Patterns of plant diversity in Africa south of the Sahara and their implications for conservation management

Plant species richness and range-size rarity in Africa south of the Sahara is concentrated in centres of plant diversity and endemism. Distribution patterns of plants mapped in the Distributiones Plantarum Africanum series and selected taxonomic monographs are analysed using the computer programme WORLDMAP. The plants are divided into four groups: herbaceous geophytes, mesophytic herbs, light-demanding shrubs and woody genera. Each group has peaks of species richness and range-size rarity at locations different to the other groups. Herbaceous geophytes and mesophytic herbs have their peaks of species richness and range-size rarity in the same location, the western Cape for geophytes and the Crystal Mountain for mesophytic herbs, whereas light-demanding shrubs and woody genera have peaks in different places. The results are discussed in relation to possible factors determining species richness and endemism and their likely conservation significance.     

Plant species richness, endemism, and genetic resources in Namibia

Namibia is a floristically diverse, arid to mesic country, with several highly distinct taxa. Including naturalized plants, there are about 4334 vascular plant species and infraspecific taxa within the country's borders, a substantial increase from the existing major reference work. Dominant families are the Poaceae (422species), Fabaceae (377), Asteraceae (385) and Mesembryanthemaceae (177). Freshwater algae and most other groups of lower plants remain poorly known. Concentrations of plant species richness are found in the Succulent Karoo biome, Kaokoveld, Otavi highland/Karstveld area, Okavango Basin, and Khomas highlands. Recent studies have led to a new estimate of 687 endemic plant species, defined as those contained wholly within Namibia's borders, amounting to about 17% of the Namibian flora. At least a further 275 species are Namib Desert endemics shared between the Kaokoveld and southern Angola (75spp.) and between the Succulent Karoo and northwestern South Africa (200spp.). Research on plant genetic resources is focused on species of potential or actual agricultural importance, such as pearl millet, Pennisetum glaucum, and cucurbits. Many wild plants have considerable genetic diversity and development potential. Primary threats to plant diversity fall in the category of poor land management and inappropriate development.     

An inflated conservation load for European butterflies: increases in rarity and endemism accompany increases in species richness

The addition of species to the European butterfly list since 1983 has resulted in a number of highly significant changes. Most important are the increases in the number and proportion of endemics and of rare species, and a regional excess of species and endemics for southern Europe compared to northern Europe. There is also a surplus of Lycaenidae and Satyridae compared to other families, and an increase in species per genus associated with the reduction in genera. These additions raise two issues. First, the potential conservation load for European butterflies is inflated at species level. This is especially the case for southern Europe, which has disproportionate increases in rare and endemic species, more particularly if rarity and endemism are found to equate with threat of extinction. Second, the inflation in rarity and endemism suggests that there is a trend to promote ever more local populations (races, subspecies) to species. The taxonomic status of species being added to the list, a quarter of which are regarded as doubtful, is increasingly difficult to determine. Consequently, there is a danger that this may call into question the validity and objectivity of taxonomic practices, and of databases dependent on them, used by conservation. Revision of higher and lower butterfly taxa is urgently required.     

The relationships between net primary productivity, human population density and species conservation

Aim In this study, I determine the relationships between net primary productivity (NPP), human population density, species richness and land use. I also examine the implications of human settlement patterns for species conservation. Location Australia. Methods I document the associations between NPP, human population density and the species richness of birds, butterflies and mammals using correlations and spatial regressions. I also assess changes in land‐use with NPP and population density, focussing particularly on protected areas. An initial exploration into the implications of the NPP‐population density relationship for regional conservation strategies is provided. Results Human population density increases with NPP suggesting that available energy may be a key driving force of human settlement patterns. The species richness of each taxonomic group and geographically restricted species also increases with NPP leading to substantial overlap between species diversity and populated regions. The percentage of land designated as minimal use decreases considerably with increasing human population density and NPP, while intensive agriculture is confined entirely to areas of high NPP. There are strong negative relationships between the size of Australia's National Parks and human population density and NPP. Small parks are often surrounded by relatively dense settlements, but have high average NPP, while large parks are mostly isolated and characterized by low productivity. There are no areas in the highest quartile of NPP that also occur in the most sparsely populated regions, presenting challenges for conservation strategies wanting to protect productive areas under the least threat of human development. Main conclusions Human population density and species richness respond similarly to variation in NPP, leading to spatial congruence between human settlements and productive, species rich regions. Planning strategies are required that minimize the potential threat posed by human development to diverse ecosystems and maximize the underlying productivity of protected areas. Reducing the level of threat may require stabilizing the size of the human population, while capturing larger areas of relatively high productivity in the conservation reserve system would lead to greater protection of local diversity.     

Extending the Namibian protected area network to safeguard hotspots of endemism and diversity

Namibia's state protected area network (PAN) covers 13.8% of the country's land area, but is seriously inadequate as a basis for effective biodiversity conservation. The early parks system was not designed with biological diversity in mind, and reflects instead a history of ideological, economic and veterinary considerations. Currently, parks in the Namib Desert biome make up 69% of the PAN, while savanna and woodland biomes are somewhat underrepresented (7.5 and 8.4% of their respective land areas), and the Karoo biome is badly underrepresented (1.6%). Four of 14 desert vegetation types are comprehensively protected, with 67 to 94% representation in the PAN, yet six savanna types have 0 to 2% representation by area. Mountain Savanna, a vegetation type unique to Namibia, is wholly unprotected. The status of two marine reserves, which in theory protect only 0.01% of Namibia's marine environment, needs clarification and augmentation with new reserves. Nearly 85% of Namibia's land is zoned for agriculture, so effective biodiversity protection means working outside the PAN to improve the sustainability and diversity of farming practices. Wildlife conservancies on commercial and communal farmlands show excellent potential to mitigate the ecological skew in the state PAN, with the ecological management of large areas being decentralized to rural communities in habitats otherwise neglected for conservation. Two important endemism zones, the Kaoko escarpment and coastal plain and the Sperrgebiet succulent steppe, plus the species-rich Caprivi area, offer three valuable opportunities for regional consolidation of protected areas into transboundary 'peace parks' or biosphere reserves.     

The biogeography of the Anura of sub-equatorial Africa and the prioritisation of areas for their conservation

There is an increasing need for protected areas to conserve biodiversity efficiently. The Anura of sub-equatorial Africa have received little attention, but we quantitatively analyse a database containing presence-only data for anurans of sub-equatorial Africa to determine patterns of distribution and species richness, and discuss the roles of present and past environmental conditions in shaping these patterns. We consider the distribution of areas rich in endemic, range-restricted and Red Data Book (RDB) species to identify areas of significance to conservation. The Eastern Highlands of Zimbabwe and adjacent area in Mozambique, southeastern Malawi and the northern coast of KwaZulu/Natal are particularly species rich, whereas the southwestern Cape of South Africa and northwestern Zambia exhibit high degrees of endemism. Four major biogeographical sub-regions are identified, which can be further subdivided into provinces. All statistically significant, current environmental factors together account for 52.6% of species richness. Annual maximum rainfall, soil type variation, minimum temperature and range of elevation were all positively correlated with species richness. Thus, both habitat influences and history appear to have influenced patterns of anuran richness in the region. Generally, areas of high species richness coincide with those high in range-restricted, endemic and RDB species. In South Africa, the northeastern coast and southwestern Cape are hypothesised to have been both refugia and centres of speciation. Results suggest that the current reserve system in sub-equatorial Africa is inadequate for the conservation of the full complement of anuran species in the region.     

Avian diversity and endemism in Namibia: patterns from the Southern African Bird Atlas Project

Namibia supports a highly diverse avifauna of 644 species, including over 90 species endemic to the southern African subregion and 13 species endemic to the country. Patterns of species diversity in relation to protected areas and habitat types were analysed using data from the Southern African Bird Atlas Project. A modified Shannon index appropriate for atlas data was used to derive an index of diversity for all species, wetland, terrestrial, threatened, and regional endemic species. Species richness for Namibian endemics was mapped. Overall species diversity is highest in the northeast of Namibia where wetland and riparian habitats coincide. Both wetland and terrestrial species show highest diversity in this area. The greatest diversity of southern African endemics falls within the Savanna-Karoo systems. Several key areas are identified for red data species, including the Caprivi Strip, Kunene and Orange Rivers, coastal wetlands and ephemeral river mouths and pans. This highlights the pressures operating on wetland and riparian habitats in arid environments. Concentrations of Namibian endemics are found in the northwestern (Kaoko) escarpment of the country. Although much of the area of high diversity of wetland, terrestrial and red data species falls within protected areas, national and regional endemics are poorly represented within national parks.     

Marine macroalgal biodiversity hotspots: why is there high species richness and endemism in southern Australian marine benthic flora?

The southern Australian marine macroalgal flora has the highest levels of species richness and endemism of any regional macroalgal flora in the world. Analyses of species composition and distributions for the southern Australian flora have identified four different floristic elements, namely the southern Australian endemic element, the widely distributed temperate element, the tropical element and a cold water element. Within the southern Australian endemic element, four species distribution patterns are apparent, thought to largely result from the Jurassic to Oligocene fragmentation of East Gondwana, the subsequent migration of Tethyan ancestors from the west Australian coast and the later invasion of high latitude Pacific species. Climatic deterioration from the late Eocene to the present is thought responsible for the replacement of the previous tropical south coast flora by an endemic temperate flora which has subsequently diversified in response to fluctuating environmental conditions, abundant rocky substrata and substantial habitat heterogeneity. High levels of endemism are attributed to Australia's long isolation and maintained, as is the high species richness, by the lack of recent mass extinction events. The warm water Leeuwin Current has had profound influence in the region since the Eocene, flowing to disperse macroalgal species onto the south coast as well as ameliorating the local environment. It is now evident that the high species richness and endemism we now observe in the southern Australian marine macroalgal flora can be attributed to a complex interaction of biogeographical, ecological and phylogenetic processes over the last 160 million years.     

Multiple transgressions of Wallace's Line explain diversity of flightless Trigonopterus weevils on Bali

The fauna of Bali, situated immediately west of Wallace''s Line, is supposedly of recent Javanese origin and characterized by low levels of endemicity. In flightless Trigonopterus weevils, however, we find 100% endemism for the eight species here reported for Bali. Phylogeographic analyses show extensive in situ differentiation, including a local radiation of five species. A comprehensive molecular phylogeny and ancestral area reconstruction of Indo-Malayan–Melanesian species reveals a complex colonization pattern, where the three Balinese lineages all arrived from the East, i.e. all of them transgressed Wallace''s Line. Although East Java possesses a rich fauna of Trigonopterus, no exchange can be observed with Bali. We assert that the biogeographic picture of Bali has been dominated by the influx of mobile organisms from Java, but different relationships may be discovered when flightless invertebrates are studied. Our results highlight the importance of in-depth analyses of spatial patterns of biodiversity.     

Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: afirst overview

Abstract Aims Ants (Hymenoptera: Formicidae) of the Baja California peninsula are poorly known, with information based largely on scattered museum and literature records. We provide the first comprehensive account of ant species occurring on the peninsula, we examine distribution patterns, and we assess the ‘peninsular effect’ which predicts that species richness declines from the base to the tip of a peninsula. Location Peninsula of Baja California, Mexico. Methods Data collection involved examining, identifying and recording label data from c. 2350 series of ants. These records provide a provisional, if incomplete, species list. We applied the incidence‐based estimator, Chao‐2, to our data base of specimen records to estimate the total number of ant species on the peninsula. We assessed endemism by comparing our peninsular species list to those from adjacent states. The peninsular effect was tested by comparing genus and species level richness between the two states of Baja California, and across five latitudinal blocks. Results We document 170 native ant species in thirty‐three genera, plus six non‐native species, in Baja California. It seems likely that additional species remain to be discovered: the Chao‐2 estimator of species richness, at 206.0 species, is about 20% higher than our observed species richness. About 30% of the species and 20% of the genera are restricted within Baja California to the relatively mesic California Floristic Province of north‐western Baja California. Nearly all of these species also occur in California. Forty‐seven species (27.6%) are peninsula endemics. Using our entire data set, the peninsular effect appears to be strong, with about twice as many species in the northern state of Baja California than are recorded from the southern state of Baja California Sur; the ratio of genera is 33 to 24. However, this effect becomes weak at the species level and absent at the genus level when minimizing habitat effects by omitting species restricted to the California Floristic Province. At a finer scale, across latitudinal blocks of about 1.9°, the number of species declines towards central portions of the peninsula and then increases in the Cape Region. Nine ant species display strongly disjunct distributions, and these occur in two general patterns: peninsula disjuncts and peninsula–mainland disjuncts. Main conclusions The Baja California peninsula supports a diverse and distinctive ant fauna, with the proportion of endemic species similar to that displayed by plants. Patterns of species and genus richness across the five latitudinal blocks provide poor support for the peninsular effect. Moreover, habitat diversity, especially that related to topographic relief, appears to be the most important factor affecting the gradient of ant species richness in Baja California. Additional collections are needed to develop a more complete species list and to determine the boundaries and status of many species. Nevertheless, the present data base provides a useful starting point for understanding the evolution of ant assemblages in Baja California and for comparison with peninsular patterns in other taxa.     

Density, body mass and parasite species richness of terrestrial mammals

We investigated the relationships between helminth species richness and body mass and density of terrestrial mammals. Cross-species analysis and the phylogenetically independent contrast method produced different results. A non-phylogenetic approach (cross-species comparisons) led to the conclusion that parasite richness is linked to host body size. However, an analysis using phylogenetically independent contrasts showed no relationship between host body size and parasite richness. Conversely, a non-phylogenetic approach generated a negative relationship between parasite richness and host density, whereas the independent contrast method showed the opposite trend – that is, parasite richness is positively correlated with host density. From an evolutionary perspective, our results suggest that opportunities for parasite colonization depend more closely on how many hosts are available in a given area than on how large the hosts are. From an epidemiological point of view, our results confirm theoretical models which assume that host density is linked to the opportunity of a parasite to invade a population of hosts. Our findings also suggest that parasitism may be a cost associated with host density. Finally, we provide some support for the non-linear allometry between density and mammal body mass (Silva and Downing, 1995), and explain why host density and host body mass do not relate equally to parasite species richness.     

The macrolepidoptera of farm woodlands: determinants of diversity and community structure

The Farm Woodland Scheme, which provided incentives to convert agricultural land to timber production, contained an implicit assumption that farm woodlands produce important benefits for wildlife. The moth fauna of 18 farm woodlands in the Vale of York was surveyed between May and November 1991. The aims were twofold. The first was to determine if there were benefits for moth species diversity. The second was to ascertain whether concepts of island biogeography and the plant species richness of the woods were related to the moth species composition.Eleven families, 214 species and over 16 000 individuals of moths were recorded. Classification of the species presence/absence matrix indicated that small woods (less than 1ha) did not have characteristic woodland moth communities. Woods larger than 5ha were judged to be more valuable for the long-term conservation of woodland moth diversity. The best predictor of moth species richness was the herbaceous plant species richness within woodlands. Species richness of the family Geometridae was positively related to woodland area, as well as to woodland shape (compact shapes being preferable to elongated shapes). Characteristic woodland species are influenced by isolation (less isolated woods being richer in species). The implications of different powers of dispersal between moth families are discussed. Farm woodlands will be of more value for the conservation of the Macrolepidoptera if they are large, compact and incorporate remnants of existing woodland with extant herbaceous vegetation. These should be factors which are taken into consideration when providing incentives to establish and manage farm woodlands.     

The effects of island ontogeny on species diversity and phylogeny

A major goal of island biogeography is to understand how island communities are assembled over time. However, we know little about the influence of variable area and ecological opportunity on island biotas over geological timescales. Islands have limited life spans, and it has been posited that insular diversity patterns should rise and fall with an island''s ontogeny. The potential of phylogenies to inform us of island ontogenetic stage remains unclear, as we lack a phylogenetic framework that focuses on islands rather than clades. Here, we present a parsimonious island-centric model that integrates phylogeny and ontogeny into island biogeography and can incorporate a negative feedback of diversity on species origination. This framework allows us to generate predictions about species richness and phylogenies on islands of different ages. We find that peak richness lags behind peak island area, and that endemic species age increases with island age on volcanic islands. When diversity negatively affects rates of immigration and cladogenesis, our model predicts speciation slowdowns on old islands. Importantly, we find that branching times of in situ radiations can be informative of an island''s ontogenetic stage. This novel framework provides a quantitative means of uncovering processes responsible for island biogeography patterns using phylogenies.