Banksia Species Research Articles

Life at the conservative end of the leaf economics spectrum: intergeneric variation in the allocation of phosphorus to biochemical fractions in species of Banksia (Proteaceae) and Hakea (Proteaceae).

In severely phosphorus (P)-impoverished environments, plants have evolved to use P very efficiently. Yet, it is unclear how P allocation in leaves contributes to their photosynthetic P-use efficiency (PPUE) and position along the leaf economics spectrum (LES). We address this question in 10 species of Banksia and Hakea, two highly P-efficient Proteaceae genera. We characterised traits in leaves of Banksia and Hakea associated with the LES: leaf mass per area, light-saturated photosynthetic rates, P and nitrogen concentrations, and PPUE. We also determined leaf P partitioning to five biochemical fractions (lipid, nucleic acid, metabolite, inorganic and residual P) and their possible association with the LES. For both genera, PPUE was negatively correlated with fractional allocation of P to lipids, but positively correlated with that to metabolites. For Banksia only, PPUE was negatively correlated with residual P, highlighting a strategy contrasting to that of Hakea. Phosphorus-allocation patterns significantly explained PPUE but were not linked to the resource acquisition vs resource conservation gradient defined by the LES. We conclude that distinct P-allocation patterns enable species from different genera to achieve high PPUE and discuss the implications of different P investments. We surmise that different LES axes representing different ecological strategies coexist in extremely P-impoverished environments.

Pollination by birds, non-flying mammals, and European honeybees in a heathland shrub, Banksia catoglypta (Proteaceae)

Abstract Vertebrate pollination has evolved in many plant lineages, with pollination by birds the most prevalent strategy. However, in Banksia (Proteaceae) many species are visited by a combination of nectarivorous birds, non-flying mammals (NFMs), and insects, though the relative importance of these pollen vectors for reproduction remains poorly understood. Here, we investigate the contribution to fruit set of birds, NFMs, and insects in Banksia catoglypta—a heathland shrub from south-west Australia. This species flowers in winter and produces large, golden-yellow inflorescences that are typically presented near the ground and consist of rigid, strongly scented flowers with a long stigma-nectary distance—suggesting pollination by vertebrates. The primary floral visitors were white-cheeked honeyeaters (Phylidonyris niger), honey possums (Tarsipes rostratus), house mice (Mus musculus), and introduced European honeybees (Apis mellifera), all of which carried B. catoglypta pollen. Banksia catoglypta was found to be obligately outcrossing with no evidence of pollen limitation. Selective exclusion of flying pollinators revealed that inflorescences pollinated solely by NFMs produced 50% fewer fruit than open-pollinated controls. Therefore, while mammals are pollinators of B. catoglypta, they may contribute comparatively little to fruit set in some Banksia species displaying floral traits considered adaptations to NFM pollination. Exclusion of vertebrate pollinators had no effect on fruit set, demonstrating that European honeybees can be effective pollinators of some primarily vertebrate-pollinated plants.

The culturable seed mycobiome of two Banksia species is dominated by latent saprotrophic and multi-trophic fungi

Seed fungal endophytes play an important beneficial role in the formation of the seedling mycobiome and contribute to plant establishment, but can also occur as latent pathogens and saprotrophs. Current knowledge on the function and diversity of seed fungal endophytes has been gained through studies in agricultural systems whilst knowledge from natural systems is relatively less. We used two co-occurring species from the genus Banksia from four sites in Australia's Sydney Basin Bioregion to investigate the abundance and diversity of seed fungal endophyte communities present in natural ecosystem hosts. Based on results from culturing and DNA sequence analysis of multiple loci, we found that Banksia seeds house a diverse range of fungal endophyte species, that when assigned to functional guilds belonged to multiple trophic modes. Thirty-one of the fungal taxa identified had not been previously reported as endophytes. Amongst the 58 Operational Taxonomic Units identified, Leotiomycetes and Sordariomycetes were the dominant classes and Banksiamyces (Leotiomycetes) and Penicillium (Sordariomycetes) the dominant genera, with many of the species isolated recorded in the literature as having a limited distribution. The two Banksias shared few fungal endophyte species, which were not always present across all study sites. We revealed a ‘hidden diversity’ within seeds of Banksia from natural ecosystems and provided insights into the influence host species can have on the seed mycobiome.

Relationship of alien species continues in a foreign land: Thecase of Phytophthora and Australian Banksia (Proteaceae) in South African Fynbos.

Fungal invasions only recently started to receive more attention in invasion biology. This is largely attributed to little or non‐existent information about these inconspicuous organisms. Most invasion hypotheses focus on factors that increase invasion success; few try to explain why invasions fail. Here we hypothesize that a host–pathogen relationships can limit the invasiveness of an alien plant species in a novel range. To test this, we investigate whether the invasiveness of the Australian genus of Proteaceae, Banksia, in South Africa is determined by the alien and major invasive phytopathogen, Phytophthora cinnamomi. The presence of P. cinnamomi in Banksia root and soil was evaluated using morphological and molecular techniques. Isolates were cultured onto selective media and polymerize chain reactions and internal transcribing spacers were used for identification. Acetone leaf extracts of 11 Banksia spp. were screened for antimicrobial activity against P. cinnamomi, using the minimum inhibitory concentration assay. A total of 3840 Banksia individuals from seven localities were surveyed. Phytophthora cinnamomi was consistently isolated from Banksia species root and soil samples. Out of the 12 Banksia species that were screened for antimicrobial activity, four introduced species, B. burdettii, B. coccinea, Banksia hookeriana, and B. prionotes and the invasive B. integrifolia and B. ericifolia exhibited relatively high antimicrobial activity against P. cinnamomi (strain 696/12). We show that the phytopathogen in the native range has similar impact in the novel range and in doing so may limit invasion success of Banksia species with low antimicrobial activity.

Phosphorus toxicity, not deficiency, explains the calcifuge habit of phosphorus-efficient Proteaceae.

The calcifuge habit of plants is commonly explained in terms of high soil pH and its effects on nutrient availability, particularly that of phosphorus (P). However, most Proteaceae that occur on nutrient-impoverished soils in south-western Australia are calcifuge, despite their ability to produce cluster-roots, which effectively mobilize soil P and micronutrients. We hypothesize that the mechanism explaining the calcifuge habit in Proteaceae is their sensitivity to P and calcium (Ca), and that soil-indifferent species are less sensitive to the interaction of these nutrients. In this study, we analyzed growth, gas-exchange rate, and chlorophyll fluorescence of two soil-indifferent and four calcifuge Hakea and Banksia (Proteaceae) species from south-western Australia, across a range of P and Ca concentrations in hydroponic solution. We observed Ca-enhanced P toxicity in all analyzed species, but to different extents depending on distribution type and genus. Increasing P supply enhanced plant growth, leaf biomass, and photosynthetic rates of soil-indifferent species in a pattern largely independent of Ca supply. In contrast, positive physiological responses to increasing [P] in calcifuges were either absent or limited to low Ca supply, indicating that calcifuges were far more sensitive to Ca-enhanced P toxicity. In calcifuge Hakeas, we attributed this to higher leaf [P], and in calcifuge Banksias to lower leaf zinc concentration. These differences help to explain these species' contrasting sensitivity to Ca-enhanced P toxicity and account for the exclusion of most Proteaceae from calcareous habitats. We surmise that Ca-enhanced P toxicity is a major factor explaining the calcifuge habit of Proteaceae, and, possibly, other P-sensitive plants.

No evidence of regulation in root-mediated iron reduction in two Strategy I cluster-rooted Banksia species (Proteaceae)

AimsNon-mycorrhizal species such as Banksia (Proteaceae) that depend on root exudates to acquire phosphorus (P) are prominent in south-western Australia, a biodiversity hotspot on severely P-impoverished soils. We investigated the consequences of an exudate-releasing P-mobilising strategy related to control of iron (Fe) acquisition in two Banksia species, B. attenuata R.Br. and B. laricina C. Gardner, that differ greatly in their geographical distribution and rarity.MethodsWe undertook solution culture experiments to measure root-mediated Fe reduction (FeR) in non-cluster and cluster roots at four stages of cluster-root development, and whole root systems for plants grown at 2 to 300 μM Fe (as Fe-EDTA). As a positive control, we used Pisum sativum (cv. Dunn) to validate the FeR assay.ResultsUnlike typical Strategy I species, both Banksia species showed no significant variation in FeR, for either cluster or non-cluster roots, when grown at a wide range of Fe supply. For roots of different developmental stages, we measured a range for B. attenuata cluster roots of 0.13 ± 0.03 to 1.29 ± 0.14 μmol Fe3+ reduced g−1 FW h−1 and 0.56 ± 0.11 to 1.10 ± 0.24 μmol Fe3+ reduced g−1 FW h−1 in non-cluster roots. Similarly, for B. laricina cluster-roots, FeR ranged from 0.22 ± 0.07 to 1.21 ± 0.37 μmol Fe3+ reduced g−1 FW h−1, and in non-cluster roots from 0.56 ± 0.11 to 0.71 ± 0.08 μmol Fe3+ reduced g−1 FW h−1. We also observed only minor differences for whole-root system FeR, and even though B. attenuata showed signs of leaf Fe deficiency in the 2 μM Fe treatment, its FeR was the lowest of both species across all treatments at 0.079 ± 0.009 μmol Fe3+ reduced g−1 FW h−1, compared with the fastest rate of 0.20 ± 0.014 μmol Fe3+ reduced g−1 FW h−1 for B. laricina in the 28 μM Fe treatment. Taking plants through a pulse from low to high Fe, then back to low Fe supply did not elucidate any significant response in FeR.ConclusionsAlthough Fe acquisition is tightly controlled in the investigated Banksia species, such control is not based on regulation of FeR, which challenges the model that is commonly accepted for Strategy I species.

Conservation challenges for Victorian Banksias: Workshop May 2020

Banksias are iconic Australian plants, with a Gondwanan origin. Their specialised roots help survival on low‐phosphorus soils, and many species have reproductive cycles adapted to fire. They are trophically important in many of Australia’s native forests, woodlands and heathlands, providing nectar, pollen and seeds for many species of birds, mammals and invertebrates. While legacies of land use have reduced the historic extent of some Banksia species, more recent declines have raised additional concerns about their persistence in some landscapes. A range of factors are considered to be impacting Banksia persistence (Lamont et al. 2007), including habitat loss and fragmentation (e.g. Miller et al. 2020), increased fire frequency (e.g. Bradstock et al. 1997) and climate change (e.g. Steel et al. 2019), but there is no consensus about how these interacting factors may be driving declines, and how widespread these declines may be. The challenges facing Banksia species conservation in the south‐eastern state of Victoria were the focus of an online workshop in May 2020. The workshop brought together researchers and land managers from Victorian and interstate universities, Victorian government agencies and volunteer land management groups to share their knowledge of the threats facing banksias, and document ideas to reverse declines. It was organised by Arthur Rylah Institute for Environmental Research (Department Environment, Land, Water and Planning), La Trobe University (Department of Ecology, Environment and Evolution) and Federation University (Centre for eResearch and Digital Innovation).

Role of roots in adaptation of soil-indifferent Proteaceae to calcareous soils in south-western Australia.

Very few of the >650 Proteaceae species in south-western Australia cope with the high calcium (Ca) levels in young, calcareous soils (soil indifferent); most are Ca sensitive and occur on nutrient-impoverished, acidic soils (calcifuge). We assessed possible control points for Ca transport across roots of two soil-indifferent (Hakea prostrata and Banksia prionotes) and two calcifuge (H. incrassata and B. menziesii) Proteaceae. Using quantitative X-ray microanalysis, we investigated cell-specific elemental Ca concentrations at two positions behind the apex in relation to development of apoplastic barriers in roots of plants grown in nutrient solution with low or high Ca supply. In H. prostrata, Ca accumulated in outer cortical cells at 20 mm behind the apex, but [Ca] was low in other cell types. In H. incrassata, [Ca] was low in all cells. Accumulation of Ca in roots of H. prostrata corresponded to development of apoplastic barriers in the endodermis. We found similar [Ca] profiles in roots and similar [Ca] in leaves of two contrasting Banksia species. Soil-indifferent Hakea and Banksia species show different strategies to inhabit calcareous soils: H. prostrata intercepts Ca in roots, reducing transport to shoots, whereas B. prionotes allocates Ca to specific leaf cells.

Host-shifts at family level in the Australian Acacia-thrips lineage (Thysanoptera, Phlaeothripinae) with two new species.

The Australian Acacia-thrips lineage comprises at least 250 species in 30 genera of Phlaeothripinae, all of them known only from Acacia species in Australia. Two new species from two of these genera are described here as the first recorded instances of host-shifting within this diverse thrips lineage, with the host shifts being between unrelated angiosperm orders, from Fabales to Proteales and Myrtales. Brakothrips eucalypti sp. n. is described from the branches of a species of Eucalyptus, and Katothrips banksiae sp. n. is described forming substantial colonies within lepidopterous leaf mines on a species of Banksia. Previously these thrips genera were known only from Acacia species, and comprised seven described species of Brakothrips and 35 described species of Katothrips.

New species of xeromorphic Banksia (Proteaceae) foliage and Banksia-like pollen from the late Eocene of Western Australia

Banksia microphylla leaf fossils and Banksieaeidites zanthus pollen are newly described from late Eocene lignite of the Zanthus-11 borehole, drilled east of Norseman in Western Australia. The leaf fossils are the first known in Banksia to show extreme narrowness (<1.5 mm wide) combined with the xeromorphic trait of margins rolled onto the lower surface so that the diffusely placed stomata are exposed to the outside environment only via grooves on each side of a thick, abaxial midrib. Both this Banksia leaf type and another with encrypted stomata evolved before the widespread initiation of severe climatic aridity in the late Neogene, likely in regions of edaphic infertility and periodic water stress. New interpretations of leaf morphology and foliar evolutionary pathways in Banksia are proposed. Banksia microphylla probably belongs to subgenus Spathulatae, where it strongly resembles many species in the large, wholly Western Australian clade that includes most species in section Oncostylis, series Abietinae. Banksieaeidites zanthus is morphologically consistent with Banksia pollen, and its extremely small size also suggests placement in Spathulatae. The new fossils and other evidence from Zanthus-11 indicate the local presence of quite open, sclerophyll vegetation with conifers, which was unlikely to have been frequently burnt.

Wandering and Wildflowering: Walking with Women into Intimacy and Ecological Action

Wandering and Wildflowering: Walking with Women into Intimacy and Ecological Action

Protecting Offspring Against Fire: Lessons From Banksia Seed Pods.

Wildfires are a natural component in many terrestrial ecosystems and often play a crucial role in maintaining biodiversity, particularly in the fire-prone regions of Australia. A prime example of plants that are able to persist in these regions is the genus Banksia. Most Banksia species that occur in fire-prone regions produce woody seed pods (follicles), which open during or soon after fire to release seeds into the post-fire environment. For population persistence, many Banksia species depend on recruitment from these canopy-stored seeds. Therefore, it is critical that their seeds are protected from heat and rapid oxidation during fire. Here, we show how different species of Banksia protect their seeds inside follicles while simultaneously opening up when experiencing fire. The ability of the follicles to protect seeds from heat is demonstrated by intense 180 s experimental burns, in which the maximum temperatures near the seeds ranged from ∼75°C for B. serrata to ∼90°C for B. prionotes and ∼95°C for B. candolleana, contrasting with the mean surface temperature of ∼450°C. Many seeds of native Australian plants, including those of Banksia, are able to survive these temperatures. Structural analysis of individual follicles from these three Banksia species demonstrates that all of them rely on a multicomponent system, consisting of two valves, a porous separator and a thin layer of air surrounding the seeds. The particular geometric arrangement of these components determines the rate of heat transfer more than the tissue properties alone, revealing that a strong embedment into the central rachis can compensate for thin follicle valves. Furthermore, we highlight the role of the separator as an important thermal insulator. Our study suggests that the genus Banksia employs a variety of combinations in terms of follicle size, valve thickness, composition and geometric arrangement to effectively protect canopy-stored seeds during fire.

Implications of Banksia seed reward for conservation and management of Carnaby’s cockatoo on the Swan coastal plain, Western Australia

The food resource utilisation of six species of Banksia by the endangered Carnaby’s cockatoo (Calyptorhynchus latirostris) was investigated on the Swan coastal plain, Western Australia, over a 12-month period. The energy yield from the seeds harvested by the cockatoos was determined and the information was combined with data on the number of infructescences produced per hectare, the average seed yield per infructescence and the average rate of harvest of that species of seed by the cockatoos to calculate estimates of the number of infructescences required to support a single cockatoo per day under a range of scenarios. Over 65% of infructescences of each species of Banksia handled by the cockatoos were consumed for seed. Banksia sessilis had the largest number of infructescences and follicles manipulated by Carnaby’s cockatoos. The energy content of Banksia seed was 20–23 kJ g–1. Seed weight varied from 0.075 ± 0.016 (s.e.) g for B. attenuata to 0.007 ± 0.002 (s.e.) g for B. sessilis. The number of infructescences required to meet the birds’ daily energy intake ranged from 14 for B. grandis to 3821 for B. sessilis. The results have important implications for the continued capacity of the Swan coastal plain to support Carnaby’s cockatoos, for the future survival of obligate seeding Banksia spp. and for anthropogenic revegetation programs utilising Banksia spp.

Salt and waterlogging stress impacts on seed germination and early seedling growth of selected endemic plant species from Western Australia

Six perennial species endemic to South West Western Australia (Acacia trulliformis, Austrostipa geoffreyi, Banksia oligantha, B. mucronulata, Hakea tuberculata and Orthrosanthus muelleri) were screened for salt tolerance and recovery during seed germination. Growth and survival of 6-month old seedlings of these six plus a further vegetatively propagated species (Myoporum turbinatum) were subsequently examined in response to salt and waterlogging application. Water uptake under elevated saline conditions (200 and 400 mM NaCl) was slow, but not restrictive to germination. Moreover, a large proportion of seeds that were unable to germinate under saline conditions recovered after being transferred to non-saline conditions. Germination, growth and survival varied with species and the salt concentration used. Increasing salt concentrations tended to increase time to germination. Germination of Acacia trulliformis seeds declined exponentially with increasing salinity, and seedlings suffered reduced growth under saline and non-saline waterlogging. Austrostipa geoffreyi seeds were sensitive to saline treatments but seedlings were highly tolerant of both saline and/or waterlogged conditions. Germination of the three proteaceous species declined significantly under highly saline conditions (400 mM NaCl) with seedlings of the two Banksia species not surviving any treatment with the exception of non-saline waterlogging. Seedlings of H. tuberculata were more resilient to treatment conditions. Orthrosanthus muelleri was sensitive to salt stress during germination but was highly resistant to waterlogging, both saline and non-saline. This study provides an insight into the response and resilience of components of the vegetation understorey of saline-affected regions of Western Australia not usually evaluated allowing for more informed restoration.

Phytophthora cinnamomi as a driver of forest change: Implications for conservation and management

Abstract Phytophthora cinnamomi is a soil-borne plant pathogen of global significance, threatening many forest tree species around the world. In contrast to other well-known tree pathogens, P. cinnamomi is a generalist pathogen that, in many cases, causes less immediately obvious symptoms, making P. cinnamomi more difficult to diagnose. This creates special challenges for those trying to assess and manage diseases caused by P. cinnanmomi. P. cinnamomi affects a wide range of tree species across the world including chestnuts, particularly American and European chestnuts, Eucalyptus and Banksia species in Australia, and oaks in Mediterranean Europe. We believe that forest professionals should incorporate an understanding of the diseases caused by P. cinnamomi in conservation, management, and restoration of threatened ecosystems dominated by P. cinnamomi-affected tree species. Here we review the impact of P. cinnamomi on forest ecosystems internationally and suggest three major strategies for improving forest resilience to P. cinnamomi: (1) Improving site quality to reduce risk of P. cinnamomi-related disease, (2) Genetically improving threatened species to improve resistance to P. cinnamomi, and (3) Restricting further spread (especially by nursery trade) of P. cinnamomi.

Fire management strategies to maintain species population processes in a fragmented landscape of fire‐interval extremes

Changed fire regimes have led to declines of fire-regime-adapted species and loss of biodiversity globally. Fire affects population processes of growth, reproduction, and dispersal in different ways, but there is little guidance about the best fire regime(s) to maintain species population processes in fire-prone ecosystems. We use a process-based approach to determine the best range of fire intervals for keystone plant species in a highly modified Mediterranean ecosystem in southwestern Australia where current fire regimes vary. In highly fragmented areas, fires are few due to limited ignitions and active suppression of wildfire on private land, while in highly connected protected areas fires are frequent and extensive. Using matrix population models, we predict population growth of seven Banksia species under different environmental conditions and patch connectivity, and evaluate the sensitivity of species survival to different fire management strategies and burning intervals. We discover that contrasting, complementary patterns of species life-histories with time since fire result in no single best fire regime. All strategies result in the local patch extinction of at least one species. A small number of burning strategies secure complementary species sets depending on connectivity and post-fire growing conditions. A strategy of no fire always leads to fewer species persisting than prescribed fire or random wildfire, while too-frequent or too-rare burning regimes lead to the possible local extinction of all species. In low landscape connectivity, we find a smaller range of suitable fire intervals, and strategies of prescribed or random burning result in a lower number of species with positive growth rates after 100years on average compared with burning high connectivity patches. Prescribed fire may reduce or increase extinction risk when applied in combination with wildfire depending on patch connectivity. Poor growing conditions result in a significantly reduced number of species exhibiting positive growth rates after 100years of management. By exploring the consequences of managing fire, we are able to identify which species are likely to disappear under a given fire regime. Identifying the appropriate complementarity of fire intervals, and their species-specific as well as community-level consequences, is crucial to reduce local extinctions of species in fragmented fire-prone landscapes.

Analysing patterns of spatial and niche overlap among species at multiple resolutions

AbstractAimAnalyses of spatial overlap in species distributions are frequently used to test a range of ecological and evolutionary hypotheses, from the role of competition in community assembly to the geography of speciation. Most studies quantify overlap at one spatial resolution. Here we explore the effects of measuring spatial and niche overlap patterns for the same clade (Banksia) at multiple resolutions.LocationAustralia.MethodsWe quantify overlap among species using broad overlap of species range polygons, proximity of occurrence points and co‐occurrence within small survey plots. We compare overlap patterns with null models using age–range correlations and the frequency of sympatric sister species. We then use similar methods to examine patterns of overlap in environmental niche dimensions.ResultsBanksia species show a wide range of overlap values based on range polygons and point proximities, but very low levels of co‐occurrence at the local scale. Intercepts of age–range correlations point to higher levels of overlap among recently diverged species than expected. However, comparing the frequency of sympatric sister species with an evolutionary null model supports a prevailing allopatric mode of speciation. In many cases, niche overlap between species exceeds that expected from phylogenetic relatedness or spatial overlap alone.Main conclusionsPatterns of broad geographical overlap among Banksia species support a predominantly allopatric mode of speciation, combined with post‐speciation range drift. There is more evidence for niche conservatism than for rapid niche divergence among closely related species. This pattern is consistent with broad‐scale geomorphic and landscape complexity as a driver of plant speciation in south‐west Australia. It is less consistent with finer‐scale mechanisms of species divergence such as fire mosaics, or with ecological divergence in sympatry. Analysis of species overlap patterns at different resolutions is a useful approach for revealing the multiple ecological and historical factors that influence species distributions.

Can sensitivity to temperature during germination help predict global warming vulnerability?

AbstractSeed germination is vital for persistence in species that rely on seeds for post-disturbance regeneration. It is a high-risk phase and vulnerable to environmental parameters. Here, I assessed temperature sensitivity for germination in Banksia L.f. (Proteaceae) from south-western Australia, screening all 38 endemic obligate seeder species. A bi-directional temperature gradient plate with 49 temperature combinations (constant and fluctuating) between 5 and 40°C was used to profile germination temperature requirements and identify upper and lower temperature thresholds for germination. Using these data the impact of increasing temperatures on germination in these species was modelled under high and low greenhouse gas scenarios for 2050 and 2070. The results suggest that many Banksia species from the region have wide physiological tolerance for high germination temperatures, although a number of common, but geographically restricted species, such as B. praemorsa, B. oreophila and B. quercifolia, have more narrow temperature windows for germination than at least one of the rarer species (B. verticillata). Only B. dryandroides is expected to decline in germination in the future; however, the optimal germination timing for many species is predicted to occur later under climate warming. In conjunction with declining rainfall, this germination delay will place seedlings closer to the summer dry in this seasonal Mediterranean-climate ecosystem and thus they will be more vulnerable to desiccation. The framework developed here can be used to identify vulnerable species for monitoring of early population decline.

Fire regime, soil fertility and growth form interact to shape fire and growth traits in two co-occurring Banksia species

Inter- and intra-specific comparisons were made between several co-occurring populations of two Banksia species growing in Eastern Australian dry sclerophyll open woodland that experiences a grass-fueled surface-fire regime. The two species differ in life-history from a short basal resprouter (B. Neoanglica) to a tall fire-survivor (B. integrifolia). Growth (LMA = leaf mass per unit mass area), persistence (bark thickness) and recruitment (serotiny) traits were determined across independent gradients in soil fertility and fire frequency. Trait correlations for the two species showed distinct patterns, from each other and across environments, with the resprouter having higher LMA and infructescence retention but lower bark thickness. However, there were no consistent intra-specific patterns with variation in fire and soil nutrients. LMA only varied in B. integrifolia with soil fertility, and fire frequency had no effect on either species. Relative bark thickness varied with plant size but not among sites. There was a trend of increasing pyriscence with fire in B. neoanglica and with lower nutrients soils but not in B. integrifolia. Clearly the two species respond differently to variability in nutrients and fire. Nevertheless, growth form appears to be the strongest determinant of both growth (LMA) and fire traits (thicker bark; pyriscence) in these co-existing Banksia species.

The importance of pollinators and autonomous self-fertilisation in the early stages of plant invasions: Banksia and Hakea (Proteaceae) as case studies.

Reproduction is a crucial stage in the naturalisation of introduced plant species. Here, using breeding system experiments and observations of floral visitors, we investigate whether a lack of pollinators or an inability to autonomously self-fertilise limits naturalisation in five Australian Banksia species and the co-familial Hakea salicifolia in South Africa. Banksia species were heavily utilised by native insects and nectar-feeding birds. Although Banksia produced fruit when pollinators were excluded, pollinators significantly increased seed set in four of the five species. H.salicifolia flowers were visited by 11 insect species; honeybees (Apis mellifera) were the main visitors. Flowers in naturalised H.salicifolia populations received almost four times the number of visits as flowers in non-naturalised populations; the latter showed both pollen limitation (PLI 0.40) and partial self-incompatibility. This should not prevent invasion, since H.salicifolia produces fruits via autonomous selfing in the absence of pollinators. The results suggest a limited role of breeding systems in mediating naturalisation of introduced Proteaceae species. Other factors, such as features of the recipient environments, appear to be more important. Spatial variation in rates of reproduction might, however, explain variation in the extent and rate of naturalisation of different populations.