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 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 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, 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, August 2005, L 396 Following personal communication with of Cameroon, Mt Manengouba, near Nsoung, 04°59'N, 009°48'E, 1,420 m elevation, 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, 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 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 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, 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 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; female adult ZMB 2,045 m 71499 (40.2 -gray background mm), perreti (44.7 elevation, (843C), elevation, August 2005; ZMB 3), ZMB 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 ); head somewhat broad, head length 79% head width; snout projecting approximately 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 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) 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 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 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 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 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 Measurements (mm) of female svntypes of ArTIIROLEPTIS ADOLF// Rlf:n/:RI( 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 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 Arthroleptis dispar Peters (1875): 210, plate Arthroleptis variabilis: Matschie Syntypes: ZMB Southwest 009°14'E, Cameroon (85 specimens): Buea, Province, ~ 950 (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 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 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: 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, 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 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 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/ , 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 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 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, 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.) 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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 — 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 — Nature de la la Republique du Zaire, 22: 3-125 plates 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 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, 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, 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 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 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 Museum, 26: 503-553 and S p Loader 2008 , taxonomic consequences forest frog, 517-526 du Beige, 56: 1-264, 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,  ... 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... with descriptions of new species Memoirs of the Museum of Comparative Zoology, 50: 87-265, Drewes, R C, and plates Ernst, R., A C Agyei, and M.-O Rodel 200 8 Blackburn, D C 200 5 Cardioglossa liheriensis... Bulletin of the Africa Zoology, 41: 377^36, Museum & fourth Central of Comparative plates 1953 Zoological results of a fifth expedition to East Africa IV Amphibians from Nyasaland and Tete Bulletin of