Metapopulation vicariance explains old endemics on young volcanic islands

Terrestrial plants and animals on oceanic islands occupy zones of volcanism found at intraplate localities and along island arcs at subduction zones. The organisms often survive as metapopulations, or populations of separate sub‐populations connected by dispersal. Although the individual islands and their local subpopulations are ephemeral and unstable, the ecosystem dynamism enables metapopulations to persist in a region, more or less in situ, for periods of up to tens of millions of years. As well as surviving on systems of young volcanic islands, metapopulations can also evolve there; tectonic changes can break up widespread insular metapopulations and produce endemics restricted to fewer islands or even a single island. These processes explain the presence of old endemic clades on young islands, which is often reported in molecular clock studies, and the many distribution patterns in island life that are spatially correlated with tectonic features. Metapopulations can be ruptured by sea floor subsidence, and this occurs with volcanic loading in zones of active volcanism and with sea floor cooling following its production at mid‐ocean ridges. Metapopulation vicariance will also result if an active zone of volcanism is rifted apart. This can be caused by the migration of an arc (by slab rollback) away from a continent or from another subduction zone, by the offset of an arc at transform faults and by sea floor spreading at mid‐ocean ridges. These mechanisms are illustrated with examples from islands in the Caribbean and the Pacific. Endemism on oceanic islands has usually been attributed to chance, long‐distance dispersal, but the processes discussed here will generate endemism on young volcanic islands by vicariance.     

Mitochondrial DNA evolution and population history of the Tenerife skink Chalcides viridanus

Recent studies of island lizards have suggested that historical vicariance as a result of volcanism may have played an important role in shaping patterns of within-island genetic diversity. The skink, Chalcides viridanus, shows variation in morphology within the volcanic island of Tenerife. Two mitochondrial DNA (mtDNA) fragments (from the 12S and 16S rRNA regions) were sequenced in individuals from 17 sites to evaluate the relationship between current phylogeography and the geological history of the island. Three main clades were detected. The two most basal clades were restricted to areas representing the ancient precursor islands of Teno and Anaga in the northwest and northeast of Tenerife, respectively. The third clade showed a widespread geographical distribution and provided evidence of a recent rapid expansion after a bottleneck. Within-island cladogenesis appears to have taken place during a recent period of volcanic activity and long after the ancient islands had been united by the eruptions that led to the formation of the Canadas edifice. Evidence of similar biogeographical histories are found in other species in the Canary archipelago, supporting the volcanism scenario as a potentially widespread cause of within-island differentiation in reptiles.     

Phylogeography of the prehensile-tailed skink Corucia zebrata on the Solomon Archipelago

The biogeography of islands is often strongly influenced by prior geological events. Corucia zebrata (Squamata: Scincidae) is endemic to the geologically complex Solomon Archipelago in Northern Melanesia. We examined the level of divergence for different island populations of C. zebrata and discussed these patterns in light of Pleistocene land bridges, island isolation, and island age. Corucia zebrata was sampled from 14 locations across the Solomon Archipelago and sequenced at two mitochondrial genes (ND2 and ND4; 1697 bp in total) and four nuclear loci (rhodopsin, an unknown intron, AKAP9, and PTPN12). Measures of genetic divergence, analyses of genetic variation, and Bayesian phylogenetic inference were used and the data assessed in light of geological information. Populations of C. zebrata on separate islands were found to be genetically different from each other, with reciprocal monophyly on mitochondrial DNA. Populations on islands previously connected by Pleistocene land bridges were marginally less divergent from each other than from populations on other nearby but isolated islands. There are indications that C. zebrata has radiated across the eastern islands of the archipelago within the last 1-4 million years. Nuclear loci were not sufficiently informative to yield further information about the phylogeography of C. zebrata on the Solomon Archipelago. Analyses of the mitochondrial data suggest that dispersal between islands has been very limited and that there are barriers to gene flow within the major islands. Islands that have been isolated during the Pleistocene glacial cycles are somewhat divergent in their mitochondrial genotypes, however, isolation by distance (IBD) and recent colonization of isolated but geologically younger islands appear to have had stronger effects on the phylogeography of C. zebrata than the Pleistocene glacial cycles. This contrasts with patterns reported for avian taxa, and highlights the fact that biogeographic regions for island species cannot be directly extrapolated among taxa of differing dispersal ability.     

The dynamics of genetic and morphological variation on volcanic islands

Oceanic archipelagos of volcanic origin have been important in the study of evolution because they provide repeated natural experiments allowing rigorous tests of evolutionary hypotheses. Ongoing volcanism on these islands may, however, affect the evolutionary diversification of species. Analysis of population structure and phylogeographic patterns in island populations can provide insight into evolutionary dynamics on volcanic islands. We analysed genetic and morphological variation in the gecko Tarentola boettgeri on the island of Gran Canaria and compared it with Tarentola delalandii on Tenerife, a neighbouring volcanic island of similar age but distinctly different geological past. Intraspecific divergence of mitochondrial haplotypes indicates long-term persistence of Tarentola on each island, with a phylogeographic signal left by older volcanic events. More recent volcanic eruptions (approximately 0.2 million years ago on Tenerife, approximately 2.2 million years ago on Gran Canaria) have left a signature of population expansion in the population genetic structure, the strength of which depends on the time since the last major volcanic eruption on each island. While these stochastic events have left traces in morphological variation in Tenerife, in Gran Canaria geographical variation was solely associated with environmental variables. This suggests that historically caused patterns in morphology may be overwritten by natural selection within 2 million years.     

Phylogeographical footprints of the Strait of Gibraltar and Quaternary climatic fluctuations in the western Mediterranean: a case study with the greater white-toothed shrew, Crocidura russula (Mammalia: Soricidae)

We used mitochondrial cyt b sequences to investigate the phylogenetic relationships of Crocidura russula (sensu lato) populations across the Strait of Gibraltar, western Europe, Maghreb, and the Mediterranean and Atlantic islands. This revealed very low genetic divergence between European and Moroccan populations. The application of a molecular clock previously calibrated for shrews suggested that the separation of European from Moroccan lineages occurred less than 60 000 bp, which is at least 5 million years (Myr) after the reopening of the Strait of Gibraltar. This means that an overwater dispersal event was responsible for the observed phylogeographical structure. In contrast, genetic analyses revealed that Moroccan populations were highly distinct from Tunisian ones. According to the molecular clock, these populations separated about 2.2 million years ago (Ma), a time marked by sharp alternations of dry and humid climates in the Maghreb. The populations of the Mediterranean islands Ibiza, Pantelleria, and Sardinia were founded from Tunisian populations by overwater dispersal. In conclusion, overwater dispersal across the Strait of Gibraltar, probably assisted by humans, is possible for small terrestrial vertebrates. Moreover, as in Europe, Quaternary climatic fluctuations had a major effect on the phylogeographical structure of the Maghreb biota.     

Testing phylogeographic predictions on an active volcanic island: Brachyderes rugatus (Coleoptera: Curculionidae) on La Palma (Canary Islands)

Volcanic islands with well-characterized geological histories can provide ideal templates for generating and testing phylogeographic predictions. Many studies have sought to utilize these to investigate patterns of colonization and speciation within groups of closely related species across a number of islands. Here we focus attention within a single volcanic island with a well-characterized geological history to develop and test phylogeographic predictions. We develop phylogeographic predictions within the island of La Palma of the Canary Islands and test these using 69 haplotypes from 570 base pairs of mitochondrial DNA cytochrome oxidase II sequence data for 138 individuals of Brachyderes rugatus rugatus, a local endemic subspecies of curculionid beetle occurring throughout the island in the forests of Pinus canariensis. Although geological data do provide some explanatory power for the phylogeographic patterns found, our network-based analyses reveal a more complicated phylogeographic history than initial predictions generated from data on the geological history of the island. Reciprocal illumination of geological and phylogeographic history is also demonstrated with previous geological speculation gaining phylogeographic corroboration from our analyses.     

Phylogeography of marine mutualists: parallel patterns of genetic structure between obligate goby and shrimp partners

The survival of many organisms depends on interspecific, mutualistic interactions. Hence, it is important to assess the phylogeography of multiple mutualistic species simultaneously to gain insight into how their metapopulations persist, spread and recover from disturbance. Consequently, we sequenced mitochondrial DNA (cytochrome b) from a gobiid fish (Ctenogobiops feroculus) and its mutualistic partner, an alpheid snapping shrimp (Alpheus djeddensis). These obligate mutualists are common in shallow coral reef lagoons, and we collected individuals from 11 Indo-Pacific islands that were likely exposed to different disturbance histories due to sea level fluctuations associated with glaciation events [geological studies indicate that eastern oceanic islands (Cook Islands, French Polynesia) were more disturbed than western continental plate islands that have deeper, more extensive lagoons (Okinawa, Fiji)]. Both phylogenetic and population genetic analyses indicated that gobies from Okinawa and Fiji were genetically distinct both from each other and those in the Cook Islands and French Polynesia. Shrimp, by contrast, grouped into two distinct genetic groups that showed no geographic structure. Restricting the analysis to one clade of shrimp, which contained two-thirds of the individuals, showed that these shrimp had very similar phylogeographical structure to the gobies. Mismatch distributions demonstrated that both gobies and shrimp experienced a recent, rapid population expansion into French Polynesia. Finally, nucleotide and haplotype diversities per sample location were significantly correlated between mutualists and tended to be higher on western continental than eastern oceanic islands. Our results suggest that these mutualists recovered from a major disturbance by colonizing South Pacific islands in synchrony.     

Stand Biomass and Tree Mortality from Permanent Forest Plots on Krakatau,Indonesia, 1989–19951

Forest closure on the three original Krakatau Islands (Panjang, Rakata, and Sertung) took place ca 1930, about 50 yr after the apparent sterilization of the islands due to volcanic eruptions. Two permanent forest plots were established on each of these islands in 1989. A full enumeration of these plots, of two additional Rakata plots, and of two “mainland”plots from the Ujung Kulon National Park, West Java, was undertaken in 1992. These data provide the first estimates of aboveground biomass from Krakatau. The values reported for Krakatau are below the local West Javan figures, with considerable variation occurring within the islands. In 1992, the fourth Krakatau island, Anak Krakatau, began an eruption sequence which continued through the study period, depositing ash on the study sites of Panjang and Sertung. A further partial survey of these plots in 1994–1995 revealed a significant increase in mortality since the volcanic activity recommenced, with an increase in deaths of larger stems. Although Rakata has not been influenced directly by volcanism, three sites surveyed on Rakata in 1994–1995 have experienced increased tree mortality, in two cases as a consequence of storm damage and in particular, of lightning strikes. Stand dynamics on Krakatau thus appear to be strongly influenced by episodic environmental disturbance with varying degrees of dependence on the volcanic activity of Anak Krakatau.     

Colonization history and population genetics of the color-polymorphic hawaiian happy-face spider theridion grallator (araneae, theridiidae)

Past geological and climatological processes shape extant biodiversity. In the Hawaiian Islands, these processes have provided the physical environment for a number of extensive adaptive radiations. Yet, single species that occur throughout the islands provide some of the best cases for understanding how species respond to the shifting dynamics of the islands in the context of colonization history and associated demographic and adaptive shifts. Here, we focus on the Hawaiian happy-face spider, a single color-polymorphic species, and use mitochondrial and nuclear allozyme markers to examine (1) how the mosaic formation of the landscape has dictated population structure, and (2) how cycles of expansion and contraction of the habitat matrix have been associated with demographic shifts, including a "quantum shift" in the genetic basis of the color polymorphism. The results show a marked structure among populations consistent with the age progression of the islands. The finding of low genetic diversity at the youngest site coupled with the very high diversity of haplotypes on the slightly older substrates that are highly dissected by recent volcanism suggests that the mosaic structure of the landscape may play an important role in allowing differentiation of the adaptive color polymorphism.     

Palaeo‐islands as refugia and sources of genetic diversity within volcanic archipelagos: the case of the widespread endemic Canarina canariensis (Campanulaceae)

Geographical isolation by oceanic barriers and climatic stability has been postulated as some of the main factors driving diversification within volcanic archipelagos. However, few studies have focused on the effect that catastrophic volcanic events have had on patterns of within‐island differentiation in geological time. This study employed data from the chloroplast (cpDNA haplotypes) and the nuclear (AFLPs) genomes to examine the patterns of genetic variation in Canarina canariensis, an iconic plant species associated with the endemic laurel forest of the Canary Islands. We found a strong geographical population structure, with a first divergence around 0.8 Ma that has Tenerife as its central axis and divides Canarian populations into eastern and western clades. Genetic diversity was greatest in the geologically stable ‘palaeo‐islands’ of Anaga, Teno and Roque del Conde; these areas were also inferred as the ancestral location of migrant alleles towards other disturbed areas within Tenerife or the nearby islands using a Bayesian approach to phylogeographical clustering. Oceanic barriers, in contrast, appear to have played a lesser role in structuring genetic variation, with intra‐island levels of genetic diversity larger than those between‐islands. We argue that volcanic eruptions and landslides after the merging of the palaeo‐islands 3.5 Ma played key roles in generating genetic boundaries within Tenerife, with the palaeo‐islands acting as refugia against extinction, and as cradles and sources of genetic diversity to other areas within the archipelago.     

Mediterranean populations of the lesser white-toothed shrew (Crocidura suaveolens group): an unexpected puzzle of Pleistocene survivors and prehistoric introductions

An earlier study revealed the strong phylogeographical structure of the lesser white-toothed shrew (Crocidura suaveolens group) within the northern Palaearctic. Here, we aim to reconstruct the colonization history of Mediterranean islands and to clarify the biogeography and phylogeographical relationships of the poorly documented Middle East region with the northern Palaearctic. We performed analyses on 998-bp-long haplotypes of the mitochondrial cytochrome b gene of 143 samples collected around the Mediterranean basin, including islands and the Middle East. The analyses suggest that the Cypriot shrew belongs to the rare group of relict insular Pleistocene mammal taxa that have survived to the present day. In contrast, the Cretan, Corsican and Menorcan populations were independently introduced from the Middle East during the Holocene. The phylogeographical structure of this temperate Palaearctic species within the Middle East appears to be complex and rich in diversity, probably reflecting fragmentation of the area by numerous mountain chains. Four deeply divergent clades of the C. suaveolens group occur in the area, meaning that a hypothetical contact zone remains to be located in central western Iran.     

Glacial bottleneck and postglacial recolonization of a seed parasitic weevil, Curculio hilgendorfi, inferred from mitochondrial DNA variation

Climatic changes during glacial periods have had a major influence on the recent evolutionary history of living organisms, even in the warm temperate zone. We investigated phylogeographical patterns of a weevil Curculio hilgendorfi (Curculionidae), a host-specific seed predator of Castanopsis (Fagaceae) growing in the broadleaved evergreen forests in Japan. We examined 2709 bp of mitochondrial DNA for 204 individuals collected from 62 populations of the weevil. Four major haplogroups were detected, in southwestern and northeastern parts of the main islands and in central and southern parts of the Ryukyu Islands. The demographic population expansion was detected for the two groups in the main islands but not for the Ryukyu groups. The beginning time of the expansion was dated to 39 000–59 000 years ago, which is consistent with the end of the last glacial period. Our data also demonstrated that the southwestern population of the main islands has experienced a more severe bottleneck and more rapid population growth after glacial ages than the northeastern population. At least three refugial areas in the main islands were likely to have existed during the last glacial periods, one of which had not previously been recognized by analyses of intraspecific chloroplast DNA variation of several plant species growing in the broadleaved evergreen forests. Our results represent the first phylogeographical and population demographic analysis of an insect species associated with the broadleaved evergreen forests in Japan, and reveal more detailed postglacial history of the forests.     

Mediterranean island biogeography: Analysis of fern species distribution in the system of islets around Sicily

Abstract

The aim of this article was to test the way in which geographical factors influence island floras in the Mediterranean basin, using ferns as target organisms, and the islands surrounding Sicily as location. A matrix with presence/absence data concerning fern taxa in the 16 islands studied was compiled. Cluster analysis, principal co-ordinates analysis (PCoA), principal components analysis (PCA) and a Bayesian analysis were performed. For each island, the total number of fern taxa was regressed against three factors: island area, island elevation and isolation. All the analyses pointed to affinities between islands according to their different geological composition, independently from their geographic position. A clear positive island species/area relationship (ISAR) was shown only for the volcanic islands. The island species/(area×elevation) relationship (ISAER), on the contrary, was unsatisfactory. The main features of interest are the following: (1) the clear division of the islands into two groups, volcanic vs. sedimentary; (2) the floristic richness of the volcanic compared to sedimentary islands and (3) the uniqueness of the pteridophyte flora of Pantelleria. This seems to demonstrate that the lower number of taxa in the islands farthest away from the “mainland” (Sicily, Tunisia) is not due to isolation, but due to another factor, probably habitat availability.     

Species status and phylogeography of two closely related Coptolabrus species (Coleoptera: Carabidae) in South Korea inferred from mitochondrial and nuclear gene sequences

We investigated the species status and intraspecific phylogeography in South Korea of two ground beetle species, Coptolabrus jankowskii and Coptolabrus smaragdinus (Coleoptera: Carabidae), using statistical parsimony networks and nested clade analyses based on sequences from the mitochondrial cytochrome oxidase subunit I (COI) and nuclear phosphoenolpyruvate carboxykinase (PepCK) and wingless (Wg) genes. Although traditional parsimony tree construction generally failed to resolve interspecific relationships and construct biologically meaningful genealogies, analysis using statistical parsimony networks yielded statistically significant inter- and intraspecific genealogical structures. We found that although these two species represent a notable case of trans-species polymorphisms in both mitochondrial and nuclear gene sequences, their status as separate species was evidenced by the nonrandom association between species and nested clades at various nesting levels. The exceptional occurrence of shared identical or very similar COI sequences was considered to be the result of introgressive hybridization. In addition, range expansion and fragmentation events across the Korean Peninsula and adjacent islands were inferred from nested clade phylogeographical analyses. The COI gene revealed the geographical divergence of major eastern and western clades and historical biogeographical events within each major clade, whereas the nuclear PepCK gene, which did not reveal corresponding east-west clades, indicated past fragmentation and range expansion across wide areas that may have been the result of older biogeographical events. Thus, phylogeographical inferences drawn from analyses of mitochondrial and nuclear genes can reveal different and potentially complementary information about phylogeographical processes.     

Eastward Ho: phylogeographical perspectives on colonization of hosts and parasites across the Beringian nexus

The response of Arctic organisms and their parasites to dramatic fluctuations in climate during the Pleistocene has direct implications for predicting the impact of current climate change in the North. An increasing number of phylogeographical studies in the Arctic have laid a framework for testing hypotheses concerning the impact of shifting environmental conditions on transcontinental movement. We review 35 phylogeographical studies of trans-Beringian terrestrial and freshwater taxa, both hosts and parasites, to identify generalized patterns regarding the number, direction and timing of trans-continental colonizations. We found that colonization across Beringia was primarily from Asia to North America, with many events occurring in the Quaternary period. The 35 molecular studies of trans-Beringian organisms we examined focused primarily on the role of glacial cycles and refugia in promoting diversification. We address the value of establishing testable hypotheses related to high-latitude biogeography. We then discuss future prospects in Beringia related to coalescent theory, palaeoecology, ancient DNA and synthetic studies of arctic host–parasite assemblages highlighting their cryptic diversity, biogeography and response to climate variation.     

Recent volcanism and mitochondrial DNA structuring in the lizard Gallotia atlantica from the island of Lanzarote

The phylogeography of the lacertid lizard Gallotia atlantica from the small volcanic island of Lanzarote (Canary Islands) was analysed based on 1075 bp of mitochondrial DNA (mtDNA) sequence (partial cytochrome b and ND2) for 157 individuals from 27 sites (including three sites from neighbouring islets). Levels of sequence divergence were generally low, with the most distant haplotypes separated by only 14 mutational steps. MtDNA divergence appears to coincide with formation of the middle Pleistocene lowland that united formerly separate ancient islands to form the current island of Lanzarote, allowing rejection of a two-island model of phylogeographical structure. There was evidence of large-scale population expansion after island unification, consistent with the colonization of new areas. A nested clade phylogeographical analysis (NCPA) revealed significant phylogeographical structuring. Two-step and higher-level clades each had disjunct distributions, being found to the east and west of a common area with a north-south orientation that extends between coasts in the centre-east of the island (El Jable). Other clades were almost entirely restricted to the El Jable region alone. Bayesian Markov chain Monte Carlo analyses were used to separate ongoing gene flow from historical associations. These supported the NCPA by indicating recent (75,000-150,000 years ago) east-west vicariance across the El Jable region. Lava flows covered El Jable and other parts of the central lowland at this time and likely led to population extinctions and temporary dispersal barriers, although present-day evidence suggests some populations would have survived in small refugia. Expansion of the latter appears to explain the presence of a clade located between the eastern and western components of the disjunct clades. Direct relationships between mtDNA lineages and morphology were not found, although one of two morphological forms on the island has a disjunct distribution that is broadly concordant with east-west components of the phylogeographical pattern. This work demonstrates how recent volcanic activity can cause population fragmentation and thus shape genetic diversity on microgeographical scales.     

Colonization and diversification of Galápagos terrestrial fauna: a phylogenetic and biogeographical synthesis

Remote oceanic islands have long been recognized as natural models for the study of evolutionary processes involved in diversification. Their remoteness provides opportunities for isolation and divergence of populations, which make islands remarkable settings for the study of diversification. Groups of islands may share a relatively similar geological history and comparable climate, but their inhabitants experience subtly different environments and have distinct evolutionary histories, offering the potential for comparative studies. A range of organisms have colonized the Galápagos Islands, and various lineages have radiated throughout the archipelago to form unique assemblages. This review pays particular attention to molecular phylogenetic studies of Galápagos terrestrial fauna. We find that most of the Galápagos terrestrial fauna have diversified in parallel to the geological formation of the islands. Lineages have occasionally diversified within islands, and the clearest cases occur in taxa with very low vagility and on large islands with diverse habitats. Ecology and habitat specialization appear to be critical in speciation both within and between islands. Although the number of phylogenetic studies is continuously increasing, studies of natural history, ecology, evolution and behaviour are essential to completely reveal how diversification proceeded on these islands.     

Dispersal limitations and historical factors determine the biogeography of specialized terrestrial protists

Recent studies show that soil eukaryotic diversity is immense and dominated by micro‐organisms. However, it is unclear to what extent the processes that shape the distribution of diversity in plants and animals also apply to micro‐organisms. Major diversification events in multicellular organisms have often been attributed to long‐term climatic and geological processes, but the impact of such processes on protist diversity has received much less attention as their distribution has often been believed to be largely cosmopolitan. Here, we quantified phylogeographical patterns in Hyalosphenia papilio, a large testate amoeba restricted to Holarctic Sphagnum‐dominated peatlands, to test if the current distribution of its genetic diversity can be explained by historical factors or by the current distribution of suitable habitats. Phylogenetic diversity was higher in Western North America, corresponding to the inferred geographical origin of the H. papilio complex, and was lower in Eurasia despite extensive suitable habitats. These results suggest that patterns of phylogenetic diversity and distribution can be explained by the history of Holarctic Sphagnum peatland range expansions and contractions in response to Quaternary glaciations that promoted cladogenetic range evolution, rather than the contemporary distribution of suitable habitats. Species distributions were positively correlated with climatic niche breadth, suggesting that climatic tolerance is key to dispersal ability in H. papilio. This implies that, at least for large and specialized terrestrial micro‐organisms, propagule dispersal is slow enough that historical processes may contribute to their diversification and phylogeographical patterns and may partly explain their very high overall diversity.     

Adult habitat preferences, larval dispersal, and the comparative phylogeography of three Atlantic surgeonfishes (Teleostei: Acanthuridae)

Although many reef fishes of the tropical Atlantic are widely distributed, there are large discontinuities that may strongly influence phylogeographical patterns. The freshwater outflow of the Amazon basin is recognized as a major barrier that produces a break between Brazilian and Caribbean faunas. The vast oceanic distances between Brazil and the mid-Atlantic ridge islands represent another formidable barrier. To assess the relative importance of these barriers, we compared a fragment of the mitochondrial DNA (mtDNA) cytochrome b gene among populations of three species of Atlantic surgeonfishes: Acanthurus bahianus, A. chirurgus and A. coeruleus. These species have similar life histories but different adult habitat preferences. The mtDNA data show no population structure between Brazil and the mid-Atlantic islands, indicating that this oceanic barrier is readily traversed by the pelagic larval stage of all three surgeonfishes, which spend approximately 45-70 days in the pelagic environment. The Amazon is a strong barrier to dispersal of A. bahianus (d = 0.024, phiST = 0.724), a modest barrier for A. coeruleus (phiST = 0.356), and has no discernible effect as a barrier for A. chirurgus. The later species has been collected on soft bottoms with sponge habitats under the Amazon outflow, indicating that relaxed adult habitat requirements enable it to readily cross that barrier. A limited ability to use soft bottom habitats may also explain the low (but significant) population structure in A. coeruleus. In contrast, A. bahianus has not been collected over deep sponge bottoms, and rarely settles outside shallow reefs. Overall, adult habitat preferences seem to be the factor that differentiates phylogeographical patterns in these reef-associated species.     

Pleistocene glaciation leaves deep signature on the freshwater crab Aegla alacalufi in Chilean Patagonia

Quaternary glacial cycles have played an important role in shaping the biodiversity in temperate regions. This is well documented in Northern Hemisphere, but much less understood for Southern Hemisphere. We used mitochondrial DNA and nuclear elongation factor 1α intron sequences to examine the Pleistocene glacial impacts on the phylogeographical pattern of the freshwater crab Aegla alacalufi in Chilean Patagonia. Phylogenetic analyses, which separated the glaciated populations on eastern continent into a north group (seven populations) and a south group (one population), revealed a shallow phylogenetic structure in the north group but a deep one in the non-glaciated populations on western islands, indicating the significant influence of glaciation on these populations. Phylogenies also identified the Yaldad population on Chiloé Island as a potentially unrecognized new species. The non-glaciated populations showed higher among population genetic divergence than the glaciated ones, but lower population genetic diversity was not detected in the latter. The two glaciated groups, which diverged from the non-glaciated populations at ~96 800–29 500 years ago and ~104 200–73 800 years ago, respectively, seem to have different glacial refugia. Unexpectedly, the non-glaciated islands did not serve as refugia for them. Demographic expansion was detected in the glaciated north group, with a constant population increase after the last glacial maximum. Nested clade analyses suggest a possible colonization from western islands to eastern continent. After arriving on the continent and surviving the last glacial period there, populations likely have expanded from high to low altitude, following the flood of melting ice. Aegla alacalufi genetic diversity has been primarily affected by Pleistocene glaciation and minimally by drainage isolation.