Using taxonomic and phylogenetic evenness to compare diversification in two Island Floras

This study compares the phylogenetic structure in the Canary Islands and Hawaii by means of the distributions of the species number for plant families (Taxonomic evenness) and lineages (Phylogenetic evenness) across archipelagos and across habitats in both archipelagos using the Gini coefficient. We then investigate phylogenies to identify particular habitats contributing to such differences using Taxonomic distinctness (AvTD) and its variation (VarTD).Our results show that the distribution of species number among Hawaiian lineages is much more uneven than the Canary Islands. In contrast, Hawaii produces a more even distribution of species number by family than the Canary Islands. This may be due to the Hawaiian Flora being derived from considerably fewer colonists than the Canarian Flora as a result of its much greater degree of isolation. At the same time, Hawaii is represented by the same number of families as the Canary Islands. This may stem from Hawaii's flora being derived from a greater range of source areas despite its isolation. Finally, there is much more diversification spread across a larger number of lineages in Hawaii. The higher degree of Hawaiian diversification may be due to a greater range of habitats, more diverse and phylogenetically distinct floristic sources, and low initial species diversity resulting from extreme isolation.Two Canarian habitats (Rock communities and Thermophilous habitats) and one Hawaiian habitat (Wet communities) contribute to the differences in phylogenetic structure between the two archipelagos. These habitats exhibit disproportionate levels of unevenness and may represent centres of diversification. We propose a combination of two habitat properties, high receptivity and low stability, to explain these results.     

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.     

Out of Hawaii: the origin and biogeography of the genus Scaptomyza (Diptera: Drosophilidae)

The Hawaiian Archipelago is the most isolated island system on the planet and has been the subject of evolutionary research for over a century. The largest radiation of species in Hawaii is the Hawaiian Drosophilidae, a group of approximately 1000 species. Dispersal to isolated island systems like Hawaii is rare and the resultant flora and fauna shows high disharmony with mainland communities. The possibility that some lineages may have originated in Hawaii and subsequently 'escaped' to diversify on continental landmasses is expected to be rarer still. We present phylogenetic analysis of 134 partially sequenced mitochondrial genomes of Drosophilidae (approx. 1.3 Mb of sequence total) to address major aspects of adaptive radiation and dispersal in Hawaii. We show that the genus Scaptomyza, a group that accounts for approximately one-third of the species-level diversity of Drosophilidae in the Hawaiian Islands, originated in Hawaii, diversified there, and subsequently colonized a number of island and continental landmasses elsewhere on the globe. We propose that a combination of small body size, rapid generation time and unique ecological and physiological adaptations have allowed this genus to effectively disperse and diversify.     

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.     

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.     

Steady Plio-Pleistocene diversification and a 2-million-year sympatry threshold in a New Zealand cicada radiation

Estimation of diversification rates in evolutionary radiations requires a complete accounting of cryptic species diversity. The rapidly evolving songs of acoustically signaling insects make them good model organisms for such studies. This paper examines the timing of diversification of a large (30 taxon) group of New Zealand cicadas (genus Kikihia Dugdale). We use Bayesian relaxed-clock methods and phylogenetic trees based on nuclear and mitochondrial DNA data, and we apply alternative combinations of evolutionary rate priors and geological calibrations. The extant Kikihia taxa began to diversify near the Miocene/Pliocene boundary around the time of increased mountain-building, and both the mitochondrial and nuclear-gene trees confirm early splits of lineages currently represented by lowland forest-dwelling taxa. Most lineages originated in the Pleistocene, and sustained diversification occurred rapidly at over 0.5 lineages/my, a rate comparable to that of the Hawaiian silverswords. Diversification rate tests suggest an increase in the early to mid-Pliocene, followed by constant diversification from the Late Pliocene onward. No descendants of the many Pleistocene-age splits have evolved the ability to coexist in sympatry, and, where they do come into contact, hybrid zones have been documented based on acoustic and DNA evidence. In contrast, lineages separated in time by approximately 2Myr often overlap in distribution with no evidence of hybridization. This suggests that at least 2Myr has been required to achieve the level of divergence required for reproductive isolation.     

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.     

Origin and diversification of the endemic Hawaiian tree snails (Achatinellidae: Achatinellinae) based on molecular evidence

Tree snails of the endemic subfamily Achatinellinae comprise a diverse and important component of the Hawaiian fauna. In recent decades anthropogenic impacts have resulted in devastating extinction rates in Hawaiian tree snails. To address long-standing biogeographic, systematic, and evolutionary questions we used cytochrome c oxidase subunit I (COI) gene sequences to reconstruct the phylogeny of 23 extant species spanning the range of the subfamily from five Hawaiian Islands. To investigate family-level relationships, data were analyzed from 11 terrestrial pulmonate families. Although nodal support for monophyly of the endemic Pacific family Achatinellidae and endemic Hawaiian subfamily Achatinellinae was strong, bifurcation order among deeper ingroup nodes was not well-supported by bootstrap resampling. We hypothesize that lineage extinction and rapidity of lineage formation may have rendered evolutionary reconstruction difficult using a standard phylogenetic approach. Use of an optimized evolutionary model, however, improved resolution and recovered three main clades. The diversification pattern inferred contradicts the traditional biogeographic hypothesis of a Maui origin of the achatinelline lineage. Taxa comprising the basal ingroup clade (Achatinella spp.) and seeding lineages for subsequent clades originated on O'ahu. Therefore it appears that the ancestral colonizing species of achatinellines arrived first on O'ahu from an unknown source, and that O'ahu is the Hawaiian origin of the subfamily. Species previously defined by morphological criteria were generally found to be phylogenetically distinct, and the overall colonization pattern follows the island-age progression rule with several instances of generic polyphyly and back-colonization.     

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.     

Phylogenetics of the Antopocerus-Modified Tarsus Clade of Hawaiian Drosophila: Diversification across the Hawaiian Islands

The Hawaiian Drosophilidae radiation is an ecologically and morphologically diverse clade of almost 700 described species. A phylogenetic approach is key to understanding the evolutionary forces that have given rise to this diverse lineage. Here we infer the phylogeny for the antopocerus, modified tarsus and ciliated tarsus (AMC) clade, a lineage comprising 16% (91 of 687 species) of the described Hawaiian Drosophilidae. To improve on previous analyses we constructed the largest dataset to date for the AMC, including a matrix of 15 genes for 68 species. Results strongly support most of the morphologically defined species groups as monophyletic. We explore the correlation of increased diversity in biogeography, sexual selection and ecology on the present day diversity seen in this lineage using a combination of dating methods, rearing records, and distributional data. Molecular dating analyses indicate that AMC lineage started diversifying about 4.4 million years ago, culminating in the present day AMC diversity. We do not find evidence that ecological speciation or sexual selection played a part in generating this diversity, but given the limited number of described larval substrates and secondary sexual characters analyzed we can not rule these factors out entirely. An increased rate of diversification in the AMC is found to overlap with the emergence of multiple islands in the current chain of high islands, specifically Oahu and Kauai.     

DIVERSIFICATION OF THE AFRICAN GENUS PROTEA (PROTEACEAE) IN THE CAPE BIODIVERSITY HOTSPOT AND BEYOND: EQUAL RATES IN DIFFERENT BIOMES

The Cape region of South Africa is a hotspot of flowering plant biodiversity. However, the reasons why levels of diversity and endemism are so high remain obscure. Here, we reconstructed phylogenetic relationships among species in the genus Protea, which has its center of species richness and endemism in the Cape, but also extends through tropical Africa as far as Eritrea and Angola. Contrary to previous views, the Cape is identified as the ancestral area for the radiation of the extant lineages: most species in subtropical and tropical Africa are derived from a single invasion of that region. Moreover, diversification rates have been similar within and outside the Cape region. Migration out of the Cape has opened up vast areas, but those lineages have not diversified as extensively at fine spatial scales as lineages in the Cape. Therefore, higher net rates of diversification do not explain the high diversity and endemism of Protea in the Cape. Instead, understanding why the Cape is so diverse requires an explanation for how Cape species are able to diverge and persist at such small spatial scales.     

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.     

Interpreting colonization of the Calathus (Coleoptera: Carabidae) on the Canary Islands and Madeira through the application of the parametric bootstrap

The Canary Islands have proven to be an interesting archipelago for the phylogeographic study of colonization and diversification with a number of recent studies reporting evolutionary patterns and processes across a diversity of floral and faunal groups. The Canary Islands differ from the Hawaiian and Galapagos Islands by their close proximity to a continental land mass, being 110 km from the northwestern coast of Africa. This close proximity to a continent obviously increases the potential for colonization, and it can be expected that at the level of the genus some groups will be the result of more than one colonization. In this study we investigate the phylogeography of a group of carabid beetles from the genus Calathus on the Canary Islands and Madeira, located 450 km to the north of the Canaries and 650 km from the continent. The Calathus are well represented on these islands with a total of 29 species, and on the continent there are many more. Mitochondrial cytochrome oxidase I and II sequence data has been used to identify the phylogenetic relationships among the island species and a selection of continental species. Specific hypotheses of monophyly for the island fauna are tested with parametric bootstrap analysis. Data suggest that the Canary Islands have been colonized three times and Madeira twice. Four of these colonizations are of continental origin, but it is possible that one Madeiran clade may be monophyletic with a Canarian clade. The Calathus faunas of Tenerife and Madeira are recent in origin, similar to patterns previously reported for La Gomera, El Hierro, and Gran Canaria.     

Floristic biogeography of the Hawaiian Islands: influences of area, environment and paleogeography

Aim A detailed database of distributions and phylogenetic relationships of native Hawaiian flowering plant species is used to weigh the relative influences of environmental and historical factors on species numbers and endemism. Location The Hawaiian Islands are isolated in the North Pacific Ocean nearly 4000 km from the nearest continent and nearly as distant from the closest high islands, the Marquesas. The range of island sizes, environments, and geological histories within an extremely isolated archipelago make the Hawaiian Islands an ideal system in which to study spatial variation in species distributions and diversity. Because the biota is derived from colonization followed by extensive speciation, the role of evolution in shaping the regional species assemblage can be readily examined. Methods For whole islands and regions of each major habitat, species–area relationships were assessed. Residuals of species–area relationships were subjected to correlation analysis with measures of endemism, isolation, elevation and island age. Putative groups of descendents of each colonist from outside the Hawaiian Islands were considered phylogenetic lineages whose distributions were included in analyses. Results The species–area relationship is a prominent pattern among islands and among regions of each given habitat. Species number in each case correlates positively with number of endemics, number of lineages and number of species per lineage. For mesic and wet habitat regions, island age is more influential than area on species numbers, with older islands having more species, more single‐island endemics, and higher species : lineage ratios than their areas alone would predict. Main conclusions Because species numbers and endemism are closely tied to speciation in the Hawaiian flora, particularly in the most species‐rich phylogenetic lineages, individual islands’ histories are central in shaping their biota. The Maui Nui complex of islands (Maui, Moloka‘i, Lāna‘i and Kaho‘olawe), which formed a single large landmass during most of its history, is best viewed in terms of either the age or area of the complex as a whole, rather than the individual islands existing today.     

Origin of Hawaiian Polystichum (Dryopteridaceae) in the context of a world phylogeny

A genus-wide molecular phylogeny for Polystichum and allied genera (Dryopteridaceae) was reconstructed to address the biogeographic origin and evolution of the three Hawaiian Polystichum species, all endemic there. The analysis was based on the cpDNA sequences rbcL and the trnL-F spacer from a taxonomically and geographically diverse sample. Parsimony and Bayesian phylogenetic analyses of the combined data support a monophyletic Polystichum and corroborate recent hypotheses as to membership and sequence of origin of the major groups within the genus. The Hawaiian Polystichum species are polyphyletic; two separate lineages appear to have arrived independently from the Old World. The provenance of the diploid Polystichum hillebrandii is continental eastern Asia, while the source of the polyploid lineage comprising tetraploid P. haleakalense and octoploid P. bonseyi is likely continental Asia. From our results, the origin of the Hawaiian species of Polystichum, like many Hawaiian fern genera with several species, is the result of multiple migrations to the islands, rather than single migrations yielding nearly all the local diversity as in the angiosperms. This emerging pattern provides a modern test of the premise that propagule vagility has a central role in determining pattern of evolution.     

Recent and rapid speciation with limited morphological disparity in the genus Rattus

Recent and rapid radiations provide rich material to examine the factors that drive speciation. Most recent and rapid radiations that have been well-characterized involve species that exhibit overt ecomorphological differences associated with clear partitioning of ecological niches in sympatry. The most diverse genus of rodents, Rattus (66 species), evolved fairly recently, but without overt ecomorphological divergence among species. We used multilocus molecular phylogenetic data and five fossil calibrations to estimate the tempo of diversification in Rattus, and their radiation on Australia and New Guinea (Sahul, 24 species). Based on our analyses, the genus Rattus originated at a date centered on the Pliocene-Pleistocene boundary (1.84-3.17 Ma) with a subsequent colonization of Sahul in the middle Pleistocene (0.85-1.28 Ma). Given these dates, the per lineage diversification rates in Rattus and Sahulian Rattus are among the highest reported for vertebrates (1.1-1.9 and 1.6-3.0 species per lineage per million years, respectively). Despite their rapid diversification, Rattus display little ecomorphological divergence among species and do not fit clearly into current models of adaptive radiations. Lineage through time plots and ancestral state reconstruction of ecological characters suggest that diversification of Sahulian Rattus was most rapid early on as they expanded into novel ecological conditions. However, rapid lineage accumulation occurred even when morphological disparity within lineages was low suggesting that future studies consider other phenotypes in the diversification of Rattus.     

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.     

The assembly of montane biotas: linking Andean tectonics and climatic oscillations to independent regimes of diversification in Pionus parrots

The mechanisms underlying the taxonomic assembly of montane biotas are still poorly understood. Most hypotheses have assumed that the diversification of montane biotas is loosely coupled to Earth history and have emphasized instead the importance of multiple long-distance dispersal events and biotic interactions, particularly competition, for structuring the taxonomic composition and distribution of montane biotic elements. Here we use phylogenetic and biogeographic analyses of species in the parrot genus Pionus to demonstrate that standing diversity within montane lineages is directly attributable to events of Earth history. Phylogenetic relationships confirm three independent biogeographic disjunctions between montane lineages, on one hand, and lowland dry-forest/wet-forest lineages on the other. Temporal estimates of lineage diversification are consistent with the interpretation that the three lineages were transported passively to high elevations by mountain building, and that subsequent diversification within the Andes was driven primarily by Pleistocene climatic oscillations and their large-scale effects on habitat change. These results support a mechanistic link between diversification and Earth history and have general implications for explaining high altitudinal disjuncts and the origin of montane biotas.     

A Complex Evolutionary History in a Remote Archipelago: Phylogeography and Morphometrics of the Hawaiian Endemic Ligia Isopods

Compared to the striking diversification and levels of endemism observed in many terrestrial groups within the Hawaiian Archipelago, marine invertebrates exhibit remarkably lower rates of endemism and diversification. Supralittoral invertebrates restricted to specific coastal patchy habitats, however, have the potential for high levels of allopatric diversification. This is the case of Ligia isopods endemic to the Hawaiian Archipelago, which most likely arose from a rocky supralittoral ancestor that colonized the archipelago via rafting, and diversified into rocky supralittoral and inland lineages. A previous study on populations of this isopod from Oʻahu and Kauaʻi revealed high levels of allopatric differentiation, and suggested inter-island historical dispersal events have been rare. To gain a better understanding on the diversity and evolution of this group, we expanded prior phylogeographic work by incorporating populations from unsampled main Hawaiian Islands (Maui, Molokaʻi, Lanaʻi, and Hawaiʻi), increasing the number of gene markers (four mitochondrial and two nuclear genes), and conducting Maximum likelihood and Bayesian phylogenetic analyses. Our study revealed new lineages and expanded the distribution range of several lineages. The phylogeographic patterns of Ligia in the study area are complex, with Hawaiʻi, Oʻahu, and the Maui-Nui islands sharing major lineages, implying multiple inter-island historical dispersal events. In contrast, the oldest and most geographically distant of the major islands (Kauaʻi) shares no lineages with the other islands. Our results did not support the monophyly of all the supralittoral lineages (currently grouped into L. hawaiensis), or the monophyly of the terrestrial lineages (currently grouped into L. perkinsi), implying more than one evolutionary transition between coastal and inland forms. Geometric-morphometric analyses of three supralittoral clades revealed significant body shape differences among them. A taxonomic revision of Hawaiian Ligia is warranted. Our results are relevant for the protection of biodiversity found in an environment subject to high pressure from disturbances.