Testing for biogeographic mechanisms promoting divergence in Caribbean crickets (genus Amphiacusta)

Aim This work examines whether the history of diversification of Amphiacusta (Orthoptera, Gryllidae) in the Caribbean corresponds to a vicariant or a dispersalist model. Location The Greater Antillean islands of the Caribbean region. Methods The phylogenetic relationships among species were estimated using a procedure that directly estimates the underlying species tree from independent loci (in this case, one mitochondrial and one nuclear locus). This tree was then used to test for topological congruence with a vicariant model, and to estimate divergence times. Results The analyses based on the expected pattern of species divergence (i.e. species‐tree topology) support a vicariant model. With the notable exception of a dispersal event marking the colonization of Jamaica, the timing of the events are generally consistent with a vicariant scenario, given the current taxon sampling and potential errors with dating the divergence events. Main conclusions The tendency of species to co‐segregate by island suggests that intra‐island diversification is common. Despite their flightlessness, species of Amphiacusta are apparently capable of long‐distance dispersal, such as colonization from the Puerto Rican/Virgin Island bank to Jamaica. The topology of the species tree is consistent with a vicariant model of divergence, and the dates of divergence between island groups are generally consistent with an island–island vicariance model. A strict island–island vicariance scenario can, however, be rejected because of inferred dispersal events such as the colonization of Jamaica. Nevertheless, the biogeographic tests suggest that most of the diversity was generated under a combination of intra‐island diversification and island–island vicariance. Additional sampling of taxa will be needed to verify this hypothesized scenario. Our findings indicate that Amphiacusta presents an ideal opportunity for examining the role of sexual selection in promoting diversification, which would complement the large number of studies focused on adaptive divergence of Caribbean taxa.     

Invertébrés Sans Frontières: Large Scales of Connectivity of Selected Freshwater Species among Caribbean Islands

The freshwater fauna (crustaceans, molluscs, fish) of many tropical islands in the Caribbean and Pacific share an amphidromous life‐cycle, meaning their larvae need to develop in saline conditions before returning to freshwater as juveniles. This community dominates the freshwaters of much of the tropics, but is poorly known and at risk from development, in particular dam construction. Amphidromy can theoretically lead to dispersal between different freshwater areas, even to distant oceanic islands, via the sea. The extent and scale of this presumed dispersal, however, is largely unknown in the Caribbean. Recent genetic work in Puerto Rico has shown that many freshwater species have little or no population structure among different river catchments, implying high levels of connectivity within an island, whereas between‐island structure is unknown. We used genetic techniques to infer the geographic scales of population structure of amphidromous invertebrates (a gastropod and a number of crustacean species) between distant parts of the Caribbean, in particular Puerto Rico, Panama and Trinidad. We found virtually no geographic population structure across over 2000 km of open sea for these freshwater species. This implies that they are indeed moving between islands in sea currents as larvae, meaning that continued recruitment requires a continuum of healthy habitat from the freshwater to marine environment. We further discuss the role of amphidromy and suggest its ecological and biogeographic role may be more important than previously presumed.     

Colonization history and genetic diversity: adaptive potential in early stage invasions

The introduction of Anolis cristatellus from the multiple species anole community of Puerto Rico in the Greater Antilles to the island of Dominica in the Lesser Antilles, with its solitary endemic anole, provides an example of a very recent, timed, single colonization. We investigate the geographic origin and adaptive potential of the Dominican population using a range of methods including mtDNA phylogeography, nuclear microsatellite variation and multiple paternity studies, as well as heritability estimates, common garden experiments and comparative geographic studies of quantitative scalation traits. Phylogeographic analysis of NADH2 and microsatellite studies suggests that the Dominican population arose from a set of individuals from the central west area of Puerto Rico within their endemic range. The multiple‐individual inoculation, together with sperm storage and evidence of multiple paternity indicate genetic variability and suggest the potential for adaptation by natural selection. Estimates of heritability, common garden experiments and broad sense Q_ST/F_ST ratios, linked to replicated comparisons along elevational transects go some way to suggesting that the invasive populations may be adapting by natural selection, in parallel with the endemic anole, in the brief period since their introduction.     

Influence of a simulated hurricane on aquatic insect recolonization in the phytotelma of Heliconia caribaea (Heliconiacaea)

Disturbances like hurricanes can affect diversity and community composition, which may in turn affect ecosystem function. We examined how a simulated hurricane disturbance affected insect communities inhabiting the phytotelma (plant-held waters) of Heliconia caribaea in the Luquillo Experimental Forest of eastern Puerto Rico, a tropical island that frequently experiences hurricanes. We hypothesized that disturbance would alter diversity and that larger Heliconia would attract more species following disturbance due to the area-diversity relationship described by the Theory of Island Biogeography. Individual flower parts (bracts) of Heliconia inflorescences (racemes) were artificially disturbed via removal of existing insect communities, then after refilling with water, cohorts of Heliconia were destructively sampled biweekly for 6 weeks to assess recolonization patterns of α (bract level), β, and γ (summed across bracts; raceme level) diversity over time and across raceme sizes. Although we found no support for our hypothesis about the effect of raceme size on recolonization, our hypothesis regarding recolonization patterns over time was supported; species richness, evenness, and abundance of bracts increased directly after the disturbance and then decreased below pre-disturbance levels, and community composition at the raceme level changed significantly over time during recolonization. β Diversity was also greater in smaller racemes compared to larger racemes, suggesting high heterogeneity across bracts of Heliconia racemes exacerbated by raceme size and age. Overall, our results highlight the importance of scale and appropriate measurements of diversity (particularly α) in experiments aiming to extrapolate conclusions about the ecological impacts of disturbances across different habitats and ecosystems.     

Sea level,topography and island diversity: phylogeography of the Puerto Rican Red‐eyed Coquí, Eleutherodactylus antillensis

Quaternary climatic oscillations caused changes in sea level that altered the size, number and degree of isolation of islands, particularly in land‐bridge archipelagoes. Elucidating the demographic effects of these oscillations increases our understanding of the role of climate change in shaping evolutionary processes in archipelagoes. The Puerto Rican Bank (PRB) (Puerto Rico and the Eastern Islands, which comprise Vieques, Culebra, the Virgin Islands and associated islets) in the eastern Caribbean Sea periodically coalesced during glaciations and fragmented during interglacial periods of the quaternary. To explore population‐level consequences of sea level changes, we studied the phylogeography of the frog Eleutherodactylus antillensis across the archipelago. We tested hypotheses encompassing vicariance and dispersal narratives by sequencing mtDNA (c. 552 bp) of 285 individuals from 58 localities, and four nuDNA introns (totalling c. 1633 bp) from 173 of these individuals. We found low support for a hypothesis of divergence of the Eastern Islands populations prior to the start of the penultimate interglacial c. 250 kya, and higher support for a hypothesis of colonization of the Eastern Islands from sources in eastern Puerto Rico during the penultimate and last glacial period, when a land bridge united the PRB. The Río Grande de Loíza Basin in eastern Puerto Rico delineates a phylogeographic break. Haplotypes shared between the PRB and St. Croix (an island c. 105 km south‐east of this archipelago) likely represent human‐mediated introductions. Our findings illustrate how varying degrees of connectivity and isolation influence the evolution of tropical island organisms.     

Colonization of islands in the Mona Passage by endemic dwarf geckoes (genus Sphaerodactylus) reconstructed with mitochondrial phylogeny

Little is known about the natural history of the Sphaerodactylus species endemic to the three islands located in the Mona Passage separating the Greater Antillean islands of Hispaniola and Puerto Rico. In this study, parts of two mitochondrial genes, 16S rRNA and 12S rRNA, were sequenced to determine the relationships between the sphaerodactylids that live in the Mona Passage and other Caribbean species from the same genus. While the main goal was to identify the biogeographical origin of these species, we also identified a genetically distinct type of dwarf gecko that warrants future evaluation as a possible new species. According to the reconstructed phylogenies, we propose a stepwise model of colonization wherein S. nicholsi from southwestern Puerto Rico or a very close ancestor gave rise through a founder event to Sphaerodactylus monensis on Mona Island. In a similar fashion, S. monensis or a very close ancestor on Mona Island gave rise to S. levinsi on Desecheo Island. This study also suggests that the most recent common ancestor between the species from the islands in the Mona Passage and Puerto Rico existed approximately 3 MYA.     

Deep intra‐island divergence of a montane forest endemic: phylogeography of the Puerto Rican frog Eleutherodactylus portoricensis (Anura: Eleutherodactylidae)

Aim Hypotheses proposed for lineage diversification of tropical montane species have rarely been tested within oceanic islands. Our goal was to understand how basin barriers and Pleistocene climatic fluctuations shaped the distribution of diversity in Eleutherodactylus portoricensis (Eleutherodactylidae), a frog endemic to the montane rain forests of Puerto Rico. Location The north‐eastern (Luquillo) and south‐eastern (Cayey) mountains of Puerto Rico. Methods We generated mitochondrial DNA (mtDNA) control region sequences (c. 565 bp) from 144 individuals of E. portoricensis representing 16 localities, and sequenced 646 bp of cytochrome b and 596 bp of nuclear DNA (nDNA) rhodopsin exon and intron 1 from a subset of individuals. We conducted a phylogenetic analysis on the mtDNA sequence data and explored population substructure with maximum parsimony networks, a spatial analysis of molecular variance, and pairwise F_ST analysis. Coalescent simulations were performed to test alternative models of population divergence in response to late Pleistocene interglacial periods. Historical demography was assessed through coalescent analyses and Bayesian skyline plots. Results We found: (1) two highly divergent groups associated with the disjunct Luquillo and Cayey Mountains, respectively; (2) a shallow mtDNA genetic discontinuity across the La Plata Basin within the Cayey Mountains; (3) phylogeographic congruence between nDNA and mtDNA markers; (4) divergence dates for both mtDNA and nDNA pre‐dating the Holocene interglacial (c. 10 ka), and nDNA suggesting divergence in the penultimate interglacial (c. 245 ka); and (5) historical demographic stability in both lineages. Main conclusions The low‐elevation Caguas Basin is a long‐term barrier to gene flow between the two montane frog populations. Measures of genetic diversity for mtDNA were similar in both lineages, but lower nDNA diversity in the Luquillo Mountains lineage suggests infrequent dispersal between the two mountain ranges and colonization by a low‐diversity founder population. Population divergence began prior to the Holocene interglacial. Stable population sizes over time indicate a lack of demonstrable demographic response to climatic changes during the last glacial period. This study highlights the importance of topographic complexity in promoting within‐island vicariant speciation in the Greater Antilles, and indicates long‐term persistence and lineage diversification despite late Pleistocene climatic oscillations.     

Genomic and phenomic analysis of island ant community assembly

Island biodiversity has long fascinated biologists as it typically presents tractable systems for unpicking the eco‐evolutionary processes driving community assembly. In general, two recurring themes are of central theoretical interest. First, immigration, diversification, and extinction typically depend on island geographical properties (e.g., area, isolation, and age). Second, predictable ecological and evolutionary trajectories readily occur after colonization, such as the evolution of adaptive trait syndromes, trends toward specialization, adaptive radiation, and eventual ecological decline. Hypotheses such as the taxon cycle draw on several of these themes to posit particular constraints on colonization and subsequent eco‐evolutionary dynamics. However, it has been challenging to examine these integrated dynamics with traditional methods. Here, we combine phylogenomics, population genomics and phenomics, to unravel community assembly dynamics among Pheidole (Hymenoptera, Formicidae) ants in the isolated Fijian archipelago. We uphold basic island biogeographic predictions that isolated islands accumulate diversity primarily through in situ evolution rather than dispersal, and population genomic support for taxon cycle predictions that endemic species have decreased dispersal ability and demography relative to regionally widespread taxa. However, rather than trending toward island syndromes, ecomorphological diversification in Fiji was intense, filling much of the genus‐level global morphospace. Furthermore, while most endemic species exhibit demographic decline and reduced dispersal, we show that the archipelago is not an evolutionary dead‐end. Rather, several endemic species show signatures of population and range expansion, including a successful colonization to the Cook islands. These results shed light on the processes shaping island biotas and refine our understanding of island biogeographic theory.     

Colonization history and population differentiation of the Honey Bees (Apis mellifera L.) in Puerto Rico

Honey bees (Apis mellifera L.) are the primary commercial pollinators across the world. The subspecies A. m. scutellata originated in Africa and was introduced to the Americas in 1956. For the last 60 years, it hybridized successfully with European subspecies, previous residents in the area. The result of this hybridization was called Africanized honey bee (AHB). AHB has spread since then, arriving to Puerto Rico (PR) in 1994. The honey bee population on the island acquired a mosaic of features from AHB or the European honey bee (EHB). AHB in Puerto Rico shows a major distinctive characteristic, docile behavior, and is called gentle Africanized honey bees (gAHB). We used 917 SNPs to examine the population structure, genetic differentiation, origin, and history of range expansion and colonization of gAHB in PR. We compared gAHB to populations that span the current distribution of A. mellifera worldwide. The gAHB population is shown to be a single population that differs genetically from the examined populations of AHB. Texas and PR groups are the closest genetically. Our results support the hypothesis that the Texas AHB population is the source of gAHB in Puerto Rico.     

Chance and determinism in the development of isolated communities

• 1 In the colonization of an island by potentially interacting species both the severity of the competition between them and the length of the interval between their arrivals should affect the likelihood of the second species becoming established. Between‐island variation in priority and the interval between colonizations will depend on the dispersal rates of the two species.
• 2 We predict that early colonization should be relatively deterministic, whereas later in the colonization process both the sequence of arrivals and the interval between them will be more variable.
• 3 Possible instances of both deterministic and stochastic stages in community development are identified in the colonization of the Krakatau islands by zoochorous forest trees and large zoochorous lianes.

     

Phylogeography of an Island endemic, the Puerto Rican Freshwater Crab (Epilobocera sinuatifrons)

The endemic Puerto Rican crab, Epilobocera sinuatifrons (Pseudothelphusidae), has a freshwater-dependant life-history strategy, although the species has some capabilities for terrestrial movement as adults. In contrast to all other freshwater decapods on the island (e.g., caridean shrimp), E. sinuatifrons does not undertake amphidromous migration, and is restricted to purely freshwater habitats and adjacent riparian zones. As Puerto Rico has a dynamic geologic history, we predicted that both the life history of E. sinuatifrons and the geological history of the island would be important determinants of phylogeographic structuring in the species. Using a fragment of the cytochrome c oxidase subunit 1 mtDNA (mitochondrial DNA) gene, we tested for deviations from panmixia among and within rivers draining Puerto Rico and used statistical phylogeography to explore processes that may explain extant patterns of genetic variation in the species. While populations of E. sinuatifrons were significantly differentiated among rivers, they were likely to be recently derived because nested clade analysis (NCA) indicated evolutionarily recent restricted gene flow with isolation by distance (IBD) and contiguous range expansion at various spatial scales. Ongoing drainage rearrangements associated with faulting and land slippage were invoked as processes involved in sporadic gene flow among rivers throughout the Pleistocene. Patterns of genetic differentiation conformed to IBD and population demographic statistics were nonsignificant, indicating that although recently derived, populations from different rivers were in drift-mutation equilibrium. A shallow (0.6 million years ago), paraphyletic split was observed in the haplotype network, which NCA indicated arose via allopatric fragmentation. This split coincides with an area of high relief in central Puerto Rico that may have experienced relatively little drainage rearrangements. Shallow but significant genetic isolation of populations of E. sinuatifrons among Puerto Rican rivers suggests phylogeographic patterns that are intermediate to terrestrial habitat specialists (highly divergent populations) and other freshwater biota, such as amphidromous species and insects with aerial adult dispersal (highly connected populations).     

Microbial colonization dynamics in temporary aquatic systems

Most studies of community development in insular systems have investigated the colonization dynamics of only a portion of the biotic community using islands that have the benefit of prior biotic modification. Two experiments assessed the predictive ability of equilibrium island theory with regard to the development of the protistan community in temporary aquatic islands (100-1 plastic swimming pools) void of any prior biotic modification. The first experiment ran for 170 d (March–September 1985) and manipulated the access of islands to animals that represent potentially important dispersal pathways for microbes. A second experiment (May–June 1986) investigated in greater detail the early stages of species accrual in the absence of dispersal pathways other than via the atmosphere. Polyurethane foam substrates were used as sampling devices to increase the accuracy and replicability of sampling and provide habitat for colonists. Sampling was determined to be asymptotic. Species accrual was asymptotic in both experiments, although it initially lagged behind that predicted by theory. Autotrophs approached equilibrium faster than heterotrophs, but at a lower species richness. The predicted number of autotroph species at equilibrium was lower in islands that were excluded from contact with birds compared to less exclosed islands. The colonization dynamics of the entire community was not significantly different among islands having different degrees of exclosure. In both experiments, rates of species immigration were nonmonotonic with respect to time and species richness. This relationship appeared to reflect the importance of species interactions during the initial accrual phase before equilibrium. Rates of extinction were positively correlated with both these parameters, although they tended to decrease with time during the equilibrium period in less exclosed islands. Turnover at equilibrium was significant and resulted in directional changes in species composition over time. Assortative processes appeared to be important since later colonists exhibited greater persistence. Colonizing species generally fall into three autecological categories: 1) those that were ubiquitous and had temporally predictable patterns of immigration (successional species); 2) those possessing temporally predictable distributions but not spatially ubiquitous distributions (dispersal limited species); 3) those that showed little temporal or spatial predictability in immigration (transient or allochthonous species). Individual islands exhibited various degrees of fluctuation in species number during the predicted equilibrium which were poorly correlated with exogenous environmental variables and physicochemical habitat parameters. The presence of predacious mosquito larvae(Culex spp.) invariably resulted in a sharp decreases in microbial species richness, while documented contact with rodents was followed by an increase in species number in an island so contaminated. Several aspects of microbial colonization of temporary waters that contradict equilibrium predictions appear to be strongly influenced by microbe-microbe as well as macrobe-microbe interactions.     

Multiple source pools for Galápagos plant species richness: a critical analysis of the line of sight connectivity index

An index (C_i*E) combining the number of line‐of‐sight islands (C_i) within a radius i and target island elevation (E) has been proposed as an improved predictive model of plant species richness (S_t) in the Galápagos Archipelago. We examined this index critically and found that several major flaws preclude it from being a useful predictive tool for the archipelago. Although the number of collecting trips to an island was reported over 20 years ago to have substantial predictive value for reported plant species richness in the Galápagos Islands, this relationship was ignored in multiple regression analyses of the index. When we included the number of collecting trips in different multiple regression analyses of the index, C_i*E had less predictive power than collecting trips or ceased to be significant at all. Additionally, the strong significant relationship between elevation and area in the Galápagos Archipelago results in area having a major confounding influence on the C_i*E index. When elevation is removed from the C_i*E index, the predictive power of C_i is far less than area alone. Finally, the data used to construct and correlate the C_i*E index with (S_t) were based only on a subset of the islands and species lists that were incomplete or out of date. Species richness on islands can be related to the interaction of different factors, so development and testing of indices like C_i*E is not inappropriate. However, it is important to examine the interrelationships among the components of these indices thoroughly, and not ignore the effect of factors already known to have high predictive power. We propose several ways in which more meaningful indices of source pool(s) capacity can be constructed.     

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.     

Otolith microchemistry of tropical diadromous fishes: spatial and migratory dynamics

Otolith microchemistry was applied to quantify migratory variation and the proportion of native Caribbean stream fishes that undergo full or partial marine migration. Strontium and barium water chemistry in four Puerto Rico, U.S.A., rivers was clearly related to a salinity gradient; however, variation in water barium, and thus fish otoliths, was also dependent on river basin. Strontium was the most accurate index of longitudinal migration in tropical diadromous fish otoliths. Among the four species examined, bigmouth sleeper Gobiomorus dormitor, mountain mullet Agonostomus monticola, sirajo goby Sicydium spp. and river goby Awaous banana, most individuals were fully amphidromous, but 9–12% were semi‐amphidromous as recruits, having never experienced marine or estuarine conditions in early life stages and showing no evidence of marine elemental signatures in their otolith core. Populations of one species, G. dormitor, may have contained a small contingent of semi‐amphidromous adults, migratory individuals that periodically occupied marine or estuarine habitats (4%); however, adult migratory elemental signatures may have been confounded with those related to diet and physiology. These findings indicate the plasticity of migratory strategies of tropical diadromous fishes, which may be more variable than simple categorization might suggest.     

Phylogeography of related diadromous species in continental and island settings,and a comparison of their potential and realized dispersal patterns

Aim To compare patterns of potential and realized dispersal in ecologically similar and phylogenetically related amphidromous shrimps (Atyidae) in continental and island‐dominated landscapes. Location Eastern Australia and the Caribbean region. Methods Population genetic and phylogeographic analyses of mitochondrial DNA data for Australatya striolata from eastern Australia (a continental landscape) and Atya scabra from the Caribbean (an island‐dominated landscape). Results Australatya striolata contained two highly divergent genetic lineages in eastern Australia, corresponding to the disjunct northern and southern populations, respectively. These lineages probably represent allopatric cryptic species, both of which were found to have genetically homogeneous population structures within their regions of occurrence. Atya scabra was genetically homogeneous throughout the Caribbean. Recent population expansions were detected for Atya scabra in the Caribbean, but not for northern or southern Australatya striolata. Main conclusions The findings of this study are consistent with previously reported patterns of genetic population structure in amphidromous species in both continental and island‐dominated landscapes, suggesting that potential for widespread dispersal is typically matched by realized patterns of panmixia. We therefore raise the hypothesis that landscape setting (i.e. continent or island‐dominated) does not influence dispersal patterns in amphidromous species. Further studies, especially of population genetic patterns of amphidromous species on continents, are needed to test this idea. Interestingly, results of the genetic neutrality tests led us to hypothesize that demographic and drift‐mutation equilibrium is attainable although not always evident for amphidromous species on continents, but is not attainable for those species distributed across island settings.     

Climate as a driver of tropical insular diversity: comparative phylogeography of two ecologically distinctive frogs in Puerto Rico

The effects of late Quaternary climate on distributions and evolutionary dynamics of insular species are poorly understood in most tropical archipelagoes. We used ecological niche models under past and current climate to derive hypotheses regarding how stable climatic conditions shaped genetic diversity in two ecologically distinctive frogs in Puerto Rico. Whereas the mountain coquí Eleutherodactylus portoricensis is restricted to montane forest in the Cayey and Luquillo Mountains, the red‐eyed coquí E. antillensis is a habitat generalist distributed across the entire Puerto Rican Bank (Puerto Rico and the Virgin Islands, excluding St Croix). To test our hypotheses, we conducted phylogeographic and population genetic analyses based on mitochondrial and nuclear loci of each species across their range in Puerto Rico. Patterns of population differentiation in E. portoricensis, but not in E. antillensis, supported our hypotheses. For E. portoricensis, these patterns include: individuals isolated by long‐term unsuitable climate in the Río Grande de Loíza Basin in eastern Puerto Rico belong to different genetic clusters; past and current climate strongly predicted genetic differentiation; and Cayey and Luquillo Mountains populations split prior to the last interglacial. For E. antillensis, these patterns include: genetic clusters did not fully correspond to predicted long‐term unsuitable climate; and past and current climate weakly predicted patterns of genetic differentiation. Genetic signatures in E. antillensis are consistent with a recent range expansion into western Puerto Rico, possibly resulting from climate change and anthropogenic influences. As predicted, regions with a large area of long‐term suitable climate were associated with higher genetic diversity in both species, suggesting larger and more stable populations. Finally, we discussed the implications of our findings for developing evidence‐based management decisions for E. portoricensis, a taxon of special concern. Our findings illustrate the role of persistent suitable climatic conditions in promoting the persistence and diversification of tropical island organisms.     

Re‐colonization by common eiders Somateria mollissima in the Aleutian Archipelago following removal of introduced arctic foxes Vulpes lagopus

Islands provide refuges for populations of many species where they find safety from predators, but the introduction of predators frequently results in elimination or dramatic reductions in island‐dwelling organisms. When predators are removed, re‐colonization for some species occurs naturally, and inter‐island phylogeographic relationships and current movement patterns can illuminate processes of colonization. We studied a case of re‐colonization of common eiders Somateria mollissima following removal of introduced arctic foxes Vulpes lagopus in the Aleutian Archipelago, Alaska. We expected common eiders to resume nesting on islands cleared of foxes and to re‐colonize from nearby islets, islands, and island groups. We thus expected common eiders to show limited genetic structure indicative of extensive mixing among island populations. Satellite telemetry was used to record current movement patterns of female common eiders from six islands across three island groups. We collected genetic data from these and other nesting common eiders at 14 microsatellite loci and the mitochondrial DNA control region to examine population genetic structure, historical fluctuations in population demography, and gene flow. Our results suggest recent interchange among islands. Analysis of microsatellite data supports satellite telemetry data of increased dispersal of common eiders to nearby areas and little between island groups. Although evidence from mtDNA is suggestive of female dispersal among island groups, gene flow is insufficient to account for recolonization and rapid population growth. Instead, near‐by remnant populations of common eiders contributed substantially to population expansion, without which re‐colonization would have likely occurred at a much lower rate. Genetic and morphometric data of common eiders within one island group two and three decades after re‐colonization suggests reduced movement of eiders among islands and little movement between island groups after populations were re‐established. We predict that re‐colonization of an island group where all common eiders are extirpated could take decades.     

Biogeographical kinetics on mainland and island volcanoes

Aim To compare post‐eruption biotic recolonization times on mainland and island volcanoes. Location The research involved the study of the recolonization kinetics of Mt Vesuvius (a mainland volcano in southern Italy) and the Island of Vulcano (southern Italy). Comparisons were also made with Jorullo Volcano (Mexico) and Mount St Helens (USA) (two mainland volcanoes) and with Krakatau (Indonesia) (an island volcano). Methods Island volcanoes are expected to possess inherently impoverished faunas and floras, and recolonization after eruption is expected to occur to a low level. In comparison, the recolonization kinetics for a mainland volcano should be characterized by a higher plateau of species, and by species with low dispersal ability. To test this model, recolonization times after the small‐scale Plinian eruption of 1631 were calculated for various insect groups of Mt Vesuvius and compared with recolonization times calculated for the Island of Vulcano, which erupted dramatically in 1888. For this purpose, thorough insect checklists, based on exhaustive samplings, were extracted from the literature. Results obtained from Mt Vesuvius and Vulcano were also compared with recolonization times calculated for biotas of other mainland and island volcanoes. Results Insect recolonization times from the 1631 Vesuvius eruption varied according to the ecology of the animal group considered and appeared very long when compared with those obtained for the island volcano of Vulcano. Results obtained from Vulcano also suggest the possibility that this island hosts more species than expected at equilibrium, a state of affairs also found for the butterflies of Krakatau. Main conclusions In keeping with the predictions of multi‐phase models developed in island biogeography theory, island volcanoes have a lower species richness at equilibrium than do mainland volcanoes, but might host, in a first phase of recolonization, more species than expected at equilibrium, because ecological space is unsaturated and inter‐specific interactions are limited. The lower isolation of mainland volcanoes (allowing higher immigration rates) leads to higher rates and longer periods of recolonization and hence to higher species richness at equilibrium.     

Insular black files (Diptera: Simuliidae) of North America: tests of colonization hypotheses

Aim To understand factors that facilitate insular colonization by black flies, we tested six hypotheses related to life‐history traits, phylogeny, symbiotes, island area, and distance from source areas. Location Four northern islands, all within 150 km of the North American mainland, were included in the study: Isle Royale, Magdalen Islands, Prince Edward Island, and Queen Charlotte Islands. Methods Immature black flies and their symbiotes were surveyed in streams on the Magdalen Islands, and the results combined with data from similar surveys on Isle Royale, Prince Edward Island, and the Queen Charlotte Islands. Black flies were analysed chromosomally to ensure that all sibling species were revealed. Tests of independence were used to examine the frequency of life‐history traits and generic representation of black flies on islands vs. source areas. Results A total of 13–20 species was found on each of the islands, but no species was unique to any of the islands. The simuliid faunas of the islands reflected the composition of their source areas in aspects of voltinism (univoltine vs. multivoltine), blood feeding (ornithophily vs. mammalophily), and phylogeny (genus Simulium vs. other genera). Five symbiotic species were found on the most distant island group, the Magdalen Islands, supporting the hypothesis that obligate symbiotes are effectively transported to near‐mainland islands. An inverse relationship existed between the number of species per island and distance from the source. The Queen Charlotte Islands did not conform to the species–area relationship. Main conclusions The lack of precinctive insular species and an absence of life‐history and phylogenetic characteristics related to the presence of black flies on these islands argue for gene flow and dispersal capabilities of black flies over open waters, possibly aided by winds. However, the high frequency of precinctive species on islands 500 km or more from the nearest mainland indicates that at some distance beyond 100 km, open water provides a significant barrier to colonization and gene exchange. An inverse relationship between number of species and distance from the source suggests that as long as suitable habitat is present, distance plays an important role in colonization. Failure of the Queen Charlotte Islands to conform to an area–richness trend suggests that not all resident species have been found.