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Background and AimsHybridization is an important evolutionary process that can have a significant impact on natural plant populations. Eucalyptus species are well known for weak reproductive barriers and extensive hybridization within subgenera, but there is little knowledge of whether patterns of hybridization differ among subgenera. Here, we examine eucalypts of Western Australia’s Stirling Range to investigate how patterns of hybridization are associated with landscape and taxon age between the two largest Eucalyptus subgenera: Eucalyptus and Symphyomyrtus. In doing so, we tested a hypothesis of OCBIL (old, climatically buffered, infertile landscape) theory that predicts reduced hybridization on older landscapes.MethodsSingle nucleotide polymorphism markers were applied to confirm the hybrid status, parentage and genetic structure of five suspected hybrid combinations for subg. Eucalyptus and three combinations for subg. Symphyomyrtus.Key ResultsEvidence of hybridization was found in all combinations, and parental taxa were identified for most combinations. The older parental taxa assessed within subg. Eucalyptus, which are widespread on old landscapes, were identified as well-defined genetic entities and all hybrids were exclusively F1 hybrids. In addition, many combinations showed evidence of clonality, suggesting that the large number of hybrids recorded from some combinations is the result of long-term clonal spread following a few hybridization events rather than frequent hybridization. In contrast, the species in subg. Symphyomyrtus, which typically occur on younger landscapes and are more recently evolved, showed less distinction among parental taxa, and where hybridization was detected, there were high levels of introgression.ConclusionsReduced hybridization in subg. Eucalyptus relative to extensive hybridization in subg. Symphyomyrtus affirmed the hypothesis of reduced hybridization on OCBILs and demonstrate that clade divergence times, landscape age and clonality are important drivers of differing patterns of speciation and hybridization in Eucalyptus.

Carnivorous plants allocate more resources to carnivorous structures under nutrient-limited conditions, and relative investment can also be influenced by animals (infauna) that live in association with these plants and feed on their prey. We investigated these effects within a population of the pitcher plant Cephalotus follicularis containing varying densities of larvae of the fly Badisis ambulans . For plants with a relatively high proportion of adult pitchers, increasing larval density was associated with lower relative leaf allocation to new pitcher buds. For plants with relatively few adult pitchers, however, there was greater relative leaf allocation to pitcher buds with increasing larval density. In a field experiment, there was no significant effect of experimental larval presence or absence on the change in carbon-to-nitrogen (C/N) ratio of plants. Although the direction of the correlation between B. ambulans larvae and relative investment in carnivorous and non-carnivorous structures depends on the relative number of mature structures, whether the larvae enhance or reduce nutrient stress under different conditions remains unclear. The change in C/N was, however, less variable for pitchers that contained larvae, suggesting a stabilizing effect. Eighteen of 52 experimental pitchers were damaged by an unknown species, causing the pitcher fluid to drain. These pitchers were significantly more likely to survive if they contained larvae. These results suggest that the relationship between infauna and host varies with the initial resource status and environmental context of the host plant.

Aim Granite outcrops may be able to act as refugia for species during adverse climate change, owing to their topographic complexity. We assessed this hypothesis by examining phylogeographical patterns in a common, geographically widespread granite endemic, Stypandra glauca (Hemerocallidaceae). Location Granite outcrops of the Southwest Australian Floristic Region, Western Australia. Main conclusions The levels of divergence and low diversity revealed in S. glauca support a scenario of prolonged isolation and persistence on granite outcrops may result from the maintenance of a larger effective population size in southern regions, which retained more mesic climates during drier glacial periods. Although the climatic conditions differ between outcrops in this study, our results indicate that outcrops in both regions have harboured S. glauca throughout climatic changes, accentuating the value of these habitats to biodiversity conservation under future changing climate. Methods Twenty-four tetraploid individuals of the granite endemic Stypandraglauca were sampled from each of 12 granite outcrops: 7 from a mesic environment and 5 from the semi-arid region. Phylogenetic reconstruction and divergence-dating was achieved using Bayesian and parsimony analyses of chloroplast haplotypes from 90 individuals. Nuclear diversity and population differentiation were analysed across all individuals using 10 microsatellite loci. Results Stypandra glauca exhibited high (chloroplast) or moderate (nuclear)levels of divergence among, and low diversity within, outcrops. Haplotype diversity was high in both sampling regions, and each haplotype was unique to one outcrop. There was little correlation between geographical and genetic distance.Both nuclear and chloroplast diversity were higher in southern (mesic)outcrops than in northern (semi-arid) outcrops, although the level of chloroplast divergence among outcrops was similar for both climatic regions. Refereed/Peer-reviewed

Identification of refugia is an increasingly important adaptation strategy in conservation planning under rapid anthropogenic climate change. Granite outcrops (GOs) provide extraordinary diversity, including a wide range of taxa, vegetation types and habitats in the Southwest Australian Floristic Region (SWAFR). However, poor characterization of GOs limits the capacity of conservation planning for refugia under climate change. A novel means for the rapid identification of potential refugia is presented, based on the assessment of local-scale environment and vegetation structure in a wider region. This approach was tested on GOs across the SWAFR. Airborne discrete return Light Detection And Ranging (LiDAR) data and Red Green and Blue (RGB) imagery were acquired. Vertical vegetation profiles were used to derive 54 structural classes. Structural vegetation types were described in three areas for supervised classification of a further 13 GOs across the region. Habitat descriptions based on 494 vegetation plots on and around these GOs were used to quantify relationships between environmental variables, ground cover and canopy height. The vegetation surrounding GOs is strongly related to structural vegetation types (Kappa = 0.8) and to its spatial context. Water gaining sites around GOs are characterized by taller and denser vegetation in all areas. The strong relationship between rainfall, soil-depth, and vegetation structure (R2 of 0.8-0.9) allowed comparisons of vegetation structure between current and future climate. Significant shifts in vegetation structural types were predicted and mapped for future climates. Water gaining areas below granite outcrops were identified as important putative refugia. A reduction in rainfall may be offset by the occurrence of deeper soil elsewhere on the outcrop. However, climate change interactions with fire and water table declines may render our conclusions conservative. The LiDAR-based mapping approach presented enables the integration of site-based biotic assessment with structural vegetation types for the rapid delineation and prioritization of key refugia. Refereed/Peer-reviewed

Aim Granite outcrops are prominent throughout the world and harbour many endemic species. Their topographic complexity and range of environments have led to the hypothesis that they act as refugia facilitating the persistence of species through climate change. We evaluate this hypothesis by investigating the phylogeographic patterns in a common granite endemic shrub. Location Granite outcrops of the Southwest Australian Floristic Region. Methods Chloroplast haplotypes of 89 Kunzea pulchella individuals from 16 granite outcrops were determined from sequences of three chloroplast intergenic spacer regions. Phylogenetic reconstruction and divergence dating was inferred using Bayesian and Parsimony analyses and phylogenetic relationships between haplotypes were examined in relation to geographic distributions. Nuclear diversity and differentiation of populations were assessed through analysis of 11 nuclear microsatellite loci across 384 individuals from the 16 granite outcrops. Results Kunzea pulchella exhibited low haplotype and allelic diversity within outcrops and high levels of divergence among outcrops, indicating an ancient restriction to specific outcrops with genetic drift as the main driver of evolution. Two divergent lineages were revealed in the chloroplast phylogeny dating to the Pliocene and potentially reflecting the initial impact of increased aridity prior to isolation on individual outcrops. Main conclusions Rather than uncovering the typical pattern for Pleistocene refugia with contraction to, and expansion from particular granite outcrops, we observed persistence, prolonged isolation and divergence of populations. We suggest the persistence of K. pulchella on multiple outcrops through a period of considerable climatic change may be a result of broad climatic tolerances or contraction and expansion dynamics operating at microrefugial scales within outcrops. Our observations of low haplotype and allelic diversity within populations of K. pulchella provide some support for the latter. The enduring nature of K. pulchella and evolutionary potential of populations on individual outcrops accentuates the value of these environments for biodiversity conservation planning in a changing climate.

BACKGROUND AND AIMS Worldwide, many plant species are confined to open, shallow-soil, rocky habitats. Although several hypotheses have been proposed to explain this habitat specificity, none has been convincing. We suggest that the high level of endemism on shallow soils is related to the edaphic specialization needed to survive in these often extremely drought-prone habitats. Previous research has shown that species endemic to ironstone communities in SW Australia have a specialized root morphology that enhances their chance to access fissures in the underlying rock. Here we test the generality of these findings for species that are confined to a shallow-soil habitat that is of much greater global significance: granite outcrops. METHODS We compared temporal and spatial root growth and allocation of three endemic woody perennials of SW Australian granite outcrop communities with those of congeners occurring on nearby deeper soils. Seedlings of all species were grown in 1·2 m long custom-made containers with a transparent bottom that allowed monitoring of root growth over time. KEY RESULTS The granite outcrop endemics mostly differed in a predictable way from their congeners from deeper soils. They generally invested a larger portion of their biomass in roots, distributed their roots faster and more evenly over the container and had a lower specific root length. In different species pairs the outcrop endemics achieved their apparent advantage by a different combination of the aforementioned traits. CONCLUSIONS Our results are consistent with earlier work, indicating that species restricted to different types of drought-prone shallow-soil communities have undergone similar selection pressures. Although adaptive in their own habitat in terms of obtaining access to fissures in the underlying rock, these root system traits are likely to be maladaptive in deeper soil habitats. Therefore, our results may provide an explanation for the narrow endemism of many shallow-soil endemics.

Recent phylogenetic analysis has placed the aquatic family Hydatellaceae as an early-divergent angiosperm. Understanding seed dormancy, germination and desiccation tolerance of Hydatellaceae will facilitate ex situ conservation and advance hypotheses regarding angiosperm evolution.Seed germination experiments were completed on three species of south-west Australian Hydatellaceae, Trithuria austinensis, T. bibracteata and T. submersa, to test the effects of temperature, light, germination stimulant and storage. Seeds were sectioned to examine embryo growth during germination in T. austinensis and T. submersa.Some embryo growth and cell division in T. austinensis and T. submersa occurred prior to the emergence of an undifferentiated embryo from the seed coat ('germination'). Embryo differentiation occurred later, following further growth and a 3- to 4-fold increase in the number of cells. The time taken to achieve 50 % of maximum germination for seeds on water agar was 50, 35 and 37 d for T. austinensis, T bibracteata and T. submersa, respectively.Seeds of Hydatellaceae have a new kind of specialized morphophysiological dormancy in which neither root nor shoot differentiates until after the embryo emerges from the seed coat. Seed biology is discussed in relation to early angiosperm evolution, together with ex situ conservation of this phylogenetically significant group.