Tuesday, November 14, 2023

[Herpetology • 2023] Poyntonophrynus fernandae & P. nambensis • Two New Poyntonophrynus Species (Anura: Bufonidae) highlight the importance of Angolan Centers of Endemism


Poyntonophrynus fernandae 
Baptista, Vaz Pinto, Keates, Lobón-Rovira, Edwards & Rödel, 2023
  
Namba pygmy toad | sapo pigmeu da Namba ||  DOI: 10.3897/vz.73.e103935

The pygmy toad genus Poyntonophrynus is endemic to southern Africa. The morphology of these small toads is conserved. They are usually dully colored, and are predominately adapted to arid conditions. During recent surveys in Angola we found Poyntonophrynus specimens that were not assignable to known species. Using an integrative approach, based on mitochondrial and nuclear DNA, morphology, osteology, biogeography and ecology, we identified three new lineages, and describe two of them as new species. All three lineages are closely related to P. pachnodes, an Angolan endemic species, but they are geographically isolated from it. The new species are morphologically distinguishable, and are associated with two of the most important Angolan centers of endemism: the western escarpment and the central highlands. In order to get a more comprehensive understanding of the osteology of the genus, we also provide an osteological characterization of P. dombensis, which was not available to date. Our findings i) increase the number of earless species in the genus Poyntonophrynus, ii) emphasize southwestern Africa as the cradle of diversification in this genus, iii) report the occurrence of Poyntonophrynus in humid environments, thus showing that these toads are ecologically more variable than previously thought, and iv) underline the importance of further biodiversity studies in Angolan centers of endemism.

Keywords: Amphibia, Angolan escarpment, Angolan highlands, character displacement, pygmy toad, sexual dichromatism, speciation, syntopy

Poyntonophrynus habitats.
A, B Habitat of Poyntonophrynus fernandae sp. nov. at Congulo forest, Cuanza-Sul Province.
C, D Habitat of P. cf. fernandae and P. nambensis sp. nov. at Namba highlands, Cuanza-Sul Province.
E, F Habitat of P. pachnodes at Serra da Neve, Namibe Province.
Photos by K. Luchansky (A) and W.R. Branch (D).

Pictures of live Poyntonophrynus fernandae sp. nov.
 A Female holotype (ZMB 91791). B, C Male paratype (ZMB 91790) with yellow color when collected, and with fading coloration after three days. D Female paratype (BMNH 2021.7534), darker individual collected in forest. E Female paratype (FKH-1086) collected on red gravel, typical of Cuanza-Sul Province ferralitic soils. F Pair in axillary amplexus from Gabela (not collected).
Photos by L. Verburgt (E) and N.T. Ngan Thanh (F).

Poyntonophrynus fernandae sp. nov.

Diagnosis: Poyntonophrynus fernandae sp. nov. sensu lato differs from all Poyntonophrynus species except P. pachnodes in lacking a tympanum and a columella. It differs from P. pachnodes in having a better-developed neopalatine. Males differ from P. pachnodes and P. nambensis sp. nov. in dorsal coloration (partially or plain bright yellow vs. complex dull patterns in P. pachnodes, and P. nambensis sp. nov.). It differs from P. nambensis sp. nov. in being smaller, less sturdy, and having relatively longer fore and hindlimbs, in ventral coloration (speckles vs. nearly immaculate). It differs from P. beiranus in parotoid glands conspicuousness (conspicuous, with clearly demarcated margins vs. inconspicuous). It differs from P. damaranus in ventral patterning (speckles vs. immaculate). It differs from P. dombensis in tympanum (not visible vs. conspicuous), and ventral patterning (speckles vs. immaculate). It differs from P. fenoulheti in parotoid glands width (thinner or around same width of eye diameter vs. wider than eye diameter). It differs from P. grandisonae in tympanum (not visible vs. conspicuous) and skin texture (rough vs. leathery). It differs from P. grindleyi in pale occipital and sacral patches (present vs. absent) and dorsal spines (small vs. large). It differs from P. hoeschi in ventral patterning (speckles vs. immaculate). It differs from P. jordani in shape of parotoid glands (kidney-shaped vs. a cluster of glands) and pale occipital patch (present vs. absent). It differs from P. lughensis in conspicuousness of parotoid glands (conspicuous vs. inconspicuous) and tympanum (not visible vs. conspicuous). It differs from P. kavangensis in foot webbing (non-serrated vs. serrated), dorsal patterning (absent vertebral line vs. present), and ventral patterning (speckles vs. immaculate). It differs from P. parkeri in ventral patterning (speckles vs. immaculate). It differs from P. vertebralis in dorsal patterning (vertebral line absent vs. present), and conspicuousness of parotoid glands (conspicuous vs. inconspicuous).

Etymology: The specific epithet fernandae (Pt.) is a tribute to Fernanda Lages, a researcher and professor of Genetics based in Lubango, Angola. Her continuous investment in capacity building over the last decades and dedication to various research projects and international collaborations gave opportunities and transformed the professional paths of several young Angolan biologists, and thus of research in Biology in the country. The name, built in the feminine singular genitive, also pays homage to women in science. We suggest “Fernanda’s pygmy toad” and “sapo pigmeu da Fernanda” as English and Portuguese common names, respectively.


Pictures of live Poyntonophrynus nambensis sp. nov. A Female paratype (FKH-0458) with greyish coloration. B, C Female holotype (ZMB 91787) with dorsal and ventral coloration, respectively. D Pair in axillary amplexus [female (CHL0326), and male (BMNH 2021.7540) paratypes]. E Female paratype (BMNH 2021.7539) with brighter shades of brown and orange. F Eggs (CHL0472). Photos by W.R. Branch (B–F).

Poyntonophrynus nambensis sp. nov.

Diagnosis: Poyntonophrynus nambensis sp. nov. differs from P. pachnodes and P. fernandae sp. nov. sensu lato in having a columella. It differs from P. pachnodes in having a better-developed neopalatine. Differs from P. fernandae sp. nov. sensu lato in ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. scattered speckles at least on the chest), and dorsal coloration (dorsal pale hourglass-shaped blotch almost always present vs. absent), and breeding male coloration (dulls, similar to females vs. partially or completely bright yellow). It differs from P. beiranus in parotoid glands conspicuousness (conspicuous, elevated, with clearly demarcated margins vs. inconspicuous), and dorsal patterning (vertebral line absent vs. present). It differs from P. damaranus in ventral patterning (few dark speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. dombensis in tympanum size (when visible, between 0.5 and 0.6 times internarial distance, vs. conspicuous and around 0.7 times internarial distance), and ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. fenoulheti in parotoid glands width (thinner or around same width of eye diameter vs. wider than eye diameter). It differs from P. grandisonae in tympanum size (when visible, between 0.5 and 0.6 times internarial distance, vs. same width or wider than internarial distance) and skin texture (rough vs. leathery). It differs from P. grindleyi in pale occipital and sacral patches (present vs. absent), dorsal spines (small vs. large), and ventral coloration (few dark speckles along midline of chest, and dark line in front of arm insertion vs. dark thick ventral marbling). It differs from P. hoeschi in ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. jordani in shape of parotoid glands (kidney-shaped vs. a cluster of glands) and pale occipital patch (present vs. absent). It differs from P. lughensis in conspicuousness of parotoid glands (conspicuous vs. inconspicuous). It differs from P. kavangensis in foot webbing (non-serrated vs. serrated), dorsal patterning (absent vertebral line vs. present), and ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. immaculate). It differs from P. parkeri in development of parotoid glands (elevated vs. flattened). It differs from P. vertebralis in dorsal patterning (absent vertebral line vs. present), ventral patterning (few speckles along midline of chest, and dark line in front of arm insertion vs. distinct thick dark blotches), and conspicuousness of parotoid glands (conspicuous vs. inconspicuous).

Etymology: The specific epithet nambensis (Pt.) is a reference to the Namba mountains. This is the largest and more preserved relic of Afromontane forest and montane grasslands in Angola. However, it lacks official protection. We suggest “Namba pygmy toad” and “sapo pigmeu da Namba” as English and Portuguese common names respectively.
 
Ninda L. Baptista, Pedro Vaz Pinto, Chad Keates, Javier Lobón-Rovira, Shelley Edwards and Mark-Oliver Rödel. 2023. Two New Poyntonophrynus Species (Anura: Bufonidae) highlight the importance of Angolan Centers of Endemism. Vertebrate Zoology. 73: 991-1031. DOI: 10.3897/vz.73.e103935


Resumo: O sapo-pigmeu do género Poyntonophrynus é endémico da África Austral. A morfologia destes pequenos sapos é pouco variável. Estes têm uma coloração geralmente parda, e estão predominantemente adaptados a ambientes áridos. Durante pesquisas recentes em Angola, encontrámos Poyntonophrynus que não correspondiam a nenhuma espécie conhecida. Usando uma abordagem integrativa baseada em DNA mitocondrial e nuclear, morfologia, osteologia, biogeografia e ecologia, identificámos três novas linhagens, duas das quais foram descritas como espécies novas. As três linhagens estão geneticamente próximas de P. pachnodes, uma espécie endémica de Angola, mas estão geograficamente isoladas desta espécie. As novas espécies são morfologicamente distintas, e estão associadas a dois dos mais importantes centros de endemismo angolanos: a escarpa e as terras altas. Para contribuir para o conhecimento da osteologia do género, fizemos também uma caracterização osteológica de P. dombensis, inexistente até à data. Os nossos resultados i) aumentam o número de espécies sem aparelho auditivo no género Poyntonophrynus, ii) reforçam o sudoeste de África como o centro de diversificação deste género, iii) relatam a ocorrência de Poyntonophrynus em floresta húmida, revelando que estes sapos são ecologicamente mais variáveis do que se pensava, e iv) realçam a importância de estudos mais aprofundados nos centros de endemismo angolanos.

Palavras-chave:Amphibia, deslocamento de caracteres, dicromatismo sexual, escarpa de Angola, especiação, sapo pigmeu, sintopia, terras altas de Angola