Panbiogeography of New Caledonia,south‐west Pacific: basal angiosperms on basement terranes,ultramafic endemics inherited from volcanic island arcs and old taxa endemic to young islands

Aim To investigate areas of endemism in New Caledonia and their relationship with tectonic history. Location New Caledonia, south‐west Pacific. Methods Panbiogeographical analysis. Results Biogeographical patterns within New Caledonia are described and illustrated with reference to eight terranes and ten centres of endemism. The basement terranes make up a centre of endemism for taxa including Amborella, the basal angiosperm. Three of the terranes that accreted to the basement in the Eocene (high‐pressure metamorphic terrane, ultramafic nappe and Loyalty Ridge) have their own endemics. Main conclusions New Caledonia is not simply a fragment of Gondwana but, like New Zealand and New Guinea, is a complex mosaic of allochthonous terranes. The four New Caledonian basement terranes were all formed from island arc‐derived and arc‐associated material (including ophiolites) which accumulated in the pre‐Pacific Ocean, not in Gondwana. They amalgamated and were accreted to Gondwana (eastern Australia) in the Late Jurassic/Early Cretaceous, but in the Late Cretaceous they separated from Australia with the opening of the Tasman Sea and break‐up of Gondwana. An Eocene collision of the basement terranes with an island arc to the north‐east – possibly the Loyalty Ridge – is of special biogeographical interest in connection with New Caledonia–central Pacific affinities. The Loyalty–Three Kings Ridge has had a separate history from that of the Norfolk Ridge/New Caledonia, although both now run in parallel between Vanuatu and New Zealand. The South Loyalty Basin opened between Grande Terre and the Loyalty Ridge in the Cretaceous and attained a width of 750 km. However, it was almost completely destroyed by subduction in the Eocene which brought the Loyalty Ridge and Grande Terre together again, after 30 Myr of separation. The tectonic history is reflected in the strong biogeographical differences between Grande Terre and the Loyalty Islands. Many Loyalty Islands taxa are widespread in the Pacific but do not occur on Grande Terre, and many Grande Terre/Australian groups are not on the Loyalty Islands. The Loyalty Islands are young (2 Myr old) but they are merely the currently emergent parts of the Loyalty Ridge whose ancestor arcs have a history of volcanism dating back to the Cretaceous. Old taxa endemic to the young Loyalty Ridge islands persist over geological time as a dynamic metapopulation surviving in situ on the individually ephemeral islands and atolls found around subduction zones. The current Loyalty Islands, like the Grande Terre terranes, have inherited their biota from previous islands. On Grande Terre, the ultramafic terrane was emplaced on Grande Terre in the Eocene (about the same time as the collision with the island arc). The very diverse endemic flora on the ultramafics may have been inherited by the obducting nappe from prior base‐rich habitat in the region, including the mafic Poya terrane and the limestones typical of arc and intraplate volcanic islands.     

Globally basal centres of endemism: the Tasman-Coral Sea region (south-west Pacific), Latin America and Madagascar/South Africa

The New Zealand wrens (Acanthisittidae) are basal in passerine birds and in New Caledonia, the closest country to New Zealand, Amborella is basal in angiosperms. A review of molecular studies indicates that 29 other locally or regionally endemic clades around the Tasman and Coral Sea basins have cosmopolitan or globally widespread sister groups. Other areas that have local endemics basal to diverse global groups include Borneo, Madagascar/South Africa/Tanzania, southern China–Taiwan–Japan, and different parts of Latin America, especially the Guayana Plateau and western Mexico. Basal clades are widely interpreted as ancestral and their location is generally taken to represent a centre of origin for the group as a whole. In the present study, basal groups are treated simply as small (less speciose) sister groups. The Tasman and western Mexico/Caribbean regions have important biogeographic and tectonic ties with each other and with the central Pacific. Large igneous provinces (Ontong Java, Hikurangi‐Manihiki, and Gorgona Plateaus) formed in the central Pacific in the Cretaceous. Fossil wood is found on the Ontong Java Plateau, and formerly emergent seamounts up to 24 km across occur on Hikurangi Plateau. Many terranes in New Zealand, New Caledonia, New Guinea and western America represent former island arcs (or their products) that formed in the central Pacific and later accreted to the Pacific margins. Long‐term survival of taxa as metapopulations on the ephemeral volcanic islands and atolls of plate margins and fissures may explain the biogeographical connections among the Tasman region, the central Pacific, and the accreted terranes of western America. Branching sequences in cladograms and phylogenetic trees have been interpreted as dispersal events, but instead probably indicate the sequence of differentiation in an already widespread ancestor. Major disjunctions of tens of thousands of kilometres often occur between taxa at consecutive nodes on a tree and dispersal by physical movement is unlikely. The break between the globally basal centres and the rest of the world marks the initial site of differentiation of a widespread ancestor, with subsequent or more or less simultaneous differentiation occurring in other areas. Differentiation of the modern angiosperms, passerines and other groups first took place around the Tasman region, or at least the terranes now accumulated there, and then around other centres, especially Madagascar–South Africa and Mexico–north‐west South America. Angiosperms are now recognized as basal to extant gymnosperms and major tectonic dynamism around the globally basal centres during the Mesozoic, involving terrane accretion, orogeny, and rifting could have been involved with the last important modernization of angiosperms, birds and other groups. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96 , 222–245.     

Biogeography of the Monimiaceae (Laurales): a role for East Gondwana and long‐distance dispersal,but not West Gondwana

Aim The biogeography of the tropical plant family Monimiaceae has long been thought to reflect the break‐up of West and East Gondwana, followed by limited transoceanic dispersal. Location Southern Hemisphere, with fossils in East and West Gondwana. Methods We use phylogenetic analysis of DNA sequences from 67 of the c. 200 species, representing 26 of the 28 genera of Monimiaceae, and a Bayesian relaxed clock model with fossil prior constraints to estimate species relationships and divergence times. Likelihood optimization is used to infer switches between biogeographical regions on the highest likelihood tree. Results Peumus from Chile, Monimia from the Mascarenes and Palmeria from eastern Australia/New Guinea form a clade that is sister to all other Monimiaceae. The next‐deepest split is between the Sri Lankan Hortonia and the remaining genera. The African Monimiaceae, Xymalos monospora, then forms the sister clade to a polytomy of five clades: (I) Mollinedia and allies from South America; (II) Tambourissa and allies from Madagascar and the Mascarenes; (III) Hedycarya, Kibariopsis and Leviera from New Zealand, New Caledonia and Australia; (IV) Wilkiea, Kibara, Kairoa; and (V) Steganthera and allies, all from tropical Australasia. Main conclusions Tree topology, fossils, inferred divergence times and ances‐tral area reconstruction fit with the break‐up of East Gondwana having left a still discernible signature consisting of sister clades in Chile and Australia. There is no support for previous hypotheses that the break‐up of West Gondwana (Africa/South America) explains disjunctions in the Monimiaceae. The South American Mollinedia clade is only 28–16 Myr old, and appears to have arrived via trans‐Pacific dispersal from Australasia. The clade apparently spread in southern South America prior to the Andean orogeny, fitting with its first‐diverging lineage (Hennecartia) having a southern‐temperate range. The crown ages of the other major clades (II–V) range from 20 to 29 Ma, implying over‐water dispersal between Australia, New Caledonia, New Zealand, and across the Indian Ocean to Madagascar and the Mascarenes. The endemic genus Monimia on the Mascarenes provides an interesting example of an island lineage being much older than the islands on which it presently occurs.     

Phylogeography of the whelk genus Cominella (Gastropoda: Buccinidae) suggests long‐distance counter‐current dispersal of a direct developer

The gastropod genus Cominella Gray, 1850 consists of approximately 20 species that inhabit a wide range of marine environments in New Zealand and Australia, including its external territory, the geographically isolated Norfolk Island. This distribution is puzzling, however, with apparently closely‐related species occurring either side of the Tasman Sea, even though all species are considered to have limited dispersal abilities. To determine how Cominella attained its current distribution, we derived a dated molecular phylogeny, which revealed a clade comprising all the Australian and Norfolk Island species nested within four clades of solely New Zealand species. This Australian clade diverged well after the vicariant separation of New Zealand from Australia, and implies two long‐distance dispersal events: a counter‐current movement across the Tasman Sea from New Zealand to Australia, occurring at the origination of the clade, followed by the colonization of Norfolk Island. The biology of Cominella suggests that the most likely method of long‐distance dispersal is rafting as egg capsules. Our robust phylogeny also means that the current Cominella classification requires revision. We propose that our clades be recognized as subgenera: Cominella (s.s.), Cominista, Josepha, Cominula, and Eucominia, with each subgenus comprising only of New Zealand or Australian species. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 315–332.     

The endemic plant families and the palms of New Caledonia: a biogeographical analysis

This paper provides a panbiogeographical analysis of the endemic plant families and the palms of New Caledonia. There are three endemic plant families in New Caledonia and several genera that were previously recognized as endemic families. Of these taxa, some are sister to widespread Northern Hemisphere or global groups (Canacomyrica, Austrotaxus, Amborella). The others belong to trans‐Indian Ocean groups (Strasburgeria), trans‐tropical Pacific groups (Oncotheca) or Tasman Sea/Coral Sea groups (Phelline, Paracryphia) that are sister to widespread Northern Hemisphere or global groups. In palms, the four clades show allopatric regional connections in, respectively: (1) western Indonesia, Malaysia and Thailand; (2) Vanuatu/Fiji and the southern Ryukyu Islands near Taiwan; (3) the western Tasman/Coral Sea (eastern Australia, New Guinea and the Solomon Islands); and (4) the eastern Tasman/Coral Sea (Lord Howe and Norfolk Islands, New Zealand, Vanuatu, Fiji and the Solomon Islands). The four clades thus belong to different centres of endemism that overlap in New Caledonia. The patterns are attributed not to chance dispersal and adaptive radiation but to the different histories of the eight terranes that fused to produce modern New Caledonia. Trans‐tropical Pacific connections can be related to the Cretaceous igneous plateaus that formed in the central Pacific and were carried, with plate movement, west to the Solomon Islands and New Zealand, and east to Colombia and the Caribbean.     

Biogeography of temperate Australasian Polystichum ferns as inferred from chloroplast sequence and AFLP

Aim and location New Zealand began to separate from Gondwana c. 85 Ma, and has been isolated from the nearest large landmass, Australia, by some 2000 km of the Tasman Sea since c. 60 Ma. Given New Zealand's long geographical isolation, there has been considerable interest in explaining the origins of its different biotic elements. Here we investigate the biogeography of the fern genus Polystichum from temperate Australasia. Six species are found in New Zealand, four in Australia, and two on Lord Howe Island. Methods The evolutionary relationships between the twelve Polystichum species found in temperate Australasia were inferred from phylogenetic analyses of two molecular data sets: DNA sequence from the chloroplast rps4–trnS spacer locus; and AFLP DNA‐fingerprinting. The timing of the separation between Australian and New Zealand Polystichum was estimated by using the fossil record to temporally calibrate the rbcL sequence differentiation between representative species from these regions. Results Species of Polystichum from New Zealand appear to comprise a monophyletic group. This suggests that Polystichum crossed the Tasman only once. Temporal calibration of the rbcL sequence differentiation between Australian and New Zealand Polystichum indicates that a vicariant explanation for their separation can be rejected in favour of trans‐oceanic dispersal. Main conclusions The extant diversity within New Zealand Polystichum appears to have been derived from a single, trans‐oceanic dispersal event (within the last c. 20 Myr), followed by a relatively extensive in situ ecological radiation.     

The trans-Pacific zipper effect: disjunct sister taxa and matching geological outlines that link the Pacific margins

Aim To combine analyses of trans‐Pacific sister taxa with geological evidence in order to test the hypothesis of the existence of a Panthalassa superocean. Location The study is concerned with taxa, both fossil and extant, from East Asia, Australia, New Zealand, South America and North America. Methods Phylogenetic and distributional analyses of trans‐Pacific biota were integrated with geological evidence from the Pacific and circum‐Pacific regions. Results A series of recent biogeographical analyses delineates a zipper‐like system of sister areas running up both margins of the Pacific, with each section of western North and South America corresponding to a particular section from East Asia/Australia/New Zealand. These sister areas coincide neatly with a jigsaw‐like fit provided by the matching Mesozoic coastlines that bracket the Pacific. Main conclusions The young age (<200 Myr) of oceanic crust, the matching Mesozoic circum‐Pacific outlines, and a corresponding system of interlocking biogeographical sister areas provide three independent avenues of support for a closed Pacific in the Upper Triassic–Lower Jurassic. The hypothesis of the existence and subsequent subduction of the pre‐Pacific superocean Panthalassa is not only unnecessary, it conflicts with this evidence. Panthalassa‐based paleomaps necessitate the invention of dozens of additional hypotheses of species‐dependent, trans‐oceanic dispersal events, often involving narrow‐range taxa of notoriously limited vagility, in order to explain repeated examples of the same biogeographical pattern. Removing the vanished‐superocean hypothesis reunites both the matching geological outlines and all the disjunct sister taxa. In brief, what appears to be a multi‐era tangle of convoluted, trans‐oceanic distributions on Panthalassa‐based paleomaps is actually a relatively simple biogeographical pattern that is explainable by a single vicariant event: the opening and expansion of the Pacific.     

Gondwanan radiation of the Southern Hemisphere crayfishes (Decapoda: Parastacidae): evidence from fossils and molecules

Aim The sequential break‐up of Gondwana is thought to be a dominant process in the establishment of shared biota across landmasses of the Southern Hemisphere. Yet similar distributions are shared by taxa whose radiations clearly post‐date the Gondwanan break‐up. Thus, determining the contribution of vicariance versus dispersal to seemingly Gondwanan biota is complex. The southern freshwater crayfishes (family Parastacidae) are distributed on Australia and New Guinea, South America, Madagascar and New Zealand and are unlikely to have dispersed via oceans, owing to strict freshwater limitations. We test the hypotheses that the break‐up of Gondwana has led to (1) a predominately east–west (((Australia, New Zealand: 80 Ma) Madagascar: 160–121 Ma) South America: 165–140 Ma), or (2) a southern (((Australia, South America: 52–35 Ma) New Zealand: 80 Ma) Madagascar: 160–121 Ma) pattern for parastacid crayfish. Further, we examine the evidence for a complete drowning of New Zealand and subsequent colonization by freshwater crayfish. Location Southern Hemisphere. Methods The evolutionary relationships among the 15 genera of Parastacidae were reconstructed using mitochondrial [16S, cytochrome c oxidase subunit I (COI)] and nuclear (18S, 28S) sequence data and maximum likelihood and Bayesian methods of phylogenetic reconstruction. A Bayesian (multidivtime ) molecular dating method using six fossil calibrations and phylogenetic inference was used to estimate divergence time among crayfish clades on Gondwanan landmasses. Results The South American crayfish are monophyletic and a sister group to all other southern crayfish. Australian crayfish are not monophyletic, with two Tasmanian genera, Spinastacoides and Ombrastacoides, forming a clade with New Zealand and Malagasy crayfish (both monophyletic). Divergence of crayfish among southern landmasses is estimated to have occurred around the Late Jurassic to Early Cretaceous (109–178 Ma). Main conclusions The estimated phylogenetic relationships and time of divergence among the Southern Hemisphere crayfishes were consistent with an east–west pattern of Gondwanan divergence. The divergence between Australia and New Zealand (109–160 Ma) pre‐dated the rifting at around 80 Ma, suggesting that these lineages were established prior to the break‐up. Owing to the age of the New Zealand crayfish, we reject the hypothesis that there was a complete drowning of New Zealand crayfish habitat.     

The importance of looking at small-scale patterns when inferring Gondwanan biogeography: a case study of the centipede Paralamyctes (Chilopoda, Lithobiomorpha, Henicopidae)

Gondwanan biogeography has fascinated zoologists and botanists for over a century, but most biogeographical work has used continent-scale areas as analytical units. More finely resolved patterns, as can be obtained from small invertebrates with limited dispersal abilities, will be obscured in those studies. A common case is treating Australia as a single biogeographical region. In the present study, the necessity of splitting Australia into multiple microareas is demonstrated using centipedes as an example. The lithobiomorph centipede Paralamyctes is distributed on fragments of Gondwana, with species in southern Africa, Madagascar, southern India, Patagonia, eastern Australia, and New Zealand. A cladogram for Paralamyctes is based on morphology and sequences for four molecular markers for 30 terminals that sample 20 of 26 known ingroup species and four outgroups. Analysis with direct optimization across a range of indel costs and transversion : transition cost ratios identifies two main clades: Paralamyctes ( Paralamyctes ) unites species from southern Africa, Madagascar, tropical and warm temperate Australia, and New Zealand. The other group includes the temperate Australian/New Zealand Paralamyctes ( Haasiella ) and Paralamyctes ( Thingathinga ) and a Chilean clade. Subtree analysis finds that different parts of Australia have closest affinities to other Gondwanan fragments, and some of these relationships (such as that between north Queensland and New Zealand) are based on taxonomically stable clades. Area delimitation for large continental fragments should use sufficiently fine resolution to test the 'monophyly' of those fragments and attempt to eliminate spurious geographical paralogy.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 65–78.     

Frugivore gape size and the evolution of fruit size and shape in southern hemisphere floras

Abstract The present study uses differences among frugivore faunas of the southern hemisphere landmasses to test whether frugivore characteristics have influenced the evolution of fruit traits. Strong floristic similarities exist among southern landmasses; for example, 75% of New Zealand vascular plant genera also have species in Australia. However, plants in Australia and South America have evolved in the presence of a range of mammalian frugivores, whereas those in New Zealand, New Caledonia and the Pacific Islands have not. In addition, the avian frugivores in New Zealand and New Caledonia are generally smaller than those of Australia. If frugivore characteristics have influenced the evolution of fruit traits, predictable differences should exist between southern hemisphere fruits, particularly fruit size and shape. Fruit dimensions were measured for 77 New Zealand species and 31 Australian species in trans‐Tasman genera. New Zealand fruits became significantly more ellipsoid in shape with increasing size. This is consistent with frugivore gape size imposing a selective pressure on fruit ingestability. This result is not a product of phylogenetic correlates, as fruit length and width scaled isometrically for Australian species in genera shared with New Zealand. Within‐genus contrasts between New Zealand and Australian species in 20 trans‐Tasman genera showed that New Zealand species have significantly smaller fruits than their Australian counterparts. Within‐genus contrasts between New Zealand and South American species in nine genera gave the same result; New Zealand species had significantly smaller fruits than their South American counterparts. No difference was found in fruit size or shape between New Zealand and New Caledonia congeneric species from 12 genera. These results are consistent with the broad characteristics of the frugivore assemblage influencing the evolution of fruit size and shape in related species. The smaller‐sized New Zealand frugivore assemblage has apparently influenced the evolution of fruit size of colonizing taxa sometimes within a relatively short evolutionary timeframe.     

East meets west: biogeology of the Campbell Plateau

The New Zealand Subantarctic Islands, emergent remnants of the Campbell Plateau, were given World Heritage status in 1998 in recognition of their importance to global biodiversity. We describe the flora and fauna of these islands and discuss the results of recent phylogenetic analyses. Part of the New Zealand Subantarctic biota appears to be relictual and to be derived from west Gondwana. The relictual element is characterized by genera endemic to the Campbell Plateau that show relationships with taxa of the southern South Island, New Zealand, southern South America, and the north Pacific. In contrast, a younger, east Gondwanan element is composed of species that are either taxonomically identical to widespread mainland species, or endemic species with close New Zealand relatives. Area cladograms support the inclusion of the southern South Island, New Zealand and Macquarie Island (although this is separate geologically) as parts of the Campbell Plateau, but suggest the Chatham Rise and Torlesse terranes of the eastern South Island, New Zealand were originally parts of east Gondwana. East and west Antarctica acted as independent plates during the breakup of Gondwana, and were separated by oceanic crust until a compressive phase sutured them along the trace of the trans‐Antarctic mountains during the early Tertiary. The Campbell Plateau microcontinent was connected to west Antarctica until its separation at 80 Mya, contemporaneous with the separation of the southern portion of the Melanesian rift from east Gondwana. Presently the Campbell Plateau is joined to the Melanesian Rift along the Alpine Fault. Cenozoic plate tectonic reconstructions place the Campbell Plateau adjacent to the Melanesian Rift throughout the rift–drift phase, relative motion being confined to strike–slip movement over the last 20 Myr. Our synthesis of phylogenetic and plate tectonic evidence suggests that the Alpine Fault is the most recent development of a much older extensional rift/basin boundary originally separating west and east Gondwana. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86 , 95–115.     

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.     

Biogeographical and geological evidence for a smaller, completely-enclosed Pacific Basin in the Late Cretaceous

Aim To use biogeographical, palaeomagnetic, palaeosedimentary, and plate circuit data from Late Cretaceous regions in and around the Pacific to test the plate tectonic hypothesis of a pre‐Pacific superocean. Location East Asia, Australia, Antarctica, the western Americas, and the Pacific. Methods Literature surveys of the distributions of Cretaceous, circum‐Pacific taxa were compared with palaeomagnetic and palaeosedimentary data. Uncontroversial plate motions based on seafloor spreading data were also used to test the results of the biogeographical and palaeomagnetic analyses. Results The distributions of Cretaceous terrestrial taxa, mostly dinosaurs, imply direct, continental connections between Australia and East Asia, East Asia and North America, North America and South America, South America and Antarctica, and Antarctica and Australia. Palaeomagnetic, palaeosedimentary, and basic plate circuit analyses require little to no latitudinal motion of the Pacific plate with respect to the surrounding continents. Specifically, the data implies that western North America, East Asia, and the Pacific plate all increased in latitude by roughly the same amount (c. 11 ± 5°) since the Campanian – and that the Pacific Ocean Basin has increased in length north‐to‐south. Main conclusions Each of the analyses provides independent corroboration for the same conclusion: the Late Cretaceous Pacific plate was completely enclosed by the surrounding continents and has not experienced significant latitudinal motion with respect to North America, East Asia, or the Bering land bridge. This contrasts significantly with the plate tectonic history of the Pacific, implying instead that the Pacific plate formed in situ, pushing the continents apart as the plate and basin expanded. These results also substantiate recent biogeographical analyses that have concluded that a narrower Pacific Ocean Basin in the Mesozoic and early Tertiary provides the most reasonable explanation for the great number of trans‐Pacific disjunctions of poor dispersing taxa.     

Late Cenozoic diversification of the austral genus Lagenophora (Astereae,Asteraceae)

Lagenophora (Astereae, Asteraceae) has 14 species in New Zealand, Australia, Asia, southern South America, Gough Island and Tristan da Cunha. Phylogenetic relationships in Lagenophora were inferred using nuclear and plastid DNA regions. Reconstruction of spatio‐temporal evolution was estimated using parsimony, Bayesian inference and likelihood methods, a Bayesian relaxed molecular clock and ancestral area and habitat reconstructions. Our results support a narrow taxonomic concept of Lagenophora including only a core group of species with one clade diversifying in New Zealand and another in South America. The split between the New Zealand and South American Lagenophora dates from 11.2 Mya [6.1–17.4 95% highest posterior density (HPD)]. The inferred ancestral habitats were openings in beech forest and subalpine tussockland. The biogeographical analyses infer a complex ancestral area for Lagenophora involving New Zealand and southern South America. Thus, the estimated divergence times and biogeographical reconstructions provide circumstantial evidence that Antarctica may have served as a corridor for migration until the expansion of the continental ice during the late Cenozoic. The extant distribution of Lagenophora reflects a complex history that could also have involved direct long‐distance dispersal across southern oceans. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177 , 78–95.     

A new Transantarctic relationship: morphological evidence for a Rheidae–Dromaiidae–Casuariidae clade (Aves,Palaeognathae, Ratitae)

Although ratites have been studied in considerable detail, avian systematists have been unable to reach a consensus regarding their relationships. Morphological studies indicate a basal split separating Apterygidae from all other extant ratites, and a sister‐group relationship between Rheidae and Struthionidae. Molecular studies have provided evidence for the paraphyly of the Struthionidae and Rheidae, with respect to a clade of Australasian extant ratites. The position of the extinct Dinornithidae and Aepyornithidae also remains hotly debated. A novel pattern of diversification of ratites is presented herein. The phylogenetic analysis is based on 17 taxa and 129 morphological characters, including 77 new characters. The resultant tree yields a sister‐group relationship between New Zealand ratites (Apterygidae plus Dinornithidae) and all other ratites. Within this clade, the Aepyornithidae and Struthionidae are successive sister taxa to a new, strongly supported clade comprising the Rheidae, Dromaiidae, and Casuariidae. The link between South American and Australian biotas proposed here is congruent with numerous studies that have evidenced closely related taxa on opposite sides of the Southern Pacific. These repeated patterns of area relationships agree with current knowledge on Gondwana break‐up, which indicates that Australia and South America remained in contact across Antarctica until the earliest Tertiary. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 641–663.     

Historical biogeography of Hexabathynella , a cosmopolitan genus of groundwater Syncarida (Crustacea, Bathynellacea, Parabathynellidae)

Hexabathynella is the only cosmopolitan genus of the order Bathynellacea (Crustacea). The known species number 18, found in Europe (9), Africa (1), South America (2), North America (3) and Australia and New Zealand (3). Phylogenetic analysis suggests that the least derived species are those from South America and the most derived those from the Iberian Peninsula, North America and Australia. The five species with the most plesiomorphic characters occur in salt or brackish water, which supports a marine origin for the genus. Phylogenetic and biogeographical analyses suggest that the distribution of the genus can be explained by dispersion and a double vicariant biogeographical model based on plate tectonics and the evolution of the Tethys during the Mesozoic and Cenozoic.  © 2003 The Linnean Society of London . Biological Journal of the Linnean Society , 2003, 78 , 457–466.     

Systematic review of a new orb‐weaving spider genus (Araneae: Araneidae), with special reference to the Australasian‐Pacific and South‐East Asian fauna

The Australasian‐Pacific and South‐East Asian species of the new orb‐weaving spider genus Plebs with Plebs eburnus (Keyserling, 1886) as type species are revised. Following this study, Plebs includes a total of 22 species of which seven are here described new. Seven species are found in Australia, two in the Pacific region (New Caledonia, Vanuatu), and two in South‐East Asia (Papua New Guinea, The Philippines). Eleven Asian species are transferred to the new genus. Plebs represent comparatively small orb‐weaving spiders of c. 1.2–15.0 mm body length with a slightly elongated abdomen and humeral (shoulder) humps. Males of most species have two to three stout setae on the ventral side of their fourth coxae. Male pedipalps are characterized by the presence of a single macroseta on the patella, the presence of a paramedian apophysis as basal extension of the conductor, and an apical tegular protrusion. The female epigyne has a scape that is generally much longer than wide. It does not have a terminal pocket and is frequently broken off in a number of species. A phylogenetic analysis of 15 species of Plebs (those for which both sexes are known), 13 Australian/Pacific orb‐weaving spider species representing the most commonly collected clades with paramedian apophysis, three species of Nearctic Eriophora Simon, 1864, and Araneus diadematus Clerck, 1758, as outgroup, identified a single synapomorphy of Plebs based on 35 morphological and three behavioural characters: a distinct, inverted U‐shaped light pattern on the ventral side of the abdomen with two additional white spots anterolateral to the spinnerets. This analysis recovered a monophyletic clade of all Asian Plebs, suggesting a single colonization event of the genus that putatively originated in Australia. Most Plebs species appear to be active during the day. They build a regular orb‐web with vertical stabilimentum in grass and low shrubs. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166 , 279–341.     

Historical biogeography of Australian Rhamnaceae, tribe Pomaderreae

Aim To discover the pattern of relationships of areas of endemism for Australian genera in the plant family Rhamnaceae tribe Pomaderreae for comparison with other taxa and interpretation of biogeographical history. Location Australian mainland, Tasmania and New Zealand. Methods A molecular phylogeny and geographic distribution of species within four clades of Pomaderreae are used as a basis for recognition of areas of endemism and analysis of area relationships using paralogy‐free subtrees. The taxon phylogeny is the strict consensus tree from a parsimony analysis of 54 taxa, in four clades, and sequence data for the internal transcribed spacer regions of ribosomal DNA (ITS1‐5.8S‐ITS2) and the plastid DNA region trnL‐F. Results The biogeographical analysis identified five subtrees, which, after parsimony analysis, resulted in a minimal tree with 100% consistency and seven resolved nodes. Three sets of area relationships were identified: the areas of Arnhem and Kimberley in tropical north Australia are related based on the phylogeny of taxa within Cryptandra; the moister South‐west of Western Australia, its sister area the coastal Geraldton Sandplains, the semi‐arid Interzone region and arid Western Desert are related, based on taxa within Cryptandra, Spyridium, Trymalium and Pomaderris; and the eastern regions of Queensland, McPherson‐Macleay, south‐eastern New South Wales (NSW), Victoria, southern Australia, Tasmania and New Zealand are related based on Cryptandra, Pomaderris and Spyridium. Tasmania and NSW are related based entirely on Cryptandra, but the position of New Zealand relative to the other south‐eastern Australian regions is unresolved. Main conclusions The method of paralogy‐free subtrees identified a general pattern of geographic area relationships based on Australian Pomaderreae. The widespread distribution of clades, the high level of endemicity and the age of fossils for the family, suggest that the Pomaderreae are an old group among the Australian flora. Their biogeographical history may date to the early Palaeogene with subsequent changes through to the Pleistocene.     

Phylogeny and historical biogeography of Gondwanan moss-bugs (Insecta: Hemiptera: Coleorrhyncha: Peloridiidae)

The moss bugs of the Peloridiidae, a small group of cryptic and mostly flightless insects, is the only living family in Coleorrhyncha (Insecta: Hemiptera). Today 37 species in 17 genera are known from eastern Australia, New Zealand, New Caledonia and Patagonia, and the peloridiids are thereby a group with a classical southern Gondwanan distribution. To explicitly test whether the present-day distribution of the Peloridiidae actually results from the sequential breakup of southern Gondwana, we provide the first total-evidence phylogenetic study based on morphological and molecular characters sampled from about 75% of recognized species representing 13 genera. The results largely confirm the established morphological phylogenetic context except that South American Peloridium hammoniorum constitutes the sister group to the remaining peloridiids. A timescale analysis indicates that the Peloridiidae began to diversify in the land mass that is today's Patagonia in the late Jurassic (153 Ma, 95% highest posterior density: 78–231 Ma), and that splitting into the three extant well-supported biogeographical clades (i.e. Australia, Patagonia and New Zealand/New Caledonia) is consistent with the sequential breakup of southern Gondwana in the late Cretaceous, indicating that the current transoceanic disjunct distributions of the Peloridiidae are best explained by a Gondwanan vicariance hypothesis.     

Tethyan vicariance, relictualism and speciation: evidence from a global molecular phylogeny of the opisthobranch genus Bulla

Aim Our aims were: (1) to reconstruct a molecular phylogeny of the cephalaspidean opisthobranch genus Bulla, an inhabitant of shallow sedimentary environments; (2) to test if divergence times are consistent with Miocene and later vicariance among the four tropical marine biogeographical provinces; (3) to examine the phylogenetic status of possible Tethyan relict species; and (4) to infer the timing and causes of speciation events. Location Tropical and warm‐temperate regions of the Atlantic, Indo‐West Pacific, Australasia and eastern Pacific. Methods Ten of the 12 nominal species of Bulla were sampled, in a total sample of 65 individuals, together with cephalaspidean outgroups. Phylogenetic relationships were inferred by Bayesian analysis of partial sequences of the mitochondrial cytochrome c oxidase I (COI) and 16S rRNA and nuclear 28S rRNA genes. Divergence times and rates of evolution were estimated using uncorrelated relaxed‐clock Bayesian methods with fossil calibrations (based on literature review and examination of fossil specimens), implemented in beast . The geographical pattern of speciation was assessed by estimating the degree of overlap between sister lineages. Results Four clades were supported: Indo‐West Pacific (four species), Australasia (one species), Atlantic plus eastern Pacific (three species) and Atlantic (two species), with estimated mean ages of 35–46 Ma. Nominal species were monophyletic, but deep divergences were found within one Indo‐West Pacific and one West Atlantic species. Species‐level divergences occurred in the Miocene or earlier. The age of a sister relationship across the Isthmus of Panama was estimated at 7.9–32.1 Ma, and the divergence of a pair of sister species on either side of the Atlantic Ocean occurred 20.4–27.2 Ma. Main conclusions Fossils suggest that Bulla originated in the Tethys realm during the Middle Eocene. Average ages of the four main clades fall in the Eocene, and far pre‐date the 18–19 Ma closure of the Tethys Seaway. This discrepancy could indicate earlier vicariant events, selective extinction or errors of calibration. Similarly, the transisthmian divergence estimate far pre‐dates the uplift of the Panamanian Isthmus at about 3 Ma. Speciation events occurred in the Miocene, consistent with tectonic events in the central Indo‐West Pacific, isolation of the Arabian Sea by upwelling and westward trans‐Atlantic dispersal. Differences in habitat between sister species suggest that ecological speciation may also have played a role. The basal position of the Australasian species supports its interpretation as a Tethyan relict.