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 theWet 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.

Evolutionary biology of Gondwanan non-biting midges (Diptera: Chironomidae)

The potential restriction to effective dispersal and gene flow caused by habitat fragmentation can apply to multiple levels of evolutionary scale; from the fragmentation of ancient supercontinents driving diversification and speciation on disjunct landmasses, to the isolation of proximate populations as a result of their inability to cross intervening unsuitable habitat. Investigating the role of habitat fragmentation in driving diversity within and among taxa can thus include inferences of phylogenetic relationships among taxa, assessments of intraspecific phylogeographic structure and analyses of gene flow among neighbouring populations. The proposed Gondwanan clade within the chironomid (non-biting midge) subfamily Orthocladiinae (Diptera: Chironomidae) represents a model system for investigating the role that population fragmentation and isolation has played at different evolutionary scales. A pilot study by Krosch et al (2009) indentified several highly divergent lineages restricted to ancient rainforest refugia and limited gene flow among proximate sites within a refuge for one member of this clade, Echinocladius martini Cranston. This study provided a framework for investigating the evolutionary history of this taxon and its relatives more thoroughly. Populations of E. martini were sampled in the Paluma bioregion of northeast Queensland to investigate patterns of fine-scale within- and among-stream dispersal and gene flow within a refuge more rigorously. Data was incorporated from Krosch et al (2009) and additional sites were sampled up- and downstream of the original sites. Analyses of genetic structure revealed strong natal site fidelity and high genetic structure among geographically proximate streams. Little evidence was found for regular headwater exchange among upstream sites, but there was distinct evidence for rare adult flight among sites on separate stream reaches. Overall, however, the distribution of shared haplotypes implied that both larval and adult dispersal was largely limited to the natal stream channel. Patterns of regional phylogeographic structure were examined in two related austral orthoclad taxa – Naonella forsythi Boothroyd from New Zealand and Ferringtonia patagonica Sæther and Andersen from southern South America – to provide a comparison with patterns revealed in their close relative E. martini. Both taxa inhabit tectonically active areas of the southern hemisphere that have also experienced several glaciation events throughout the Plio-Pleistocene that are thought to have affected population structure dramatically in many taxa. Four highly divergent lineages estimated to have diverged since the late Miocene were revealed in each taxon, mirroring patterns in E. martini; however, there was no evidence for local geographical endemism, implying substantial range expansion post-diversification. The differences in pattern evident among the three related taxa were suggested to have been influenced by variation in the responses of closed forest habitat to climatic fluctuations during interglacial periods across the three landmasses. Phylogeographic structure in E. martini was resolved at a continental scale by expanding upon the sampling design of Krosch et al (2009) to encompass populations in southeast Queensland, New South Wales and Victoria. Patterns of phylogeographic structure were consistent with expectations and several previously unrecognised lineages were revealed from central- and southern Australia that were geographically endemic to closed forest refugia. Estimated divergence times were congruent with the timing of Plio-Pleistocene rainforest contractions across the east coast of Australia. This suggested that dispersal and gene flow of E. martini among isolated refugia was highly restricted and that this taxon was susceptible to the impacts of habitat change. Broader phylogenetic relationships among taxa considered to be members of this Gondwanan orthoclad group were resolved in order to test expected patterns of evolutionary affinities across the austral continents. The inferred phylogeny and estimated divergence times did not accord with expected patterns based on the geological sequence of break-up of the Gondwanan supercontinent and implied instead several transoceanic dispersal events post-vicariance. Difficulties in appropriate taxonomic sampling and accurate calibration of molecular phylogenies notwithstanding, the sampling regime implemented in the current study has been the most intensive yet performed for austral members of the Orthocladiinae and unsurprisingly has revealed both novel taxa and phylogenetic relationships within and among described genera. Several novel associations between life stages are made here for both described and previously unknown taxa. Investigating evolutionary relationships within and among members of this clade of proposed Gondwanan orthoclad taxa has demonstrated that a complex interaction between historical population fragmentation and dispersal at several levels of evolutionary scale has been important in driving diversification in this group. While interruptions to migration, colonisation and gene flow driven by population fragmentation have clearly contributed to the development and maintenance of much of the diversity present in this group, long-distance dispersal has also played a role in influencing diversification of continental biotas and facilitating gene flow among disjunct populations.

Limestone and speleothem trace element geochemistry as tools for palaeoclimatic reconstruction, Mount Etna region, central-coastal Queensland

This study investigated potential palaeoclimate proxies provided by rare earth element (REE) geochemistry in speleothems and in clay mineralogy of cave sediments. Speleothem and sediment samples were collected from a series of cave fill deposits that occurred with rich vertebrate fossil assemblages in and around Mount Etna National Park, Rockhampton (central coastal Queensland). The fossil deposits range from Plio- Pleistocene to Holocene in age (based on uranium/thorium dating) and appear to represent depositional environments ranging from enclosed rainforest to semi-arid grasslands. Therefore, the Mount Etna cave deposits offer the perfect opportunity to test new palaeoclimate tools as they include deposits that span a known significant climate shift on the basis of independent faunal data. The first section of this study investigates the REE distribution of the host limestone to provide baseline geochemistry for subsequent speleothem investigations. The Devonian Mount Etna Beds were found to be more complex than previous literature had documented. The studied limestone massif is overturned, highly recrystallised in parts and consists of numerous allochthonous blocks with different spatial orientations. Despite the complex geologic history of the Mount Etna Beds, Devonian seawater-like REE patterns were recovered in some parts of the limestone and baseline geochemistry was determined for the bulk limestone for comparison with speleothem REE patterns. The second part of the study focused on REE distribution in the karst system and the palaeoclimatic implications of such records. It was found that REEs have a high affinity for calcite surfaces and that REE distributions in speleothems vary between growth bands much more than along growth bands, thus providing a temporal record that may relate to environmental changes. The morphology of different speleothems (i.e., stalactites, stalagmites, and flowstones) has little bearing on REE distributions provided they are not contaminated with particulate fines. Thus, baseline knowledge developed in the study suggested that speleothems were basically comparable for assessing palaeoclimatically controlled variations in REE distributions. Speleothems from rainforest and semi-arid phases were compared and it was found that there are definable differences in REE distribution that can be attributed to climate. In particular during semiarid phases, total REE concentration decreased, LREE became more depleted, Y/Ho increased, La anomalies were more positive and Ce anomalies were more negative. This may reflect more soil development during rainforest phases and more organic particles and colloids, which are known to transport REEs, in karst waters. However, on a finer temporal scale (i.e. growth bands) within speleothems from the same climate regime, no difference was seen. It is suggested that this may be due to inadequate time for soil development changes on the time frames represented by differences in growth band density. The third part of the study was a reconnaissance investigation focused on mineralogy of clay cave sediments, illite/kaolinite ratios in particular, and the potential palaeoclimatic implications of such records. Although the sample distribution was not optimal, the preliminary results suggest that the illite/kaolinite ratio increased during cold and dry intervals, consistent with decreased chemical weathering during those times. The study provides a basic framework for future studies at differing latitudes to further constrain the parameters of the proxy. The identification of such a proxy recorded in cave sediment has broad implications as clay ratios could potentially provide a basic local climate proxy in the absence of fossil faunas and speleothem material. This study suggests that REEs distributed in speleothems may provide information about water throughput and soil formation, thus providing a potential palaeoclimate proxy. It highlights the importance of understanding the host limestone geochemistry and broadens the distribution and potential number of cave field sites as palaeoclimate information no longer relies solely on the presence of fossil faunas and or speleothems. However, additional research is required to better understand the temporal scales required for the proxies to be recognised.

The Palu Metamorphic Complex, NW Sulawesi, Indonesia: Origin and evolution of a young metamorphic terrane with links to Gondwana and Sundaland

The Palu Metamorphic Complex (PMC) is exposed in a late Cenozoic orogenic belt in NW Sulawesi, Indonesia. It is a composite terrane comprising a gneiss unit of Gondwana origin, a schist unit composed of meta-sediments deposited along the SE Sundaland margin in the Late Cretaceous and Early Tertiary, and one or more slivers of amphibolite with oceanic crust characteristics. The gneiss unit forms part of the West Sulawesi block underlying the northern and central sections of the Western Sulawesi Province. The presence of Late Triassic granitoids and recycled Proterozoic zircons in this unit combined with its isotopic signature suggests that the West Sulawesi block has its origin in the New Guinea margin from which it rifted in the late Mesozoic. It docked with Sundaland sometime during the Late Cretaceous. U–Th–Pb dating results for monazite suggest that another continental fragment may have collided with the Sundaland margin in the earliest Miocene. High-pressure (HP) and ultrahigh-pressure (UHP) rocks (granulite, peridotite, eclogite) are found as tectonic slices within the PMC, mostly along the Palu–Koro Fault Zone, a major strike-slip fault that cuts the complex. Mineralogical and textural features suggest that some of these rocks resided at depths of 60–120 km during a part of their histories. Thermochronological data (U–Th–Pb zircon and 40Ar/39Ar) from the metamorphic rocks indicate a latest Miocene to mid-Pliocene metamorphic event, which was accompanied by widespread granitoid magmatism and took place in an extensional tectonic setting. It caused recrystallization of, and new overgrowths on, pre-existing zircon crystals, and produced andalusite–cordierite–sillimanite–staurolite assemblages in pelitic protoliths, indicating HT–LP (Buchan-type) metamorphism. The PMC was exhumed as a core complex at moderate rates (c. 0.7–1.0 mm/yr) accompanied by rapid cooling in the Plio-Pleistocene. Some of the UHP rocks were transported to the surface at significantly higher rates (⩾16 mm/yr). The results of our study do not support recent plate tectonic reconstructions that propose a NW Australia margin origin for the West Sulawesi block (e.g. Hall et al., 2009).

Pleistocene climate fluctuations influence phylogeographical patterns in Melomys cervinipes across the mesic forests of eastern Australia

Aim Our aim was to clarify the lineage-level relationships for Melomys cervinipes and its close relatives and investigate whether the patterns of divergence observed for these wet-forest-restricted mammals may be associated with recognized biogeographical barriers. Location Mesic closed forest along the east coast of Australia, from north Queensland to mid-eastern New South Wales. Methods To enable rigorous phylogenetic reconstruction, divergence-date estimation and phylogeographical inference, we analysed DNA sequence and microsatellite data from 307 specimens across the complete distribution of M. cervinipes (45 localities). Results Three divergent genetic lineages were found within M. cervinipes, corresponding to geographically delineated northern, central and southern clades. Additionally, a fourth lineage, comprising M. rubicola and M. capensis, was identified and was most closely related to the northern M. cervinipes lineage. Secondary contact of the northern and central lineages was identified at one locality to the north of the Burdekin Gap. Main conclusions Contemporary processes of repeated habitat fragmentation and contraction, local extinction events and subsequent re-expansion across both small and large areas, coupled with the historical influence of the Brisbane Valley Barrier, the St Lawrence Gap and the Burdekin Gap, have contributed to the present phylogeographical structure within M. cervinipes. Our study highlights the need to sample close to the periphery of putative biogeographical barriers or risk missing vital phylogeographical information that may significantly alter the interpretation of biogeographical hypotheses.

Late Pleistocene stratigraphy of IODP Site U1396 and compiled chronology offshore of south and south west Montserrat, Lesser Antilles

Marine sediments around volcanic islands contain an archive of volcaniclastic deposits, which can be used to reconstruct the volcanic history of an area. Such records hold many advantages over often incomplete terrestrial datasets. This includes the potential for precise and continuous dating of intervening sediment packages, which allow a correlatable and temporally-constrained stratigraphic framework to be constructed across multiple marine sediment cores. Here, we discuss a marine record of eruptive and mass-wasting events spanning ~250 ka offshore of Montserrat, using new data from IODP Expedition 340, as well as previously collected cores. By using a combination of high-resolution oxygen isotope stratigraphy, AMS radiocarbon dating, biostratigraphy of foraminifera and calcareous nannofossils and clast componentry, we identify five major events at Soufriere Hills volcano since 250 ka. Lateral correlation of these events across sediment cores collected offshore of the south and south west of Montserrat, have improved our understanding of the timing, extent and associations between events in this area. Correlations reveal that powerful and potentially erosive density-currents travelled at least 33 km offshore, and demonstrate that marine deposits, produced by eruption-fed and mass-wasting events on volcanic islands, are heterogeneous in their spatial distribution. Thus, multiple drilling/coring sites are needed to reconstruct the full chronostratigraphy of volcanic islands. This multidisciplinary study will be vital to interpreting the chaotic records of submarine landslides at other sites drilled during Expedition 340 and provides a framework that can be applied to the stratigraphic analysis of sediments surrounding other volcanic islands.

Prehistoric hand stencils at Fern Cave, North Queensland (Australia): environmental and chronological implications of Raman spectroscopy and FT-IR imaging results

Raman spectroscopy and FT-IR imaging analyses of cave wall pigment samples from north Queensland (Australia) indicate that some hand stencils were undertaken during a dry environmental phase indicating late Holocene age. Other, earlier painting episodes also took place during dry environmental periods of the terminal Pleistocene and/or early Holocene. These results represent a rare opportunity to attain chronological information for rock art in conditions where insufficient carbon is present for radiocarbon dating.

Palaeoecology of the Mount Etna bat fauna, coastal Eastern Queensland

Global warming is already threatening many animal and plant communities worldwide, however, the effect of climate change on bat populations is poorly known. Understanding the factors influencing the survival of bats is crucial to their conservation, and this cannot be achieved solely by modern ecological studies. Palaeoecological investigations provide a perspective over a much longer temporal scale, allowing the understanding of the dynamic patterns that shaped the distribution of modern taxa. In this study twelve microchiropteran fossil assemblages from Mount Etna, central-eastern Queensland, ranging in age from more than 500,000 years to the present day, were investigated. The aim was to assess the responses of insectivorous bats to Quaternary environmental changes, including climatic fluctuations and recent anthropogenic impacts. In particular, this investigation focussed on the effects of increasing late Pleistocene aridity, the subsequent retraction of rainforest habitat, and the impact of cave mining following European settlement at Mount Etna. A thorough examination of the dental morphology of all available extant Australian bat taxa was conducted in order to identify the fossil taxa prior to their analysis in term of species richness and composition. This detailed odontological work provided new diagnostic dental characters for eighteen species and one genus. It also provided additional useful dental characters for three species and seven genera. This odontological analysis allowed the identification of fifteen fossil bat taxa from the Mount Etna deposits, all being representatives of extant bats, and included ten taxa identified to the species level (i.e., Macroderma gigas, Hipposideros semoni, Rhinolophus megaphyllus, Miniopterus schreibersii, Miniopterus australis, Scoteanax rueppellii, Chalinolobus gouldii, Chalinolobus dwyeri, Chalinolobus nigrogriseus and Vespadelus troughtoni) and five taxa identified to the generic level (i.e., Mormopterus, Taphozous, Nyctophilus, Scotorepens and Vespadelus). Palaeoecological analysis of the fossil taxa revealed that, unlike the non-volant mammal taxa, bats have remained essentially stable in terms of species diversity and community membership between the mid-Pleistocene rainforest habitat and the mesic habitat that occurs today in the region. The single major exception is Hipposideros semoni, which went locally extinct at Mount Etna. Additionally, while intensive mining operations resulted in the abandonment of at least one cave that served as a maternity roost in the recent past, the diversity of the Mount Etna bat fauna has not declined since European colonisation. The overall resilience through time of the bat species discussed herein is perhaps due to their unique ecological, behavioural, and physiological characteristics as well as their ability to fly, which have allowed them to successfully adapt to their changing environment. This study highlights the importance of palaeoecological analyses as a tool to gain an understanding of how bats have responded to environmental change in the past and provides valuable information for the conservation of threatened modern species, such as H. semoni.