Orchid biogeography and factors associated with rarity in a biodiversity hotspot,the Southwest Australian Floristic Region

Aim The causes of orchid diversification and intrinsic rarity are poorly resolved. The Orchidaceae of the Southwest Australian Floristic Region use a diversity of pollination strategies and sites of mycorrhizal infection, and occupy a diversity of habitats. We combined a biogeographic analysis with analysis of factors associated with rarity to establish: (1) the landscape features correlated with taxon turnover and speciation, and (2) the possible role in taxon rarity of geographic region, pollination strategy, edaphic habitat and site of mycorrhizal infection. Location Southwest Australian Floristic Region. Methods The distributions of 407 orchid taxa (species and subspecies) were mapped at the quarter‐degree scale using 13,267 collections in the Western Australian Herbarium. This database was used to map taxon richness, for a biogeographic analysis and to quantify rarity of taxa. Using herbarium records, rarity was expressed as mean abundance, mean distribution and incidence of rarity based on abundance and distribution for each genus. We tested for differences in rarity of species between pollination strategies, edaphic habitats and sites of mycorrhizal infection. Results Taxon richness was highest in the High Rainfall Province. Biogeographic provincial boundaries for orchids were aligned with rainfall, while district boundaries tended to follow geological formations. When rarity was defined as either low abundance or small distribution, the greatest number of rare taxa occurred in areas of high taxon richness and naturally fragmented edaphic environments. For both abundance and distributional extent, sexual deception had a significantly higher incidence of rarity than food‐rewarding taxa. There was no significant difference in rarity with site of mycorrhizal infection. Main conclusions While large‐scale edaphic and climatic variation are correlated with orchid taxon turnover and speciation in a similar fashion to the flora in general, the processes responsible for patterns of diversity may differ. Fragmented edaphic environments appear to be associated with a higher incidence of rare species due to limited dispersal/colonization opportunities or radiations of taxa in allopatry. The high incidence of rarity in sexually deceptive taxa could be due to either low fruit set or the risk of specializing on a single pollinator species.     

Fine‐scale patterns of species and phylogenetic turnover in a global biodiversity hotspot: Implications for climate change vulnerability

Question: What is the relative importance of environmental and spatial factors for species compositional and phylogenetic turnover? Location: High‐rainfall zone of the Southwest Australian Floristic Region (SWAFR). Methods: Correlates of species compositional turnover were assessed using quadrat‐based floristic data, and establishing relationships with environmental and spatial factors using canonical correspondence analyses and Mantel tests. Between‐quadrat phylogenetic distance measures were computed and examined for correlations with environmental and spatial attributes. Processes structuring pa2t2terns of beta diversity were also evaluated within four broad floristic assemblages defined a priori. Results: Floristic diversity was strongly related to environmental attributes. A low significance of spatial variables on assemblage patterns suggested no evident effect of dispersal limitations. Species compositional turnover was especially high within the swamp and outcrop assemblage. Phylogenetic turnover was closely coupled to species compositional turnover, implying the occurrence of many locally endemic and phylogenetically relict taxa. Beta diversity patterns within assemblages were also significantly correlated with the local environment, and relevant correlates differed between floristic assemblage types. Conclusion: Phylogenetic diversity in the SWAFR high‐rainfall zone is clustered within edaphic microhabitats in a generally subdued landscape. A clustered rather than dispersed distribution of phylogenetic diversity increases the probability of significant plant diversity loss during periods of climate change. Climate change susceptibility of the region's flora is accordingly estimated to be high. We highlight the conservation significance of swamp and outcrops that are characterized by distinct hydrological properties and may provide refugial habitat for plant diversity during periods of moderate climate change.     

Environmental niche conservatism explains the accumulation of species richness in Mediterranean‐hotspot plant genera

The causes of exceptionally high plant diversity in Mediterranean‐climate biodiversity hotspots are not fully understood. We asked whether a mechanism similar to the tropical niche conservatism hypothesis could explain the diversity of four large genera (Protea, Moraea, Banksia, and Hakea) with distributions within and adjacent to the Greater Cape Floristic Region (South Africa) or the Southwest Floristic Region (Australia). Using phylogenetic and spatial data we estimated the environmental niche of each species, and reconstructed the mode and dynamics of niche evolution, and the geographic history, of each genus. For three genera, there were strong positive relationships between the diversity of clades within a region and their inferred length of occupation of that region. Within genera, there was evidence for strong evolutionary constraint on niche axes associated with climatic seasonality and aridity, with different niche optima for hotspot and nonhotspot clades. Evolutionary transitions away from hotspots were associated with increases in niche breadth and elevated rates of niche evolution. Our results point to a process of “hotspot niche conservatism” whereby the accumulation of plant diversity in Mediterranean‐type ecosystems results from longer time for speciation, with dispersal away from hotspots limited by narrow and phylogenetically conserved environmental niches.     

Biotechnology for saving rare and threatened flora in a biodiversity hotspot

The Southwest Australian Floristic Region (SWAFR) is a plant biodiversity hotspot with a geographically isolated and predominantly endemic flora. Known threatening processes (i.e. excessive clearing of native vegetation, soil salinity, soil erosion and chronic weed infestation) combined with uncertain but potentially deleterious environmental (climate) changes pose great challenges for conservation and restoration efforts. With a paucity of nature reserves, in situ protection of species can be problematic. For many species, ex situ conservation becomes the only viable strategy for saving species from extinction via seed banking or living collections established through vegetative propagation, or tissue (in vitro) culture methods. Development of specific in vitro protocols is necessary to successfully initiate culture lines, with considerable development of nutrient media, plant growth regulator regimes and incubation conditions required to optimise shoot regeneration and multiplication, especially with woody species of the SWAFR. In addition, integration of root induction and acclimatization stages has allowed significant improvements in speed and success of plant production of both endangered and difficult-to-propagate woody species. We contend that there is also considerable potential for expansion of alternative in vitro technologies such as somatic embryogenesis for difficult taxa to complement existing ex situ conservation and restoration strategies in biodiversity hotspots such as SWAFR.     

Contrasting patterns of radiation in African and Australian Restionaceae

The floras of the Mediterranean-climate areas of southern Africa and southwestern Australia are remarkably species rich. Because the two areas are at similar latitudes and in similar positions on their respective continents, they have probably had similar Cenozoic climatic histories. Here we test the prediction that the evolution of the species richness in the two areas followed a similar temporal progression by comparing the rates of lineage accumulation for African and Australian Restionaceae. Restionaceae (Poales) are typical and often dominant elements in the fynbos vegetation of the Cape Floristic Region of southern Africa and the kwongan vegetation of the Southwestern Floristic Province of Western Australia. The phylogeny of the family was estimated from combined datasets for rbcL and trnL-F sequences and a large morphological dataset; these datasets are largely congruent. The monophyly of Restionaceae is supported and a basal division into an African clade (approximately 350 species) and an Australian clade (146 species) corroborated. There is also support for a futher subdivision of these two large sister-clades, but the terminal resolution within the African clade is very weak. Fossil pollen records provided a minimum age of the common ancestor of Australian and African Restionaceae as 64-71 million years ago, and this date was used to calibrate a molecular clock. A molecular clock was rejected by a likelihood ratio test; therefore, rate changes between the lineages were smoothed using nonparametric rate smoothing. The rate-corrected ages were used to construct a plot of lineages through time. During the Palaeogene the Australian lineage diversity increased consistent with the predictions of the constant birthrate model, while the African lineage diversity showed a dramatic increase in diversification rate in the Miocene. Incomplete sampling obscures the patterns in the Neogene, but extending the trends to the modern extant diversity suggests that this acceleration in the speciation rate continued in the African clade, whereas the Australian clade retained a constant diversification rate. The substantial morphological and anatomical similarity between the African and Australian Restionaceae appear to preclude morphological innovations as possible explanations for the intercontinental differences. Most likely these differences are due to the greater geographical extent and ecological variation in temperate Australia than temperate Africa, which might have provided refugia for basal Restionaceae lineages, whereas the more mountainous terrain of southern Africa might have provided the selective regimes for a more rapid, recent speciation.     

贡嘎山蕨类植物区系的特点

贡嘎山蕨类植物区系共含４０科,９３属,３９９种,最主要的是耳蕨属Ｐｏｌｙｓｔｉｃｈｕｍ,鳞毛蕨属Ｄｒｙｏｐｔｅｒｉｓ,蹄盖蕨属Ａｔｈｙｒｉｕｍ以及水龙骨科Ｐｏｌｙｐｏｄｉａｃｅａｅ等系统演化上较高级的类群,所含的种子是喜马拉雅和中国西南当地分化的种;其特有成分多为新特有属,可认为它是随青藏高原隆起而形成的较年青的蕨类区系,热带属在本区数量多而种类少,在山上河可与北温带种类并存。贡嘎山处于蕨类物种东     

湖南桃源洞自然保护区植物区系初步研究

湖南桃源洞自然保护区地处湘东边陲,罗霄山脉中段,与江西井冈山毗邻。通过植物区系调查研究表明,全区有种子植物153科、620属、1270种(包括变种),占湖南省全部种子植物种的31.9%,其中木本植物93科、285属、748种,为湖南木本植物种的33.9%,新分布记录12种、3变种。地理成分中,热带成分306属、温带成分269属、分别为本区种子植物总属的53.2%和46.8%,表明本区具有明显的中亚热带区系性质。植物区系基本属华东区系,由于地理位置特殊,华中和华南区系的影响也非常深刻。     

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.     

Projecting climate change impacts on species distributions in megadiverse South African Cape and Southwest Australian Floristic Regions: Opportunities and challenges

Increasing evidence shows that anthropogenic climate change is affecting biodiversity. Reducing or stabilizing greenhouse gas emissions may slow global warming, but past emissions will continue to contribute to further unavoidable warming for more than a century. With obvious signs of difficulties in achieving effective mitigation worldwide in the short term at least, sound scientific predictions of future impacts on biodiversity will be required to guide conservation planning and adaptation. This is especially true in Mediterranean type ecosystems that are projected to be among the most significantly affected by anthropogenic climate change, and show the highest levels of confidence in rainfall projections. Multiple methods are available for projecting the consequences of climate change on the main unit of interest – the species – with each method having strengths and weaknesses. Species distribution models (SDMs) are increasingly applied for forecasting climate change impacts on species geographic ranges. Aggregation of models for different species allows inferences of impacts on biodiversity, though excluding the effects of species interactions. The modelling approach is based on several further assumptions and projections and should be treated cautiously. In the absence of comparable approaches that address large numbers of species, SDMs remain valuable in estimating the vulnerability of species. In this review we discuss the application of SDMs in predicting the impacts of climate change on biodiversity with special reference to the species‐rich South West Australian Floristic Region and South African Cape Floristic Region. We discuss the advantages and challenges in applying SDMs in biodiverse regions with high levels of endemicity, and how a similar biogeographical history in both regions may assist us in understanding their vulnerability to climate change. We suggest how the process of predicting the impacts of climate change on biodiversity with SDMs can be improved and emphasize the role of field monitoring and experiments in validating the predictions of SDMs.     

High species diversity and turnover in granite inselberg floras highlight the need for a conservation strategy protecting many outcrops

Determining patterns of plant diversity on granite inselbergs is an important task for conservation biogeography due to mounting threats. However, beyond the tropics there are relatively few quantitative studies of floristic diversity, or consideration of these patterns and their environmental, biogeographic, and historical correlates for conservation. We sought to contribute broader understanding of global patterns of species diversity on granite inselbergs and inform biodiversity conservation in the globally significant Southwest Australian Floristic Region (SWAFR). We surveyed floristics from 16 inselbergs (478 plots) across the climate gradient of the SWAFR stratified into three major habitats on each outcrop. We recorded 1,060 species from 92 families. At the plot level, local soil and topographic variables affecting aridity were correlated with species richness in herbaceous (HO) and woody vegetation (WO) of soil‐filled depressions, but not in woody vegetation on deeper soils at the base of outcrops (WOB). At the outcrop level, bioclimatic variables affecting aridity were correlated with species richness in two habitats (WO and WOB) but, contrary to predictions from island biogeography, were not correlated with inselberg area and isolation in any of the three habitats. Species turnover in each of the three habitats was also influenced by aridity, being correlated with bioclimatic variables and with interplot geographic distance, and for HO and WO habitats with local site variables. At the outcrop level, species replacement was the dominant component of species turnover in each of the three habitats, consistent with expectations for long‐term stable landscapes. Our results therefore highlight high species diversity and turnover associated with granite outcrop flora. Hence, effective conservation strategies will need to focus on protecting multiple inselbergs across the entire climate gradient of the region.     

South African Papilionoid Legumes Are Nodulated by Diverse Burkholderia with Unique Nodulation and Nitrogen-Fixation Loci

The root-nodule bacteria of legumes endemic to the Cape Floristic Region are largely understudied, even though recent reports suggest the occurrence of nodulating Burkholderia species unique to the region. In this study, we considered the diversity and evolution of nodulating Burkholderia associated with the endemic papilionoid tribes Hypocalypteae and Podalyrieae. We identified distinct groups from verified rhizobial isolates by phylogenetic analyses of the 16S rRNA and recA housekeeping gene regions. In order to gain insight into the evolution of the nodulation and diazotrophy of these rhizobia we analysed the genes encoding NifH and NodA. The majority of these 69 isolates appeared to be unique, potentially representing novel species. Evidence of horizontal gene transfer determining the symbiotic ability of these Cape Floristic Region isolates indicate evolutionary origins distinct from those of nodulating Burkholderia from elsewhere in the world. Overall, our findings suggest that Burkholderia species associated with fynbos legumes are highly diverse and their symbiotic abilities have unique ancestries. It is therefore possible that the evolution of these bacteria is closely linked to the diversification and establishment of legumes characteristic of the Cape Floristic Region.     

OCBIL theory: towards an integrated understanding of the evolution, ecology and conservation of biodiversity on old, climatically buffered, infertile landscapes

OCBIL theory aims to develop an integrated series of hypotheses explaining the evolution and ecology of, and best conservation practices for, biota on very old, climatically buffered, infertile landscapes (OCBILs). Conventional theory for ecology and evolutionary and conservation biology has developed primarily from data on species and communities from young, often disturbed, fertile landscapes (YODFELs), mainly in the Northern Hemisphere. OCBILs are rare, but are prominent in the Southwest Australian Floristic Region, South Africa’s Greater Cape, and Venezuela’s Pantepui Highlands. They may have been more common globally before Pleistocene glaciations. Based on the premise that natural selection has favoured limited dispersability of sedentary organisms, OCBILs should have elevated persistence of lineages (Gondwanan Heritage Hypothesis) and long-lived individuals (Ultimate Self Hypothesis), high numbers of localised rare endemics and strongly differentiated population systems. To counter such natural fragmentation and inbreeding due to small population size, ecological, cytogenetic and genetic mechanisms selecting for the retention of heterozygosity should feature (the James Effect). The climatic stability of OCBILs should be paralleled by persistence of adjacent semi-arid areas, conducive to speciation (Semiarid Cradle Hypothesis). Special nutritional and other biological traits associated with coping with infertile lands should be evident, accentuated in plants, for example, through water-foraging strategies, symbioses, carnivory, pollination and parasitism. The uniquely flat landscapes of southwestern Australia have had prolonged presence of saline lakes along palaeoriver systems favouring evolution of accentuated tolerance to salinity. Lastly, unusual resiliences and vulnerabilities might be evident among OCBIL organisms, such as enhanced abilities to persist in small fragmented populations but great susceptibility to major soil disturbances. In those places where it is most pertinent, OCBIL theory hopefully lays a foundation for future research and for better informed conservation management.     

Ants, altitude and change in the northern Cape Floristic Region

Aim Climate‐modelling exercises have demonstrated that the Cape Floristic Region is highly sensitive to climate change and will apparently lose much of its northern limits over the next few decades. Because there is little monitoring of diversity in this area, ant assemblage structure was investigated within the main vegetation types in the Greater Cederberg Biodiversity Corridor. In particular, we sought to determine how ant assemblage structure differs between the main vegetation types, how restricted ants – and in particular the major myrmecochores – are to the major vegetation types, and which environmental variables might underlie differences in the ant assemblages and in the specificity of species to particular areas. Location Northern Cape Floristic Region, Western Cape, South Africa. Methods Sampling was undertaken during October 2002 and March 2003 across an altitudinal gradient ranging from sea level (Lambert's Bay) to c. 2000 m a.s.l. (Sneeukop, Cederberg) and down again to 500 m a.s.l. (Wupperthal) in the Western Cape, South Africa. Pitfall traps were used to sample ants at 17 altitudinal bands, stretching over three vegetation types (Strandveld, Mountain Fynbos and Succulent Karoo). Biotic and abiotic environmental variables were collected at each sampling site. Generalized linear models were used to determine the relationships between species richness, density, abundance and the abundance of the major myrmecochores, and the environmental variables. Redundancy analysis was used to determine the relationship between ant assemblage structure and the environmental variables. The Indicator Value Method was used to identify characteristic ant species for each vegetation type and altitudinal site. Results Temperature explained significant proportions of the variation in species density and abundance, and, together with area and several vegetation variables, contributed significantly to the separation of the assemblages in the major vegetation types and biomes. Four major myrmecochores were identified [Anoplolepis sp. (cf. custodiens), Anoplolepis sp. (cf. steinergroeveri), Camponotus niveosetosus, Tetramorium quadrispinosum]. The abundances of the two Anoplolepis species were related to vegetation variables, while the abundance of the other two species showed opposite relationships with temperature variables. Fourteen ant species were characteristic of certain vegetation types and altitudes. Several of these species contributed to the differences between the assemblages. Main conclusions There are likely to be substantial and complex changes to ant assemblages as climates change in the northern Cape Floristic Region. Moreover, the importance of ants for ecosystem functioning suggests that these responses are not only likely to be a response solely to vegetation changes, but might also precipitate vegetation changes. The changes that are predicted to take place in the next 50 years in the Cape Floristic Region could be substantially exacerbated by such synergistic effects, which have major implications for long‐term conservation plans. Ongoing monitoring of this transect will reveal the nature and pace of the change as it unfolds.     

Community‐level changes in Banksia woodland following plant pathogen invasion in the Southwest Australian Floristic Region

Question: Does the introduced pathogen Phytophthora cinnamomi change Banksia woodland α‐ or β‐diversity and what are the implications for species re‐colonization? Location: High rainfall zone of the Southwest Australian Floristic Region (SWAFR). Methods: We measured pathogen‐induced floristic change along a disease chronosequence, and re‐sampled historic quadrats in Banksia attenuata woodlands of the SWAFR. The chronosequence represents three disease stages: (1) healthy vegetation with no disease expression; (2) the active disease front; and (3) diseased vegetation infected for at least 15 years. Comparative data were obtained by resampling diseased plots that were historically disease‐free when established in 1990. Results: β‐diversity differed substantially for both chronosequence and historic data, while α‐diversity was maintained, as measured by plot species richness and Simpson's reciprocal index. Species of known pathogen susceptibility were significantly reduced in cover–abundance, including the structurally dominant species; Banksia attenuata, B. ilicifolia and Allocasuarina fraseriana. Although these species remained present on diseased sites, there were overall reductions in canopy closure, leaf litter and basal area. These declines were coupled with an increase of species with unknown susceptibility, suggesting potential resistance and capacity to take advantage of altered site conditions. Conclusions: This study highlights the ability of an introduced plant pathogen to alter community floristics and associated stand variables. Species cover–abundances are unlikely to recover due to a reduced seed source, altered site conditions and pathogen persistence at the landscape level. However, maintenance of α‐diversity suggests continued biological significance of Phytophthora‐affected sites and the formation of novel ecosystems, themselves worthy of conservation.     

The Greater Cape Floristic Region

Aim The Cape Floristic Region (CFR) (Cape Floristic Kingdom) is currently narrowly delimited to include only the relatively mesic Cape fold mountains and adjacent intermontane valleys and coastal plains. We evaluate the floristic support for expanding the delimitation to include the whole winter‐rainfall area (arid and mesic climates) into a Greater CFR. Location Southern Africa, particularly the south‐western tip. Methods The initial divisive hierarchical classification analysis twinspan used the presence/absence of vascular plant genera to obtain major floristic groupings in southern Africa. For the more detailed analyses, we scored the flora as present/absent within a set of centres, among which the floristic relationships were investigated (agglomerative methods, upgma and minimum spanning trees). These analyses were conducted with species, genera and families separately. The centres were grouped into five regions. The species richness and endemism was calculated for the centres, regions and combination of regions. The dominant floristic components of each region were sought by calculating the percentage contribution of each family to the flora. Results The divisive method showed that the winter‐rainfall areas are floristically distinct from the rest of southern Africa. The species‐ and generic‐level analyses revealed five regions: CFR, Karoo Region, Hantam‐Tanqua‐Roggeveld Region, Namaqualand Region and Namib‐Desert Region. The CFR has the highest endemism and richness. However, the combination of the CFR, the Hantam‐Tanqua‐Roggeveld Region and the Namaqualand Region results in a higher total endemism. Combined, these three regions almost match the region delimited by the twinspan analysis, and together constitute the Greater CFR. Main conclusions The CFR constitutes a valid floristic region. This is evident from the endemism and the distinctive composition of the flora. However, the total endemism is higher for the whole winter‐rainfall area, and this supports the recognition of the larger unit. If floristic regions are to be delimited only on endemism, then the Greater CFR is to be preferred. If floristic regions are delimited on the composition of their floras at family level, then the support for such a grouping is weaker.     

Conochironomus Freeman: an Afro-Australian Chironomini genus revised (Diptera: Chironomidae)

The chironomid genus Conochironomus is revised, with diagnoses provided for both sexes of adults and, for the first time, the immature stages. The Afrotropical genotype C.acutistilus Freeman and the newly described Australian species C.australiensis (for misidentified acutistilus in Australia), C.cygnus and C.kakadu are described in all stages. C.avicula Freeman is redescribed from the male, and C.deemingi is described as new from the male alone, both from the Afrotropical Region. The Australian species C.cervus is described as new, based on the pupa alone. Comments are made on the ecology and distribution. A previously little-recognized 'tropical' Gondwanan biogeographic explanation for Conochironomus distribution is postulated.