Ancient forest fragmentation or recent radiation? Testing refugial speciation models in chameleons within an African biodiversity hotspot

Aim East Africa is one of the most biologically diverse regions, especially in terms of endemism and species richness. Hypotheses put forward to explain this high diversity invoke a role for forest refugia through: (1) accumulation of new species due to radiation within refugial habitats, or (2) retention of older palaeoendemic species in stable refugia. We tested these alternative hypotheses using data for a diverse genus of East African forest chameleons, Kinyongia. Location East Africa. Methods We constructed a dated phylogeny for Kinyongia using one nuclear and two mitochondrial markers. We identified areas of high phylogenetic diversity (PD) and evolutionary diversity (ED), and mapped ancestral areas to ascertain whether lineage diversification could best be explained by vicariance or dispersal. Results Vicariance best explains the present biogeographic patterns, with divergence between three major Kinyongia clades (Albertine Rift, southern Eastern Arc, northern Eastern Arc) in the early Miocene/Oligocene (> 20 Ma). Lineage diversification within these clades pre‐dates the Pliocene (> 6 Ma). These dates are much older than the Plio‐Pleistocene climatic shifts associated with cladogenesis in other East African taxa (e.g. birds), and instead point to a scenario whereby palaeoendemics are retained in refugia, rather than more recent radiations within refugia. Estimates of PD show that diversity was highest in the Uluguru, Nguru and East Usambara Mountains and several lineages (from Mount Kenya, South Pare and the Uluguru Mountains) stand out as being evolutionarily distinct as a result of isolation in forest refugia. PD was lower than expected by chance, suggesting that the phylogenetic signal is influenced by an unusually low number of extant lineages with long branch lengths, which is probably due to the retention of palaeoendemic lineages. Main conclusions The biogeographic patterns associated with Kinyongia are the result of long evolutionary histories in isolation. The phylogeny is dominated by ancient lineages whose origins date back to the early Miocene/Oligocene as a result of continental wide forest fragmentation and contraction due to long term climatic changes in Africa. The maintenance of palaeoendemic lineages in refugia has contributed substantially to the remarkably high biodiversity of East Africa.     

Rapid radiation of Impatiens (Balsaminaceae) during Pliocene and Pleistocene: Result of a global climate change

Impatiens comprises more than 1000 species and is one of the largest genera of flowering plants. The genus has a subcosmopolitan distribution, yet most of its evolutionary history is unknown. Diversification analyses, divergence time estimates and historical biogeography, illustrated that the extant species of Impatiens originated in Southwest China and started to diversify in the Early Miocene. Until the Early Pliocene, the net diversification rate within the genus was fairly slow. Since that time, however, approximately 80% of all Impatiens lineages have originated. This period of rapid diversification coincides with the global cooling of the Earth’s climate and subsequent glacial oscillations. Without this accelerated diversification rate, Impatiens would only have contained 1/5th of its current number of species, thereby indicating the rapid radiation of the genus.     

Palynological evidence for the impact of colonial settlement within lowland fynbos: A high‐resolution study from the Verlorenvlei,southwestern Cape Province,South Africa

The present distribution of lowland fynbos of the southwestern Cape coastal forelands occupies a unique yet threatened existence. Long renowned for its extraordinary species richness, the lowland fynbos now reveals a decline in diversity following the impact of recent European settlement. Fine resolution pollen analysis, conducted on a dated sediment core from the Verlorenvlei, reveals a 250 year pattern of debilitating vegetation transformations. The arrival of European colonists in the area circa 1700 is noted in the sedimentary sequence, not only with respect to changes in the pollen spectra, but also through concomitant changes in the nature and timing of sedimentation. The rapid extermination of large mammalian fauna, overgrazing by domestic stock and decades of agricultural mismanagement in a marginal landscape are considered to be the principal factors responsible for deliterious alterations of the natural disturbance regime.     

THE EVOLUTION OF REPRODUCTIVE EFFORT IN LIZARDS AND SNAKES

Life history theory suggests that the optimal evolved level of reproductive effort (RE) for an organism depends upon the degree to which additional current reproductive investment reduces future reproductive output. Future reproduction can be decreased in two ways, through (i) decreases in the organism's survival rate, and/or (ii) decreases in the organism's growth (and hence subsequent fecundity). The latter tradeoff–that is, the “potential fecundity cost”—should affect the evolution of RE only in species with relatively high survival rate, a relatively high rate of fecundity increase with body size, or a relatively high reproductive frequency per annum. Unless these conditions are met, the probable benefit in future fecundity obtained from decreasing present reproductive output is too low for natural selection to favor any reduction in RE below the maximum physiologically possible. Published data on survival rate, reproductive frequency and relative clutch mass (RCM) suggest that many lizard species fall well below the level at which natural selection can be expected to influence RE through such “potential fecundity” tradeoffs. Hence, the relative allocation of resources between growth and reproduction is unlikely to be directly optimized by natural selection in these animals. Instead, energy allocation should influence the evolution of RE only indirectly, via effects on an organism's probability of survival during reproduction. Survival costs of reproduction may be the most important evolutionary determinants of RE in many reptiles, and information on the nature and extent of such costs is needed before valid measures of reptilian RE can be constructed.     

Causes of perceived faunal change in the later Neogene of East Africa

African terrestrial vertebrate faunas change over the time range from 14 my to 4 my, the period during which hominids are presumed to have diverged from other hominoids. The most complete local sequence showing this is provided by the Tugen Hills, west of Lake Baringo in Kenya. A large component of the pattern of faunal changes seen there, and at other relevant sites, could be explained by local environmental alterations, or by sampling properties of fossil assemblages and collections. It is hard to justify global climatic change as the sole explanation of observed faunal shifts, although such external factors cannot be ruled out and on general grounds are likely to be involved. As yet there is no convincing demonstration of synchronicity in faunal turnover. Evidence of a simple shift from forest conditions to grasslands, an event that has been correlated with the origins of hominid bipedalism, is hard to detect. Existing data suggest a more patchy distribution of forest and grassland in both space and time.     

A partial systematic revision of the day geckos, Phelsuma Gray, of Madagascar (Reptilia: Squamata: Gekkonidae)

Recent survey work in Madagascar has led to significant changes in the systematics of Madagascan Phelsuma. A new species from the Masoala Peninsula in northeastern Madagascar is described, which has a nostril position typical of Phelsuma from Mauritius, Reunion and Rodriguez islands. Phelsuma breviceps was rediscovered in extreme southern Madagascar and is resurrected from the synonymy of P. mutabilis. Phelsuma breviceps has fragile skin, a probable predator escape mechanism, similar to the condition found in geckos of the genera Ailuronyx and Geckokpis. Three new synonyms are recognized. Phelsuma befotahensis and P. checki are junior synonyms of P. abbotti , and P. minuthi is a junior synonym of P. lineata. Twenty species of Phelsuma occur in Madagascar, of which 16 are endemic. A checklist and identification key are provided. Conservation problems of Phelsuma in the coastal regions of Madagascar are briefly discussed.