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The African anuran genus Cardioglossa contains 19 described species, most of which are distinguished from one another by striking patterns and colors. We present a well-resolved phylogeny based on analyses of mitochondrial and nuclear loci for 18 species of Cardioglossa. This provides the basis for species-delimitation analyses and interpreting historical biogeography in the genus. Whereas much of the diversification within the genus occurred among Central African lineages during the Miocene following the origin of Cardioglossa in the latest Oligocene or earliest Miocene, most species-pairs in the genus diverged more recently during the Plio-Pleistocene. The two most geographically peripheral species—C. cyaneospila in the Albertine Rift Mountains and C. occidentalis in the Upper Guinean Forests of West Africa—both diverged from other lineages during the mid-late Miocene. Because our analyses do not support C. manengouba and C. oreas as distinct species, we recognize these geographically separate and phenotypically distinct populations as subspecies of C. oreas that diverged subsequent to the origin of Mount Manengouba during the past 1.5 million years. In contrast, we find that C. leucomystax likely represents two species found in the Lower Guinean and Congolian forests, respectively. We find recent divergences between several allopatric lineages (either species or populations) that differ in coloration and pattern, including in C. nigromaculata which varies in color across its range in Central Africa and Bioko Island. These recent divergences among allopatric lineages with distinctive coloration and pattern raise new questions about the significance of these traits in this genus for which little is known of its natural history and biology.

Arthroleptis carquejai Ferreira, 1906 Material (one specimen): One adult female specimen (MHNCUP/ ANF 188, formerly UP-MHNFCP-018586; Fig. S19), collected from Cambondo [-9.16349º, 14.66432º, 394 m a.s.l.] Kwanza Norte Province, date unknown. Comments: Arthroleptis carquejai is an Angolan endemic species, found in the northern part of the country (Marques et al. 2018). The specific epithet carquejai was in honor of Bento Carqueja (1860–1935), a businessman, writer, and professor at APP. The single specimen collected by Newton was used by Ferreira (1906) as the holotype for the species. Ceríaco et al. (2014b) recognized A. carquejai as a valid species belonging to the same group of species as Arthroleptis variabilis Matschie, 1893. The authors also analyzed specimens from the Zoologisches Museum Hamburg collected from ““Roca Nova” (Roça Novo Mundo)”, approximately 135 km from the type locality, and determined them to also be A. carquejai. The species was recently collected in Serra do Pingano, Uíge Province, by Ceríaco et al. (Fig. 4) and Ernst et al. (2020). Marques et al. (2018) suggested that A. carquejai most likely also occurs in the Democratic Republic of the Congo and possibly Gabon., Published as part of Santos, Bruna S., Marques, Mariana P., Bauer, Aaron M. & Ceríaco, Luis M. P., 2021, Herpetological results of Francisco Newton's Zoological Expedition to Angola (1903 – 1906): a taxonomic revision and new records of a forgotten collection, pp. 1-80 in Zootaxa 5028 (1) on page 15, DOI: 10.11646/zootaxa.5028.1.1, http://zenodo.org/record/5453967, {"references":["Ferreira, J. B. (1906) Algumas especies novas ou pouco conhecidas de amphibios e reptis de Angola (Coleccao Newton - 1903 - 1904). Jornal de Sciencias Mathematicas Physicas e Naturaes, Segunda Serie, 7 (26), 159 - 171.","Marques, M. P., Ceriaco, L. M. P., Blackburn, D. C. & Bauer, A. M. (2018) Diversity and distribution of the amphibians and terrestrial reptiles of Angola atlas of historical and bibliographic records (1840 - 2017). Proceedings of the California Academy of Sciences, Series 4, 65 (Supplement 2), 1 - 501.","Ceriaco, L. M. P., Blackburn, D. C. Marques, M. P. & Calado, F. M. (2014 b) Catalogue of the amphibian and reptile type specimens of the Museu de Historia Natural da Universidade do Porto in Portugal, with some comments on problematic taxa. Alytes, 31, 13 - 36."]}

Francisco Newton’s zoological expedition to Angola undertaken between 1903 and 1906 is one of the least studied of the naturalist’s life. Only three major papers regarding the herpetofauna collected in this expedition have been published, and a significant part of the specimens remains unstudied since the 1900’s. Here we review the extant herpetological specimens of this expedition, present an updated taxonomic revision, and provide new insights on their taxonomic status. The extant collection is constituted by 329 specimens (155 amphibians and 174 reptiles), corresponding to 73 species, 39 genera and 22 families, and it is presently housed in the Museu de História Natural e da Ciência da Universidade do Porto (MHNC-UP).

The Central African treefrog Leptopelis brevirostris has a characteristically short and vertically truncated snout, a tooth-like process at the centre of the mandible, and unlike most species of Le...

The tadpole of poorly studied Leptopelisragazzii (Boulenger, 1896), a high-altitude tree frog species from the Ethiopian highlands, is described for the first time and compared with closely related sympatric species – L.gramineus (Boulenger, 1898) and L.vannutellii (Boulenger, 1898).

We report the first known occurrence of the Foulassi Screeching Frog, Arthroleptis adelphus (Perret, 1966), from Nigeria. A specimen of A. adelphus was collected during herpetological survey work conducted in Cross River National Park, south-eastern Nigeria. Morphometrics and mitochondrial 16S rRNA gene confirm identity of the specimen. Matrilineal genealogy reveals a sister relationship of A. adelphus from Nigeria with individuals from south-western Cameroon. Genetic analysis further shows geographic structuring and divergence among populations of A. adelphus from the Guineo–Congolian forest region. We offer updates to the IUCN geographic range of A. adelphus.

A new ground-dwelling species of treefrog in the genus Leptopelis is described from the Harenna Forest in south-eastern Ethiopia. The description is based on morphology and acoustics and is supported by molecular data. The new species has a small body size, and the digital discs on fingers and toes are significantly more conspicuous than in other semi-fossorial members of the L. gramineus complex. It occupies forest habitats at lower altitudes and is separated ecologically and geographically from high-altitude species of the complex. One of them, a parapatric cryptic species from Bale and Arsi Mountains, is resurrected from synonymy of L. gramineus and given a new name, L. montanus. Genetic barcoding of specimens from both populations showed that they belong to two distinct lineages that had been revealed by recent phylogenetic research. To confirm the geographic separation of the studied populations, the collection area of L. gramineus types was verified through analysis of the diary and the final report of the 2nd expedition of V. Bottego, and through matching of the route described in it with modern maps. The type locality of L. gramineus sensu stricto is restricted to Gamo Gofa, Ethiopia. Following the results of recent phylogenetic studies, the range of L. gramineus is limited to west of the Great Rift Valley. An identification key to the named Ethiopian species of the genus is provided.

Developmental dynamics 250(4), 584 - 600 (2021). doi:10.1002/dvdy.275, BackgroundCases of parallel evolution offer the possibility to identify adaptive traits and to uncover developmental constraints on the evolutionary trajectories of these traits. The independent evolution of direct development from the ancestral biphasic life history in frogs is such a case of parallel evolution. In frogs, aquatic larvae (tadpoles) differ profoundly from their adult forms and exhibit a stunning diversity regarding their habitats, morphology and feeding behaviors. The transition from the tadpole to the adult is a climactic, thyroid hormone (TH)-dependent process of profound and fast morphological rearrangement called metamorphosis. One of the organ systems that experiences the most comprehensive metamorphic rearrangements is the skin. Direct-developing frogs lack a free-swimming tadpole and hatch from terrestrial eggs as fully formed froglets. In the few species examined, development is characterized by the condensed and transient formation of some tadpole-specific features and the early formation of adult-specific features during a “cryptic” metamorphosis.ResultsWe show that skin in direct-developing African squeaker frogs (Arthroleptis) is also repatterned from a tadpole-like to an adult-like histology during a cryptic metamorphosis. This repatterning correlates with histological thyroid gland maturation. A comparison with data from the Puerto Rican coqui (Eleutherodactylus coqui) reveals that the evolution of direct development in these frogs is associated with a comparable heterochronic shift of thyroid gland maturation.ConclusionThis suggests that the development of many adult features is still dependent on, and possibly constrained by, the ancestral dependency on thyroid hormone signaling., Published by Wiley, New York, NY [u.a.]

Leptopelis viridis (G��nther, 1869) (Fig. 3D) COLLECTION MATERIAL. ��� Burkina Faso ��� 2♀ (SVL 38.7-41.7 mm); Kadiogo province, Pabr��; 12.51602��N, ��� 1.59691��W; 300 m a.s.l.; M. Lamotte leg.; MNHN-RA-1999.7060, 1999.7061. FIELD MATERIAL. ��� Burkina Faso ��� 10 ♂ (SVL 30.5-37.9 mm); Kadiogo province, Pabr�� dam; 12.51605��N, ��� 1.59689��W; 290 m a.s.l.; 12.VII.2017; H. J. Ayoro & A. Ohler leg.; bank of Pabr�� dam; hand capture ��� 3 ♂ (SVL 36.1-38.7 mm); Oubritenga province, Loumbila dam; 12.49829��N, ��� 1.40310��W; 275 m a.s.l.; 14.VII.2017; same collectors as for preceding; around a plantain field downstream from the Loumbila dam; hand capture ��� 5 ♂ (SVL 32.9-36.1 mm); Koubri, ���Ancien barrage���; 12.22883��N, ��� 1.34522��W; 290 m a.s.l.; 16.VII.2017; same collectors as for preceding; in a maize field on the bank of ���Ancien barrage��� ��� 1 ♀ (SVL 54.5 mm); Koubri, the inn���s court of Monastery; 12.21685��N, ��� 1.35041��W; 285 m a.s.l.; 17.VII.2017; same collectors as for preceding; under shrubs; hand capture ��� 1 ♂; Kadiogo province, town hall of Saaba; 12.37209��N, ��� 1.40374��W; 305 m a.s.l.; 30.VII.2018; H. J. Ayoro leg.; on shrubs near habitations; acoustic recording ��� 2 ♂ (SVL 33.9-36.5 mm); Arly National Park, Bouliabou pond; 11.57890��N, 1.46393��E; 170 m a.s.l.; 12. VI.2018; same collector as for preceding; on a shrub; hand capture ��� 6 ♂ (SVL 32.2- 33.8 mm); Arly National Park, Bongola swampy valley; 11.53255��N, 1.48244��E; 170 m a.s.l.; 13. VI.2018; same collector as for preceding; on a shrub; hand capture ��� 3 ♀ (SVL 39.3-42.3 mm); Arly National Park, a floodplain (500 m north of Bouliabou pond); 11.57852��N, 1.46000��E; 170 m a.s.l.; 16. VI.2018; same collector as for preceding; on the edge of a small water body; hand capture ��� 1 ♂ (SVL 33.8 mm), 2 ♀ (SVL 45.3-45.7 mm); W National Park, Kabougou River (500 m north-west of Kabougou forest post office); 11.97062��N, 2.01624��E; 280 m a.s.l.; 18. VI.2018; same collector as for preceding; on the edge of a pond; hand capture ��� 1 ♂ (SVL 34.4 mm), 1 ♀ (SVL 39.8 mm); W National Park, Nangouli swampy valley; 11.93663��N, 1.99461��E; 310 m a.s.l.; 21. VI.2018; same collector as for preceding; around a small pond; hand capture ��� 1 ♂; Deux Bal��s National Park, downstream from the Soumbou dam; 11.69831��N, ��� 2.79705��W; 260 m a.s.l.; 8.XI.2017; same collector as for preceding; hand capture ��� 2 ♂ (SVL 30. 8-31.2 mm), 1 ♀ (SVL 40.1 mm); Tuy province, Koti; 11.38783��N, ��� 3.06404��W; 300 m a.s.l.; 23.VIII.2018; same collector as for preceding; near a maize field; hand capture ��� 12 ♂ (SVL 31.3-36.4 mm), 2 ♀ (SVL 43.1-43.6 mm); Como��-L��raba Forest, Baboro swampy valley; 9.92523��N, ��� 4.57562��W; 240 m a.s.l.; 4.VIII.2018; H. J. Ayoro & A. Ohler leg.; on shrubs; hand capture ��� 3 ♂ (SVL 29.5-33.5 mm); Como�� province, bank of Folonzo village pond; 9.95775��N, ��� 4.67451��W; 240 m a.s.l.; 6.VIII.2018; same collectors as for preceding; on grasses, hand capture ��� 1 ♂ (SVL 35.6 mm), 1 ♀ (SVL 48.7 mm); Nasso, Kou Forest; 11.18812��N, ��� 4.44390��W; 320 m a.s.l.; 9.VIII.2018; same collectors as for preceding; near a small water body; hand capture ��� Ioba province, Bontioli reserve; 10.93685��N, ��� 3.13801��W; 260 m a.s.l.; 2011; J. Penner leg.; field direct observation. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Yako; 12.96740��N, ��� 2.26151��W; 330 m a.s.l.; 1961 (Schi��tz 1967; B��hme et al. 1996) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke 2011; Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke 2011; Mohneke et al. 2011). DESCRIPTION. ��� 58 specimens examined. Size of this species 33- 35 mm for males and 42-48 mm for females, according to Schi��tz (1967; 1999) and R��del (2000). Size slightly larger in our voucher specimens. Leptopelis with smoother skin, longer extremities and larger discs on toes and fingers (Fig. 3C, D). Dorsal colouration brownish or reddish brown with darker spots forming a mark of the shape of a reversed ���V���. Venter whitish. juveniles colour mostly green. For detailed descriptions, see Lamotte & Xavier (1981), Schi��tz (1967, 1999), R��del (2000) and Amiet (2012). HABITAT. ��� The species was fossorial during the day and arboreal at night. Leptopelis viridis was found in herbaceous and wooded savannahs (Fig. 2C, E). Males were caught on trees and shrubs around ponds and rivers at heights varying between 0.5- 3 m. In contrast to males, females were collected on the ground of the same habitats., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on pages 552-554, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","BOHME W., MEINIG H. & RODEL M. - O. 1996. - New records of amphibians and reptiles from Burkina Faso and Mali. British herpetological Society Bulletin 56: 7 - 26.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","SCHIOTZ A. 1999. - Treefrogs of Africa. Edition Chimaira, Germany, Frankfurt am Main, 350 p.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","LAMOTTE M. & XAVIER F. 1981. - Les Amphibiens de l'Afrique de l'Ouest, in DURAND J. R. & LEVEQUE C. (eds), Faune et Flore aquatiques de l'Afrique Sahelo-soudanienne, Editions de l'Office de la Recherche Scientifique et Technique d'Outre-Mer 2: 773 - 816.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p."]}

Arthroleptis poecilonotus Peters, 1863 (Fig. 3A, B) COLLECTION MATERIAL. ��� Burkina Faso ��� 10 subadults (SVL 12.1- 20.1 mm); Bobo-Dioulasso, near Nasso; 11.18803��N, ��� 4.29741��W; 420 m a.s.l.; VII.1970; M. Lamotte leg.; MNHN-RA-2008.1055, 2008.1056, 2008.1057, 2008.1058, 2008.1059, 2008.1060, 2008.1061, 2008.1062, 2008.1063, 2008.1064. FIELD MATERIAL. ��� Burkina Faso ��� 2 ♀ (SVL 27.6-28.1 mm), 5 ♂ (SVL 24.4-26.5 mm), 3 subadults (SVL 16.6-18.3 mm); Nasso, Kou Forest; 11.18888��N, ��� 4.43935��W; 320 m a.s.l.; 7-9.VIII. 2018; H. J. Ayoro & A. Ohler leg.; gallery forest; hand capture. DESCRIPTION. ��� 20 specimens examined. Small frogs with a blunt snout, short head and a concave tympanum; slender limbs, the inner metatarsal tubercle long and prominent. Adult males and females with protruding palmar and supernumerary tubercles on fingers. Tubercles absent on toes. Dorsum feebly granulated. In life, dorsum grey, brown or reddish-brown coloured. Sometimes an hourglass pattern present on back (Fig. 3A, B). Venter with whitish or brownish, smooth skin. As other Arthroleptis species (e.g. Blackburn 2009; Blackburn et al. 2010), presence of an elongated third finger, and transparent spines on second and third fingers in adult males. HABITAT. ��� The historical specimens and the recent samples of A. poecilonotus were from Kou Forest (Fig. 2E). At this site, males were seen at night on leaves of shrub and grass at heights varying from 0.2- 1 m. Females and subadults were seen hiding in leaflitter. The species may occur in other gallery forests along the major rivers (Mouhoun, Como�� and Pendjari rivers) in the western and southern part of Burkina Faso as well (Lamotte 1967b). It was found in a forest close to a waterfall near Pendjari National Park in Benin (Nago et al. 2006), in Togo (Segniagbeto et al. 2007), in Ivory Coast (Lamotte & Perret 1963; Lamotte 1967b; Barbault & Trefaut-Rodrigues 1979; R��del 2000; R��del 2003; R��del & Ernst 2003) and in Nigeria (Schi��tz 1964, 1967; Leach�� et al. 2006; Onadeko 2016). TAXONOMIC REMARK. ��� As stated in R��del & Bangoura (2004) and Blackburn (2010), it is currently not possible to assign these West African Arthroleptis taxa with certainty to one of the various available names, as intra-specific variability equals or surpasses interspecific differences. As actually understood, A. poecilonotus has an extended distribution range, including West, Central and parts of East Africa (Frost 2019) and probably recovers a group of species. Type locality is in West Africa, in south-eastern Ghana. Further studies, in particular including topotypical material, are needed for taxonomic decisions., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 552, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["BLACKBURN D. C. 2009. - Diversity and evolution of male secondary sexual characters in African squeakers and long-fingered frogs. Biological Journal of the Linnean Society 96: 553 - 573. https: // doi. org / 10.1111 / j. 1095 - 8312.2008.01138. x","BLACKBURN D. C., GVOZDIK V. & LEACHE A. D. 2010. - A new squeaker frog (Arthroleptidae: Arthroleptis) from the mountains of Cameroon and Nigeria. Herpetologica 66: 335 - 348. https: // doi. org / 10.1655 / HERPETOLOGICA-D- 10 - 00015.1","LAMOTTE M. 1967 b. - Les batraciens de la region de Gbakobo (Cote d'Ivoire). Memoire de l'Institut fondamental de l'Afrique noire (A) 29: 218 - 294.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90.","LAMOTTE M. & PERRET J. L. 1963. - Contribution a l'etude des batraciens de l'Ouest Africain XV - Le developpement direct de l'espece Arthroleptis poecilonotus Peters. Bulletin de l'Institut fondamental d'Afrique noire (A) 25: 277 - 284.","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","RODEL M. - O. & ERNST R. 2003. - The amphibians of Marahoue and Mont Peko National Parks, Ivory Coast. Herpetozoa 16: 23 - 39.","SCHIOTZ A. 1964. - A preliminary list of amphibians collected in Ghana. Videnskabelige Meddelelser fra dansk Naturhistorik Forening 127: 1 - 17.","SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","LEACHE A., RODEL M. - O., LINKEM C. W., DIAZ R. E., HILLERS A. & FUJITA M. K. 2006. - Biodiversity in a forest island: reptiles and amphibians of the West African Togo hills. Amphibian and Reptile Conservation 4: 22 - 45. https: // doi. org / 10.1514 / journal. arc. 0040018","ONADEKO B. A. 2016. - Distribution, diversity and abundance of anuran species in three different vegetations habitats in southwestern Nigeria. Ethiopian Journal of environmental Studies and Management 9: 22 - 34. http: // dx. doi. org / 10.4314 / ejesm. v 9 i 1.3","FROST D. R. 2019. - Amphibian Species of World: an online reference. Version 6.0. Available at htt: // research. amnh. org / herpetology / amphibia / index. html. Accessed on 15 February 2019 and 2 November 2020."]}

Leptopelis bufonides Schi��tz, 1967 (Fig. 3C) FIELD MATERIAL. ��� Burkina Faso ��� 8 ♀ (SVL 40.2-44.9 mm), 10 ♂ (SVL 30.5-37.5 mm); W National Park; 11.93664��N, 1.9946��E; 310 m a.s.l.; 21.VI.2018; H. J. Ayoro leg.; edge of savannah pond; hand capture ��� 1 ♀ (SVL 41.2 mm), 1♂ (SVL 32.1 mm); Tuy province, Koti; 11.38785��N, ��� 3.06404��W; 295 m a.s.l.; 23.VIII.2018; H. J. Ayoro leg.as for preceding;near small pond in bush; hand capture. LITERATURE REPORTS. ��� Burkina Faso ��� Passor�� province, Samba; 12.70143��N, ��� 2.40389��W; 300 m a.s.l.; 18.VIII.1961 (Schi��tz 1967) ��� Gourma province, Fada N���gourma; 12.06051��N, 0.34913��E; 300 m a.s.l. (Mohneke et al. 2011) ��� Ganzourgou province, Zorgho; 12.2483��N, ��� 0.6278��W; 300 m a.s.l.; I-III.2008 (Mohneke et al. 2011). DESCRIPTION. ��� 20 specimens examined. Leptopelis bufonides with small to moderate size and warty skin, stocky body shape, and short legs. In contrast to most other members of that genus, the fingers with small discs on their tips. In life colouration greyish brown with dark spots, including a dark interorbital spot and elongated spots or bands on the back (Fig. 3C). These markings making almost a ���m��� pattern. The venter light to dark grey, with a dark throat in both sexes. For detailed descriptions, see Schi��tz (1967), R��del (2000) and Amiet (2012). HABITAT. ��� This species was usually found in grassy and wooded savannahs (Fig. 2C, D). Specimens were collected from the ground around temporary ponds with grass banks and shrubs. It was also Evans from dry savannahs in Senegal (Schi��tz 1967), Mali (B��hme & Heath 2018), Ghana (Schi��tz 1967), Benin (Nago et al. 2006), Togo (Segniagbeto et al. 2007) and Cameroon (Amiet 2012)., Published as part of Ayoro, Halamoussa Jo��lle, Segniagbeto, Gabriel Hoinsoud��, Hema, Emmanuel Midibahaye, Penner, Johannes, Oueda, Adama, Dubois, Alain, R��del, Mark-Oliver, Kabr��, Gustave Boureima & Ohler, Annemarie, 2020, List of amphibian species (Vertebrata, Tetrapoda) of Burkina Faso, pp. 547-582 in Zoosystema 42 (28) on page 552, DOI: 10.5252/zoosystema2020v42a28, http://zenodo.org/record/4265439, {"references":["SCHIOTZ A. 1967. - The treefrogs (Rhacophoridae) of West Africa. Spolia zoologica Musei hauniensis 25: 1 - 346.","MOHNEKE M., ONADEKO A. B. & RODEL M. - O. 2011. - Medicinal and dietary uses of amphibians in Burkina Faso. African Journal of Herpetology 60: 78 - 83. https: // doi. org / 10.1080 / 21 564574.2011. 564660","RODEL M. - O. 2000. - Herpetofauna of West Africa: Amphibians of the West African savannah. Edition Chimaira. Vol. 1. Germany, Frankfurt am Main, 300 p.","AMIET J. L. 2012. - Les rainettes du Cameroun. Edition J. L. Amiet et La Nef des Livres, Nyons & Saint Nazaire, France, 591 p.","BOHME W. & HEATH J. 2018. - Amphibian and reptilian records from south-central Mali and western Burkina Faso. Bonn zoological Bulletin 67: 59 - 69.","NAGO A. G. S., GRELL O., SINSIN B. & RODEL M. - O. 2006. - The amphibian fauna of Pendjari National Park and surroundings, northern Benin. Salamandra 42: 93 - 108.","SEGNIAGBETO G. H., BWOSSIDJAOU J. E., DUBOIS A. & OHLER A. 2007. - Les amphibiens du Togo: etat actuel de connaissance. Alytes 24: 72 - 90."]}

We related the first commented list of the amphibian species of Burkina Faso. To obtain contemporary data, we investigated six sites from July 2017 to September 2018. The survey is unique for West Africa in combining a variety of different habitat types, e.g. Sahelian areas, grassland and woodland savannahs, floodplains, gallery forests and agricultural sites. Animals were found at varying spatial scales by opportunistic visual encounter surveys after sunset supplemented with acoustic searching. Further data were based on museum vouchers collected between the years 1959 and 2011 and deposited in European and American museums, and the literature review focusing on the country species. We listed 36 anuran species from 11 families. Surprisingly seven taxa, namely Arthroleptis poecilonotus Peters, 1863, Afrixalus vittiger (Peters, 1876), Afrixalus weidholzi (Mertens, 1938), Kassina cassinoides (Boulenger, 1903), Ptychadena schillukorum (Werner, 1908), Hyperolius lamottei Laurent, 1958 and Amnirana albolabris (west) (Jongsma et al. 2018), were new country records. A few rare, hard to record or little known species, including Arthroleptis poecilonotus, Hyperolius lamottei, Xenopus tropicalis (Gray, 1864) and Amnirana albolabris (west) can be confirmed. The origin of a specimen of Sclerophrys mauritanica (Schlegel, 1841) mentioned as coming from Burkina Faso deposited in the collections of MNHN must be considered as an error. We briefly discussed the taxonomy and indicated the habitats of all species mentioned in this country so far. The discovery of several new country records highlights that the amphibian fauna of Burkina Faso is still incompletely known and that many poorly explored sites may still harbour a high diversity.

Cases of parallel evolution offer the possibility to identify adaptive traits and to uncover developmental constraints on the evolutionary trajectories of these traits. The independent evolution of direct development, from the ancestral biphasic life history in frogs is such a case of parallel evolution. In frogs, aquatic larvae (tadpoles) differ profoundly from their adult forms and exhibit a stunning diversity regarding their habitats, morphology and feeding behaviors. The transition from the tadpole to the adult is a climactic, thyroid hormone (TH)-dependent process of profound and fast morphological rearrangement called metamorphosis. One of the organ systems that experiences the most comprehensive metamorphic rearrangements is the skin. Direct-developing frogs lack a free-swimming tadpole and hatch from terrestrial eggs as fully formed froglets. In the few species examined, development is characterized by the condensed and transient formation of some tadpole-specific features and the early formation of adult-specific features during a “cryptic” metamorphosis. In this study we show that skin in direct-developing African squeaker frogs (Arthroleptis) is also repatterned from a tadpole-like to an adult-like histology during a cryptic metamorphosis. This repatterning correlates with an increase of thyroid gland activity. A comparison with data from the Puerto Rican coqui (Eleutherodactylus coqui) reveals that direct development might have evolved in parallel in these frogs by a comparable heterochronic shift of thyroid gland activity. This suggests that the development of many adult-features is still constrained by the ancestral dependency on thyroid hormone signaling.

Adult males of the African treefrog species Leptopelis flavomaculatus occur in either brown or green color morphs. In this study, we investigated whether the two color morphs of breeding males of L. flavomaculatus differ in traits other than color. We examined call differences (dominant frequencies and call durations), call-site selection, and body size. Our results show differences in the call durations, dominant frequencies, mean call intensities, as well as in size (body length and body mass) of the two vocally active color morphs of this species. The two morphs were similar in their choice of plant species used as calling sites and the heights of those sites. Given the sensitivity of mate recognition systems for most anurans, our results provide evidence of both call and morphological variation between the two color morphs of this species. Our findings establish a foundation for future phylogenetic and mating system studies to support the contention that the two color morphs of L. flavomaculatus might warrant taxonomic recognition as separate species.

Direct development has evolved independently several times in anurans and direct-developing species are characterized by large-scale developmental repatterning and a complete, or near complete, absence of most tadpole-specific structures. Earlier studies stressed the similarities among different direct-developing species, but more recent studies have indicated differences in the reduction of tadpole-specific structures among different taxa. Here, we describe egg deposition, clutch characteristics and embryonic development of the direct-developing squeaker frogs of the genus Arthroleptis, providing the first detailed description of direct development in Arthroleptidae. Embryonic development in Arthroleptis is characterized by the presence of an opercular fold that still encloses the developing forelimbs, the absence of external gills and an only moderately extended tail. A comparison with published information on other direct-developing anurans reveals broad dissimilarities in the formation of an opercular fold and very different tail morphology among different taxa. An egg tooth, often considered characteristic of direct-developing anurans, seems to be restricted to New World Terrarana. The embryonic diversity seen in direct-developing anuran taxa argues against simplistic assumptions about the evolution of direct development.

Herein, we describe the tadpoles of six Astylosternus species, A.fallax, A.cf.fallax, A.laurenti, A.montanus, A.perreti, A.ranoides, and Scotoblepsgabonicus, and redescribe the tadpoles of A.batesi, A.diadematus, A.laticephalus, A.occidentalis, A.rheophilus, and Nyctibatescorrugatus. All Astylosternus tadpoles are adapted to torrent currents and share a long, oval body, slightly flattened in lateral view, with very long muscular tails with narrow fins. The jaws are massive, serrated, and often show a tooth-like medial projection (fang) in the upper jaw. Body proportions of Astylosternus tadpoles are extremely similar. The best characters to distinguish species might be life coloration and potentially the shape of labial papillae. The tadpole of Scotoblepsgabonicus is similar to Astylosternus and differs only slightly by a narrower body with a shorter and rounder head. The upper jaw of Scotobleps carries two or three lateral fangs instead of one medial one. The tadpole of Nyctibatescorrugatus is easily distinguishable from the other two genera on the basis of their very long, eel-shaped body and tail and the bluish-black color.

Objective: The aim of this study was to make a systematic inventory of Anuran species in three agroindustrial zones of South-East Cote d'Ivoire, with a view to making a contribution to the knowledge of amphibians in Cote d’Ivoire. Method and Results: Anurans were captured using catch boxes and dip nets. Species identification was done using the determination keys proposed by Rodel (2000a) and Fretey et al . (2011). Twenty-eight (28) species belonging to 12 genus and 10 families have been inventoried.The family Ptychadenidae is the most diversified with 6 species. They are followed by Hyperoliidae and Phrynobatrachidae both having 5 species each. Arthroleptidae and Bufonidae contain 4 species and 3 species respectively. Dicroglossidae, Hemisotidae, Pipidae, Pyxicephaliidae and Ranidae are the least diversified with one (1) species each. Three unidentified species have also been inventoried: Artroleptis sp, Sclerophrys sp and Phrynobatrachus sp. It appears also 6 species are common to all the three plantations while 1, 4 and 8 species are specific to coconut plantation, palm plantations and banana plantations respectively. Conclusion and application: This study has made it possible to make a qualitative inventory of the Anura in three Agro-industrial zones of the South-eastern Cote d’Ivoire. These results will serve as a reference for further investigations of the amphibians and the development of appropriate measures to ensure better protection and sustainable exploitation of Anuras in Cote d’Ivoire. Keywords: Anurans, qualitative inventory, diagnosis, agro-industrial zone, Cote d’Ivoire

We studied the effects of forest management on amphibian communities in the Budongo Central Forest Reserve, Uganda. We sampled amphibians from May to August of 2012 in four compartments with different logging and arboricide-treatment histories. We used pitfall traps with drift fences combined with visual encounter surveys to sample amphibians from 36 plots in four 1-km long transects along the Sonso River. From 126 encounters across plots, we recorded 25 frog species belonging to six families and eight genera. Arthroleptidae was the most diverse family represented by 10 species within two genera. Arthroleptis had the highest number of species (six), Ptychadena the second most (five), followed by Leptopelis (four) and Sclerophrys (four). Species composition differed across transects. The unlogged study site possessed the highest species richness, diversity, and evenness, and had the greatest frequency of species encounters. The most heavily logged site had the lowest species diversity and fewest amount of species encounters. This site also had the most dissimilar species composition among sites and was significantly different in species richness compared to the unlogged site. The two moderately logged sites had the second and third most species, and had the most similar species composition to each other. Our study provides data on the amphibian species of a protected site in the Albertine Rift, part of the Eastern Afromontane biodiversity hotspot, and results suggest that the forest management regimes in Budongo have exerted an influence over the amphibian communities after more than 50 years of forest recovery.

We describe two new frog species of Cardioglossa (Family Arthroleptidae) from Central Africa. The new species are found in the rainforests of western-central Democratic Republic of the Congo and the Republic of the Congo near the Gabonese border, respectively. We demonstrate that these species are morphologically and genetically distinct from each other and all other species of Cardioglossa. Both new species lack the dorsal hour-glass pattern present in many species of the genus, but they can be distinguished from each other and related species by distinctive colour patterns on their lateral surfaces and extremities. Both new species most closely resemble C. gratiosa, which occurs in the Atlantic coastal forests extending from Cameroon through Gabon. The new species can be differentiated from C. gratiosa by the absence of black transverse bars on all limbs or by distinctive lateral colouration. Analysis of mitochondrial ribosomal 16S DNA sequences reveals low to moderate levels (1.9–7.5%) of diver...

We describe a new species of Leptopelis from the L. modestus complex. The new species is described from forest and lacustrine habitats of the Albertine Rift of Central Africa, is superficially similar to topotypic L. modestus from Cameroon, but it is distinguished by advertisement call, morphology, and sequences of 16S, cyt b, and RAG1. The new species is yet another amphibian that occurs in the Itombwe Plateau, further solidifying the position of the plateau as an area of conservation concern.

We describe the tadpoles of two closely related frog species of the genus Leptodactylodon (family Arthroleptidae), L. axillaris and L. perreti, that are restricted to the mountains of Cameroon. Tadpoles were collected from near the type locality of each species and identified using DNA sequence data. The tadpoles of both species are stream-dwelling and cryptic, often found in clumps of debris collected in the course of the stream, and orient roughly vertically in the water column with their mouths at or near the surface. As in L. ventrimarmoratus, the only other species in the genus described in detail, the tadpoles of these two species have a prominent umbelliform and up-turned oral disc, robust keratinous jaw sheaths, large lateral sacs along the elongate and dorsoventrally depressed body, and a robust elongate tail with short fins. Tadpoles of the two species are broadly similar morphologically. However, differences in the jaw sheaths may enable discrimination, including more pronounced laterally placed caniniform projections and more small needle-like serrations in the central region of the lower jaw sheath in L. perreti.

The species diversity of the frog genus Cardioglossa (family Arthroleptidae) is concentrated in the Lower Guinean Forest Zone of Central Africa with most of the 19 species occurring in Cameroon and neighboring countries (Amiet 1972a,b; Blackburn 2008; Hirschfeld et al. 2015). These small leaf-litter frogs are typically found in primary or secondary forest, have shrill whistling calls, are characterized by a variety of color patterns, and lay terrestrial eggs that hatch and develop into elongate, stream-adapted tadpoles (Amiet 1972a,b, 1973; Rödel et al. 2001; Hirschfeld et al. 2012). One of the most poorly known species—the Bururi Long-fingered Frog Cardioglossa cyaneospila Laurent, 1950—is also among the most geographically peripheral to the rest of the species diversity. To date, it is known only from two locations in Burundi and four in eastern Democratic Republic of Congo, regions in which armed conflicts have long hampered scientific research. In this short contribution, we (1) document both new and long unpublished records of C. cyaneospila, associate these with known museum records, and extend its geographic range, (2) highlight fruitful areas for future field surveys based on predicting an environmental envelope for this species, and (3) summarize what little is known of its natural history Fil: Blackburn, David C.. University of Florida; Estados Unidos Fil: Boix, Christian. Africa Geographic; Sudáfrica Fil: Greenbaum, Eli. University of Texas at El Paso; Estados Unidos Fil: Fabrezi, Marissa. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Salta. Instituto de Bio y Geociencias del NOA. Universidad Nacional de Salta. Facultad de Ciencias Naturales. Museo de Ciencias Naturales. Instituto de Bio y Geociencias del NOA; Argentina Fil: Meirte, Danny. Royal Museum of Central Africa; Bélgica Fil: Plumptre, Andrew J.. Wildlife Conservation Society; Estados Unidos. Conservation Science Group; Reino Unido Fil: Stanley, Edward L.. University of Florida; Estados Unidos

Herein is presented a list of the reptiles and anurans from the Parc National des Volcans (PNV) (01°43’ S, 29°52’ W), an area in the west and north provinces of the Republic of Rwanda in the Albertine Rift region of Africa. Fieldwork was conducted between two and six days per week from June through August 2007 and 2008. We also conducted literature searches of all historical expeditions within the park for species records. Seventeen species of reptiles and anurans are recorded from the PNV. Nine of the species were anurans, distributed in five families: Arthroleptidae (3), Bufonidae (1), Hyperoliidae (3), Phrynobatrachidae (1), and Pipidae (1). Eight species of reptiles were recorded from five families: Chamaeleonidae (1), Lacertidae (2), Scincidae (2), Colubridae (2), and Viperidae (1). Eight of the seventeen species found in the PNV are endemic to the Albertine Rift.

A new species of squeaker frog (Arthroleptis) is described from Ngozi Crater in the Poroto Mountains of southwestern Tanzania. The holotype, and only specimen, was collected 80 years ago but remained undescribed because of past taxonomic confusion relating to large-bodied species of Arthroleptis from eastern Africa. Morphological study, including multivariate analysis of measurement data, indicates that this new species is distinguishable from other species of Arthroleptis, including large-bodied species such as Arthroleptis adolfifriederici, Arthroleptis affinis, Arthroleptis nikeae, Arthroleptis nguruensis, and Arthroleptis tanneri. The new species differs from other described species of Arthroleptis by the combination of its large body size, relatively long toes, and relatively small tympanum. The description of this new species further highlights the biological diversity of the Southern Rift Mountains.

We examined the systematics of two frequently confused Albertine Rift endemic treefrogs, Leptopelis karissimbensis and Leptopelis kivuensis. One mitochondrial (16S) and one nuclear (BDNF) gene were sequenced to demonstrate genetic differences between the two species. Although we did not find morphometric differences between the two taxa, several characters can be used to diagnose each species, including coloration (adult male throat), ventral pigmentation, presence/absence of a slight heel spur in large individuals, and male advertisement calls. The geographic distribution of each species is revised based on our new locality records and reconsideration of historical records from the literature. We recommend that L. karissimbensis be downgraded in conservation status to near threatened, because it does not meet the IUCN restricted geographic distribution criteria of an endangered species.

Amphibians of the Eastern Arc Mountains are poorly known, apart from those of the East Usambara, Uluguru, and Udzungwa Mountains. Here, we detail specimens collected in the northwestern Eastern Arc Mountains (West Usambara, South Pare, and North Pare) over the past 100 years. Although not comprehensive, the list is compiled from specimens held at institutions in Europe and the United States and in personal collections; it includes most of the specimens known from this area. From these specimens, we describe one new species of the frog genus Arthroleptis and discuss other specimens that may represent new species. We add substantially to the known diversity in these areas. Using the compiled species list, we assess species diversity and relate the differences to sampling bias and possible biogeographic differences among the mountains. The future preservation of amphibians across this area is of concern given the increasing loss of forest.

A new species of squeaker frog (genus Arthroleptis) is described from Pico Basile on Bioko Island (Equatorial Guinea), which lies just off the coast of Cameroon in the Gulf of Guinea. A previous molecular phylogenetic study indicates that this new species is nested within a species complex—Arthroleptis poecilonotus—that comprises morphologically indistinguishable populations that represent several cryptic species. The new species from Bioko is, however, easily distinguishable from both closely related and geographically proximate Arthroleptis by a combination of its moderately large body size (approximately 25% larger than frogs in the remaining poecilonotus species complex) and unique ventral color pattern. It is further distinguished from all other Arthroleptis by the presence of spines on the crus in large males and from other populations in the poecilonotus species complex by the occasional presence of a supernumerary tubercle at the base of pedal digit II. Statistical analyses of measurement...

A new species of squeaker frog (genus Arthroleptis) is described from the northern mountains of the Cameroon Volcanic Line, a species-rich region within the West African biodiversity hotspot. This new species is associated with human-altered landscapes, including farms and grasslands, and is known from the highland areas near Bamenda, including Mt. Oku, Cameroon, as well as the Obudu and Mambilla Plateaus of eastern Nigeria. The known distribution suggests that it may eventually also be found throughout the Acha- Tugi Ridge that traverses the Cameroon-Nigeria border. More than 40 years ago, a specimen from one of these populations was recognized as probably representing a new species, but no further work was published on this material. We posit that the taxonomy of this new species has proven problematic because of morphological similarity to Arthroleptis poecilonotus. Although in some ways resembling A. poecilonotus ,a molecular phylogeny based on mitochondrial gene sequence data demonstrates that the new species is not closely related to A. poecilonotus and, instead, forms a clade with A. krokosua, A. perreti, and A. variabilis. The new species is diagnosable by the combination of medium-small body size (24-29 mm snout-vent length in adult females), the color patterns of the venter and head, and a relatively stout hindlimb with a prominent inner metatarsal tubercle. The description of this new species further highlights these mountains as an important region of endemism within the Cameroon Volcanic Line.

Ethiopia harbours 9 species of threatened amphibians, all of which inhabit the high- lands. Over the last decades, there has been a rapid increase in deforestation and habitat degrada- tion in Ethiopia, with the result that the last undisturbed refuges for forest-dwelling anurans in the mountain areas have shrunk noticeably, especially outside of protected areas. In the present study, we used a maximum entropy ecological niche modelling approach to model the potential distribution of 2 poorly known Leptopelis species. The models were derived from climate and land cover data at known occurrences of the species (i.e. data from a number of natural history museums). These mod- els can be used to select the priority areas most critical for future protection and may guide further field surveys to accelerate the discovery of unknown populations.

A new small West African Arthroleptis from the rainforest at the south-eastern tip of Mount Nimba, Liberia is described. The new species differs from all known congeners by the combination of a unique color pattern, a rounded canthal region and a completely smooth skin. The comparison of 592 base pairs of the mitochondrial 16S rRNA from the new species with Arthroleptis samples from all over West Africa, revealed genetic distances between 13.4–17.9%. From the morphologically similar A. aureoli the new species differed by 17.9%. Future research may reveal the necessity to establish a new genus for this peculiar new frog.

Neobatrachia is a phylogenetically-nested anuran group and contains approximately 85% of amphibian species. It is difficult to analyze the neobatrachian mitochondrial (mt) genome due to accelerated nucleotide substitution rates and occasional gene rearrangements. To overcome these problems, we designed 15 PCR primers. These primers, together with two primers from a previous study, could amplify almost the entire mt genome in all neobatrachians tested (seven species from seven different families) without optimizing PCR conditions. Among the tested taxa, Trichobatrachus robustus (Arthroleptidae) and Hyperolius viridiflavus (Hyperoliidae) showed unique amplification patterns, suggesting occurrences of gene duplication and/or rearrangements in these mt genomes.

The previously unreported tadpole of Leptopelis karissimbensis, an endangered treefrog from Rwanda, is described. Tadpoles were collected, photographed, measured, and examined for standard metrics of tadpole morphology. Larvae of L. karissimbensis resemble other tadpoles in the genus Leptopelis in being muscular, elongate, and eel-like. The lateral tooth row formula for L. karissimbensis is 4/3. In late stage larvae, tadpoles of L. karissimbensis exhibit a prominent white spot below the eye. Larvae of this species were often abundant in suitable habitat at approximately 2,800 m in elevation. Larval habitat for L. karissimbensis includes seasonally flooded marshes, forest pools, and permanent ponds in the Virunga Mountains, the only known range of this species.

Biodiversity hotspots are centres of endemism and thus contain many range-restricted species. In addition, within these hotspots occur widespread species that might have originated within a hotspot before dispersing to neighbouring or distant regions. We test this hypothesis with a phylogeographic analysis of a miniature leaf litter frog, Arthroleptis xenodactyloides, that has a large distribution throughout the Eastern Arc biodiversity hotspot and other regions in East Africa. Maximum-likelihood and Bayesian estimates of the mitochondrial gene phylogeny are used as a proxy for understanding the evolutionary history of diversification and the historical relationships between populations. The north-south range of this species extends for approximately 1900 km; our sampling covers approximately 85% of this range. Using phylogenetic comparative methods, we estimate the region of origin and direction of dispersal within A. xenodactyloides. We compare contrasting hypotheses of latitudinal range expansion using bayes factors. The ancestral region of origin of A. xenodactyloides is reconstructed as having occurred within the Eastern Arc before dispersing southwards into the southern Rift Mountains, probably in the Pleistocene. The phylogeographic structure within this leaf litter frog is surprisingly similar to that of forest birds, revealing that similar geographic features might have had a driving role in diversification of these very dissimilar taxa. Latitudinal expansion occurred early in the evolutionary history of A. xenodactyloides, which may indicate that physiological adaptation facilitated its wide geographic distribution.

The African frog genera Arthroleptis and Cardioglossa are unique among vertebrates in having males with extremely long third fingers. In some species, this sexual dimorphism is impressive, with male third fingers approaching 40% of body length. However, the diversity of this trait has not been documented thoroughly and several species appear to lack this trait. The present study documents the diversity of male secondary sexual traits in Arthroleptis and Cardioglossa, including elongate third fingers and digital and inguinal spines. Furthermore, it explores hypotheses of trait evolution, including explanations for the absence of male traits. Analyses of covariance suggest that the functional relationship between finger length and snout–vent length (SVL), both within and among species, is different for male finger III than for male fingers I, II, and IV, or for female finger III. Ancestral state reconstruction suggests that all male traits were present in the most recent common ancestor of Arthroleptis and Cardioglossa and that reduction or loss of traits occurred later. Across species, independent contrast analyses find no correlation between SVL and either male relative third digit length or dimorphism in relative third digit length. The number of spines on male fingers II and III are positively correlated but spine number is not correlated with SVL and only weakly correlated with relative third digit length. The diversity of male traits is evolutionarily labile and is not explained by simple hypotheses of character evolution. Arthroleptis and Cardioglossa may thus provide an interesting study system for understanding how changes in sexual selection forces produce male trait diversity. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96, 553–573.

The taxonomic status of Hylambates rufus aubryioides Andersson, 1907, a taxon rarely treated in the literature and considered to represent a junior synonym of Leptopelis modestus, is re-evaluated. The taxon is redescribed and a lectotype is designated. Morphological comparisons revealed that it is neither conspecific with L. modestus, nor with L. calcaratus, as suggested by an unpublished identification tag. Instead, no morphological characters distinguish the taxa L. aubryioides and L. omissus. Leptopelis aubryioides (Andersson, 1907) is considered a valid species and Leptopelis omissus Amiet, 1992 is placed as its junior synonym.

We describe a new species of squeaker frog (Arthroleptis) from Mt. Manengouba in southwestern Cameroon. The new species is distinguished from other Cameroonian Arthroleptis by moderately larger body size; a darkened throat and posterior thigh, both with many white spots; and, in females, a fourth finger longer than the first and second fingers. This species corresponds to a Cameroonian taxon previously identified as Arthroleptis adolfifriederici 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.

A new frog species of the genus Cardioglossa, that is probably restricted to less than 10 km2 of gallery forest near the summit of Mount Manengouba in the Republic of Cameroon, is described. Unlike all other Cardioglossa, in life the coloration of the new species is typically brownish red, with no dorsal markings, but with a pair of thin, golden white lines that extend posteriorly from the tip of the rostrum and terminate just above and behind the tympanum. This new species is morphologically similar to C. oreas, another high-altitude species endemic to the Cameroonian mountains: both have small tympana and lack the hypertrophied third manual digit typical of males of other Cardioglossa. The tadpole of the new species is similar to that of C. occidentalis, the only other Cardioglossa for which the tadpole is fully described, but differs in having a broad anterolaterally projecting supraoral labium, lacking ventral pigmentation, and having a relatively longer spiracular tube. Several features of tadpole morphology are convergent with the unusual fossorial tadpole of the microhylid Otophryne and suggest a similar convergence in ecology. Principal components analysis of morphometric data, followed by discriminant function analysis, was used to explore patterns of morphological similarity within Cardioglossa. These analyses provide support for five species groups: (1) C. aureoli (distinct from other Cardioglossa); (2) C. escalerae, C. gratiosa and C. nigromaculata; (3) C. gracilis, C. melanogaster and C. schioetzi; (4) C. sp. nov. and C. oreas; and (5) C. cyaneospila, C. pulchra, C. venusta and C. trifasciata. The inclusion of C. cyaneospila in the last group supports previous suggestions of past biotic connections between the Albertine Rift mountains and those of the Cameroon Volcanic Line. The description of this new species further emphasizes Mt Manengouba as a centre of anuran endemism within the Cameroonian mountains and reaffirms the importance of this ecoregion within sub-Saharan Africa. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154, 611–630.

Typically small body size and similar external anatomy have made it difficult for researchers to distinguish between species of two distantly related, but often sympatric, African ranoid frog genera: Arthroleptis (Arthroleptidae) and Phrynobatrachus (Phrynobatrachidae). We define a suite of external morphological characters, extending beyond the traditionally used secondary sexual characteristics, that can be used to definitively distinguish between adult Arthroleptis and Phrynobatrachus. Photographs comparing absence, presence, and variation of morphological characters are included for clarification. Significant differences between body proportions (head width/snout–vent length) are also observed between these genera. Although smaller species might be more difficult to identify, larger species (> 30 mm snout–vent length) of Arthroleptis can be distinguished from Phrynobatrachus because of their relatively wider heads. We intend this to serve as a heuristic guide for both field- and museum-based ...

The validity of the name Hylambates hyloides is being examined. This name was commonly used for a small West African forest Leptopelis since the 1960s. In this paper it is shown that Leptopelis hyloides is a synonym of L. viridis, a widespread West African savanna frog. Consequently this forest taxon is being described as a new Leptopelis species. It ranges throughout the whole Upper Guinean forest zone from the Niger Delta to western Guinea. It inhabits primary and partly degraded evergreen and semi-deciduous forests. It is compared to all valid West African Leptopelis species, as well as to all other available names. The new species is characterized by the combination of a compact body shape with a head wider than the body in males, small size, large distinctively coloured protruding eyes, large tympanum, almost smooth skin, comparatively long legs, a white line on the distal parts of arms and feet, webbing between the 2–4 fingers, well developed webbing on the feet, and a buzzing advertisement call. (© 2007 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Leptopelis spiritusnoctis R��del 2007 Mitteilungen aus dem Museum f��r in Berlin, Supplement, 83 (Supplement): 91 ��� 00; Fig. 1, 2 A���E. Paratype: PEM A 7917; Haute Dodo Forest Reserve (04�� 59 ' 14 " N, 0 7 �� 19 ' 30 " W), Cote d'Ivoire; W.R. Branch and M.- O. R��del, 15 June 2002. Remarks. Left fourth and fifth toe removed as genetic sample. Holotype in ZMB (699511)., Published as part of Conradie, Werner, Branch, William R. & Watson, Gillian, 2015, Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 1: Amphibians in Zootaxa 3936 (1), DOI: 10.11646/zootaxa.3936.1.2, http://zenodo.org/record/237278, {"references":["Rodel, M. - O. (2007) The identity of Hylambates hyloides Boulenger, 1906 and description of a new small species of Leptopelis from West Africa. Mitteilungen aus dem Museum fur Naturkunde in Berlin, Zoologische Reihe, 83 (Supplement), 90 - 100. http: // dx. doi. org / 10.1002 / mmnz. 200600031"]}

Arthroleptis bequaerti Barbour and Loveridge 1929 a Proceedings of the New England Zo��logical Club, 11: 25. Current name: Phrynobatrachus bequaerti (Barbour and Loveridge, 1929) Paratypes (2): PEM A 8518���8519 (formerly AMG 6741); Mt. Vissoke, Belgian Congo, Democratic Republic of the Congo; J. Dencherd, 1927. Remarks. The original type series consisted of 25 specimens: The holotype and 10 remaining paratypes remain in MCZ. Numerous paratypes have been exchanged: three to NMB, five to FMNH (Marx 1958), one each to ZMB, UMMZ, UIMNH (Smith et al. 1964), and BMNH. No reference is made in Barbour and Loveridge���s (1946) catalogue on the exchange of two paratypes (MCZ A- 14771 and A- 14772) to AMG or, for that matter, to any other museum. No documentation relating to the transfer survives from AMG. Transferred from Arthroleptis to Pararthroleptis by Deckert (1938), and subsequently to Phrynobatrachus by De Witte (1941)., Published as part of Conradie, Werner, Branch, William R. & Watson, Gillian, 2015, Type specimens in the Port Elizabeth Museum, South Africa, including the historically important Albany Museum collection. Part 1: Amphibians in Zootaxa 3936 (1), DOI: 10.11646/zootaxa.3936.1.2, http://zenodo.org/record/237278, {"references":["Barbour, T. & Loveridge, A. (1929 a) A new frog of the genus Arthroleptis from the Belgian Congo. Proceedings of the New England Zoological Club, Cambridge, Massachusetts, 11, 25 - 26.","Marx, H. (1958) Catalogue of the type specimens of reptiles and amphibians in the Chicago Natural History Museum. Fieldiana, Zoology, 36 (4), 409 - 496.","Smith, H. M., Langebartel, D. A. & Williams, K. L. (1964) Herpetological type-specimens in the University of Illinois Museum of Natural History. Illinois biological monographs, 32, 1 - 80.","Deckert, K. (1938) Beitrage zur Osteologie und Systematik ranider Froschlurche. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin, 1938, 127 - 184.","De Witte, G. F. (1941) Batraciens et reptiles. Exploration du Parc National Albert. Fascicule 33. Mission G. F. de Witte (1933 - 1935), Bruxelles, v + 261 pp."]}