots and
base color
into

much

lighter than

creamy white of ventral

dorsum grading
surface; dorsal

surface of hind limbs grayish brown; very

with

white

distinct

spots

and laterally bordering omosternum, darker than rest of throat; medial
surfaces of arm and forearm unpigmented
and creamy; ventral surface of forearm very
dark brown with several prominent white
spots; ventral surface of hand and fingers
brown; palmar, metacarpal, and subarticular
clavicles

tubercles

unpigmented;

lighter in color

than

ventral

rest

fingertips

of finger; brown

pigmentation of throat broken into mottling

over pectoral girdle extending ventrally as
diffuse

brown

pigmentation

poorly

defined

white

ventral thigh both
scattered

with

large,

and
cream colored with few
spots;

brown melanocytes;

venter

ventral surface

of crus mottled brown and cream; plantar
surface

homogeneous dark brown; subaron toes and inner metatarsal

ticular tubercles

tubercle lighter brown, tending to gray, than

surrounding skin but not unpigmented.
Coloration

in Life.

Based on D. C. Black-

burn's field notes and photographs (Fig.

2).

Dorsal coloration ranging from dark gray

transverse dark-brown stripes; feet mottled

brown markings (MCZ A- 137980) to
brown with red tones (MCZ A- 137978;
Fig. 2A); darker markings on dorsal and
lateral surfaces ranging from light to dark
brown and even black; lighter markings on

gray and brown with poorly defined medi-

lateral surface

indistinct,

broken,

dark-brown

band

on

dorsal surface of thigh; posterior surface of
thigh dark

brown with prominent

spots (Fig. 3A); crus with

large white

two prominent

with

ranging from white to gray;

NEW CAMEROONIAN

2009

\ciitial surface o\' lliroal generally

gray

vvilli

ARTIIROI.EPTIS

Herrmann

et

Fven

2005).

al..

if

creamy xellow lones; venter with more
pronounced creamy yellow lones; bright

these t)ther localities, A. perreti

yellow splotches in inguinal region (Fig. 2B).

2().()0()

tainl

Wtiicition.

ed

in

the

Meristic \arialion

Online Table

tongue

I.

more

is

document-

is

The median

conical

137980. Sexual dimorphism

papilla of

MCZ

in

in

have

A-

A. pcrrcti

is

an

present at

would

of occurrence

extent

less

still

than

km^.

Coiiservaiioii.

(iiven the small

number of

possible localities, extent of occurrence (pre-

<

sumably

20.000 km-), and that both the

extent and quality of forest habitats on Mt.

similar to that of other species oWlnhroleptis

Manengouba, and probably other nearby

and includes larger
body size in females and the presence of an

elongate third finger and both digital and
inguinal spines in males, in male A. pcrreti
(e.g., ZMB 71472 and 71474). small spines

mountains, are declining (Gartshore, 1986;

(Blackburn,

press)

in

medial surface of both the second

line the

(MDIl) and
number of

(MDlll)

third

spines

digital

fingers.

is

comparable

between the male paratypes

(ZMB

MDII

right

18:

MDII

MDIIl

right 6. left-8,

ZMB
I

MDII

71474,

right

18, left

20).

(ZMB

male paratype

white spines are found

right

The

13,

71472,
17, left12,

left

In the oiie larger

71474),

pronounced

in the inguinal

region

and extend both forward along the lateral

surface of the body and dorsally onto the
posterior dorsal surface of the body and the
proximal dorsal surface of the hind limbs.
Unlike females, the fourth finger of males of
A. pervt'ti

is

similar in length to that of the

and second fingers.
Habitat and Range. All specimens were
collected in montane forest during the day
first

when they were active in
litter. The type series comes from

Global Amphibian Assessment, 2006; Gon-

wouo

et

2006; Stuart, 1984), A. perreti

al.,

should be considered Vulnerable according
to

lUCN

(2001) criteria.

Etymology. This species
of Dr. Jean- Luc Perret

Morphological Comparison with Arthrovariabilis. As indicated in previous

leptis

literature

(Perret,

prominent

the

Mt. Manengouba
perreti might

also

1).

elevation)

on

occur on several other

nearby mountains, including Mt. Nlonako,

Mt. Kupe, and

in the

Rumphi

Hills (Gart-

et al., 2005; Lawson,
specimens
However,
1993; Perret, 1966).
studied by Lawson (1993) and Herrmann et
al. (2005) do not correspond to A. perreti and
represent A. variabilis (i.e., Lawson, 1993) or

shore, 1986;

possibly

Herrmann

another undescribed species

(i.e..

species

is

white

gular

stripe.

These

species appear to be osteologically indistin-

guishable

PCA

on

the

indicates

of

basis

that

radiographic

unpublished

(Blackburn,

analysis

several

data).

morphological

characters differentiate A. perreti from A.

The

variabilis (Fig. 4).

first

principal

compo-

nent axis (PCI) accounts for the majority

(52.7%) of the variance

in the

data and

taken as a general measurement of body

moist leaf

m

this

1966),

extremely similar to A. variabilis yet lacks

(Table

Populations of ^.

honor

comments on this new
more than 40 years ago (Perret, 1966).

differentiated along

(Fig.

in

provided prelim-

inary, but accurate,

species

(1000-1400 h)

high elevations (1,400-2,200

named

is

who

1);

A. perreti

is

size

and A. variabilis are not
PCI. The second princi-

components axis (PC2) accounts for

19.4% of the variance and plots of PC2
scores reveal that A. perreti and A. variabilis

pal

are

differentiate along this axis (Fig.

Plots of

PC3

scores

do not

4).

reveal differences

between these two species. PC2 loads most
strongly, and positively, on inner metatarsal
tubercle length, eye diameter, and tympanum
height

(Table

loadings for

finger length,

The

I).

PC2

strongest

negative

are on snout length, fourth

and

fifih

toe length. Because


BREVIORA

10

Components Analysis

Principal

No. 515

Table 1. Principal components analysis comparing
Arthroleptis PERRETI and Arthroleptis variabilis.
Eigenvalues, percent variance, cumulative variance,

and loadings for the

first three principal

(PC) AXES.

Figure

is

Scatteqjlot of the

4.

component

scores.

The second

plotted against the

indicator of
variabilis,

PC2

body

first

principal

first

Circles

size.

and second principal

component (PC2)
component (PCI), an

principal

represent

Arthrolcptis

squares represent Arthwieptis pencti.

scores for A. perreti are lower than

one specimen of A.

all

but

these results

variabilis,

together indicate that the length of the inner

metatarsal tubercle, eye diameter, and tym-

panum

height

variabilis

and

fifth

and

are

relatively

greater in A.

that the snout, fourth finger,

toe are relatively longer in A. perreti.

Following

this

PCA,

visual inspection of the

specimens indicated that only relative fourth
finger length

is

a reliable diagnostic without

actually taking measurements.

REDESCRIPTIONS
Arthroleptis perreti

has been previously

confused with A. adolfifriederici
1975; Gartshore, 1986;

(e.g.,

Herrmann

Amiet,

et al.,

2005;

Lawson, 1993; Ferret, 1966; Plath et al.,
2004). Although the features given by Perret
(1966) are sufficient to distinguish A. perreti

from both

A. adolfifriederiei

this species

mid

A. variabilis,

remained undescribed for more

than 40 years. This
part, to the general

is

likely due, at least in

inadequacy of the original

component


NEW CAMHROONIAN

2009

ARTIlROLEPriS

Arthroleptis adolfifriederici: Lovcridge (1942):

429

Hyphen removed from name.

(part).

A rthrolcpis
Lovcridge

adolfifriederici

387;

(1953):

adolfijriederiei:

Skelton-Bourgcois

(1961): 323.

Ahroscapluis adolfi-friederici: Laurent

(

1957):

275.

Arthroleptis adoljifriderici: Lovcridge (1957):

352 (part). Incorrect subsequent spelling.
Reference Sample. Type material, sex, and

SVL (mm)
cratic

indicated in parentheses.

Republic of Congo:

(female).

Rwanda:

[formerly

ZMB

FMNH

MCZ

Demo-

A- 14696

73836 (syntype

21789]; female, 40.1);

ZMB

21787 (syntype; female, 42.0); KU 154322
(male),
154323 325 (females, 30.1, 38.0,
39.1). Uganda: CAS 177029 (female, 35.8),
177030 (cleared and stained; female, SVL
undocumented),
177031
(female,
43.4),
177032 (juvenile, 22.2), 177033 (male, 27.6).
Diagnosis.

A

medium-large

Arthroleptis

characterized by long, slender hind limbs, a

fourth finger as long as or
first

and second

fingers,

much

longer than the

and generally glandu-

lar skin. Arthroleptis adolfifriederici differs in

ways from other medium to large
(maximum SVL > 35 mm): from

the following
Arthroleptis
A. affinis

by a

less

prominent and

tympanum and by
tubercles

on the

lacking

feet;

Figure
in dorsal

5.

Anliroleptis cuhlfijrieclerici

{CAS

m

elevation

(lat.,

long., elev., estimated).

Arthroleptis adolfi-friderici:
165. Incorrect

A rthroleptis

(

Nieden (1913):

subsequent spelling.

A hroscaplms

Laurent (1940): 82.

)

lighter

on the lateral
by a more
globose inner metatarsal tubercle, more grandular skin, and less pigmentation on throat;
from A. nikeae by smaller adult body size (>
50 mm in A. nikeae); from A. stenodactylns by

Nyungwe; Gartshore, 1986)

1,800-2,400

by

mm.

and Bugoie (= Bugoya?) Forests; Nyungwe
Forest, ca. 02°20'S^to 02°45'S, 029°05'E to
ca.

greater than

pigmentation on throat and vent and lacking
large well-defined black spots

surface of the body;

029°25'E,

much

krokosita

177031)

(A) and ventral (B) views. Scale bar, 10

(presently called

A.

round

from A. francei and A.

stenodactylns by crus length

50% SVL; from

less

supernumerary

adolfi-friederici:

from

A. perreti

an inner metatarsal tubercle length less than
80% first toe length; from A. tanneri by a
relatively

narrower head; from A.

variabilis

by

on the midline of throat;

from both A. stenodactylns and A. variabilis by
lacking a white stripe

having a globose inner metatarsal tubercle.


BREVIORA

12

Table

2.

Measurements (mm) of female svntypes of
ArTIIROLEPTIS ADOLF//

Rlf:n/:RI(

7.

No. 515


NEW CAMEROONIAN

2009

FMNH
is

73836. This description of coloralion

based on

a siininiar\

of the reference

all

specimens.

Dorsal base color

light to

medium brown

with small dark-brown spots (Fig. 5A);

dark

gray

with

brown than
brown

white

pupil:

snout

iris

lighter

interorbital bar but grading to

dark

near

and
suborbital regions light brown: dark-brown
supratympanic band extending from posterodorsal margin of orbit, extending posteroventrally and temiinating just posterior to the
ventral
lucent

rostral

tip;

loreal

tympanum margin: tympanum

and

light

trans-

brown: dorsal surface of amis

brown to tan: single, somewhat broken
brown band on forearm: dorsal surface of

ARIII ROLEPTIS

/hi/vtat

ami

13

Specimens similar to

Ran}y,c.

the type specimens have been collected

in

mountainous areas to the north and east of
Lake Kivu in the Democratic Republic of
Congo, Rwanda, and southwestern Uganda

(Drewes and Vindum, 1994: Nieden, 1911.

We

1913).
explicit

have not examined, nor found

reference to, specimens from

although
occurring there
undi,

has

it

been

(Channing

Bur-

reported

and

as

Howell.

Amphibian
Assessment,
examined from Tanzania,
Malawi, and Kenya previously referred to as
Global

2006:

2006). Specimens

A. adoljifriederici invariably represent differ-

Remarks

light

ent species (see

hands mottled

Remarks. Nieden (1913) referred Tanzanian specimens from Tanga and Amani to A.

bar

bital

light

medium

and dark brown: interorbrown: posterior to

to dark

two dark-brown, highly fragmented
chevrons (e.g., MCZ A- 14696) with apices
directed anteriorly, separated by light-brown
region: scattered medium- to dark-brown
small spots on dorsal and lateral surfaces of
body: lateral surface of body with many small
medium- and dark-brown spots and base color
much lighter than dorsum: dorsal surface of
hind limbs light brown with medium- to darkbrown mottling: crus with two variably
skull,

developed transverse brown

Throat very

medium-brown

light

scattered

spots especially concentrated

Given that the material

later

used by Ahl (1939) to describe A. ajfinis
single

large

specimen collected

Awerinzew in Amani, it seems
Nieden (1913) had examined

by

S.

likely

the

is

a

G.
that

same

specimen but referred xiio A. adolfifriederici.
Following Nieden's (1913) precedent, Bar-

bour and Loveridge (1928) assigned many
specimens from eastern Tanzania to A.
adolfifriederici and Loveridge (1942, 1957)
recognized

later

synonym of A.

stripes.

brown with

adolfifriederici.

section).

A.

affinis

adolfifriederici.

as

a

junior

These records

extend

the
range of A. adolfifriederici
throughout much of the Eastern Arc Moun-

margins of lower jaw (Fig. 5B): venter

tains.

Later authors recognized A. adolfifrie-

brown with few to no dark
spots: medial surfaces of arm and forearm
unpigmented and creamy: ventral surface of

derici
(e.g.,

and A. affinis as two distinct species
Channing and Howell. 2006: Grand-

ison,

1983: Poynton, 2003b: Skelton-Bour-

at

mostly very

light

fore limbs very light

brown: palmar, meta-

less

and subarticular tubercles generally
pigmented than surrounding skin: ven-

tral

surface

carpal,

o'i

thigh very light brown, with

no markings: ventral surface of
crus very light brown to weakly mottled with
essentially

medium brown:
medium brown.

light to

to

Variation.

umented

in

plantar surface light

Measurement variation
Table

2.

geois, 1961), but records o{ A. adolfifriederici

from throughout the Eastern Arc Mountains
(e.g., Channing and
Howell, 2006). Examination of MCZ specimens from Tanzania and Kenya that were
assigned to A. adolfifriederici (Barbour and
continue to be reported

Loveridge, 1928: Loveridge. 1942) indicates
that these should be referred to A. affinis. A.

is

doc-

stenodactyliis, A. tanneri,

and possibly

a large

undescribed species from southwestern Tan-


BREVIORA

14

zania

unpublished

(Blackburn,

(MCZ

specimens

data;

One

Material Examined section).

No. 515

see

of these

A-13166) was designated a

paratype oi A. tawieri by Grandison (1983).
In

morphological

of

study

recent

their

variation in populations of A. affinis from

throughout much of the Eastern Arc

Moun-

Loader (2008) were
unable to identify any large Arthroleptis
from eastern Tanzania as morphologically
tains,

Poynton and

similar to the type specimen of A. adolfifriederici that they

examined

(ZMB

21787).

Etymology. Based on the "stately form" of
(Nieden, 1913), it was named in
honor of German explorer Adolf Friedrich,
Duke of Mecklenburg, who led an expedition in the mountains of Central Africa from
this species

1907 to 1908.
Arthroleptis variabilis Matschie, 1893

Variable Squeaker Frog

Figure 6
Arthroleptis dispar. Peters (1875): 210, plate

Arthroleptis variabilis: Matschie

Syntypes:

ZMB

Southwest
009°14'E,

Cameroon.

(85 specimens):

Buea,

Province,

~

950

3.

(1893): 173.

ca.

04°09'N.

m elevation.

Arthroleptis Seimiindi: Boulenger (1905): 180.
Arthroleptis variabilis: Boulenger (1906): 320.
Arthroleptis Seinnindi recognized as a junior

synonym of

Arthroleptis variabilis.

Arthroleptis (Arthroleptis) variabilis: Laurent

Abroscaphus

By

variabilis:

35.0),

Reference Sample. Type material, sex, and

mm)

(in

MCZ

indicated

in

parentheses.

AA-46985
(male?, 35.0) A-1 36744 (female, 35.8), A136775 (female, 28.7), A- 136777-79 (females.
29.0, 29.0, 29.4), A- 136820 (female, 34.6), A136823-25 (females, 31.8, 29.6. 29.4). A136827 (female, 33.5). A-1 36830 (female.

Cameroon:
3428-29

6.

Arthroleptis vanuhilis

(MCZ A- 136830)

A-2654 (female,

(females,

in

mm.

Laurent (1957): 275.

implication.

SVL

Figure

dorsal (A) and ventral (B) views. Scale bar, 10

(1940): 85.

30.5,

29.4),

34.6),

A-137297

563684 (female,

(female,

35.6);

USNM

563685 (male?, 26.3),
563686 Guvenile, 20.8). 563688 (male. 31.0)
563689 (female, 37.8); UTA A-35924 (female, 32.3), A-35933 (male, 32.1), A-35940
37.8),

36.0), A-35949-50 (females, 34.7,
A-44447 (female. 35.4), A-44451 (fe-

(female,
37.4),

male, 37.7);
36.7),
32.1).

ZMB

70085-86

15206 (syntype; female,

(syntypes;

females,

30.8,


NEW CAMEROONIAN

2009

A

Diagnosis.

medium-sized

Avthrolcptis

characterized by a dark-gray throat thai

is

bisected by a prominent, white gular stripe

6B) and a fourth finger that

(Fig.

is

approx-

imately the same length as or shorter than

and second fingers. This species
from nearly all other similarly sized or
larger Avthrolcptis by the presence o{ the
the

fust

differs

white gular stripe (incipient
stripe

can be obscured

in A. nikcucY. this

in A.

variahilis

males

with darkened throats. Arthroleptis variahilis
differs

SVL >

following ways

the

in

medium

from other

Avthrolcptis

large

to

mm): from

(maximum

by lacking
supernumerary tubercles on the feet; from A.
fvancci and A. stenodactylus by crus length
much greater than 50% SVL; from A. nikeac
35

A. ajfiuis

by smaller adult body size (> 50 mm SVL in
A. nikeac); from A. kvokosua by smaller body
size and generally lacking large well-defined
black spots on the lateral surface of the
body; from A. pcvvcti by relatively shorter
snout and fourth finger and by lacking the
distinctive pattern of many white spots on
the darkened posterior thigh; from A.
stenodactylus by an inner metatarsal tubercle
length usually less than

80%

first

toe length;

from A. tanncvi by a relatively narrower head.
Descviption. Medium-sized, robust species
with moderately robust limbs (Fig.

mean male SVL 31.1
mean female SVL 33.1

ble 3);
3.1),

Ta-

6;

mm (n = 4;
mm (n = 27;

±
±

head somewhat broad; head length 80
width; head width 38^5% SVL;
posterior margin of head not always distinct
mm
from body; snout projecting less than
beyond lower jaw; rostral tip rounded in
3.0);

85% head

1

dorsal and lateral views; in dorsal view, eyes
projecting

beyond

visible;

eyes

beyond

lateral

eyelids such that pupil just

close

to

jecting slightly or well

of head
larger

rounded

in

than

or just

projecting

margins of head; eyes pro-

lateral

above dorsal surface

view; eye diameter just

interorbital

distance;

pupil

to horizontally elliptical in preser-

ARl IIROLEniS

TaMI.I: 3.

15

MEASUKrMr.NTS (mm)

A nillHOI V.I'IIS

I

I

OI- II-MAI.I- SYNT'il'l-S oi-

Rl

)

nil. IS.


A

B RE 10 R

16

No. 515

I

Skin of dorsal surface of head and body
smooth, sometimes weakly granular posteriof fore and hind limbs smooth;
skin of lateral body glandular; most of
ventral surface very smooth, but tending
orly; skin

toward glandular posteriorly; median skin
raphe ranging from indistinct to indiscernible in preservation; smooth skin surround-

length

of

fingertips

uniformly

regions

suborbital

darker

than dorsal surfaces; variably present dark
band extending from ventral eye to lip of

upper jaw, with loreal region much lighter;
prominent dark-brown or black supratympanic band extending from posterodorsal

margin of

orbit,

extending posteroventrally

and terminating posterior

to

tympanum or
tympanum

even extending posterior to arm;

ing cloacal region.

Limbs and

and

developed; relative

digits well

fingers:

III

>

I

-=

II

>

IV;

rounded and often very swollen
with

rounded,

not expanded;
prominent, globular, single subarticular tubercles; palmar and metacarpal tubercles
finger

but

translucent and typically lighter than supra-

tympanic

band;

coloration

markings

dorsal

dorsal

as

base

similar

but

coloration

shades darker; some specimens with scat-

and projecting from surface of hand; webbing on manual digits absent; thigh length
ranges from 78 to 102% of crus length (90-

spots; usually two fairly
dark-brown bands on forearm;
dorsal surface of hands mottled light and
dark brown or gray; variably present color
morph in which all lateral surfaces distinctly
darker and different color than dorsal
with sharp boundary between
surfaces,

98%

regions; variably present, usually continuous,

ranging from weakly developed (mostly in
old preserved material) to very prominent

thigh length in majority of specimens);

>

relative length of toes: IV
I;

toe tips slightly

expanded

III

>V>

II

>

to just greater

than width of toe at interphalangeal joints;
toes with prominent, single, rounded, and
globular

subarticular

tubercles;

webbing

small light

tered

complete

white vertebral line extending from snout
to vent; other dorsal markings,

if

tip

present,

comprising dark triangle between

usually

eyes (apex directed posteriorly) often, but

not necessarily, confluent with broad circuor diamond-shaped dark mid-

between pedal digits absent; very prominent,

lar, elliptical,

flange-like inner metatarsal tubercle, length

dorsal marking located over suprascapulars;

ranging from 55 to 120% of

mid-dorsal marking often, but not necessar-

Measurements. Table

first

toe length.

ily,

3.

confluent with circular or elliptical dark

Coloration in Alcohol. This species exhibits

posterior dorsal marking, located over sa-

remarkable diversity of coloration and
patterns that was first noted long ago (e.g.,
Andersson, 1905; Matschie, 1893; Noble,

crum, poorly defined along most posterior
margins; dorsal markings sometimes discontinuous or reduced to poorly defined or even

The following description of colorais a summary based on the reference

paired blotches symmetrical across midline;

a

1924).

tion

specimens, but

it

is

premature to consider

it

exhaustive.

less

Base color of dorsal and
limbs,

grayish

(Fig. 6A); iris

body with scattered well-defined
dark-brown spots and, in some cases, small,
lateral

brown

to

lateral

body, and

brownish

gray

dark gray or black with white

defined light spots;

uniform
usually

coloration

in

with

hind limbs

fairly

and pattern; thigh

two dark transverse

stripes;

dorsal and posterior thigh fairly uniform in

or creamy pupil; usually solid interorbital

coloration

and

bar medium to dark brown; snout usually
uniformly lighter than interorbital bar and
typically similar to remaining dorsum; loreal

sometimes

three,

brown

stripes;

of foot

at

pattern;

crus

with

two,

pronounced transverse
dark spot on lateral margin

base of metatarsal V; dark region


NEW CAMEROONIAN

20(W

ARTHROl.EPTIS

covering cloaca, well defined dorsally and

patterns

extending onto and becoming

ventral

less distinct

on

Throat

light to

dark gray with very small,

and coloration with that of the
which is fairly uniform

surface,

among

posterior thigh.

specimens.

MCZ

(e.g.,

17

One unusual

A- 136827)

morph

color

exhibits

large

a

irregularly placed light spots (Fig. 6B); light

immaculate

spots at jaw margin very irregularly placed;

posterior to the eye, that extends posterior

well-defined, usually continuous, white stripe

from

extending

rostral

along

tip

throat

becomes dominat-

ed by large light spots with dark pigmentation

becoming

lighter

posterior

posteriorly;

more

and

venter

diffuse

mostly white

or very light gray with no markings; medial
surfaces of

or

light

arm and forearm

white, creamy,

dark

surface

gray;

ventral

of

forearm; palmar surface and fingers light to

dark gray; palmar, metacarpal, and subarticular tubercles unpigmented; ventral fingertips lighter in color

than

rest

of finger; venter

just

below

and

onto the fore limb along the dorsal surface to
just distal to the elbow.

midline to level of pectoral girdle; posterior
to pectoral girdle pattern

band,

white

Habitat and Range.

common

lowland forests

in

This typically

known from through-

out Central Africa, with localities

Amiet,

(e.g.,

Bohmeand
1905,
al.,

a

Amiet, 1975). Ar-

(e.g.,

throlepti.s variai^ilis is

oon

is

species that occurs predominantly

Nieden,

Lawson,
1908;

Mertens,

1905;

Schneider, 1987; Boulenger 1900,

1906; Gartshore,

2005;

Camer-

in

Andersson,

1975;

Herrmann

Matschie.

1966;

Perret,

et

1893;

and

Perret

Central African

1957),

(Joger, 1990),

1986;

1993;

Republic

Bioko Island (e.g., Boulenger,
mainland Equatorial Guinea

and ventral thigh both cream colored with
few scattered brown melanocytes; lightcreamy or light-gray ventral surface of thigh

1900,

1905),

(e.g.,

De

with few markings; ventral surface of crus

Nigeria

mottled dark gray and cream; plantar surface

Schiotz, 1963), the Democratic Republic of

homogeneous dark gray or black; subarticular tubercles on toes and inner metatarsal

Congo

tubercle lighter shades than surrounding skin.

of

Coloration
burn's field

Based on D. C. Blacknotes and photographs. Dorsal
in Life.

base coloration black, dark gray, dark,

light,

Riva,

la

et al.,

(Kinshasa;

Laurent, 1972; No-

e.g.,

Congo

(Brazzaville;

e.g.,

Remarks. Arthro/eptis

variabilis

generally dark gray to black with white or

are

(Rodel and Branch, 2002).

light- to

dark-gray spots; inguinal

region ranging from pale yellow to orangey red.
Variation.

umented

in

Measurement variation
Table

3.

No

is

doc-

relationship

is

obvious between the great diversity of dorsal

and

is

some-

times stated to occur in the forests of western

creamy

gray with

Largen

Dowsett-Lemaire, 1991).

Africa

along midline; venter white or

data;

de Witte, 1934), and the Republic

and ruddy brown, red, or olive green;
markings on dorsal and lateral surfaces
ranging from tan, orange, light or dark
brown, and even black; mid-vertebral stripe,
when present, ranging from gray to orange;
lighter markings on lateral surface ranging
from white to gray; ventral surface of throat
stripe

(e.g.,

2006), eastern

unpublished

(Blackburn,

ble, 1924;

Gabon

1994),

Boulenger, 1905; Burger

Global Amphibian Assessment,

(i.e.

2006). However, the western African populations

probably

several

undescribed

sp. 2":

Ernst

et al.,

correspond

to

one

2008; Rodel and Branch,

2002; Rodel and Ernst, 2004; Rodel
2005). Populations
similar

to

southwestern
described

A.

et

al.,

from western Ivory Coast

A.

variabilis,

A

but

smaller

specimen from

Ghana (Rodel et al., 2005) was
as a new species, A.

recently

krokosua, by Ernst
perreti,

or

{^"Arthroleptis

species

et

al.

(2008).

Like A.

krokosua also lacks the white

gular stripe present in A. variabilis and has a


BREVIORA

18

fourth

shorter

finger

second fingers (Ernst

than

the

et al., 2008).

first
and
However,

No. 515

countries
1973,

morphological and molecular data indicate

2004a,b;

that these taxa are neither conspecific nor

al.,

sister

taxa

Ernst

et

unpublished data;

(Blackburn,

al.,

2008).

The many taxonomic

problems of Art/iroleptis from western Africa
are detailed by Rodel and Bangoura (2004).
Etymology.

The name

derives

from the

Latin word varius meaning different, and in
the

adjectival

presumably

form

variabilis,

in recognition

changeable,

of the wide range

of coloration and pattern exhibited by this
species.

Amiet,

(e.g.,

Blackburn,

2000,

Differences

between A.

and

is

1980;

et al.,

200 described species.
concentrated

Much

of this diversity

mountains of the
Line, especially Mt.

the

in

Cameroon Volcanic
Manengouba (Amiet,

1971a,b,

1973,

documented in this study are
found by Perret (1966). In
general, he found A. variabilis to be more
robust and to have a relatively shorter snout,
variabilis

similar to those

relatively shorter crus, relatively larger inner

metatarsal tubercle, a prominent white stripe

on the midline of the throat, and gray or
brown coloration on the posterior surface of
the

Arthroleptis

thigh.

adolfifriederici,

amphibians

Gartshore,

that

is

brown with

with A. perreti as de-

scribed here. Although superficially similar,
A. perreti

and A.

cally distinct.

morphologi-

Molecular phylogenetic data

indicates that,

two

variabilis are

while closely related, these

species are not sister taxa (see Arthro-

leptis

sp.

nov.

1

in

Blackburn,

of

1986;

et

(e.g.,

2006).

al.,

Manen-

the need for conservation

measures focused on the preservation of the
unique biodiversity found on this and other
mountains of the Cameroon Volcanic Line.

as

light spots. Perret's (1966)

agrees

attention

Gonwouo

Description of A. perreti from Mt.

MATERIAL EXAMINED

on the throat and has a posterior thigh

description

especially

1986), but has received

conservation

gouba highlights

1977,
1980).

Amiet, 1975; Blackburn,

(e.g.,

limited

described by Perret (1966), lacks the white
stripe

Cameroon,

2008b; Gartshore,
only

al.,

an important center of

is

within

diversity

A.

et

Lawson, 2000;
Loumont and Kobel, 1991; Rodel et al.,
2004). This region is a hotspot of amphibian
diversity within sub-Saharan Africa (Global

Amphibian Assessment, 2006), and the
Cameroonian amphibian fauna is one of
the most diverse in the world, with nearly
2004; Kobel

This mountain

perreti

2001,

Herrmann

2008b;

1980a; Blackburn, 2008b; Kobel et

DISCUSSION

1972a, b,

1971a, b,
1981,

1980a,b,

1977.

2008a).

Type

material,

sex,

and

SVL

(in

mm)

indicated in parentheses. Arthroleptis affinis:

Tanzania:

MCZ A-13145-146 (females,

37.2,

A-13150-151 (males, 23.4, 25.3), A13152-153 (females, 35.8, 40.5), A-13157160 (females, 32.8, 33.7, 34.7, 36.6), A13162-163 (females, 36.6, 35.7), A-13165
(female, 35.6), A-13167 (male, 27.8), A37.6),

Guided by the description and redescriptions

13169 (male, 29.2), A-25400^01

presented here, future studies should focus

35.5,

30.5),

on determining or re-evaluating the presence
of A. perreti at other montane localities in
Cameroon.
The past four decades have witnessed a
dramatic increase in the described amphibian
diversity in Cameroon and neighboring

33.8,

42.9,

37.4);

38.8);

ZMB

23093 (holotype; female,

A-138241-244

TNHC

(females,

(female,

38577

36.2,

(female,
35.6).

BMNH 1954.1.
32.4); MCZ A-

Arthroleptis francei: Malawi:

13.74

(paratype;

27474-476
28.0),

female,

(paratypes;

A-27477^78

males,

25.4,

(paratypes;

28.2,

females.


NEW CAMEROONIAN

2009

38.7, 41.0), A-27479 (holotype; female, 41.9),
A-137038 (female, 37.3): TMP 48089 (female, 43.0), 48092 (male, 27.8). Arthrolcplis
kwkosua: Ghana: SMNS 12555 (hololype:

male, 43.3).

BMNH

Arthrolcptis iilkeac:

2002.101

(hololype:

2002.102 (paratype: female,

Malawi:

stemxiactylus:

(female,

BMNH

(female,

52.5),

53.7). Arthrolcptis

MMB

HA2()()2.1.13

HA2002.1.60

37.0),

HA2002.4.17

Tanzania:

female,

(male,

38.4).

2002.596 (female, 33.0);

32.7),

Tanzania:

CAS

168455

MCZ

ARIII KOLIiniS

CO/M N F F/SG/D FA P/S AN 05121 6/PR BS/
1

,

Summer Research Funds, a Putnam
Museum of

Goelet

Expeditionary Grant from the

Comparative Zoology (Cambridge, MassaU.S.A.), NSF grant EF-0334939
(AmphibiaTree) to D. Cannatella and J.
Hanken, and Project Exploration, a not-for-

chusetts,

educational

profit

(female 40.0), A/4251/4 (male, 31.0), A/4401/3

with

Bonneaud

Tanzania:

41.9),

CAS

168825 (female; 54.7);

168823 (male;

MCZ

A-13166

(paratype; female; 36.5). Arthrolcptis variabihs

(specimens not included

Guinea:

CAS

in

PCA): Equatorial

DC,

207824-826 (females,

32.7, 33.4, 31.0),

207827

(female?, 26.9), 207828 (female, 35.9). Arthrolcptis sp.:

Tanzania:

MCZ

(Washington,

International

translated

French. C.

German, and

from

translations

the

abstract

Walker (Harvard

S.

Map

1.

Collec-

J.

draft of this manuscript.

LITERATURE CITED
Ahl, E. 1939 "1938". Beschreibung neuer afrikanischer
Frosche der Gattung Arthroleptis. Sitzungsberichte
der Gesellschaft naturforschender Freunde Berlin.
1938: 303-310.

A- 16952

(female,

44.4).

Amiet.

Especes nouvelles ou mal connues de

J.-L. 1971a.

Leptodactylodon (Amphibiens Anoures) de
sale

camerounaise.

Annales

de

la

ACKNOWLEDGMENTS

Sciences du

Many

people assisted with the fieldwork,

(Amphibiens Anoures). Annales de

logistics,

or both needed to collect specimens

Sciences

.

for this study: K. S. Blackburn,

Che,

J.

L. Diffo,

C.
into

Hanken and R. C.
Drewes provided valuable comments on a

tion) created Figure

207817-819 (females, 34.0, 30.6,
(female?, 26.2), 207821-822

(female, 33.3, 36.1), 207823 (juvenile, 25.3),

(Chicago,

was supported by

U.S.A.). D. Hewitt provided assistance

207820

35.4),

organization

LNG

U.S.A.).

Illinois,

(female, 25.7), A/4401/6 (female, 26.2). Arthroleptis tanneri:

was

funded by the Department of Organismic and
Evolutionary Biology (Harvard University),

Conservation

NMK

DCB

MINFOF/SG/DFAP/SDVEF/SC).

A-21715 (male, 27.6), A25387 (female, 28.6), A-27447 (female, 39.5).
A/4251 (male. 33.0), A/4251/3
Kenya:
(male, 35.6);

19

M. Che, V.

D. Fotibu, N. Gawani,

P.

Huang, M. LeBreton, and M. T. Kouete.
The people of Nsoung and the Fulbespeaking community in the Mt. Manengouba crater deserve our sincere gratitude
for their hospitality and allowing use of their
buildings. The Cameroonian ministry of
forestry and wildlife (MINFOF, formerly
MINEF) supplied research and collection

.

Cameroun,

5:

la

Faculte

Dordes

57-81.

1971b. Leptodactylodon nouveaux du
la

Cameroun

Faculte des

du Cameroun, 7-8: 141-172.

1972a. Description de cinq nouvelles especes

camerounaises de Cardioglossa (Amphibiens Anoures). Biologia
.

Gabonica,

8:

201-231.

1972b. Description de trois Bufonides orophiles

du Cameroun appartenant au groupe de Bufu
preussi Matschie (Amphibiens Anoures). Annales

de

la

Faculte

des

Sciences

du Cameroun.

11:

121-140.
.

1973. Caracteres diagnostiques de Petropedetes

peneti. nov.

sp.

et

notes sur

camerounaises. Bulletin de

les

autres especes

I'lnstitut Franijais Afri-

que Noire. 35: 462-474.
.

1975. Ecologie et distribution des amphibiens

(0173PR/MINEF/SG/DFAP, 0588/
PRBS/MINFOF/SG/DFAP/SDVEF/SC)and

anoures de

la

region de

Annales de

la

Faculte des Sciences de Yaounde, 20:

permissions to export specimens (0850 54/

33-107.

permits

Nkongsamba (Cameroun).


BREVIORA

20

—

1977. Les Astylostennis

.

du Cameroun (Amphib-

Anura, Astylosterninae). Annales de

ia

des Sciences de Yaounde, 23-24: 99-227.

—

Revision

1980a.

.

and long-fingered
Linnean Society.

Faculte

la

No. 515

27: 69-224.

plates.

Un

1980b.

Hyperoliiis

Hyperolius endjaiui
peroliidae).

—

.

sp.

Dorsale camerounaise:

Anura,

phibia,

.

Annales

de

.

.

1987. Aires disjointes et taxons vicariants chez

Anoures du Cameroun: implications

les

matiques. Alytes,

—
—

.

6:

Un nouveau

2001.

.

.

Une

nistik

Cameroun (Amphibia,
17: 125-164.

de

Lepiopelis

A

2004b.

pour

la

zone

la

Hyperoliidae). Re-

S.

Barbour,

T..

F. Dallmeier (eds.). Bulletin

Channing,

Washington no.

and K. M. Howell.

a.,

New

of East Africa. Ithaca,

G. 1905. Batrachians from Cameroon

collected by Dr. Y. Sjostedt in the years 1890-1892.

Arkiv for Zoologie,

&

G. Pauwels,

Biological Society of

propos de deux Leptopelis nouveaux

faune du Cameroun (Anura. Hyperoliidae).

L.

mit Beschreibung einer neuen

(III)

Complex of protected areas. Gabon, pp. 297-307.
//; Gamba. Gabon: Biodiversite d'une foret equatoriale africaine. A. Alonso. M. E. Lee. P. Campbell, O.

Alytes. 21: 111-170.

Andersson,

Schneider. 1987. Zur Herpetofau-

B.

Kameruns

ologische Beitrage, 38: 241-263.

vue Suisse de Zoologie, 11: 567-583.
.

2:

1-29.

1

and A. Loveridge.

1928.

la

A

comparative

12.

2006. Amphibians

York, Cornell Univer-

Riva,

1994.

I.

anuros del Parque
Rio Muni, Guinea
Espanola Herpetologie, 8:

Anfibios

Monte

de

Alen,

Revista

Ecuatorial.

123-139.

and Usambara Mountains, Tanganyika Territory
with descriptions of new species. Memoirs of the
Museum of Comparative Zoology, 50: 87-265, 4

Drewes, R. C, and

plates.

Ernst, R., A. C. Agyei, and M.-O. Rodel. 2008.

Blackburn, D. C. 2005. Cardioglossa liheriensis Barbour & Loveridge 1927 is a junior synonym of

(Chabanaud

frateradus

1921).

African Journal of Herpetology, 54: 171-179.
.

and

history

squeakers

of African frogs:

(Arthroleptis)

and

phylogeny of

long-fingered

frogs

(Cardioglossa) estimated from mitochondrial data.

Molecular Phylogenetics and Evolution, 49: 806-826.
.

2008b.

A new

Anura:

ia:

species of Cardioglossa

Arthroleptidae)

Manengouba

in

endemic

the Republic of

Journal of the

Mount

Cameroon, with

an analysis of morphological diversity
Zoological

(Amphib-

to

in the

genus.

Linnean Society, 54:

611-630.
.

In

press.

V. Vindum. 1994. Amphibians of

J.

nal of African Zoology, 108: 55-70.

giant

species

and evolution of male

secondary sexual characters

in

A new

of Arthroleptis (Amphibia: Anura:

Arthroleptidae) from the Krokosua Hills Forest

south-western

Reserve,

Ghana.

Frost, D. R. (ed.)

World:

Zootaxa,

1697:

Lawrence,

Amphibian Species of

1985.

A Taxonomic
Kansas,

African squeakers

the

and Geographical Reerence.
Association

of

Systematics

Collections and Allen Press. 732 pp.

Frost, D. R. (ed.) 2007. Amphibian species of the world:

an online reference

[Internet].

Version

5.

1

.

New York,

American Museum of Natural Histoi7 [cited
August 2008]. Available from: h.
1

org/herpetology/amphibia/index.php.

Frost. D. R.. T. Grant.

J.

Faivovich, R. H. Bain, A.

Haas, C. F. B. Haddad, R. O. de
Diversity

Uganda. Jour-

the Impenetrable Forest, southwest

58-68.

2008a. Biogeography and evolution of body size
life

of the

436 pp.

418 pp.

sity Press.

De

Nacional

plate.

study of the herpetological faunae of the Uluguru

Phryuohatrachus

17:

Burger, M., O. S. G. Pauwels, W. R. Branch, E. Tobi,
J. -A. Yoga, and E.-N. Mirolo. 2006. An assessment of the amphibian fauna of the Gamba

nouvelle espece d' Hyperolius du

Cameroun (Amphibia, Anura,

—

Mr. G. L.

Magazine of Natural History,

BoHME, W., AND

paleocli-

camerounaise (Amphibia, Anura, Hyper-

2004a.

16: 180-184.

new batrachians discovBates in South Cameroon.

317-323.

oliidae). Alytes, 19: 29-44.

—

batrachians in

tailless

Museum. Annals and

Cardioglossa (Anura: Arthroleptidae). Bonner Zo-

Anura, Hyperoliidae). Alytes,
forestiere

new

Annals and

99-115.

2000. Les Alexteroon du

with

species. Proceedings

1906. Descriptions of

ered by

la

Faculte des Sciences de Yaounde, 28: 117-131.

—

1905. Descriptions of

Magazine of Natural History,

la

(Am-

schioetzi nov. sp.

C.

Arthroleptinae).

new genera and

the collection of the British

87: 445-460.

nouvelle Cardioglossa orophile de

of the batrachians and

of the Zoological Society, London, 1900: 433^56, 6

nouveau du Cameroun:
(Amphibia Anura. Hy-

Revue Suisse de Zoologie,

Une

1981.

n.

list

Gaboon (French Congo),

of the

reptiles

descriptions of

.

A

BouLENGER, G. A. 1900.

du genre Leptudactyloclcm

Andersson (Amphibia. Anura, Astylosterninae).
Annales de la Faculte des Sciences de Yaounde,

—

frogs. Biological Journal of the

S.4, A. Channing,
Donnellan, C. J. RaxA. Campbell, B. L. Blotto, P. Moler,

M. Wilkinson,
woRTH'i

,

J.

S.

C.


NEW CAMhROONlAN

2009

R. C. Drivvis. R. a. Nussbaum,

M. Green, and W.
amphibian

Museum

of

tree

C.

life.

D. L>nc

J.

Whehii:r.

Bullclin of the

of Natural History. 297:

I).

ii.

2006.

Tlic

American

291.

1

204-240.

1986.

///

Stuart, S.

Cameroon Montane

N.

(ed.).

Forests.

Conservation of

London. Internation-

Council for Bird Preservation. 263 pp.
Global Ami'human Assessment. 2006. Washington,
al

D.C.,

IUCN/SSC-CT/CABS; c2006

[cited

—

1

August

2008] Available from: htlp://wvvw.globalamphibians.

Remarques sur le genre Scliouieilenella
du Musee Royal du Congo Beige,

1954.

.

21

Witte. Annales

Tervuren.

1:

34-40.

Notes sur les lyperoliidae. Re\ue de
Zoologie et de Botanique Africaines. 56: 274 282.
1957.

.

The status ol" the montane
herpetofauna of the Cameroon highlands, pp.
M.

GartshorI',

ARTHROl.EPTIS

—

I

Amphibians. Exploration du Pare Na-

1972.

.

tional des Virunga. institul National Conservation

de
1

—

Nature de

la

la

Republique du Zaire, 22: 3-125.

plates.

1

1973.

.

The

natural classification of the Arthro-

ogie et de Botanique Africaines, 87: 666-678.
p. 1993. The
Korup National Park

Lawson, D.

org.

GoNwouo,

M. LeBreton, C. Wild, L. Chirio, P.
Ngassam, and M. N. Tchamba. 2006. Geographic
L. N.,

and ecological distribution of the endemic montane
chameleons along the Cameroon mountain range.

A new species

of Arihro/cpil.s

(Anura: Ranidae) from the West Usambara
tains,

Moun-

Museum

Tanzania. Bulletin of the British

of

W. Bohme.

A.

P.

A

2004.

Cardioglossa

from

Herrmann, A. Schmitz. and
new frog species of the genus

Mbabo

Tchabal

the

Project.

tological Natural History,

1:

and amphibians of
Cameroon. Ilerpe-

27-90.

A

new eaecilian from Cameroon, Africa
(Amphibia:
Gymnophiona: Scolecomorphidae).
.

2000.

Leviton, a. E.. R. H. Gibbs. Jr.. E. Heal,

Dawson.

Standards

1985.

ichthyology: part

I.

Mtns,

Cameroon. Herpetozoa, 17: 119-125.
W. Bohme, P. A. Herrmann, M. Plath, A.
Schmitz, and M. Solbach. 2005. African biodiversity hotspots: the amphibians of Mt. Nlonako,

Cameroon. Salamandra,

lUCN

red

categories

and

criteria

Version 3.1. Gland. Switzerland, lUCN
August 2008]. Available from: http://www.

[Internet].
[cited

1

Loveridge,

a.

1942.

Scientific

results

of a

1990.

The herpetofauna of

the

African Republic, with description of a
Rhinotyphlops

of

85-102.

//;

(Serpentes:

G. Peters,

&

new

Central
species

Typhlopidae),

pp.

R. Hutterer (eds.), Verte-

brates of the Tropics. Berlin,

Museum Alexander

Koenig.

KoBEL. H.

R..

L.

Dv

Pasql'ier,

M. Fischberg. and

H. Gloor. 1980. Xenopiis amieti
Pipidae)

sp.

nov. (Anura:

from the Cameroons, another case of
Revue Suisse de Zoologie, 87: 919-

tetraploidy.

926.

Bulletin of the

Africa.

Zoology, 41: 377^36, 4

Museum

&

fourth

Central

of Comparative

plates.

1953. Zoological results of a fifth expedition to

East Africa. IV. Amphibians from Nyasaland and
Tete.

Bulletin

of the

Museum

Zoology, 110: 325-406, 4
.

U.

East

expedition to forested areas in

ucnredlist.org.

JoGER,

and

Loumont, C, and H. R. Kobel. 1991. Xeiwpiis longipes
sp. nov., a new polyploid pidid from western
Cameroon. Revue Suisse de Zoologie, 98: 731-738.

41: 61-81.

list

and

ichthyology. Copeia, 1985: 802-832.

.

2001.

C. E.

Standard symbolic codes for

,

lUCN.

and

herpetology

in

institutional resource collections in herpetology

Natural History (Zoology), 45: 77-84.

Herrmann. H.-W.,

reptiles

Herpetologica. 56: 77-80.

Salamandra, 42: 213-230.

Grandison, A. G. C. 1983.

Revue de Zool-

leptinae (Amphibia, Hyperoliidac).

1957.

Check

East

Africa

of

list

of Comparative

plates.

of the reptiles and amphibians

(Uganda;

Zanzibar). Bulletin of the

Kenya;

Museum

Tanganyika;

of Comparative

Zoology, 117: 153-362, i-xxxvi.

Matschie,

p.

1893. Einige anscheinend neue Reptilien

und Amphibien aus West-Afrika. Sitzungsberichte
der Gesellschaft Naturforschender Freunde zu

Berlin, 1893: 170-175.

Matsui, M. 1984. Morphometric variation analysis and
revision

of

Bufonidae).

the

Japanese

Contributions

toads

from

(genus
the

Biifn.

Biological

Laboratory, Kyoto University, 26: 209^28.

M. J., AND F. Dowsett-Lemaire.
Amphibians (Anura) from the Kouilou River

Largen,

1991.
basin,

Myers. N., R. A. Mittermeier, C. G. Mittermeier, G.
A. B. da Fonseca, and J. Kent. 2000. Biodiversity

Republique du Congo. Tauraco Research Report.

hotspots for conservation priorities. Nature, 403:

4:

145-168.

Laurent. R.

853-858.

F. 1940. Contribution a I'osteologie et a la

Nieden, F. 1908. Die Amphibienfauna von Kamerun.

systematique des ranides africains - Premiere note.

mit

Revue de Zoologie

dem

74-97. 3 plates.

et

de Botanique Africaines, 34:

einer

Bestimmungstabelle.

Zoologischen

491-518.

Museum

Mitteilungen
in

Berlin,

aus
1908:


BREVIORA

22

1911 "1910".

.

dem

Kgl. Zool.

Neue

ostafrikanische Frosche aus

Museum

Freunde

Nalurforschender

Gesellschaft

Sitzungsberichte

in Berlin.

Berlin,

Amphibia. Wissenschaftliche Ergebnisse

1913.

.

RoDEL, M.-O., AND M. A. Bangoura. 2004. A conservation assessment of amphibians in the Foret

Zentral-Afrika-Expedition,

Deutschen

4:

165-195.

a

new Amnirana

AND W. R. Branch.

of the Haute
245-268.

Museum

amphibian

1966.

amphibiens du Cameroun.

Les

Zoologische Jahrbucher (Systematik),

Le

1991.

.

8:

289-464.

AND R. Ernst.

I:

2004. Measuring and monitoring

diversity

tropical

in

F.

Arthroleptidae).

de

la

Bulletin

Salamandra, 41: 107-127.

AND R. Mertens.

herpetologique

1957. Etude d'une collection
au Cameroun de 1952 a 1955.

faite

Bulletin de Tlnstitut Frangais Afrique Noire, 49:

548-601.

W.

1875.
in

An

M. Gil, A. C. Agyei. A. D. Leache, R. E. Diaz,

M. K. FujiTA, AND R. Ernst. 2005. The amphibians
of the

Buchholz

I.

standardization. Ecotropica, 10: 1-14.

hivit tarns

71-76.

Peters,

forests.

evaluation of methods with recommendations for

Societe neuchateloise des Sciences naturelles, 114:

,

Cavally forests, western

amphibians. Salamandra, 38:

,

(Anura,

Miiller

d^Arthroleptis

statut

,

2002. Herpetological survey

Dodo and

Ivory Coast, Part

American Museum Congo Expedition, 1909-1915.
Part III. Amphibia. Bulletin of the American
of Natural History, 49: 147-347, 20 plates.

(Amphibia Anura Rani-

species

dae). Tropical Zoology, 17: 201-232.
,

Noble, G. K. 1924. Contributions to the herpetology of
the Belgian Congo based on the collection of the

Perret, J.-L.

Simandou Range, south-

Classee du Pic de Eon,

eastern Republic oi Guinea, with the description of

1910:436-441.
der

No. 515

Uber

die

von Hrn. Professor Dr. R.

Westafrika gesammelten Amphibien.

Monatsberichte der Koniglichen Preussische Aka-

demie des Wissenschaften zu

Berlin, 1875: 196-212,

A. ScHMiTZ, O.

,

of south-western

parts

forested

S.

Ghana.

G. Pauwels, and W. Bohme.

2004. Revision of the genus Werneria Poche, 1903,

new species from
Cameroon and Gabon (Amphibia: Anura: Bufoniincluding descriptions of two

dae).

Zootaxa, 720: 1-28.
A.

ScHiOTZ,

from

1963.

Nigeria.

On

a

collection

Amphibia

of

meddelelser

Videnskabelige

fra

Kjobenhaven,
Dansk naturhistorisk forening
129: 43^8.
Schmidt, K., and R. F. Inger. 1959. Amphibians,
i

3 plates.

Plath, M., M. Solbach, and H.-W. Herrmann. 2004.
Anuran habitat selection and temporal partitioning

montane

a

in

and

submontane

Southwestern Cameroon

—

first

rainforest

results.

Salaman-

nae.
.

J.

C. 1976. Classification and the Arthrolepti-

Revue de Zoologie

2003a. Arthroleptis troglodytes and the content

A new

2003b.

giant

species

of Arthroleptis

(Amphibia: Anura) from the Rubeho Mountains,
Tanzania.

African

Journal

of Herpetology,

52:

,

and D. G. Broadley.

1985.

Amphibia Zambe-

Scolecomorphidae, Pipidae, Microhylidae,

Hemisidae, Arthroleptidae. Annals of the Natal
1.

Museum, 26: 503-553.
and S. p. Loader. 2008.
,

taxonomic consequences
forest

frog,

517-526.

du

Beige, 56: 1-264, 9 plates.

J.

E.

collections

2002. Herpetological collecting and
management. Herpetological Circular,

31: 1-153.

Skelton-Bourgeois, M.

d'Afrique

orientale.

1961.

batraciens

Reptiles et

Revue de Zoologie

et

de

Botanique Africaines, 63: 309-338.
Stuart,

S.

N. 1984. Conservation recommendations and
(ed.).
/// Stuart, S. N.

conclusions, pp. 241-245.

107-112.

siaca

Congo
Simmons,

Africaine, 90: 215-220.

of Schoutedenella (Amphibia: Anura: Arthroleptidae). African Journal of Herpetology, 52: 49-51.
.

Exploration du Pare National de I'Upemba, Mission G.F. de Witte, Institut des Pares Nationaux

dra, 40: 239-260.

Poynton,

exclusive of the genera Afrixalus and Hyperolius.

in

Clinal variation and

in the

Arthroleptis

its

common Tanzanian

affinis.

Copeia,

2008:

Conservation

of

Cameroon

Montane

London, International Council
tion.

Forests.

for Bird Preserva-

263 pp.

DE Witte, G.-F. 1934. Batraciens recoltes au Congo
Beige par le Dr. H. Schouteden et par M. G.-F. de
Witte. Annales

du Musee Royal du Congo

Tervuren, C. - Zoologie, Serie
plates.

I,

3:

Beige,

157-188, 7