Evolutionary biogeography of the terrestrial biota of the Marquesas Islands,one of the world's remotest archipelagos

Aim

Here I review phylogenetic studies concerning the biogeography of the Marquesas Islands, an oceanic hotspot archipelago in the Pacific Ocean formed <5.5 Ma, and compare patterns (particularly pertaining to colonization and diversification) within the archipelago to those reported from the Hawaiian and Society Islands.

Location

Marquesas Islands, French Polynesia (Pacific Ocean).

Methods

I reviewed 37 phylogenetic studies incorporating Marquesas‐endemic taxa. I asked the following questions: (a) where are the sister‐groups of Marquesas lineages distributed? (b) are Marquesas‐endemic “radiations” monophyletic or polyphyletic? (c) what major between‐island phylogeographic barriers are seen in the Marquesas? (d) what evidence exists for diversification within islands? (e) how old is the Marquesas biota compared to the archipelago's age? Finally, these patterns are compared with those seen in the Society Islands and Hawaii.

Results

Most Marquesan lineages have their closest known relatives on other Pacific plate archipelagos (particularly the Society, Hawaiian, and Austral islands). Most Marquesas‐endemic radiations are found to be monophyletic, and among‐island diversification appears to be common. There is limited evidence for within‐island diversification. Some radiations may be consistent with a weak progression rule in which younger lineages are on younger islands. Crown ages of no Marquesas radiations appear to be older than the age of the archipelago (with one exception).

Main conclusions

Diversification of the Marquesas biota resembles that of the Hawaiian Islands more than that of the Society Islands. Many radiations are monophyletic and some appear to diversify in parallel with the formation of the archipelago.

     

Microevolutionary patterns and processes of the Native Hawaiian colonizing fern Odontosoria chinensis (Lindsaeaceae)

Abstract.— The vascular‐plant flora of the Hawaiian Islands is characterized by one of the highest rates of species endemism in the world. Among flowering plants, approximately 89% of species are endemic, and among pteridophytes, about 76% are endemic. At the single‐island level, however, rates of species endemism vary dramatically between these two groups with 80% of angiosperms and only 6% of pteridophytes being single‐island endemics. Thus, in many groups of Hawaiian angiosperms, it is possible to link studies of phylogeny, evolution, and biogeographic history at the interspecific and interisland levels. In contrast, the low level of single‐island species endemism among Hawaiian pteridophytes makes similar interspecific and interisland studies nearly impossible. Higher levels of interisland gene flow may account for the different levels of single‐island endemism in Hawaiian pteridophytes relative to angiosperms. The primary question we addressed in the present study was: Can we infer microevolutionary patterns and processes among populations within widespread species of Hawaiian pteridophytes wherein gene flow is probably common? To address this broad question, we conducted a population genetic study of the native Hawaiian colonizing species Odontosoria chinensis. Data from allozyme analyses allowed us to infer: (1) significant genetic differentiation among populations from different islands; (2) historical patterns of dispersal between particular pairs of islands; (3) archipelago‐level patterns of dispersal and colonization; (4) founder effects among populations on the youngest island of Hawaii; and, (5) that this species primarily reproduces via outcrossing, but may possess a mixed‐mating system.     

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.     

The Hawaiian Archipelago is a stepping stone for dispersal in the Pacific: an example from the plant genus Melicope (Rutaceae)

Aim Pacific biogeographical patterns in the widespread plant genus Melicope J.R. Forst. & G. Forst. (Rutaceae) were examined by generating phylogenetic hypotheses based on chloroplast and nuclear ribosomal sequence data. The aims of the study were to identify the number of colonization events of Melicope to the Hawaiian Islands and to reveal the relationship of Hawaiian Melicope to the Hawaiian endemic genus Platydesma H. Mann. The ultimate goal was to determine if the Hawaiian Islands served as a source area for the colonization of Polynesia. Location Nineteen accessions were sampled in this study, namely eight Melicope species from the Hawaiian Islands, four from the Marquesas Islands, one species each from Tahiti, Australia and Lord Howe Island, two Australian outgroups and two species of the Hawaiian endemic genus Platydesma. To place our results in a broader context, 19 sequences obtained from GenBank were included in an additional analysis, including samples from Australia, Papua New Guinea, New Zealand, Southeast Polynesia and Asia. Methods DNA sequences were generated across 19 accessions for one nuclear ribosomal and three chloroplast gene regions. Maximum parsimony analyses were conducted on separate and combined data sets, and a maximum likelihood analysis was conducted on the combined nuclear ribosomal and chloroplast data set. A broader nuclear ribosomal maximum parsimony analysis using sequences obtained from GenBank was also performed. Geographic areas were mapped onto the combined chloroplast and nuclear ribosomal tree, as well as onto the broader tree, using the parsimony criterion to determine the dispersal patterns. Results Phylogenetic analyses revealed that Platydesma is nested within Melicope and is sister to the Hawaiian members of Melicope. The Hawaiian Melicope + Platydesma lineage was a result of a single colonization event, probably from the Austral region. Finally, Marquesan Melicope descended from at least one, and possibly two, colonization events from the Hawaiian Islands. Main conclusions These data demonstrate a shifting paradigm of Pacific oceanic island biogeography, in which the patterns of long‐distance dispersal and colonization in the Pacific are more dynamic than previously thought, and suggest that the Hawaiian Islands may act as a stepping stone for dispersal throughout the Pacific.     

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.     

Biogeography and the evolution of flightlessness in a radiation of Hawaiian moths (Xyloryctidae: Thyrocopa)

Aim Although the ability to fly confers benefits to most insects, some taxa have become secondarily flightless. Insect flightlessness may be more likely to evolve in environments such as islands and other windswept and alpine areas, but this prediction has rarely been tested while controlling for phylogenetic effects. Here we present a phylogeny for the endemic Hawaiian Lepidoptera genus Thyrocopa, which has two flightless species that occur in alpine areas on Maui and Hawaii islands, in order to determine whether the flightless species are sister to each other or represent separate losses of flight. We also explore divergence times and biogeographic patterns of inter‐island colonization in Thyrocopa, and present the first Hawaiian study to sample a genus from nine islands. Location The Hawaiian Islands. Methods The phylogeny is composed of 70 individuals (including 23 Thyrocopa species and 7 outgroup species) sequenced for portions of cytochrome c oxidase subunit I, elongation factor 1α and wingless genes, for a total of 1964 base pairs, and was estimated using both parsimony (paup *) and Bayesian inference (Mr Bayes ). Divergence times were estimated using the beast software package. Results Our results indicate that two independent invasions of alpine habitats with concomitant loss of flight have occurred in Thyrocopa. Based on current taxon sampling, Thyrocopa colonized the Hawaiian Islands slightly before the formation of Kauai. In terms of overall patterns of diversification, subclades generally follow a progression from older to younger islands. The genus has the greatest number of species on Kauai, with species numbers generally decreasing with decreasing island age. Main conclusions Loss of flight ability has evolved twice in a short period of geological time in Thyrocopa, perhaps as a result of low temperatures, high winds and/or a lack of predation pressure. However, several other Thyrocopa species that live on small islands with consistently high winds, such as Necker and Nihoa islands, retain the ability to fly.     

Area and the rapid radiation of Hawaiian Bidens (Asteraceae)

Aim To estimate the rate of adaptive radiation of endemic Hawaiian Bidens and to compare their diversification rates with those of other plants in Hawaii and elsewhere with rapid rates of radiation. Location Hawaii. Methods Fifty‐nine samples representing all 19 Hawaiian species, six Hawaiian subspecies, two Hawaiian hybrids and an additional two Central American and two African Bidens species had their DNA extracted, amplified by polymerase chain reaction and sequenced for four chloroplast and two nuclear loci, resulting in a total of approximately 5400 base pairs per individual. Internal transcribed spacer sequences for additional outgroup taxa, including 13 non‐Hawaiian Bidens, were obtained from GenBank. Phylogenetic relationships were assessed by maximum likelihood and Bayesian inference. The age of the most recent common ancestor and diversification rates of Hawaiian Bidens were estimated using the methods of previously published studies to allow for direct comparison with other studies. Calculations were made on a per‐unit‐area basis. Results We estimate the age of the Hawaiian clade to be 1.3–3.1 million years old, with an estimated diversification rate of 0.3–2.3 species/million years and 4.8 × 10^?5 to 1.3 × 10^?4 species Myr^?1 km^?2. Bidens species are found in Europe, Africa, Asia and North and South America, but the Hawaiian species have greater diversity of growth form, floral morphology, dispersal mode and habitat type than observed in the rest of the genus world‐wide. Despite this diversity, we found little genetic differentiation among the Hawaiian species. This is similar to the results from other molecular studies on Hawaiian plant taxa, including others with great morphological variability (e.g. silverswords, lobeliads and mints). Main conclusions On a per‐unit‐area basis, Hawaiian Bidens have among the highest rates of speciation for plant radiations documented to date. The rapid diversification within such a small area was probably facilitated by the habitat diversity of the Hawaiian Islands and the adaptive loss of dispersal potential. Our findings point to the need to consider the spatial context of diversification – specifically, the relative scale of habitable area, environmental heterogeneity and dispersal ability – to understand the rate and extent of adaptive radiation.     

The Hawaiian amphibious caterpillar guild: new species of Hyposmocoma (Lepidoptera: Cosmopterigidae) confirm distinct aquatic invasions and complex speciation patterns

Although aquatic caterpillars are a globally rare lifestyle, we have found them in multiple, independent lineages of the endemic moth genus Hyposmocoma across all of the Hawaiian Islands with flowing water. We formally describe 11 new species of Hyposmocoma that belong to four different larval case types: cone, bugle, medium burrito, and large burrito: Hyposmocoma kahamanoa sp. nov. from Oahu Island, Hyposmocoma kamakou sp. nov. from Molokai Island, Hyposmocoma kahaiao sp. nov. , Hyposmocoma waihohonu sp. nov. , and Hyposmocoma moopalikea sp. nov. from Maui Island, and Hyposmocoma aumakuawai sp. nov. , Hyposmocoma eepawai sp. nov. , Hyposmocoma ipowainui sp. nov. , Hyposmocoma kawaikoi sp. nov. , Hyposmocoma uhauiole sp. nov. , and Hyposmocoma wailua sp. nov. from Kauai Island. We also illustrate and describe in detail the aquatic case‐bearing larva of Hyposmocoma kahamanoa. Despite having similar ecologies as algae and lichen grazers at and below the water line of streams, prior research indicates that species with each case type constitute an independent lineage, with terrestrial sister taxa, and therefore the different groups of species bearing unique case types each represent an independent aquatic invasion. The case‐bearing larvae often occur sympatrically, and on Kauai even species with similar case‐types occur together, suggesting complex patterns of speciation and either past periods of isolation or sympatric speciation. Phylogenetic analysis of 2243 base pairs from two nuclear and one mitochondrial gene for 18 species confirm that each species is endemic to a single volcano, and that morphological divergence within case‐types has not been dramatic. Diversification has been complex, and superficially similar case type lineages are not all monophyletic. Kauai, the oldest but smallest of the major high islands, supports more species in the aquatic guild than any other island, thus island age, rather than size, may be important in generating diversity in this group. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 15–42.     

Speciation and phylogeography of Hawaiian terrestrial arthropods

The Hawaiian archipelago is arguably the world's finest natural laboratory for the study of evolution and patterns of speciation. Arthropods comprise over 75% of the endemic biota of the Hawaiian Islands and a large proportion belongs to species radiations. We classify patterns of speciation within Hawaiian arthropod lineages into three categories: (i) single representatives of a lineage throughout the islands; (ii) species radiations with either (a) single endemic species on different volcanoes or islands, or (b) multiple species on each volcano or island; and (iii) single widespread species within a radiation of species that exhibits local endemism. A common pattern of phylogeography is that of repeated colonization of new island groups, such that lineages progress down the island chain, with the most ancestral groups (populations or species) on the oldest islands. While great dispersal ability and its subsequent loss are features of many of these taxa, there are a number of mechanisms that underlie diversification. These mechanisms may be genetic, including repeated founder events, hybridization, and sexual selection, or ecological, including shifts in habitat and/or host affiliation. The majority of studies reviewed suggest that natural selection is a primary force of change during the initial diversification of taxa.     

Dispersal and diversification: macroevolutionary implications of the MacArthur–Wilson model, illustrated by Simulium (Inseliellum) Rubstov (Diptera: Simuliidae)

Aim Provide an empirical test of the ‘radiation zone’ hypothesis of the MacArthur–Wilson theory of island biogeography using the taxon‐pulse hypothesis of Erwin and Brooks Parsimony Analysis (BPA) on Simulium (Inseliellum) Rubstov. Location Micronesia, Cook Islands, Austral Islands, Society Islands, Marquesas Islands, Fiji and New Caledonia. Methods Primary and secondary BPA of the phylogeny of Inseliellum. Results Primary BPA showed that 15% of the taxon area cladogram contained area reticulations. Secondary BPA (invoking the area duplication convention) generated a clear sequence of dispersal for Inseliellum. The sequence follows a Micronesia – Cook Islands – Marquesas Islands – Society Islands dispersal, with a separate dispersal from the Cook Islands to the Austral Islands less than 1 Ma. A radiation in the island of Tahiti (Society Islands) produced numerous dispersals from Tahiti to other islands within the Society Islands system. Islands close to Tahiti (source island) have been colonized from Tahiti more often than islands far from Tahiti, but a higher proportion of those species colonizing distant islands have become distinct species. Main conclusions The dispersal sequence of Inseliellum exhibits both old to young island dispersal and young to old island dispersal. This is due to habitat availability on each island. Inseliellum is a model system in exemplifying the ‘radiation zone’ hypothesis of MacArthur and Wilson. As well, islands close to the source are colonized more often that those far from the source, but colonization of islands far away from the source results in a higher proportion of speciation events than for islands close to the source. The diversification of Inseliellum corresponds to a taxon‐pulse radiation, with a centre of diversification on Tahiti resulting from its large area and abundant freshwater habitats. This study illustrates the utility of BPA in identifying complex scenarios that can be used to test theories about the complementary roles of ecology and phylogeny in historical biogeography.     

Ecomorphological convergence in Eleutherodactylus frogs: a case of replicate radiations in the Caribbean

Replicate radiations, the repeated multiplication of species associated with ecological divergence, have attracted much attention and generated as much debate. Due to the few well‐studied cases, it remains unclear whether replicate radiations are an exceptional result of evolution or a relatively common example of the power of adaptation by natural selection. We examined the case of Eleutherodactylus frogs, which radiated in the Caribbean islands resulting in more than 160 species that occupy very diverse habitats. A time‐calibrated phylogeny revealed that these frogs independently diversified on all larger islands producing species that occupy a broad range of microhabitats in different islands. Using phylogenetic comparative methods, we found an association between morphological traits and particular microhabitats, and for most microhabitats detected significant morphological convergence. Our results indicate Caribbean Eleutherodactylus are a novel example of replicate radiations, and highlight the predictability of evolutionary processes, as similar ecological opportunities can lead to similar outcomes.     

SEQUENTIAL RADIATIONS AND PATTERNS OF SPECIATION IN THE HAWAIIAN CRICKET GENUS LAUPALA INFERRED FROM DNA SEQUENCES

The tremendous diversity of endemic Hawaiian crickets is thought to have originated primarily through intraisland radiations, in contrast to an interisland mode of diversification in the native Hawaiian Drosophila. The Hawaiian cricket genus Laupala (family Gryllidae) is one of several native genera of flightless crickets found in rain-forest habitat across the Hawaiian archipelago. I examined the phylogenetic relationships among mitochondrial DNA (mtDNA) sequences sampled from 17 species of Laupala, including the 12S ribosomal RNA (rRNA), transfer RNA (RNA)^val and 16S rRNA regions. The distribution of mtDNA variants suggests that species within Laupala are endemic to single islands. The phylogenetic estimate produced from both maximum likelihood and maximum parsimony supports the hypothesis that speciation in Laupala occurred mainly within islands. The inferred biogeographical history suggests that diversification in Laupala began on Kauai, the oldest rain-forested Hawaiian island. Subsequently, colonization to younger islands in the archipelago resulted in a radiation of considerable phylogenetic diversity. Phylogenetic patterns in mtDNA are not congruent with prior systematic or taxonomic hypotheses. Hypotheses that may explain the conflict between the phylogenetic patterns of mtDNA variation and the species taxonomy are discussed.     

PACT in practice: comparative historical biogeographic patterns and species–area relationships of the Greater Antillean and Hawaiian Island terrestrial biotas

Aim To compare the evolutionary and ecological patterns of two extensively studied island biotas with differing geological histories (the Hawaiian Islands and the Greater Antilles). We evaluated the results from PACT (phylogenetic analysis for comparing trees), an innovative approach that has been proposed to reveal general patterns of biotic expansion (between regions) and in situ (within a region) diversification, as well as species–area relationships (SAR) and the taxon pulse dynamic. Location The Hawaiian Islands and Greater Antilles. Methods We used the PACT algorithm to construct general area cladograms and identified biotic expansion and in situ nodes. We analysed the power‐law SAR and relative contribution of biotic expansion and in situ diversification events using power‐law and linear regression analyses. Results Both biotic expansion and in situ nodes were prevalent throughout the PACT general area cladograms (Greater Antilles, 55.9% biotic expansion, 44.1% in situ; Hawaiian Islands, 40.6% biotic expansion, 59.4% in situ). Of the biotic expansion events, both forward and backward events occurred in both regions (Greater Antilles, 85.1% forward, 14.9% backward; Hawaiian Islands, 65% forward, 35% backward). Additionally, there is a power‐law SAR for the Greater Antilles but not for the Hawaiian Islands. However, exclusion of Hawai'i (the youngest, largest Hawaiian Island) produced a power‐law SAR for the Hawaiian Islands. Main conclusions The prevalence of in situ events as well as forward and backward biotic expansion events reveals that both Hawaiian and Greater Antillean biotas have evolved through alternating episodes of biotic expansion and in situ diversification. These patterns are characteristic of the taxon pulse dynamic, for which few data have previously been recorded on islands. Additionally, our analysis revealed that historical influences on the power‐law SARs are pronounced in both assemblages: old, small islands are relatively species rich and young, large islands are relatively species poor. Thus, our PACT results are consistent with hypotheses of geological influence on the evolution of island biotas and also provide greater insight into the role of the taxon pulse dynamic in the formation of island equilibria.     

ACCELERATED EVOLUTION OF LAND SNAILS MANDARINA IN THE OCEANIC BONIN ISLANDS: EVIDENCE FROM MITOCHONDRIAL DNA SEQUENCES

An endemic land snail genus Mandarina of the oceanic Bonin (Ogasawara) Islands shows exceptionally rapid evolution not only of morphological and ecological traits, but of DNA sequence. A phylogenetic relationship based on mitochondrial DNA (mtDNA) sequences suggests that morphological differences equivalent to the differences between families were produced between Mandarina and its ancestor during the Pleistocene. The inferred phylogeny shows that species with similar morphologies and life habitats appeared repeatedly and independently in different lineages and islands at different times. Sequential adaptive radiations occurred in different islands of the Bonin Islands and species occupying arboreal, semiarboreal, and terrestrial habitat arose independently in each island. Because of a close relationship between shell morphology and life habitat, independent evolution of the same life habitat in different islands created species possesing the same shell morphology in different islands and lineages. This rapid evolution produced some incongruences between phylogenetic relationship and species taxonomy. Levels of sequence divergence of mtDNA among the species of Mandarina is extremely high. The maximum level of sequence divergence at 16S and 12S ribosomal RNA sequence within Mandarina are 18.7% and 17.7%, respectively, and this suggests that evolution of mtDNA of Mandarina is extremely rapid, more than 20 times faster than the standard rate in other animals. The present examination reveals that evolution of morphological and ecological traits occurs at extremely high rates in the time of adaptive radiation, especially in fragmented environments.     

Parallel habitat specialization within the wolf spider genus Hogna from the Galápagos

Within most island archipelagos, such as the Galápagos, similar ecological gradients are found on geographically isolated islands. Species radiations in response to these ecological gradients may follow different scenarios being (i) a single habitat specialization event followed by secondary colonization of each ecotype on the different islands or (ii) repeated and parallel habitat specialization on each island separately. This latter scenario has been considered less likely as gene flow might hamper such ecotypic differentiation. At least for the Galápagos, the extent to which this process is involved in species radiations remains yet poorly understood. Within the wolf spider genus Hogna, seven species are described that can be divided into three different ecotypes based on general morphology and habitat preference i.e. species that inhabit the pampa vegetation in the highlands, species that occur in coastal dry habitats and one generalist species. Comparison of the species phylogeny based on one mitochondrial (COI) and one nuclear (28S) gene fragment convincingly demonstrates that ‘pampa’ and ‘coastal dry’ species evolved in parallel on the islands Santa Cruz and San Cristóbal. Despite the observation that allozymes analysis indicated that each species forms a distinct genetic cluster, phylogenetic divergence within these species complexes was very low and paraphyletic and most likely due to hybridization rather than incomplete lineage sorting, as demonstrated for the Santa Cruz species complex. This suggests that within‐island speciation occurred under low levels of gene flow. Species phylogeny in general did not follow the progression of island emergence as a molecular clock analysis suggested that island endemic species may have diverged after as well as before the emergence of the islands. This represents the first clear example of parallel and within‐island speciation because of habitat specialization on the Galápagos and that such divergence most likely occurred under historic gene flow.     

Origins,ages and population histories: comparative phylogeography of endemic Hawaiian butterflyfishes (genus Chaetodon)

Aim To investigate phylogeographic relationships, genetic connectivity and potential colonization routes for Hawaii’s endemic butterflyfishes (family Chaetodontidae). Location The Hawaiian Archipelago (central Pacific Ocean). Methods Molecular genetic analysis of mitochondrial DNA from three species of endemic Hawaiian butterflyfishes (Chaetodon multicinctus, n = 280; Chaetodon miliaris, n = 408; Chaetodon fremblii, n = 358) sampled from across the Hawaiian Archipelago was used in a suite of population genetic analyses to examine population histories and calculate coalescence times. We review a recent phylogenetic hypothesis for the Chaetodontidae and optimize ancestral distributions to nodes as a means of inferring colonization pathways to Hawaii. Results We found no evidence for population subdivisions across their ranges for any of the three endemic Hawaiian butterflyfish species (Φ_ST ≈ 0; P > 0.05 in each case). Coalescence analyses revealed that C. multicinctus, C. miliaris and C. fremblii date to genetic bottlenecks of c. 12 ka (95% confidence interval of τ, 0–46,732 years ago), c. 74 ka (95% confidence interval of τ, 62,918–105,699 years ago) and c. 301 ka (95% confidence interval of τ, 88,981–478,495 years ago), respectively. We found that the West Pacific appears to be the ancestral source for two of three independent colonization events to Hawaii (leading to the speciation of C. miliaris and C. fremblii). The third colonization of Hawaii (leading to the divergence of C. multicinctus) is ambiguous, but may have involved island‐hopping from the South Pacific via the Line Islands. Main conclusions Our results are consistent with the growing body of data indicating that ecological specialists may be more susceptible to severe population bottlenecks during large‐scale climatic variation on evolutionary time‐scales. The isolation of the Hawaiian Archipelago presents challenges for colonization by even the most highly dispersive marine organisms, and routes of colonization by endemic butterflyfishes show a variable pattern, indicating that there may be several pathways, both spatially and temporally, for marine fauna to colonize Hawaii. Hawaiian endemic butterflyfishes appear to represent genetically homogeneous populations across the archipelago.     

Diversity despite dispersal: colonization history and phylogeography of Hawaiian crab spiders inferred from multilocus genetic data

The Hawaiian archipelago is often cited as the premier setting to study biological diversification, yet the evolution and phylogeography of much of its biota remain poorly understood. We investigated crab spiders (Thomisidae, Mecaphesa ) that demonstrate contradictory tendencies: (i) dramatic ecological diversity within the Hawaiian Islands, and (ii) accompanying widespread distribution of many species across the archipelago. We used mitochondrial and nuclear genetic data sampled across six islands to generate phylogenetic hypotheses for Mecaphesa species and populations, and included penalized likelihood molecular clock analyses to estimate arrival times on the different islands. We found that 17 of 18 Hawaiian Mecaphesa species were monophyletic and most closely related to thomisids from the Marquesas and Society Islands. Our results indicate that the Hawaiian species evolved from either one or two colonization events to the archipelago. Estimated divergence dates suggested that thomisids may have colonized the Hawaiian Islands as early as ~10 million years ago, but biogeographic analyses implied that the initial diversification of this group was restricted to the younger island of Oahu, followed by back-colonizations to older islands. Within the Hawaiian radiation, our data revealed several well-supported genetically distinct terminal clades corresponding to species previously delimited by morphological taxonomy. Many of these species are codistributed across multiple Hawaiian Islands and some exhibit genetic structure consistent with stepwise colonization of islands following their formation. These results indicate that dispersal has been sufficiently limited to allow extensive ecological diversification, yet frequent enough that interisland migration is more common than speciation.     

Patterns of ecomorphological convergence among mainland and island Anolis lizards

Differing selective pressures on islands versus the mainland may produce alternative evolutionary outcomes among closely related lineages. Conversely, lineages may be constrained to produce similar outcomes in different mainland and island environments, or mainland and island environments may not differ significantly. Among the best‐studied island radiations are Caribbean Anolis lizards. Distinct morphotypes, or ‘ecomorphs’, have been described, and the same ecomorphs have evolved independently on each Greater Antillean island. The mainland Anolis radiation has received much less attention. We use a large morphological data set and a novel phylogenetic hypothesis to show that mainland Anolis did not evolve the same morphotypes as island Anolis, despite some island species being more closely related to mainland species than to island species that share their morphotype. A maximum of four of the six Caribbean ecomorphs were found to exist on the mainland, and just 15 of 123 mainland species are assignable to a Caribbean ecomorph. This result was insensitive to differing taxon samples and alternative phylogenetic hypotheses. Mainland convergence to a Caribbean ecomorph occurs only among species assigned to the grass‐bush ecomorph. Thus, the ecomorphs that have evolved convergently multiple times in the Caribbean have not evolved in parallel on the mainland. These results are consistent with the hypothesis that mainland and island environments offer different selective pressures. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101 , 852–859.     

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.