Decline of a biome: evolution,contraction, fragmentation,extinction and invasion of the Australian mesic zone biota

Aim The mesic biome, encompassing both rain forest and open sclerophyllous forests, is central to understanding the evolution of Australia’s terrestrial biota and has long been considered the ancestral biome of the continent. Our aims are to review and refine key hypotheses derived from palaeoclimatic data and the fossil record that are critical to understanding the evolution of the Australian mesic biota. We examine predictions arising from these hypotheses using available molecular phylogenetic and phylogeographical data. In doing so, we increase understanding of the mesic biota and highlight data deficiencies and fruitful areas for future research. Location The mesic biome of Australia, along the eastern coast of Australia, and in the south‐east and south‐west, including its rain forest and sclerophyllous, often eucalypt‐dominated, habitats. Methods We derived five hypotheses based on palaeoclimatic and fossil data regarding the evolution of the Australian mesic biota, particularly as it relates to the mesic biome. We evaluated predictions formulated from these hypotheses using suitable molecular phylogenies of terrestrial plants and animals and freshwater invertebrates. Results There was support for the ancestral position of mesic habitat in most clades, with support for rain forest habitat ancestry in some groups, while evidence of ancestry in mesic sclerophyllous habitats was also demonstrated for some plants and herpetofauna. Contraction of mesic habitats has led to extinction of numerous lineages in many clades and this is particularly evident in the rain forest component. Species richness was generally higher in sclerophyllous clades than in rain forest clades, probably due to higher rates of net speciation in the former and extinction in the latter. Although extinction has been prominent in rain forest communities, tropical rain forests appear to have experienced extensive immigration from northern neighbours. Pleistocene climatic oscillations have left genetic signatures at multiple levels of divergence and with complex geographical structuring, even in areas with low topographical relief and few obvious geographical barriers. Main conclusions Our review confirms long‐held views of the ancestral position of the Australian mesic biome but also reveals new insights into the complexity of the processes of contraction, fragmentation, extinction and invasion during the evolution of this biome.     

Contemporary landscape burning patterns in the far North Kimberley region of north-west Australia: human influences and environmental determinants

Aim This study of contemporary landscape burning patterns in the North Kimberley aims to determine the relative influences of environmental factors and compare the management regimes occurring on Aboriginal lands, pastoral leases, national park and crown land. Location The study area is defined at the largest scale by Landsat Scene 108–70 that covers a total land area of 23,134 km² in the North Kimberley Bioregion of north‐west Australia, including the settlement of Kalumburu, coastline between Vansittart Bay in the west and the mouth of the Berkeley River in the east, and stretching approximately 200 km inland. Methods Two approaches are applied. First, a 10‐year fire history (1990–1999) derived from previous study of satellite (Landsat‐MSS) remote sensing imagery is analysed for broad regional patterns. And secondly, a 2‐year ground‐based survey of burning along major access roads leading to an Aboriginal community is used to show fine‐scale burning patterns. anova and multiple regression analyses are used to determine the influence of year, season, geology, tenure, distance from road and distance from settlement on fire patterns. Results Satellite data indicated that an average of 30.8% (±4.4% SEM) of the study area was burnt each year with considerable variability between years. Approximately 56% of the study area was burnt on three or more occasions over the 10‐year period. A slightly higher proportion of burning occurred on average in the late dry season (17.2 ± 3.6%), compared with the early dry season (13.6 ± 3.3%). The highest fire frequency occurred on basalt substrates, on pastoral tenures, and at distances 5–25 km from roads. Three‐way anova demonstrated that geological substrate and land use were the most significant factors influencing fire history, however a range of smaller interactions were also significant. Analysis of road transects, originating from an Aboriginal settlement, showed that the timing of fire and geology type were the most significant factors affecting the pattern of area burnt. Of the total transect area, 28.3 ± 2.9% was burnt annually with peaks in burning occurring into the dry season months of June, August and September. Basalt uplands (81.2%) and lowlands (30.1%) had greater areas burnt than sandstone (12.3%) and sands (17.7%). Main conclusions Anthropogenic firing is constrained by two major environmental determinants; climate and substrate. Seasonal peaks in burning activity in both the early and late dry season relate to periods of optimal fire‐weather conditions. Substrate factors (geology, soils and physiognomy) influence vegetation‐fuel characteristics and the movement of fire in the landscape. Basalt hills overwhelmingly supported the most frequent wildfire regime in the study region because of their undulating topography and relatively fertile soils that support perennial grasslands. Within these spatial and temporal constraints people significantly influenced the frequency and extent of fire in the North Kimberley thus tenure type and associated land uses had a significant influence on fire patterning. Burning activity is high on pastoral lands and along roads and tracks on some tenure types. While the state government uses aerial control burning and legislation to try to restrict burning to the early dry season across all geology types, in practice burning is being conducted across the full duration of the dry season with early dry season burning focused on sandstone and sand substrates and late dry season burning focused on basalt substrates. There is greater seasonal and spatial variation in burning patterns on landscapes managed by Aboriginal people.     

Small snails in a big place: a radiation in the semi‐arid rangelands in northern Australia (Eupulmonata,Camaenidae, Nanotrachia gen. nov.)

Continuing the revision of the Camaenidae in the Australian Monsoon Tropics, we employed comparative analyses of morphological features (with a focus on shell and penial anatomy) and genetic markers (with a focus on mitochondrial COI and 16S sequences) to address the systematic relationships of land snails from the Victoria River District, Northern Territory, and adjacent East Kimberley (Western Australia). These analyses revealed that the species under study represented the previously undescribed genus Nanotrachia. This genus differs from all other camaenid genera known from north‐western Australia most conspicuously by its small, flat, and ribbed shell. Six species are identified as members of the new genus, four of them new species ( Nanotrachia costulata sp. nov. , Nanotrachia carinata sp. nov. , Nanotrachia coronata sp. nov. , Nanotrachia levis sp. nov. ). Two further species have already been described previously but assigned to different genera. These species, Ordtrachia intermedia (as the type species of Nanotrachia) and Mouldingia orientalis, are here transferred to Nanotrachia. Like other camaenids from the Australian Monsoon Tropics, species of Nanotrachia are characterized by essentially allopatric distributions, regional endemism, and a patchy distribution across their range. © 2013 The Linnean Society of London     

Arnhemtrachia ramingining: a new genus and species of land snail from Arnhem Land,Australia (Stylommatophora: Camaenidae)

The new, monotypic camaenid genus Arnhemtrachia is described for the new species Arnhemtrachia ramingining from Ramingining, Arnhem Land, Northern Territory, Australia with additional records from the Gove Peninsula in eastern Arnhem Land. This camaenid is characterised by a combination of distinct morphological features, such as a small, discoidal, keeled shell with regular axial ribs, open umbilicus and simple, rounded aperture as well as a reproductive system with a long epiphallus possessing a long flagellum and absence of a penial sheath, with a flap on the penial wall which transverses longitudinal pilasters, and a long, simple bursa copulatrix. Based on comparative morphology, Arnhemtrachia reveals affinities with Trozena morata Iredale, 1838 from NE Queensland.     

Rapid recovery of mammal fauna in the central Kimberley,northern Australia,following the removal of introduced herbivores

Australia has lost more native mammal species than any other country in the past two centuries, and this record of loss looks likely to worsen over the next few decades. Small‐ to medium‐sized mammals are declining in both distribution and density across large tracts of northern Australia's tropical savannas, including within protected areas. The most likely causes are a combination of changed fire patterns, the impacts of introduced herbivores and predation by feral cats. Here, in contrast to the prevailing trend across northern Australia, we report the recovery of native mammals in response to a large‐scale (>40 000 ha) destocking experiment carried out at Mornington Wildlife Sanctuary in the central Kimberley, north‐west Australia. Following the removal of introduced herbivores from 2004, the species richness and abundance of small native rodents and dasyurids increased significantly across all sampled habitats over the next 3 years. We discuss the implications of these results for guiding land management and applied research to help to reduce the impending risk of mammalian extinctions in northern Australia.     

Patterns of molecular diversity in wild stocks of the redclaw crayfish (Cherax quadricarinatus) from northern Australia and Papua New Guinea: impacts of Plio-Pleistocene landscape evolution

1. Analysis of mitochondrial and nuclear DNA (microsatellites) in 379 individuals, collected from 15 localities in northern Australia and Papua New Guinea (PNG), demonstrated that wild redclaw crayfish ( Cherax quadricarinatus ) populations consist of two highly divergent Australian lineages and two PNG lineages.
2. The disjunction between the two Australian lineages occurs over a distance of approximately 200 km in the south-western corner of the Gulf of Carpenteria. These data conflict with an earlier study that detected no significant differentiation in 23 variable allozyme loci in redclaw sampled from northern Australia, but concur broadly with the previous recognition of two morphologically distinct species ( C. quadricarinatus and C. bicarinatus ) across northern Australia, and a third species in PNG ( C. albertsii ).
3. The inferred timing and patterns of divergence evident in the molecular data presented here closely align with a similar pattern reported in a co-distributed freshwater decapod crustacean, and broadly reflect patterns in some vertebrate taxa with similar distributions across northern Australia and PNG.
4. These congruent patterns most probably reflect periodic Plio-Pleistocene land and freshwater connections between Australia and New Guinea.     

Chloroplast DNA diversity associated with protected slopes and valleys for hybridizing Eucalyptus species on isolated ranges in south‐eastern Australia

Aim To relate genetic diversity to topographic features and to investigate genetic interactions between Eucalyptus species in a local centre of endemism and diversity in south‐eastern Australia. Location Grampian Ranges, Victoria, Australia. Methods We documented chloroplast DNA (cpDNA) variation for a group of endemic Eucalyptus species (E. serraensis, E. verrucata and E. victoriana) that dominate rocky, high‐elevation ridgelines of the Grampian Ranges and for one closely‐related, widespread species (E. baxteri) occupying flanking slopes and valleys. We documented genetic patterns across the landscape using cpDNA microsatellites, and related them to topographic features (exposed west‐facing versus protected east‐facing slopes and valleys). We also determined the extent of local haplotype sharing between populations of endemic species and neighbouring E. baxteri downslope with cpDNA microsatellites, and haplotype sharing between the endemic group and more distantly related species (E. obliqua, E. pauciflora and E. willisii) with sequences of the J_LA+ chloroplast region. Results We detected 26 cpDNA microsatellite haplotypes in a relatively small area of c. 20 km × 50 km. Populations of E. baxteri on east‐facing slopes and valleys had greater cpDNA microsatellite diversity than E. baxteri and endemic species on exposed west‐facing slopes. Endemic species frequently shared chloroplast haplotypes with E. baxteri downslope. Sharing of J_LA+ haplotypes with species outside the endemic group was mostly restricted to E. victoriana, which had cpDNA more similar to the species from other sections of Eucalyptus (E. obliqua, E. willisii and E. pauciflora). Main conclusions Intensive sampling of related species on small isolated mountain ranges allowed us to relate genetic diversity to fine‐scale habitats and to document extensive local haplotype sharing between species. This study contributes to a general understanding of the environmental conditions that enable plant population persistence by linking concentrations of genetic diversity to particular habitats.     

Aboriginal Resource Utilization and Fire Management Practice in Western Arnhem Land, Monsoonal Northern Australia: Notes for Prehistory, Lessons for the Future

This paper considers traditional resources and fire management practices of Aboriginal people living in a near-coastal region of western Arnhem Land, monsoonal northern Australia. The data illustrate that before the arrival of Europeans freshwater floodplains and riverine habitats provided the major proportion of food resources over much of the seasonal cycle. By contrast, the extensive lowland woodlands and open forests, the sparser vegetation of the Arnhem Land escarpment and plateau, and the generally small patches of rain forest (jungle), provided relatively few resources, although jungle yams were of critical importance through the relatively lean wet season. The paper then considers burning as a management tool through the seasonal cycle. In broad terms, burning commenced in the early dry season and was applied systematically and purposefully over the landscape. Burning in the late dry season was undertaken with care, and resumed in earnest with the onset of the first storms of the new wet season, particularly on floodplains. These general patterns of resource use and fire management are shown to have applied widely over much of near-coastal northern Australia. The implications of these data for prehistory and for contemporary land management practices in the region, are considered. It is suggested that pre-European patterns of fire management in the region are likely to have been practiced only over the past few thousand years, given the development of abundant food resources in the late Holocene. It is shown that traditional burning practice offers a generally useful, conservative model for living in and managing a highly fire-prone savanna environment.     

Biogeography and ecology of freshwater diatoms in Subantarctica: a review

Subantarctica is situated between the Antarctic and the Subtropical Convergence and consists of the islands in the southern Atlantic, Indian and Pacific Oceans. Diatoms are an important component of all Subantarctic aquatic, moss and soil habitats. Taxonomic studies reveal a high diversity and species richness in both the present-day and the fossil diatom flora. Planktonic diatoms are almost completely absent. A similarity analysis of the diatom composition from different localities in the southern zone (below 40°S) resulted in a biogeographical zonation. Three regions could be formed, based on their diatom composition: Subantarctica, Maritime Antarctica and the Antarctic Continent. The diatom communities in the different regions are all characterized by a high proportion of cosmopolitan species. A second feature of the southern diatom floras is the decreasing diversity when moving southwards.     

Notes on the vegetation of the Bougainville Peninsula,Osborn and Institut Islands,North Kimberley District,Western Australia

The Bougainville Peninsula is formed of basalt capped with mesaform remnants of a bauxitic duricrust. On the plateau so formed there is little soil and the vegetation comprises scattered stunted eucalypts with grasses and hummock grasses or an understorey of Acacia and other shrubs. Locally there are patches of woodland of Eucalyptus nesophila and E. miniata. On the basalt country below the plateau there is deciduous vine thicket in the less accessible parts, a patchy mixture of vine thicket and savanna elsewhere including communities of vine thicket with scattered emergent eucalypts. The Osborn Islands are similar except that the mesaform capping is generally of sandstone. It is concluded that fire is the principal factor regulating the balance between vine thicket and savanna. Up to 40 years ago the area was populated by aborigines and regularly burnt off but this has now ceased so that colonisation by vine thicket is in progress, in spite of the low rainfall of the area.These authors are indebted to the Amax Exploration Company for providing helicopter transport to the localities concerned and for approval to publish this paper: also to the staff of the Western Australian Herbarium, Mr M. I. Brooker, Mr J. Macorochie and Dr V. Semeniuk for the identification of botanical specimens.This author is indebted to the Department of Defence (Navy Office) for permission to join a naval patrol boat. The assistance of Lieutenants J. Wells, R. Gates and the crew of HMAS Attack is gratefully acknowledged. The co-operation of Mr T. P. Farrell (CRA Services) and the Mitchell Plateau Bauxite Company for providing helicopter transport is also appreciated.     

Biogeography and endemism of ants (Hymenoptera: Formicidae) in Baja California, Mexico: afirst overview

Abstract Aims Ants (Hymenoptera: Formicidae) of the Baja California peninsula are poorly known, with information based largely on scattered museum and literature records. We provide the first comprehensive account of ant species occurring on the peninsula, we examine distribution patterns, and we assess the ‘peninsular effect’ which predicts that species richness declines from the base to the tip of a peninsula. Location Peninsula of Baja California, Mexico. Methods Data collection involved examining, identifying and recording label data from c. 2350 series of ants. These records provide a provisional, if incomplete, species list. We applied the incidence‐based estimator, Chao‐2, to our data base of specimen records to estimate the total number of ant species on the peninsula. We assessed endemism by comparing our peninsular species list to those from adjacent states. The peninsular effect was tested by comparing genus and species level richness between the two states of Baja California, and across five latitudinal blocks. Results We document 170 native ant species in thirty‐three genera, plus six non‐native species, in Baja California. It seems likely that additional species remain to be discovered: the Chao‐2 estimator of species richness, at 206.0 species, is about 20% higher than our observed species richness. About 30% of the species and 20% of the genera are restricted within Baja California to the relatively mesic California Floristic Province of north‐western Baja California. Nearly all of these species also occur in California. Forty‐seven species (27.6%) are peninsula endemics. Using our entire data set, the peninsular effect appears to be strong, with about twice as many species in the northern state of Baja California than are recorded from the southern state of Baja California Sur; the ratio of genera is 33 to 24. However, this effect becomes weak at the species level and absent at the genus level when minimizing habitat effects by omitting species restricted to the California Floristic Province. At a finer scale, across latitudinal blocks of about 1.9°, the number of species declines towards central portions of the peninsula and then increases in the Cape Region. Nine ant species display strongly disjunct distributions, and these occur in two general patterns: peninsula disjuncts and peninsula–mainland disjuncts. Main conclusions The Baja California peninsula supports a diverse and distinctive ant fauna, with the proportion of endemic species similar to that displayed by plants. Patterns of species and genus richness across the five latitudinal blocks provide poor support for the peninsular effect. Moreover, habitat diversity, especially that related to topographic relief, appears to be the most important factor affecting the gradient of ant species richness in Baja California. Additional collections are needed to develop a more complete species list and to determine the boundaries and status of many species. Nevertheless, the present data base provides a useful starting point for understanding the evolution of ant assemblages in Baja California and for comparison with peninsular patterns in other taxa.     

Dry-season Mosquito Breeding Associated with Irrigation in the Northeast Kimberley Region of Western Australia: Potential Impact on Mosquito-borne Disease Transmission

Recent Ross River virus activity prior to the onset of the wet season in the Kimberley region of Western Australia points to an increased dry season mosquito-borne disease risk at Kununurra, compared to other Kimberley towns. This study describes a preliminary investigation into the role of the Ord River Irrigation Area at Kununurra in mosquito production during the dry season. Specifically, the study sought to determine whether the irrigation area has provided the opportunity for year-round breeding of arbovirus vector mosquitoes. A 10-day adult and larval mosquito sampling survey, in August 2003, revealed that mosquito breeding was occurring during the driest month of the year at Kununurra, supporting the hypothesis that mosquitoes can breed year-round. Importantly, significantly larger numbers and percentage of adult Culex annulirostris, an important disease vector in Australia, were collected within the irrigation area (44.6% of total catch) compared to nearby reference (nonirrigated) breeding sites (9.8%) (mean difference 76.2 per trap per night; 95% CI 38.6, 113.7; P <0.001). Larval Cx. annulirostris were also collected at several sites within the irrigation area, whereas none were collected at reference sites. These results indicate that mosquito breeding associated with anthropogenic environmental changes may be responsible for an increased health risk at the end of the dry season. Mosquito management needs to be given a high priority to ensure this potential health risk is not further exacerbated. Several control strategies to reduce breeding of disease vector mosquitoes are identified, primarily focusing on modification of breeding habitats and alteration of irrigation protocols.     

Natural 15N abundance of vegetation and soil in the Kapalga savanna,Australia

Abstract The natural abundance of the stable isotope ¹⁵N was measured in different vegetation components and in the soil of a northern Australian savanna. Most of the vegetation was found to be ¹⁵N-depleted compared to atmospheric N₂. Herbaceous legumes, perennial grasses, tree legumes, non-legume trees and annual grasses exhibited mean δ¹⁵N of ? 1.7, ? 0.8, ? 0.7, 0.0 and + 0.3‰, respectively. These results are in good agreement with previous studies. Legumes exhibit slightly negative values, indicating that they are likely to be nitrogen-fixing plants. Non-legume plants have a δ¹⁵N close to zero, which could equally result from non-symbiotic fixation, soil organic matter mineralization, or fresh root litter mineralization. In contrast, soil organic matter was ¹⁵N-enriched. Values of δ¹⁵N increased with depth and were + 2.5, + 5.2 and +6.1‰ in the 0–10, 10–20 and 20–40cm layers, respectively. Soil organic matter δ¹⁵N shows a typical profile of mature soils.     

The evolution of sexual and parthenogenetic Warramaba: a window onto Plio–Pleistocene diversification processes in an arid biome

Environmental changes over the Plio‐Pleistocene have been key drivers of speciation patterns and genetic diversification in high‐latitude and mesic environments, yet comparatively little is known about the evolutionary history of species in arid environments. We applied phylogenetic and phylogeographic analyses to understand the evolutionary history of Warramaba grasshoppers from the Australian arid zone, a group including sexual and parthenogenetic lineages. Sequence data (mitochondrial COI) showed that the four major sexual lineages within Warramaba most likely diverged in the Pliocene, around 2–7 million years ago. All sexual lineages exhibited considerable phylogenetic structure. Detailed analyses of the hybrid parthenogenetic species W. virgo and its sexual progenitors showed a pattern of high phylogenetic diversity and phylogeographic structure in northern lineages, and low diversity and evidence for recent expansion in southern lineages. Northern sexual lineages persisted in localized refugia over the Pleistocene, with sustained barriers promoting divergence over this period. Southern parts of the present range became periodically unsuitable during the Pleistocene, and it is into this region that parthenogenetic lineages have expanded. Our results strongly parallel those for sexual and parthenogenetic lineages of the gecko Heteronotia from the same region, indicating a highly general effect of Plio‐Pleistocene environmental change on diversification processes in arid Australia.