Atlantic reef fish biogeography and evolution

Aim To understand why and when areas of endemism (provinces) of the tropical Atlantic Ocean were formed, how they relate to each other, and what processes have contributed to faunal enrichment. Location Atlantic Ocean. Methods The distributions of 2605 species of reef fishes were compiled for 25 areas of the Atlantic and southern Africa. Maximum‐parsimony and distance analyses were employed to investigate biogeographical relationships among those areas. A collection of 26 phylogenies of various Atlantic reef fish taxa was used to assess patterns of origin and diversification relative to evolutionary scenarios based on spatio‐temporal sequences of species splitting produced by geological and palaeoceanographic events. We present data on faunal (species and genera) richness, endemism patterns, diversity buildup (i.e. speciation processes), and evaluate the operation of the main biogeographical barriers and/or filters. Results Phylogenetic (proportion of sister species) and distributional (number of shared species) patterns are generally concordant with recognized biogeographical provinces in the Atlantic. The highly uneven distribution of species in certain genera appears to be related to their origin, with highest species richness in areas with the greatest phylogenetic depth. Diversity buildup in Atlantic reef fishes involved (1) diversification within each province, (2) isolation as a result of biogeographical barriers, and (3) stochastic accretion by means of dispersal between provinces. The timing of divergence events is not concordant among taxonomic groups. The three soft (non‐terrestrial) inter‐regional barriers (mid‐Atlantic, Amazon, and Benguela) clearly act as ‘filters’ by restricting dispersal but at the same time allowing occasional crossings that apparently lead to the establishment of new populations and species. Fluctuations in the effectiveness of the filters, combined with ecological differences among provinces, apparently provide a mechanism for much of the recent diversification of reef fishes in the Atlantic. Main conclusions Our data set indicates that both historical events (e.g. Tethys closure) and relatively recent dispersal (with or without further speciation) have had a strong influence on Atlantic tropical marine biodiversity and have contributed to the biogeographical patterns we observe today; however, examples of the latter process outnumber those of the former.     

Marine longitudinal biodiversity: causes and conservation

The horizontal temperature zones of the earth tend to restrict the latitudinal ranges of species but allow the possibility of exceedingly broad longitudinal dispersals. In the Tropical Zone, biodiversity on the continental shelves is not homogeneous but is concentrated in two conspicuous peaks, one in the Indo‐Pacific Ocean and the other in the Atlantic. The Indo‐Pacific biodiversity peak is located within a relatively small area called the East Indies Triangle. The Atlantic peak is located in the southern Caribbean Sea. Evidence that has been accumulated over the years indicates that each area functions as a centre of origin and evolutionary radiation. What are the causes of these concentrations and their present functions? A newly published model indicates a positive relationship between environmental temperature and the rate of speciation. While this helps to explain the generally high tropical diversity, and the negative relationship between diversity and latitude, it does not provide a reason for the longitudinal concentrations. But, other new research serves to substantiate previous indications of a positive relationship between speciation rate and species diversity. The existence of this positive feedback, together with some contributory factors, provides the reason why concentrations occur. The evolutionary radiation probably begins when the build‐up of species diversity reaches a critical level. The warm‐temperate biotas are derived from the tropics. Their northern longitudinal relationships tend to be minor but, in the southern hemisphere, the West Wind Drift is an important dispersal mechanism for both warm‐temperate and cold‐temperate species. The cold‐temperate biotas peaked in two areas, the North Pacific and the Antarctic; each has developed into a centre of origin. The continuous dispersal of well‐adapted species from the centres helps peripheral communities maintain diversity.     

A realignment of marine biogeographic provinces with particular reference to fish distributions

Marine provinces, founded on contrasting floras or faunas, have been recognized for more than 150 years but were not consistently defined by endemism until 1974. At that time, provinces were based on at least a 10% endemism and nested within biogeographic regions that covered large geographic areas with contrasting biotic characteristics. Over time, some minor adjustments were made but the overall arrangement remained essentially unaltered. In many provinces, data on endemism were still not available, or were available only for the most widely studied vertebrates (fishes), a problem that is ongoing. In this report we propose a realignment for three reasons. First, recent works have provided new information to modify or redefine the various divisions and to describe new ones, including the Mid‐Atlantic Ridge, Southern Ocean, Tropical East Pacific and Northeast Pacific. Second, phylogeographic studies have demonstrated genetic subdivisions within and between species that generally corroborated provinces based on taxonomic partitions, with a notable exception at the Indian–Pacific oceanic boundary. Third, the original separation of the warm‐temperate provinces from the adjoining tropical ones has distracted from their close phylogenetic relationships. Here we propose uniting warm‐temperate and tropical regions into a single warm region within each ocean basin, while still recognizing provinces within the warm‐temperate and tropical zones. These biogeographic subdivisions are based primarily on fish distribution but utilize other marine groups for comparison. They are intended to demonstrate the evolutionary relationships of the living marine biota, and to serve as a framework for the establishment of smaller ecological units in a conservation context.     

What biogeography is: a place for process

The search for understanding of the past and present processes that have and/or continue to generate observed biotic distribution patterns substantially involves historical reconstruction based on present patterns (both phylogenetic and geographical). How this should be undertaken has been a cause for major debate over many decades. Residual patterns do not always provide explicit pointers to the causal processes, and in addition to applying our understanding of earth history, we need also to carefully explore the implications of contemporary processes as a means for unravelling pattern. Some biogeographers assert that earth and life evolve together, but knowledge of distributions and ecologies indicate that this is sometimes true and sometimes false. Just as general patterns may sometimes indicate a commonality of the means that generated the patterns so, too, do observable processes sometimes indicate commonalities. Vicariance and dispersal are fundamental attributes of biotic distributions. Phylogeography has the potential to assist us in determining which of these mechanisms has generated observable patterns.     

Minute moss beetles in the Southern Hemisphere: Molecular phylogeny,historical biogeography and habitat shifts (Coleoptera: Hydraenidae)

Minute moss beetles (Hydraenidae) are one of the most speciose and widespread families of aquatic Coleoptera, with an estimated 4000 extant species, found in the majority of aquatic habitats from coastal rock pools to mountain streams and from the Arctic Circle to the Antarctic islands. Molecular phylogenetic works have improved our understanding of the evolutionary history of the megadiverse Hydraena, Limnebius and Ochthebius in recent years, but most genera in the family have not yet been included in any phylogenetic analyses, particularly most of those which are restricted to the Southern Hemisphere. Using a multimarker molecular matrix, sampling over 40% of described species richness and 75% of currently recognized genera, we infer a comprehensive molecular phylogeny of these predominantly Gondwanan Hydraenidae. Whilst the genera we focus on are morphologically diverse, and currently classified across all four hydraenid subfamilies, our phylogenetic analyses suggest that these Gondwanan genera may instead constitute a single clade. As a result of our findings, the African genus Oomtelecopon Perkins syn.n. is shown to nest within Coelometopon Janssens, the New Zealand Homalaena Ordish syn.n. and Podaena Ordish syn.n. are synonymised with Orchymontia Broun, and the South African Pterosthetops Perkins syn.n. is synonymised with Prosthetops Waterhouse, resulting in Pterosthetopini Perkins syn.n. being synonymised with Prosthetopini Perkins. Mesoceratops Bilton & Jäch gen.n. is erected to accommodate six former members of Mesoceration Janssens, which is shown to be polyphyletic. We propose the replacement name Orchymontia ordishi Jäch & Bilton nom.n. for Homalaena dilatata Ordish, 1984 (now a junior homonym); altogether 39 new combinations are proposed. Our Bayesian divergence times infer an origin for this ‘Gondwana group’ of genera in Africa plus Madagascar in the mid-Cretaceous and suggest that both vicariant and dispersal processes, together with extinctions, have shaped the biogeographic history of these beetles in the Southern Hemisphere during the Cretaceous, resulting in geographically conserved extant lineages. Finally, we reconstruct ancestral habitat shifts across our phylogeny, revealing numerous changes in habitat occupancy in these genera, including multiple origins of fully terrestrial, humicolous taxa in different regions.     

Origin and post-colonization evolution of the Chatham Islands skink (Oligosoma nigriplantare nigriplantare)

Island ecosystems provide an opportunity to examine a range of evolutionary and ecological processes. The Chatham Islands are an isolated archipelago situated approximately 800 km east of New Zealand. Geological evidence indicates that the Chatham Islands re-emerged within the last 1-4 million years, following a prolonged period of marine inundation, and therefore the resident flora and fauna is the result of long-distance overwater dispersal. We examine the origin and post-colonization evolution of the Chatham Islands skink, Oligosoma nigriplantare nigriplantare, the sole reptile species occurring on the archipelago. We sampled O. n. nigriplantare from across nine islands within the Chatham Islands group, and representative samples from across the range of its closest relative, the New Zealand mainland common skink (Oligosoma nigriplantare polychroma). Our mitochondrial sequence data indicate that O. n. nigriplantare diverged from O. n. polychroma 5.86-7.29 million years ago. This pre-dates the emergence date for the Chatham Islands, but indicates that O. n. nigriplantare colonized the Chatham Islands via overwater dispersal on a single occasion. Despite the substantial morphological variability evident in O. n. nigriplantare, only relatively shallow genetic divergences (maximum divergence approximately 2%) were found across the Chatham Islands. Our analyses (haplotypic diversity, Phi(ST), analysis of molecular variance, and nested clade phylogeographical analysis) indicated restricted gene flow in O. n. nigriplantare resulting in strong differentiation between islands. However, the restrictions to gene flow might have only arisen recently as there was also a significant pattern of isolation by distance, possibly from when the Chatham Islands were a single landmass during Pleistocene glacial maxima when sea levels were lower. The level of genetic and morphological divergence between O. n. nigriplantare and O. n. polychroma might warrant their recognition as distinct species.     

The other face of Lyell: historical biogeography in his Principles of geology

Although some excellent articles about Lyell's work have been published, they do not explicitly deal with Lyell's biogeographical conceptions. The purpose of this paper is to analyse Lyell's biogeographical model in terms of its own internal structure. Lyell tried to explain the distribution of organisms by appealing to a real cause (climate). However, he was aware that environmental conditions were clearly insufficient to explain the existence of biogeographical regions. Lyell's adherence to ecological determinism generated strong tensions within his biogeographical model. He shifted from granting a secondary weight to dispersal to assigning it a major role. By doing so, Lyell was led into an evident contradiction. A permanent tension in Lyell's ideas was generated by the prevalent explanatory pattern of his time. The explanatory model based on laws did not produce satisfactory results in biology because it did not deal with historical processes. We may conclude that the knowledge of organic distribution interested Lyell as long as it could be explained by the uniformitarian principles of his geological system. The importance of the second volume of the Principles of geology lies in its ample and systematic argumentation about the geographical distribution of organisms. Lyell established, independently from any theory about organic change, the first version of dispersalist biogeography.     

Dispersal is fundamental to biogeography and the evolution of biodiversity on oceanic islands

Vicariance biogeography emerged several decades ago from the fusion of cladistics and plate tectonics, and quickly came to dominate historical biogeography. The field has since been largely constrained by the notion that only processes of vicariance and not dispersal offer testable patterns and refutable hypotheses, dispersal being a random process essentially adding only noise to a vicariant system. A consequence of this thinking seems to have been a focus on the biogeography of continents and continental islands, considering the biogeography of oceanic islands less worthy of scientific attention because, being dependent on stochastic dispersal, it was uninteresting. However, the importance of dispersal is increasingly being recognized, and here we stress its fundamental role in the generation of biodiversity on oceanic islands that have been created in situ , never connected to larger land masses. Historical dispersal patterns resulting in modern distributions, once considered unknowable, are now being revealed in many plant and animal taxa, in large part through the analysis of polymorphic molecular markers. We emphasize the profound evolutionary insights that oceanic island biodiversity has provided, and the fact that, although small in area, oceanic islands harbour disproportionately high biodiversity and numbers of endemic taxa. We further stress the importance of continuing research on mechanisms generating oceanic island biodiversity, especially detection of general, non-random patterns of dispersal, and hence the need to acknowledge oceanic dispersal as significant and worthy of research.     

Proximate sources of marine biodiversity

When temperature and other kinds of barrier divide formerly continuous populations and confine them to more restricted geographical areas, there is an evolutionary reaction that will, over time, result in the formation of endemic species. In such cases, an allopatric speciation process is considered to have taken place because reproductive isolation was caused by physical means instead of by natural selection. In contrast, when populations exist in a very high-diversity area and remain undivided by physical events, they exhibit a tendency to speciate by means of sympatry (or parapatry). This process, sometimes called competitive or ecological speciation, does involve reproductive isolation by means of natural selection. Populations that exist in geographical provinces bounded by physical barriers add to the overall diversity through the production of endemic species. This increase by species packing is relatively slow due to the very gradual tempo of the allopatric speciation process. Populations existing in centres of origin add to the general diversity through the production of species that are dominant in terms of their ability to spread over large parts of the world. It is proposed that such species are usually formed by sympatric speciation, a process that can be c. 20 times faster than species formation by allopatry. It is not suggested that sympatry is exclusive to centres of origin, nor that allopatry is confined to peripheral provinces. Both processes are widespread, but there do appear to be distinctive geographical concentrations. Considering that numbers of widespread species produced by centres of origin may eventually become subdivided by barriers, and thus give rise to descendants by allopatry, it is difficult to say how much of our present species diversity has come from one source or the other. Both speciation by sympatry from centres of origin and speciation by allopatry in peripheral provinces appear to be important sources of marine biodiversity.     

Diversity and biogeography of the Antarctic flora

Aim To establish how well the terrestrial flora of the Antarctic has been sampled, how well the flora is known, and to determine the major patterns in diversity and biogeography. Location Antarctica south of 60° S, together with the South Sandwich Islands, but excluding South Georgia, Bouvetøya and the periantarctic islands. Methods Plant occurrence data were collated from herbarium specimens and literature records, and assembled into the Antarctic Plant Database. Distributional patterns were analysed using a geographic information system. Biogeographical patterns were determined with a variety of multivariate statistics. Results Plants have been recorded from throughout the Antarctic, including all latitudes between 60° S and 86° S. Species richness declines with latitude along the Antarctic Peninsula, but there was no evidence for a similar cline in Victoria Land and the Transantarctic mountains. Multi‐dimensional scaling ordinations showed that the species compositions of the South Orkney, South Shetland Islands and the north‐western Antarctic Peninsula are very similar to each other, as are the floras of different regions in continental Antarctica. They also suggest, however, that the eastern Antarctic Peninsula flora is more similar to the flora of the southern Antarctic Peninsula than to the continental flora (with which it has traditionally been linked). The South Sandwich Islands have a flora that is very dissimilar to that in all Antarctic regions, probably because of their isolation and volcanic nature. Main conclusions The Antarctic flora has been reasonably well sampled, but certain areas require further floristic surveys. Available data do, however, allow for a number of robust conclusions. A diversity gradient exists along the Antarctic Peninsula, with fewer species (but not fewer higher taxa) at higher latitudes. Multi‐dimensional scaling ordination suggests three major floral provinces within Antarctica: northern maritime, southern maritime, and continental. Patterns of endemism suggest that a proportion of the lichen flora may have an ancient vicariant distribution, while most bryophytes are more recent colonists.     

Marine biogeography and ecology: invasions and introductions

Although biogeography and ecology had previously been considered distinct disciplines, this outlook began to change in the early 1990s. Several people expressed interest in creating a link that would help ecologists become more aware of external influences on communities and help biogeographers realize that distribution patterns had their genesis at the community level. They proposed an interdisciplinary approach called macroecology. This concept has been aided by the advent of phylogeography, for a better knowledge of genetic relationships has had great interdisciplinary value. Two areas of research that should obviously benefit from a macroecological approach are: (1) the question of local vs. regional diversity and (2) the question of whether invader species pose a threat to biodiversity. The two questions are related, because both deal with the vulnerability of ecosystems to penetration by invading species. Biogeographers, who have studied the broad oceanic patterns of dispersal and colonization, tend to regard isolated communities as being open to invasion from areas with greater biodiversity. It became evident that many wide‐ranging species were produced in centres of origin, and that the location of communities with respect to such centres had a direct effect on the level of species diversity. Ecologists, in earlier years, thought that a community could become saturated with species and would thereafter be self‐sustaining. But recent research has shown that saturation is probably never achieved and that the assembly of communities and their maintenance is more or less dependent on the invasion of species from elsewhere. The study of invasions that take place in coastal areas, usually the result of ship traffic and/or aquaculture imports, has special importance due to numerous opinions expressed by scientists and policy‐makers that such invasions are a major threat to biodiversity. However, none of the studies so far conducted has identified the extinction of a single, native marine species due to the influence of an exotic invader. Furthermore, fossil evidence of historical invasions does not indicate that invasive species have caused native extinctions or reductions in biodiversity.     

Endemism in Namibia: patterns, processes and predictions

Sandwiched between the Namib and Kalahari Deserts of southwestern Africa are the karooid and escarpment biotopes of Namibia which are rich in endemics of many taxa. Most plant, invertebrate, amphibian, reptile, mammal and bird species endemic to Namibia are found in a zone running through, and to the west of, Namibia's escarpment region. There is also an important region of endemism for succulent plants, reptiles and invertebrates in the Succulent Karoo biome. Congruence between endemism hotspots, particularly on rocky substrates, is remarkably high for most taxa, implying broadly similar speciation processes. Possible speciation mechanisms in different parts of the country include the spatial isolation of rupicolous taxa such as insects and reptiles by the formation of large coastal dune fields; the expansion and contraction of wooded savannas during pluvial and interpluvial periods; and global temperature shifts which created highland refugia for frost-susceptible plants and poikilotherms. Areas of endemism and species richness overlap poorly for Namibia's mainly arid-dwelling endemic vertebrates, as richness is highest in the mesic wetlands and woodlands of northeast Namibia. The overlap for succulent plants, insects and arachnids, however, is relatively high. Centres of endemism for plants and vertebrates fall mainly outside protected areas, as few parks were established with biodiversity indices in mind. Our analysis of endemism congruence provides a strong platform for the promulgation of new protected areas to safeguard Namibia's unique biota. Furthermore, analysis of speciation patterns and processes is a useful predictive tool for the identification of other biotically important sites.     

Phylogeny and biogeography in the evolution of migration: shorebirds of the Charadrius complex

Aim We explore whether molecular phylogeny and biogeography can complement evolutionary ecology in developing a method to address a long-standing issue in the evolution of migration: have migrations between breeding and non-breeding grounds, which may be on different continents, evolved through origins in the breeding grounds with successive shifts of the non-breeding distribution or vice versa? Methods To accommodate the biology of migration, we treated breeding and non-breeding distributions as characters to be mapped onto a phylogeny derived from mitochondrial DNA sequence data and so examined the ancestral home issue as a study in the direction of character evolution. Results Our main findings from applying this approach to a subset of the Charadrius complex of shorebirds (Aves: Charadriinae) are that a case can be made for shifts of breeding distributions having occurred in the ancestries of C. alexandrinus and C. veredus as those species evolved their present migration patterns. Our results also argue for a southern hemisphere origin (specifically South America) for the Charadrius complex as a whole. A South American origin implies other shifts in breeding distributions having occurred in the evolution of the species C. semipalmatus and C. vociferus. On applying the methods we developed for dealing with phylogenetic uncertainty, these results are reinforced and the merit of testing them further is suggested. Conclusions By way of a new approach to the evolution of migration, our study adds to a consensus emerging from the evolutionary ecology of migrant birds, arguing that shifts of breeding distributions are commonly, though not necessarily exclusively, involved in the evolution of migration.     

New perspectives on the origin and diversification of Africa's forest avifauna

The use of DNA sequence data in systematic studies has brought about a revolution in our understanding of avian relationships and when combined with digitized distributional data, has facilitated new interpretations about the origins of diverse clades of the African avifauna including its diversification up through the Tertiary until the present. Here we review recent studies with special reference to Africa's forest avifauna and specifically comment on the putative origins of 'hotspots' of endemism in the Eastern Arc Mountains of Tanzania and in the Cape Region of South Africa. Intriguingly, both these areas appear to have retained populations of relict taxa since the mid-tertiary thermal optimum and at the same time have been centres of recent species differentiation.     

Historical biogeography and ecology of a Continental Antarctic mite genus, Maudheimia (Acari, Oribatida): evidence for a Gondwanan origin and Pliocene-Pleistocene speciation

Of the eight genera (30 species) of extant Acari in Continental (= East) Antarctica, the genus Maudheimia Dalenius & Wilson, 1958 (Oribatida; Maudheimiidae) is uniquely endemic. A Gondwanan origin is proposed for the genus based on antiquity, inferred from endemism, a widespread distribution throughout Continental Antarctica and a limited dispersal capacity. Adaptation for a high montane epilithic existence, a necessity for the origination and long-term persistence in Antarctica, is inferred from the life history, physiology and ecology of Maudheimia. Phylogenetic analysis placed all four Maudheimia species in a single (generic) clade with the following structure (M. pelronia Wallwork, 1962 (M. tanngardenensis Coctzee, 1997 (M. manhalli Coetzee, 1997 (M. wilsoni Dalenius & Wilson, 1958)))). Geographical distributions of the Maudheimia species, in relation to their phylogenetic relationships, support the hypothesis that post-Gondwanan speciation occurred as a consequence of isolation during glaciation of Antarctica.     

Diversity, endemism and evolution in the Coral Triangle

In a recent paper by D. R. Bellwood and C. P. Meyer ('Searching for heat in a marine biodiversity hotspot', Journal of Biogeography , 2009, 36 , 569–576), the authors had two evident objectives: (1) to disprove the theory that the geographical origins of reef organisms could be determined by locating concentrations of endemic species, and (2) to emphasize that the high diversity of the Coral Triangle was due to an accumulation of species from outside that area. With regard to the first point, no such theory had previously been proposed to my knowledge. Second, the accumulation theory was promoted without consideration of the facts supporting the centre of origin hypothesis, except to dismiss it by saying that it had its origin in pre-continental drift ideas. This short response outlines the properties and evidence for the operation of centres of origin in this region.     

The West Indies as a laboratory of biogeography and evolution

Islands have long provided material and inspiration for the study of evolution and ecology. The West Indies are complex historically and geographically, providing a rich backdrop for the analysis of colonization, diversification and extinction of species. They are sufficiently isolated to sustain endemic forms and close enough to sources of colonists to develop a dynamic interaction with surrounding continental regions. The Greater Antilles comprise old fragments of continental crust, some very large; the Lesser Antilles are a more recent volcanic island arc, and the low-lying Bahama Islands are scattered on a shallow oceanic platform. Dating of island lineages using molecular methods indicates over-water dispersal of most inhabitants of the West Indies, although direct connections with what is now southern Mexico in the Early Tertiary, and subsequent land bridges or stepping stone islands linking to Central and South America might also have facilitated colonization. Species-area relationships within the West Indies suggest a strong role for endemic radiations and extinction in shaping patterns of diversity. Diversification is promoted by opportunities for allopatric divergence between islands, or within the large islands of the Greater Antilles, with a classic example provided by the Anolis lizards. The timing of colonization events using molecular clocks permits analysis of colonization-extinction dynamics by means of species accumulation curves. These indicate low rates of colonization and extinction for reptiles and amphibians in the Greater Antilles, with estimated average persistence times of lineages in the West Indies exceeding 30Myr. Even though individual island populations of birds might persist an average of 2Myr on larger islands in the Lesser Antilles, recolonization from within the archipelago appears to maintain avian lineages within the island chain indefinitely. Birds of the Lesser Antilles also provide evidence of a mass extinction event within the past million years, emphasizing the time-heterogeneity of historical processes. Geographical dynamics are matched by ecological changes in the distribution of species within islands over time resulting from adaptive radiation and shifts in habitat, often following repeatable patterns. Although extinction is relatively infrequent under natural conditions, changes in island environments as a result of human activities have exterminated many populations and others--especially old, endemic species--remain vulnerable. Conservation efforts are strengthened by recognition of aesthetic, cultural and scientific values of the unique flora and fauna of the West Indies.     

Congruent phylogeographic patterns in a young radiation of live‐bearing marine snakes: Pleistocene vicariance and the conservation implications of cryptic genetic diversity

Aim

To investigate phylogeographic patterns among and within co‐occurring sea snake species from Australia's endemic viviparous Aipysurus lineage, which includes critically endangered species, and evaluate the conservation implications of geographically structured patterns of genetic divergence and diversity.

Location

Australia's tropical shallow water marine environments spanning four regions: Great Barrier Reef (GBR), Gulf of Carpentaria (GoC), Timor Sea (TS) and coastal WA (WAC).

Methods

Samples from >550 snakes representing all nine nominal Aipysurus group species were obtained from throughout their known Australian ranges. Coalescent phylogenetic analyses and Bayesian molecular dating of mitochondrial DNA, combined with Bayesian and traditional population genetic analyses of 11 microsatellite loci, were used to evaluate genetic divergence and diversity.

Results

Mitochondrial DNA revealed highly congruent phylogeographic breaks among co‐occurring species, largely supported by nuclear microsatellites. For each species, each region was characterized by a unique suite of haplotypes (phylogroups). Divergences between the TS, GoC and/or GBR were invariably shallow and dated as occurring 50,000–130,000 years ago, coinciding with the cyclic Pleistocene emergence of the Torres Strait land bridge. By contrast, sea snakes from coastal WA were consistently highly divergent from other regions and dated as diverging 178,000–526,000 years ago, which was not associated with any known vicariant events.

Main Conclusions

Previously unappreciated highly divergent sea snake lineages in coastal WA potentially represent cryptic species, highlighting this region as a high‐priority area for conservation. The cyclic emergence of the Torres Strait land bridge is consisted with observed divergences between the TS, GoC and/or GBR; however, processes involved in the earlier divergences involving the WAC remain to be determined. The observed strong population genetic structures (as surrogates for dispersal) indicate that sea snakes have limited potential to reverse population declines via replenishment from other sources over time frames relevant to conservation.

     

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.