Out of southern Africa: Origins and cryptic speciation in Chamaeleo, the most widespread chameleon genus.

Molecular phylogenetics and the application of species delimitation methods have proven useful in addressing limitations associated with morphology based taxonomy and have highlighted the inconsistencies in the current taxonomy for many groups. For example, the genus Chamaeleo, which comprises 14 species with large distributions across mainland Africa and parts of Eurasia, exhibits relatively minor phenotypic differentiation between species, leading to speculation regarding the presence of cryptic diversity in the genus. Therefore, the aims of the present study were to construct a robust and comprehensive phylogeny of the genus and highlight potential species-level cryptic diversity. Additionally, we sought to ascertain the most likely biogeographic origin of the genus and understand its spatio-temporal diversification. Accordingly, we made use of species delimitation methods (Bayesian and divergence based) to investigate the extent of cryptic diversity in Chamaeleo, and applied an ancestral area reconstruction to examine the biogeographic origin of the group. Our phylogenetic analyses suggested the presence of at least 18 taxa within Chamaeleo. Notably, three taxa could be recognised within C. dilepis, none of which are equivalent in context with any of the currently described subspecies. There were also three taxa within C. gracilis and two within C. anchietae. The single available tissue specimen identified as C. necasi was embedded within the C. gracilis clade. Our ancestral area reconstruction points to a southern African/Zambezian origin for Chamaeleo, with diversification beginning during the cooling and aridification of Africa that characterised the Oligocene Epoch, ca. 34-23 Mya (Million years ago). Species-level diversification began in the Miocene Epoch (ca. 23-5 Mya), possibly tracking the aridification that triggered the shift from forest to more open, mesic savanna for most clades, but with tectonic events influencing speciation in a Palearctic clade. These findings lay the foundation for a future integrative taxonomic re-evaluation of Chamaeleo, which will be supported with additional lines of evidence before implementing any taxonomic changes.

Envenomation by the spotted harlequin snake, Homoroselaps lacteus (Linnaeus) 1754 (Serpentes: Atractaspidinae).

Two cases of envenomation from the small, semi-fossorial, front-fanged colubrid Homoroselaps lacteus are presented and the recent literature reviewed. In addition to local oedema and lymphadenopathy, the development of significant subcutaneous loco-regional bruising suggest that damage to capillary basement membrane and possibly other underlying anti-haemostatic effects may pertain to the clinical syndrome. Although local clinical effects may appear dramatic, due to the small volumes of venom that can be injected, it is considered that envenomation is unlikely to cause life-threatening effects.

Cryptic diversity in Rhampholeon boulengeri (Sauria: Chamaeleonidae), a pygmy chameleon from the Albertine Rift biodiversity hotspot.

Several biogeographic barriers in the Central African highlands have reduced gene flow among populations of many terrestrial species in predictable ways. Yet, a comprehensive understanding of mechanisms underlying species divergence in the Afrotropics can be obscured by unrecognized levels of cryptic diversity, particularly in widespread species. We implemented a multilocus phylogeographic approach to examine diversity within the widely distributed Central African pygmy chameleon, Rhampholeon boulengeri. Gene-tree analyses coupled with a comparative coalescent-based species delimitation framework revealed R. boulengeri as a complex of at least six genetically distinct species. The spatiotemporal speciation patterns for these cryptic species conform to general biogeographic hypotheses supporting vicariance as the main factor behind patterns of divergence in the Albertine Rift, a biodiversity hotspot in Central Africa. However, we found that parapatric species and sister species inhabited adjacent habitats, but were found in largely non-overlapping elevational ranges in the Albertine Rift, suggesting that differentiation in elevation was also an important mode of divergence. The phylogeographic patterns recovered for the genus-level phylogeny provide additional evidence for speciation by isolation in forest refugia, and dating estimates indicated that the Miocene was a significant period for this diversification. Our results highlight the importance of investigating cryptic diversity in widespread species to improve understanding of diversification patterns in environmentally diverse regions such as the montane Afrotropics.

A fatal bite from the burrowing asp Atractaspis corpulenta (Hallowell 1854).

Bites from the various species of Atractaspis are a common occurrence in Africa but deaths are very unusual. Of the 19 described species, the clinical effects of the bite of only seven have been described, and in only three (Atractaspis irregularis, Atractaspis microlepidota and Atractaspis engaddensis) have fatalities been documented. A case of envenomation is described following a bite to a finger by Atractaspis corpulenta, which resulted in sudden death approximately two and a half hours later. The victim received antivenom and although anaphylaxis to this cannot be ruled out, we consider it to be unlikely to be the cause of death. A late autopsy was performed and the findings and their interpretation are discussed. The previous case fatalities, toxic fractions and clinical effects of Atractaspis venom are briefly reviewed.

Contributions to the herpetofauna of the Albertine Rift: Two new species of chameleon (Sauria: Chamaeleonidae) from an isolated montane forest, south eastern Democratic Republic of Congo.

Two new species of chameleons from the genera Rhampholeon and Kinyongia are described from an isolated montane forest remnant situated toward the southern end of the Albertine Rift bordering Lake Tanganyika. The closest known localities of species from these genera are 200km and 400km to the north respectively, separated by large intervening tracts of lowland savannah and Brachystegia (Miombo) woodland - habitats not normally inhabited by species of these genera. Rhampholeon hattinghi sp. nov. and Kinyongia mulyai sp. nov. bear superficial resemblances to previously described species (Rh. boulengeri Steindachner and K. adolfifriderici  (Sternfeld)). Rhampholeon hattinghi sp. nov. has a relatively smooth supra-orbital ridge, deep axillary but absent inguinal mite pockets, prominent white spots on the base of the tail and a uniquely derived hemipenal morphology with billowing parasulcal evaginations. Like K. adolfifriderici, Kinyongia mulyai sp. nov. is devoid of a rostral appendage but differs in having a longer and narrower head, a higher upper labial scale count and by the absence of a dorsal crest in the male. To place these new chameleons within the context of their respective genera, Bayesian and maximum likelihood phylogenetic analyses were carried out utilising two mitochondrial (ND2 and 16S) and one nuclear marker (RAG1).  Both chameleons were found to have morphological features that distinguish them from other congeners. Based on phylogenetic analysis they are clearly separate evolutionary lineages and are described as new species. 

Evolutionary relationships, species delimitation and biogeography of Eastern Afromontane horned chameleons (Chamaeleonidae: Trioceros).

The Eastern Afromontane Region (EAR) contains numerous endemic species, yet its reptile diversity remains relatively poorly understood. We used molecular data to examine species diversity of the Sub-Saharan chameleon genus Trioceros. In particular, we focus on establishing species boundaries for taxa with disjunct distributions across the fragmented mountains of the EAR, including T. affinis, T. balebicornutus, T. deremensis, T. harennae, T. tempeli and T. werneri. We applied three species-delimiting approaches, General Mixed Yule-Coalescent (GMYC), a Bayesian implementation of the GMYC, and Bayes Factor Delimitation to estimate species diversity. Using a dated phylogeny, we also examined spatial and temporal diversification patterns in Trioceros. We found strong congruence between different species delimitation approaches, with all methods suggesting that species diversity is currently underestimated. In particular, T. werneri consists of at least four candidate species (i.e. species awaiting description) with some mountain ranges (Uluguru and Udzungwa) having potentially more than one species. Most interspecific divergences between extant Trioceros lineages are estimated to be >5Mya, consistent with a Pliocene origin of the endemic montane fauna, as exhibited in other taxonomic groups. Multiple, overlapping geographic events (climate and/or geomorphological changes) might account for speciation patterns in Trioceros given the dating results.

A phylogenetic review of the African leaf chameleons: genus Rhampholeon (Chamaeleonidae): the role of vicariance and climate change in speciation.

The phylogenetic associations among 13 currently recognized African leaf chameleon species were investigated by making use of mitochondrial and nuclear DNA sequence data (44 taxa and 4145 characters). The gene tree indicates two divergent clades within Rhampholeon; this finding is congruent with previous morphological suggestions. The first clade (I) comprises three taxa (R. kerstenii, R. brevicaudatus and R. brachyurus) and is widely distributed in lowland forest and or non-forest biomes. The second clade (II) comprises the remaining Rhampholeon species and can be subdivided into three subclades. By contrast, most taxa belonging to clade II are confined to relict montane forest biotopes. Based on geographical, morphological and molecular evidence, it is suggested that the taxonomy of Rhampholeon be revised to include two genera (Rieppeleon and Rhampholeon) and three subgenera (Rhampholeon, Bicuspis and Rhinodigitum). There is a close correlation between geographical distribution and phylogenetic relatedness among Rhampholeon taxa, indicating that vicariance and climate change were possibly the most influential factors driving speciation in the group. A relaxed Bayesian clock suggests that speciation times coincided both with the northern movement of Africa, which caused the constriction of the pan African forest, and to rifting in east Africa ca. 20 Myr ago. Subsequent speciation among taxa was probably the result of gradual desiccation of forests between 20 and 5 Myr ago.

Phylogenetics of the southern African dwarf chameleons, Bradypodion (Squamata: Chamaeleonidae).

The taxonomic relationships within the dwarf chameleons (Bradypodion) of southern Africa have long been controversial. Although informal phenotypic groups have been suggested, the evolutionary relationships among the 15 recognised species in southern Africa have not been previously investigated. To investigate the relationships among species within this genus, fragments of two mitochondrial genes (16S ribosomal RNA and ND2) were sequenced and analysed using maximum parsimony, maximum likelihood and Bayesian inference. All analyses showed congruent topologies, revealing at least 5 well-supported clades distributed across distinct geographic regions. The mtDNA gene tree indicated that in many instances, geographic location has played a role in shaping the evolution of this group, and that the previously suggested phenotypic groupings do not adequately reflect evolutionary relationships. Furthermore, it appears that some of the currently recognised species (described on morphology) are polyphyletic for mitochondrial sequences, most notably those occurring in the isolated forest patches of north-eastern South Africa, near the Drakensberg Escarpment.