Diversity and Evolution of Frog Visual Opsins: Spectral Tuning and Adaptation to Distinct Light Environments.

Visual systems adapt to different light environments through several avenues including optical changes to the eye and neurological changes in how light signals are processed and interpreted. Spectral sensitivity can evolve via changes to visual pigments housed in the retinal photoreceptors through gene duplication and loss, differential and coexpression, and sequence evolution. Frogs provide an excellent, yet understudied, system for visual evolution research due to their diversity of ecologies (including biphasic aquatic-terrestrial life cycles) that we hypothesize imposed different selective pressures leading to adaptive evolution of the visual system, notably the opsins that encode the protein component of the visual pigments responsible for the first step in visual perception. Here, we analyze the diversity and evolution of visual opsin genes from 93 new eye transcriptomes plus published data for a combined dataset spanning 122 frog species and 34 families. We find that most species express the four visual opsins previously identified in frogs but show evidence for gene loss in two lineages. Further, we present evidence of positive selection in three opsins and shifts in selective pressures associated with differences in habitat and life history, but not activity pattern. We identify substantial novel variation in the visual opsins and, using microspectrophotometry, find highly variable spectral sensitivities, expanding known ranges for all frog visual pigments. Mutations at spectral-tuning sites only partially account for this variation, suggesting that frogs have used tuning pathways that are unique among vertebrates. These results support the hypothesis of adaptive evolution in photoreceptor physiology across the frog tree of life in response to varying environmental and ecological factors and further our growing understanding of vertebrate visual evolution.

A biogeographical appraisal of the threatened South East Africa Montane Archipelago ecoregion.

Recent biological surveys of ancient inselbergs in southern Malawi and northern Mozambique have led to the discovery and description of many species new to science, and overlapping centres of endemism across multiple taxa. Combining these endemic taxa with data on geology and climate, we propose the 'South East Africa Montane Archipelago' (SEAMA) as a distinct ecoregion of global biological importance. The ecoregion encompasses 30 granitic inselbergs reaching > 1000 m above sea level, hosting the largest (Mt Mabu) and smallest (Mt Lico) mid-elevation rainforests in southern Africa, as well as biologically unique montane grasslands. Endemic taxa include 127 plants, 45 vertebrates (amphibians, reptiles, birds, mammals) and 45 invertebrate species (butterflies, freshwater crabs), and two endemic genera of plants and reptiles. Existing dated phylogenies of endemic animal lineages suggests this endemism arose from divergence events coinciding with repeated isolation of these mountains from the pan-African forests, together with the mountains' great age and relative climatic stability. Since 2000, the SEAMA has lost 18% of its primary humid forest cover (up to 43% in some sites)-one of the highest deforestation rates in Africa. Urgently rectifying this situation, while addressing the resource needs of local communities, is a global priority for biodiversity conservation.

A new species of tree snake (Dipsadoboa, Serpentes: Colubridae) from 'sky island' forests in northern Mozambique, with notes on other members of the Dipsadoboa werneri group.

A new species of tree snake Dipsadoboa montisilva Branch, Conradie Tolley sp. nov. (Serpentes: Colubridae) is described from the 'sky islands' of Mount Mabu and Mount Ribáuè in northern Mozambique. Features of scalation, colour, body form and habitat distinguish the new species from other Dipsadoboa. This is supported by a phylogenetic analysis using one mitochondrial marker (cytochrome b) that shows the new Mozambican species is divergent from other sampled Dipsadoboa, including D. flavida and D. aulica, the only congeners known to occur in Mozambique. Morphologically, the new Dipsadoboa forms part of the D. werneri-shrevei complex from east and southeast Africa, but differs in having higher subcaudal counts, a different temporal pattern and only two supralabials entering the orbit. Phylogenetically, it occurs in a clade with D. shrevei and D. werneri. The status of D. shrevei in East Africa is reassessed, particularly in terms of the poorly-known Dipsadoboa shrevei kageleri from northern Tanzania. It is morphologically well defined from D. shrevei shrevei and utilises a different habitat. Although based on limited genetic data, it appears to be well-defined from typical D. shrevei and is accordingly raised to specific status. The only Tanzanian record for typical D. shrevei from Mtene, Rondo Plateau in southeast Tanzania is well isolated from the species' range to the west (e.g. Zambia, Angola) and the published scalation features, particularly ventral counts, do not fully accord with D. shrevei. The Rondo Plateau population is treated as Dipsadoboa incerta sedis, and because we return D. shrevei to its binomial status, we can no longer consider D. shrevei as occurring in Tanzania. Biogeographically, the Rondo Plateau population may have a stronger affinity to the new Mozambican species. The discovery of isolated populations of the new species in mid-altitude forest remnants on Mt Mabu and Mt Ribáuè emphasizes the high conservation importance of the Mozambique forest 'sky islands' from which numerous other endemic new species have been recently discovered. These species are impacted by ongoing habitat destruction through slash and burn clearing for subsistence agriculture.

Impact of species delimitation and sampling on niche models and phylogeographical inference: A case study of the East African reed frog Hyperolius substriatus Ahl, 1931.

Ecological niche models (ENMs) have been used in a wide range of ecological and evolutionary studies. In biogeographic studies these models have, among other things, helped in the discovery of new allopatric populations, and even new species. However, small sample sizes and questionable taxonomic delimitation can challenge models, often decreasing their accuracy. Herein we examine the sensitivity of ENMs to the addition of new, geographically isolated populations, and the impact of applying different taxonomic delimitations. The East African reed frog Hyperolius substriatus Ahl, 1931 was selected as a case study because it has been the subject of previous ENM predictions. Our results suggest that addition of new data and reanalysis of species lineages of H. substriatus improved our understanding of the evolutionary history of this group of frogs. ENMs provided robust predictions, even when some populations were deliberately excluded from the models. Splitting the lineages based on genetic relationships and analysing the ENMs separately provided insights about the biogeographical processes that led to the current distribution of H. substriatus.

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.

Effects of fragmentation and sea-level changes upon frog communities of land-bridge islands off the southeastern coast of Brazil.

We investigate the composition of anuran communities of land-bridge islands off the southeastern coast of Brazil. These islands provide natural long-term experiments on the effects of fragmentation in the Brazilian Atlantic Forest (BAF). We hypothesize that Pleistocene sea-level changes, in combination with other abiotic variables such as area and habitat diversity, has affected anuran species richness and community composition on these islands. Data from the literature and collections databases were used to produce species lists for eight land-bridge islands and for the mainland adjacent to the islands. We assess the effects of area, number of breeding habitats and distance to the mainland upon anuran species richness on land-bridge islands. Additionally we use nestedness analysis to quantify the extent to which the species on smaller and less habitat-diverse islands correspond to subsets of those on larger and more diverse ones. We found that area has both direct and indirect effects on anuran species richness on land-bridge islands, irrespective of distance to the mainland. However, on islands with comparable sizes, differences in species richness can be attributed to the number and quality of breeding habitats. Anuran communities on these islands display a nested pattern, possibly caused by selective extinction related to habitat loss. Common lowland pond-breeders were conspicuous by their absence. In the BAF, the conservation of fragments with a high diversity of breeding habitats could compensate for the generally negative effect of small area upon species richness. We suggest that sea-level changes have an important role in shaping composition of anuran species on coastal communities.