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B RE VI O R A 



M 



HI s e HI iin o 



I v^oimpaFattive 



ogy 



us ISSN 0006-9698 



CAMBRiDCiE, Mass. 



2 February 2009 



Number 515 



A 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 

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 basses plaines environnantes. Du fait de la confusion anterieure entre la nouvelle 
espece et A. adolfifriederici, et du fait de sa ressemblance a A. variabilis, nous decrivons a nouveau ces dernieres deux 



' Natural History Museum and Biodiversity Research Center, University of Kansas. Lawrence, Kansas 66045, 

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

"Cameroon Biodiversity Conservation Society, Yaounde, Cameroon; e-mail: lgonwouo(a yahoo.com 

'Department of Biodiversity Dynamics, Technische Universitiit Berlin, D-12165, Berlin, Germany; e-mail: 

raffael.ernst(a tu-berlin.de 

■* Humboldt University. Museum of Natural History, 10115 Berlin. Germany; e-mail: mo. roedelf« museum. 

hu-berlin.de 



© The President and Fellows of Harvard College 2009. 



BREVIORA 



No. 515 



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



Squeaker frogs {ArtJiroleptis sensii Frost, 
2007) are distributed throughout much of 
sub-Saharan Africa. This genus occurs in 
two biodiversity hotspots deemed conserva- 
tion priorities (Myers et al., 2000) and hkely 
contains high levels of cryptic species diver- 
sity (e.g., Poynton, 2003b; Rodel and Ban- 
goura, 2004). The taxonomic history of 
Artlvoleptis is characterized by long-stand- 
ing disagreements between researchers about 
the status of both species and supraspecific 
taxa (e.g., Laurent, 1940, 1954, 1973; Perret, 
1991; Poynton, 1976, 2003a; Poynton and 
Broadley, 1985; Schmidt and Inger, 1959). 
Typically, these disagreements focus on the 
smallest species, which previously were 
placed in Schoutedenella but now are includ- 
ed within Arthroleptis (Frost et al., 2006; see 
also Blackburn, 2008a). One large species 
that has caused significant taxonomic con- 
fusion 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 estab- 
lished. Over the past century, A. adolfifrie- 
derici has been claimed to occur in Camer- 
oon at the western extent of its distribution 
(e.g., Noble, 1924; Perret, 1966) to the 
Eastern Arc Mountains of Tanzania and 
Kenya at its eastern extent (e.g., Barbour 
and Loveridge, 1928; Channing and Howell, 
2006; Loveridge, 1942. 1957). Specimens 
from Cameroon have long been recognized 
as distinct from A. adolfifriederici; however, 
this taxon has remained undescribed (e.g., 
Amiet, 1987; Frost, 1985, 2007; Gartshore, 



1986; Global Amphibian Assessment, 2006; 
Herrmann et al., 2005). Tanzanian popula- 
tions previously assigned to A. adolfifrieder- 
ici correspond to several currently recognized 
taxa, including Arthroleptis affiuis Ahl, 1939 
"1938," Arthroleptis fa77«^r/ Grandison, 1983, 
and possibly Arthroleptis leleupi Skelton- 
Bourgeois, 1961 (Grandison, 1983; Poynton, 
2003b; Poynton and Loader, 2008), and it 
is unlikely that A. adolfifriederici occurs in 
eastern Tanzania (e.g., Poynton and Loader, 
2008). 

During recent fieldwork on Mt. Manen- 
gouba in southwestern Cameroon, specimens 
of an undescribed Arthroleptis species were 
found. These specimens are distinct from all 
other described Cameroonian species. The 
similarity between these specimens and 
Perret's (1966) description of A. adolfifrie- 
derici from near Nkongsamba, approximate- 
ly 10 km southeast of Mt. Manengouba, 
initiated this study, which describes these 
specimens as a new species. 

MATERIALS AND METHODS 

Specimens were collected during visual 
encounter surveys on Mt. Manengouba in 
the Republic of Cameroon (Fig. 1). Surveys 
were conducted in and near the village of 
Nsoung, as well as near Moabi and Ebone- 
min and in forests near the summit. Animals 
were euthanatized in an aqueous solution of 
chloretone following standard collections 
procedures (Simmons, 2002). Liver tissue 
samples were preserved in 95% ethanol; 
remaining voucher specimens were preserved 
overnight in 10% neutral buffered formalin 
before storage in 70% ethanol. Type material 
was examined of all species discussed, except 
those of Arthroleptis stenodactylus Pfeffer, 
1893, which were destroyed during World 



2009 



NEW CAMEROONIAN ARTIIROLEPTIS 




War II (Frost, 2007). All measurements (± 
0.1 mm) were taken with digital calipers and 
a dissecting microscope. Limb measurements 
were taken on the right side. Measurements 
follow Blackburn (2005), which is a modifi- 
cation 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 
(Synoptics). Museum abbreviations follow 
Leviton et al. (1985), with the addition of 
Museums of Malawi, Blantyre (MMB). 

To determine whether the new taxon 
differs morphologically from the superficial- 
ly similar Arthroleptis variabilis, a multivar- 
iate analysis based on the type specimens was 
used for ordination. Twenty-seven adult 
specimens (including three syntypes) of A. 
variabilis were measured (Appendix 1). Be- 
cause few male specimens have been collect- 
ed of the new species, this analysis is 
restricted to female A. variabilis. Females 
were identified by large body size, the 
presence of ova (visible either in dissection 
or through 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 
by R 2.7.1 for Mac OS X (The R Foundation 
for Statistical Computing, http://www. 
R-project.org). A multivariate ordination 
technique, principal component analysis 
(PCA), was used to analyze patterns of 
variation and covariation within the mea- 
surement 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 of Anlirolcpiis perreti on 
Mt. Manengouba in southwest Cameroon. Star desig- 
nates type locality near village of Nsoung; circles 
indicate paratype localities. 



BREVIORA 



No. 515 



counting for 85% of the cumulative variance 
were retrieved from the analysis. The rela- 
tionship between the two species in morpho- 
space was evaluated by plotting principal 
component (PC) scores. 

DESCRIPTION OF NEW SPECIES 

Arthroleptis perreti^ new species 

Perret's Squeaker Frog 
Figures 2, 3 

Arthroleptis adolfifriederici: Perret (1966): 

396. 

Arthroleptis sp. 7: Gartshore (1986): 220. 

Following personal communication with J.- 

L. Amiet; Amiet (1987): 100. 

Hohtype. MCZ A- 137978 (field number 
DCB 34368), adult female. Republic of 
Cameroon, Southwest Province, near village 
of Nsoung, southwest slope of Mt. Manen- 
gouba,04°59'05.5"N,009°48'41.0"E, 1,400 m 
elevation, 13 July 2006, D. C. Blackburn, K. 
S. Blackburn, and M. T. Kouete. 

Paratypes. MCZ A- 136931 (DCB 34208), 
A-1 36932 (DCB 34209), subadult female and 
juvenile, respectively. Republic of Camer- 
oon, Littoral Province, Mt. Manengouba, 
05°00'38.9"N, 009°51'24.8"E, 2,160 m eleva- 
tion, type locality 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 
2006, D. C. Blackburn, K. S. Blackburn, M. 
T. Kouete; MCZ 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 Prov- 
ince, Mt. Manengouba, near summit, adult 
female (42.0 mm), 05°0rN, 009°52'E, 22 
March 2006, L. N. Gonwouo; ZMB 71472 



(705G), adult male (27.4 mm). Republic of 
Cameroon, Littoral Province, Mt. Manen- 
gouba, 05°00'N, 009°49'E, 2,045 m eleva- 
tion, 6 August 2005, L. N. Gonwouo; ZMB 
71473 (560G), aduU female (39.6 mm), Re- 
public of Cameroon, Southwest Province, 
Mt. Manengouba, near Nsoung, 04°59'N, 
009°48'E, 1,420 m elevation, 4 July 2004, 
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. 
N. Gonwouo; ZMB 71475 (0883N), adult 
female (34.7 mm). Republic of Cameroon, 
Littoral Province, Mt. Manengouba, near 
summit, 05°00'N, 009°51 'E, 2,200 m elevation, 
17 March 2006, L. N. Gonwouo; ZMB 71476 
(0023LG), adult female (39.3 mm). Republic of 
Cameroon, Littoral Province, Mt. Manen- 
gouba, 05°00'N, 009°51'E, 2,185 m elevation, 
28 September 2006, L. N. Gonwouo. 

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

ZMB 71484 (0263LG), juvenile (22.9 mm), 
western fiank, no data available, ~ 1,500 m 
elevation, 20 December 2006; ZMB 71478 
(0899N), aduh female (41.5 mm), near Man- 
engouba lakes. 05°0r49"N, 009°50'39"E, 
2,155 m elevation, 20 March 2006; ZMB 
71479 (282C), juvenile (22.6 mm), near 
Nsoung, 04°59'N, 009°48'E, 1,420 m eleva- 
tion, 4 July 2005; ZMB 71485 (0393N), adult 
female (37.9 mm), near Moabi, 05°10'N, 
009°50'E, 1,980 m elevation, 12 August 
2005; ZMB 71486 (0396N). juvenile (23.6 
mm), near Moabi, 05°10'N, 009°50'E, 1,980 
m elevation, 12 August 2005; ZMB 71487 
(0804N), juvenile (23.1 mm), near Ebonemin, 
05°00'N, 009°44'E, 1,650 m elevation, 18 
November 2005; ZMB 71488 (0992N), juve- 
nile (25.1 mm), near Ebonemin, 05°01'N. 
()09°46'E, 1.465 m elevation, 16 November 



2009 



NEW CAMHROONIAN ARI II ROI.EI'TIS 




Figure 2. Arihrulepii.s pcrrcii in life: holotype (MCZ A-137978) in roslrokitcral view (A), and paratype (MCZ A- 
137980) in ventral view (B). Scale bar, 10 mm. 



2005; ZMB 71489 (0945N), juvenile (25.1 
mm), no data available. ~ 2.085 m elevation. 
13 August 2005; ZMB 71490 (0963N), adult 
female (41.1 mm), no data available, ~ 2,085 
m elevation, 5 September 2005; ZMB 71491 
(88G), near summit, adult female (44.7 mm), 
05°0rN. 009°52'E, 22 March 2006; ZMB 



71492 (0865N). adult female (42.7 mm), near 
summit. ()5°00'N. 0()9°5rE, 2,185 m eleva- 
tion. 15 March 2006; ZMB 71494 (0884N), 
juvenile (21.7 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 



BREVIORA 



No. 515 




Figure 3. Artliroleptis perreti holotype (MCZ A- 
137978) in dorsal (A) and ventral (B) views. Scale bar, 
10 mm. 



elevation, 17 March 2006; ZMB 71496 (31G), 
adult female (35.5 mm), 4°59'N, 009°48'E, 
2,045 m elevation, 6 August 2005; ZMB 
71498 (553G), adult female (40.2 mm), 
05°00'N, 009M9'E, 2,045 m elevation, 6 
August 2005; ZMB 71499 (843C), adult 
female (34.2 mm), 05°00'N, 0()9°49'E, 2,045 
m elevation, 6 August 2005; ZMB 71500 



(0006LG), adult female (38.3 mm), 04°59'N, 
009°50'E, 2,010 m elevation, 25 September 
2006; ZMB 71501 (0007LG), adult male 
(30.4 mm), 04°59'N, 009°50'E, 2,010 m ele- 
vation, 25 September 2006; ZMB 71502 
(0020LG), adult female (37.2 mm), 04°59'N, 
009°50'E, 2,010 m elevation, 25 September 
2006; ZMB 71503 (0022LG), adult female 
(37.6 mm), 05°00'N, 009°5rE, 2,185 m ele- 
vation, 28 September 2006; ZMB 71504 
(0024LG), adult female (35.5 mm), 05°00'N, 
009°51'E, 2,185 m elevation, 28 September 
2006; ZMB 71505 (0027LG), juvenile (22.0 
mm), 05°00'N, 009°51'E, 2,185 m elevation, 
28 September 2006; ZMB 71506 (0044LG), 
juvenile (27.2 mm), 05°02'N, 009°86'E, 2,066 
m elevation, 28 September 2006; ZMB 71507 
(0045LG), adult female (36.8 mm), 05°02'N, 
009°86'E, 2,066 m elevation, 28 September 
2006; ZMB 71508 (0048 LG), adult female 
(43.8 mm), 05°00'N, 009M9'E, 2,184 m 
elevation, 30 September 2006; ZMB 71509 
(0053LG), adult female (27.4 mm), 05°00'N, 
009°49'E, 2,184 m elevation, 30 September 
2006; ZMB 71510 (0182LG), adult female 
(36.8 mm), 04°59'N, 009M7'E, 1,393 m ele- 
vation, 19 November 2006. 

Diagnosis. A medium-sized Arthroleptis 
that is very similar to A. variabilis but readily 
distinguished from that species by lacking a 
midline gular stripe (Fig. 2) and by having a 
relatively longer snout and, in females, a 
fourth fmger that is longer than the first and 
second fingers (see Variation section). In 
addition, the posterior surface of the thigh of 
A. perreti is dark gray with many small white 
spots (Fig. 3), whereas in A. variabilis, the 
posterior surface of the thigh exhibits poorly 
defined light mottling on a dark-brown or 
-gray background. Maximum body size in A. 
perreti (44.7 mm SVL, ZMB 71491) is larger 
than that of other described species of 
Cameroonian Arthroleptis. including A. var- 
iabilis (37.8 mm SVL; Blackburn, 2008a); 
mean male SVL of A. perreti is 29.5 mm (n = 



2(HW 



NEW CAMEROONIAN ARI II ROLIIPTIS 



3; ± 1.8). whereas mean female SVL is 
37.5 mm (n = 25; ± 4.3). Anhroleptis perreti 
differs in the following ways from other 
medium to large Arihrolcpiis (maximum 
SVL > 35 mm): from A. culolfifricdcrici by 
less granular skin, more extensive pigmenta- 
tion on throat, and less globular inner 
metatarsal tubercle; from A. a/finis by 
lacking supernumerary tubercles on the feet; 
from Avthroleptis francei and A. steuodacty- 
his by 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 
metatarsal tubercle length less than 80% of 
first toe length; 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 
globose inner metatarsal tubercle. 

Description of Holotype. Medium-sized 
(37.4 mm SVL), moderately robust female 
with slender limbs (Figs. 2A, 3; Online 
Table 1 ); head somewhat broad, head length 
79% head width; snout projecting approxi- 
mately 1 mm beyond lower jaw; rostral tip 
rounded in dorsal view, blunt and only 
slightly curving posteroventrally in lateral 
view; eyes barely project beyond eyelids in 
dorsal view; eyes placed well medial of, and 
not projecting laterally beyond, margins of 
head; eyes fiush with dorsal margin of head 
in lateral view; eye diameter approximately 
equal to interorbital distance; pupil round to 
slightly horizontally elliptical in preservative; 
loreal region weakly concave; naris small, 
rounded, facing laterally, and barely visible 
in dorsal view; canthus rostralis short, 
indistinct, rounded; eye diameter 1.5 times 



eye-narial distance; eye diameter 2.1 times 
distance from naris to rostral tip; internarial 
region fiat; internarial distance 90% interor- 
bital distance; tympanum distinctly ovoid, 
height slightly less than half the diameter of 
eye; tympanic annulus poorly defined; su- 
pratympanic fold absent; tongue very broad, 
heart-shaped, covered with many small 
pustules; anterior attachment of tongue 
narrow; posterior notch of tongue approxi- 
mately one fourth anteroposterior length of 
tongue; prominent, fiat, fiap-like median 
papilla on dorsal tongue surface near ante- 
rior attachment; choana hidden by maxillary 
shelf in ventral view; premaxillary and 
maxillary teeth present, but hidden by lips; 
vomerine teeth absent. 

Skin of limbs, dorsal surface of head and 
body, and ventral head smooth (more 
tuberculate in Hfe; Fig. 2); skin of lateral 
body glandular, becoming more indistinct on 
ventral surface; median skin raphe promi- 
nent in life (Fig. 2), but very indistinct in 
preservative; cloacal region weakly glandu- 
lar. 

Limbs and digits well developed but 
slender; digits of both manus and pes 
slender; relative length of fingers: III > IV 
> II > I; fingertips rounded, slightly swollen, 
approximately same width as rest of finger; 
finger with rounded, prominent, globular, 
single subarticular tubercles; palmar and 
metacarpal tubercles present but weakly 
developed and barely projecting from surface 
of hand; webbing between manual digits 
absent; thigh length 93% of crus length; 
relative length of toes: IV > III > V > II > 
I; toe tips pointed and slightly expanded to 
approximately same width as interphalange- 
al regions; toes with prominent, single, 
elongate, and slightly conical subarticular 
tubercles; webbing between pedal digits 
absent; prominent fiange-like inner metatar- 
sal tubercle, length 72% of first toe length. 

Measurements. See Online Table I . 



BREMORA 



No. 515 



Coloration of Holotype (in Alcohol). Dor- 
sal ground color silvery brown with small, 
weakly defined gray and well-defined dark- 
brown 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 poste- 
rior to eye; interorbital bar medium brown 
but broken and poorly delimited from other 
brown spots on posterodorsal surface of 
head; dark-brown supratympanic band ex- 
tending from posterodorsal margin of orbit, 
slightly continuous with dark mask of 
suborbital region, extending posteroventrally 
and terminating anterior to and just dorsal 
of arm; tympanum translucent, with silver 
flecks, and 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 anteropos- 
terior 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 much lighter than dorsum grading 
into creamy white of ventral surface; dorsal 
surface of hind limbs grayish brown; very 
indistinct, broken, dark-brown band on 
dorsal surface of thigh; posterior surface of 
thigh dark brown with prominent large white 
spots (Fig. 3A); crus with two prominent 
transverse dark-brown stripes; feet mottled 
gray and brown with poorly defined medi- 



um-brown spot on lateral margin of foot at 
base of metatarsal V; base color of dorsal 
foot silvery and creamy with many small 
brown well-defined melanocytes; general 
trend of lighter and less pigmentation from 
lateral to medial toes; dorsal surface of toe 
tips dark brown; ultimate phalangeal joints 
of toes I-III unpigmented and creamy; 
ultimate phalangeal joints of toes IV V 
lighter in pigmentation than surrounding 
skin; dark-brown triangle with apex directed 
dorsally centered on cloaca. 

Lateral margin of lower jaw dark brown 
with small white spots at regular intervals; 
throat brown with distinct white spots 
(Fig. 3B); posterior throat, just anterior to 
clavicles and laterally bordering omoster- 
num, 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 
tubercles unpigmented; ventral fingertips 
lighter in color than rest of finger; brown 
pigmentation of throat broken into mottling 
over pectoral girdle extending ventrally as 
diffuse brown pigmentation with large, 
poorly defined white spots; venter and 
ventral thigh both cream colored with few 
scattered brown melanocytes; ventral surface 
of crus mottled brown and cream; plantar 
surface homogeneous dark brown; subar- 
ticular tubercles on toes and inner metatarsal 
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 
with 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 
lateral surface ranging from white to gray; 



2009 



NEW CAMEROONIAN ARTIIROI.EPTIS 



\ciitial surface o\' lliroal generally gray vvilli 
tainl creamy xellow lones; venter with more 
pronounced creamy yellow lones; bright 
yellow splotches in inguinal region (Fig. 2B). 

Wtiicition. Meristic \arialion is document- 
ed in Online Table I. The median papilla of 
the tongue is more conical in MCZ A- 
137980. Sexual dimorphism in A. pcrrcti is 
similar to that of other species oWlnhroleptis 
(Blackburn, in press) 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 
line the medial surface of both the second 
(MDIl) and third (MDlll) fingers. The 
number of digital spines is comparable 
between the male paratypes (ZMB 71472, 
MDII right 6. left-8, MDIIl right 17, left- 
18: ZMB 71474, MDII right 13, left 12, 
MDII I right 18, left 20). In the oiie larger 
male paratype (ZMB 71474), pronounced 
white spines are found 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 
first and second fingers. 

Habitat and Range. All specimens were 
collected in montane forest during the day 
(1000-1400 h) when they were active in 
moist leaf litter. The type series comes from 
high elevations (1,400-2,200 m elevation) on 
Mt. Manengouba (Fig. 1). Populations of ^. 
perreti might also occur on several other 
nearby mountains, including Mt. Nlonako, 
Mt. Kupe, and in the Rumphi Hills (Gart- 
shore, 1986; Herrmann et al., 2005; Lawson, 
1993; Perret, 1966). However, specimens 
studied by Lawson (1993) and Herrmann et 
al. (2005) do not correspond to A. perreti and 
represent A. variabilis (i.e., Lawson, 1993) or 
possibly another undescribed species (i.e.. 



Herrmann et al.. 2005). Fven if present at 
these t)ther localities, A. perreti would still 
have an extent of occurrence less than 
2().()0() 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. 
Manengouba, and probably other nearby 
mountains, are declining (Gartshore, 1986; 
Global Amphibian Assessment, 2006; Gon- 
wouo et al., 2006; Stuart, 1984), A. perreti 
should be considered Vulnerable according 
to lUCN (2001) criteria. 

Etymology. This species is named in honor 
of Dr. Jean- Luc Perret who provided prelim- 
inary, but accurate, comments on this new 
species more than 40 years ago (Perret, 1966). 

Morphological Comparison with Arthro- 
leptis variabilis. As indicated in previous 
literature (Perret, 1966), this species is 
extremely similar to A. variabilis yet lacks 
the prominent white gular stripe. These 
species appear to be osteologically indistin- 
guishable on the basis of radiographic 
analysis (Blackburn, unpublished data). 
PCA indicates that several morphological 
characters differentiate A. perreti from A. 
variabilis (Fig. 4). The first principal compo- 
nent axis (PCI) accounts for the majority 
(52.7%) of the variance in the data and is 
taken as a general measurement of body size 
(Table 1); A. perreti and A. variabilis are not 
differentiated along PCI. The second princi- 
pal components axis (PC2) accounts for 
19.4% of the variance and plots of PC2 
scores reveal that A. perreti and A. variabilis 
are differentiate along this axis (Fig. 4). 
Plots of PC3 scores do not reveal differences 
between these two species. PC2 loads most 
strongly, and positively, on inner metatarsal 
tubercle length, eye diameter, and tympanum 
height (Table I). The strongest negative 
loadings for PC2 are on snout length, fourth 
finger length, and fifih toe length. Because 



10 



BREVIORA 



No. 515 



Principal Components Analysis 




Figure 4. Scatteqjlot of the first and second principal 
component scores. The second principal component (PC2) 
is plotted against the first principal component (PCI), an 
indicator of body size. Circles represent Arthrolcptis 
variabilis, squares represent Arthwieptis pencti. 



PC2 scores for A. perreti are lower than all but 
one specimen of A. variabilis, these results 
together indicate that the length of the inner 
metatarsal tubercle, eye diameter, and tym- 
panum height are relatively greater in A. 
variabilis and that the snout, fourth finger, 
and fifth 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 (e.g., Amiet, 
1975; Gartshore, 1986; Herrmann 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 mid A. variabilis, 
this species remained undescribed for more 
than 40 years. This is likely due, at least in 
part, to the general inadequacy of the original 



Table 1. Principal components analysis comparing 

Arthroleptis PERRETI and Arthroleptis variabilis. 
Eigenvalues, percent variance, cumulative variance, 
and loadings for the first three principal component 

(PC) AXES. 





PCI 


PC2 


PC3 


Eigenvalue 


0.278 


0.038 


0.023 


Percent variance 


52.68 


19.44 


15.13 


Cumulative variance 


52.68 


72.12 


87.25 


Loadings 








Snout-vent length 


0.171 


0.117 


-0.219 


Head width 


0.150 


0.121 


-0.171 


Tympanum height 


0.220 


0.224 


-0.446 


Eye diameter 


0.119 


0.166 


-0.552 


Snout length 


0.266 


-0.245 


-0.107 


Forearm length 


0.163 


0.174 


-0.153 


Manual digit 1 


0.205 


0.327 


0.293 


Manual digit II 


0.252 


0.166 


0.135 


Manual digit III 


0.246 


0.072 


0.107 


Manual digit IV 


0.335 


-0.231 


0.053 


Thigh length 


0.210 


-0.061 


-0.235 


Crus length 


0.195 


-0.086 


-0.125 


Pedal digit I 


0.275 


-0.099 


0.315 


Pedal digit 11 


0.256 


-0.124 


0.117 


Pedal digit III 


0,270 


-0.151 


0.069 


Pedal digit IV 


0.256 


-0.177 


-0.007 


Pedal digit V 


0.314 


-0.293 


0.073 


Inner metatarsal length 


0.224 


0.659 


0.267 



descriptions of A. adolfifriederici and A. 
variabilis. Furthermore, recent studies de- 
scribing new species of large Arthroleptis from 
the Eastern Arc Mountains have had to rely 
on only a single specimen oi A. adolfifriederici 
from near the type locality for this species 
(Poynton, 2003b; Poynton and Loader, 2008). 
Redescriptions of A. adolfifriederici and A. 
variabilis are thus provided here. 

Arthroleptis adolfifriederici Nieden, 
1911 "1910" 

Rugege Forest Squeaker Frog 

Adolf Friedrich's Squeaker Frog 

Figure 5 

Arthroleptis adolfifriederici: Nieden (1911): 
440. Syntypes: ZMB 21787. FMNH 73836 
(formerly ZMB 21789): Rwanda. Rugege 



2009 



NEW CAMHROONIAN ARTIlROLEPriS 




Figure 5. Anliroleptis cuhlfijrieclerici {CAS 177031) 
in dorsal (A) and ventral (B) views. Scale bar, 10 mm. 



(presently called Nyungwe; Gartshore, 1986) 

and Bugoie (= Bugoya?) Forests; Nyungwe 

Forest, ca. 02°20'S^to 02°45'S, 029°05'E to 

029°25'E, ca. 1,800-2,400 m elevation (lat., 

long., elev., estimated). 

Arthroleptis adolfi-friderici: Nieden (1913): 

165. Incorrect subsequent spelling. 

A rthroleptis ( A hroscaplms ) adolfi-friederici: 

Laurent (1940): 82. 



Arthroleptis adolfifriederici: Lovcridge (1942): 

429 (part). Hyphen removed from name. 

A rthrolcpis adolfifriederici adolfijriederiei: 

Lovcridge (1953): 387; 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) indicated in parentheses. Demo- 
cratic Republic of Congo: MCZ A- 14696 
(female). Rwanda: FMNH 73836 (syntype 
[formerly ZMB 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 much longer than the 
first and second fingers, and generally glandu- 
lar skin. Arthroleptis adolfifriederici differs in 
the following ways from other medium to large 
Arthroleptis (maximum SVL > 35 mm): from 
A. affinis by a less prominent and less round 
tympanum and by lacking supernumerary 
tubercles on the feet; from A. francei and A. 
stenodactylns by crus length much greater than 
50% SVL; from A. krokosita by lighter 
pigmentation on throat and vent and lacking 
large well-defined black spots on the lateral 
surface of the body; from A. perreti by a more 
globose inner metatarsal tubercle, more grand- 
ular skin, and less pigmentation on throat; 
from A. nikeae by smaller adult body size (> 
50 mm in A. nikeae); from A. stenodactylns by 
an inner metatarsal tubercle length less than 
80% first toe length; from A. tanneri by a 
relatively narrower head; from A. variabilis by 
lacking a white stripe on the midline of throat; 
from both A. stenodactylns and A. variabilis by 
having a globose inner metatarsal tubercle. 



12 



BREVIORA 



No. 515 



Table 2. Measurements (mm) of female svntypes of 

ArTIIROLEPTIS ADOLF// Rlf:n/:RI( 7. 





ZMB 21787 


FMNH 73836 


Snout-vent length 


42,0 


40. 1 


Head width 


14.7 


14.0 


Tympanum height 


2.4 


2.1 


Eye diameter 


4.9 


4.6 


Snout length 


3.8 


4.0 


Forearm length 


10.5 


9.8 


Manual digit I 


3.6 


3.9 


Manual digit II 


4.3 


4.1 


Manual digit III 


6.7 


6.1 


Manual digit IV 


4.5 


4.1 


Thigh length 


24.2 


22.3 


Crus length 


26.6 


23.5 


Pedal digit I 


3.1 


2.8 


Pedal digit II 


5.3 


4.6 


Pedal digit III 


9.6 


7.8 


Pedal digit IV 


14.7 


12.8 


Pedal digit V 


8.1 


7.1 


Inner metatarsal length 


1.9 


1.8 



Description. Based on syntype ZMB 21787: 
large (42.0 mm SVL), moderately robust 
female with long, slender limbs (Table 2); 
dissected and missing left forelimb beyond 
distal two thirds of humerus; head somewhat 
broad; head length 79% head width; head 
width 35% SVL; snout projecting approxi- 
mately 1 mm beyond lower jaw; rostral tip 
rounded in dorsal view, blunt and essentially 
straight in lateral view; eyes projecting be- 
yond eyelids in dorsal view, such that pupil 
just visible; eyes close to but not projecting 
beyond lateral margins of head; eyes project- 
ing well above dorsal surface of head in lateral 
view; eye diameter approximately equal to 
interorbital distance; pupil horizontally ellip- 
tical in preservative; loreal region essentially 
flat; naris large, rounded, facing laterally, and 
obvious from above; distinct triangular pa- 
pilla at ventral narial rim; canthus rostralis 
very indistinct and rounded; eye diameter 1.5 
times eye-narial distance; eye diameter 2.9 
times distance from naris to rostral tip; 
internarial region flat; internarial distance 
91% interorbital distance; tympanum distinct. 



round, and tending toward ovoid, height just 
less than half diameter of eye; tympanic 
annulus poorly defined; supratympanic fold 
absent; tongue lozenge-shaped, width approx- 
imately half of head width; tongue covered 
with many small pustules, posterior notch 
approximately one eighth anteroposterior 
length of tongue; anterior attachment of 
tongue narrow; prominent, flat, fiap-like 
median papilla on dorsal anterior tongue 
surface; choana visible behind maxillary shelf 
in ventral view; premaxillary and maxillary 
teeth present, but hidden by lips; vomerine 
teeth absent. 

Skin of dorsal surface of head and body 
very weakly tuberculate; skin of fore and 
hind limbs smooth; skin of lateral body 
glandular, becoming more distinct extending 
onto ventrolateral surface; ventral surface 
weakly glandular and somewhat wrinkled; 
median skin raphe indistinct in preservative; 
cloacal region very glandular. 

Limbs and digits well developed; digits of 
manus slightly robust and of pes more 
slender; relative length of fingers; III > IV 

> II > I; fingertips rounded, slightly swollen, 
approximately same width as rest of finger; 
finger with rounded, prominent, globular, 
single subarticular tubercles; palmar and 
metacarpal tubercles present but very weakly 
developed and barely projecting from surface 
of hand; webbing on manual digits absent; 
hind limbs long and slender; thigh length 
91% of crus length; relative length of toes: IV 

> III > V > II > I; toe tips swollen but 
damaged from past dehydration; toes with 
prominent, single, rounded, and globular 
subarticular tubercles; webbing between ped- 
al digits absent; prominent rounded and 
ovoid inner metatarsal tubercle, length 61% 
of first toe length. 

Measurements. Table 2. 

Co/oration in Alcohol. Little of the original 
pigmentation is preserved on ZMB 217897, 
the skin of which is mostly transparent, and 



2009 



NEW CAMEROONIAN ARIII ROLEPTIS 



13 



FMNH 73836. This description of coloralion 
is a siininiar\ based on all of the reference 
specimens. 

Dorsal base color light to medium brown 
with small dark-brown spots (Fig. 5A); iris 
dark gray with white pupil: snout lighter 
brown than interorbital bar but grading to 
dark brown near rostral tip; loreal and 
suborbital regions light brown: dark-brown 
supratympanic band extending from postero- 
dorsal margin of orbit, extending posteroven- 
trally and temiinating just posterior to the 
ventral tympanum margin: tympanum trans- 
lucent and light brown: dorsal surface of amis 
light brown to tan: single, somewhat broken 
brown band on forearm: dorsal surface of 
hands mottled light and dark brown: interor- 
bital bar medium to dark brown: posterior to 
skull, 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 dark- 
brown mottling: crus with two variably 
developed transverse brown stripes. 

Throat very light brown with scattered 
medium-brown spots especially concentrated 
at margins of lower jaw (Fig. 5B): venter 
mostly very light brown with few to no dark 
spots: medial surfaces of arm and forearm 
unpigmented and creamy: ventral surface of 
fore limbs very light brown: palmar, meta- 
carpal, and subarticular tubercles generally 
less pigmented than surrounding skin: ven- 
tral surface o'i thigh very light brown, with 
essentially no markings: ventral surface of 
crus very light brown to weakly mottled with 
light to medium brown: plantar surface light 
to medium brown. 

Variation. Measurement variation is doc- 
umented in Table 2. 



/hi/vtat ami Ran}y,c. Specimens similar to 
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. 
1913). We have not examined, nor found 
explicit reference to, specimens from Bur- 
undi, although it has been reported as 
occurring there (Channing and Howell. 
2006: Global Amphibian Assessment, 
2006). Specimens examined from Tanzania, 
Malawi, and Kenya previously referred to as 
A. adoljifriederici invariably represent differ- 
ent species (see Remarks section). 

Remarks. Nieden (1913) referred Tanza- 
nian specimens from Tanga and Amani to A. 
adolfifriederici. Given that the material later 
used by Ahl (1939) to describe A. aj finis is a 
single large specimen collected by S. G. 
Awerinzew in Amani, it seems likely that 
Nieden (1913) had examined the 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) 
later recognized A. affinis as a junior 
synonym of A. adolfifriederici. These records 
extend the range of A. adolfifriederici 
throughout much of the Eastern Arc Moun- 
tains. Later authors recognized A. adolfifrie- 
derici and A. affinis as two distinct species 
(e.g., Channing and Howell. 2006: Grand- 
ison, 1983: Poynton, 2003b: Skelton-Bour- 
geois, 1961), but records o{ A. adolfifriederici 
from throughout the Eastern Arc Mountains 
continue to be reported (e.g., Channing and 
Howell, 2006). Examination of MCZ speci- 
mens from Tanzania and Kenya that were 
assigned to A. adolfifriederici (Barbour and 
Loveridge, 1928: Loveridge. 1942) indicates 
that these should be referred to A. affinis. A. 
stenodactyliis, A. tanneri, and possibly a large 
undescribed species from southwestern Tan- 



14 



BREVIORA 



No. 515 



zania (Blackburn, unpublished data; see 
Material Examined section). One of these 
specimens (MCZ A-13166) was designated a 
paratype oi A. tawieri by Grandison (1983). 
In their recent study of morphological 
variation in populations of A. af finis from 
throughout much of the Eastern Arc Moun- 
tains, Poynton and Loader (2008) were 
unable to identify any large Arthroleptis 
from eastern Tanzania as morphologically 
similar to the type specimen of A. adolfifrie- 
derici that they examined (ZMB 21787). 

Etymology. Based on the "stately form" of 
this species (Nieden, 1913), it was named in 
honor of German explorer Adolf Friedrich, 
Duke of Mecklenburg, who led an expedi- 
tion in the mountains of Central Africa from 
1907 to 1908. 

Arthroleptis variabilis Matschie, 1893 

Variable Squeaker Frog 
Figure 6 

Arthroleptis dispar. Peters (1875): 210, plate 3. 
Arthroleptis variabilis: Matschie (1893): 173. 
Syntypes: ZMB (85 specimens): Cameroon. 
Southwest Province, Buea, ca. 04°09'N. 
009°14'E, ~ 950 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 
(1940): 85. 

Abroscaphus variabilis: Laurent (1957): 275. 
By implication. 

Reference Sample. Type material, sex, and 
SVL (in mm) indicated in parentheses. 
Cameroon: MCZ A-2654 (female, 34.6), A- 
3428-29 (females, 30.5, 29.4), A-46985 
(male?, 35.0) A-1 36744 (female, 35.8), A- 
136775 (female, 28.7), A- 136777-79 (females. 
29.0, 29.0, 29.4), A- 136820 (female, 34.6), A- 
136823-25 (females, 31.8, 29.6. 29.4). A- 
136827 (female, 33.5). A-1 36830 (female. 




Figure 6. Arthroleptis vanuhilis (MCZ A- 136830) in 
dorsal (A) and ventral (B) views. Scale bar, 10 mm. 

35.0), A-137297 (female, 35.6); USNM 
563684 (female, 37.8), 563685 (male?, 26.3), 
563686 Guvenile, 20.8). 563688 (male. 31.0) 
563689 (female, 37.8); UTA A-35924 (fe- 
male, 32.3), A-35933 (male, 32.1), A-35940 
(female, 36.0), A-35949-50 (females, 34.7, 
37.4), A-44447 (female. 35.4), A-44451 (fe- 
male, 37.7); ZMB 15206 (syntype; female, 
36.7), 70085-86 (syntypes; females, 30.8, 
32.1). 



2009 



NEW CAMEROONIAN ARl IIROLEniS 



15 



Diagnosis. A medium-sized Avthrolcptis 
characterized by a dark-gray throat thai is 
bisected by a prominent, white gular stripe 
(Fig. 6B) and a fourth finger that is approx- 
imately the same length as or shorter than 
the fust and second fingers. This species 
differs from nearly all other similarly sized or 
larger Avthrolcptis by the presence o{ the 
white gular stripe (incipient in A. nikcucY. this 
stripe can be obscured in A. variahilis males 
with darkened throats. Arthroleptis variahilis 
differs in the following ways from other 
medium to large Avthrolcptis (maximum 
SVL > 35 mm): from A. ajfiuis by lacking 
supernumerary tubercles on the feet; from A. 
fvancci and A. stenodactylus by crus length 
much greater than 50% SVL; from A. nikeac 
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. 6; Ta- 
ble 3); mean male SVL 31.1 mm (n = 4; ± 
3.1), mean female SVL 33.1 mm (n = 27; ± 
3.0); head somewhat broad; head length 80 
85% head width; head width 38^5% SVL; 
posterior margin of head not always distinct 
from body; snout projecting less than 1 mm 
beyond lower jaw; rostral tip rounded in 
dorsal and lateral views; in dorsal view, eyes 
projecting beyond eyelids such that pupil just 
visible; eyes close to or just projecting 
beyond lateral margins of head; eyes pro- 
jecting slightly or well above dorsal surface 
of head in lateral view; eye diameter just 
larger than interorbital distance; pupil 
rounded to horizontally elliptical in preser- 



TaMI.I: 3. MEASUKrMr.NTS (mm) OI- II-MAI.I- SYNT'il'l-S oi- 
A nillHOI V.I'IIS I I Rl ) nil. IS. 





ZMB 


ZMB 


ZMB 




1 5206 


70085 


70086 


Snoul vcnl length 


36.7 


30.8 


32.1 


Head vvidlh 


14.1 


12.8 


12.7 


rympaniim height 


2.6 


2.1 


2.1 


Eye diameter 


5.1 


5.0 


4.9 


Snout length 


2.7 


2.4 


2.8 


iHireaim length 


9.5 


7.5 


7.9 


Manual digit I 


3.8 


3.1 


3.3 


Manual digit II 


3.9 


3.0 


3.4 


Manual digit III 


5.5 


4.6 


4.7 


Manual digit IV 


3.1 


2.9 


2.8 


Thigh length 


18.9 


15.9 


15.8 


Crus length 


19.5 


16.3 


17.0 


Pedal digit I 


2.6 


2.3 


2.2 


Pedal digit II 


3.8 


3.2 


3.3 


Pedal digit III 


6.7 


5.5 


6.0 


Pedal digit IV 


10.4 


8.7 


8.7 


Pedal digit V 


5.4 


4.3 


4.6 


Inner metatarsal length 


2.2 


1.7 


1.7 



vative; loreal region ranging from flat to 
slightly concave; naris small and rounded, 
directed laterally, but visible from above; 
distinct small, triangular papilla at ventral 
narial rim; canthus rostralis indistinct, 
rounded; eye diameter 1.8-2.3 times eye- 
narial distance; eye diameter 1.2-1.5 times 
distance from naris to rostral tip; internarial 
region slightly convex; internarial distance 
85-92% interorbital distance; tympanum 
distinct, ovoid, height ranging from 40-50% 
diameter of eye; tympanic annulus usually 
poorly defined; supratympanic fold absent; 
tongue broad and heart shaped, width 
approximately half of head width; tongue 
covered with many small pustules, posterior 
notch ranging from very shallow to less than 
one fifth of anteroposterior length of tongue; 
anterior attachment of tongue narrow; 
prominent, conical, median papilla on dorsal 
anterior tongue surface; choana hidden by 
maxillary shelf in ventral view; premaxillary 
and maxillary teeth present, but hidden by 
lips; vomerine teeth absent. 



16 



B RE I 10 R A 



No. 515 



Skin of dorsal surface of head and body 
smooth, sometimes weakly granular posteri- 
orly; skin of fore and hind limbs smooth; 
skin of lateral body glandular; most of 
ventral surface very smooth, but tending 
toward glandular posteriorly; median skin 
raphe ranging from indistinct to indiscern- 
ible in preservation; smooth skin surround- 
ing cloacal region. 

Limbs and digits well developed; relative 
length of fingers: III > I -= II > IV; 
fingertips rounded and often very swollen 
but not expanded; finger with rounded, 
prominent, globular, single subarticular tu- 
bercles; palmar and metacarpal tubercles 
ranging from weakly developed (mostly in 
old preserved material) to very prominent 
and projecting from surface of hand; web- 
bing on manual digits absent; thigh length 
ranges from 78 to 102% of crus length (90- 
98% thigh length in majority of specimens); 
relative length of toes: IV > III > V > II > 
I; toe tips slightly expanded to just greater 
than width of toe at interphalangeal joints; 
toes with prominent, single, rounded, and 
globular subarticular tubercles; webbing 
between pedal digits absent; very prominent, 
flange-like inner metatarsal tubercle, length 
ranging from 55 to 120% of first toe length. 

Measurements. Table 3. 

Coloration in Alcohol. This species exhibits 
a remarkable diversity of coloration and 
patterns that was first noted long ago (e.g., 
Andersson, 1905; Matschie, 1893; Noble, 
1924). The following description of colora- 
tion is a summary based on the reference 
specimens, but it is premature to consider it 
exhaustive. 

Base color of dorsal and lateral body, and 
limbs, grayish brown to brownish gray 
(Fig. 6A); iris dark gray or black with white 
or creamy pupil; usually solid interorbital 
bar medium to dark brown; snout usually 
uniformly lighter than interorbital bar and 
typically similar to remaining dorsum; loreal 



and suborbital regions uniformly 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 supratym- 
panic band extending from posterodorsal 
margin of orbit, extending posteroventrally 
and terminating posterior to tympanum or 
even extending posterior to arm; tympanum 
translucent and typically lighter than supra- 
tympanic band; dorsal markings similar 
coloration as dorsal base coloration but 
shades darker; some specimens with scat- 
tered small light spots; usually two fairly 
complete 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 
surfaces, with sharp boundary between 
regions; variably present, usually continuous, 
white vertebral line extending from snout tip 
to vent; other dorsal markings, if present, 
usually comprising dark triangle between 
eyes (apex directed posteriorly) often, but 
not necessarily, confluent with broad circu- 
lar, elliptical, or diamond-shaped dark mid- 
dorsal marking located over suprascapulars; 
mid-dorsal marking often, but not necessar- 
ily, confluent with circular or elliptical dark 
posterior dorsal marking, located over sa- 
crum, poorly defined along most posterior 
margins; dorsal markings sometimes discon- 
tinuous or reduced to poorly defined or even 
paired blotches symmetrical across midline; 
lateral body with scattered well-defined 
dark-brown spots and, in some cases, small, 
less defined light spots; hind limbs fairly 
uniform in coloration and pattern; thigh 
usually with two dark transverse stripes; 
dorsal and posterior thigh fairly uniform in 
coloration and pattern; crus with two, 
sometimes three, pronounced transverse 
brown stripes; dark spot on lateral margin 
of foot at base of metatarsal V; dark region 



20(W 



NEW CAMEROONIAN ARTHROl.EPTIS 



17 



covering cloaca, well defined dorsally and 
extending onto and becoming less distinct on 
posterior thigh. 

Throat light to dark gray with very small, 
irregularly placed light spots (Fig. 6B); light 
spots at jaw margin very irregularly placed; 
well-defined, usually continuous, white stripe 
extending from rostral tip along throat 
midline to level of pectoral girdle; posterior 
to pectoral girdle pattern becomes dominat- 
ed by large light spots with dark pigmenta- 
tion becoming lighter and more diffuse 
posteriorly; posterior venter mostly white 
or very light gray with no markings; medial 
surfaces of arm and forearm white, creamy, 
or light gray; dark ventral surface of 
forearm; palmar surface and fingers light to 
dark gray; palmar, metacarpal, and subartic- 
ular tubercles unpigmented; ventral finger- 
tips lighter in color than rest of finger; venter 
and ventral thigh both cream colored with 
few scattered brown melanocytes; light- 
creamy or light-gray ventral surface of thigh 
with few markings; ventral surface of crus 
mottled dark gray and cream; plantar surface 
homogeneous dark gray or black; subarticu- 
lar tubercles on toes and inner metatarsal 
tubercle lighter shades than surrounding skin. 

Coloration in Life. Based on D. C. Black- 
burn's field notes and photographs. Dorsal 
base coloration black, dark gray, dark, light, 
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 
generally dark gray to black with white or 
creamy stripe along midline; venter white or 
gray with light- to dark-gray spots; inguinal 
region ranging from pale yellow to orangey red. 

Variation. Measurement variation is doc- 
umented in Table 3. No relationship is 
obvious between the great diversity of dorsal 



patterns and coloration with that of the 
ventral surface, which is fairly uniform 
among specimens. One unusual color morph 
(e.g., MCZ A- 136827) exhibits a large 
immaculate white band, just below and 
posterior to the eye, that extends posterior 
onto the fore limb along the dorsal surface to 
just distal to the elbow. 

Habitat and Range. This typically is a 
common species that occurs predominantly 
in lowland forests (e.g., Amiet, 1975). Ar- 
throlepti.s variai^ilis is known from through- 
out Central Africa, with localities in Camer- 
oon (e.g., Amiet, 1975; Andersson, 1905; 
Bohmeand Schneider, 1987; Boulenger 1900, 
1905, 1906; Gartshore, 1986; Herrmann et 
al., 2005; Lawson, 1993; Matschie. 1893; 
Nieden, 1908; Perret, 1966; Perret and 
Mertens, 1957), Central African Republic 
(Joger, 1990), Bioko Island (e.g., Boulenger, 
1900, 1905), mainland Equatorial Guinea 
(e.g., De la Riva, 1994), Gabon (e.g., 
Boulenger, 1905; Burger et al., 2006), eastern 
Nigeria (Blackburn, unpublished data; 
Schiotz, 1963), the Democratic Republic of 
Congo (Kinshasa; e.g., Laurent, 1972; No- 
ble, 1924; de Witte, 1934), and the Republic 
of Congo (Brazzaville; e.g., Largen and 
Dowsett-Lemaire, 1991). 

Remarks. Arthro/eptis variabilis is some- 
times stated to occur in the forests of western 
Africa (i.e. Global Amphibian Assessment, 
2006). However, the western African popu- 
lations probably correspond to one or 
several undescribed species {^"Arthroleptis 
sp. 2": Ernst et al., 2008; Rodel and Branch, 
2002; Rodel and Ernst, 2004; Rodel et al., 
2005). Populations from western Ivory Coast 
are similar to A. variabilis, but smaller 
(Rodel and Branch, 2002). A specimen from 
southwestern Ghana (Rodel et al., 2005) was 
described recently as a new species, A. 
krokosua, by Ernst et al. (2008). Like A. 
perreti, A. krokosua also lacks the white 
gular stripe present in A. variabilis and has a 



18 



BREVIORA 



No. 515 



fourth finger shorter than the first and 
second fingers (Ernst et al., 2008). However, 
morphological and molecular data indicate 
that these taxa are neither conspecific nor 
sister taxa (Blackburn, unpublished data; 
Ernst et 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 form variabilis, changeable, 
presumably in recognition of the wide range 
of coloration and pattern exhibited by this 
species. 

DISCUSSION 

Differences between A. perreti and A. 
variabilis documented in this study are 
similar to those found by Perret (1966). In 
general, he found A. variabilis to be more 
robust and to have a relatively shorter snout, 
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 thigh. Arthroleptis adolfifriederici, as 
described by Perret (1966), lacks the white 
stripe on the throat and has a posterior thigh 
that is brown with light spots. Perret's (1966) 
description agrees with A. perreti as de- 
scribed here. Although superficially similar, 
A. perreti and A. variabilis are morphologi- 
cally distinct. Molecular phylogenetic data 
indicates that, while closely related, these 
two species are not sister taxa (see Arthro- 
leptis sp. nov. 1 in Blackburn, 2008a). 
Guided by the description and redescriptions 
presented here, future studies should focus 
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 



countries (e.g., Amiet, 1971a, b, 1972a, b, 
1973, 1977. 1980a,b, 1981, 2000, 2001, 
2004a,b; Blackburn, 2008b; Herrmann et 
al., 2004; Kobel et al., 1980; 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 
200 described species. Much of this diversity 
is concentrated in the mountains of the 
Cameroon Volcanic Line, especially Mt. 
Manengouba (Amiet, 1971a,b, 1973, 1977, 
1980a; Blackburn, 2008b; Kobel et al., 1980). 
This mountain is an important center of 
diversity within Cameroon, especially of 
amphibians (e.g., Amiet, 1975; Blackburn, 
2008b; Gartshore, 1986), but has received 
only limited conservation attention (e.g., 
Gartshore, 1986; Gonwouo et al., 2006). 
Description of A. perreti from Mt. Manen- 
gouba highlights the need for conservation 
measures focused on the preservation of the 
unique biodiversity found on this and other 
mountains of the Cameroon Volcanic Line. 

MATERIAL EXAMINED 

Type material, sex, and SVL (in mm) 
indicated in parentheses. Arthroleptis affinis: 
Tanzania: MCZ A-13145-146 (females, 37.2, 
37.6), A-13150-151 (males, 23.4, 25.3), A- 
13152-153 (females, 35.8, 40.5), A-13157- 
160 (females, 32.8, 33.7, 34.7, 36.6), A- 
13162-163 (females, 36.6, 35.7), A-13165 
(female, 35.6), A-13167 (male, 27.8), A- 
13169 (male, 29.2), A-25400^01 (females, 
35.5, 30.5), A-138241-244 (female, 36.2, 
33.8, 42.9, 37.4); TNHC 38577 (female, 
38.8); ZMB 23093 (holotype; female, 35.6). 
Arthroleptis francei: Malawi: BMNH 1954.1. 
13.74 (paratype; female, 32.4); MCZ A- 
27474-476 (paratypes; males, 25.4, 28.2, 
28.0), A-27477^78 (paratypes; females. 



2009 



NEW CAMEROONIAN ARIII KOLIiniS 



19 



38.7, 41.0), A-27479 (holotype; female, 41.9), 
A-137038 (female, 37.3): TMP 48089 (fe- 
male, 43.0), 48092 (male, 27.8). Arthrolcplis 
kwkosua: Ghana: SMNS 12555 (hololype: 
male, 43.3). Arthrolcptis iilkeac: Tanzania: 
BMNH 2002.101 (hololype: female, 52.5), 
2002.102 (paratype: female, 53.7). Arthrolcptis 
stemxiactylus: Malawi: MMB HA2()()2.1.13 
(female, 37.0), HA2002.1.60 (male, 32.7), 
HA2002.4.17 (female, 38.4). Tanzania: 
BMNH 2002.596 (female, 33.0); CAS 168455 
(male, 35.6); MCZ A-21715 (male, 27.6), A- 
25387 (female, 28.6), A-27447 (female, 39.5). 
Kenya: NMK A/4251 (male. 33.0), A/4251/3 
(female 40.0), A/4251/4 (male, 31.0), A/4401/3 
(female, 25.7), A/4401/6 (female, 26.2). Arthro- 
leptis tanneri: Tanzania: CAS 168823 (male; 
41.9), 168825 (female; 54.7); MCZ A-13166 
(paratype; female; 36.5). Arthrolcptis variabihs 
(specimens not included in PCA): Equatorial 
Guinea: CAS 207817-819 (females, 34.0, 30.6, 
35.4), 207820 (female?, 26.2), 207821-822 
(female, 33.3, 36.1), 207823 (juvenile, 25.3), 
207824-826 (females, 32.7, 33.4, 31.0), 207827 
(female?, 26.9), 207828 (female, 35.9). Arthro- 
lcptis sp.: Tanzania: MCZ A- 16952 (female, 
44.4). 

ACKNOWLEDGMENTS 

Many people assisted with the fieldwork, 
logistics, or both needed to collect specimens 
for this study: K. S. Blackburn, M. Che, V. 
Che, J. L. Diffo, D. Fotibu, N. Gawani, P. 
Huang, M. LeBreton, and M. T. Kouete. 
The people of Nsoung and the Fulbe- 
speaking community in the Mt. Manen- 
gouba 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 
permits (0173PR/MINEF/SG/DFAP, 0588/ 
PRBS/MINFOF/SG/DFAP/SDVEF/SC)and 
permissions to export specimens (0850 54/ 



CO/M 1 N F F/SG/D FA P/S AN , 05121 6/PR BS/ 
MINFOF/SG/DFAP/SDVEF/SC). DCB was 
funded by the Department of Organismic and 
Evolutionary Biology (Harvard University), 
Goelet Summer Research Funds, a Putnam 
Expeditionary Grant from the Museum of 
Comparative Zoology (Cambridge, Massa- 
chusetts, U.S.A.), NSF grant EF-0334939 
(AmphibiaTree) to D. Cannatella and J. 
Hanken, and Project Exploration, a not-for- 
profit educational organization (Chicago, 
Illinois, U.S.A.). LNG was supported by 
Conservation International (Washington, 
DC, U.S.A.). D. Hewitt provided assistance 
with translations from German, and C. 
Bonneaud translated the abstract into 
French. C. S. Walker (Harvard Map Collec- 
tion) created Figure 1. J. Hanken and R. C. 
Drewes provided valuable comments on a 
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. 

Amiet. J.-L. 1971a. Especes nouvelles ou mal connues de 
Leptodactylodon (Amphibiens Anoures) de la Dor- 
sale camerounaise. Annales de la Faculte des 
Sciences du Cameroun, 5: 57-81. 

. 1971b. Leptodactylodon nouveaux du Cameroun 

(Amphibiens Anoures). Annales de la Faculte des 
Sciences du Cameroun, 7-8: 141-172. 

. 1972a. Description de cinq nouvelles especes 

camerounaises de Cardioglossa (Amphibiens An- 
oures). 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 les autres especes 
camerounaises. Bulletin de I'lnstitut Franijais Afri- 
que Noire. 35: 462-474. 

. 1975. Ecologie et distribution des amphibiens 

anoures de la region de Nkongsamba (Cameroun). 
Annales de la Faculte des Sciences de Yaounde, 20: 
33-107. 



20 



BREVIORA 



No. 515 



— . 1977. Les Astylostennis du Cameroun (Amphib- 
ia Anura, Astylosterninae). Annales de la Faculte 
des Sciences de Yaounde, 23-24: 99-227. 

— . 1980a. Revision du genre Leptudactyloclcm 
Andersson (Amphibia. Anura, Astylosterninae). 
Annales de la Faculte des Sciences de Yaounde, 
27: 69-224. 

— . 1980b. Un Hyperoliiis nouveau du Cameroun: 
Hyperolius endjaiui n. sp. (Amphibia Anura. Hy- 
peroliidae). Revue Suisse de Zoologie, 87: 445-460. 

— . 1981. Une nouvelle Cardioglossa orophile de la 
Dorsale camerounaise: C. schioetzi nov. sp. (Am- 
phibia, Anura, Arthroleptinae). Annales de la 
Faculte des Sciences de Yaounde, 28: 117-131. 

— . 1987. Aires disjointes et taxons vicariants chez 
les Anoures du Cameroun: implications paleocli- 
matiques. Alytes, 6: 99-115. 

— . 2000. Les Alexteroon du Cameroun (Amphibia, 
Anura, Hyperoliidae). Alytes, 17: 125-164. 

— . 2001. Un nouveau Lepiopelis de la zone 
forestiere camerounaise (Amphibia, Anura, Hyper- 
oliidae). Alytes, 19: 29-44. 

— . 2004a. Une nouvelle espece d' Hyperolius du 



Cameroun (Amphibia, Anura, Hyperoliidae). Re- 
vue Suisse de Zoologie, 11: 567-583. 
— . 2004b. A propos de deux Leptopelis nouveaux 



pour la faune du Cameroun (Anura. Hyperoliidae). 
Alytes. 21: 111-170. 

Andersson, L. G. 1905. Batrachians from Cameroon 
collected by Dr. Y. Sjostedt in the years 1890-1892. 
Arkiv for Zoologie, 2: 1-29. 1 plate. 

Barbour, T.. and A. Loveridge. 1928. A comparative 
study of the herpetological faunae of the Uluguru 
and Usambara Mountains, Tanganyika Territory 
with descriptions of new species. Memoirs of the 
Museum of Comparative Zoology, 50: 87-265, 4 
plates. 

Blackburn, D. C. 2005. Cardioglossa liheriensis Bar- 
bour & Loveridge 1927 is a junior synonym of 
Phryuohatrachus frateradus (Chabanaud 1921). 
African Journal of Herpetology, 54: 171-179. 

. 2008a. Biogeography and evolution of body size 

and life history of African frogs: phylogeny of 
squeakers (Arthroleptis) and long-fingered frogs 
(Cardioglossa) estimated from mitochondrial data. 
Molecular Phylogenetics and Evolution, 49: 806-826. 

. 2008b. A new species of Cardioglossa (Amphib- 
ia: Anura: Arthroleptidae) endemic to Mount 
Manengouba in the Republic of Cameroon, with 
an analysis of morphological diversity in the genus. 
Zoological Journal of the Linnean Society, 54: 
611-630. 

. In press. Diversity and evolution of male 

secondary sexual characters in African squeakers 



and long-fingered frogs. Biological Journal of the 
Linnean Society. 

BouLENGER, G. A. 1900. A list of the batrachians and 
reptiles of the Gaboon (French Congo), with 
descriptions of new genera and species. Proceedings 
of the Zoological Society, London, 1900: 433^56, 6 
plates. 

. 1905. Descriptions of new tailless batrachians in 

the collection of the British Museum. Annals and 
Magazine of Natural History, 16: 180-184. 

. 1906. Descriptions of new batrachians discov- 



ered by Mr. G. L. Bates in South Cameroon. 
Annals and Magazine of Natural History, 17: 
317-323. 

BoHME, W., AND B. Schneider. 1987. Zur Herpetofau- 
nistik Kameruns (III) mit Beschreibung einer neuen 
Cardioglossa (Anura: Arthroleptidae). Bonner Zo- 
ologische Beitrage, 38: 241-263. 

Burger, M., O. S. G. Pauwels, W. R. Branch, E. Tobi, 
J. -A. Yoga, and E.-N. Mirolo. 2006. An assess- 
ment of the amphibian fauna of the Gamba 
Complex of protected areas. Gabon, pp. 297-307. 
//; Gamba. Gabon: Biodiversite d'une foret equator- 
iale africaine. A. Alonso. M. E. Lee. P. Campbell, O. 
S. G. Pauwels, & F. Dallmeier (eds.). Bulletin of the 
Biological Society of Washington no. 12. 436 pp. 

Channing, a., and K. M. Howell. 2006. Amphibians 
of East Africa. Ithaca, New York, Cornell Univer- 
sity Press. 418 pp. 

De la Riva, I. 1994. Anfibios anuros del Parque 
Nacional de Monte Alen, Rio Muni, Guinea 
Ecuatorial. Revista Espanola Herpetologie, 8: 
123-139. 

Drewes, R. C, and J. V. Vindum. 1994. Amphibians of 
the Impenetrable Forest, southwest Uganda. Jour- 
nal of African Zoology, 108: 55-70. 

Ernst, R., A. C. Agyei, and M.-O. Rodel. 2008. A new 
giant species of Arthroleptis (Amphibia: Anura: 
Arthroleptidae) from the Krokosua Hills Forest 
Reserve, south-western Ghana. Zootaxa, 1697: 
58-68. 

Frost, D. R. (ed.) 1985. Amphibian Species of the 
World: A Taxonomic and Geographical Reerence. 
Lawrence, Kansas, 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 1 
August 2008]. Available from: http://research.amnh. 
org/herpetology/amphibia/index.php. 

Frost. D. R.. T. Grant. J. Faivovich, R. H. Bain, A. 
Haas, C. F. B. Haddad, R. O. de S.4, A. Channing, 
M. Wilkinson, S. C. Donnellan, C. J. Rax- 
woRTH'i , J. A. Campbell, B. L. Blotto, P. Moler, 



2009 



NEW CAMhROONlAN ARTHROl.EPTIS 



21 



R. C. Drivvis. R. a. Nussbaum, J. D. L>nc ii. I). 
M. Green, and W. C. Whehii:r. 2006. Tlic 
amphibian tree of life. Bullclin of the American 
Museum of Natural History. 297: 1 291. 

GartshorI', M. 1986. The status ol" the montane 
herpetofauna of the Cameroon highlands, pp. 
204-240. /// Stuart, S. N. (ed.). Conservation of 
Cameroon Montane Forests. London. Internation- 
al Council for Bird Preservation. 263 pp. 

Global Ami'human Assessment. 2006. Washington, 
D.C., IUCN/SSC-CT/CABS; c2006 [cited 1 August 
2008] Available from: htlp://wvvw.globalamphibians. 
org. 

GoNwouo, L. N., M. LeBreton, C. Wild, L. Chirio, P. 
Ngassam, and M. N. Tchamba. 2006. Geographic 
and ecological distribution of the endemic montane 
chameleons along the Cameroon mountain range. 
Salamandra, 42: 213-230. 

Grandison, A. G. C. 1983. A new species of Arihro/cpil.s 
(Anura: Ranidae) from the West Usambara Moun- 
tains, Tanzania. Bulletin of the British Museum of 
Natural History (Zoology), 45: 77-84. 

Herrmann. H.-W., P. A. Herrmann, A. Schmitz. and 
W. Bohme. 2004. A new frog species of the genus 
Cardioglossa from the Tchabal Mbabo Mtns, 
Cameroon. Herpetozoa, 17: 119-125. 

, W. Bohme, P. A. Herrmann, M. Plath, A. 

Schmitz, and M. Solbach. 2005. African biodiver- 
sity hotspots: the amphibians of Mt. Nlonako, 
Cameroon. Salamandra, 41: 61-81. 

lUCN. 2001. lUCN red list categories and criteria 
[Internet]. Version 3.1. Gland. Switzerland, lUCN 
[cited 1 August 2008]. Available from: http://www. 
ucnredlist.org. 

JoGER, U. 1990. The herpetofauna of the Central 
African Republic, with description of a new species 
of Rhinotyphlops (Serpentes: Typhlopidae), pp. 
85-102. //; G. Peters, & 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 sp. nov. (Anura: 
Pipidae) from the Cameroons, another case of 
tetraploidy. Revue Suisse de Zoologie, 87: 919- 
926. 

Largen, M. J., AND F. Dowsett-Lemaire. 1991. 
Amphibians (Anura) from the Kouilou River basin, 
Republique du Congo. Tauraco Research Report. 
4: 145-168. 

Laurent. R. F. 1940. Contribution a I'osteologie et a la 
systematique des ranides africains - Premiere note. 
Revue de Zoologie et de Botanique Africaines, 34: 
74-97. 3 plates. 



— . 1954. Remarques sur le genre Scliouieilenella 
Witte. Annales du Musee Royal du Congo Beige, 
Tervuren. 1: 34-40. 

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

— . 1972. Amphibians. Exploration du Pare Na- 
tional des Virunga. institul National Conservation 
de la Nature de la Republique du Zaire, 22: 3-125. 
1 1 plates. 

— . 1973. The natural classification of the Arthro- 



leptinae (Amphibia, Hyperoliidac). Revue de Zool- 
ogie et de Botanique Africaines, 87: 666-678. 

Lawson, D. p. 1993. The reptiles and amphibians of 
Korup National Park Project. Cameroon. Ilerpe- 
tological Natural History, 1: 27-90. 

. 2000. A new eaecilian from Cameroon, Africa 

(Amphibia: Gymnophiona: Scolecomorphidae). 
Herpetologica. 56: 77-80. 

Leviton, a. E.. R. H. Gibbs. Jr.. E. Heal, and C. E. 
Dawson. 1985. Standards in herpetology and 
ichthyology: part I. Standard symbolic codes for 
institutional resource collections in herpetology and 
ichthyology. Copeia, 1985: 802-832. 

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. 

Loveridge, a. 1942. Scientific results of a fourth 
expedition to forested areas in East & Central 
Africa. Bulletin of the Museum of Comparative 
Zoology, 41: 377^36, 4 plates. 

. 1953. Zoological results of a fifth expedition to 

East Africa. IV. Amphibians from Nyasaland and 
Tete. Bulletin of the Museum of Comparative 
Zoology, 110: 325-406, 4 plates. 

. 1957. Check list of the reptiles and amphibians 



of East Africa (Uganda; Kenya; Tanganyika; 

Zanzibar). Bulletin of the Museum 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 the Japanese toads (genus Biifn. 

Bufonidae). Contributions from the Biological 

Laboratory, Kyoto University, 26: 209^28. 
Myers. N., R. A. Mittermeier, C. G. Mittermeier, G. 

A. B. da Fonseca, and J. Kent. 2000. Biodiversity 

hotspots for conservation priorities. Nature, 403: 

853-858. 
Nieden, F. 1908. Die Amphibienfauna von Kamerun. 

mit einer Bestimmungstabelle. Mitteilungen aus 

dem Zoologischen Museum in Berlin, 1908: 

491-518. 



22 



BREVIORA 



No. 515 



. 1911 "1910". Neue ostafrikanische Frosche aus 

dem Kgl. Zool. Museum in Berlin. Sitzungsberichte 
Gesellschaft Nalurforschender Freunde Berlin, 
1910:436-441. 

. 1913. Amphibia. Wissenschaftliche Ergebnisse 

der Deutschen Zentral-Afrika-Expedition, 4: 
165-195. 

Noble, G. K. 1924. Contributions to the herpetology of 
the Belgian Congo based on the collection of the 
American Museum Congo Expedition, 1909-1915. 
Part III. Amphibia. Bulletin of the American 
Museum of Natural History, 49: 147-347, 20 plates. 

Perret, J.-L. 1966. Les amphibiens du Cameroun. 
Zoologische Jahrbucher (Systematik), 8: 289-464. 

. 1991. Le statut d^Arthroleptis hivit tarns F. 

Miiller (Anura, Arthroleptidae). Bulletin de la 
Societe neuchateloise des Sciences naturelles, 114: 
71-76. 

, AND R. Mertens. 1957. Etude d'une collection 



herpetologique faite au Cameroun de 1952 a 1955. 
Bulletin de Tlnstitut Frangais Afrique Noire, 49: 
548-601. 

Peters, W. 1875. Uber die von Hrn. Professor Dr. R. 
Buchholz in Westafrika gesammelten Amphibien. 
Monatsberichte der Koniglichen Preussische Aka- 
demie des Wissenschaften zu Berlin, 1875: 196-212, 
3 plates. 

Plath, M., M. Solbach, and H.-W. Herrmann. 2004. 
Anuran habitat selection and temporal partitioning 
in a montane and submontane rainforest in 
Southwestern Cameroon — first results. Salaman- 
dra, 40: 239-260. 

Poynton, J. C. 1976. Classification and the Arthrolepti- 
nae. Revue de Zoologie Africaine, 90: 215-220. 

. 2003a. Arthroleptis troglodytes and the content 

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

. 2003b. A new giant species of Arthroleptis 

(Amphibia: Anura) from the Rubeho Mountains, 
Tanzania. African Journal of Herpetology, 52: 
107-112. 

, and D. G. Broadley. 1985. Amphibia Zambe- 

siaca 1. Scolecomorphidae, Pipidae, Microhylidae, 
Hemisidae, Arthroleptidae. Annals of the Natal 
Museum, 26: 503-553. 

, and S. p. Loader. 2008. Clinal variation and its 

taxonomic consequences in the common Tanzanian 
forest frog, Arthroleptis affinis. Copeia, 2008: 
517-526. 



RoDEL, M.-O., AND M. A. Bangoura. 2004. A conser- 
vation assessment of amphibians in the Foret 
Classee du Pic de Eon, Simandou Range, south- 
eastern Republic oi Guinea, with the description of 
a new Amnirana species (Amphibia Anura Rani- 
dae). Tropical Zoology, 17: 201-232. 

, AND W. R. Branch. 2002. Herpetological survey 

of the Haute Dodo and Cavally forests, western 
Ivory Coast, Part I: amphibians. Salamandra, 38: 
245-268. 

, AND R. Ernst. 2004. Measuring and monitoring 

amphibian diversity in tropical forests. I. An 
evaluation of methods with recommendations for 
standardization. Ecotropica, 10: 1-14. 

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

M. K. FujiTA, AND R. Ernst. 2005. The amphibians 
of the forested parts of south-western Ghana. 
Salamandra, 41: 107-127. 

, A. ScHMiTZ, O. S. G. Pauwels, and W. Bohme. 



2004. Revision of the genus Werneria Poche, 1903, 
including descriptions of two new species from 
Cameroon and Gabon (Amphibia: Anura: Bufoni- 
dae). Zootaxa, 720: 1-28. 

ScHiOTZ, A. 1963. On a collection of Amphibia 
from Nigeria. Videnskabelige meddelelser fra 
Dansk naturhistorisk forening i Kjobenhaven, 
129: 43^8. 

Schmidt, K., and R. F. Inger. 1959. Amphibians, 
exclusive of the genera Afrixalus and Hyperolius. 
Exploration du Pare National de I'Upemba, Mis- 
sion G.F. de Witte, Institut des Pares Nationaux du 
Congo Beige, 56: 1-264, 9 plates. 

Simmons, J. E. 2002. Herpetological collecting and 
collections management. Herpetological Circular, 
31: 1-153. 

Skelton-Bourgeois, M. 1961. Reptiles et batraciens 
d'Afrique orientale. Revue de Zoologie et de 
Botanique Africaines, 63: 309-338. 

Stuart, S. N. 1984. Conservation recommendations and 
conclusions, pp. 241-245. /// Stuart, S. N. (ed.). 
Conservation of Cameroon Montane Forests. 
London, International Council for Bird Preserva- 
tion. 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 Beige, 
Tervuren, C. - Zoologie, Serie I, 3: 157-188, 7 
plates.