Coastal Refugia Research Articles

Wildlife following people: A multidisciplinary assessment of the ancient colonization of the Mediterranean Basin by a long‐lived raptor

Abstract Modern humans widely shaped present ecosystems through intentional and unintentional geographical redistribution of wildlife, both in historical and pre‐historical times. However, the patterns of ancient human‐mediated indirect changes in wildlife range are largely unknown, and the mechanisms behind them remain obscure. We used a multidisciplinary approach to (a) reconstruct the process of colonization of the Mediterranean Basin by a long‐lived bird of prey, the Bonelli's eagle (Aquila fasciata), and (b) test the hypothesis that this colonization was unintentionally favoured by anatomically modern humans through a release of competition by dominant species, primarily golden eagles (A. chrysaetos). The fossil record of Bonelli's eagles in the Mediterranean Basin was restricted to the last c. 50 ky. This timing matches the period of modern human presence in Europe. Distribution modelling showed that Bonelli's eagles find more suitable conditions in interglacial periods, while glacial maxima are largely unfavourable unless in coastal refugia. In agreement with this, all Bonelli's eagle's fossils were found in coastal areas, and demographic inference from genetic data revealed a drop in the effective population size by around the last glacial maximum. In today's communities, we found a strongly asymmetric competitive relationship between (subordinate) Bonelli's and (dominant) golden eagles, with the former occupying far more humanized areas than the latter both at the landscape scale and the local (i.e. nesting cliff) scale. Moreover, the nesting habitat overlap analysis indicated that, in the absence of the other species, a notably higher population of Bonelli's eagle, but not of golden eagle, could be expected. Our findings are consistent with the human‐mediated competitor release hypothesis, by which anatomically modern humans could have unintentionally favoured the large‐scale colonization by Bonelli's eagles of a previously competitively hostile Mediterranean Basin. Reconstructing the role of ancient humans in shaping present ecosystems may help to understand the historical, current and future population trajectories of competing species of conservation concern under the ongoing scenario of global environmental change. It also illustrates how human‐mediated apparent competition may promote large‐scale redistribution and colonization of wildlife, including long‐lived species. Read the free Plain Language Summary for this article on the Journal blog.

Improved eDNA assay evidences further refugia for critically endangered smalltooth sawfish (Pristis pectinata) in Mexico

Sawfishes are considered one of the most threatened marine fish taxa globally, with major contractions in their geographic range documented over the last 50 years across all oceans. Two sawfish species used to be found in Mexico, but a historical lack of research and first-hand baseline information recently raised concerns about the fate of both species there. Recent Local Ecological Knowledge (LEK) and environmental DNA (eDNA) research has shown that: a) Pristis pectinata and P. pristis were formerly abundant and widely distributed in the Mexican Atlantic coast, and the latter also in the Mexican Pacific coast, and b) sawfishes (at least P. pectinata) are still found in some coastal refugia in central Veracruz state and Campeche state on the Atlantic coast. In the present research we developed improved primers that allow for better species ID resolution for Pristis pectinata and P. pristis through successful sequencing of eDNA samples by amplifying a 270 bp fragment of the CO1 gene. Our results based on the analysis of 305 eDNA samples showed an improved positive species identification rate through sequencing, demonstrate that P. pectinata occurs in other coastal refugia along the Yucatán Peninsula in the Mexican Caribbean, and suggest this species might be extirpated from southern Veracruz, while no P. pristis eDNA has been found along the Atlantic coast.

Postglacial phylogeography, admixture, and evolution of red spruce (Picea rubens Sarg.) in Eastern North America.

Climate change is a major evolutionary force that can affect the structure of forest ecosystems worldwide. Red spruce (Picea rubens Sarg.) has recently faced a considerable decline in the Southern Appalachians due to rapid environmental change, which includes historical land use, and atmospheric pollution. In the northern part of its range, red spruce is sympatric with closely related black spruce (Picea mariana (Mill.) B.S.P.), where introgressive hybridization commonly occurs. We investigated range-wide population genetic diversity and structure and inferred postglacial migration patterns and evolution of red spruce using nuclear microsatellites. Moderate genetic diversity and differentiation were observed in red spruce. Genetic distance, maximum likelihood and Bayesian analyses identified two distinct population clusters: southern glacial populations, and the evolutionarily younger northern populations. Approximate Bayesian computation suggests that patterns of admixture are the result of divergence of red spruce and black spruce from a common ancestor and then introgressive hybridization during post-glacial migration. Genetic diversity, effective population size (Ne) and genetic differentiation were higher in the northern than in the southern populations. Our results along with previously available fossil data suggest that Picea rubens and Picea mariana occupied separate southern refugia during the last glaciation. After initial expansion in the early Holocene, these two species faced a period of recession and formed a secondary coastal refugium, where introgressive hybridization occurred, and then both species migrated northward. As a result, various levels of black spruce alleles are present in the sympatric red spruce populations. Allopatric populations of P. rubens and P. mariana have many species-specific alleles and much fewer alleles from common ancestry. The pure southern red spruce populations may become critically endangered under projected climate change conditions as their ecological niche may disappear.

Ice and ocean constraints on early human migrations into North America along the Pacific coast

Founding populations of the first Americans likely occupied parts of Beringia during the Last Glacial Maximum (LGM). The timing, pathways, and modes of their southward transit remain unknown, but blockage of the interior route by North American ice sheets between ~26 and 14 cal kyr BP (ka) favors a coastal route during this period. Using models and paleoceanographic data from the North Pacific,we identify climatically favorable intervals when humans could have plausibly traversed the Cordilleran coastal corridor during the terminal Pleistocene. Model simulations suggest that northward coastal currents strengthened during the LGM and at times of enhanced freshwater input, making southward transit by boat more difficult. Repeated Cordilleran glacial-calving events would have further challenged coastal transit on land and at sea. Following these events, ice-free coastal areas opened and seasonal sea ice was present along the Alaskan margin until at least 15 ka. Given evidence for humans south of the ice sheets by 16 ka and possibly earlier, we posit that early people may have taken advantage of winter sea ice that connected islands and coastal refugia. Marine ice-edge habitats offer a rich food supply and traversing coastal sea ice could have mitigated the difficulty of traveling southward in watercraft or on land over glaciers.We identify 24.5 to 22 ka and 16.4 to 14.8 ka as environmentally favorable time periods for coastal migration, when climate conditions provided both winter sea ice and ice-free summer conditions that facilitated year-round marine resource diversity and multiple modes of mobility along theNorth Pacific coast.

Whole genome population structure of North Atlantic kelp confirms high-latitude glacial refugia.

Coastal refugia during the Last Glacial Maximum (~21,000 years ago) have been hypothesized at high latitudes in the North Atlantic, suggesting marine populations persisted through cycles of glaciation and are potentially adapted to local environments. Here, whole-genome sequencing was used to test whether North Atlantic marine coastal populations of the kelp Alaria esculenta survived in the area of southwestern Greenland during the Last Glacial Maximum. We present the first annotated genome for A. esculenta and call variant positions in 54 individuals from populations in Atlantic Canada, Greenland, Faroe Islands, Norway and Ireland. Differentiation across populations was reflected in ~1.9 million single nucleotide polymorphisms, which further revealed mixed ancestry in the Faroe Islands individuals between putative Greenlandic and European lineages. Time-calibrated organellar phylogenies suggested Greenlandic populations were established during the last interglacial period more than 100,000 years ago, and that the Faroe Islands population was probably established following the Last Glacial Maximum. Patterns in population statistics, including nucleotide diversity, minor allele frequencies, heterozygosity and linkage disequilibrium decay, nonetheless suggested glaciation reduced Canadian Atlantic and Greenlandic populations to small effective sizes during the most recent glaciation. Functional differentiation was further reflected in exon read coverage, which revealed expansions unique to Greenland in 337 exons representing 162 genes, and a modest degree of exon loss (103 exons from 56 genes). Altogether, our genomic results provide strong evidence that A. esculenta populations were resilient to past climatic fluctuations related to glaciations and that high-latitude populations are potentially already adapted to local conditions as a result.

Ecosystem engineer morphological traits and taxon identity shape biodiversity across the euphotic-mesophotic transition.

The euphotic-mesophotic transition is characterized by dramatic changes in environmental conditions, which can significantly alter the functioning of ecosystem engineers and the structure of their associated communities. However, the drivers of biodiversity change across the euphotic-mesophotic transition remain unclear. Here, we investigated the mechanisms affecting the biodiversity-supporting potential of free-living red coralline algae-globally important habitat creators-towards mesophotic depths. Across a 73 m depth gradient, we observed a general decline in macrofaunal biodiversity (fauna abundance, taxon richness and alpha diversity), but an increase in beta-diversity (i.e. variation between assemblages) at the deepest site (86 m depth, where light levels were less than 1% surface irradiance). We identified a gradient in abundance decline rather than distinct ecological shifts, driven by a complex interaction between declining light availability, declining size of the coralline algal host individuals and a changing host taxonomy. However, despite abundance declines, high between-assemblage variability at deeper depths allowed biodiversity-supporting potential to be maintained, highlighting their importance as coastal refugia.

Genomic footprints of Quaternary colonization and population expansion in the Patagonian‐Fuegian region rules out a separate southern refugium in Tierra del Fuego

AbstractAimDuring the glacial phases, the Andean slopes of southern South America were covered by ice sheets, causing the isolation of many species in glacial refugia. This work aimed to test the number and putative location of glacial refugia in southern South America and establish the sources, timing and direction of post‐glacial colonization events.LocationSouthern South America, Patagonia.TaxonOlive mouse (Abrothrix olivacea).MethodsUsing exome capture, we genotyped 7339 SNPs in 172 specimens of A. olivacea collected at 21 localities distributed across the steppe, grasslands, and forests in the Patagonian‐Fuegian region and seven specimens from two other species of Abrothrix. Based on the effective migration rate, principal component analysis, and admixture composition, we first study the population structure of A. olivacea. The source and direction of colonization events were inferred using the directionality index and standard population genetic analyses. To study the role of the glacial phases, we performed a bayesian demographic analysis to infer divergences times and ancestral population sizes.ResultsPopulation genomic analyses based on SNPs of more than 7000 exons identified three Patagonian genetic clusters: Valdivian forests, continental Magellan forests, and a combination of continental open biomes and Tierra del Fuego. The directionality index supports a single source in northwestern forests, in line with a Valdivian forest refugium, from which the post‐glacial colonization may have started. The effective diversity rate, the onset of population expansions and divergence times also support a Valdivian forest refugium and rule out a separate southern refugium in Tierra del Fuego.Main conclusionsOur results support a coastal refugium located in or near the previously proposed Valdivian refugium and reject in situ differentiation in Tierra del Fuego. More in general, our demographical analysis reveals the role of the Last Glacial Maximum in affecting the population dynamics (i.e., contraction and expansion) of A. olivacea and in fostering the geographical diversification of populations.

Whole‐genome resequencing reveals persistence of forest‐associated mammals in Late Pleistocene refugia along North America’s North Pacific Coast

AbstractAimNumerous glacial refugia have been hypothesized along North America's North Pacific Coast that may have increased divergence of refugial taxa, leading to elevated endemism and subsequently clustered hybrid zones following deglaciation. The locations and community composition of these ice‐free areas remains controversial, but whole‐genome sequences now enable detailed analysis of the demographic and evolutionary histories of refugial taxa. Here, we use genomic data to test spatial and temporal processes of diversification among martens with respect to the Coastal Refugium Hypothesis, to understand the role of climate cycling in shaping diversity across complex landscapes.LocationNorth America and North Pacific Coast archipelagos.TaxonNorth American martens (Martes).MethodsShort‐read whole‐genome resequencing data were generated for 11 martens: four M. americana, four M. caurina, two hybrids, and one outgroup (Martes zibellina). Sampling was representative of known genetic clades within New World martens, including sampling within insular and continental hybrid zones and along the North Pacific Coast (five island populations). ADMIXTURE, F‐statistics, and D‐statistics (ABBA‐BABA) were used to identify introgression and infer directionality. Heterozygosity densities, estimated via PSMC, were used to characterize historical demography at and below the species level to infer refugial and colonization processes.ResultsForest‐associated Pacific martens (M. caurina) are divided into distinct insular and continental clades consistent with the Coastal Refugium Hypothesis. There was no evidence of introgression on islands that received historical translocations of American pine martens (M. americana), but introgression was detected in two active zones of secondary contact: one insular and one continental. Only early‐generational hybrids were identified across multiple hybrid zones, a pattern consistent with potential genetic swamping of M. caurina by M. americana.Main conclusionsDespite an incomplete fossil record, genomic evidence supports the persistence of forest‐associated martens, likely the insular Pacific marten lineage, along the western edges of the Alexander Archipelago during the Last Glacial Maximum. This discovery informs our understanding of refugial paleoenvironments, critical to interpreting refugial timing, duration, and community composition. Genomic reevaluations of other taxa along North America's North Pacific Coast may yield new and deeper perspectives on the history of refugial forest communities and the role of dynamic climate shifts in shaping high‐latitude diversity across complex insular landscapes.

A new pollen sequence from southern Iberia suggesting coastal Pleistocene phytodiversity hotspot

This paper presents a palynological study of the archeological layers from the Neanderthal site Abrigo 3 del Complejo del Humo, in southern Spain (Málaga), with the aim of reconstructing the environmental conditions in the vicinity of this hominin site. The Upper Pleistocene vegetation and its variability are described, revealing a high diversity of thermophilous plant taxa throughout the cold dry phases, together with a long-term persistence of woody taxa, including Mediterranean, mesophytes, xerothermics and conifers. With the pollen records of Maytenus senegalensis as an outstanding finding, this study demonstrates the co-existence of temperate, Mediterranean and Ibero-Maghrebian angiosperms on the southern coastal plains of the Iberian Pleistocene where Neanderthals survived for a long time. It is therefore clear that Neanderthals and early Upper Paleolithic modern humans lived in a litoral refugium, which was a propitious environment for maintaining a high biodiversity, including potentially edible plant species. Besides, this coastal refugium offers broad possibilities for hunting, and interpopulational relationships through coastal platforms.

Late Pleistocene marine resources from the Bering Glacier Foreland and human coastal migration in the northern Gulf of Alaska region

AbstractRecent research on the Bering Glacier forelands in the northern Gulf of Alaska provides new insights into late Pleistocene/early Holocene shorelines, providing a favorable route for human migration as early as ~16,000 cal yr BP. This route included an irregular coastline with embayments and islands offering protection from the open ocean; edible marine invertebrates dating from 15,000 to 5,500 cal yr BP; and marine vertebrates dating as early as 16,000 cal yr BP. The latter included walrus (Odobenus rosmarus), bearded seal (Erignathus barbatus), and ringed seal (Phoca cf. hispida), all associated with pack ice conditions unlike those present today. While this ecosystem could have supported humans migrating along the coastline, and coastal refugia may have existed elsewhere in the region, coastal archaeological sites in the northern Gulf of Alaska and southwest Alaska are no older than ~9,500 cal yr BP. This suggests that the earliest sites have been eroded or destroyed, that the earliest migrants ignored available marine resources, and/or that these migrants did not use a coastal route. In contrast, the earliest archaeological sites in southeast Alaska date to ~12,500 cal yr BP, suggesting migration from interior Alaska to the coast somewhere east of the Copper River delta.

Thamnolia tundrae sp. nov., a cryptic species and putative glacial relict

AbstractThe lichen species of the genus Thamnolia, with their striking wormlike thalli and frequent occurrence in arctic and tundra environments, have often been debated with regard to the use of chemistry in lichen taxonomy. Phylogenetic studies have arrived at different conclusions as to the recognition of species in the genus, but in a recent study based on the analyses of six nuclear markers (genes or noncoding regions) of a worldwide sample of Thamnolia, we showed the existence of three well-supported lineages with two different chemistries and geographical distributions. Here, we present two analyses based on ITS and three markers, respectively, which were extended from the study mentioned above to include type specimens and additional Thamnolia strains and taxa. In these analyses the same three clades were retrieved. A putative DEAD-box helicase is used here for the first time as an informative phylogenetic marker to provide taxonomic resolution at species level. The distribution of morphological and chemical characters across the phylogeny was analyzed and it was concluded that three morphologically cryptic, but genetically well supported, species occur: T. vermicularis s. str., T. subuliformis s. str. and T. tundrae sp. nov. Thamnolia vermicularis s. str. contains individuals with uniform secondary chemistry (producing thamnolic acid) and a rather limited distribution in the European Alps, Tatra Mts and the Western Carpathians, a distribution which might result from glacial survival in an adjacent refugium/refugia. Thamnolia subuliformis s. str. is widely distributed in all hemispheres and the samples contain two chemotypes (either with thamnolic or squamatic acids). Thamnolia tundrae is described as new; it produces baeomycesic and squamatic acids, and has a distribution limited to the arctic tundra of Eurasia extending to the Aleutian Islands in North America. It may have survived the latest glaciation in coastal refugia near its present distribution. Thus, secondary chemistry alone is not suitable for characterizing species in Thamnolia, secondary chemistry and geographical origin are informative, and the ITS region can be confidently used for species recognition. Nomenclatural notes are given on several other names that have been used in Thamnolia.

Quaternary range-shift history of Japanese wingnut (Pterocarya rhoifolia) in the Japanese Archipelago evidenced from chloroplast DNA and ecological niche modeling

ABSTRACTBased on organelle DNA phylogeographic analyses and ecological niche modeling (ENM), we investigated the range-shift history of the Japanese wingnut (Pterocarya rhoifolia) during the Quaternary climatic oscillations with particular emphasis on the Last Glacial Maximum (LGM). Phylogeographic patterns of this species were determined using 376 individuals from 53 populations for chloroplast DNA sequencing of three spacers. Spatial analysis of molecular variance revealed that the current phylogeographic structure would be sculptured by multiple range shifts from each glacial refugium, which would have been repeated several times during the Quaternary climatic oscillations. High haplotype diversity and private haplotypes were detected in southwestern Japan, where wingnut is currently infrequent and found mainly in high mountains, whereas in northernmost Japan, haplotype diversity was low though this plant is quite common at present. According to ENM approach, during the LGM, the climatically suitable distribution areas were not recovered in northeastern Japan but in lowlands of southwestern Japan. Our combined results suggest that Japanese wingnut primarily persisted in the lowlands of southwestern Japan and coastlines below 36.5°N latitude during the LGM, having led to the postglacial range expansion from the refugia in each area of southwestern and lower latitudinal northeastern Japan, and to the wide-range recolonization from the southerly refugium/refugia to the north in northernmost Japan. The southwestern and coastal refugia have played a role in shaping the current haplotype diversity and phylogeographic structure, whereas some rear edge populations, harboring unique haplotypes, have been also maintained.

Phylogeographic evidence for two mesic refugia in a biodiversity hotspot.

Phylogeographic studies of flora in species-rich south-western Australia point to complex evolutionary histories, reflecting patterns of persistence and resilience to climatic changes during the Pleistocene. We asked whether coastal areas of the mid-west and south, as well as granite outcrops and inland ranges, have acted as major refugia within this region during Pleistocene climatic fluctuations by analysing phylogeographic patterns in the shrub Calothamnus quadrifidus R.Br. (Myrtaceae). We determined variation in chloroplast DNA data for 41 populations across the geographic range. Relationships and major clades were resolved using parsimony and Bayesian analyses. We tested for demographic and spatial expansion of the major clades and estimated clade divergence dates using an uncorrelated, lognormal relaxed clock based on two conservative chloroplast mutation rates. Two distinct phylogeographic clades were identified showing divergence during the Pleistocene, consistent with other phylogeographic studies of south-west Australian flora, emphasising the impact of climatic oscillations in driving divergence in this landscape. The southern clade was more diverse, having higher haplotype diversity and greater genetic structure, while the northern clade showed evidence of fluctuation in population size. Regions of high haplotype diversity with adjacent areas of low diversity observed in each clade indicated the locations of two coastal refugia: one on the south coast and another along the mid-west coast. This is the first evidence for major Pleistocene refugia using chloroplast genetic data in a common, widespread species from this region.

Morphological Analysis and Phytogeography of Native Calamagrostis (Poaceae) from British Columbia, Canada and Adjacent Regions

The taxonomically difficult and ecologically and phytogeographically important genus, Calamagrostis, was examined for British Columbia (BC). Morphological characters were analyzed by Principal Components Analysis (PCA) to characterize taxa and to aid in the development of a new key. Eight native species (Calamagrostis canadensis, C. lapponica, C. montanensis, C. nutkaensis, C. purpurascens, C. rubescens, C. sesquiflora, and C. stricta) are confirmed to occur in British Columbia, of which C. montanensis, C. nutkaensis, C. purpurascens, C. rubescens, and C. sesquiflora are reliably distinguishable. Comparison of species distribution to regional climatic and vegetation history suggests that Calamagrostis nutkaensis and C. sesquiflora likely survived in coastal refugia during late Wisconsin glaciations. Calamagrostis purpurascens likely persisted beyond the glacial limits or within nunataks and then spread into previously glaciated sites. Two interior continental species, C. montanensis and C. rubescens, probably spread north and west from the unglaciated zone south of the Cordilleran and Laurentide ice sheets. Calamagrostis lapponica likely persisted north of the ice sheets, and then spread southward into high-elevation sites in northern and eastern BC. Calamagrostis canadensis and C. stricta probably survived south and north of the ice sheets, and then spread into the previously glaciated terrain.

Phylogeography of a Holarctic nematode, Soboliphyme baturini, among mustelids: climate change, episodic colonization, and diversification in a complex host-parasite system

Phylogeography of Soboliphyme baturini, a nematode parasite in mustelids, is explored across Beringia. Sequences of the mitochondrial cytochrome c oxidase subunit I and nicotinamide adenine dinucleotide dehydrogenase subunit 4 genes were evaluated from 37 S. baturini, representing 19 localities throughout Alaska, Canada, and Siberia. A total of 30 haplotypes was recovered and maximum parsimony and Bayesian phylogenetic analyses support the recognition of a single species with a distribution extending from the Palearctic to the Nearctic. Within S. baturini, a host-specific partition in North America between Martes caurina and Martes americana was not identified. Instead, substantial geographic structure within S. baturini relates to the dynamic geological history of this northern region and especially the North Pacific Coast. Beringia and other coastal refugia along the western margin of North America played a large role during stadial maxima in the persistence and divergence of the parasite. Repeated events for biotic expansion and geographic colonization across the Bering Land Bridge and the Holarctic during glacial maxima in the Pleistocene appear to have facilitated at least two episodes of host-switching of this nematode among mustelids in populations now distributed in eastern Beringia.

Postglacial colonization and population genetic relationships in thePinus contortacomplex

To explore the evolutionary relationships among the subspecies of lodgepole pine ( Pinus contorta Dougl. ex Loud.) and the possible existence of a northern glacial refugium for Pinus contorta Dougl. ex Loud. subsp. latifolia Engelm. and a coastal refugium for Pinus contorta Dougl. ex Loud. subsp. contorta , 31 populations were analyzed using RAPD markers to determine whether a pattern of isolation by distance exists. A total of 52 markers from 10 primers were scored. A significant pattern of isolation by distance was detected when all 31 populations were analyzed together and for the 19 populations of subsp. latifolia. A phylogenetic tree constructed from all 31 populations does not show an unambiguous division of the subspecies of P. contorta into distinct clades. Our results provide no support for the separation of lodgepole pine into various subspecies and show that the subspecies of lodgepole pine have not been sufficiently isolated from one another for a large amount of genetic divergence to occur between them. For subsp. latifolia, the pattern of isolation by distance is consistent with the hypothesis of a single refugium south of the glacial maximum with population expansion following glacial retreat. The amount of genetic distance and the pattern of differentiation in populations of subsp. contorta suggest a more complex history associated with fluctuation in ocean levels and glacial margins.

Rapid divergence and postglacial colonization in western North American Steller's jays (Cyanocitta stelleri)

Post-Pleistocene avian colonization of deglaciated North America occurred from multiple refugia, including a coastal refugium in the northwest. The location of a Pacific Coastal refugium is controversial; however, multiple lines of evidence suggest that it was located near the Queen Charlotte Islands (also known as Haida Gwaii). The Queen Charlotte Islands contain a disproportionately large number of endemic plants and animals including the Steller's jay Cyanocitta stelleri carlottae. Using five highly variable microsatellite markers, we studied population structure among eight populations of Steller's jay (N = 150) from geographical areas representing three subspecies in western North America: C. s. carlottae, C. s. stelleri and C. s. annectens. Microsatellite analyses revealed genetic differentiation between each of the three subspecies, although more extensive sampling of additional C. s. annectens populations is needed to clarify the level of subspecies differentiation. High levels of population structure were found among C. s. stelleri populations with significant differences in all but two pairwise comparisons. A significant isolation by distance pattern was observed amongst populations in the Pacific Northwest and Alaska. In the C. s. carlottae population, there was evidence of reduced genetic variation, higher number of private alleles than northern C. s. stelleri populations and higher levels of divergence between Queen Charlotte Island and other populations. We were unable to reject the hypothesis that the Queen Charlotte Islands served as a refugium during the Pleistocene. Steller's jay may have colonized the Queen Charlotte Islands near the end of the last glaciation or persisted throughout the Pleistocene, and this subspecies may thus represent a glacial relic. The larger number of private alleles, despite reduced genetic variation, morphological distinctiveness and high divergence from other populations suggests that the Queen Charlotte Island colonization pre-dates that of the mainland. Furthermore, our results show rapid divergence in Steller's jay populations on the mainland following the retreat of the ice sheets.

TREES BY THE SEA: ADVANTAGES AND DISADVANTAGES OF OCEANIC CLIMATES

Proximity to the ocean can have both positive and negative effects for tree survival. Across the world relict forests that were once trans-continental in their distribution depend for their continued survival on coastal refugia. The tree species that dominate the cloud-zone forests of Macaronesia, the coastal redwoods of California, the Atlantic forests of Brazil, and the Podocarp forests of New Zealand's South Island are all examples of palaeoendemic species that once had a much wider distribution and appear to owe their survival to the particular environmental conditions provided by coastal sites and oceanic islands. By contrast, in the islands of the North Atlantic, oceanic conditions appear to limit tree regeneration and make forests vulnerable to human disturbance. Paradoxically, winter warmth appears to be harmful to trees in northern cool and moist oceanic conditions. There may be many reasons why the warm winters of maritime environments can be detrimental for woody species. Insect attack, pathogenicity and metabolic dysfunction with loss of frost hardiness and over-wintering carbohydrate reserves are all possibilities. Where long, mild winters are combined with wet soil conditions, metabolic dysfunction brought about by prolonged periods of oxygen deprivation can deplete root meristems of carbohydrate reserves that are essential for avoiding postanoxic injury when the soil profile once again becomes aerated in spring. In areas where the temperature remains below zero for the greater part of the winter, this is not a danger. However, in oceanic habitats there is always the risk, even in the far north, of temperatures rising above zero. Model predictions and field observations confirm the potential dangers of warm winters by suggesting that any significant degree of winter warming will cause a retreat of Pinus sylvestris and other woody species from oceanic regions at northern latitudes.

Trees by the Sea: Advantages and Disadvantages of Oceanic Climates

Proximity to the ocean can have both positive and negative effects for tree survival. Across the world relict forests that were once trans-continental in their distribution depend for their continued survival on coastal refugia. The tree species that dominate the cloud-zone forests of Macaronesia, the coastal redwoods of California, the Atlantic forests of Brazil, and the Podocarp forests of New Zealand's South Island are all examples of palaeoendemic species that once had a much wider distribution and appear to owe their survival to the particular environmental conditions provided by coastal sites and oceanic islands. By contrast, in the islands of the North Atlantic, oceanic conditions appear to limit tree regeneration and make forests vulnerable to human disturbance. Paradoxically, winter warmth appears to be harmful to trees in northern cool and moist oceanic conditions. There may be many reasons why the warm winters of maritime environments can be detrimental for woody species. Insect attack, pathogenicity and metabolic dysfunction with loss of frost hardiness and over-wintering carbohydrate reserves are all possibilities. Where long, mild winters are combined with wet soil conditions, metabolic dysfunction brought about by prolonged periods of oxygen deprivation can deplete root meristems of carbohydrate reserves that are essential for avoiding post anoxic injury when the soil profile once again becomes aerated in spring. In areas where the temperature remains below zero for the greater part of the winter, this is not a danger. However, in oceanic habitats there is always the risk, even in the far north, of temperatures rising above zero. Model predictions and field observations confirm the potential dangers of warm winters by suggesting that any significant degree of winter warming will cause a retreat of Pinns sylvestris and other woody species from oceanic regions at northern latitudes.

PHYLOGEOGRAPHY OF KEEN'S MOUSE (PEROMYSCUS KEENI) IN A NATURALLY FRAGMENTED LANDSCAPE

Phylogenetic methods were used to analyze cytochrome-b sequences (n ¼ 257; 560 base pairs) from Peromyscus keeni (Keen’s mouse) collected from 23 islands of the Alexander Archipelago and 6 mainland locations in southeast Alaska and western Canada. Although P. keeni is ubiquitous across this region, island populations are genetically distinctive. Genetic structure of extant populations of P. keeni appears to have been more heavily influenced by vicariance than post-glacial colonization. Populations of P. keeni might have survived in coastal refugia during the last glacial maximum (15 � 10 3 years ago). Island area was significantly correlated with 2 of 3 genetic diversity measures whereas island isolation was not. Areas with divergent populations were discovered, but were largely inconsistent with 3 of 5 currently recognized subspecies. Cryptic variation was detected in 8 areas not previously identified by morphologic analyses.