Resilient forest faunal communities in South Africa: a legacy of palaeoclimatic change and extinction filtering?

Aim To examine the influence of climatic extinction filtering during the last glacial maximum (LGM; c. 18,000 yr bp ) and of the subsequent recolonization of forest faunas on contemporary assemblage composition in southern African forests. Location South Africa, Mozambique, Swaziland, Zimbabwe. Methods Data comprised presence/absence by quarter‐degree grid cell for forest‐dependent and forest‐associated birds, non‐volant mammals and frogs. Twenty‐one forest subregions were assigned to one of three previously identified forest types: Afrotemperate, scarp, and Indian Ocean coastal belt. Differences among forest types were examined through patterns and gradients of species richness and endemism, assemblage similarity, species turnover, and coefficients of species dispersal direction. The influence of contemporary environment on assemblage composition was investigated using partial canonical correspondence analysis. Several alternative biogeographical hypotheses for the recolonization of forest faunas were tested. Results Afrotemperate faunas are relatively species‐poor, have low species turnover, and are unsaturated and infiltrated by generalist species. In northern and central regions, communities are supplemented by recolonization from scarp forest refugia, and among frogs by autochthanous speciation in localized refugia. Scarp faunas are relatively species‐rich, contain many forest‐dependent species, have high species turnover, and overlap with coastal and Afrotemperate faunas. Coastal forests are relatively species‐rich with high species turnover. Main conclusions Afrotemperate communities were affected most by climatic extinction filtering events. Scarp forests were Afrotemperate refugia during the LGM and are a contemporary overlap zone between Afrotemperate and coastal forest. Coastal faunas derive from post‐LGM colonization along the eastern seaboard from tropical East African refugia. The greatest diversity is achieved in scarp and coastal forest faunas in northern KwaZulu–Natal province. This historical centre of diversity has influenced the faunal diversity of nearly all other forests in South Africa. The response of vertebrate taxa to large‐scale, historical processes is dependent on their relative mobility: forest birds best illustrate patterns resulting from post‐glacial faunal dispersal, while among mammals and frogs the legacy of climatic extinction filtering remains stronger.     

Angulate and leopard tortoises in the Thicket Biome, Eastern Cape, South Africa: populations and biomass estimates

The population parameters (morphometrics, population size and biomass) were determined for leopard and angulate tortoises in the Thicket Biome, Eastern Cape, South Africa. Leopard tortoises had a mean total length of 306.47mm (SD = 94.65mm, range 59–496mm, n = 89). Males were smaller (mean total length 277.31mm, SD = 71.85mm, range 73–377mm, n = 36) than females (mean total length = 332.33mm, SD = 97.64mm, range 59–496mm, n = 53). Angulate tortoises had a mean total length of 155.17mm (SD = 33.89mm, range 59–194mm, n = 18). Males were larger (mean total length = 186mm, SD = 8.40mm, range 174–194mm, n = 5) than females (mean total length = 143.31mm, SD = 32.48mm, range 59–178mm, n = 13). The estimated population density for leopard tortoises was 0.85 tortoises ha⁻¹ with an estimated biomass of 6.02kg ⁻¹. Angulate tortoise population density was estimated at 0.12 tortoises ha⁻¹ with a mean biomass of 0.06kg ha⁻¹. The possible consequences of this high density and biomass are discussed.     

Comparative molecular phylogeography of two Xenopus species, X. gilli and X. laevis, in the south-western Cape Province, South Africa

Xenopus gilli is a vulnerable anuran with a patchy distribution along the south-western coast of the Cape Province, South Africa. This species is sympatric with Xenopus laevis laevis , a widespread relative found over much of southern Africa. We examined the molecular phylogeography and population structure of the contact zone between these species to obtain information about historical biogeography and conservation management of this region. Analyses of the distribution, frequency, and cladistic and phenetic relationships among mitochondrial DNA haplotypes indicate that population subdivision is present in both taxa but that long-term isolation of sets of populations has occurred in X. gilli only. Haplotype and nucleotide diversity are also considerably higher within and among X. gilli ponds than X. l. laevis ponds in this region. We attribute the genetic segregation of X. gilli populations to ancient habitat fragmentation by ocean transgression into X. gilli habitat and to continued habitat alteration by human activity. The lower level of genetic diversity in X. l. laevis in this region is likely a result of a recent arrival of this taxon to the south-western Cape region relative to X. gilli . Population structure in X. l. laevis may be a result of isolation by distance. Clear evidence exists for at least two management units within X. gilli and strongly supports the establishment of protective measures east of False Bay in order to conserve a substantial portion of this species' extant genetic diversity.     

Phylogeography of the Cape velvet worm (Onychophora: Peripatopsis capensis) reveals the impact of Pliocene/Pleistocene climatic oscillations on Afromontane forest in the Western Cape, South Africa

Habitat specialists such as soft-bodied invertebrates characterized by low dispersal capability and sensitivity to dehydration can be employed to examine biome histories. In this study, the Cape velvet worm (Peripatopsis capensis) was used to examine the impacts of climatic oscillations on historical Afromontane forest in the Western Cape, South Africa. Divergence time estimates suggest that the P. capensis species complex diverged during the Pliocene epoch. This period was characterized by dramatic climatic and topographical change. Subsequently, forest expansion and contraction cycles led to diversification within P. capensis. Increased levels of genetic differentiation were observed along a west-to-south-easterly trajectory because the south-eastern parts of the Cape Fold Mountain chain harbour larger, more stable fragments of forest patches, have more pronounced habitat heterogeneity and have historically received higher levels of rainfall. These results suggest the presence of three putative species within P. capensis, which are geographically discreet and genetically distinct.     

Distribution and population dynamics of Porphyra (Bangiales, Rhodophyta) in the southern Western Cape, South Africa

Although Porphyra is commercially farmed in many countries, in South Africa only small harvests of wild populations for sale as nori have been carried out. The discovery that Porphyra improves growth of South African abalone (Haliotis midae) farmed inland-based tanks has led to increased pressure to harvest wild populations. This paper reports on a survey of the distribution and seasonality of Porphyra in the southern Western Cape. Porphyrawas present at all sites surveyed, and showed considerable temporal variation. A significant amount of the Porphyra present is in reserves and therefore protected from harvesting. Close rexamination of one site revealed seasonal populations of Porphyra that occupied different niches dependent on season. Recruitment peaked in spring and autumn, leading to dense summer and winter populations. Summer populations generally grew lower in the eulittoral than winter populations. No pattern in the mortality of larger thalli wasde tected, though sporeling mortality was high following recruitment peaks. Although it seems that most sites in the southern Western Cape are suitable for harvesting, the taxonomy of the genus in the region urgently needs revision if populations are to be appropriately managed. This revised version was published online in August 2006 with corrections to the Cover Date.     

A new species of Arctotis (Compositae, Arctotideae) from kommetjie grassland in Eastern Cape Province, South Africa

Arctotis debensis R.J.McKenzie, a new species from the Albany Centre of Floristic Endemism, South Africa, is described and illustrated. It grows almost exclusively in grassland on kommetjies, a distinctive landform of depressions and mounds associated with giant earthworms, in a restricted area west of King William's Town, Eastern Cape Province. It is the first plant taxon reported to be endemic or near-endemic to this unusual habitat. The new species appears to have been collected rarely and previous collections examined have remained undetermined to species level. The new species is distinguished from closely allied species by the combination of its usually larger dimensions in all parts, prostrate habit, coriaceous leaves that are green, scabrous, and eglandular on the adaxial surface and densely lanate on the abaxial surface, and involucral-bract morphology.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 581–588.     

Waterbird assemblages and associated habitat characteristics of farm ponds in the Western Cape, South Africa

Natural freshwater wetlands are among the most threatened habitats on Earth. Effective wetland biodiversity conservation can not, however, be evaluated without fully understanding the roles of artificial waterbodies as refuges for water-dependent plants and animals. Waterbird assemblages were examined on 59 farm ponds in the Elgin and Caledon districts of the Western Cape, South Africa. This study examines the relationship between waterbird use and habitat characteristics of farm ponds. Patterns of temporal and spatial variation of waterbird species richness and abundance were quantified in relation to the habitat characteristics of each pond. Cluster analysis and multiple regression analyses identified surface area of the farm ponds as an important variable determining the presence and abundance of many waterbird species. Structural diversity in terms of vegetation in and around the ponds was especially important in determining their usage by waterbirds. These variables were evaluated in terms of creating a mosaic of habitat types (by varying vegetation structure and pond topography), which may be a useful way to enhance waterbird diversity at farm ponds. This study concludes that the high number of farm ponds in the transformed habitat matrix of the Western Cape plays an important role in conserving waterbirds.     

Climate change drives speciation in the southern rock agama (Agama atra) in the Cape Floristic Region, South Africa

Aim Vicariance has played a major role in the evolution of the southern rock agama, Agama atra (Reptilia: Agamidae), and it is hypothesized that habitat shifts will affect small‐scale patterns of gene flow. The Cape Floristic Region (CFR) is known for high levels of diversity and endemism; thus we set out to investigate whether genetic structuring of CFR populations of A. atra corresponds to regional environmental shifts. Location Cape Fold Mountains and the Cape Floristic Region of South Africa. Methods The phylogeographical structure of 116 individuals of A. atra was determined by making use of 988 characters derived from two mitochondrial DNA fragments (control region and the NADH dehydrogenase subunit 2 coding region, ND2). Most animals originated from the CFR, but to gain a better understanding of the processes and patterns of dispersal within the species, 17 additional specimens from outside the CFR were also included and analysed in a phylogenetic context. Results Parsimony and Bayesian analyses revealed four distinct CFR clades (Cape clades) associated with geography. Phylogenetic analyses suggest that populations of A. atra in the CFR region are not entirely isolated from other populations, because some individuals from outside the CFR were nested within the four main Cape clades. The combined mitochondrial DNA data set revealed 59 distinct haplotypes in the CFR. Analysis of molecular variance (amova ) confirmed the high degree of genetic structure among the Cape clades, with more than 75% of the genetic variation found among the geographical areas. A spatial amova suggested that a ‘central clade’ originally defined as one of the four Cape clades may contain several additional populations. The main cladogenesis of A. atra within the CFR is estimated to have taken place c. 0.64–2.36 Ma. Main conclusions Agama atra shows at least four distinct genetic provinces within the CFR region, which highlights the conservation importance of this biologically diverse area. The dates of separation among the clades coincide well with the documented Pleistocene climate fluctuations, which might have contributed towards the isolation among lineages; the congruent genetic structure of A. atra with other CFR taxa further supports vicariance as a main isolating factor.     

Significant population structure and asymmetric gene flow patterns amidst expanding populations of Clinus cottoides (Perciformes,Clinidae): application of molecular data to marine conservation planning in South Africa

Clinus cottoides is a fish endemic to the coast of South Africa, predominantly inhabiting rock pools. All South African clinids are viviparous, but probably breed throughout the year; as such, their dispersal may be limited, unlike species with pelagic larval stages. We analysed 343 fish from 14 localities on the west, south and east coasts using two mitochondrial genes and the second intron of the S7 ribosomal gene. Mitochondrial DNA analyses recovered significant genetic differentiation between fish populations from the east coast and other sampling locations, with a second break found between Gansbaai and Cape Agulhas on the south coast. Nuclear DNA recovered shallower, but significant, levels of population structure. Coalescent analyses suggested remarkably asymmetrical gene flow between sampling locations, suggesting that the cold Atlantic Benguela Current and Indian Ocean Agulhas counter‐current play important roles in facilitating dispersal. There was no gene flow between the east coast and the other sites, suggesting that these populations are effectively isolated. Divergence times between them were estimated to at least 68 000 years. Neutrality tests and mismatch distributions suggest recent population expansions, with the exception of peripheral western and eastern populations (possibly a consequence of environmental extremes at the edge of the species distribution). Analyses of the current South African marine protected areas network show that it is not connected and that De Hoop, one of South Africa's largest marine reserves, appears to be an important source population of recruits to both the south and southwest coasts.     

Seasonal abundance, reproduction and hunting of common quail Coturnix coturnix in the Eastern Cape Province, South Africa

This study investigated the potential overlap between breeding and hunting seasons for a population of common quail Coturnix coturnix in the Alexandria District. Seasonal abundance (assessed as the number of birds flushed per unit effort), sex ratios and reproductive condition (based on gonad sizes) of the quail were measured over a period of 16 months, including two hunting seasons. Hunting impacts were assessed in terms of the number of birds killed per unit of hunting effort. Quail exhibited a spring peak in abundance and reproduction, which coincided with the legal hunting season, and bird numbers varied markedly between years. Sex ratio varied, with males dominating early in spring. Hunters killed nearly 60% of the birds flushed, and hunter efficiency was positively correlated with the abundance of quail. These results confirm the dilemma for managers, as quail are currently hunted during the main breeding season and offtakes are high. The development of management strategies is hampered by a lack of understanding of the population biology and ecology, and impacts of hunting. A number of possible hunting strategies are discussed. Since current harvests may not be sustainable, further research is necessary.     

Phylogeography of the Alpine salamander, Salamandra atra (Salamandridae) and the influence of the Pleistocene climatic oscillations on population divergence

Fifty individuals of the endemic Alpine salamander, Salamandra atra, representing 13 populations throughout the range of the two currently recognized subspecies, atra and aurorae, were examined for sequence variation in a large portion (1050 bp) of the mitochondrial cytochrome b gene. We revealed a large number of mitochondrial DNA (mtDNA) haplotypes (10). Interpopulation sequence divergence was very low, ranging from 0 to 3.1%. The relationships among haplotypes were poorly resolved. The divergence time estimate between several mtDNA haplotypes suggested a pre-Pleistocene differentiation approximately 3 million years ago. Moreover, the impact of the Pleistocene glaciations on the phylogeographical patterns appears to have been secondary, although a somewhat reduced genetic variability was found in populations living in areas that were directly affected by the glaciation.     

Genetic diversity,evolutionary history and implications for conservation of the lion (Panthera leo) in West and Central Africa

Aim In recent decades there has been a marked decline in the numbers of African lions (Panthera leo), especially in West Africa where the species is regionally endangered. Based on the climatological history of western Africa, we hypothesize that West and Central African lions have a unique evolutionary history, which is reflected by their genetic makeup. Location Sub‐Saharan Africa and India, with special focus on West and Central Africa. Method In this study 126 samples, throughout the lion’s complete geographic range, were subjected to phylogenetic analyses. DNA sequences of a mitochondrial region, containing cytochrome b, tRNAPro, tRNAThr and the left part of the control region, were analysed. Results Bayesian, maximum likelihood and maximum parsimony analyses consistently showed a distinction between lions from West and Central Africa and lions from southern and East Africa. West and Central African lions are more closely related to Asiatic lions than to the southern and East African lions. This can be explained by a Pleistocene extinction and subsequent recolonization of West Africa from refugia in the Middle East. This is further supported by the fact that the West and Central African clade shows relatively little genetic diversity and is therefore thought to be an evolutionarily young clade. Main conclusions The taxonomic division between an African and an Asian subspecies does not fully reflect the overall genetic diversity within lions. In order to conserve genetic diversity within the species, genetically distinct lineages should be prioritized. Understanding the geographic pattern of genetic diversity is key to developing conservation strategies, both for in situ management and for breeding of captive stocks.     

Population genetics and conservation of critically small cycad populations: a case study of the Albany Cycad,Encephalartos latifrons (Lehmann)

Declining populations of less than 250 mature individuals are symptomatic of many Critically Endangered cycads, which, globally, comprise the most threatened group of organisms as a result of collecting and habitat loss. Survival plans focus on law enforcement, reintroduction, and augmentation programmes using plants from the wild and botanical gardens. Augmentation is one of the few remaining options for cycad populations, although the assumed benefits remain untested and there is a possibility that augmentation from different sources could compromise the genetic integrity of existing populations, especially when garden plants have no provenance data. We studied Encephalartos latifrons, a South African endemic, which is a typical Critically Endangered cycad. We studied the extent and structure of genetic diversity in wild and ex situ populations to assess the potential benefits and risks associated with augmentation programmes. We examined 86 plants using amplified fragment length polymorphisms (AFLPs). The 417 AFLP markers thus generated yielded a unique DNA ‘fingerprint’ for each plant. Wild populations retain high levels of genetic diversity and this is reflected among the ex situ holdings at the Kirstenbosch Botanical Garden. No population differentiation is evident, indicating a single panmictic population, consistent with moderately high levels of gene flow between subpopulations and a sexual mode of reproduction. Bayesian clustering identified four genotype groups in the wild, as well as a genotype group only found in ex situ collections. Our results indicate that E. latifrons would benefit from augmentation programmes, including the use of undocumented collections, and careful management of breeding plants would increase the heterogeneity of propagules. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 293–308.     

Estimating population structure under nonequilibrium conditions in a conservation context: continent-wide population genetics of the giant Amazon river turtle, Podocnemis expansa (Chelonia; Podocnemididae)

Giant Amazon river turtles, Podocnemis expansa, are indigenous to the Amazon, Orinoco, and Essequibo River basins, and are distributed across nearly the entire width of the South American continent. Although once common, their large size, high fecundity, and gregarious nesting, made P. expansa especially vulnerable to over-harvesting for eggs and meat. Populations have been severely reduced or extirpated in many areas throughout its range, and the species is now regulated under Appendix II of the Convention on International Trade in Endangered Species. Here, we analyse data from mitochondrial DNA sequence and multiple nuclear microsatellite markers with an array of complementary analytical methods. Results show that concordance from multiple data sets and analyses can provide a strong signal of population genetic structure that can be used to guide management. The general lack of phylogeographic structure but large differences in allele and haplotype frequencies among river basins is consistent with fragmented populations and female natal-river homing. Overall, the DNA data show that P. expansa populations lack a long history of genetic differentiation, but that each major tributary currently forms a semi-isolated reproductive population and should be managed accordingly.     

Diet of Anguilla mossambica (Teleostei,Anguillidae) elvers in the Sundays River,Eastern Cape,South Africa

The southern African crab Hymenosoma orbiculare was recently split into five distinct species, of which three are estuarine/coastal and have peripatric distributions that are linked to temperature-defined marine bioregions. This suggests that the species’ ranges may be limited by physiological adaptations to their thermal environment. We explored this hypothesis by rearing the larvae of the warm-temperate lineage of H. orbiculare and the warm-temperate/subtropical H. longicrure at a range of water temperatures, and found clear temperature-dependent differences in the duration of larval development. Our study contributes to the growing body of evidence that stresses the importance of adaptation to regional environmental conditions, rather than physical dispersal barriers on their own, in limiting the mixing of marine species between temperature-defined biogeographic regions.     

Notes on the Habitat and Adult Behaviour of Three Red-Listed <Emphasis Type="Italic">Colophon</Emphasis> spp. (Coleoptera: Lucanidae) of the Cape Floristic Region,South Africa

The habitat restrictions of three species of Colophon (C. stokoei, C. neli, C. westwoodi) in the Cape Floristic Province are discussed, together with their patterns of seasonal activity and biological features in captivity, to augment the biological background needed for their effective conservation management.     

Contrasting genetic patterns and population histories in three threatened redfin species (Cyprinidae) from the Olifants River system, western South Africa

Differentiation occurred in different areas of the same river system in Pseudobarbus phlegethon and two sister species of the serrated redfin lineage ( Barbus calidus and Barbus erubescens ) of the Western Cape region of South Africa. In an analysis of 27 allozyme loci a deep divergence was found within P. phlegethon between the Olifants and Doring catchments (seven fixed allelic differences; 0·338 <  D  < 0·366; F _ST = 0·925). In contrast, speciation in the serrated redfin lineage occurred within the Doring catchment between B. calidus and B. erubescens (one fixed allelic difference; 0·008 <  D  < 0·052; F _ST = 0·760). Different niche preferences, behaviour and morphology probably played an important role in the formation of these contrasting patterns of genetic structure and suggest that B. calidus may have been better able to disperse between the Olifants and Doring catchments (no fixed allelic differences; 0 <  D  < 0·040; F _ST = 0·281) than P. phlegethon .     

Mitochondrial DNA sequence analysis reveals multiple Pleistocene glacial refugia for the Yellow‐spotted mountain newt,Neurergus derjugini (Caudata: Salamandridae) in the mid‐Zagros range in Iran and Iraq

Phylogeography is often used to investigate the effects of glacial cycles on current genetic structure of various plant and animal species. This approach can also identify the number and location of glacial refugia as well as the recolonization routes from those refugia to the current locations. To identify the location of glacial refugia of the Yellow‐spotted mountain newt, Neurergus derjugini, we employed phylogeography patterns and genetic variability of this species by analyzing partial ND4 sequences (867 bp) of 67 specimens from 15 sampling localities from the whole species range in Iran and Iraq. Phylogenetic trees concordant with haplotype networks showed a clear genetic structure among populations as three groups corresponding to the populations in the north, center, and south. Evolutionary ages of clades north and south ranging from 0.15 to 0.17 Myr, while the oldest clade is the central clade, corresponding to 0.32 Myr. Bayesian skyline plots of population size change through time show a relatively slight increase until about 25 kyr (around the last glacial maximum) and a decline of population size about 2.5 kyr. The presence of geographically structured clades in north, center, and south sections of the species range signifies the disjunct populations that have emerged in three different refugium. This study illustrates the importance of the effect of previous glacial cycles in shaping the genetic structure of mountain species in the Zagros range. These areas are important in terms of long‐term species persistence and therefore valuable areas for conservation of biodiversity.