Patterns of endemism in the limestone flora of South African lowland fynbos

Taxonomie and biological aspects of endemism and Red Data Book status were studied amongst the limestone endemics of the lowland fynbos in the Cape Floristic Region, South Africa. Of the 110 limestone endemics, 1.8% are widely distributed in the Cape Floristic Region and 56.4% are regional endemics. Relative to flora of non-limestone lowland fynbos (n=538 species), the families which were overrepresented in terms of limestone endemics included the Ericaceae, Fabaceae, Polygalaceae, Rutaceae and Sterculiaceae. The Restionaceae was the only underrepresented family. The local limestone endemics were not significantly different from regional endemics in terms of their biological attributes. An analysis of the frequency of the biological traits associated with the limestone-endemic flora established a biological profile for a limestone endemic: a dwarf-to-low shrub with soil-stored seeds which are ant or wind dispersed. In terms of the species richness of limestone endemics, the De Hoop Nature Reserve was the hotspot within the region. Relative to the total species richness, the Hagelkraal and Stilbaai areas contained higher-than-predicted numbers of rare species. These areas require urgent attention if the unique floral diversity associated with limestone substrata within the Bredasdorp-Riversdale centre of endemism is to be conserved.     

Species richness and endemism in the Western Australian flora

Aim Estimates of endemic and non‐endemic native vascular plant species in each of the three Western Australian Botanical Provinces were made by East in 1912 and Beard in 1969. The present paper contains an updated assessment of species endemism in the State. Location Western Australia comprises one third of the continental Australian land mass. It extends from 13° to 35° S and 113° to 129° W. Methods Western Australia is recognized as having three Botanical Provinces (Northern, Eremaean and South‐West) each divided into a number of Botanical Districts. Updated statistics for number of species and species endemism in each Province are based on the Census of Western Australian Plants data base at the Western Australian Herbarium ( Western Australian Herbarium, 1998 onwards). Results The number of known species in Western Australia has risen steadily over the years but reputed endemism has declined in the Northern and Eremaean Provinces where cross‐continental floras are common. Only the isolated South‐West Province retains high rates of endemism (79%). Main conclusions With 5710 native species, the South‐West Province contains about the same number as the California Floristic Province which has a similar area. The Italian mediterranean zone also contains about this number but in a smaller area, while the much smaller Cape Floristic Region has almost twice as many native species. The percentage of endemic species is highest at the Cape, somewhat less in south‐western Australia and less again in California. Italy, at 12.5%, has the lowest value. Apart from Italy, it is usual for endemism to reach high values in the largest plant families. In Western Australia, these mainly include woody sclerophyll shrubs and herbaceous perennials with special adaptations to environmental conditions. While those life forms are prominent in the Cape, that region differs in the great importance of herbaceous families and succulents, both of which are virtually absent from Western Australia. In California and Italy, most endemics are in families of annual, herbaceous perennial and soft shrub plants. It is suggested that the dominant factor shaping the South‐West Province flora is the extreme poverty of the area’s soils, a feature that emphasizes sclerophylly, favours habitat specialization and ensures relatively many local endemic species.     

Montane Refugia for Endemic and Red Listed Dragonflies in the Cape Floristic Region Biodiversity Hotspot

One of the features of many endemic organisms is that they are highly spatially restricted, and habitat specialists. The Kogelberg Biosphere Reserve (KBR) is a major centre of plant endemism within a global hotspot, the Cape Floristic Region (CFR). Dragonflies in this botanical hotspot have a range of habitat specialization from narrow-range specialists to widespread generalists, with an unusually strong bias towards the specialists. A huge 53% of dragonfly individuals and 26% of taxa recorded are national endemics, and three species are Red Listed. Thus, a group of predatory insects, which are largely not dependent on plant composition, mirrors the level of habitat specialization and restricted distributions of the plants at the spatial scale of the whole reserve. Although some studies caution the use of one taxon as a surrogate for another, the results here show that at the reserve scale in this global hotspot there can be remarkable concordance, suggesting further studies on other taxa should be carried out to determine the full extent of taxonomic concordance in this irreplaceable area.     

What can phylogenetics tell us about speciation in the Cape flora?

The spectacular diversity of the Cape flora has promoted wide speculation on the evolutionary processes behind its origins, but until recently these ideas could not be tested rigorously due to the almost complete absence of a fossil record for the region. Now, molecular phylogenetic approaches, combined with analyses of ecological and biogeographical information, offer the potential to test key hypotheses about speciation of so-called Cape clades of flowering plants. We outline the main theories and how they might be tested by phylogenetic approaches. One conclusion is that population level studies of particular species complexes are now needed to complement the growing volume of phylogenetic information for Cape clades and to provide better understanding of mechanisms of population divergence in the Cape. Another is that comparisons between Cape and non-Cape clades are needed to confirm whether speciation is indeed faster in the Cape region. An alternative possibility, that extinction rates are lower, should also be considered in these comparisons. By virtue of the ongoing, coordinated efforts by a global team of botanists, the Cape is now uniquely placed for exploring the origins and assembly of a regional assemblage or biome.     

Impact of a dam on benthic macroinvertebrates in a small river in a biodiversity hotspot: Cape Floristic Region, South Africa

Suitable reservoirs and monitoring methods are needed to manage scarce water supplies in dry countries. We assessed here the impact on aquatic macroinvertebrates of the only dam on the Eerste River, which runs through the heart of a biodiversity hotspot, the Cape Floristic Region, South Africa. The dam and associated activities, were the only forms of disturbance in this otherwise pristine area. We sampled over 20,000 macroinvertebrate individuals and illustrated some categorical effects of the impoundment and its effects on macroinvertebrate assemblages. Macroinvertebrate species diversity below the dam was only half of that in the pristine catchment area above the dam. Furthermore, Ephemeroptera, Plecoptera and Trichoptera diversity and abundance dropped to almost zero as a result of the impoundment. In contrast, the abundance of the Diptera family Chironomidae increased substantially below the dam. These changes in macroinvertebrate diversity mirrored those recorded in biologically less diverse areas, but are of major concern in this biodiversity hotspot with its rich endemic fauna. We conclude that such an impoundment, while important for human welfare, results in a high price being paid in terms of loss of local biodiversity.     

Floristic diversity in the Cape Flora of South Africa

Comprising a land area of ca 90000 km2, less than 4% of the total land area for the Southern African subcontinent, the Cape Floristic Province is one of the world's richest areas in terms of botanical diversity for its size. An estimated 8650 species of vascular plants occur in this area, about 65% of which are endemic. This is about 42% of the estimated total for all of southern Africa. The number of species packed into so small an area is remarkable for the temperate zone, and compares closely with species totals for areas of comparable size in the wet tropics. The Cape Region consists of a mosaic of sandstone and shale substrates that give rise to soils of quite different types, and in addition local areas of limestone add to the edaphic diversity. Climates across the region are extremely variable, and the predominant orographic rainfall pattern ranges from 2000mm locally to less than 100mm, often with extremely steep gradients, the result of a mountainous landscape. The edaphic diversity resulting from a mosaic of different soils is compounded by sharp local gradients in precipitation that creates an unusual number of local habitats. A feature of the some of the soils in the Region is low nutrient levels and many of the plants on such soils have low seed dispersal capabilities, a factor important in explaining the high levels of local endemism. Species richness in the Cape Region is hypothesized to have resulted from the presence of a complex mosaic of diverse habitats and steep ecological gradients against a background of relatively stable climate and geology after the mediterranean climate was established there sometime after the beginning of the Pliocene. A local or ecological mode of speciation may have been more important under these conditions than allopatric speciation.     

The Cape Peninsula, South Africa: physiographical, biological and historical background to an extraordinary hot-spot of biodiversity

The Cape Peninsula, a 470 km² area of rugged scenery and varied climate, is located at the southwestern tip of the Cape Floristic Region, South Africa. The Peninsula is home to 2285 plant species and is a globally important hot-spot of biodiversity for higher plants and invertebrates. This paper provides a broad overview of the physiography, biological attributes and history of human occupation of the Peninsula. The Peninsula is characterized physiographically by extremely high topographical heterogeneity, very long and steep gradients in annual rainfall, and a great diversity of nutrient-poor soils. Thus, the Peninsula supports a high number of habitats and ecological communities. The predominant vegetation is fynbos, a fire-prone shrubland, and 12 broadly characterized fynbos types have been described on the Peninsula. Animal community structure, especially with regard to invertebrates, is poorly known. Vertebrate community structure is probably strongly influenced by nutrient poverty and recurrent fire. Generally, most vertebrates are small and typically occur in low numbers. Some invertebrates play keystone roles in facilitating ecological processes. Human occupation of the Peninsula was limited, until relatively recently, by nutrient poverty. After Dutch colonization in 1652, direct and indirect impacts on the natural ecosystems of the Peninsula escalated dramatically, and by 1994, some 65% of original natural habitat was either transformed by urbanization and agriculture, or invaded by alien plants. Nonetheless, there is still excellent potential to conserve the Cape Peninsula's remaining biodiversity.     

Biogeographic patterns and phylogeography of dwarf chameleons (Bradypodion) in an African biodiversity hotspot

The southern African landscape appears to have experienced frequent shifts in vegetation associated with climatic change through the mid-Miocene and Plio-Pleistocene. One group whose historical biogeography may have been affected by these fluctuations are the dwarf chameleons (Bradypodion), due to their associations with distinct vegetation types. Thus, this group provides an opportunity to investigate historical biogeography in light of climatic fluctuations. A total of 138 dwarf chameleons from the Cape Floristic Region of South Africa were sequenced for two mitochondrial genes (ND2 and 16S), and resulting phylogenetic analyses showed two well-supported clades that are distributed allopatrically. Within clades, diversity among some lineages was low, and haplotype networks showed patterns of reticulate evolution and incomplete lineage sorting, suggesting relatively recent origins for some of these lineages. A dispersal-vicariance analysis and a relaxed Bayesian clock suggest that vicariance between the two main clades occurred in the mid-Miocene, and that both dispersal and vicariance have played a role in shaping present-day distributions. These analyses also suggest that the most recent series of lineage diversification events probably occurred within the last 3-6 million years. This suggests that the origins of many present-day lineages were founded in the Plio-Pleistocene, a time period that corresponds to the reduction of forests in the region and the establishment of the fynbos biome.     

Profiling a besieged flora: endemic and threatened plants of the Cape Peninsula, South Africa

The Cape Peninsula (area: 471 km²), situated at the south-western extremity of the Cape Floristic Region, has exceptionally high plant species richness (2285 species and infraspecific taxa) and numbers of endemic (90; 88 species and two infraspecific) and threatened (141; 138 species and three infraspecific) taxa (termed species from here on). This biodiversity is threatened by urban development and the spread of invasive alien plants. Peninsula endemics are concentrated in a few, predominantly species-rich families and these correspond well with endemic-rich families in other areas of the Cape Floristic Region. A high level of similarity exists between families with threatened and families with endemic species. A frequency analysis of the biological traits of both endemic and threatened species shows that low growing, ant-dispersed shrubs are over-represented in both groups. Endemics are most likely to be non-sprouters, but threatened plants do not have a specific post-fire regeneration strategy. Threatened species have higher frequencies of geophytes, sprouters and wind-dispersed species compared to endemic species. Numbers of endemic and threatened species are not randomly distributed with regard to occurrence in vegetation types and patterns are similar for both groups. The habitat and biological profiles of both endemic and threatened species suggest that they are highly vulnerable to extinction as a result of increasing rates of alien plant infestation, urbanization and inappropriate fire regimes.     

A pragmatic approach to estimating the distributions and spatial requirements of the medium- to large-sized mammals in the Cape Floristic Region, South Africa

Conservation planning in the Cape Floristic Region, a recognized world plant diversity hotspot, required systematic information on the estimated distributions and spatial requirements of the medium- to large-sized mammals within each of 102 Broad Habitat Units delineated according to key biophysical parameters. As a consequence of a general lack of data, we derived a pragmatic approach for obtaining estimates of these two parameters. Distribution estimates were based on a combination of a literature survey (with emphasis on early texts) and the ecological requirements of the species. Spatial requirement estimates were derived from a simple spreadsheet model that is based on forage availability estimates and the metabolic requirements of the mammals in question. Our analysis incorporated adaptations of the agriculture-based Large Stock Unit or Animal Unit approach. The predictions of the model were tested by comparing them with actual density data. The outcomes provided realistic estimates of the two parameters. However, they should be considered as testable hypotheses and the concept of adaptive management — or management by hypothesis — must apply. Examples of the outcomes are provided in the form of maps and tables.     

Insect diversity in the Cape Floristic Region

The Cape Floristic Region (CFR) is characterized by exceptionally high plant species richness and it is yet to be determined whether this is matched by an equally high diversity of insect fauna associated with these plants. In an attempt to do this, data from the literature on the number of insects for various taxa found at different sites in the CFR were compared with equivalent data from other areas in South Africa and elsewhere. Results indicate that the herbivorous insect fauna of the CFR is not particularly rich in species, perhaps except for the small guild of endophagous insects. It is speculated that this is the result of CFR vegetation being a poor food source for herbivores, particularly in view of its sclerophyllous nature. Plant defence mechanisms, such as a pronounced cyanogenic ability of the leaves coupled with the relatively simple architecture of the plants are possible contributing factors.     

Current and future threats to plant biodiversity on the Cape Peninsula, South Africa

The biodiversity of the Cape Peninsula (49127 ha in extent) has been considerably affected by various factors since European settlement in 1652. Urbanization and agriculture have transformed 37% of the original area of natural vegetation. Lowland vegetation types have been worst affected, with almost half of the transformation occurring in one of the 15 recognized vegetation types. Vegetation at high altitudes has been little affected by urbanization and agriculture, but alien trees and shrubs are now threatening biodiversity in these areas. Of the area not affected by urbanization and agriculture 10.7% is currently under dense stands (>25% canopy cover) of alien plants and another 32.9% is lightly invaded. Dense stands of Acacia cyclops, the most widespread invader, cover 2510 ha, 76% of the total area under dense alien stands. This paper assesses the impacts of these factors on aspects of the plant biodiversity of the area, namely, the distribution of major vegetation types and of endemic, rare and threatened plant taxa and of taxa in the Proteaceae (a prominent element in almost all communities, taken as an indicator of overall plant biodiversity).Possible future impacts on biodiversity are assessed by considering the effects of several scenarios involving increased urbanization and changes to alien plant control strategies and further spread over the next 50–100 years. The worst-case scenario for urbanization sees the area under natural vegelation reduced to 12163 ha (39.3% of its extent in 1994, or 24.8% of its original extent). Under this scenario almost a quater of the 161 endemic, rare and threatened (special) taxa are confined totally to urban areas; 57.4% of the known localities of these taxa, and 40.1% of the remaining localities of Proteaceae taxa are transformed. Dense alien stands currently affect 29.8% of the localities of special taxa known from herbarium records and 8.4% of these taxa currently occur only in areas at present affected by aliens. Clearing all dense stands would result in 62.9% of special taxa having less than half of their known localities affected (49.1% at present). Under this scenario, 92% of Proteaceae taxa have more than 75% of their localities unaffected by aliens. If clearing is confined to specific areas (the Cape Peninsula Protected Natural Environment or all publicly-owned land) and the aliens spread further outside these areas, the area of natural vegetation remaining shrinks (to 82.4% of the current extent if control is confined to public land). The further losses in biodiversity associated with these scenarios are described. If control programmes collapse and all potentially invadable land is occupied by dense alien stands, only 407 ha of natural vegetation would remain (1.5% of the current extent).The probability of the various scenarios materializing is discussed. To minimize further losses in biodiversity it is essential that: (1) a major initiative is launched immediately to clear (firstly) the 10184 ha of lightly invaded vegetation and then the 3313 ha of densely invaded vegetation; (2) no urban development be permitted within the boundaries of the Cape Peninsula Protected Natural Environment; (3) a systematic programme of prescribed burning (linked to the alien control programme) is initiated; and (4) contingency measures are implemented to improve the status of seriously threatened taxa, habitats and vegetation types.     

High ploidy diversity and distinct patterns of cytotype distribution in a widespread species of Oxalis in the Greater Cape Floristic Region

Background and Aims

Genome duplication is widely acknowledged as a major force in the evolution of angiosperms, although the incidence of polyploidy in different floras may differ dramatically. The Greater Cape Floristic Region of southern Africa is one of the world''s biodiversity hotspots and is considered depauperate in polyploids. To test this assumption, ploidy variation was assessed in a widespread member of the largest geophytic genus in the Cape flora: Oxalis obtusa.

Methods

DNA flow cytometry complemented by confirmatory chromosome counts was used to determine ploidy levels in 355 populations of O. obtusa (1014 individuals) across its entire distribution range. Ecological differentiation among cytotypes was tested by comparing sets of vegetation and climatic variables extracted for each locality.

Key Results

Three majority (2x, 4x, 6x) and three minority (3x, 5x, 8x) cytotypes were detected in situ, in addition to a heptaploid individual originating from a botanical garden. While single-cytotype populations predominate, 12 mixed-ploidy populations were also found. The overall pattern of ploidy level distribution is quite complex, but some ecological segregation was observed. Hexaploids are the most common cytotype and prevail in the Fynbos biome. In contrast, tetraploids dominate in the Succulent Karoo biome. Precipitation parameters were identified as the most important climatic variables associated with cytotype distribution.

Conclusions

Although it would be premature to make generalizations regarding the role of genome duplication in the genesis of hyperdiversity of the Cape flora, the substantial and unexpected ploidy diversity in Oxalis obtusa is unparalleled in comparison with any other cytologically known native Cape plant species. The results suggest that ploidy variation in the Greater Cape Floristic Region may be much greater than currently assumed, which, given the documented role of polyploidy in speciation, has direct implications for radiation hypotheses in this biodiversity hotspot.     

西倾山地区植物区系研究

西倾山地区位于青海省东南部 ,北纬 33°80′～ 35°50′ ,东经 10 0°10′～ 10 2°4 0′。本地区共有种子植物 56科、 2 57属、 711种。区系特征如下 :(1)温带成分占 96 88% ,以绝对优势确定了本区系明显的温带性质。 (2 )世界成分多为衍生 ,更兼所含的中国特有属亦多为衍生 ,表明本区系的年轻性和衍生的性质。 (3)主要植被均属典型的高寒类型 ,它们的建群种亦多为典型的高寒植被的特征种。所以 ,本区系高寒植被典型 ,特征种典型 ,高寒区系特色明显。(4 )生态环境对本区系特征的塑造表现为高寒生态因子的选择和高山特化的作用强烈 ,而在一定程度上湿冷生性质和寒旱生性质的高山特化作用更为明显。西倾山地区的植物区系是唐古特地区的代表区系之一。但在中国特有种的分析中 ,本区系却与横断山和甘肃南部区系的联系最为密切 ,三者间的共有种最为丰富 ,有 36 3种 ,占本区中国特有种的 6 6 73%。如此大的相似性 ,无疑且有力地支持了李锡文等 (1993)关于横断山地区的“川西北、甘西南、青东南小区”划分的观点。     

Endemism in Namibia: patterns, processes and predictions

Sandwiched between the Namib and Kalahari Deserts of southwestern Africa are the karooid and escarpment biotopes of Namibia which are rich in endemics of many taxa. Most plant, invertebrate, amphibian, reptile, mammal and bird species endemic to Namibia are found in a zone running through, and to the west of, Namibia's escarpment region. There is also an important region of endemism for succulent plants, reptiles and invertebrates in the Succulent Karoo biome. Congruence between endemism hotspots, particularly on rocky substrates, is remarkably high for most taxa, implying broadly similar speciation processes. Possible speciation mechanisms in different parts of the country include the spatial isolation of rupicolous taxa such as insects and reptiles by the formation of large coastal dune fields; the expansion and contraction of wooded savannas during pluvial and interpluvial periods; and global temperature shifts which created highland refugia for frost-susceptible plants and poikilotherms. Areas of endemism and species richness overlap poorly for Namibia's mainly arid-dwelling endemic vertebrates, as richness is highest in the mesic wetlands and woodlands of northeast Namibia. The overlap for succulent plants, insects and arachnids, however, is relatively high. Centres of endemism for plants and vertebrates fall mainly outside protected areas, as few parks were established with biodiversity indices in mind. Our analysis of endemism congruence provides a strong platform for the promulgation of new protected areas to safeguard Namibia's unique biota. Furthermore, analysis of speciation patterns and processes is a useful predictive tool for the identification of other biotically important sites.     

Non-linearities, synergisms and plant extinctions in South African fynbos and Australian kwongan

This paper explores the determinants of extinction of endemic plant taxa in mediterranean-climate regions in South Africa and southwestern Australia. Major threats to biodiversity in these areas include agriculture, deforestation, fragmentation, invasive alien organisms and urbanization. Case studies from the two regions show that synergisms between factors can lead to discontinuous, or non-linear, responses that have increased extinction rates (or threaten to) beyond predictions based on simple deterministic processes.     

巴颜喀拉山地区植物区系研究

巴颜喀拉山地区位于青海省南部 ,北纬 32°2 0′～ 35°15′ ,东经 94°5 0′～ 10 1°4 5′ ,面积约84 0 0 0km2 ,海拔高程介于 36 0 0m至 5 36 9m之间。本地区拥有种子植物 1116种 ,分属于 6 4科 ,2 95属。区系特征概述如下 :1)对属的分布区类型的分析表明 ,本地区植物区系以北温带成分为主 ,仅有 9个分布区从热带延伸到温带的热带属 ,且在本区所含种类极少或仅有 1种。 2 )对种的分布区类型的分析表明 ,大多数种 (不包括中国特有种 )属于中亚成分和东亚成分 ,且多呈中亚 -喜马拉雅 -中国西南或中国 -喜马拉雅分布式样。以此为据所得结论是 ,本地区的种以温带成分为优势 ,具高原、高山分布的特点。 3)本区东南部的生态环境和区系成分的来源均不同于其余地区。东亚分布型及其变型中国 -喜马拉雅的种聚集在东南部。这里被认为是那些来源于横断山和西秦岭的区系成分的一个通道。 4 )对中国特有种的分析表明 ,本区系与横断山区系和甘肃南部区系联系最为密切。 5 )在高山特化作用和高山生态因子的选择之下 ,本区植物获得了适应寒冷和干旱的特性。 6 )本区植物的耐寒性常常伴随着耐旱和耐湿的双重特性 ,植物的这些特性也反映在它们的地理分布上。 7)本区的区系性质和特点与唐古特地区基本一致 ,因此本植物区     

New species and combinations in the African Restionaceae

Eight new species of the African Restionaceae (Restionoideae) are described, viz.: Cannomois anfracta, Cannomois arenicola, Cannomois grandis, Nevillea vlokii, Thamnochortus kammanassiae, Willdenowia pilleata, Restio uniflorus and Restio mkambatiae. A key to the species of Cannomois is provided, as well as a table comparing the characters of the three species in Nevillea. For all new species, notes on the affinities of the species and their habitats are provided. Two new combinations, Cannomois primosii (Pillans) H.P. Linder and Cannomois robusta (Kunth) H.P. Linder, are made.