South-western Australian Biodiversity Hotspot Research Articles

A revision of the white-headed spiny trapdoor spiders of the genus Euoplos (Mygalomorphae: Idiopidae: Arbanitinae): a remarkable lineage of rare mygalomorph spiders from the south-western Australian biodiversity hotspot

The ‘white-headed’ spiny trapdoor spiders of the Euoplos mcmillani-group are revised, and three new species are described from the biodiversity hotspot of south-western Western Australia: E. cornishi sp. nov., E. kalbarri sp. nov. and E. saplan sp. nov. Two previously described species, E. ballidu (Main, 2000) and E. mcmillani (Main, 2000), are further re-described and re-diagnosed. The mcmillani-group of Euoplos are remarkable for the unique ‘bright white’ carapace coloration of males, and we here document the known diversity and conservation status of these rare and enigmatic spiders.

Cryptic species in an ancient flowering‐plant lineage (Hydatellaceae, Nymphaeales) revealed by molecular and micromorphological data

The flora of the southwestern Australian biodiversity hotspot is rich in endemic species, many of which remain to be discovered or properly described; estimates of species diversity and levels of endemism should take into account the possible occurrence of cryptic species. Here we explore taxonomic diversity in a Western Australian lineage belonging to the primarily Australian genus Trithuria, the sole genus of Hydatellaceae (Nymphaeales). Recent molecular evidence supports the existence of cryptic species in self‐pollinating members of section Trithuria. We investigate Western Australian plants currently classified as T. australis s.l., a self‐pollinating member of the section Hydatella. Using evidence from microsatellite data (SSRs), an expanded molecular phylogenetic analysis based on four plastid markers, and fruit micromorphology, we suggest that material traditionally classified as T. australis s.l. belongs to at least four species. Two species occur in the northern part of the distribution range of the group (31° S to 33°27′ S), and two in the southern part (33°27′ S to 35° S). Each northern species has distinctive fruit micromorphology not recorded in other members of the genus. The two southern species are well characterized by molecular characters and seem to be allopatric, but lack obvious morphological differences from each other. We describe one of the northern species as T. fitzgeraldii sp. nov. However, clarifying the names of the other three species is currently problematic as T. australis and another available name are based on collections made 117 years ago, from localities distant from any subsequent records of Hydatellaceae. Based on genome size estimations, we also demonstrate two ploidy levels in the T. australis complex. Our study supports the view that species diversity in Hydatellaceae is strongly underestimated.

Evolutionary and natural history of the turtle frog, Myobatrachus gouldii, a bizarre myobatrachid frog in the southwestern Australian biodiversity hotspot.

Southwest Australia (SWA) is a global biodiversity hotspot and a centre of diversity and endemism for the Australo-Papuan myobatrachid frogs. Myobatrachus gouldii (the turtle frog) has a highly derived morphology associated with its forward burrowing behaviour, largely subterranean habit, and unusual mode of reproduction. Its sister genera Metacrinia and Arenophryne have restricted distributions in Western Australia with significant phylogeographic structure, leading to the recent description of a new species in the latter. In contrast, Myobatrachus is distributed widely throughout SWA over multiple climatic zones, but little is known of its population structure, geographic variation in morphology, or reproduction. We generated molecular and morphological data to test for genetic and morphological variation, and to assess whether substrate specialisation in this species may have led to phylogeographic structuring similar to that of other plant and animal taxa in SWA. We assembled sequence data for one mitochondrial and four nuclear DNA loci (3628 base pairs) for 42 turtle frogs sampled throughout their range. Likelihood phylogenetic analyses revealed shallow phylogeographic structure in the mtDNA locus (up to 3.3% genetic distance) and little variation in three of the four nDNA loci. The mtDNA haplotype network suggests five geographically allopatric groups, with no shared haplotypes between regions. These geographic patterns are congruent with several other SWA species, with genetic groups restricted to major hydrological divisions, the Swan Coastal Plain, and the Darling Scarp. The geographically structured genetic groups showed no evidence of significant morphological differentiation (242 individuals), and there was little sexual size dimorphism, but subtle differences in reproductive traits suggest more opportunistic breeding in lower rainfall zones. Call data were compared to sister genera Metacrinia and Arenophryne and found to be highly conservative across the three genera. Like many taxa in SWA, topographic variation and Plio-Pleistocene arid fluctuations likely were historic drivers of diversification in M. gouldii.

Fossil evidence for open, Proteaceae-dominated heathlands and fire in the Late Cretaceous of Australia.

The origin of biomes is of great interest globally. Molecular phylogenetic and pollen evidence suggest that several plant lineages that now characterize open, burnt habitats of the sclerophyll biome, became established during the Late Cretaceous of Australia. However, whether this biome itself dates to that time is problematic, fundamentally because of the near-absence of relevant, appropriately aged, terrestrial plant macro- or mesofossils. We recovered, identified, and interpreted the ecological significance of fossil pollen, foliar and other remains from a section of core drilled in central Australia, which we dated as Late Campanian-Maastrichtian. The sediments contain plant fossils that indicate nutrient-limited, open, sclerophyllous vegetation and abundant charcoal as evidence of fire. Most interestingly, >30 pollen taxa and at least 12 foliage taxa are attributable to the important Gondwanan family Proteaceae, including several minute, amphistomatic, and sclerophyllous foliage forms consistent with subfamily Proteoideae. Microfossils, including an abundance of Sphagnales and other wetland taxa, provided strong evidence of a fenland setting. The local vegetation also included diverse Ericaceae and Liliales, as well as a range of ferns and gymnosperms. The fossils provide strong evidence in support of hypotheses of great antiquity for fire and open vegetation in Australia, point to extraordinary persistence of Proteaceae that are now emblematic of the Mediterranean-type climate southwestern Australian biodiversity hotspot and raise the profile of open habitats as centers of ancient lineages.

Disjunct, highly divergent genetic lineages within two rareEremophila(Scrophulariaceae: Myoporeae) species in a biodiversity hotspot: implications for taxonomy and conservation

Effective conservation management should target appropriate conservation units, but evolutionarily and genetically divergent lineages within nominal taxa are often unrecognized. The south-western Australian biodiversity hotspot may harbour many cryptic taxa, as it contains many plant species with naturally fragmented population distributions. Using microsatellite markers, we tested the hypothesis that disjunct population groups in the rare species Eremophila microtheca and E. rostrata (Scrophulariaceae: Myoporeae) are highly genetically divergent and represent separate evolutionarily significant units (ESUs). Chromosome counts indicated that all individuals assessed were diploid (2n = 36). Genetic differentiation among disjunct population groups was highly significant (P < 0.001) for both E. microtheca (FST = 0.301–0.383; Dest = 0.756–0.774) and E. rostrata (FST = 0.325–0.346; Dest = 0.628–0.660), and was similar to their differentiation from allied species. These results, including high incidences of private alleles, suggest historical divergence among cryptic taxa within E. microtheca and E. rostrata. Population groups in E. rostrata have recently been taxonomically recognized as two subspecies. Our study suggests that E. microtheca should also be reassessed as two taxa or considered as two ESUs, and the southern occurrence should be listed as Critically Endangered. We suggest a precautionary approach for flora in this and similar landscapes, whereby historically wide geographical disjunctions are assumed to indicate separate units for conservation. © 2014 State of Western Australia. Botanical Journal of the Linnean Society © 2014 The Linnean Society of London, 2015, 177, 96–111.

Phosphorus nutrition of phosphorus-sensitive Australian native plants: threats to plant communities in a global biodiversity hotspot.

South-western Australia harbours a global biodiversity hotspot on the world's most phosphorus (P)-impoverished soils. The greatest biodiversity occurs on the most severely nutrient-impoverished soils, where non-mycorrhizal species are a prominent component of the flora. Mycorrhizal species dominate where soils contain slightly more phosphorus. In addition to habitat loss and dryland salinity, a major threat to plant biodiversity in this region is eutrophication due to enrichment with P. Many plant species in the south-western Australian biodiversity hotspot are extremely sensitive to P, due to a low capability to down-regulate their phosphate-uptake capacity. Species from the most P-impoverished soils are also very poor competitors at higher P availability, giving way to more competitive species when soil P concentrations are increased. Sources of increased soil P concentrations include increased fire frequency, run-off from agricultural land, and urban activities. Another P source is the P-fertilizing effect of spraying natural environments on a landscape scale with phosphite to reduce the impacts of the introduced plant pathogen Phytophthora cinnamomi, which itself is a serious threat to biodiversity. We argue that alternatives to phosphite for P. cinnamomi management are needed urgently, and propose a strategy to work towards such alternatives, based on a sound understanding of the physiological and molecular mechanisms of the action of phosphite in plants that are susceptible to P. cinnamomi. The threats we describe for the south-western Australian biodiversity hotspot are likely to be very similar for other P-impoverished environments, including the fynbos in South Africa and the cerrado in Brazil.

Systematics of Diplodactylus (Squamata: Diplodactylidae) from the south-western Australian biodiversity hotspot: redefinition of D. polyophthalmus and the description of two new species

– The Australian gecko genus Diplodactylus is characterised by morphologically similar but genetically divergent lineages and taxa. Recent molecular analysis indicated the presence of an undescribed Diplodactylus from the Geraldton Sandplain on the western coast of Australia, and a relatively deep divergence between populations of D.

Molecular phylogeny and morphological revision of the Ctenotus labillardieri (Reptilia: Squamata: Scincidae) species group and a new species of immediate conservation concern in the southwestern Australian biodiversity hotspot

Molecular phylogeny and morphological revision of the Ctenotus labillardieri (Reptilia: Squamata: Scincidae) species group and a new species of immediate conservation concern in the southwestern Australian biodiversity hotspot

Molecular phylogeny and morphological revision of the Ctenotus labillardieri (Reptilia: Squamata: Scincidae) species group and a new species of immediate conservation concern in the southwestern Australian biodiversity hotspot

Ctenotus is the largest and most diverse genus of skinks in Australia with at least 97 described species. We generated largemitochondrial and nuclear DNA data sets for 70 individuals representing all available species in the C. labillardieri species-group to produce the first comprehensive phylogeny for this clade. The widespread C. labillardieri was sampled extensively toprovide the first detailed phylogeographic data set for a reptile in the southwestern Australian biodiversity hotspot. Wesupplemented our molecular data with a comprehensive morphological dataset for the entire group, and together these data areused to revise the group and describe a new species. The morphologically highly variable species C. labillardieri comprisesseven well-supported genetic clades that each occupy distinct geographic regions. The phylogeographic patterns observed inthis taxon are consistent with studies of frogs, plants and invertebrates, adding strength to emerging biogeographic hypothesesin this iconic region. The species C. catenifer, C. youngsoni, and C. gemmula are well supported, and despite limited samplingboth C. catenifer and C. gemmula show substantial genetic structure. The threatened C. lancelini from Lancelin Island and theadjacent mainland is the sister taxon to a new species from the Swan Coastal Plain, which we describe as C. ora sp. nov. Thisspecies is a habitat specialist, occurring primarily in sandy regions south of Perth that currently are under intense development. Ctenotus ora sp. nov. should be considered for conservation attention immediately.

Effects of vicariant barriers, habitat stability, population isolation and environmental features on species divergence in the south‐western Australian coastal reptile community

Identifying explicit hypotheses regarding the factors determining genetic structuring within species can be difficult, especially in species distributed in historically dynamic regions. To contend with these challenges, we use a framework that combines species distribution models, environmental data and multi-locus genetic data to generate and explore phylogeographic hypotheses for reptile species occupying the coastal sand-dune and sand-plain habitats of the south-western Australian biodiversity hotspot, a community which has both a high diversity of endemics and has varied dramatically in spatial extent over time. We use hierarchical amova, summary statistic and distance-based analyses to explicitly test specific phylogeographic hypotheses. Namely, we test if biogeographic vicariance across barriers, habitat stability, population isolation along a linear habitat or fragmentation across different environments can explain genetic divergence within five co-distributed squamate reptile species. Our results show that patterns of genetic variation reflect complex and species-specific interactions related to the spatial distribution of habitats present currently and during repeated glacial minima, as opposed to being associated with historical factors such as habitat stability between glacial and inter-glacial periods or vicariant barriers. We suggest that the large impact of habitat characteristics over time (i.e. relative levels of habitat connectivity, climatic gradients and spatial heterogeneity of soil types) reflects the ecological restrictions of the sand-dune and sand-plain reptile communities and may explain the lack of concordance across taxa. The study demonstrates the general utility of the approach for assemblage-level, as well as single species, phylogeographic study, including its usefulness for exploring biologically informed hypotheses about what factors have influenced patterns of genetic variation.

Phylogenetic relationships of the heath dragons (Rankinia adelaidensis and R. parviceps) from the south-western Australian biodiversity hotspot

Although the south-western Australian region is recognised as a global biodiversity hotspot, there are still significant gaps in our understanding of the biodiversity of this region. We present a phylogenetic study of the heath dragons (Rankinia adelaidensis and R. parviceps) from this region, incorporating a 1612-bp section of mtDNA and two nuclear introns, Gapdh (~244 bp) and Enol (~330 bp). In addition, we present a generic-level analysis of three gene regions (mtDNA, Gapdh, BDNF), which provides clear evidence that Rankinia adelaidensis and R. parviceps are not closely related to Rankinia diemensis from eastern Australia. Instead, the heath dragons are strongly supported as forming a clade with the genus Ctenophorus. In addition, we find that there are significant levels of haplotype divergence between currently recognised subspecies of the heath dragons (R. a. adelaidensis, R. a. chapmani, R. p. parviceps, R. p. butleri). We suggest that the genetic divergences between subspecies result from geographic isolation in allopatry owing to habitat preferences, followed by drift and/or selection. On the basis of these deep divergences and consistent morphological differences between subspecies, we recommend elevating all taxa to full species, and provide a taxonomic revision of the genera Rankinia and Ctenophorus.

Biogeography and speciation of a direct developing frog from the coastal arid zone of Western Australia

Within the southwestern Australian biodiversity hotspot, the Shark Bay region displays high levels of plant and animal endemism, particularly in the herpetofauna. The region has been subjected to dramatic climatic fluctuations and has been geologically active from the Late Miocene to the present. The myobatrachid frog Arenophryne rotunda, a Shark Bay endemic, provides an ideal opportunity to examine the relative effects of fluctuating climates and geological activity on the biota of Shark Bay. A comprehensive phylogeographic analysis of A. rotunda, based on data comprising 1154bp of the mitochondrial gene ND2, is presented. My results demonstrate a major genetic break that divides this species at the northern edge of the Victorian Plateau into northern and southern species lineages, dating to the Late Miocene, with a further division of the southern species lineage across the Murchison Gorge dating to the Plio-Pleistocene border. Both of these periods are related to prominent geological activity and climatic shifts in the Shark Bay region. Interpretation of phylogeographic results point to the prominent role of fluctuating Pleistocene climates and associated coastal landscape evolution in the generation of phylogeographic structure within the distinct A. rotunda species lineages. Similar processes have been invoked to explain the diversity of other Shark Bay biota.