Historical transoceanic dispersal of a freshwater shrimp: the colonization of the South Pacific by the genus Paratya (Atyidae)

Aim To infer the phylogenetic relationships within the freshwater shrimp genus Paratya Miers, 1882 (Atyidae) and to use these data to answer biogeographical questions about the location, timing and form of evolution of this genus in the South Pacific. Location Paratya are spread throughout various freshwater habitats in the western Pacific, with a disjunct northern range in the North Pacific (Japan, Korea, Ryukyu Islands, Siberia) and South Pacific (Australia, New Zealand, New Caledonia, Lord Howe, Norfolk Island). Methods Specimens were obtained from throughout its range. Mitochondrial sequences of cytochrome oxidase subunit I and 16S ribosomal DNA were analysed using phylogenetic techniques to identify whether landmasses are monophyletic and what the relationships are between landmasses. Molecular clock dating methods were used to date divergences between taxa. Results Each landmass was recovered as monophyletic. Japan/Ryukyu Islands is the most basal group, followed by New Zealand. Australian specimens form a sister group to a clade made up of two groups (New Caledonia and Lord Howe/Norfolk Island). The oldest divergence within the genus (between North and South Pacific) took place 12–19 Ma. Main conclusions The geographical origin of the genus (either Gondwana or Laurasia) is unclear. Dispersal occurred between the North and South Pacific long after the split up of Gondwana. Dispersal likely explains the presence of Paratya on each landmass in the South Pacific, from continent to isolated oceanic island. This dispersal is conjectured to have taken place through oceanic currents because of the amphidromous life cycle of some taxa of Paratya, given that amphyidromy is plesiomorphic in atyid shrimp.     

Molecular phylogeny and biogeography of spring-associated hydrobiid snails of the Great Artesian Basin, Australia

The Great Artesian Basin (GAB) of Australia underlies some of the driest parts of South Australia and Queensland and feeds numerous freshwater springs. Prominent and endangered components of the GAB spring community are snails of the family Hydrobiidae. This paper examines the evolutionary relationships of the entire hydrobiid fauna associated with the GAB, and includes appropriate non-GAB species to place the GAB fauna in a broader phylogenetic context. The Queensland genus Jardinella is a focus of this paper, providing a fine scale examination of relationships between spring supergroups in the northeastern regions of the GAB. Maximum parsimony and Bayesian analyses performed on 16S, CO1, and combined sequence data from 40 hydrobiid taxa found four major clades of Australian taxa. The analysis revealed that at least three separate colonization events of the GAB spring fauna have occurred. Two of these are represented by considerable radiations, (1) Jardinella to the north and east and (2) Caldicochlea, Fonscochlea, and possibly Trochidrobia in South Australia. The phylogenetic position of the latter is uncertain so it may represent yet another invasion. The third definite invasion is represented by a single species of the speciose SE Australian genus Austropyrgus in the Dalhousie Springs in South Australia. Jardinella is found to be monophyletic, and with one exception, its members in each of the Queensland spring supergroups are found to be monophyletic.     

Examining the phylogeny of the Australasian Lymnaeidae (Heterobranchia: Pulmonata: Gastropoda) using mitochondrial, nuclear and morphological markers

We examined the species groups relationships of the freshwater snail genus Austropeplea using mitochondrial, nuclear and morphological markers in addition to traditional methods of shell shape analysis. Based primarily on the results of a combined molecular and morphological analysis, samples of the nominal species A. tomentosa form distinct lineages. The New Zealand populations of A. tomentosa are a very distinct lineage from any of the Australian populations attributed to A. tomentosa. Furthermore, within the Australian group, three lineages, south Australia, Tasmania and eastern Australia, appear to have undergone recent and/or rapid speciation events. Samples assigned to A. lessoni were resolved as two distinct lineages, representing the eastern and northern Australian populations. Kutikina hispida was resolved within the Australian A. tomentosa clade. Molecular results for A. viridis suggests that it is also composed of at least two distinct lineages that could be treated as species. Incongruence observed between the single mitochondrial, nuclear and morphological topologies highlight the importance of using a number of different datasets in the delimitation of species-group taxa.     

Molecular phylogeny and zoogeography of the freshwater crayfish genus Cherax Erichson (Decapoda: Parastacidae) in Australia

The evolutionary history and biogeography of freshwater-dependent taxa in Australia is of intrinsic interest given the present-day aridity of this continent. Cherax is the most widespread and one of the most species-rich of Australia's nine freshwater crayfish genera. The phylogenetic relationships amongst 19 of the 23 Australian Cherax were established from mitochondrial DNA sequences representing the 12S rRNA and 16S rRNA gene regions. The relationships among species support an initial east–west separation, followed by a north–south divergence in eastern Australia. Molecular clock estimations suggest that these divergences date back to the Miocene. The phylogenetic relationships support endemic speciation within geographical regions and indicate that long-distance dispersal has not led to recent speciation as previously hypothesized. This new evolutionary scenario is consistent with the climatic history of Australia and the evolutionary history of other similarly distributed freshwater-dependent organisms in Australia.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 553–563.     

Phylogeny and historical biogeography of ancient assassin spiders (Araneae: Archaeidae) in the Australian mesic zone: evidence for Miocene speciation within Tertiary refugia

The rainforests, wet sclerophyll forests and temperate heathlands of the Australian mesic zone are home to a diverse and highly endemic biota, including numerous old endemic lineages restricted to refugial, mesic biomes. A growing number of phylogeographic studies have attempted to explain the origins and diversification of the Australian mesic zone biota, in order to test and better understand the mode and tempo of historical speciation within Australia. Assassin spiders (family Archaeidae) are a lineage of iconic araneomorph spiders, characterised by their antiquity, remarkable morphology and relictual biogeography on the southern continents. The Australian assassin spider fauna is characterised by a high diversity of allopatric species, many of which are restricted to individual mountains or montane systems, and all of which are closely tied to mesic and/or refugial habitats in the east and extreme south-west of mainland Australia. We tested the phylogeny and vicariant biogeography of the Australian Archaeidae (genus Austrarchaea Forster & Platnick), using a multi-locus molecular approach. Fragments from six mitochondrial genes (COI, COII, tRNA-K, tRNA-D, ATP8, ATP6) and one nuclear protein-coding gene (Histone H3) were used to infer phylogenetic relationships and to explore the phylogeographic origins of the diverse Australian fauna. Bayesian analyses of the complete molecular dataset, along with differentially-partitioned Bayesian and parsimony analyses of a smaller concatenated dataset, revealed the presence of three major Australian lineages, each with non-overlapping distributions in north-eastern Queensland, mid-eastern Australia and southern Australia, respectively. Divergence date estimation using mitochondrial data and a rate-calibrated relaxed molecular clock revealed that major lineages diverged in the early Tertiary period, prior to the final rifting of Australia from East Antarctica. Subsequent speciation occurred during the Miocene (23-5.3 million years ago), with tropical and subtropical taxa diverging in the early-mid Miocene, prior to southern and temperate taxa in the mid-late Miocene. Area cladograms reconciled with Bayesian chronograms for all known Archaeidae in southern and south-eastern Australia revealed seven potentially vicariant biogeographic barriers in eastern Queensland, New South Wales and southern Australia, each proposed and discussed in relation to other mesic zone taxa. Five of these barriers were inferred as being of early Miocene age, and implicated in the initial vicariant separation of endemic regional clades. Phylogeographic results for Australian Archaeidae are congruent with a model of sequential allopatric speciation in Tertiary refugia, as driven by the contraction and fragmentation of Australia’s mesic biomes during the Miocene. Assassin spiders clearly offer great potential for further testing historical biogeographic processes in temperate and eastern Australia, and are a useful group for better understanding the biology and biogeography of the Australian mesic zone.     

Phylogenetic relationships of the globally distributed freshwater prawn genus Macrobrachium (Crustacea: Decapoda: Palaemonidae): biogeography, taxonomy and the convergent evolution of abbreviated larval development

There has hitherto been little research into evolutionary and taxonomic relationships amongst species of the freshwater prawn genus Macrobrachium Bate across its global distribution. Previous work by the authors demonstrated that the endemic Australian species did not evolve from a single ancestral lineage. To examine whether other regional Macrobrachium faunas also reflect this pattern of multiple origins, the phylogeny of 30 Macrobrachium species from Asia, Central/South America and Australia was inferred from mitochondrial 16S rRNA sequences. Phylogenetic relationships demonstrate that, despite some evidence for regional diversification, Australia, Asia and South America clearly contain Macrobrachium species that do not share a common ancestry, suggesting that large-scale dispersal has been a major feature of the evolutionary history of the genus. The evolution of abbreviated larval development (ALD), associated with the transition from an estuarine into a purely freshwater lifecycle, was also mapped onto the phylogeny and was shown to be a relatively homoplasious trait and not taxonomically informative. Other taxonomic issues, as well as the evolutionary origins of Macrobrachium , are also discussed.     

Flying with the birds? Recent large-area dispersal of four Australian Limnadopsis species (Crustacea: Branchiopoda: Spinicaudata)

Temporary water bodies are important freshwater habitats in the arid zone of Australia. They harbor a distinct fauna and provide important feeding and breeding grounds for water birds. This paper assesses, on the basis of haplotype networks, analyses of molecular variation and relaxed molecular clock divergence time estimates, the phylogeographic history, and population structure of four common temporary water species of the Australian endemic clam shrimp taxon Limnadopsis in eastern and central Australia (an area of >1,350,000 km(2)). Mitochondrial cytochrome c oxidase subunit I sequences of 413 individuals and a subset of 63 nuclear internal transcribed spacer 2 sequences were analyzed. Genetic differentiation was observed between populations inhabiting southeastern and central Australia and those inhabiting the northern Lake Eyre Basin and Western Australia. However, over large parts of the study area and across river drainage systems in southeastern and central Australia (the Murray-Darling Basin, Bulloo River, and southern Lake Eyre Basin), no evidence of population subdivision was observed in any of the four Limnadopsis species. This indicates recent gene flow across an area of ～800,000 km(2). This finding contrasts with patterns observed in other Australian arid zone taxa, particularly freshwater species, whose populations are often structured according to drainage systems. The lack of genetic differentiation within the area in question may be linked to the huge number of highly nomadic water birds that potentially disperse the resting eggs of Limnadopsis among temporary water bodies. Genetically undifferentiated populations on a large geographic scale contrast starkly with findings for many other large branchiopods in other parts of the world, where pronounced genetic structure is often observed even in populations inhabiting pools separated by a few kilometers. Due to its divergent genetic lineages (up to 5.6% uncorrected p-distance) and the relaxed molecular clock divergence time estimates obtained, Limnadopsis parvispinus is assumed to have inhabited the Murray-Darling Basin continuously since the mid-Pliocene (～4 million years ago). This means that suitable temporary water bodies would have existed in this area throughout the wet-dry cycles of the Pleistocene.     

Botryocladius gen.n.: a new transantarctic genus of orthocladiine midge (Diptera: Chironomidae)

A new genus of orthocladiine Chironomidae, Botryocladius (type species B. grapeth sp.n. from eastern Australia) is described and illustrated in all life history stages. All thirteen included species are described as new, six from eastern Australia (B. grapeth, B. brindabella, B. mdfrc, B. collessi, B. tasmania, B. australoalpinus), two from Western Australia (B. bibulmun and B. freemani), one from ephemeral streams in Australia (B. petrophilus) and four from Patagonian Argentina and Chile (B. edwardsi, B. glacialis, B. mapuche and B. tronador). All Australian species are known from at least pupal exuviae, most from adult males and several from larvae. In contrast, only B. edwardsi amongst Neotropical species is known from the adult male; all others are described from pupal exuviae. The immature stages are lotic in Australian permanent and temporary streams and Patagonian glacial streams and rivers, and lentic in Neotropical glacial-fed and Australian subalpine lakes. Botryocladius appears to belong with a grouping centred on two formally undescribed taxa from Australia. The genus evidently demonstrates a vicariant distribution with at least two sister-group relationships between South American and Australian taxa, providing a minimum dating for the clade of 38 Ma., with apparent absence from New Zealand indicating a maximum date of 80 Ma.     

Phylogenetic relationships and remarkable radiation in Parartemia (Crustacea: Anostraca), the endemic brine shrimp of Australia: evidence from mitochondrial DNA sequences

The Australian continent is notable for the faunal radiations which have occurred in its saline inland waters. The endemic brine shrimp genus Parartemia inhabits many of these habitats. The complex pattern of morphological variation in parartemiids has impeded the establishment of a sound scheme of species relationships. The present study provides an explicit hypothesis of relationships for the genus based on nucleotide sequence data from a segment of mitochondrial DNA coding for the large subunit rRNA gene. Phylogenetic analyses indicated that the eight known species are genetically distinct, and revealed the existence of at least two new species. The molecular data support certain morphology-based relationships among species, but are inconsistent with other hypotheses. There is evidence that most members of the genus arose in a short interval, followed by remarkable genetic divergence. Comparisons of levels of mt DNA sequence divergence between lineages from saline inland waters and freshwaters using representative crustacean groups from Australia that included parartemiids indicated profound differences in rates of evolution, with halophiles exhibiting greater rates of change than their counterparts from freshwaters.     

A multigene molecular assessment of cryptic biodiversity in the iconic freshwater blackfishes (Teleostei: Percichthyidae: Gadopsis) of south‐eastern Australia

Freshwater biodiversity is under ever increasing threat from human activities, and its conservation and management require a sound knowledge of species‐level taxonomy. Cryptic biodiversity is a common feature for aquatic systems, particularly in Australia, where recent genetic assessments suggest that the actual number of freshwater fish species may be considerably higher than currently listed. The freshwater blackfishes (genus Gadopsis) are an iconic group in south‐eastern Australia and, in combination with their broad, naturally divided distribution and biological attributes that might limit dispersal, as well as ongoing taxonomic uncertainty, they comprise an ideal study group for assessing cryptic biodiversity. We used a multigene molecular assessment including both nuclear (51 allozyme loci; two S7 introns) and matrilineal markers (cytb) to assess species boundaries and broad genetic substructure within freshwater blackfishes. Range‐wide examination demonstrates the presence of at least six candidate species across two nominal taxa, Gadopsis marmoratus and Gadopsis bispinosus. Phylogeographical patterns often aligned to purported biogeographical provinces but occasionally reflected more restricted and unexpected relationships. We highlight key issues with taxonomy, conservation, and management for a species group in a highly modified region. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 521–540.     

Molecular evidence suggests an ancient radiation for the fairy shrimp genus Streptocephalus (Branchiopoda: Anostraca)

Phylogenetic relationships among assumed Gondwanan aquatic inland invertebrate fauna are generally largely neglected, and biogeographical hypotheses for these organisms are generally inferred from historic (palaeogeographical) and contemporary distribution patterns. The distribution of the monogeneric thermophilic freshwater fairy shrimp family Streptocephalidae ( Streptocephalus ) provides a particularly useful framework to test the three contrasting biogeographical scenarios proposed for the evolution of this group: (1) the genus evolved in Laurasia and subsequently dispersed into Africa and North America; (2) the genus evolved and dispersed out of Africa and (3) the current distribution of the genus is the result of vicariance following the fragmentation of Gondwana. In the present study, the phylogenetic relationships of species in this genus are examined with the use of two mitochondrial genes (12S rRNA and COI mtDNA), while the phylogenetic relationships among the North American species and selected African taxa was investigated using the nuclear fragment (5.8S-ITS-1-18S). Phylogenetic results indicate that Streptocephalus probably evolved in Gondwana and that the current distribution patterns are a consequence of a combination of vicariance and limited dispersal. The implications for the evolution of continental freshwater crustaceans are discussed.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 313–327.     

What lies beneath: molecular phylogenetics and ancestral state reconstruction of the ancient subterranean Australian Parabathynellidae (Syncarida, Crustacea)

The crustacean family Parabathynellidae is an ancient and significant faunal component of subterranean ecosystems. Molecular data were generated in order to examine phylogenetic relationships amongst Australian genera and assess the species diversity of this group within Australia. We also used the resultant phylogenetic framework, in combination with an ancestral state reconstruction (ASR) analysis, to explore the evolution of two key morphological characters (number of segments of the first and second antennae), previously used to define genera, and assess the oligomerization principle (i.e. serial appendage reduction over time), which is commonly invoked in crustacean systematics. The ASR approach also allowed an assessment of whether there has been convergent evolution of appendage numbers during the evolution of Australian parabathynellids. Sequence data from the mtDNA COI and nDNA 18S rRNA genes were obtained from 32 parabathynellid species (100% of described genera and ~25% of described species) from key groundwater regions across Australia. Phylogenetic analyses revealed that species of each known genus, defined by traditional morphological methods, were monophyletic, suggesting that the commonly used generic characters are robust for defining distinct evolutionary lineages. Additionally, ancestral state reconstruction analysis provided evidence for multiple cases of convergent evolution for the two morphological characters evaluated, suggesting that caution needs to be shown when using these characters for elucidating phylogenetic relationships, particularly when there are few morphological characters available for reconstructing relationships. The ancestral state analysis contradicted the conventional view of parabathynellid evolution, which assumes that more simplified taxa (i.e. those with fewer-segmented appendages and setae) are derived and more complex taxa are primitive.     

Molecular systematics of the Kakaducarididae (Crustacea: Decapoda: Caridea)

The systematic relationships of the freshwater shrimp family, Kakaducarididae, were examined using mitochondrial and nuclear DNA sequences. Combined nuclear (18S rDNA, 28S rDNA, Histone) and mitochondrial (16S rDNA) analyses placed the kakaducaridid genera, Kakaducaris and Leptopalaemon, as a strongly supported clade within the Palaemonidae, in a close relationship with the genus Macrobrachium. Monophyly of the Australian Kakaducarididae was strongly supported by the molecular data. Estimated net divergence times between Kakaducaris and Leptopalaemon using mitochondrial 16S rDNA equate to a late Miocene/Pliocene split. Within Leptopalaemon, each locality was distinct for mitochondrial COI haplotypes, suggesting long-term isolation or recent genetic bottlenecks, a lack of contemporary gene flow amongst sites and a small Ne. Mitochondrial groupings within Leptopalaemon were largely congruent with several previously recognised morphotypes. Estimated net divergence times between L. gagadjui and the new Leptopalaemon morphotypes equate to a split in the late Pliocene/early Pleistocene. The hypothesis that the Kakaducarididae is comprised of relict species in specialised ecological niches is not supported by the molecular data, which instead suggest a relatively recent origin for the group in northern Australia, sometime in the late Miocene or Pliocene.     

Phylogeny of the cave shrimp Troglocaris: Evidence of a young connection between Balkans and Caucasus

The remarkably discontinuous distribution of the cave shrimp genus Troglocaris in South France, West Balkans, and West Caucasus has long been considered a biogeographic enigma. To solve it, its phylogeny was reconstructed by analyzing sequences from two mitochondrial (cytochrome oxidase I and 16S rRNA) and one nuclear gene (28S rRNA) using maximum likelihood, parsimony and Bayesian inference. The genus was found to be polyphyletic because the French taxon T. inermis had no direct common ancestry with other Troglocaris taxa but was sister to the epigean freshwater atyid Dugastella valentina. All other Troglocaris species constituted a well-supported monophylum, the second cave shrimp genus Spelaeocaris nested within. The monophylum had a well-defined structure: (1) a clade restricted to the Dinaric area of the Western Balkans containing the type species T. anophthalmus along with some unnamed species, and (2) a geographically mixed clade split between the Caucasian T. kutaissiana species complex on one, and T. hercegovinensis, S. pretneri, plus an unnamed taxon on the other side. It was surprising to find the dichotomy between the Caucasian and one of the West-Balkan lineages so low in the phylogenetic hierarchy of the genus. Taking into account molecular rates of other decapods, we tentatively dated this split at 6-11 Myr. This time is in agreement with the brackish and freshwater phase of the Paratethys thus allowing for a freshwater common ancestor of Caucasian and Dinaric cave shrimps. This would weaken the marine relicts hypothesis that has often been invoked to explain the distribution of freshwater cave species with close marine relatives.     

Assessing the ecological values of rivers: an application of a multi-criteria approach to rivers of the South Coast Region,Western Australia

There is a growing awareness that in the light of increased threats, Australia’s freshwater ecosystems of highest value need to be protected. This implies that a consistent and comprehensive system for the classification of freshwater systems in terms of their ecological values is required for conservation planning. Despite earlier steps taken to develop appropriate methodologies for prioritising Australian waterways for protection, there is, as yet, no nationally agreed method for prioritising waterways in Australia based on their ecological values. Using the core criteria of naturalness, representativeness, diversity, rarity and special features, this paper describes the development of a framework of criteria, indicators and measures for the identification of high conservation value aquatic ecosystems, and trials this system by undertaking a comparative assessment of the ecological values of river systems in the South Coast region, Western Australia. A total of 33 river systems located in two aquatic bioregions were successfully ranked according to their ecological value, confirming that the development and implementation of a relatively simple system for assessing ecological values of Australian rivers shows promise, and that core criteria identified previously in the literature provide a good starting point for assessing ecological values of Australian aquatic ecosystems.     

Molecular identification of the precise geographic origins of an invasive shrimp species in a globally significant Australian biodiversity hotspot

This report details the first detection of invasive freshwater shrimp in the south-west of Western Australia, a region recognised for its biodiversity and the vulnerability of its endemic species. The species was detected as part of biosecurity surveillance of Perth’s freshwater lakes. The introduced specimens were tentatively identified morphologically as Caridina indistincta, and then confirmed by molecular analysis as “species B” of that taxon, part of a species-complex native to eastern Australia. Phylogeographic analyses were then used to narrow down the likely source population to a small area of south-eastern Queensland, over 3500 kms across the continent from the invasion sites. The potential source area is heavily involved in the trade in freshwater species for aquaria and recreational fish stocking. This information will help in identifying the precise invasion vector and could thus allow more targeted management measures to prevent future imports of exotic aquatic invasive species.     

Chromosome numbers in AustralianEuphrasia (Scrophulariaceae)

Chromosome numbers for six Australian taxa ofEuphrasia have been determined. Improved staining techniques have shown that numbers for four of the taxa published previously by the first author were incorrect. The investigated taxa show high ploidy levels with an apparent base number of x = 11, the same as for the genus outside Australia.Dedicated to Prof. DrElisabeth Tschermak-Woess on the occasion of her 70th birthday.     

Desert vines: a comparison of Australia with other areas

Aim To characterize the Australian desert vine flora and to compare it with that of deserts in other continents. Location The Stony Deserts, the Simpson Desert and the four main deserts of Western Australia. Methods The Western Australian Herbarium data base and published papers were used to develop vine checklists for each Australian desert studied. A literature search was used to classify the families and genera into phytogeographical elements and to provide data for intercontinental comparisons. Results Thirty‐seven vine species are listed for the six Australian deserts studied. They constitute from 0.8 to 2.7% of the vascular flora, which is within the normal range for arid zone floras. Comparing Australian data with those from the Sonoran and Chihuahuan deserts (North America), for non‐vine taxa, very different phylogenetic lines are present. However, for vines, three of the four most important families are the same in each case, viz. Convolvulaceae, Fabaceae and Asclepiadaceae. This reflects large pantropical and cosmopolitan families shared between all three data sets. At the generic level, in Australia pantropical taxa and taxa from the Old World tropics far outnumber endemic ones, as do pantropical and neotropical genera in North America. Herbaceous vines predominate in Australian deserts as they do in North American ones, but nevertheless, the percentage of woody vines is higher in Australia (32%) than in North America (highest value of 24%). Earlier views that Australian deserts are rich in annual vines are not supported. Main conclusions For many life‐forms, the Australian flora is composed of very different phylogenetic lines to the floras of other continents. However, for desert vines, at the level of family and even of genus, there are surprising similarities between Australia and even a continent as distant as North America, because of shared pantropical and cosmopolitan taxa.     

Evolutionary divergence in freshwater insects with contrasting dispersal capacity across a sea of desert

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