Nick Porch

Birth of a biome: insights into the assembly and maintenance of the Australian arid zone biota

The integration of phylogenetics, phylogeography and palaeoenvironmental studies is providing major insights into the historical forces that have shaped the Earths biomes. Yet our present view is biased towards arctic and temperate/tropical forest regions, with very little focus on the extensive arid regions of the planet. The Australian arid zone is one of the largest desert landform systems in the world, with a unique, diverse and relatively well-studied biota. With foci on palaeoenvironmental and molecular data, we here review what is known about the assembly and maintenance of this biome in the context of its physical history, and in comparison with other mesic biomes. Aridification of Australia began in the Mid-Miocene, around 15 million years, but fully arid landforms in central Australia appeared much later, around 1-4 million years. Dated molecular phylogenies of diverse taxa show the deepest divergences of arid-adapted taxa from the Mid-Miocene, consistent with the onset of desiccation. There is evidence of arid-adapted taxa evolving from mesic-adapted ancestors, and also of speciation within the arid zone. There is no evidence for an increase in speciation rate during the Pleistocene, and most arid-zone species lineages date to the Pliocene or earlier. The last 0.8 million years have seen major fluctuations of the arid zone, with large areas covered by mobile sand dunes during glacial maxima. Some large, vagile taxa show patterns of recent expansion and migration throughout the arid zone, in parallel with the ice sheet-imposed range shifts in Northern Hemisphere taxa. Yet other taxa show high lineage diversity and strong phylogeographical structure, indicating persistence in multiple localised refugia over several glacial maxima. Similar to the Northern Hemisphere, Pleistocene range shifts have produced suture zones, creating the opportunity for diversification and speciation through hybridisation, polyploidy and parthenogenesis. This review highlights the opportunities that development of arid conditions provides for rapid and diverse evolutionary radiations, and re-enforces the emerging view that Pleistocene environmental change can have diverse impacts on genetic structure and diversity in different biomes. There is a clear need for more detailed and targeted phylogeographical studies of Australias arid biota and we suggest a framework and a set of a priori hypotheses by which to proceed.

Ecological niche modelling and nDNA sequencing support a new, morphologically cryptic beetle species unveiled by DNA barcoding.

Background DNA sequencing techniques used to estimate biodiversity, such as DNA barcoding, may reveal cryptic species. However, disagreements between barcoding and morphological data have already led to controversy. Species delimitation should therefore not be based on mtDNA alone. Here, we explore the use of nDNA and bioclimatic modelling in a new species of aquatic beetle revealed by mtDNA sequence data. Methodology/Principal Findings The aquatic beetle fauna of Australia is characterised by high degrees of endemism, including local radiations such as the genus Antiporus. Antiporus femoralis was previously considered to exist in two disjunct, but morphologically indistinguishable populations in south-western and south-eastern Australia. We constructed a phylogeny of Antiporus and detected a deep split between these populations. Diagnostic characters from the highly variable nuclear protein encoding arginine kinase gene confirmed the presence of two isolated populations. We then used ecological niche modelling to examine the climatic niche characteristics of the two populations. All results support the status of the two populations as distinct species. We describe the south-western species as Antiporus occidentalis sp.n. Conclusion/Significance In addition to nDNA sequence data and extended use of mitochondrial sequences, ecological niche modelling has great potential for delineating morphologically cryptic species.

Pliocene reversal of late Neogene aridification.

Significance The warm climates of the Pliocene epoch are considered our best analog for a future anthropogenic greenhouse world. However, understanding of the nature of Pliocene climate variability and change on land is currently limited by the poor age control of most existing terrestrial climate archives. We present a radiometrically dated history of the evolution of Southern Hemisphere vegetation and hydroclimate from the latest Miocene to the middle Pliocene. These data reveal a sharp increase in precipitation in the Early Pliocene, which drove complete vegetation turnover. The development of warm, wet early Pliocene climates clearly reversed a long-term Southern Hemisphere trend of late Neogene cooling and aridification, highlighting the question of what initiated this sustained, ∼1.5-My-long interval of warmth. The Pliocene epoch (5.3–2.6 Ma) represents the most recent geological interval in which global temperatures were several degrees warmer than today and is therefore considered our best analog for a future anthropogenic greenhouse world. However, our understanding of Pliocene climates is limited by poor age control on existing terrestrial climate archives, especially in the Southern Hemisphere, and by persistent disagreement between paleo-data and models concerning the magnitude of regional warming and/or wetting that occurred in response to increased greenhouse forcing. To address these problems, here we document the evolution of Southern Hemisphere hydroclimate from the latest Miocene to the middle Pliocene using radiometrically-dated fossil pollen records preserved in speleothems from semiarid southern Australia. These data reveal an abrupt onset of warm and wet climates early within the Pliocene, driving complete biome turnover. Pliocene warmth thus clearly represents a discrete interval which reversed a long-term trend of late Neogene cooling and aridification, rather than being simply the most recent period of greater-than-modern warmth within a continuously cooling trajectory. These findings demonstrate the importance of high-resolution chronologies to accompany paleoclimate data and also highlight the question of what initiated the sustained interval of Pliocene warmth.

Unveiling the Diversification Dynamics of Australasian Predaceous Diving Beetles in the Cenozoic

During the Cenozoic, Australia experienced major climatic shifts that have had dramatic ecological consequences for the modern biota. Mesic tropical ecosystems were progressively restricted to the coasts and replaced by arid-adapted floral and faunal communities. Whilst the role of aridification has been investigated in a wide range of terrestrial lineages, the response of freshwater clades remains poorly investigated. To gain insights into the diversification processes underlying a freshwater radiation, we studied the evolutionary history of the Australasian predaceous diving beetles of the tribe Hydroporini (147 described species). We used an integrative approach including the latest methods in phylogenetics, divergence time estimation, ancestral character state reconstruction, and likelihood-based methods of diversification rate estimation. Phylogenies and dating analyses were reconstructed with molecular data from seven genes (mitochondrial and nuclear) for 117 species (plus 12 outgroups). Robust and well-resolved phylogenies indicate a late Oligocene origin of Australasian Hydroporini. Biogeographic analyses suggest an origin in the East Coast region of Australia, and a dynamic biogeographic scenario implying dispersal events. The group successfully colonized the tropical coastal regions carved by a rampant desertification, and also colonized groundwater ecosystems in Central Australia. Diversification rate analyses suggest that the ongoing aridification of Australia initiated in the Miocene contributed to a major wave of extinctions since the late Pliocene probably attributable to an increasing aridity, range contractions and seasonally disruptions resulting from Quaternary climatic changes. When comparing subterranean and epigean genera, our results show that contrasting mechanisms drove their diversification and therefore current diversity pattern. The Australasian Hydroporini radiation reflects a combination of processes that promoted both diversification, resulting from new ecological opportunities driven by initial aridification, and a subsequent loss of mesic adapted diversity due to increasing aridity.

Tasmania: archaeological and palaeo-ecological perspectives

Tasmania, at the south of the land-mass, experienced the Glacial Maximum as a properly cold affair. Recent archaeological work, some in country now difficult of human access, has developed an intricate story of changing adaptations. At the Pleistocene-Holocene boundary, a major reorganization of Aboriginal adaptation strategies is seen in the archaeological record, argued to follow late-Pleistocene environmental amelioration.

Mid- to late Holocene landscape change and anthropogenic transformations on Mo'orea, Society Islands: A multi-proxy approach

Archaeology’s ability to generate long-term datasets of natural and human landscape change positions the discipline as an inter-disciplinary bridge between the social and natural sciences. Using a multi-proxy approach combining archaeological data with palaeoenvironmental indicators embedded in coastal sediments, we outline millennial timescales of lowland landscape evolution in the Society Islands. Geomorphic and cultural histories for four coastal zones on Mo‘orea are reconstructed based on stratigraphic records, sedimentology, pollen analysis, and radiocarbon determinations from mid- to late Holocene contexts. Prehuman records of the island’s flora and fauna are described utilizing landsnail, insect, and botanical data, providing a palaeo-backdrop for later anthropogenic change. Several environmental processes, including sea level change, island subsidence, and anthropogenic alterations, leading to changes in sedimentary budget have operated on Mo‘orea coastlines from c. 4600 to 200 BP. We document significant transformation of littoral and lowland zones which obscured earlier human activities and created significant changes in vegetation and other biota. Beginning as early as 440 BP (1416–1490 cal. ad), a major phase of sedimentary deposition commenced which can only be attributed to anthropogenic effects. At several sites, between 1.8 and 3.0 m of terrigenous sediments accumulated within a span of two to three centuries due to active slope erosion and deposition on the coastal flats. This phase correlates with the period of major inland expansion of Polynesian occupation and intensive agriculture on the island, indicated by the presence of charcoal throughout the sediments, including wood charcoal from several economically important tree species.

The Early Shorebird Will Catch Fewer Invertebrates on Trampled Sandy Beaches

Many species of birds breeding on ocean beaches and in coastal dunes are of global conservation concern. Most of these species rely on invertebrates (e.g. insects, small crustaceans) as an irreplaceable food source, foraging primarily around the strandline on the upper beach near the dunes. Sandy beaches are also prime sites for human recreation, which impacts these food resources via negative trampling effects. We quantified acute trampling impacts on assemblages of upper shore invertebrates in a controlled experiment over a range of foot traffic intensities (up to 56 steps per square metre) on a temperate beach in Victoria, Australia. Trampling significantly altered assemblage structure (species composition and density) and was correlated with significant declines in invertebrate abundance and species richness. Trampling effects were strongest for rare species. In heavily trafficked plots the abundance of sand hoppers (Amphipoda), a principal prey item of threatened Hooded Plovers breeding on this beach, was halved. In contrast to the consistently strong effects of trampling, natural habitat attributes (e.g. sediment grain size, compactness) were much less influential predictors. If acute suppression of invertebrates caused by trampling, as demonstrated here, is more widespread on beaches it may constitute a significant threat to endangered vertebrates reliant on these invertebrates. This calls for a re-thinking of conservation actions by considering active management of food resources, possibly through enhancement of wrack or direct augmentation of prey items to breeding territories.

Evolutionary history of a secondary terrestrial Australian diving beetle (Coleoptera, Dytiscidae) reveals a lineage of high morphological and ecological plasticity

The evolution of a secondary terrestrial lifestyle in diving beetles (Coleoptera: Dytiscidae) has never been analysed in a phylogenetic framework before. Here we study Terradessus caecus Watts, a terrestrial species of the subfamily Hydroporinae endemic to Australia. We infer its phylogenetic placement using Bayesian inference and maximum‐likelihood methods based on a multilocus molecular dataset. We also investigate the divergence time estimates of this lineage using a Bayesian relaxed clock approach. Finally, we infer ancestral ecological preferences using a likelihood approach. We recover T. caecus nested in the genus Paroster Sharp with strong support. Therefore, we establish a synonymy for both species of Terradessus with Paroster: Paroster caecus (Watts) n.comb. and Paroster anophthalmus (Brancucci & Monteith) n.comb. Paroster is an endemic Australian genus that has a remarkable number of subterranean species in underground aquifers with highly derived morphologies. Our results highlight one of the most remarkable radiations of aquatic beetles with independent ecological pathways likely linked to palaeoclimatic disruptions in the Neogene. Paroster caecus (Watts) n.comb. originated from a mid‐Miocene split following the onset of an aridification episode that has been ongoing to the present day. The deep changes in ecological communities in association with the drying‐out of palaeodrainage systems might have pushed this lineage to colonize a new niche in terrestrial habitats.

Reassembling a lost lowland carabid beetle assemblage (Coleoptera) from Kauai, Hawaiian Islands

Abstract. A late Holocene but prehistoric carabid beetle fauna from the lowland Makauwahi Cave, Kauai, is characterised. Seven extinct species – Blackburnia burneyi, B. cryptipes, B. godzilla, B. menehune, B. mothra, B. ovata and B. rugosa, spp. nov. (tribe Platynini) – represent the first Hawaiian insect species to be newly described from subfossil specimens. Four extant Blackburnia spp. – B. aterrima (Sharp), B. bryophila Liebherr, B. pavida (Sharp), and B. posticata (Sharp) – and three extant species of tribe Bembidiini – Bembidion ignicola Blackburn, B. pacificum Sharp and Tachys oahuensis Blackburn – are also represented. All subfossil fragments are disarticulated, with physical dimensions and cladistic analysis used to associate the major somites – head, prothorax and elytra – for description of the new species. The seven new Makauwahi Cave species support recognition of a lowland area of endemism adjoining Haupu, a low-stature 700 m elevation ridgeline in southern Kauai. Four of the extinct Blackburnia are adelphotaxa to extant species currently found at higher elevations in Kauai. Addition of these lowland specialists to the phylogenetic hypothesis undercuts applicability of the taxon cycle for interpreting evolutionary history of these taxa. Two of the extinct species are Kauai representatives in clades that subsequently colonised younger Hawaiian Islands, enhancing support for the progressive biogeographic colonisation of the archipelago by this lineage. And three of the extinct Blackburnia species comprised larger beetles than those of any extant Kauai Blackburnia, consistent with the evolution of island gigantism in the lowland habitats of Kauai.

STRATIGRAPHY AND CHRONOLOGY OF KARST FEATURES ON RODRIGUES ISLAND, SOUTHWESTERN INDIAN OCEAN

6 Abstract: The remote Indian Ocean island of Rodrigues, while largely of volcanic origin, also contains a large body of eolian calcarenite with over thirty surveyed caves and many other karst features. Little is known, however, regarding the age and stratigraphy of the clastic deposits in the caves and the associated fossils of the highly endemic, now mostly extinct, fauna. On the Plaine Caverne and Plaine Corail of the southwestern part of the island, we obtained sediment cores up to 10 m in length and excavated bones of the extinct fauna from caves in the vicinity. Stratigraphic description and radiocarbon dating revealed that sediments in Canyon Tiyel, a collapsed-cave feature, primarily accumulated during the early and middle Holocene. Sedimentation in the canyon and adjacent caves has slowed in recent millennia, with the result that many bones of fauna that went extinct after human arrival in recent centuries are on or near the surface. The chemistry of the sediments and the alternate wet and dry regime of the cave and canyon surfaces are often not conducive to preservation of bone collagen and plant microfossils. Grotte Fougere, with an apparently unique anchialine pond inside a collapsing cave, however, contains over one meter of highly organic sediment with excellent preservation of plant and animal remains.