Repeated evolution of carnivory among Indo‐Australian rodents

Convergent evolution, often observed in island archipelagos, provides compelling evidence for the importance of natural selection as a generator of species and ecological diversity. The Indo‐Australian Archipelago (IAA) is the world's largest island system and encompasses distinct biogeographic units, including the Asian (Sunda) and Australian (Sahul) continental shelves, which together bracket the oceanic archipelagos of the Philippines and Wallacea. Each of these biogeographic units houses numerous endemic rodents in the family Muridae. Carnivorous murids, that is those that feed on animals, have evolved independently in Sunda, Sulawesi (part of Wallacea), the Philippines, and Sahul, but the number of origins of carnivory among IAA murids is unknown. We conducted a comprehensive phylogenetic analysis of carnivorous murids of the IAA, combined with estimates of ancestral states for broad diet categories (herbivore, omnivore, and carnivore) and geographic ranges. These analyses demonstrate that carnivory evolved independently four times after overwater colonization, including in situ origins on the Philippines, Sulawesi, and Sahul. In each biogeographic unit the origin of carnivory was followed by evolution of more specialized carnivorous ecomorphs such as vermivores, insectivores, and amphibious rats.     

Does biogeographical history matter? Diversity and distribution of lotic midges (Diptera: Chironomidae) in the Australian Wet Tropics

Abstract We examined broad scale patterns of diversity and distribution of lotic Chironomidae (Diptera) within the Wet Tropics bioregion of northern Queensland, Australia. Field surveys across broad latitudinal and altitudinal gradients within the Wet Tropics revealed a fauna of 87 species‐level taxa in 49 genera comprising three main elements: a small genuinely tropical fraction, and larger cosmopolitan and Gondwanan components. The latter group originated when Australia, as part of the ancient Gondwana supercontinent, was situated over Antarctic latitudes with a cooler, wetter climate than today. In the Wet Tropics, cool Gondwanan taxa occurred predominantly in upland and shaded lowland sites, but no species appeared narrowly temperature restricted, and there was no faunal zonation with altitude. Most chironomid species occurred at all latitudes within the Wet Tropics, with no evidence for an enduring effect of the historical rainforest contractions on current‐day distribution patterns. These findings contrast with those for aquatic faunas elsewhere in the world and for the terrestrial Wet Tropics fauna. We relate this to the generally broad environmental tolerances of Australian chironomids, and comment on why the latitudinal diversity gradient does not apply to the Australian chironomid fauna.     

Australian Tropical and Subtropical Rain Forest Community Assembly: Phylogeny,Functional Biogeography,and Environmental Gradients

To compare community assemblage patterns in tropical northeastern and subtropical central eastern Australia across selected gradients and scales, we tested the relationship of species traits with phylogenetic structure, and niche breadth. We considered phylogenetic relationships across current‐day species in assemblages in relation to rain forest species pool sizes, and trait values along gradients including elevation and latitude. Trait values were quantified across scales for seed size, leaf area, wood density and maximum height at maturity for 1137 species and 596 assemblages using trait gradient analysis (TGA). Local assemblages of subtropical species had narrower trait ranges, and higher niche breadth values than corresponding assemblages of tropical species. Leaf size and seed size increased at low latitudes, and community phylogenetic structure was most strongly correlated with seed traits in the subtropics, reflecting dispersal and re‐colonization processes. Elevation accounted for little of the variance in community phylogenetic structure or trait variation across local and regional scales. Stable moist forest areas retained many species from ancestral rain forest lineages across a range of temporally conserved habitats; species within assemblages were less related; and rain forest assemblages had higher functional diversity, but lower niche breadth. This suggests that on average, assemblages of species in stable areas had greater trait variation and narrower distributions. Historic and recent rain forest contraction and re‐expansion can result in recolonized areas that are dominated by species that are more related (phylogenetically) than by chance, have smaller, widely dispersed seeds, and greater niche breadth (broader distributions).     

Deeply divergent mitochondrial lineages reveal patterns of local endemism in chironomids of the Australian Wet Tropics

The Wet Tropics bioregion of north‐eastern Australia has been subject to extensive fluctuations in climate throughout the late Pliocene and Pleistocene. Cycles of rainforest contraction and expansion of dry sclerophyll forest associated with such climatic fluctuations are postulated to have played a major role in driving geographical endemism in terrestrial rainforest taxa. Consequences for the distributions of aquatic organisms, however, are poorly understood. The Australian non‐biting midge species Echinocladius martini Cranston (Diptera: Chironomidae), although restricted to cool, well‐forested freshwater streams, has been considered to be able to disperse among populations located in isolated rainforest pockets during periods of sclerophyllous forest expansion, potentially limiting the effect of climatic fluctuations on patterns of endemism. In this study, mitochondrial COI and 16S data were analysed for E. martini collected from eight sites spanning the Wet Tropics bioregion to assess the scale and extent of phylogeographic structure. Analyses of genetic structure showed several highly divergent cryptic lineages with restricted geographical distributions. Within one of the identified lineages, strong genetic structure implied that dispersal among proximate (<1 km apart) streams was extremely restricted. The results suggest that vicariant processes, most likely due to the systemic drying of the Australian continent during the Plio‐Pleistocene, might have fragmented historical E. martini populations and, hence, promoted divergence in allopatry.     

Recent colonization and expansion through the Lesser Sundas by seven amphibian and reptile species

The Lesser Sundas Archipelago is comprised of two parallel chains of islands that extend between the Asian continental shelf (Sundaland) and Australo‐Papuan continental shelf (Sahul). These islands have served as stepping stones for taxa dispersing between the Asian and Australo‐Papuan biogeographical realms. While the oceanic barriers have prevented many species from colonizing the archipelago, a number of terrestrial vertebrate species have colonized the islands either by rafting/swimming or by human introduction. Here, we examine phylogeographic structure within the Lesser Sundas for three snake, two lizard and two frog species that each has a Sunda Shelf origin. These species are suspected to have recently colonized the archipelago, though all have inhabited the Lesser Sundas for over 100 years. We sequenced mtDNA from 231 samples to test whether there is sufficiently deep genetic structure within any of these taxa to reject human‐mediated introduction. Additionally, we tested for genetic signatures of population expansion consistent with recent introduction and estimated the ages of Lesser Sundas clades, if any exist. Our results show little to no genetic structure between populations on different islands in five species and moderate structure in two species. Nucleotide diversity is low for all species, and the ages of the most recent common ancestor for species with monophyletic Lesser Sundas lineages date to the Holocene or late Pleistocene. These results support the hypothesis that these species entered the archipelago relatively recently and either naturally colonized or were introduced by humans to most of the larger islands in the archipelago within a short time span.     

Phylogeny and spatio‐temporal diversification of Prunus subgenus Laurocerasus section Mesopygeum (Rosaceae) in the Malesian region

The goals of this study were to reconstruct the phylogeny of Prunus subgenus Laurocerasus section Mesopygeum and to provide a preliminary assessment of its spatio‐temporal diversification in the Malesian region. We inferred the phylogeny using nuclear ITS and ETS and plastid psbA‐trnH, rps16, rpl16, and trnC‐petN sequences. Our analyses support the monophyly of sect. Mesopygeum. Within sect. Mesopygeum, we identified four main subclades: (i) Prunus lancilimba from continental Asia; (ii) Prunus ruthii from Malay Peninsula; (iii) a subclade comprising species from areas centered on the Sunda shelf and also a few species from continental Asia and Wallacea; and (iv) a subclade composed of species from areas of the Sahul shelf, with a small number of taxa also from areas of the Sunda shelf, continental Asia, and the Philippines. We estimated that sect. Mesopygeum originated in continental Asia at c. 44.71 Mya (95% HPD: 31.66–46.90). Nine dispersals between major geographic areas were inferred. From continental Asia, three and two dispersals were inferred to the Sunda shelf and the Sahul shelf in the mid‐Oligocene, respectively. Two dispersals were inferred from the Sahul shelf region to the Sunda shelf in late Oligecene and early Miocene, respectively. There were also two dispersals inferred from the Sunda shelf region, one to the Philippines and one to Wallacea, in the middle and late Miocene, respectively. The diversification in sect. Mesopygeum was likely driven by active geologic events and orogenies in the Neogene in the Malesian region.     

Exotic vine invasions following cyclone disturbance in Australian Wet Tropics rainforests: A review

The Australian Wet Tropics region extends for almost 900 000 ha along the coastline of north‐east Queensland. The rainforests in this region have a rich and unique biodiversity and are World Heritage listed by UNESCO. Disturbance from tropical cyclones is a significant driver of the rainforest dynamics in this area, and when frequent or intense can facilitate the recruitment and expansion of exotic invasive species. Exotic vines are of particular concern for forest conservation as they can be highly competitive with native vegetation and may prevent forest regeneration. This literature review found evidence that fragmented forests, which are very common in the Australian Wet Tropics, are vulnerable to post‐cyclone vine invasion. In particular, although the diversity and abundance of herbaceous vines tend to decline as the canopy closes 2 years post‐cyclone disturbance, woody exotic vines and scramblers may persist for much longer or even increase in numbers. Since forest recovery in these systems is influenced by the severity and recurrence of disturbance, an increase in cyclone intensity under climate change may cause rapid changes in rainforest structure, composition and diversity, and increase exotic vine invasion. Post‐cyclone management of vines appears to require direct intervention, with manual cutting being currently the most effective method. However, there are a number of difficulties to its wide implementation in Australia, and further study on options for control is needed. Abstract in Spanish is available with online material.     

Monsoon rain forest seedling dynamics, northern Australia: contrasts with regeneration in eucalypt-dominated savannas

Aim To explore: (1) the relative influences of site conditions, especially moisture relations, on pathways and rates of monsoon rain forest seedling and sapling regeneration, especially of canopy dominants, in northern Australia; and (2) contrasts between regeneration syndromes of dominant woody taxa in savannas and monsoon rain forest. Location Four monsoon rain forest sites, representative of regional major habitat and vegetation types, in Kakadu National Park, northern Australia. Methods A decadal study involved: (1) initial assessment over 2.5 years to explore within‐year variability in seed rain, dormant seed banks and seedling (< 50 cm height) dynamics; and (2) thereafter, monitoring of seedling and sapling (50 cm height to 5 cm d.b.h.) dynamics undertaken annually in the late dry season. On the basis of observations from this and other studies, regeneration syndromes of dominant monsoon rain forest taxa are contrasted with comparable information for dominant woody savanna taxa, Eucalyptus and Corymbia especially. Results Key observations from the monsoon rain forest regeneration dynamics study component are that: (1) peak seed rain inputs of rain forest taxa were observed in the wet season at perennially moist sites, whereas inputs at seasonally dry sites extended into, or peaked in, the dry season; (2) dormant soil seed banks of woody rain forest taxa were dominated by pioneer taxa, especially figs; (3) longevity of dormant seed banks of woody monsoon rain forest taxa, including figs, was expended within 3 years; (4) seedling recruitment of monsoon rain forest woody taxa was derived mostly from wet season seed rain with limited inputs from soil seed banks; (5) at all sites rain forest seedling mortality occurred mostly in the dry season; (6) rain forest seedling and sapling densities were consistently greater at moist sites; (7) recruitment from clonal reproduction was negligible, even following unplanned low intensity fires. Main conclusions By comparison with dominant savanna eucalypts, dominant monsoon rain forest taxa recruit substantially greater stocks of seedlings, but exhibit slower aerial growth and development of resprouting capacity in early years, lack lignotubers in mesic species, and lack capacity for clonal reproduction. The reliance on sexual as opposed to vegetative reproduction places monsoon rain forest taxa at significant disadvantage, especially slower growing species on seasonally dry sites, given annual–biennial fires in many north Australian savannas.     

Extent and timing of floristic exchange between Australian and Asian rain forests

Aim We tested an entrenched concept – that the Australian rain forest flora is essentially a Gondwanan relict. We also assessed the role of regional‐level source–sink dynamics in the assembly of this flora. Location Eastern Australia. Methods To avoid potential biases inherent in selective studies undertaken to date, we used an analytical, whole‐of‐flora approach integrated with the fossil record. We identified disjunctions between woody Australian rain forest plant taxa and relatives on other land masses. To test the strength of the fossil evidence for the regional antiquity of this flora, we evaluated the proportion of these disjunct clades represented in the Australian fossil record, and to minimize the effects of biases in this record, we compared late Quaternary (i.e. late Pleistocene and Holocene, 126–0 ka), Pliocene and late Oligocene–early Miocene Australian pollen records interpreted as tropical rain forest. Using within‐species disjunctions as a proxy, we assessed the role of recent immigration from Asia into Australia. To assess the role of source–sink dynamics, we performed comparative analyses of disjunctions in major rain forest categories representing a north–south/climatic gradient. Results Southern Australian, cool temperate (microthermal) rain forests contain many floristic disjunctions with Gondwanan fragments and most of these clades have Gondwanan fossils. Disjunct clades in Australian mesothermal rain forest mostly occur in Asia/Malesia and a low proportion of these clades show pre‐Neogene records. Many clades in lowland tropical and ‘dry’ rain forest show disjunctions with Asia/Malesia and few have Australian fossil records. Rates of recent immigration from Asia/Malesia are high in these northern forests, and outweigh rates of recent emigration approximately nine‐fold. The late Quaternary fossil record has many more rain forest angiosperms than Oligocene–Miocene and Pliocene floras, consistent with extensive late Cenozoic immigration. Main conclusions The microthermal rain forests are largely Gondwanan relicts, but there is progressively greater, and more recent contribution from Asia/Malesia into more northern, and more lowland tropical rain forests. This variation reflects a strong gradient in geographic and ecological proximity between these forests and source floras in Asia/Malesia, and is consistent with a source–sink size model of immigration driven by late Cenozoic contractions and expansions of Australian rain forest.     

Amphidromy links a newly documented fish community of continental Australian streams, to oceanic islands of the west Pacific

Background

Indo-Pacific high island streams experience extreme hydrological variation, and are characterised by freshwater fish species with an amphidromous life history. Amphidromy is a likely adaptation for colonisation of island streams following stochastic events that lead to local extirpation. In the Wet Tropics of north-eastern Australia, steep coastal mountain streams share similar physical characteristics to island systems. These streams are poorly surveyed, but may provide suitable habitat for amphidromous species. However, due to their ephemeral nature, common non-diadromous freshwater species of continental Australia are unlikely to persist. Consequently, we hypothesise that coastal Wet Tropics streams are faunally more similar, to distant Pacific island communities, than to nearby faunas of large continental rivers.

Methods/Principal Findings

Surveys of coastal Wet Tropics streams recorded 26 species, 10 of which are first records for Australia, with three species undescribed. This fish community is unique in an Australian context in that it contains mostly amphidromous species, including sicydiine gobies of the genera Sicyopterus, Sicyopus, Smilosicyopus and Stiphodon. Species presence/absence data of coastal Wet Tropics streams were compared to both Wet Tropics river networks and Pacific island faunas. ANOSIM indicated the fish fauna of north-eastern Australian coastal streams were more similar to distant Pacific islands (R = 0.76), than to nearby continental rivers (R = 0.98).

Main Conclusions/Significance

Coastal Wet Tropics streams are faunally more similar to distant Pacific islands (79% of species shared), than to nearby continental fauna due to two factors. First, coastal Wet Tropics streams lack many non-diadromous freshwater fish which are common in nearby large rivers. Second, many amphidromous species found in coastal Wet Tropics streams and Indo-Pacific islands remain absent from large rivers of the Wet Tropics. The evolutionary and conservation significance of this newly discovered Australian fauna requires clarification in the context of the wider amphidromous fish community of the Pacific.     

Mechanisms of rainforest persistence and recruitment in frequently burnt wet tropical eucalypt forests

The role of fire in governing rainforest–eucalypt forest ecotone dynamics is of theoretical interest and has conservation management implications. Several eucalypt forests in the Wet Tropics of Australia have an endangered status due to extensive conversion to rainforest. Rainforest plants are known to survive occasional low intensity fires in the eucalypt forest ecotone. However, the ability of rainforest plants to survive frequent fires remains untested. The timing of rainforest expansion is also a subject of interest, and is generally considered to be delayed until fire has been absent for several years. We used 14 years of data collected across 13 plots in the Wet Tropics of north‐eastern Australia to test predictions regarding rainforest seedling recruitment and post‐fire regenerative capacity. The 13 plots received different numbers of fires, between zero and five, over the 14‐year study. The recruitment of new rainforest plants in the ecotone was most abundant in the initial year after fire. If this post‐fire pulse of recruitment is left undisturbed, it can facilitate the subsequent germination of additional rainforest species. The removal of grass cover, whether temporarily in the immediate post‐fire environment or once a developing rainforest mid strata shades out grasses, appears crucial to abundant rainforest recruitment. A variety of tropical rainforest species can persist under a frequent fire regime through resprouting. The difference in the mode of resprouting, between ground‐level coppicing rainforest plants and canopy resprouting eucalypt forest trees, is the critical mechanism that causes regular fire to maintain an open structure in eucalypt forests. The inability of rainforest species to maintain their height when fires fully scorch their crowns, temporarily resets the forest's open structure and delays the rainforest's ability to dominate through shading out grasses to transform the ecosystem into a closed forest.     

Laying low: Rugged lowland rainforest preferred by feral cats in the Australian Wet Tropics

Invasive mesopredators are responsible for the decline of many species of native mammals worldwide. Feral cats have been causally linked to multiple extinctions of Australian mammals since European colonization. While feral cats are found throughout Australia, most research has been undertaken in arid habitats, thus there is a limited understanding of feral cat distribution, abundance, and ecology in Australian tropical rainforests. We carried out camera‐trapping surveys at 108 locations across seven study sites, spanning 200 km in the Australian Wet Tropics. Single‐species occupancy analysis was implemented to investigate how environmental factors influence feral cat distribution. Feral cats were detected at a rate of 5.09 photographs/100 days, 11 times higher than previously recorded in the Australian Wet Tropics. The main environmental factors influencing feral cat occupancy were a positive association with terrain ruggedness, a negative association with elevation, and a higher affinity for rainforest than eucalypt forest. These findings were consistent with other studies on feral cat ecology but differed from similar surveys in Australia. Increasingly harsh and consistently wet weather conditions at higher elevations, and improved shelter in topographically complex habitats may drive cat preference for lowland rainforest. Feral cats were positively associated with roads, supporting the theory that roads facilitate access and colonization of feral cats within more remote parts of the rainforest. Higher elevation rainforests with no roads could act as refugia for native prey species within the critical weight range. Regular monitoring of existing roads should be implemented to monitor feral cats, and new linear infrastructure should be limited to prevent encroachment into these areas. This is pertinent as climate change modeling suggests that habitats at higher elevations will become similar to lower elevations, potentially making the environment more suitable for feral cat populations.     

Vagility and the Monsoon Rain Forest Archipelago of Northern Australia: Patterns of Genetic Diversity in Syzygium nervosum (Myrtaceae)1

Syzygium nervosum is a common monsoon rain forest tree. Its habitat in Australia consists of small rain forest patches that are scattered through a savanna matrix. It is a mast flowering canopy species that produces large quantities of fruits fed on by mobile frugivores such as birds and fruit bats. The genetic diversity of this species was investigated, especially in relation to rain forest patch size, geographic isolation, and geographic distribution. Syzygium nervosum was found to have high levels of genetic diversity within populations (H_e= 0.307). Diversity among populations, however, was relatively low (Fsr = 0.118), and was not spatially structured across its geographic range in Australia. This is thought to have been caused by relatively frequent gene flow among populations (N_m= 1.67), mediated primarily by mobile frugivores. Genetic diversity was not correlated with patch size or isolation. It is thought that seed dispersal by frugivores has acted to expand the effective population size of this species beyond the individual rain forest patch, and thus has prevented the substantial loss of genetic diversity that otherwise would have been observed. Thus this species is dependent upon these frugivores for the maintenance of its genetic diversity and hence its long-term viability. These results lend support to theories of post-Holocene expansion of rain forest by vagile species in northern Australia.     

Humid tropical rain forest has expanded into eucalypt forest and savanna over the last 50 years

Tropical rain forest expansion and savanna woody vegetation thickening appear to be a global trend, but there remains uncertainty about whether there is a common set of global drivers. Using geographic information techniques, we analyzed aerial photography of five areas in the humid tropics of northeastern Queensland, Australia, taken in the 1950s and 2008, to determine if changes in rain forest extent match those reported for the Australian monsoon tropics using similar techniques. Mapping of the 1950s aerial photography showed that of the combined study area (64,430 ha), 63% was classified as eucalypt forests/woodland and 37% as rain forest. Our mapping revealed that although most boundaries remained stable, there was a net increase of 732 ha of the original rain forest area over the study period, and negligible conversion of rain forest to eucalypt forest/woodland. Statistical modeling, controlling for spatial autocorrelation, indicated distance from preexisting rain forest as the strongest determinant of rain forest expansion. Margin extension had a mean rate across the five sites of 0.6 m per decade. Expansion was greater in tall open forest types but also occurred in shorter, more flammable woodland vegetation types. No correlations were detected with other local variables (aspect, elevation, geology, topography, drainage). Using a geographically weighted mean rate of rain forest margin extension across the whole region, we predict that over 25% of tall open forest (a forest type of high conservation significance) would still remain after 2000 years of rain forest expansion. This slow replacement is due to the convoluted nature of the rain forest boundary and the irregular shape of the tall open forest patches. Our analyses point to the increased concentration of atmospheric CO(2) as the most likely global driver of indiscriminate rain forest expansion occurring in northeastern Australia, by increasing tree growth and thereby overriding the effects of fire disturbance.     

Whole-Genome Genetic Diversity in a Sample of Australians with Deep Aboriginal Ancestry

Australia was probably settled soon after modern humans left Africa, but details of this ancient migration are not well understood. Debate centers on whether the Pleistocene Sahul continent (composed of New Guinea, Australia, and Tasmania) was first settled by a single wave followed by regional divergence into Aboriginal Australian and New Guinean populations (common origin) or whether different parts of the continent were initially populated independently. Australia has been the subject of relatively few DNA studies even though understanding regional variation in genomic structure and diversity will be important if disease-association mapping methods are to be successfully evaluated and applied across populations. We report on a genome-wide investigation of Australian Aboriginal SNP diversity in a sample of participants from the Riverine region. The phylogenetic relationship of these Aboriginal Australians to a range of other global populations demonstrates a deep common origin with Papuan New Guineans and Melanesians, with little evidence of substantial later migration until the very recent arrival of European colonists. The study provides valuable and robust insights into an early and important phase of human colonization of the globe. A broader survey of Australia, including diverse geographic sample populations, will be required to fully appreciate the continent''s unique population history and consequent genetic heritage, as well as the importance of both to the understanding of health issues.     

The effect of habitat fragmentation on the bee visitor assemblages of three Australian tropical rainforest tree species

Tropical forest loss and fragmentation can change bee community dynamics and potentially interrupt plant–pollinator relationships. While bee community responses to forest fragmentation have been investigated in a number of tropical regions, no studies have focused on this topic in Australia. In this study, we examine taxonomic and functional diversity of bees visiting flowers of three tree species across small and large rainforest fragments in Australian tropical landscapes. We found lower taxonomic diversity of bees visiting flowers of trees in small rainforest fragments compared with large forest fragments and show that bee species in small fragments were subsets of species in larger fragments. Bees visiting trees in small fragments also had higher mean body sizes than those in larger fragments, suggesting that small‐sized bees may be less likely to persist in small fragments. Lastly, we found reductions in the abundance of eusocial stingless bees visiting flowers in small fragments compared to large fragments. These results suggest that pollinator visits to native trees living in small tropical forest remnants may be reduced, which may in turn impact on a range of processes, potentially including forest regeneration and diversity maintenance in small forest remnants in Australian tropical countryside landscapes.     

Geographical variation in the morphology of four snake species from the Lesser Sunda Islands, eastern Indonesia

The morphology of three colubrid snakes, Elaphe subradiata, Psammodynastes pulverulentus, Lycodon aulicus capucinus , and the viper Trimeresurus albolabris , were examined from populations throughout the Lesser Sunda islands using canonical variate analysis. The Lesser Sundas form the western part of two extensive Island chains, the Banda Arcs, that lie between the large islands on the Sunda (Asian) and Sahul (Australian) continental shelves. The snakes of the Lesser Sunda islands show considerable intraspecific variation in morphology. Populations of Trimeresurus albolabris from Wetar and Psammodynastes pulverulentus from Alor show the greatest morphological separation from conspecifics. The most pronounced morphological differentiation in multidimensional space occurs between populations on islands that remained separated throughout the Pleistocene, when sea levels were about 120 m lower than present. Consensus tree evaluation of Mahalanobis distance for populations of these four species and Dendrelaphis pubis on the islands of Lombok, Sumba, Flores, Lembata and Alor, supports the finding that populations from islands that remained isolated throughout the Pleistocene by sea barriers show the greatest morphological divergence.     

Landscape Patterns in Rainforest Phylogenetic Signal: Isolated Islands of Refugia or Structured Continental Distributions?

Objectives

Identify patterns of change in species distributions, diversity, concentrations of evolutionary history, and assembly of Australian rainforests.

Methods

We used the distribution records of all known rainforest woody species in Australia across their full continental extent. These were analysed using measures of species richness, phylogenetic diversity (PD), phylogenetic endemism (PE) and phylogenetic structure (net relatedness index; NRI). Phylogenetic structure was assessed using both continental and regional species pools. To test the influence of growth-form, freestanding and climbing plants were analysed independently, and in combination.

Results

Species richness decreased along two generally orthogonal continental axes, corresponding with wet to seasonally dry and tropical to temperate habitats. The PE analyses identified four main areas of substantially restricted phylogenetic diversity, including parts of Cape York, Wet Tropics, Border Ranges, and Tasmania. The continental pool NRI results showed evenness (species less related than expected by chance) in groups of grid cells in coastally aligned areas of species rich tropical and sub-tropical rainforest, and in low diversity moist forest areas in the south-east of the Great Dividing Range and in Tasmania. Monsoon and drier vine forests, and moist forests inland from upland refugia showed phylogenetic clustering, reflecting lower diversity and more relatedness. Signals for evenness in Tasmania and clustering in northern monsoon forests weakened in analyses using regional species pools. For climbing plants, values for NRI by grid cell showed strong spatial structuring, with high diversity and PE concentrated in moist tropical and subtropical regions.

Conclusions/Significance

Concentrations of rainforest evolutionary history (phylo-diversity) were patchily distributed within a continuum of species distributions. Contrasting with previous concepts of rainforest community distribution, our findings of continuous distributions and continental connectivity have significant implications for interpreting rainforest evolutionary history and current day ecological processes, and for managing rainforest diversity in changing circumstances.     

Recruitment dynamics in a rainforest seedling community: context-independent impact of a keystone consumer

The influence of keystone consumers on community structure is frequently context-dependent; the same species plays a central organising role in some situations, but not others. On Christmas Island, in the Indian Ocean, a single species of omnivorous land crab, Gecarcoidea natalis, dominates the forest floor across intact rainforest. We hypothesised that this consumer plays a key role in regulating seedling recruitment and in controlling litter dynamics on the island, independent of the type of vegetation in which it occurred. To test this hypothesis, we conducted crab exclusion experiments in two forest types on the island and followed the dynamics of seedling recruitment and litter processing for six years. To determine if these effects were likely to be general across the island, we compared land crab densities and seedling abundance and diversity at ten sites across island rainforest. Surveys across island rainforest showed that seedlings of species susceptible to predation by land crabs are consistently rare. Abundance and diversity of these species were negatively correlated to red crab abundance. Although red land crabs may be important determinants of seedling recruitment to the overstorey, differences in overstorey and seedling composition at the sites suggests that recruitment of vulnerable trees still occurs at a temporal scale exceeding that of this study. These “windows” of recruitment may be related to infrequent events that reduce the effects of land crabs. Our results suggest that unlike the context dependence of most keystone consumers in continental systems, a single consumer, the red land crab, consistently controls the dynamics of seedling recruitment across this island rainforest.     

Phylogeography and divergence in the chloroplast genome of Western Australian Sandalwood (Santalum spicatum)

Western Australian sandalwood (Santalum spicatum) is widespread throughout Western Australia across the semiarid and arid regions. The diversity and phylogeographic patterns within the chloroplast genome of S. spicatum were investigated using restriction fragment length polymorphism analysis of 23 populations. The chloroplast diversity was structured into two main clades that were geographically separated, one centred in the southern (semiarid region) and the other in the northern (arid) region. Fragmentation due to climatic instability was identified as the most likely influence on the differentiation of the lineages. The lineage in the arid region showed a greater level of differentiation than that in the southern region, suggesting a higher level of gene flow or a more recent range expansion of sandalwood in the southern region. The phylogeographic pattern in the chloroplast genome is congruent with that detected in the nuclear genome, which identified different genetic influences between the regions and also suggested a more recent expansion of sandalwood in the southern region.