Werner Conradie

Revised phylogeny of African sand lizards (Pedioplanis), with the description of two new species from south-western Angola

Abstract Although reptile diversity in Africa is high, it is poorly represented in Angola, with just 257 species known. Despite its greater surface area and habitat diversity Angola has significantly lower lacertid lizard diversity than adjacent Namibia. This is particularly notable in African sand lizards (Pedioplanis), where 10 species (two endemic) are known from Namibia but only two are recorded from adjacent Angola. Pedioplanis benguellensis was described from Angola, but its taxonomic status is problematic and it was previously synonymised with P. namaquensis. All other Angolan Pedioplanis were referred to Namibian P. undata, although this taxon is now known to comprise a complex of at least five different species and the relationship of Angolan material to this complex has not been assessed. In this study, we investigated the phylogenetic placement of Angolan Pedioplanis using two mitochondrial (ND2 and 16S) and one nuclear (RAG-1) markers. A Bayesian analysis was conducted on 21 samples from Angola, combined with existing data for 45 individuals from GenBank and three additional samples from central Namibia. The phylogeny demonstrates that P. benguellensis is a valid species and that it is not the sister taxon to P. namaquensis with which it has been morphologically confused. In addition, Angolan lacertids previously referred to P. undata are not conspecific with any of the Namibian or South African species in that complex. Rather, there is strong support for the presence in Angola of additional species of Pedioplanis, which form a well-supported sister clade to the P. undata complex (sensu stricto) of Namibia and two of which are described herein. These discoveries highlight the need for further biodiversity surveys in Angola, as similar increases in species diversity in other Angolan taxa might be found given sufficient investment in biodiversity surveys.

Molecular phylogeny of Panaspis and Afroablepharus skinks (Squamata: Scincidae) in the savannas of sub-Saharan Africa

African snake-eyed skinks are relatively small lizards of the genera Panaspis and Afroablepharus. Species allocation of these genera frequently changed during the 20th century based on morphology, ecology, and biogeography. Members of these genera occur primarily in savanna habitats throughout sub-Saharan Africa and include species whose highly conserved morphology poses challenges for taxonomic studies. We sequenced two mitochondrial (16S and cyt b) and two nuclear genes (PDC and RAG1) from 76 Panaspis and Afroablepharus samples from across eastern, central, and southern Africa. Concatenated gene-tree and divergence-dating analyses were conducted to infer phylogenies and biogeographic patterns. Molecular data sets revealed several cryptic lineages, with most radiations occurring during the mid-Miocene to Pliocene. We infer that rifting processes (including the formation of the East African Rift System) and climatic oscillations contributed to the expansion and contraction of savannas, and caused cladogenesis in snake-eyed skinks. Species in Panaspis and Afroablepharus used in this study, including type species for both genera, formed a monophyletic group. As a result, the latter genus should be synonymized with the former, which has priority. Conservatively, we continue to include the West African species P. breviceps and P. togoensis within an expanded Panaspis, but note that they occur in relatively divergent clades, and their taxonomic status may change with improved taxon sampling. Divergence estimates and cryptic speciation patterns of snake-eyed skinks were consistent with previous studies of other savanna vertebrate lineages from the same areas examined in this study.

The phylogenetic position and diversity of the enigmatic mongrel frog Nothophryne Poynton, 1963 (Amphibia, Anura)

The phylogenetic relationships of the African mongrel frog genus Nothophryne are poorly understood. We provide the first molecular assessment of the phylogenetic position of, and diversity within, this monotypic genus from across its range-the Afromontane regions of Malawi and Mozambique. Our analysis using a two-tiered phylogenetic approach allowed us to place the genus in Pyxicephalidae. Within the family, Nothophryne grouped with Tomopterna, a hypothesis judged significantly better than alternative hypotheses proposed based on morphology. Our analyses of populations across the range of Nothophryne suggest the presence of several cryptic species, at least one species per mountain. Formal recognition of these species is pending but there is a major conservation concern for these narrowly distributed populations in an area impacted by major habitat change. The phylogenetic tree of pyxicephalids is used to examine evolution of life history, ancestral habitat, and biogeography of this group.

The King of the Dwarves: a new cryptic species of Dainty Frog (Anura: Pyxicephalidae: Cacosternum ) from the eastern Great Escarpment of South Africa

Phylogenetic reconstruction using the mitochondrial 16S marker shows the presence of a cryptic species of Cacosternum (Anura: Pyxicephalidae) from the eastern Great Escarpment of South Africa, supporting the Greater Maputaland-Pondoland-Albany region of vertebrate endemism. Bioacoustic and morphological characteristics, in conjunction with colouration differences, allow the description of this cryptic species. Tadpoles and details of life history are described.

A single origin of extreme matrotrophy in African mabuyine skinks.

Most mammals and approximately 20% of squamates (lizards and snakes) are viviparous, whereas all crocodilians, birds and turtles are oviparous. Viviparity evolved greater than 100 times in squamates, including multiple times in Mabuyinae (Reptilia: Scincidae), making this group ideal for studying the evolution of nutritional patterns associated with viviparity. Previous studies suggest that extreme matrotrophy, the support of virtually all of embryonic development by maternal nutrients, evolved as many as three times in Mabuyinae: in Neotropical Mabuyinae (63 species), Eumecia (2 species; Africa) and Trachylepis ivensii (Africa). However, no explicit phylogenetic hypotheses exist for understanding the evolution of extreme matrotrophy. Using multilocus DNA data, we inferred a species tree for Mabuyinae that implies that T. ivensii (here assigned to the resurrected genus Lubuya) is sister to Eumecia, suggesting that extreme matrotrophy evolved only once in African mabuyine skinks.

Designation and description of a neotype of Sclerophrys maculata (Hallowell, 1854), and reinstatement of S. pusilla (Mertens, 1937) (Amphibia: Anura: Bufonidae)

Molecular analysis indicates that African material previously referred to Amietophrynus maculatus (Hallowell, 1854; now Sclerophrys maculata), is divisible into two distinct clades: a Western Clade from Cameroon westwards and an Eastern Clade from Central African Republic eastwards, and Uganda southwards to South Africa, extending to Angola-Namibia. Preliminary morphological and bioacoustic data support this division. The two clades are recognised here as two separate species. The Western species retains the name S. maculata, with Hallowells designated type locality of Liberia. The Eastern Clade retains the name published by Mertens (1937), S. pusilla. It is noted that a type specimen of S. maculata cannot be traced and is presumed lost; the so-called syntypes in the Philadelphia Academy of Natural Sciences are not the material described by Hallowell. None of these have been designated as a neotype, consequently a specimen from Liberia in the collection of the Natural History Museum, London, is designated here as the neotype of S. maculata.

Unmasking evolutionary diversity among two closely related South African legless skink species (Acontinae: Acontias) using molecular data

We examined species boundaries among two phylogenetically closely related and morphologically similar South African fossorial legless skink species, Acontias breviceps and Acontias gracilicauda. Samples of these two species were collected throughout their distribution ranges and sequenced for three DNA loci (two mitochondrial loci, 16S rRNA and cytochrome b (Cyt b), plus the nuclear locus prolactin). Phylogenetic relationships were determined using maximum parsimony, Bayesian inference and maximum likelihood analyses of the combined DNA sequence data set. The total evidence topology retrieved two paraphyletic clades in both Acontias species with strong statistical support. The phylogenetic results revealed that A. breviceps specimens from the Eastern Cape Province were basal (Clade 1), while the Highveld specimens of A. breviceps from the Mpumalanga Province (Clade 2) were retrieved as sister to A. gracilicauda (Clade 1). In addition, the A. gracilicauda specimens from the interior of the Northern Cape Province (Clade 2) were found embedded within the A. occidentalis species complex. These clades were characterised by marked sequence divergence for the Cyt b locus. Furthermore, no maternal or nuclear haplotypes were shared between clades within both A. breviceps and A. gracilicauda, alluding to genetic and reproductive isolation. The results provide overwhelming evidence to assign A. breviceps from the Mpumalanga Highveld to a novel species. Further sampling is required to accurately delineate species boundaries within A. gracilicauda. The conservation implications of our results are briefly discussed.

Sexual Dichromatism Drives Diversification Within a Major Radiation of African Amphibians

Theory predicts that sexually dimorphic traits under strong sexual selection, particularly those involved with intersexual signaling, can accelerate speciation and produce bursts of diversification. Sexual dichromatism (sexual dimorphism in color) is widely used as a proxy for sexual selection and is associated with rapid diversification in several animal groups, yet studies using phylogenetic comparative methods to explicitly test for an association between sexual dichromatism and diversification have produced conflicting results. Sexual dichromatism is rare in frogs, but it is both striking and prevalent in African reed frogs, a major component of the diverse frog radiation termed Afrobatrachia. In contrast to most other vertebrates, reed frogs display female-biased dichromatism in which females undergo color transformation, often resulting in more ornate coloration in females than in males. We produce a robust phylogeny of Afrobrachia to investigate the evolutionary origins of sexual dichromatism in this radiation and examine whether the presence of dichromatism is associated with increased rates of net diversification. We find that sexual dichromatism evolved once within hyperoliids and was followed by numerous independent reversals to monochromatism. We detect significant diversification rate heterogeneity in Afrobatrachia and find that sexually dichromatic lineages have double the average net diversification rate of monochromatic lineages. By conducting trait simulations on our empirical phylogeny, we demonstrate our inference of trait-dependent diversification is robust. Although sexual dichromatism in hyperoliid frogs is linked to their rapid diversification and supports macroevolutionary predictions of speciation by sexual selection, the function of dichromatism in reed frogs remains unclear. We propose that reed frogs are a compelling system for studying the roles of natural and sexual selection on the evolution of sexual dichromatism across both micro- and macroevolutionary timescales.

Two new species of Acontias (Acontinae, Scincidae) from the Mpumalanga Highveld escarpment of South Africa

The African genus of fossorial legless lizards (Acontias Cuvier) currently comprises 26 species and subspecies. In a recent study on the two disjunct populations of Acontias breviceps Essex, the presence of cryptic species was discovered. Here, we increase the sampling size and describe these disjunct populations from the Mpumalanga Escarpment of South Africa as new species. The new species differ from congeners based on a combination of factors, including the number of midbody, ventral, and subcaudal scale counts, ventral pigmentation, allopatric distributions, and genetic divergences. The new species are genetically distant from nominal A. breviceps, with which it shares overall pigmentation and scalation. The new description adds to the growing number of Mpumalanga escarpment endemic reptiles, and highlights the area as a biodiversity hotspot. The use of vertebral counts as a distinguishing character between species is briefly discussed.

A new Reed Frog (Hyperoliidae: Hyperolius ) from coastal northeastern Mozambique

A new species of African reed frog (genus Hyperolius Rapp, 1842) is described from the Coastal Forests of the Eastern Africa Biodiversity Hotspot in northeastern Mozambique. It is currently only known from less than ten localities associated with the Mozambican coastal pans system, but may also occur in the southeastern corner of Tanzania. Phylogenetic reconstructions using the mitochondrial 16S marker revealed that it is the sister taxon of Hyperolius mitchelli (>5.6% 16S mtDNA sequence divergence) and forms part of a larger H. mitchelli complex with H. mitchelli and H. rubrovermiculatus. The new species is distinguished from other closely related Hyperolius species by genetic divergence, morphology, vocalisation, and dorsal colouration.