Southwest Australian Biodiversity Hotspot Research Articles

A cool spot in a biodiversity hotspot: why do tall Eucalyptus forests in Southwest Australia exhibit low diversity?

Background and aimsSouthwest Australia is a biodiversity hotspot, with greatest plant species diversity on the most severely phosphorus (P)-impoverished soils. Here, non-mycorrhizal species with highly-effective carboxylate-releasing P-acquisition strategies coexist with mycorrhizal species that are less effective at accessing P on these soils. Non-mycorrhizal carboxylate-releasing species facilitate P acquisition of mycorrhizal neighbours that are better defended against pathogens. In the Southwest Australian Biodiversity Hotspot, there are also ‘cool spots’ of low-diversity tall mycorrhizal Eucalyptus communities on P-impoverished soils. These Eucalyptus trees obviously do not require facilitation of their P acquisition by carboxylate-releasing neighbours, because these are only a minor component of the low-diversity communities. We hypothesised that in low-diversity tall Eucalyptus forests, mycorrhizal species release carboxylates to acquire P. Thus, they would not depend on facilitation, and must be strong competitors. However, because they would not depend on external mycorrhizal hyphae to acquire P, they would also not be able to access soil organic nitrogen (N), for which they would need external hyphae.MethodsSince carboxylates not only mobilise P, but also manganese (Mn), we used leaf Mn concentrations ([Mn]) in the natural habitat to proxy rhizosphere carboxylates. To verify this proxy, we also measured carboxylate exudation of targeted species with high leaf [Mn] using seedlings grown in low-P nutrient solutions.ResultsUsing these complementary approaches, we confirmed our hypothesis that dominant Eucalyptus species in ‘cool spots’ release carboxylates. Since mineralisation of organic N is associated with fractionation of N, enriching organic N with 15N while nitrate is depleted in 15N, we measured the stable N isotope composition of leaf material. The results show that dominant Eucalyptus species did not access organic N, despite being ectomycorrhizal.ConclusionsThe low diversity of tall Eucalyptus forests in southwest Australia can be explained by dominant mycorrhizal species exhibiting a carboxylate-releasing strategy. The tall eucalypts are therefore strong competitors that do not require facilitation, but also do not access organic N.

Urban native vegetation remnants support more diverse native bee communities than residential gardens in Australia's southwest biodiversity hotspot

Native bees are declining in many regions, often associated with loss of natural habitat. Urbanisation replaces natural vegetation with a highly-modified landscape, where residential gardens are a major component of urban greenspace. While many cities retain native vegetation remnants within the urban matrix, these are often small, isolated and degraded. However, there is little empirical evidence on the capacity of residential gardens to provide equivalent or beneficial habitat for native bees, and which local and landscape factors influence bee assemblages. We surveyed bee assemblages in the southwest Australian biodiversity hotspot at seven residential gardens and seven bushland remnants over two years. We recorded 153 species/morphospecies of native bees. Native bees were more abundant in bushland remnants than residential gardens. Abundance of the introduced honeybee Apis mellifera was generally high, and did not differ between habitats. Bushland remnants hosted more species, and rare and unique species, than did residential gardens. Native bee body-size and nesting guilds varied in their response to habitat type. Native bee abundance and richness increased with abundance of native plant species, but decreased with total flower species richness. Native bee species richness was negatively impacted by urbanisation (built space and isolation from bushland reserves). There were no significant relationships between honeybee abundance and local and landscape factors. Our study demonstrates that while residential gardens can host native bees, urban bushland remnants harbour a more comprehensive suite of species and are key for the conservation of native bee populations.

A bee's eye view of remarkable floral colour patterns in the south-west Australian biodiversity hotspot revealed by false colour photography.

Colour pattern is a key cue of bee attraction selectively driving the appeal of pollinators. It comprises the main colour of the flower with extra fine patterns, indicating a reward focal point such as nectar, nectaries, pollen, stamens and floral guides. Such advertising of floral traits guides visitation by the insects, ensuring precision in pollen gathering and deposition. The study, focused in the Southwest Australian Floristic Region, aimed to spot bee colour patterns that are usual and unusual, missing, accomplished by mimicry of pollen and anthers, and overlapping between mimic-model species in floral mimicry cases. Floral colour patterns were examined by false colour photography in 55 flower species of multiple highly diverse natural plant communities in south-west Australia. False colour photography is a method to transform a UV photograph and a colour photograph into a false colour photograph based on the trichromatic vision of bees. This method is particularly effective for rapid screening of large numbers of flowers for the presence of fine-scale bee-sensitive structures and surface roughness that are not detectable using standard spectrophotometry. Bee- and bird-pollinated flowers showed the expected but also some remarkable and unusual previously undetected floral colour pattern syndromes. Typical colour patterns include cases of pollen and flower mimicry and UV-absorbing targets. Among the atypical floral colour patterns are unusual white and UV-reflecting flowers of bee-pollinated plants, bicoloured floral guides, consistently occurring in Fabaceae spp., and flowers displaying a selective attractiveness to birds only. In the orchid genera (Diuris and Thelymitra) that employ floral mimicry of model species, we revealed a surprising mimicry phenomenon of anthers mimicked in turn by model species. The study demonstrates the applicability of 'bee view' colour imaging for deciphering pollinator cues in a biodiverse flora with potential to be applied to other eco regions. The technique provides an exciting opportunity for indexing floral traits on a biome scale to establish pollination drivers of ecological and evolutionary relevance.

Ecological interactions shape the evolution of flower color in communities across a temperate biodiversity hotspot.

Processes driving the divergence of floral traits may be integral to the extraordinary richness of flowering plants and the assembly of diverse plant communities. Several models of pollinator‐mediated floral evolution have been proposed; floral divergence may (i) be directly involved in driving speciation or may occur after speciation driven by (ii) drift or local adaptation in allopatry or (iii) negative interactions between species in sympatry. Here, we generate predictions for patterns of trait divergence and community assembly expected under these three models, and test these predictions in Hakea (Proteaceae), a diverse genus in the Southwest Australian biodiversity hotspot. We quantified functional richness for two key floral traits (pistil length and flower color), as well as phylogenetic distances between species, across ecological communities, and compared these to patterns generated from null models of community assembly. We also estimated the statistical relationship between rates of trait evolution and lineage diversification across the phylogeny. Patterns of community assembly suggest that flower color, but not floral phenology or morphology, or phylogenetic relatedness, is more divergent in communities than expected. Rates of lineage diversification and flower color evolution were negatively correlated across the phylogeny and rates of flower colour evolution were positively related to branching times. These results support a role for diversity‐dependent species interactions driving floral divergence during the Hakea radiation, contributing to the development of the extraordinary species richness of southwest Australia.

Interactions between the introduced European honey bee and native bees in urban areas varies by year, habitat type and native bee guild

AbstractEuropean honey bees have been introduced across the globe and may compete with native bees for floral resources. Compounding effects of urbanization and introduced species on native bees are, however, unclear. Here, we investigated how honey bee abundance and foraging patterns related to those of native bee abundance and diversity in residential gardens and native vegetation remnants for 2 years in urbanized areas of the Southwest Australian biodiversity hotspot and assessed how niche overlap influenced these relationships. Honey bees did not overtly suppress native bee abundance; however, complex relationships emerged when analysing these relationships according to body size, time of day and floral resource levels. Native bee richness was positively correlated with overall honeybee abundance in the first year, but negatively correlated in the second year, and varied with body size. Native bees that had higher resource overlap with honey bees were negatively associated with honey bee abundance, and resource overlap between honey bees and native bees was higher in residential gardens. Relationships with honey bees varied between native bee taxa, reflecting adaptations to different flora, plus specialization. Thus, competition with introduced bees varies by species and location, mediated by dietary breadth and overlap and by other life-history traits of individual bee species.

Plant-pollinator networks in Australian urban bushland remnants are not structurally equivalent to those in residential gardens

Urbanisation is a prominent and increasing form of land-use change, with the potential to disrupt the interactions between pollinators such as bees and the flowering plants that they visit. This in turn may cause cascading local extinctions and have consequences for pollination services. Network approaches go beyond simple metrics of abundance and species richness, enabling understanding of how the structure of plant-pollinator communities are affected by urbanisation. Here we compared pollination networks between native vegetation (bushland) remnants and residential gardens in the urbanised region of the southwest Australian biodiversity hotspot. Across fourteen sites, seven per habitat, plant-bee visitor networks were created from surveys conducted monthly during the spring-summer period over two years. Extinction slope (a measure of how extinctions cascade through the network), and network robustness and nestedness were higher for bushland remnants, suggesting that networks in bushland remnants had greater functional integrity, but if disrupted, more cascading extinctions could occur. In contrast, niche overlap between pollinators was higher in residential gardens, suggesting greater competition for resources. Most species-level properties did not differ between habitats, except for normalised degree, which was higher in bushland remnants. In conclusion, it appears that pollination networks in managed residential gardens are not structurally equivalent with those in bushland remnants. This has implications for conservation of wild bee assemblages in this biodiversity hotspot, and suggests removal of remnant native vegetation for residential development could disrupt the integrity of plant-pollinator assemblages.

Xylomelum occidentale (Proteaceae) accesses relatively mobile soil organic phosphorus without releasing carboxylates

Abstract Hundreds of Proteaceae species in Australia and South Africa typically grow on phosphorus (P)‐impoverished soils, exhibiting a carboxylate‐releasing P‐mobilizing strategy. In the Southwest Australian Biodiversity Hotspot, two Xylomelum (Proteaceae) species are widely distributed, but restricted within that distribution. We grew Xylomelum occidentale in hydroponics at 1 μM P. Leaves, seeds, rhizosheath and bulk soil were collected in natural habitats. Xylomelum occidentale did not produce functional cluster roots and occupied soils that are somewhat less P‐impoverished than those in typical Proteaceae habitats in the region. Based on measurements of foliar manganese concentrations (a proxy for rhizosphere carboxylate concentrations) and P fractions in bulk and rhizosheath soil, we conclude that X. occidentale accesses organic P, without releasing carboxylates. Solution 31P‐NMR spectroscopy revealed which organic P forms X. occidentale accessed. Xylomelum occidentale uses a strategy that differs fundamentally from that typical in Proteaceae, accessing soil organic P without carboxylates. We surmise that this novel strategy is likely expressed also in co‐occurring non‐Proteaceae that lack a carboxylate‐exuding strategy. These co‐occurring species are unlikely to benefit from mycorrhizal associations, because plant‐available soil P concentrations are too low. Synthesis. Our findings show the first field evidence of effectively utilizing soil organic P by X. occidentale without carboxylate exudation and explain their relatively restricted distribution in an old P‐impoverished landscape, contributing to a better understanding of how diverse P‐acquisition strategies coexist in a megadiverse ecosystem.

Pronounced differences in visitation by potential pollinators to co-occurring species of Fabaceae in the Southwest Australian biodiversity hotspot

AbstractDespite their diversity and the potential for specialized pollination systems, Australian Fabaceae have received little attention in pollination studies. In the Southwest Australian Floristic Region (SWAFR), a recognized biodiversity hotspot, co-occurring and abundant species of Faboideae exhibit a range of floral colours and forms, suggestive of adaptation to different groups of pollinators. For four communities of Fabaceae in the SWAFR we investigated whether co-occurring species overlap in pollinator genera, whether these pollinators show differences in behaviour on the pea flower and whether variations in stamen length and nectar composition among species are associated with different pollinator types. Species of Fabaceae were visited by one to four genera of native bees, suggesting varying levels of ecological specialisation. In Fabaceae with more specialized interactions, co-occurring species showed marked differences in the bee genera attracted. Unexpectedly, some Fabaceae frequently attracted beetles, which may play an important role in their pollination. There was no evidence for an association between stamen length or nectar composition and the type of pollinator. The introduced honeybee, visited all studied species of Fabaceae, suggesting that they may act both as a pollinator and a potential competitor with native pollinators.

The relative performance of sampling methods for native bees: an empirical test and review of the literature

AbstractMany bee species are declining globally, but to detect trends and monitor bee assemblages, robust sampling methods are required. Numerous sampling methods are used, but a critical review of their relative effectiveness is lacking. Moreover, evidence suggests the relative effectiveness of sampling methods depends on habitat, yet efficacy in urban areas has yet to be evaluated. This study compared the bee community documented using observational records, targeted netting, mobile gardens, pan traps (blue and yellow), vane traps (blue and yellow), and trap‐nests. The comparative surveys of native bees and honeybees were undertaken in an urbanized region of the southwest Australian biodiversity hot spot. The outcomes of the study were then compared to a synthesis based on a comprehensive literature review of studies where two or more bee sampling methods were conducted. Observational records far exceeded all other methods in terms of abundance of bees recorded, but were unable to distinguish finer taxonomic levels. Of methods that captured individuals, thereby permitting taxonomic identification, targeted sweep netting vastly outperformed the passive sampling methods, yielding a total of 1324 individuals, representing 131 taxonomic units—even when deployed over a shorter duration. The relative effectiveness of each method differed according to taxon. From the analysis of the literature, there was high variability in relative effectiveness of methods, but targeted sweep netting and blue vane traps tended to be most effective, in accordance with results from this study. However, results from the present study differed from most previous studies in the extremely low catch rates in pan traps. Species using trap‐nests represented only a subset of all potential cavity‐nesters, and their relative abundances in the trap‐nests differed from those in the field. Mobile gardens were relatively ineffective at attracting bees. For urbanized habitat within this biodiversity hot spot, targeted sweep netting is indispensable for obtaining a comprehensive indication of native bee assemblages; passive sampling methods alone recorded only a small fraction of the native bee community. Overall, a combination of methods should be used for sampling bee communities, as each has their own biases, and certain taxa were well represented in some methods, but poorly represented in others.

Extinction pulse at Eocene-Oligocene boundary drives diversification dynamics of two Australian temperate floras.

The diversification dynamics of the Australian temperate flora remains poorly understood. Here, we investigate whether differences in plant richness in the southwest Australian (SWA) biodiversity hotspot and southeast Australian (SEA) regions of the Australian continent can be attributed to higher net diversification, more time for species accumulation, or both. We assembled dated molecular phylogenies for the 21 most species-rich flowering plant families found across mesic temperate Australia, encompassing both SWA and SEA regions, and applied a series of diversification models to investigate responses across different groups and timescales. We show that the high richness in SWA can be attributed to a higher net rate of lineage diversification and more time for species accumulation. Different pulses of diversification were retrieved in each region. A decrease in diversification rate across major flowering plant lineages at the Eocene-Oligocene boundary (ca 34 Ma) was witnessed in SEA but not in SWA. Our study demonstrates the importance of historical diversification pulses and differential responses to global events as drivers of present-day diversity. More broadly, we show that diversity within the SWA biodiversity hotspot is not only the result of recent radiations, but also reflects older events over the history of this planet.

Floral display and habitat fragmentation: Effects on the reproductive success of the threatened mass-flowering Conospermum undulatum (Proteaceae).

Fragmentation of natural vegetation is currently one of the largest threats to plant populations and their interactions with pollinators. Plant reproductive susceptibility to habitat fragmentation has been investigated in many species; however, the response of wild mass‐flowering species is poorly known, with research limited to mainly boreal plant species.Here, we studied twelve remnant populations of the threatened mass‐flowering shrub Conospermum undulatum in the southwest Australian biodiversity hotspot, each presenting different population size, level of isolation, and floral display. We assessed the impact of fragmentation on (a) fruit and seed production; and (b) seed germination. To gain a deeper understanding of factors influencing the reproductive success of C. undulatum, we performed pollinator exclusion and self‐pollination treatments to experimentally assess the mating system of this threatened shrub.We found C. undulatum to be strictly self‐incompatible and totally reliant on pollinators visiting with an outcrossed pollen load to complete the reproductive cycle. Further, we found that fruit production dropped from 35% to <20% as a result of decreasing floral display. A reduction in population size from 880 to 5 plants and from ~700 to 0.21 in the floral display index led to a decrease in seed output, while a similar reduction in seed output, from 6% to 3%, was observed as a result of increasing isolation index from −21.41 to −0.04. Overall, seed germination was positively related to population size, and a negative relationship was found between germination and isolation. Synthesis and applications. Our results demonstrate the important relationship between pollinators and floral morphology in plants of southwest Australia that have coevolved with native pollinators and developed characteristic flower morphologies over long time frames. Indeed, due to its characteristic pollination mechanism, the self‐incompatible C. undulatum can only rely on specialized native pollinators for pollen flow and cannot rely on its mass‐flowering trait to attract generalist pollinators from coflowering species; neither can it compensate for the lack of visitors by promoting geitonogamy. Consequently, fragmentation has a significant effect on the reproductive output of C. undulatum, and size, isolation, and floral display of populations are important factors to be considered when planning conservation actions for the species.

Seed Treatment Optimizes Benefits of Seed Bank Storage for Restoration‐Ready Seeds: The Feasibility of Prestorage Dormancy Alleviation for Mine‐Site Revegetation

AbstractDormant seeds of 18 species from 9 families covering a diverse range of seed dormancy syndromes and life histories from the southwest Australian biodiversity hotspot were assessed for germinability following storage at 15–25°C for 36 months. A total of 10 species with physical dormancy (PY) and 8 with either physiological dormancy (PD) or morphophysiological dormancy (MPD) were assessed as part of the study. Prior to storage, germination from dormant seeds was 1–27%, rising to 41–100% following specific dormancy‐breaking treatments. When seed dormancy was removed prior to storage for 36 months seeds from all species were found to maintain a nondormant state and germinate to a similar level to that observed at the beginning of the experiment (44–100%). Likewise, seeds that did not receive a prestorage dormancy‐breaking treatment maintained a dormant state (0–50% germination) and subsequently responded well to a dormancy‐breaking treatment immediately prior to germination assessment (49–99%). There were minimal differences in response to dormancy‐breaking treatments before and after 36 months storage (average 4–6% difference) and in the germination responses observed between both storage environments assessed (15°C/15% eRH or 15–25°C air dried). Based on these findings, storing seeds in a nondormant state does not alter germinability and this approach provides significant benefits to current seed‐based restoration programs through reduction of double handling and improved seed use efficiency.

Plant diversity is linked to nutrient limitation of dominant species in a world biodiversity hotspot

AbstractQuestionsWhat is the role of nutrient limitation of dominant species in maintaining plant species diversity at small spatial scales, particularly in biodiversity hotspots?LocationSouthwest of Western Australia.MethodsThe extent of nutrient limitation was determined by measuring variation in the foliar nutrients of dominant plant species and total above‐ground biomass in relation to soil N and P across 16 plots. Plant species richness and evenness for the same plots were then regressed against foliar N:P and δ15N of the dominant species.ResultsFoliar N and P content as well as above‐ground biomass increased, while foliar N:P decreased with increasing soil N and P, suggesting that dominant species are limited by both N and P in southwest Australia. In contrast, foliar δ15N values only increased with decreasing soil P, indicating that δ15N enrichment reflects low P availability. Species richness increased at sites where foliar N:P was higher and δ15N more enriched, suggesting that a relatively greater number of rare plant species co‐exist where the growth of dominant vegetation is more nutrient‐limited.ConclusionsThe mechanisms that allow an exceptionally high number of plant species to co‐exist at small spatial scales in a southwest Australian biodiversity hotspot are linked to the extent to which the dominant plant species are limited in N and P. In addition, on nutrient‐impoverished soils, a foliar N:P ratio &gt;16 can reflect N as well as P limitation. While foliar δ15N also reflects availability of soil P, it is not necessarily a reliable indicator of nutrient limitation in southwest Australia, and perhaps in other environments where plants are highly adapted to low‐nutrient conditions.

Multi-scale patterns in the host specificity of plant-dwelling arthropods: the influence of host plant and temporal variation on species richness and assemblage composition of true bugs (Hemiptera)

The influence of temporal variation in the host specificity of invertebrates to estimates of biodiversity is rarely considered. While patterns at large spatial scales have stimulated much attention, such comparisons are constrained for southern-hemisphere biomes because the patterning of invertebrates on plants is largely unknown. Here, we analyse variation of plant-dwelling Hemiptera from 15 understorey plant species over 18 months in the south-west Australian biodiversity hotspot. Analyses showed significant interactions in species composition between sampling period and plant species. Fauna that were “effectively specialized” (host-specificity index) to plants changed with season, although this was also related to the number of singletons and overall species richness. Sampling from a single season also overestimated the degree of host specificity by 52% and underestimated the perception of species richness when an outbreak of a particular herbivore occurred. High host-specificity values (12.7 hemipteran species per plant) support the case for high estimates of global arthropod richness.

Comparative longevity of Australian orchid (Orchidaceae) seeds under experimental and low temperature storage conditions

Seeds from ten terrestrial orchid species, nine from the south-west Australian biodiversity hotspot (Caladenia arenicola, Caladenia flava, Caladenia huegelii, Diuris laxiflora, Microtis media ssp. media, Pterostylis recurva, Pterostylis sanguinea, Thelymitra crinita and Thelymitra macrophylla) and one from south-east Australia (Diuris fragrantissima), were placed into experimental storage to assess their relative longevity and likely optimal conditions for long-term conservation seed banking. Seeds from all species were desiccation tolerant, germinating after drying at 23% relative humidity (C. arenicola, C. huegelii, P. sanguinea and T. macrophylla) or 5% relative humidity (C. flava, D. laxiflora, M. media ssp. media, P. recurva and T. crinita) at 23 °C. From automatedly determined moisture adsorption and desorption isotherms at 23 °C, these equate to tolerance of drying to 0.03–0.06 g water g−1 dry weight or 0.013–0.028 g water g−1 dry weight, respectively. Results of storage experiments at a range of moisture contents and temperatures suggest conventional seed bank storage at −18 °C after equilibration at c. 23% relative humidity (at 23 °C) may be suitable for most of the species, although there was higher germination of P. recurva seeds stored at −80 °C and of M. media ssp. media seeds equilibrated at 75% relative humidity. However, there was considerable variation in germination of seeds sampled after different storage periods, making it difficult to identify optimal storage conditions definitively. Results of comparative longevity storage experiments at 60% relative humidity and 40 °C suggest seeds from these orchid species are short-lived compared with non-orchid species, and with Australian species in particular. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164, 26–41.

Pollination ecology and the possible impacts of environmental change in the Southwest Australian Biodiversity Hotspot.

The Southwest Australian Biodiversity Hotspot contains an exceptionally diverse flora on an ancient, low-relief but edaphically diverse landscape. Since European colonization, the primary threat to the flora has been habitat clearance, though climate change is an impending threat. Here, we review (i) the ecology of nectarivores and biotic pollination systems in the region, (ii) the evidence that trends in pollination strategies are a consequence of characteristics of the landscape, and (iii) based on these discussions, provide predictions to be tested on the impacts of environmental change on pollination systems. The flora of southwestern Australia has an exceptionally high level of vertebrate pollination, providing the advantage of highly mobile, generalist pollinators. Nectarivorous invertebrates are primarily generalist foragers, though an increasing number of colletid bees are being recognized as being specialized at the level of plant family or more rarely genus. While generalist pollination strategies dominate among insect-pollinated plants, there are some cases of extreme specialization, most notably the multiple evolutions of sexual deception in the Orchidaceae. Preliminary data suggest that bird pollination confers an advantage of greater pollen movement and may represent a mechanism for minimizing inbreeding in naturally fragmented populations. The effects of future environmental change are predicted to result from a combination of the resilience of pollination guilds and changes in their foraging and dispersal behaviour.