Phylogeny and biogeography of the sandalwoods (Santalum, Santalaceae): repeated dispersals throughout the Pacific

Results of the first genus-wide phylogenetic analysis for Santalum (Santalaceae), using a combination of 18S-26S nuclear ribosomal (ITS, ETS) and chloroplast (3' trnK intron) DNA sequences, provide new perspectives on relationships and biogeographic patterns among the widespread and economically important sandalwoods. Congruent trees based on maximum parsimony, maximum likelihood, and Bayesian methods support an origin of Santalum in Australia and at least five putatively bird-mediated, long-distance dispersal events out of Australia, with two colonizations of Melanesia, two of the Hawaiian Islands, and one of the Juan Fernandez Islands. The phylogenetic data also provide the best available evidence for plant dispersal out of the Hawaiian Islands to the Bonin Islands and eastern Polynesia. Inability to reject rate constancy of Santalum ITS evolution and use of fossil-based calibrations yielded estimates for timing of speciation and colonization events in the Pacific, with dates of 1.0-1.5 million yr ago (Ma) and 0.4-0.6 Ma for onset of diversification of the two Hawaiian lineages. The results indicate that the previously recognized sections Polynesica, Santalum, and Solenantha, the widespread Australian species S. lanceolatum, and the Hawaiian species S. freycinetianum are not monophyletic and need taxonomic revision, which is currently being pursued.     

Dispersal and adaptive radiation of Bidens (Compositae) across the remote archipelagoes of Polynesia

The genus Bidens (Compositae) comprises c. 230 species distributed across five continents, with the 41 Polynesian species displaying the greatest ecomorphological variation in the group. However, the genus has had a long and complicated taxonomic history, and its phylogenetic and biogeographic history are poorly understood. To resolve the evolutionary history of the Polynesian Bidens, 152 individuals representing 91 species were included in this study, including 39 of the 41 described species from Polynesia. Four chloroplast and two nuclear DNA markers were utilized to estimate phylogenetic relationships, divergence times, and biogeographic history. Bidens was found to be polyphyletic within Coreopsis, consistent with previous assessments. The Polynesian radiation was resolved as monophyletic, with the initial dispersal into the Pacific possibly from South America to either the Hawaiian or Marquesas Islands. From the Marquesas, Bidens dispersed to the Society Islands, and ultimately to the Austral Islands. The initial diversification of the crown group in the Pacific is estimated to have occurred ~1.63 mya (0.74–2.72, 95% HPD), making Polynesian Bidens among the youngest and most rapid plant diversification events documented in the Pacific. Our findings suggest that relatively rare long‐distance dispersal and founder‐event speciation, coupled with subsequent loss of dispersal potential and within‐island speciation, can explain the repeated and explosive adaptive radiation of Bidens throughout the archipelagoes of Polynesia.     

Island hopping across the central Pacific: mitochondrial DNA detects sequential colonization of the Austral Islands by crab spiders (Araneae: Thomisidae)

Aim Phylogenetic studies concerning island biogeography have been concentrated in a fraction of the numerous hot‐spot archipelagos contained within the Pacific Ocean. In this study we investigate relationships among island populations of the thomisid spider Misumenops rapaensis Berland, 1934 across the Austral Islands, a remote and rarely examined southern Pacific hot‐spot archipelago. We also assess the phylogenetic position of M. rapaensis in relation to thomisids distributed across multiple Polynesian archipelagos in order to evaluate the proposed hypothesis that thomisid spiders colonized Polynesia from multiple and opposing directions. The data allow an examination of genetic divergence and species accumulation in closely related lineages distributed across four Polynesian archipelagos. Location The study focused on four Polynesian hot‐spot archipelagos: the Austral, Hawaiian, Marquesan and Society islands. Methods Mitochondrial DNA sequences comprising c. 1400 bp (portions of cytochrome oxidase subunit I, ribosomal 16S and NADH dehydrogenase subunit I) were obtained from thomisid spiders (64 specimens, representing 33 species) collected in the Australs, the Hawaiian Islands, the Society Islands, the Marquesas, Tonga, Fiji, New Zealand, New Caledonia and North and South America. Phylogenetic analyses using parsimony, maximum‐likelihood and Bayesian approaches were employed to resolve relationships of M. rapaensis to other Polynesian Misumenops and across the Austral Islands. Results Rather than grouping with other Misumenops spp. from the archipelagos of the Society Islands, Marquesas and Hawaiian Islands, M. rapaensis appears more closely related to Diaea spp. from Tonga, Fiji, New Zealand and New Caledonia. Phylogenetic analyses strongly support M. rapaensis as monophyletic across the Austral Islands. Misumenops rapaensis sampled from the two older islands (Rurutu and Tubuai) form reciprocally monophyletic groups, while individuals from the younger islands (Raivavae and Rapa) are paraphyletic. Across the Austral Islands, M. rapaensis exhibits a surprising level of genetic divergence (maximally 11.3%), an amount nearly equivalent to that found across the 16 examined Hawaiian species (14.0%). Main conclusions Although described as a single morphologically recognized species, our results suggest that M. rapaensis comprises multiple genetically distinct lineages restricted to different Austral Islands. Phylogenetic relationships among the island populations are consistent with sequential colonization of this lineage down the Austral archipelago toward younger islands. Analyses support the hypothesis that thomisid spiders colonized the central Pacific multiple times and suggest that M. rapaensis arrived in the Austral Islands from a westward direction, while Misumenops found in neighbouring archipelagos appear to be more closely related to New World congeners to the east. Finally, our data detect asymmetrical rates of morphological evolution and species diversification following colonization of four different Polynesian archipelagos.     

Convergent evolution of morphology and habitat use in the explosive Hawaiian fancy case caterpillar radiation

Species occurring in unconnected, but similar habitats and under similar selection pressures often display strikingly comparable morphology, behaviour and life history. On island archipelagos where colonizations and extinctions are common, it is often difficult to separate whether similar traits are a result of in situ diversification or independent colonization without a phylogeny. Here, we use one of Hawaii's most ecologically diverse and explosive endemic species radiations, the Hawaiian fancy case caterpillar genus Hyposmocoma, to test whether in situ diversification resulted in convergence. Specifically, we examine whether similar species utilizing similar microhabitats independently developed largely congruent larval case phenotypes in lineages that are in comparable, but isolated environments. Larvae of these moths are found on all Hawaiian Islands and are characterized by an extraordinary array of ecomorphs and larval case morphology. We focus on the ‘purse cases’, a group that is largely specialized for living within rotting wood. Purse cases were considered a monophyletic group, because morphological, behavioural and ecological traits appeared to be shared among all members. We constructed a phylogeny based on nuclear and mitochondrial DNA sequences from 38 Hyposmocoma species, including all 14 purse case species and 24 of non‐purse case congeners. Divergence time estimation suggests that purse case lineages evolved independently within dead wood and developed nearly identical case morphology twice: once on the distant Northwest Hawaiian Islands between 15.5 and 9 Ma and once on the younger main Hawaiian Islands around 3.0 Ma. Multiple ecomorphs are usually found on each island, and the ancestral ecomorph of Hyposmocoma appears to have lived on tree bark. Unlike most endemic Hawaiian radiations that follow a clear stepwise progression of colonization, purse case Hyposmocoma do not follow a pattern of colonization from older to younger island. We postulate that the diversity of microhabitats and selection from parasitism/predation from endemic predators may have shaped case architecture in this extraordinary endemic radiation of Hawaiian insects.     

Progressive island colonization and ancient origin of Hawaiian Metrosideros (Myrtaceae)

Knowledge of the evolutionary history of plants that are ecologically dominant in modern ecosystems is critical to understanding the historical development of those ecosystems. Metrosideros is a plant genus found in many ecological and altitudinal zones throughout the Pacific. In the Hawaiian Islands, Metrosideros polymorpha is an ecologically dominant species and is also highly polymorphic in both growth form and ecology. Using 10 non-coding chloroplast regions, we investigated haplotype diversity in the five currently recognized Hawaiian Metrosideros species and an established out-group, Metrosideros collina, from French Polynesia. Multiple haplotype groups were found, but these did not match morphological delimitations. Alternative morphologies sharing the same haplotype, as well as similar morphologies occurring within several distinct island clades, could be the result of developmental plasticity, parallel evolution or chloroplast capture. The geographical structure of the data is consistent with a pattern of age progressive island colonizations and suggests de novo intra-island diversification. If single colonization events resulted in a similar array of morphologies on each island, this would represent parallel radiations within a single, highly polymorphic species. However, we were unable to resolve whether the pattern is instead explained by ancient introgression and incomplete lineage sorting resulting in repeated chloroplast capture. Using several calibration methods, we estimate the colonization of the Hawaiian Islands to be potentially as old as 3.9 (-6.3) Myr with an ancestral position for Kaua'i in the colonization and evolution of Metrosideros in the Hawaiian Islands. This would represent a more ancient arrival of Metrosideros to this region than previous studies have suggested.     

Multiple origins of Hawaiian drosophilids: Phylogeography of Scaptomyza Hardy (Diptera: Drosophilidae)

Scaptomyza is a highly diversified genus in the family Drosophilidae, having undergone an explosive radiation, along with the Hawaiian‐endemic genus Idiomyia in the Hawaiian Islands: about 60% of 269 Scaptomyza species so far described are endemic to the Hawaiian Islands. Two hypotheses have been proposed for the origin and diversification of Hawaiian drosophilids. One is the “single Hawaiian origin” hypothesis: Scaptomyza and Idiomyia diverged from a single common ancestor that had once colonized the Hawaiian Islands, and then non‐Hawaiian Scaptomyza migrated back to continents. The other is the “multiple origins” hypothesis: Hawaiian Scaptomyza and Idiomyia derived from different ancestors that independently colonized the Hawaiian Islands. A key issue for testing these two hypotheses is to clarify the phylogenetic relationships between Hawaiian and non‐Hawaiian species in Scaptomyza. Toward this goal, we sampled additional non‐Hawaiian Scaptomyza species, particularly in the Old World, and determined the nucleotide sequences of four mitochondrial and seven nuclear genes for these species. Combining these sequence data with published data for 79 species, we reconstructed the phylogeny and estimated ancestral distributions and divergence times. In the resulting phylogenetic trees, non‐Hawaiian Scaptomyza species were interspersed in two Hawaiian clades. From a reconstruction of ancestral biogeography, we inferred that Idiomyia and Scaptomyza diverged outside the Hawaiian Islands and then independently colonized the Hawaiian Islands, twice in Scaptomyza, thus supporting the “multiple origins” hypothesis.     

Historical biogeography of Melicope (Rutaceae) and its close relatives with a special emphasis on Pacific dispersals

The genus Melicope (Rutaceae) occurs on most Pacific archipelagos and is perfectly suited to study Pacific biogeography. The main goal was to infer the age, geographic origin and colonization patterns of Melicope and its relatives. We sequenced three nuclear and two plastid markers for 332 specimens that represent 164 species in 16 genera of Rutaceae. Phylogenetic reconstruction, molecular dating, ancestral area reconstruction and diversification analyses were carried out. The two main clades (Acronychia‐Melicope and Euodia) originated in Australasia and their crown ages are dated to the Miocene. Diversification rates differed among the subclades and were lowest in the Euodia lineage and highest in the Hawaiian Melicope lineage. The Malagasy and Mascarene species form a clade, which split from its SE Asian relatives in the Pliocene/Pleistocene. At least eight colonizations to the Pacific islands occurred. The timing of all colonizations except for the Hawaiian group is congruent with age of the island ages. Australia, New Guinea and New Caledonia have been the source of colonizations into the Pacific islands in the Melicope clade. Melicope shows high dispersability and has colonized remote archipelagos such as the Austral and Marquesas Islands each twice. Colonization of islands of the Hawaiian‐Emperor seamount chain likely predates the ages of the current main islands, and the initial colonization to Kaua'i occurred after the splitting of the Hawaiian lineage into two subclades. Wider ecological niches and adaptations to bird‐dispersal likely account for the much higher species richness in the Acronychia‐Melicope clade compared to the Euodia clade.     

Radiation following long‐distance dispersal: the contributions of time,opportunity and diaspore morphology in Sicyos (Cucurbitaceae)

Aim  To infer the most plausible explanations for the presence of 14 species of the Neotropical cucurbit genus Sicyos on the Hawaiian Islands, two on the Galápagos Islands, two in Australia, and one in New Zealand. Location  Neotropics, the Hawaiian and Galápagos archipelagos, Australia and New Zealand. Methods  We tested long‐problematic generic boundaries in the tribe Sicyoeae and reconstructed the history of Sicyos using plastid and nuclear DNA sequences from 87 species (many with multiple accessions) representing the group’s generic and geographic diversity. Maximum likelihood and Bayesian approaches were used to infer relationships, divergence times, biogeographic history and ancestral traits. Results  Thirteen smaller genera, including Sechium, are embedded in Sicyos, which when re‐circumscribed as a monophyletic group comprises 75 species. The 14 Hawaiian species of Sicyos descended from a single ancestor that arrived c. 3 million years ago (Ma), Galápagos was reached twice at c. 4.5 and 1 Ma, the species in Australia descended from a Neotropical ancestor (c. 2 Ma), and New Zealand was reached from Australia. Time since arrival thus does not correlate with Sicyos species numbers on the two archipelagos. Main conclusions  A plausible mechanism for the four trans‐Pacific dispersal events is adherence to birds of the tiny hard fruit with retrorsely barbed spines found in those lineages that underwent long‐distance migrations. The Hawaiian clade has lost these spines, resulting in a lower dispersal ability compared with the Galápagos and Australian lineages, and perhaps favouring allopatric speciation.     

The origin and number of introductions of the Hawaiian endemic Silene species (Caryophyllaceae)

The Hawaiian endemic Silene are a small group of woody or semiwoody representatives from a large, predominantly herbaceous, species-rich genus. We here investigated the origin and number of introductions of the endemic Hawaiian Silene based on phylogenetic relationships inferred from DNA sequences from both the plastid (the rps16 intron) and the nuclear (ribosomal internal transcribed sequences, ITS, and intron 23 of the RPB2 gene) genomes. Silene antirrhina, a widespread weedy American annual, is strongly supported as sister to a monophyletic group consisting of the Hawaiian Silene, indicating a single colonization event. There are no obvious morphological similarities between S. antirrhina and any of the species of Hawaiian Silene. Our results suggest an American origin for the Hawaiian endemics because that would require only a single trans-ocean dispersal. Two of the Hawaiian endemics (S. struthioloides and S. hawaiiensis) that form a subclade in the analyses have evolved woodiness after introduction to the Hawaiian Islands. Our results contribute to other recent results based on molecular phylogenetics that emphasize the American continent as a source area for the Hawaiian flora and support a striking morphological radiation and evolution of woodiness from a single introduction to the archipelago.     

Evolution of Cyrtandra (Gesneriaceae) in the Pacific Ocean: the origin of a supertramp clade

Cyrtandra comprises at least 600 species distributed throughout Malesia, where it is known for many local endemics and in Polynesia and Micronesia, where it is present on most island groups, and is among the most successfully dispersing genera of the Pacific. To ascertain the origin of the oceanic Pacific island species of Cyrtandra, we sequenced the internal transcribed spacers of nuclear ribosomal DNA of samples from throughout its geographical range. Because all oceanic Pacific island species form a well-supported clade, these species apparently result from a single initial colonization into the Pacific, possibly by a species from the eastern rim of SE Asia via a NW-to-SE stepping stone migration. Hawaiian species form a monophyletic group, probably as a result of a single colonization. The Pacific island clade of Cyrtandra dispersed across huge distances, in contrast to the apparent localization of the SE Asian clades. Although highly vagile, the Pacific clade is restricted to oceanic islands. Individual species are often endemic to a single island, characteristic of the "supertramp" life form sensu Diamond (1974, Science 184: 803-806). The evolution of fleshy fruit within Cyrtandra provided an adaptation for colonization throughout the oceanic Pacific via bird dispersal from a single common ancestor.     

BIOGEOGRAPHY OF CLADOPHOROPSIS MEMBRANACEA (CHLOROPHYTA) BASED ON COMPARISONS OF NUCLEAR rDNA ITS SEQUENCES1

Nucleotides were compared at 988 sites, spanning both internal transcribed spacers (ITS1 and ITS2) of the nuclear ribosomal DNA, among 17 isolates of the green alga Cladophoropsis membranacea (Hofman Bang ex C. Agardh) Boergesen and two isolates of Struvea anastomosans (Harvey) Piccone and Grunow. Collections were made from Bonaire, Curaçao, St. Croix, the Canary Islands, the Cape Verde Islands, Mauritania, Syria, the Red Sea, Okinawa, and Hawaii. Two nucleotide substitutions were found between the sequenced coding regions of C. membranacea and S. anastomosans. Of the 720 nucleotides compared in ITS1 and ITS2, an average of 6.7% sequence divergence was found within C. membranacea and 17.4% between C. membranacea and S. anastomosans. Sequences were analyzed using maximum parsimony. Phylogenetic hypotheses were compared with the biogeographic hypothesis of an east-west Tethyan vicariance. Results of the study allow the identification of widely dispersed biogeographic populations, the identification of an underlying Tethyan imprint, and support for the hypothesis that dispersal has occurred between the Caribbean and the tropical eastern Atlantic. These results demonstrate the usefulness of ITS sequences at the sub-specific level in C. membranacea.     

Four new species of Rhodophyceae from Fiji,Polynesia and Vanuatu,South Pacific

Four new species of Rhodophyceae are described from the South Pacific, with type localities in Fiji, French Polynesia and Vanuatu. Chondria bullata from the Tuamotus (French Polynesia), Vanuatu, Palmerston Atoll (Cook Islands) and Fiji is unique owing to its non‐constricted axes with markedly protruding, bubble‐like cortical cells. Halymenia nukuhivensis, from the Marquesas Islands in French Polynesia, is distinguished from others in the genus by its dichotomous, papery blades issued from a strap‐shaped basal region, and the equal proportion of anti‐clinal, periclinal and oblique filaments in its medullary layer. Jania articulata, so far known only from the Tuamotus in French Polynesia and Manihiki in the Northern Cook Islands, superficially resembles the genus Amphiroa with its articulated branches with numerous genicula between successive dichotomies, and its large axis diameter. Meristotheca peltata from the Fiji Islands is unique among the genus by its distinctly peltate, erect habit. The recent high number of newly described species from the South Pacific region emphasizes the need for more in‐depth surveys, particularly in deeper outer reef slope habitats, which remain for the most part unexplored and could yield particularly interesting new taxa or distributional records.     

Evolution of insular Pacific Pittosporum (Pittosporaceae): origin of the Hawaiian radiation.

We investigated the origin of Hawaiian Pittosporum and their relationship to other South Pacific Pittosporum species using internal transcribed spacer sequences of nuclear ribosomal DNA. We performed both maximum-parsimony and maximum-likelihood analyses, which produced congruent results. Sequence divergence was 0.0% between Hawaiian members of Pittosporum. These taxa formed a strongly supported clade, suggesting a single colonization event followed by phyletic radiation. Sister to the Hawaiian clade were two South Pacific species, P. yunckeri from Tonga and P. rhytidocarpum from Fiji. This result presents convincing evidence for a South Pacific origin of Hawaiian Pittosporum. Our results also identify a monophyletic group comprising three species representing the Fijian Province and East Polynesia, two introductions onto New Caledonia, and at least one (but possibly two) introduction(s) onto New Zealand. Whether the New Zealand taxa form a monophyletic group is unclear from these data. Previous morphologically based hypotheses, however, suggest the presence of four different lineages occupying New Zealand. The nonmonophyly of the New Caledonian species was not surprising based on the extent of their morphological diversity. Although this latter result is not strongly supported, these species are morphologically complex and are currently the subject of taxonomic revision and molecular systematic analyses.     

Origin of the Hawaiian endemic mints within North American Stachys (Lamiaceae)

The Hawaiian endemic mints constitute a major island radiation, displaying a remarkable diversity of floral, fruit, and vegetative features. Haplostachys and Phyllostegia have flowers associated with insect pollination, whereas Stenogyne has flowers typical of bird pollination. The three genera had been thought to be closely related to East Asian members of Lamioideae tribe Prasieae because of the fleshy nutlets borne by Phyllostegia and Stenogyne. We evaluated the origins of the Hawaiian mints using phylogenetic analyses of DNA sequence data from the plastid rbcL and trnL intron loci and the nuclear ribosomal 5S nontranscribed spacer. The Hawaiian genera were found to be monophyletic but deeply nested inside another lamioid genus, Stachys. In particular, they were found to be most closely related to a group of temperate North American Stachys from the Pacific coast, suggesting that the Hawaiian mints derived from a single colonization event from western North America to the Hawaiian Islands. Furthermore, Stachys, which contains amphiatlantic and transberingian clades, was found to be polyphyletic, with some species more closely related to Gomphostemma, Phlomidoschema, Prasium, and Sideritis than to other species of Stachys. Based on chromosomal evidence and our phylogenetic analyses, we hypothesize that the Hawaiian mints may be polyploid hybrids whose reticulate genomes predate the Hawaiian dispersal event and are derived from Stachys lineages with flowers exhibiting insect- vs. bird-pollination characteristics. Thus, the Hawaiian endemic mints may provide yet another insular system for the combined study of polyploidy, hybrid cladogenesis, and adaptive radiation.     

Molecular phylogenetic relationships reveal taxonomic and biogeographic clades in Dianella (flax lilies; Asphodelaceae,Hemerocallidoideae)

A phylogeny of Dianella is presented based on Bayesian and maximum parsimony analyses of a combined molecular data set using three chloroplast markers (trnQ^UUG–5'rps16, 3'rps16–5'trnK^(UUU) and rpl14–rps8–infA–rpl36) and two nuclear markers (ITS and ETS). Accessions included most Dianella species, including all species from Australia, the centre of diversity for the genus, and related outgroup genera Eccremis, Stypandra, Thelionema and Herpolirion. The phylogeny showed Stypandra sister to Herpolirion + Thelionema, and confirmed the monophyly of Dianella. Within Dianella, a number of clades were resolved that revealed biogeographic relationships. Accessions from south-western Australia (extending into South Australia) formed the earliest diverging clade, followed by D. serrulata from New Guinea, sister to all other clades of Dianella from Australia and other regions. Tropical North Queensland species, including the D. pavopennacea complex, were related to a clade of accessions from New Caledonia and the Hawaiian Islands in the Pacific, and a clade that included samples of D. carolinensis (Caroline Islands) and the widespread D. ensifolia from South-East Asia and across the Indian Ocean to Mauritius and Madagascar. However, D. ensifolia is not monophyletic, with accessions from Japan and Taiwan related to a clade of Queensland samples that are part of the D. revoluta complex. Three New Zealand species (diploid, 2n?=?16) were found to be related to Norfolk Island D. intermedia (type locality; octoploid, 2n?=?64). In contrast ‘D. intermedia’ from Lord Howe Island was resolved as sister to the eastern Australian D. caerulea complex. The phylogenetic results indicate the need for taxonomic revision, particularly revision of the species ‘complexes’ D. longifolia and D. caerulea in Australia, and recognition of more than one species within D. ensifolia and within D. sandwicensis on the Hawaiian Islands.     

Phylogeny and historical biogeography of the cave-adapted shrimp genus Typhlatya (Atyidae) in the Caribbean Sea and western Atlantic

Aim To infer phylogenetic relationships among five species of the cave‐adapted shrimp genus Typhlatya in order to test competing hypotheses of dispersal and colonization of the disjunct cave localities occupied by these five species. Location Typhlatya species are found in caves and anchialine ponds across the northern margin of the Caribbean Sea, along the Mediterranean and Adriatic coasts and on oceanic islands in the Atlantic and eastern Pacific oceans. This study focuses on five species, one from Bermuda, one from the Caicos Islands and three from the Yucatan Peninsula of Mexico. Methods Partial sequences (c. 1400 bp) from the mitochondrial cytochrome b, 16S rDNA and COI genes were obtained from representative samples of the five species. Phylogenetic inference was carried out with maximum parsimony and maximum likelihood analyses. Parsimony networks were constructed for the Bermudian species Typhlatya iliffei and one Yucatan species Typhlatya mitchelli, to determine the degree of connectivity among populations inhabiting different cave systems. Results All three land masses were recovered as monophyletic. The two insular marine species from Bermuda and the Caicos Islands formed a clade, while the three continental freshwater species from the Yucatan Peninsula formed another. Within both Bermuda and the Yucatan, shared haplotypes were found in different cave systems, suggesting recent or ongoing gene flow among populations in both locales. Main conclusions The two insular marine Typhlatya species originated from an ancestral marine population, possibly already cave‐adapted, that is suggested to have colonized the Caicos Islands and subsequently dispersed to Bermuda via the Gulf Stream. Divergence estimates suggest that colonization occurred before the formation of present‐day anchialine cave habitat, which did not form on either island until the late Pliocene to early Pleistocene. Divergence estimates also indicate that the Yucatan freshwater species split before the formation of freshwater cave habitat in the Yucatan. These species could have inhabited crevicular marine habitats before the late Pliocene/early Pleistocene in the Yucatan or elsewhere in the Caribbean, and subsequently migrated to freshwater caves once they formed.     

Phylogenetic biogeography of the leafy liverwort Herbertus (Jungermanniales, Herbertaceae) based on nuclear and chloroplast DNA sequence data: correlation between genetic variation and geographical distribution

Aim The cosmopolitan genus Herbertus is notorious for having a difficult taxonomy and for the fact that there is limited knowledge of species ranges and relationships. Topologies generated from variable molecular markers are used to discuss biogeographical patterns in Herbertus and to compare them with the geological history of continents and outcomes reported for other land plants. Location Africa, Asia, Azores, Europe, southern South America, northern South America, North America, New Zealand. Methods Phylogenetic analyses of nuclear ribosomal internal transcribed spacer and chloroplast (cp) trnL–trnF sequences of 66 accessions of Herbertus and the outgroup species Triandrophyllum subtrifidum and Mastigophora diclados were used to investigate biogeographical patterns in Herbertus. Areas of putative endemism were defined based on the distribution of species included in the analyses. Maximum parsimony analyses were undertaken to reconstruct ancestral areas and intraspecies migration routes. Results The analyses reveal species‐level cladograms with a correlation between genetic variation and the geographical distribution of the related accessions. The southern South American Herbertus runcinatus is sister to the remainder of the genus, which is split into two main clades. One contains the Neotropical–African Herbertus juniperoideus and the New Zealand/Tasmanian Herbertus oldfieldianus. An African accession of H. juniperoideus is nested within Neotropical accessions. The second main clade includes species that inhabit Asia, the Holarctic, Africa, and northern South America. Maximum parsimony analyses indicate that this clade arose in Asia. Herbertus sendtneri originated in Asia and subsequently colonized the Holarctic and northern South America. An Asian origin and colonization into Africa is indicated for H. dicranus. Main conclusions The current distribution of Herbertus cannot be explained by Gondwanan vicariance. A more feasible explanation of the range is a combination of short‐distance dispersal, rare long‐distance dispersal events (especially into regions that faced floral displacements as a result of climatic changes) extinction, recolonization, and diversification. The African Herbertus flora is a mixture of Asian and Neotropical elements. Southern South America harbours an isolated species. The molecular data indicate partial decoupling of molecular and morphological variation in Herbertus. Biogeographical patterns in Herbertus are not dissimilar to those of other groups of bryophytes, but elucidation of the geographical ranges requires a molecular approach. Some patterns could be the result of maintenance of Herbertus in the inner Tropics during glacial maxima, and dispersal into temperate regions in warm phases.     

Molecular phylogeography of the fruit bat genus Melonycteris in northern Melanesia

Aim To investigate patterns of genetic divergence between populations of the fruit bat genus Melonycteris Dobson 1877 in relation to the possible effects on dispersal of the geological history of water barriers within and between northern Melanesian archipelagos. Location The genus is found only in the Bismarck Archipelago and Solomon Islands of northern Melanesia. Methods Up to 935 aligned bases of cytochrome b and cytochrome c oxidase subunit I DNA sequences were determined for specimens of most species and subspecies of Melonycteris. Measures of genetic distance, analysis of molecular variation and phylogenetic investigations (using maximum parsimony, maximum likelihood and Bayesian approaches) were conducted to assess the evolutionary relationships amongst populations. Results The deepest divergences within Melonycteris separate the genus into two reciprocally monophyletic clades from first, the Bismarck Archipelago, and secondly, the Solomon Islands. Within the Solomon Islands, five major clades received strong support. Listed in a generally north‐western to south‐eastern direction these were: (1) specimens from Choiseul and Santa Isabel; (2) specimens from New Georgia and Kolombangara; (3) specimens from Malaita; (4) specimens from Guadalcanal; and (5) specimens from Makira. Outgroup rooting suggested that the clade from Makira was the most basal within the Solomon Islands, being shown as the sister group to all other Melonycteris from this archipelago. Main conclusions Patterns of genetic variation within Melonycteris were generally consistent, given current knowledge of northern Melanesian geological history, with the hypothesis that the dispersal of these fruit bats is strongly inhibited by water barriers. Within the Solomon Islands the main genetic clades were each restricted to a single island or to a group of islands that are thought to have belonged to larger landmasses (Greater Gatumbangara and Greater Bukida) formed by land bridges during the Pleistocene. The high genetic distance between specimens from the Bismarck Archipelago and from the Solomon Islands reflects the persistently large geographic distance between these archipelagos. The unexpected phylogenetic position of the Makira specimens suggests either that this island was the first colonized by Melonycteris in the Solomon Islands or that this population is the relict of a clade that was previously more widely distributed.     

Repeated range expansion and niche shift in a volcanic hotspot archipelago: Radiation of C4 Hawaiian Euphorbia subgenus Chamaesyce (Euphorbiaceae)

Woody perennial plants on islands have repeatedly evolved from herbaceous mainland ancestors. Although the majority of species in Euphorbia subgenus Chamaesyce section Anisophyllum (Euphorbiaceae) are small and herbaceous, a clade of 16 woody species diversified on the Hawaiian Islands. They are found in a broad range of habitats, including the only known C₄ plants adapted to wet forest understories. We investigate the history of island colonization and habitat shift in this group. We sampled 153 individuals in 15 of the 16 native species of Hawaiian Euphorbia on six major Hawaiian Islands, plus 11 New World close relatives, to elucidate the biogeographic movement of this lineage within the Hawaiian island chain. We used a concatenated chloroplast DNA data set of more than eight kilobases in aligned length and applied maximum likelihood and Bayesian inference for phylogenetic reconstruction. Age and phylogeographic patterns were co‐estimated using BEAST. In addition, we used nuclear ribosomal ITS and the low‐copy genes LEAFY and G3pdhC to investigate the reticulate relationships within this radiation. Hawaiian Euphorbia first arrived on Kaua`i or Ni`ihau ca. 5 million years ago and subsequently diverged into 16 named species with extensive reticulation. During this process Hawaiian Euphorbia dispersed from older to younger islands through open vegetation that is disturbance‐prone. Species that occur under closed vegetation evolved in situ from open vegetation of the same island and are only found on the two oldest islands of Kaua`i and O`ahu. The biogeographic history of Hawaiian Euphorbia supports a progression rule with within‐island shifts from open to closed vegetation.     

A review of Polynesian Carposina Herrich‐Schäffer (Lepidoptera: Carposinidae), with descriptions of four new species

Hawaiian Carposina represent over 17% of the known world fauna of Carposinidae. In contrast, only two species are known for all of French Polynesia in the South Pacific. Here we describe four new species: two from the Hawaiian Islands, C arposina urbanae sp. nov. and C . gagneorum sp. nov. , and two from the Society Islands, C . longignathosa sp. nov. and C . brevinotata sp. nov. We further recognize another new Hawaiian species too worn to describe. Additionally, we present the first phylogeny for Polynesian Carposina, including 19 taxa, using one mitochondrial and two nuclear gene regions. The Hawaiian Carposina sampled thus far form a monophyletic clade. Lastly, we provide a framework to better understand the diversification and phylogeography of this group, and provide a summary of currently known host plant associations. Diversification appears to have resulted from interplay between host switching and geographic isolation across the Hawaiian Archipelago.