Trans‐Atlantic,trans‐Pacific and trans‐Indian Ocean dispersal in the small Gondwanan Laurales family Hernandiaceae

Aim To investigate the historical biogeography of the pantropical flowering plant family Hernandiaceae (Laurales), which today comprises 62 species in five genera. Location Hernandiaceae occur in Africa (9 species), Madagascar (4), the Neotropics (25), Australia (3), southern China, Indochina, Malesia, and on numerous Pacific Islands (32). These numbers include two widespread species, Hernandia nymphaeifolia, which ranges from East Africa to the Ogasawara Islands and New Caledonia, and Gyrocarpus americanus, thought to have a pantropical range. Methods We sampled 37 species from all genera, the widespread ones with multiple accessions, for a chloroplast DNA matrix of 2210 aligned nucleotides, and used maximum likelihood to infer species relationships. Divergence time estimation relied on an uncorrelated‐rates relaxed molecular clock calibrated with outgroup fossils of Lauraceae and Monimiaceae. Results The deepest split in the family is between a predominantly African–Madagascan–Malesian lineage comprising Hazomalania, Hernandia and Illigera, and an African–Neotropical lineage comprising Gyrocarpus and Sparattanthelium; this split may be 122 (110–134) Myr old. The stem lineages of the five genera date back at least to the Palaeocene, but six splits associated with transoceanic range disjunctions date only to the Oligocene and Miocene, implying long‐distance dispersal. It is inferred that Hernandia beninensis reached the West African islands of São Tomé and Bioko from the West Indies or the Guianas; Hernandia dispersed across the Pacific; and Illigera madagascariensis reached Madagascar from across the Indian Ocean. Main conclusions The disjunct ranges and divergence times of sister clades in the Hernandiaceae are partly congruent with the break‐up of West Gondwana, but mostly with later transoceanic dispersal. An exceptional ability to establish following prolonged oceanic dispersal may be largely responsible for the evolutionary persistence of this small clade.     

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.     

Australian biogeographical connections and the phylogeny of large genera in the plant family Myrtaceae

Aim To compare the phylogeny of the eucalypt and melaleuca groups with geological events and ages of fossils to discover the time frame of clade divergences. Location Australia, New Caledonia, New Guinea, Indonesian Archipelago. Methods We compare published molecular phylogenies of the eucalypt and melaleuca groups of the plant family Myrtaceae with geological history and known fossil records from the Cretaceous and Cenozoic. Results The Australasian eucalypt group includes seven genera, of which some are relictual rain forest taxa of restricted distribution and others are species‐rich and widespread in drier environments. Based on molecular and morphological data, phylogenetic analyses of the eucalypt group have identified two major clades. The monotypic Arillastrum endemic to New Caledonia is related in one clade to the more species‐rich Angophora, Corymbia and Eucalyptus that dominate the sclerophyll vegetation of Australia. Based on the time of rifting of New Caledonia from eastern Gondwana and the age of fossil eucalypt pollen, we argue that this clade extends back to the Late Cretaceous. The second clade includes three relictual rain forest taxa, with Allosyncarpia from Arnhem Land the sister taxon to Eucalyptopsis of New Guinea and the eastern Indonesian archipelago, and Stockwellia from the Atherton Tableland in north‐east Queensland. As monsoonal, drier conditions evolved in northern Australia, Arnhem Land was isolated from the wet tropics to the east and north during the Oligocene, segregating ancestral rain forest biota. It is argued also that the distribution of species in Eucalyptopsis and Eucalyptus subgenus Symphyomyrtus endemic in areas north of the stable edge of the Australian continent, as far as Sulawesi and the southern Philippines, is related to the geological history of south‐east Asia‐Australasia. Colonization (dispersal) may have been aided by rafting on micro‐continental fragments, by accretion of arc terranes onto New Guinea and by land brought into closer proximity during periods of low sea‐level, from the Late Miocene and Pliocene. The phylogenetic position of the few northern, non‐Australian species of Eucalyptus subgenus Symphyomyrtus suggests rapid radiation in the large Australian sister group(s) during this time frame. A similar pattern, connecting Australia and New Caledonia, is emerging from phylogenetic analysis of the Melaleuca group (Beaufortia suballiance) within Myrtaceae, with Melaleuca being polyphyletic. Main conclusion The eucalypt group is an old lineage extending back to the Late Cretaceous. Differentiation of clades is related to major geological and climatic events, including rifting of New Caledonia from eastern Gondwana, development of monsoonal and drier climates, collision of the northern edge of the Australian craton with island arcs and periods of low sea level. Vicariance events involve dispersal of biota.     

From East Gondwana to Central America: historical biogeography of the Alstroemeriaceae

Aim The Alstroemeriaceae is among 28 angiosperm families shared between South America, New Zealand and/or Australia; here, we examine the biogeography of Alstroemeriaceae to better understand the climatic and geological settings for its diversification in the Neotropics. We also compare Alstroemeriaceae with the four other Southern Hemisphere families that expanded from Patagonia to the equator, to infer what factors may have permitted such expansions across biomes. Location South America, Central America, Australia and New Zealand. Methods Three chloroplast genes, one mitochondrial gene and one nuclear DNA region were sequenced for 153 accessions representing 125 of the 200 species of Alstroemeriaceae from throughout the distribution range; 25 outgroup taxa were included to securely infer evolutionary directions and be able to use both ingroup and outgroup fossil constraints. A relaxed‐clock model relied on up to three fossil calibrations, and ancestral ranges were inferred using statistical dispersal–vicariance analysis (S‐DIVA). Southern Hemisphere disjunctions in the flowering plants were reviewed for key biological traits, divergence times, migration directions and habitats occupied. Results The obtained chronogram and ancestral area reconstruction imply that the most recent common ancestor of Colchicaceae and Alstroemeriaceae lived in the Late Cretaceous in southern South America/Australasia, the ancestral region of Alstroemeriaceae may have been South America/Antarctica, and a single New Zealand species is due to recent dispersal from South America. Chilean Alstroemeria diversified with the uplift of the Patagonian Andes c. 18 Ma, and a hummingbird‐pollinated clade (Bomarea) reached the northern Andes at 11–13 Ma. The South American Arid Diagonal (SAAD), a belt of arid vegetation caused by the onset of the Andean rain shadow 14–15 Ma, isolated a Brazilian clade of Alstroemeria from a basal Chilean/Argentinean grade. Main conclusions Only Alstroemeriaceae, Calceolariaceae, Cunoniaceae, Escalloniaceae and Proteaceae have expanded and diversified from Patagonia far into tropical latitudes. All migrated northwards along the Andes, but also reached south‐eastern Brazil, in most cases after the origin of the SAAD. Our results from Alstroemeria now suggest that the SAAD may have been a major ecological barrier in southern South America.     

Phylogenetic systematics of the gondwanan Empis macrorrhyncha group (Diptera,Empididae, Empidinae)

The Empis macrorrhyncha group (Diptera: Empididae) from cool to warm temperate areas of South America and Australia is diagnosed and cladistically analysed, and five new species, Empis animosa sp.n. , E. austera sp.n. , E. maculosa sp.n. , E. occidentalis sp.n. and E. pedivillosula sp.n. , are described. Cladistic analysis of 23 adult morphological characters for 14 species of the group generated a single tree of 28 steps (CI = 0.82; RI = 0.93). Monophyly was established on the basis of a single apomorphy, possession of a bilobed cercus of the male hypopygium. Three main clades were inferred: clade 1 included three Patagonian and a single southwestern Australian species; clade 2 included two species from southeastern Australia; clade 3 included a large Patagonian group of five species and a single southeastern Australian species. The E. fulvicollis complex (clade 1) is a sister‐group of the E. macrorrhyncha complex (clades 2 + 3). A provisional historical biogeographic hypothesis is advanced correlating the appearance of the South American and Australian sister lineages with the timing of the break‐up of Gondwana.     

Molecular evidence for long distance dispersal across the Southern Hemisphere in the Ganoderma applanatum-australe species complex (Basidiomycota)

We examined phylogeographic relationships in the cosmopolitan polypore fungus Ganoderma applanatum and allies, and conservatively infer a possible age of origin for these fungi. Results indicate that it is very unlikely that members of this species complex diversified before the break-up of Gondwana from Laurasia ca 120 M years ago, and also before the final separation of the Gondwanan landmasses from each other that was achieved about 66 M years ago. An earliest possible age of origin of 30 M years was estimated from nucleotide substitution rates in the 18S rDNA gene. Phylogenetic reconstruction of a worldwide sampling of ITS rDNA sequences reveals at least eight distinct clades that are strongly correlated with the geographic origin of the strains, and also correspond to mating groups. These include one Southern Hemisphere clade, one Southern Hemisphere–Eastern Asia clade, two temperate Northern Hemisphere clades, three Asian clades, and one neotropical clade. Geographically distant collections from the Southern Hemisphere shared identical ITS haplotypes, and an ITS recombinant was noted. Nested clade analysis of a parsimony network among isolates of the Southern Hemisphere clade indicated restricted gene flow with isolation-by-distance among the New Zealand, Australia–Tasmania, Chile–Argentine, and South Africa populations, suggesting episodic events of long-distance dispersal within the Southern Hemisphere. This study indicates that dispersal bias plays a more important role than generally admitted to explain the Southern Hemisphere distribution of many taxa, at least for saprobic fungi.     

Molecular dating of the 'Gondwanan' plant family Proteaceae is only partially congruent with the timing of the break-up of Gondwana

Aim  The flowering plant family Proteaceae is putatively of Gondwanan age, with modern and fossil lineages found on all southern continents. Here we test whether the present distribution of Proteaceae can be explained by vicariance caused by the break-up of Gondwana.
Location  Africa, especially southern Africa, Australia, New Zealand, South America, New Caledonia, New Guinea, Southeast Asia, Sulawesi, Tasmania.
Methods  We obtained chloroplast DNA sequence data from the rbc L gene, the rbc L- atp B spacer, and the atp B gene from leaf samples of forty-five genera collected from the field and from living collections. We analysed these data using Bayesian phylogenetic and molecular dating methods, with five carefully selected fossil calibration points to obtain age estimates for the nodes within the family.
Results  Four of eight trans-continental disjunctions of sister groups within our sample of the Proteaceae post-date the break-up of Gondwana. These involve independent lineages, two with an Africa-Australia disjunction, one with an Africa–South America disjunction, and one with a New Zealand–Australasia disjunction. The date of the radiation of the bird-pollinated Embothriinae corresponds approximately to the hypothesized date of origin of nectar-feeding birds in Australia.
Main conclusions  The findings suggest that disjunct distributions in Proteaceae result from both Gondwanan vicariance and transoceanic dispersal. Our results imply that ancestors of some taxa dispersed across oceans rather than rafting with Gondwanan fragments as previously thought. This finding agrees with other studies of Gondwanan plants in dating the divergence of Australian, New Zealand and New Caledonian taxa in the Eocene, consistent with the existence of a shared, ancestral Eocene flora but contrary to a vicariance scenario based on accepted geological knowledge.     

Craniid brachiopods: aspects of clade structure and distribution reflect continental drift (Brachiopoda: Craniiformea)

We present maximum likelihood and Bayesian inference relative time‐tree analyses of aligned gene sequences from a worldwide collection of craniiform brachiopods belonging to two genera, Novocrania and Neoancistrocrania. Sequences were obtained from one mitochondrial and three nuclear‐encoded ribosomal RNA genes from varying numbers of specimens. Data‐exploration by network (splits) analyses indicates that each gene identifies the same divergent clades and (with one minor exception) the same inter‐clade relationships. Neoancistrocrania specimens were found only in the Pacific Ocean, near Japan, on the Norfolk and Chesterfield Ridges, and near the Solomon Islands. The Novocrania clades, in approximate order of increasing distance from the root comprise 1. a ‘Northern’ clade of animals collected in the NE. Atlantic, W. Mediterranean and Adriatic; 2. a ‘Tethyan’ clade comprising animals from the E. Mediterranean, Cape Verde islands and the Caribbean (Belize and Jamaica); 3. a ‘NE. Pacific’ clade containing animals from Vancouver Island and from localities near Japan and south of Taiwan; 4. a ‘Southern’ clade that contains two widely separated subclades, one from New Zealand and the other with an extraordinarily wide distribution, ranging from near Japan in the north to the Chesterfield Ridge and Solomon Islands in the West, and in the East to the Galapagos Islands, the coast of South America (Chile) and Richardson seamount (off South Africa) in the South Atlantic. To the South, members of this clade were found in the Weddell, Scotia and Bellinghausen Antarctic Seas. The root of the extant craniid radiation was previously found (by relaxed‐clock analysis) to lie on the branch connecting the two genera so that, in effect, the one clade of Neoancistrocrania serves to polarise evolutionary relationships within the several clades of Novocrania. As previously suggested, all results confirm that Neoancistrocrania is sister to the ‘Northern’ Novocrania clade, and this leads to a proposal that Neoancistrocrania represents one extreme of a wide range of variation in ancestral ventral valve mineralisation, speciation (～90 Ma) resulting from competitive exclusion in rapidly‐growing reef environments. To the extent possible, the identified molecular clades are correlated with named species of Novocrania. The reproductive and population biology of craniid brachiopods is not well known, but from available evidence they are considered to have low‐dispersal potential and, except in enclosed localities such as cold‐water fjords, to have small effective population sizes, features which are consistent with the observed divergent populations in well‐separated localities. Exceptionally slow craniid molecular (rDNA) evolution is suggested by the short branch of Novocrania where it has been used as an outgroup for large‐scale analyses of metazoans. Slow molecular evolution is also indicated by the existence of a distinct Tethyan clade, reflecting restricted dispersal at former times, and by the uniform, short, genetic distances and exceptionally wide geographical distribution of the Southern clade. Thus, the geographical distribution and phylogenetic divergence of craniid brachiopods is an example of phylotectonics, in which relationships revealed by phylogenetic analyses reflect opportunities for dispersal and settlement that were created by tectonic plate movements associated, in this case, with opening and closure of Tethys and the breakup of Gondwana. Molecular dating of craniid divergences and radiochemical dating of tectonic events thus illuminate one another. © 2014 The Linnean Society of London     

Gondwanan evolution of the grass alliance of families (Poales)

Phylogenetic interrelationships among all 18 families of Poales were assessed by cladistic analysis of chloroplast DNA rbcL and atpB sequences from 65 species. There are two well-supported main clades; the graminoid clade with Poaceae (grasses), Anarthriaceae, Centrolepidaceae, Ecdeiocoleaceae, Flagellariaceae, Joinvilleaceae, and Restionaceae; and the cyperoid clade with Cyperaceae, Juncaceae, and Thurniaceae. A sister group relationship between Poaceae and Ecdeiocoleaceae is identified with strong support. The sister group of this pair is Joinvilleaceae. These relationships help in elucidating the evolution of grasses and the grass spikelet. Dating of the tree was done by nonparametric rate smoothing of rbcL molecular evolution. Most Poales families date back to the Cretaceous >65 million years ago (mya). Dispersal-vicariance analysis indicates that the Poales originated in South America, the cyperoid clade in West Gondwana (South America or Africa), and the graminoid clade in East Gondwana (Australia). The Trans-Antarctic connection between South America and Australia, and its breakup about 35 mya, probably influenced the evolution of the Poales and the graminoid clade in particular, leading to vicariance between the continents, but the separation of Africa from the other Gondwanan areas, completed about 105 mya, is too old for such a relation.     

The phylogenetic placement and biogeographical origins of the New Zealand stick insects (Phasmatodea)

The Lanceocercata are a clade of stick insects (Phasmatodea) that have undergone an impressive evolutionary radiation in Australia, New Caledonia, the Mascarene Islands and areas of the Pacific. Previous research showed that this clade also contained at least two of the nine New Zealand stick insect genera. We have constructed a phylogeny of the Lanceocercata using 2277 bp of mitochondrial and nuclear DNA sequence data to determine whether all nine New Zealand genera are indeed Lanceocercata and whether the New Zealand fauna is monophyletic. DNA sequence data were obtained from mitochondrial cytochrome oxidase subunits I and II and the nuclear large subunit ribosomal RNA and histone subunit 3. These data were subjected to Bayesian phylogenetic inference under a partitioned model and maximum parsimony. The resulting trees show that all the New Zealand genera are nested within a large New Caledonian radiation. The New Zealand genera do not form a monophyletic group, with the genus Spinotectarchus Salmon forming an independent lineage from the remaining eight genera. We analysed Lanceocercata apomorphies to confirm the molecular placement of the New Zealand genera and to identify characters that confirm the polyphyly of the fauna. Molecular dating analyses under a relaxed clock coupled with a Bayesian extension to dispersal‐vicariance analysis was used to reconstruct the biogeographical history for the Lanceocercata. These analyses show that Lanceocercata and their sister group, the Stephanacridini, probably diverged from their South American relatives, the Cladomorphinae, as a result of the separation of Australia, Antarctica and South America. The radiation of the New Caledonian and New Zealand clade began 41.06 million years ago (mya, 29.05–55.40 mya), which corresponds to a period of uplift in New Caledonia. The main New Zealand lineage and Spinotectarchus split from their New Caledonian sister groups 33.72 (23.9–45.62 mya) and 29.9 mya (19.79–41.16 mya) and began to radiate during the late Oligocene and early Miocene, probably in response to a reduction in land area and subsequent uplift in the late Oligocene and early Miocene. We discuss briefly shared host plant patterns between New Zealand and New Caledonia. Because Acrophylla sensu Brock & Hasenpusch is polyphyletic, we have removed Vetilia Stål from synonymy with Acrophylla Gray.     

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.     

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.     

Dispersal,vicariance, and the Late Cretaceous to early tertiary land mammal biogeography from South America to Australia

A review of paleontological, phyletic, geophysical, and climatic evidence leads to a new scenario of land mammal dispersal among South America, Antarctica, and Australia in the Late Cretaceous to early Tertiary epochs. New fossil land vertebrate material has been recovered from all three continents in recent years. As regards Gondwana, the present evidence suggests that monotreme mammals and ratite birds are of Mesozoic origin, based on both geochronological and phyletic grounds. The occurrence of monotremes in the early Paleocene (ca. 62 Ma) faunas of Patagonia and of ratites in late Eocene (ca. 41-37 m.y.) faunas of Seymour Island (Antarctic Peninsula) probably is an artifact of a much older and widespread Gondwana distribution prior to the Late Cretaceous Epoch. Except for South American microbiotheres being australidelphians, marsupial faunas of South America and Australia still are fundamentally disjunct. New material from Seymour Island (Microbiotheriidae) indicates the presence there of a derived taxon that resides in a group that is the sister taxon of most Australian marsupials. There is no compelling evidence that dispersal between Antarctica and Australia was as recent as ca. 41 Ma or later. In fact, the derived marsupial and placental land mammal fauna of Seymour Island shows its greatest affinity with Patagonian forms of Casamayoran age (ca. 51–54 m.y.). This suggests an earlier dispersal of more plesiomorphic marsupials from Patagonia to Australia via Antarctica, and vicariant disjunction subsequently. This is consistent with geophysical evidence that the South Tasman Rise was submerged by 64 Ma and with geological evidence that a shallow water marine barrier was present from then onward. The scenario above is consistent with molecular evidence suggesting that australidelphian bandicoots, dasyurids, and diprotodontians were distinct and present in Australia at least as early as the 63-Ma-old australidelphian microbiotheres and the ancient but not basal australidelphian,Andinodelphys, in the Tiupampa Fauna of Bolivia. Land mammal dispersal to Australia typically has been considered to be at a low level of probability (e.g., by sweepstakes dispersal). This study suggests that the marsupial colonizers of Australia included already recognizable members of the Peramelina, Dasyuromorphia, and Diprotodontia, at least, and entered via a filter route rather than by a sweepstakes dispersal.To whom correspondence should be addressed.     

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.     

The biogeography of Gunnera L.: vicariance and dispersal

Aim The genus Gunnera is distributed in South America, Africa and the Australasian region, a few species reaching Hawaii and southern Mexico in the North. A cladogram was used to (1) discuss the biogeography of Gunnera and (2) subsequently compare this biogeographical pattern with the geological history of continents and the patterns reported for other Southern Hemisphere organisms. Location Africa, northern South America, southern South America, Tasmania, New Zealand, New Guinea/Malaya, Hawaii, North America, Antarctica. Methods A phylogenetic analysis of twenty‐six species of Gunnera combining morphological characters and new as well as published sequences of the ITS region, rbcL and the rps16 intron, was used to interpret the biogeographical patterns in Gunnera. Vicariance was applied in the first place and dispersal was only assumed as a second best explanation. Results The Uruguayan/Brazilian Gunnera herteri Osten (subgenus Ostenigunnera Mattfeld) is sister to the rest of the genus, followed sequentially upwards by the African G. perpensa L. (subgenus Gunnera), in turn sister to all other, American and Australasian, species. These are divided into two clades, one containing American/Hawaiian species, the other containing all Australasian species. Within the Australasian clade, G. macrophylla Blume (subgenus Pseudogunnera Schindler), occurring in New Guinea and Malaya, is sister to a clade including the species from New Zealand and Tasmania (subgenus Milligania Schindler). The southern South American subgenus Misandra Schindler is sister to a clade containing the remaining American, as well as the Hawaiian species (subgenus Panke Schindler). Within subgenus Panke, G. mexicana Brandegee, the only North American species in the genus, is sister to a clade wherein the Hawaiian species are basal to all south and central American taxa. Main conclusions According to the cladogram, South America appears in two places, suggesting an historical explanation for northern South America to be separate from southern South America. Following a well‐known biogeographical pattern of vicariance, Africa is the sister area to the combined southern South America/Australasian clade. Within the Australasian clade, New Zealand is more closely related to New Guinea/Malaya than to southern South America, a pattern found in other plant cladograms, contradictory to some of the patterns supported by animal clades and by the geological hypothesis, respectively. The position of the Tasmanian G. cordifolia, nested within the New Zealand clade indicates dispersal of this species to Tasmania. The position of G. mexicana, the only North American species, as sister to the remaining species of subgenus Panke together with the subsequent sister relation between Hawaii and southern South America, may reflect a North American origin of Panke and a recolonization of South America from the north. This is in agreement with the early North American fossil record of Gunnera and the apparent young age of the South American clade.     

Fruit structure and systematics of Monimiaceae s.s. (Laurales)

Fruit structure (anatomy) was studied in 27 species of 15 genera of Monimiaceae s.s. Almost all have apocarpous gynoecia, with the carpels more or less surrounded by a floral cup. The fruitlets are presented on the opened floral cup, which, depending on its pre‐ and post‐floral development, differentially contributes to the attractive part of the mature fruit. Morphologically similar fruits may differ conspicuously in anatomical structure. Based on anatomical characters two different fruit forms were found: drupe(let)s (with compact sclerenchymatic endocarp forming a stone: putamen) and berry(let)s (with parenchymatic endocarp, and mesocarp parenchyma containing isolated sclereid nests). Four types of drupelets differing by the endocarp structure were tentatively distinguished: (1) the Monimia‐type has a many‐cell‐layered putamen of large isodiametric sclereids, interrupted on the ventral side by few radial rows of small sclereids; (2) the Hortonia‐type has a few‐cell‐layered putamen of isodiametric, especially thick‐walled sclereids – it may be composed of two lateral halves, i.e. with the sclerenchyma partially interrupted on the ventral and dorsal sides (but without rows of small sclereids); (3) the Mollinedia‐type has a few‐cell‐layered putamen, with more or less radially elongate sclereids with wavy cell walls; and (4) the Hedycarya‐type has a one‐cell‐layered putamen of pronouncedly radially elongate sclereids with wavy cell walls. Drupelets of some taxa with a single‐cell‐layered endocarp with only weakly thickened cell walls may represent a transition from drupelets to berrylets. The fruit structure supports three major clades recognized earlier by morphological studies and by molecular phylogenetic analyses: (1) Monimioideae (Monimia‐type drupelets), (2) Hortonieae of Mollinedioideae (Hortonia‐type drupelets), and (3) the remainder of Mollinedioideae (Hedycarya‐ and Mollinedia‐types) and berrylets. Fruit structure also supports the close relationship of Monimiaceae and Lauraceae. © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 153 , 265–285.     

South Pacific biogeography,tectonic calibration,and pre‐drift tectonics: cladogenesis in Abrotanella (Asteraceae)

Abrotanella is the basal genus in the large tribe Senecioneae (Asteraceae) and has a disjunct distribution in Australasia and South America. A recent molecular phylogeny of the genus was used to investigate whether the main biogeographical patterns in the group could be related to the region's tectonic history in a coherent way. The phylogenetic/biogeographical breaks and overlaps in the genus imply a series of vicariance and range expansion events. Each of these can be related to one of the main tectonic events in the region, including assembly of the New Zealand terranes, crustal extension, and magmatism in Gondwana that preceded seafloor spreading, opening of the Tasman and Pacific basins, and transcurrent movement on the New Zealand Alpine fault. The coincident sequence indicates that pre‐drift tectonics and magmatism have been more important for the origin of trans‐Tasman and trans‐Pacific groups than the final rifting of Gondwana that led to their disjunction. For example, during the pre‐drift phase of break‐up, the Whitsunday volcanic province of Australia and the Median Batholith of New Zealand formed a large, active igneous belt. Its distribution is aligned with the break between New Zealand–south‐eastern Australia clades, and New Zealand–New Guinea clades. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Historical biogeography of Southeast Asia and the West Pacific,or the generality of unrooted area networks as historical biogeographic hypotheses

Aim Unrooted area networks are perhaps a general way in which different historical biogeographical patterns may be combined. Location Southeast Asia up to the West Pacific, Australia, South America. Methods Unrooted area networks based on Primary Brooks Parsimony Analysis of different data sets of Southeast Asian–West Pacific, Australian and South American clades. Results A large Brooks Parsimony historical (cladistic) biogeographic analysis of Southeast Asia and the West Pacific gave a meaningful result when all clades (representing different historical biogeographic patterns) were united into one matrix and an unrooted area network was produced. This network showed geographically adjacent areas as neighbours, which is interpreted as clades dispersing and speciating as soon as areas rafted towards each other. This pseudo‐vicariance mechanism, together with the very limited, mainly linear dispersal possibilities, a few large, widespread clades with many endemic species, and the large overlap in distributions displayed by different patterns, may explain the peculiar result. When applied to examples from other areas (bird data from Australia and South America), unrooted area networks for all data perform very poorly. Main conclusions Unrooted historical general area networks are not universally applicable. In general, it is better to split historical patterns a priori and analyse them separately.     

Systematics, biogeography and diversification of the Indo-Australian genus Delias Hübner (Lepidoptera: Pieridae): phylogenetic evidence supports an 'out-of-Australia' origin

Abstract Two alternative hypotheses for the origin of butterflies in the Australian Region, that elements dispersed relatively recently from the Oriental Region into Australia (northern dispersal hypothesis) or descended from ancient stocks in Gondwana (southern vicariance hypothesis), were tested using methods of cladistic vicariance biogeography for the Delias group, a diverse and widespread clade in the Indo‐Australian Region. A phylogenetic hypothesis of the twenty‐four species‐groups recognized currently in Delias and its sister genus Leuciacria is inferred from molecular characters generated from the nuclear gene elongation factor‐1 alpha (EF‐1α) and the mitochondrial genes cytochrome oxidase subunits I and II (COI/COII) and NADH dehydrogenase 5 (ND5). Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined dataset (3888 bp, 1014 parsimony informative characters) confirmed the monophyly of Delias and recovered eight major lineages within the genus, informally designated the singhapura, belladonna, hyparete, chrysomelaena, eichhorni, cuningputi, belisama and nigrina clades. Species‐group relationships within these clades are, in general, concordant with current systematic arrangements based on morphology. The major discrepancies concern the placement of the aganippe, belisama and chrysomelaena groups, as well as several species‐groups endemic to mainland New Guinea. Two species (D. harpalyce (Donovan), D. messalina Arora) of uncertain group status are currently misplaced based on strong evidence of paraphyly, and are accordingly transferred to the nigrina and kummeri groups, respectively. Based on this phylogeny, a revised systematic classification is presented at the species‐group level. An historical biogeographical analysis of the Delias group revealed that the most parsimonious reconstruction is an origin in the Australian Region, with at least seven dispersal events across Wallacea to the Oriental Region. The eight major clades of Delias appear to have diverged rapidly following complete separation of the Australian plate from Gondwana and its collision with the Asian plate in the late Oligocene. Further diversification and dispersal of Delias in the Miocene–Pliocene are associated with major geological and climatic changes that occurred in Australia–New Guinea during the late Tertiary. The ‘out‐of‐Australia’ hypothesis for the Delias group supports an origin of the Aporiina in southern Gondwana (southern vicariance hypothesis), which proposes that the ancestor of Delias + Leuciacria differentiated vicariantly on the Australian plate.