Rapid parallel evolution of aberrant traits in the diversification of the Gulf of Guinea white-eyes (Aves, Zosteropidae)

Archipelago-endemic bird radiations are familiar to evolutionary biologists as key illustrations of evolutionary patterns. However, such radiations are in fact rare events. White-eyes (Zosteropidae) are birds with an exceptionally high colonization and speciation potential; they have colonized more islands globally than any other passerine group and include the most species-rich bird genus. The multiplication of white-eye island endemics has been consistently attributed to independent colonizations from the mainland; the white-eyes of the Gulf of Guinea archipelago had been seen as a classic case, spanning as great a breadth of phenotypic diversity as the family worldwide. Contrary to this hypothesis, our molecular phylogenetic analysis places the Gulf of Guinea white-eyes in just two radiations, one grouping all five oceanic island taxa and the other grouping continental island and land-bridge taxa. Numerous 'aberrant' phenotypes (traditionally grouped in the genus Speirops) have evolved independently over a short space of time from nonaberrant (Zosterops) phenotypes; the most phenotypically divergent species have separated as recently as 0.22 Ma. These radiations rival those of Darwin's finches and the Hawaiian honeycreepers in terms of the extent of adaptive radiation per unit time, both in terms of species numbers and in terms of phenotypic diversity. Tempo and patterns of morphological divergence are strongly supportive of an adaptive radiation in the oceanic islands driven by ecological interactions between sympatric white-eyes. Here, very rapid phenotypic evolution mainly affected taxa derived from the youngest wave of colonization, in accordance with the model of asymmetric divergence owing to resource competition in sympatry.     

The role of immigration and in-situ radiation in explaining blood parasite assemblages in an island bird clade

Parasite communities on islands are assembled through multiple immigrations and/or in-situ diversification. In this study, we used a phylogenetic approach to investigate the role of such processes in shaping current patterns of diversity in Leucocytozoon, a group of haemosporidian blood parasites infecting whites eyes (Zosterops) endemic to the Mascarene archipelago (south-western Indian Ocean). We found that this parasite community arose through a combination of multiple immigrations and in-situ diversification, highlighting the importance of both processes in explaining island diversity. Specifically, two highly diverse parasite clades appear to have been present in the Mascarenes for most of their evolutionary history and have diversified within the archipelago, while another lineage apparently immigrated more recently, probably with human-introduced birds. Interestingly, the evolutionary histories of one clade of parasites and Indian Ocean Zosterops seem tightly associated with a significant signal for phylogenetic congruence, suggesting that host-parasite co-divergence may have occurred in this system.     

Tracking island colonization history and phenotypic shifts in Indian Ocean bulbuls (Hypsipetes: Pycnonotidae)

Molecular phylogenies of island organisms provide useful systems for testing hypotheses of convergent or parallel evolution, since selectively neutral molecular characters are likely to be independent of phenotype, and the existence of similar environments on multiple isolated islands provides numerous opportunities for populations to evolve independently under the same constraints. Here we construct a phylogenetic hypothesis for Hypsipetes bulbuls of the western Indian Ocean, and use this to test hypotheses of colonization pattern and phenotypic change among islands of the region. Mitochondrial sequence data were collected from all extant taxa of the region, combined with sequence data from relevant lineages in Asia. Data are consistent with a single Hypsipetes colonization of the western Indian Ocean from Asia within the last 2.6 Myr. The expansion of Hypsipetes appears to have occurred rapidly, with descendants found across the breadth of its western Indian Ocean range. The data suggest that a more recent expansion of Hypsipetes madagascariensis from Madagascar led to the colonization of Aldabra and a secondary colonization of the Comoros. Groupings of western Indian Ocean Hypsipetes according to phenotypic similarities do not correspond to mtDNA lineages, suggesting that these similarities have evolved by convergence or parallelism. The direction of phenotypic change cannot be inferred with confidence, since the primary expansion occurred rapidly relative to the rate of mtDNA substitution, and the colonization sequence remains uncertain. However, evidence from biogeography and comparison of independent colonization events are consistent with the persistence of a small grey continental bulbul in India and Madagascar, and multiple independent origins of large size and green plumage in insular island populations of the Comoros, Mascarenes and Seychelles. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 271–287.     

Biogeography of Old World emballonurine bats (Chiroptera: Emballonuridae) inferred with mitochondrial and nuclear DNA

Extant bats of the genus Emballonura have a trans-Indian Ocean distribution, with two endemic species restricted to Madagascar, and eight species occurring in mainland southeast Asia and islands in the western Pacific Ocean. Ancestral Emballonura may have been more widespread on continental areas, but no fossil identified to this genus is known from the Old World. Emballonura belongs to the subfamily Emballonurinae, which occurs in the New and Old World. Relationships of all Old World genera of this subfamily, including Emballonura and members of the genera Coleura from Africa and western Indian Ocean islands and Mosia nigrescens from the western Pacific region, are previously unresolved. Using 1833 bp of nuclear and mitochondrial genes, we reconstructed the phylogenetic history of Old World emballonurine bats. We estimated that these lineages diverged around 30 million years ago into two monophyletic sister groups, one represented by the two taxa of Malagasy Emballonura, Coleura and possibly Mosia, and the other by a radiation of Indo-Pacific Emballonura, hence, rendering the genus Emballonura paraphyletic. The fossil record combined with these phylogenetic relationships suggest at least one long-distance dispersal event across the Indian Ocean, presumably of African origin, giving rise to all Indo-Pacific Emballonura species (and possibly Mosia). Cladogenesis of the extant Malagasy taxa took place during the Quaternary giving rise to two vicariant species, E. atrata in the humid east and E. tiavato in the dry west.     

Molecular phylogeography reveals island colonization history and diversification of western Indian Ocean sunbirds (Nectarinia: Nectariniidae)

We constructed a phylogenetic hypothesis for western Indian Ocean sunbirds (Nectarinia) and used this to investigate the geographic pattern of their diversification among the islands of the Indian Ocean. A total of 1309 bp of mitochondrial sequence data was collected from the island sunbird taxa of the western Indian Ocean region, combined with sequence data from a selection of continental (African and Asian) sunbirds. Topological and branch length information combined with estimated divergence times are used to present hypotheses for the direction and sequence of colonization events in relation to the geological history of the Indian Ocean region. Indian Ocean sunbirds fall into two well-supported clades, consistent with two independent colonizations from Africa within the last 3.9 million years. The first clade contains island populations representing the species Nectarinia notata, while the second includes Nectarinia souimanga, Nectarinia humbloti, Nectarinia dussumieri, and Nectarinia coquereli. With respect to the latter clade, application of Bremer's [Syst. Biol. 41 (1992) 436] ancestral areas method permits us to posit the Comoros archipelago as the point of initial colonization in the Indian Ocean. The subsequent expansion of the souimanga clade across its Indian Ocean range occurred rapidly, with descendants of this early expansion remaining on the Comoros and granitic Seychelles. The data suggest that a more recent expansion from Anjouan in the Comoros group led to the colonization of Madagascar by sunbirds representing the souimanga clade. In concordance with the very young geological age of the Aldabra group, the sunbirds of this archipelago have diverged little from the Madagascar population; this is attributed to colonization of the Aldabra archipelago in recent times, in one or possibly two or more waves originating from Madagascar. The overall pattern of sunbird radiation across Indian Ocean islands indicates that these birds disperse across ocean barriers with relative ease, but that their subsequent evolutionary success probably depends on a variety of factors including prior island occupation by competing species.     

Diversity and distribution of avian haematozoan parasites in the western Indian Ocean region: a molecular survey

The genetic diversity of haematozoan parasites in island avifauna has only recently begun to be explored, despite the potential insight that these data can provide into the history of association between hosts and parasites and the possible threat posed to island endemics. We used mitochondrial DNA sequencing to characterize the diversity of 2 genera of vector-mediated parasites (Plasmodium and Haemoproteus) in avian blood samples from the western Indian Ocean region and explored their relationship with parasites from continental Africa. We detected infections in 68 out of 150 (45·3%) individuals and cytochrome b sequences identified 9 genetically distinct lineages of Plasmodium spp. and 7 lineages of Haemoproteus spp. We found considerable heterogeneity in parasite lineage composition across islands, although limited sampling may, in part, be responsible for perceived differences. Two lineages of Plasmodium spp. and 2 lineages of Haemoproteus spp. were shared by hosts in the Indian Ocean and also on mainland Africa, suggesting that these lineages may have arrived relatively recently. Polyphyly of island parasites indicated that these parasites were unlikely to constitute an endemic radiation and instead probably represent multiple colonization events. This study represents the first molecular survey of vector-mediated parasites in the western Indian Ocean, and has uncovered a diversity of parasites. Full understanding of parasite community composition and possible threats to endemic avian hosts will require comprehensive surveys across the avifauna of this region.     

Ancient mitochondrial DNA and morphology elucidate an extinct island radiation of Indian Ocean giant tortoises (Cylindraspis).

Ancient mitochondrial DNA sequences were used for investigating the evolution of an entire clade of extinct vertebrates, the endemic tortoises (Cylindraspis) of the Mascarene Islands in the Indian Ocean. Mitochondrial DNA corroborates morphological evidence that there were five species of tortoise with the following relationships: Cylindraspis triserrata ((Cylindraspis vosmaeri and Cylindraspis peltastes) (Cylindraspis inepta and Cylindraspis indica)). Phylogeny indicates that the ancestor of the group first colonized Mauritius where speciation produced C. triserrata and the ancestor of the other species including a second sympatric Mauritian form, C. inepta. A propagule derived from this lineage colonized Rodrigues 590 km to the east, where a second within-island speciation took place producing the sympatric C. vosmaeri and C. peltastes. A recent colonization of Réunion 150 km to the southwest produced C. indica. In the virtual absence of predators, the defensive features of the shells of Mascarene tortoises were largely dismantled, apparently in two stages. 'Saddlebacked' shells with high fronts evolved independently on all three islands. This and other features, such as a derived jaw structure and small body size, may be associated with niche differentiation in sympatric species and may represent a striking example of parallel differentiation in a large terrestrial vertebrate. The history of Mascarene tortoises contrasts with that of the Galápagos, where only a single species is present and surviving populations are genetically much more similar. However, they too show some reduction in anti-predator mechanisms and multiple development of populations with saddlebacked shells.     

Biogeography and diversification of hermit spiders on Indian Ocean islands (Nephilidae: Nephilengys)

The origin of the terrestrial biota of Madagascar and, especially, the smaller island chains of the western Indian Ocean is relatively poorly understood. Madagascar represents a mixture of Gondwanan vicariant lineages and more recent colonizers arriving via Cenozoic dispersal, mostly from Africa. Dispersal must explain the biota of the smaller islands such as the Comoros and the chain of Mascarene islands, but relatively few studies have pinpointed the source of colonizers, which may include mainland Africa, Asia, Australasia, and Madagascar. The pantropical hermit spiders (genus Nephilengys) seem to have colonized the Indian Ocean island arc stretching from Comoros through Madagascar and onto Mascarenes, and thus offer one opportunity to reveal biogeographical patterns in the Indian Ocean. We test alternative hypotheses on the colonization route of Nephilengys spiders in the Indian Ocean and simultaneously test the current taxonomical hypothesis using genetic and morphological data. We used mitochondrial (COI) and nuclear (ITS2) markers to examine Nephilengys phylogenetic structure with samples from Africa, southeast Asia, and the Indian Ocean islands of Madagascar, Mayotte, Réunion and Mauritius. We used Bayesian and parsimony methods to reconstruct phylogenies and haplotype networks, and calculated genetic distances and fixation indices. Our results suggest an African origin of Madagascar Nephilengys via Cenozoic dispersal, and subsequent colonization of the Mascarene islands from Madagascar. We find strong evidence of gene flow across Madagascar and through the neighboring islands north of it, while phylogenetic trees, haplotype networks, and fixation indices all reveal genetically isolated and divergent lineages on Mauritius and Réunion, consistent with female color morphs. These results, and the discovery of the first males from Réunion and Mauritius, in turn falsify the existing taxonomic hypothesis of a single widespread species, Nephilengys borbonica, throughout the archipelago. Instead, we diagnose three Nephilengys species: Nephilengys livida (Vinson, 1863) from Madagascar and Comoros, N. borbonica (Vinson, 1863) from Réunion, and Nephilengys dodo new species from Mauritius. Nephilengys followed a colonization route to Madagascar from Africa, and on through to the Mascarenes, where it speciated on isolated islands. The related golden orb-weaving spiders, genus Nephila, have followed the same colonization route, but Nephila shows shallower divergencies, implying recent colonization, or a moderate level of gene flow across the archipelago preventing speciation. Unlike their synanthropic congeners, N. borbonica and N. dodo are confined to pristine island forests and their discovery calls for evaluation of their conservation status.     

In and out of Madagascar: Dispersal to Peripheral Islands, Insular Speciation and Diversification of Indian Ocean Daisy Trees (Psiadia, Asteraceae)

Madagascar is surrounded by archipelagos varying widely in origin, age and structure. Although small and geologically young, these archipelagos have accumulated disproportionate numbers of unique lineages in comparison to Madagascar, highlighting the role of waif-dispersal and rapid in situ diversification processes in generating endemic biodiversity. We reconstruct the evolutionary and biogeographical history of the genus Psiadia (Asteraceae), a plant genus with near equal numbers of species in Madagascar and surrounding islands. Analyzing patterns and processes of diversification, we explain species accumulation on peripheral islands and aim to offer new insights on the origin and potential causes for diversification in the Madagascar and Indian Ocean Islands biodiversity hotspot. Our results provide support for an African origin of the group, with strong support for non-monophyly. Colonization of the Mascarenes took place by two evolutionary distinct lineages from Madagascar, via two independent dispersal events, each unique for their spatial and temporal properties. Significant shifts in diversification rate followed regional expansion, resulting in co-occurring and phenotypically convergent species on high-elevation volcanic slopes. Like other endemic island lineages, Psiadia have been highly successful in dispersing to and radiating on isolated oceanic islands, typified by high habitat diversity and dynamic ecosystems fuelled by continued geological activity. Results stress the important biogeographical role for Rodrigues in serving as an outlying stepping stone from which regional colonization took place. We discuss how isolated volcanic islands contribute to regional diversity by generating substantial numbers of endemic species on short temporal scales. Factors pertaining to the mode and tempo of archipelago formation and its geographical isolation strongly govern evolutionary pathways available for species diversification, and the potential for successful diversification of dispersed lineages, therefore, appears highly dependent on the timing of arrival, as habitat and resource properties change dramatically over the course of oceanic island evolution.     

Sequential colonization and diversification of Galapágos endemic land snail genus Bulimulus (Gastropoda, Stylommatophora)

Species richness on island or islandlike systems is a function of colonization, within-island speciation, and extinction. Here we evaluate the relative importance of the first two of these processes as a function of the biogeographical and ecological attributes of islands using the Galápagos endemic land snails of the genus Bulimulus, the most species-rich radiation of these islands. Species in this clade have colonized almost all major islands and are found in five of the six described vegetation zones. We use molecular phylogenetics (based on COI and ITS 1 sequence data) to infer the diversification patterns of extant species of Bulimulus, and multiple regression to investigate the causes of variation among islands in species richness. Maximum-likelihood, Bayesian, and maximum-parsimony analyses yield well-resolved trees with similar topologies. The phylogeny obtained supports the progression rule hypothesis, with species found on older emerged islands connecting at deeper nodes. For all but two island species assemblages we find support for only one or two colonization events, indicating that within-island speciation has an important role in the formation of species on these islands. Even though speciation through colonization is not common, island insularity (distance to nearest major island) is a significant predictor of species richness resulting from interisland colonization alone. However, island insularity has no effect on the overall bulimulid species richness per island. Habitat diversity (measured as plant species diversity), island elevation, and island area, all of which are indirect measures of niche space, are strong predictors of overall bulimulid land snail species richness. Island age is also an important independent predictor of overall species richness, with older islands harboring more species than younger islands. Taken together, our results demonstrate that the diversification of Galápagos bulimulid land snails has been driven by a combination of geographic factors (island age, size, and location), which affect colonization patterns, and ecological factors, such as plant species diversity, that foster within-island speciation.     

Subantarctic flowering plants: pre‐glacial survivors or post‐glacial immigrants?

Aim The aim here was to assess whether the present‐day assemblage of subantarctic flowering plants is the result of a rapid post‐Last Glacial Maximum (LGM) colonization or whether subantarctic flowering plants survived on the islands in glacial refugia throughout the LGM. Location The circumpolar subantarctic region, comprising six remote islands and island groups between latitudes 46° and 55° S, including South Georgia in the South Atlantic Ocean, the Prince Edward Islands, Îles Crozet, Îles Kerguelen, the Heard Island group in the South Indian Ocean and Macquarie Island in the South Pacific Ocean. Methods Floristic affinities between the subantarctic islands were assessed by cluster analysis applied to an up‐to‐date dataset of the phanerogamic flora in order to test for the existence of provincialism within the subantarctic. A review of the primary literature on the palaeobotany, geology and glacial history of the subantarctic islands was carried out and supplemented with additional palaeobotanical data and new field observations from South Georgia, Île de la Possession (Îles Crozet) and Îles Kerguelen. Results First, a strong regionalism was observed, with different floras characterizing the islands in each of the ocean basins, and endemic species being present in the South Indian Ocean and South Pacific Ocean provinces. Second, the majority of the plant species were present at the onset of accumulation of post‐glacial organic sediment and there is no evidence for the natural arrival of new immigrants during the subsequent period. Third, a review of geomorphological data suggested that the ice cover was incomplete during the LGM on the majority of the islands, and ice‐free biological refugia were probably present even on the most glaciated islands. Main conclusions Several independent lines of evidence favour the survival of a native subantarctic phanerogamic flora in local refugia during the LGM rather than a post‐LGM colonization from more distant temperate landmasses in the Southern Hemisphere.     

Deciphering patterns of transoceanic dispersal: the evolutionary origin and biogeography of coastal lizards (Cryptoblepharus) in the Western Indian Ocean region

Aim Cryptoblepharus is a genus of small arboreal or rock‐dwelling scincid lizards, widespread through the Indo‐Pacific and Australian regions, with a disjunct outlier in the Malagasy region. The taxonomy within this genus is controversial, with different authors ranking the different forms (now some 36) at various levels, from different species to subspecies of a single species, Cryptoblepharus boutonii. We investigated the biogeography and genetic differentiation of the Cryptoblepharus from the Western Indian Ocean region, in order to understand their origin and history. Location Western Indian Ocean region. Methods We analysed sequences of mitochondrial DNA (partial 12s and 16s rRNA genes, 766 bp) from 48 specimens collected in Madagascar, Mauritius, the four Comoros islands and East Africa, and also in New Caledonia, representing the Australo‐Pacific unit of the distribution. Results Pairwise sequence divergences of c. 3.1% were found between the New Caledonian forms and the ones from the Western Indian Ocean. Two clades were identified in Madagascar, probably corresponding to the recognized forms cognatus and voeltzkowi, and two clades were identified in the Comoro islands, where each island population formed a distinct haplotype clade. The East African samples form a monophyletic unit, with some variation existing between Pemba, Zanzibar and continental Tanzania populations. Individuals from Mauritius form a divergent group, more related to populations from Moheli and Grand Comore (Comoros islands) than to the others. Main conclusions The level of divergence between the populations from the Western Indian Ocean and Australian regions and the geographic coherence of the variation within the Western Indian Ocean group are concordant with the hypothesis of a colonization of this region by a natural transoceanic dispersal (from Australia or Indonesia). The group then may have diversified in Madagascar, from where it separately colonized the East African coast, the Comoros islands (twice), and Mauritius. The genetic divergence found is congruent with the known morphological variation, but its degree is much lower than typically seen between distinct species of reptiles.     

The evolutionary origin of Indian Ocean tortoises (Dipsochelys)

Today, the only surviving wild population of giant tortoises in the Indian Ocean occurs on the island of Aldabra. However, giant tortoises once inhabited islands throughout the western Indian Ocean. Madagascar, Africa, and India have all been suggested as possible sources of colonization for these islands. To address the origin of Indian Ocean tortoises (Dipsochelys, formerly Geochelone gigantea), we sequenced the 12S, 16S, and cyt b genes of the mitochondrial DNA. Our phylogenetic analysis shows Dipsochelys to be embedded within the Malagasy lineage, providing evidence that Indian Ocean giant tortoises are derived from a common Malagasy ancestor. This result points to Madagascar as the source of colonization for western Indian Ocean islands by giant tortoises. Tortoises are known to survive long oceanic voyages by floating with ocean currents, and thus, currents flowing northward towards the Aldabra archipelago from the east coast of Madagascar would have provided means for the colonization of western Indian Ocean islands. Additionally, we found an accelerated rate of sequence evolution in the two Malagasy Pyxis species examined. This finding supports previous theories that shorter generation time and smaller body size are related to an increase in mitochondrial DNA substitution rate in vertebrates.     

Establishing a perimeter position: speciation around the Indian Ocean Basin

Historical biological interactions among peripheral landmasses on the periphery of the Indian Ocean Basin (IOB) are generally poorly understood. While interactions based on early Gondwanan vicariance have been used to explain present day lineage distributions, several recent studies have instead inferred dispersal across the IOB. This inference is often advanced because lineages under study have species inhabiting IOB islands. Here we examine the roles of continental vicariance vs. trans-IOB dispersal in the distribution of an avian genus found around the perimeter of the IOB. A molecular phylogeny does reveal evidence of a relationship that would require the inference of trans-IOB dispersal between eastern Africa and Sri Lanka. However, molecular clock data, ancestral area analyses and paleoclimatic reconstructions suggest that vicariance related to tropical forest expansion and retraction is more likely to have facilitated African-Asian interchange, with an initial colonization of Africa from Asia quickly followed by a recolonization of Asia. Subsequent dispersal from Asia to Sri Lanka and islands east of the Sunda Shelf are inferred; these latter islands were colonized in a stepping-stone fashion that culminated in colonization of the Sunda Shelf, and a recolonization of mainland Asia. We propose that circum-IOB distributions, which post-date early Gondwanan breakup, are most likely the result of continent-based vicariant events, particularly those events related to large-scale habitat alterations, and not trans-IOB dispersals.     

Diversification of sympatric Sapromyza (Diptera: Lauxaniidae) from Madeira: six morphological species but only four mtDNA lineages

A series of recent studies on speciation of insects within the Canary Islands have indicated considerable within-island diversification, similar to that described in the Hawaiian islands. Little work has yet been carried out on the neighboring Madeiran archipelago, which is also volcanic. This study examines relationships among all known Lauxaniid flies of the genus Sapromyza from Madeira (including six newly described morphological species) based on mitochondrial gene trees constructed from cytochrome c oxidase (subunit I) and 16S rRNA partial sequences. Phylogenies based on maximum likelihood distances, a Bayesian method based on Markov chain Monte Carlo sampling from the posterior probability distribution, and maximum parsimony show that eight of the nine Madeiran species comprise a single monophyletic group. This clade is also split into two subclades representing black- and yellow/orange-bodied forms. The latter mtDNA clade corresponds to only two species (Sapromyza imitans and Sapromyza indigena) which are not reciprocally monophyletic. Monophyly is strongly supported within four of the six black-bodied species but not for the species pair (Sapromyza inconspicua, Sapromyza laurisilvae). We discuss the double occurrence (at least) of introgressive hybridization/incomplete lineage sorting within this group and suggest that recent speciation is the most likely explanation. The remaining species on the island, Sapromyza madeirensis, is very divergent from the aforementioned group, occupying a more basal position in the tree than the other Atlantic island and continental Sapromyza that were included in the analysis. At least two speciation events for Madeiran Sapromyza appear to correspond to quite ancient periods relative to the age of the island, while others are more recent. This suggests that a combination of island colonization and within-island sympatric and/or vicariance-mediated speciation may explain the observed diversity.     

Phylogeny and colonization history of Pringlea antiscorbutica (Brassicaceae), an emblematic endemic from the South Indian Ocean Province

The origins and evolution of sub-Antarctic island floras are not well understood. In particular there is uncertainty about the ages of the contemporary floras and the ultimate origins of the lineages they contain. Pringlea R. Br. (Brassicaceae) is a monotypic genus endemic to four sub-Antarctic island groups in the southern Indian Ocean. Here we used sequences from both the chloroplast and nuclear genomes to examine the phylogenetic position of this enigmatic genus. Our analyses confirm that Pringlea falls within the tribe Thelypodieae and provide a preliminary view of its relationships within the group. Divergence time estimates and ancestral area reconstructions imply Pringlea diverged from a South American ancestor ～5Myr ago. It remains unclear whether the ancestor of Pringlea dispersed directly to the South Indian Ocean Province (SIOP) or used Antarctica as a stepping-stone; what is clear, however, is that following arrival in the SIOP several additional long-distance dispersal events must be inferred to explain the current distribution of this species. Our analyses also suggest that although Pringlea is likely to have inherited cold tolerance from its closest relatives, the distinctive morphology of this species evolved only after it split from the South American lineage. More generally, our results lend support to the hypothesis that angiosperms persisted on the sub-Antarctic islands throughout the Pliocene and Pleistocene. Taken together with evidence from other sub-Antarctic island plant groups, they suggest the extant flora of sub-Antarctic is likely to have been assembled over a broad time period and from lineages with distinctive biogeographic histories.     

Lack of premating isolation at the base of a phylogenetic tree

Darwin's finches in the Galápagos archipelago are an unusual example of adaptive radiation in that the basal split separates two lineages of warbler finches (Certhidea olivacea and Certhidea fusca) believed until recently to be only one species. The large genetic difference between them contrasts with their similarity in plumage, size, shape, and courtship behavior. They differ in song, which is a key factor in premating isolation of other sympatric Darwin's finches. We conducted playback experiments to see whether members of the population of C. olivacea on Santa Cruz Island would respond to songs of C. fusca from two islands, Genovesa and Pinta, and songs of C. olivacea from another island (Isabela). Another set of experiments was performed, using the same playback tapes, with C. fusca on Genovesa. Some members of both populations responded to all playbacks; therefore, the hypothesis of complete premating isolation on the basis of song is rejected. Discrimination between songs of the two lineages was inconsistent. We conclude that premating barriers to interbreeding among the tested populations have not arisen in the 1.5-2.0 m.yr. of their geographical isolation on different islands. This contrasts with strong premating barriers between more recently derived sympatric species. Early learning of song associated with morphology is later used in mate recognition. This explains why sympatric species that are vocally and morphologically distinct yet genetically less differentiated than Certhidea do not interbreed, whereas the Certhidea lineages that are genetically well differentiated but vocally and morphologically similar have no apparent premating barrier. We discuss this unusual situation in terms of the forces that have produced similarities and differences in song, morphology, and ecology and their relevance to phylogenetic and biological species concepts. Neither principles nor details are unique to Darwin's finches, and we conclude by pointing out strong parallels with some continental birds.     

Linking Vital Rates of Landbirds on a Tropical Island to Rainfall and Vegetation Greenness

Remote tropical oceanic islands are of high conservation priority, and they are exemplified by range-restricted species with small global populations. Spatial and temporal patterns in rainfall and plant productivity may be important in driving dynamics of these species. Yet, little is known about environmental influences on population dynamics for most islands and species. Here we leveraged avian capture-recapture, rainfall, and remote-sensed habitat data (enhanced vegetation index [EVI]) to assess relationships between rainfall, vegetation greenness, and demographic rates (productivity, adult apparent survival) of three native bird species on Saipan, Northern Mariana Islands: rufous fantail (Rhipidura rufifrons), bridled white-eye (Zosterops conspicillatus), and golden white-eye (Cleptornis marchei). Rainfall was positively related to vegetation greenness at all but the highest rainfall levels. Temporal variation in greenness affected the productivity of each bird species in unique ways. Predicted productivity of rufous fantail was highest when dry and wet season greenness values were high relative to site-specific 5-year seasonal mean values (i.e., relative greenness); while the white-eye species had highest predicted productivity when relative greenness contrasted between wet and dry seasons. Survival of rufous fantail and bridled white eye was positively related to relative dry-season greenness and negatively related to relative wet-season greenness. Bridled white-eye survival also showed evidence of a positive response to overall greenness. Our results highlight the potentially important role of rainfall regimes in affecting population dynamics of species on oceanic tropical islands. Understanding linkages between rainfall, vegetation, and animal population dynamics will be critical for developing effective conservation strategies in this and other regions where the seasonal timing, extent, and variability of rainfall is expected to change in the coming decades.     

Niche divergence promotes rapid diversification of East African sky island white‐eyes (Aves: Zosteropidae)

The Eastern Afromontane biodiversity hotspot composed of highly fragmented forested highlands (sky islands) harbours exceptional diversity and endemicity, particularly within birds. To explain their elevated diversity within this region, models founded on niche conservatism have been offered, although detailed phylogeographic studies are limited to a few avian lineages. Here, we focus on the recent songbird genus Zosterops, represented by montane and lowland members, to test the roles of niche conservatism versus niche divergence in the diversification and colonization of East Africa's sky islands. The species‐rich white‐eyes are a typically homogeneous family with an exceptional colonizing ability, but in contrast to their diversity on oceanic islands, continental diversity is considered depauperate and has been largely neglected. Molecular phylogenetic analysis of ~140 taxa reveals extensive polyphyly among different montane populations of Z. poliogastrus. These larger endemic birds are shown to be more closely related to taxa with divergent habitat types, altitudinal distributions and dispersal abilities than they are to populations of restricted endemics that occur in neighbouring montane forest fragments. This repeated transition between lowland and highland habitats over time demonstrate that diversification of the focal group is explained by niche divergence. Our results also highlight an underestimation of diversity compared to morphological studies that has implications for their taxonomy and conservation. Molecular dating suggests that the spatially extensive African radiation arose exceptionally rapidly (1–2.5 Ma) during the fluctuating Plio‐Pleistocene climate, which may have provided the primary driver for lineage diversification.     

Why does the biota of the Madagascar region have such a strong Asiatic flavour?

A corollary of island biogeographical theory is that islands are largely colonized from their nearest mainland source. Despite Madagascar’s extreme isolation from India and proximity to Africa, a high proportion of the biota of the Madagascar region has Asian affinities. This pattern has rarely been viewed as surprising, as it is consistent with Gondwanan vicariance. Molecular phylogenetic data provide strong support for such Asian affinities, but often not for their vicariant origin; most divergences between lineages in Asia and the Madagascar region post‐date the separation of India and Madagascar considerably (up to 87 Myr), implying a high frequency of dispersal that mirrors colonization of the Hawaiian archipelago in distance. Indian Ocean bathymetry and the magnitude of recent sea‐level lowstands support the repeated existence of sizeable islands across the western Indian Ocean, greatly reducing the isolation of Madagascar from Asia. We put forward predictions to test the role of this historical factor in the assembly of the regional biota. © The Willi Hennig Society 2009.