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Aim The Aleutian Islands, one of the southern margins of Beringia, were covered with ice caps during the Last Glacial Maximum (LGM). Although insular refugia for animal species have been hypothesized in this region, phylogeographical studies have never explored the biogeographical history of plant species. We aim to assess the population history of the flora of the Aleutian Islands and its biogeographical relevance to East Asia. Location The Aleutian Islands, Kamchatka, the Japanese Archipelago. Methods We sequenced seven nuclear loci from the alpine shrub Therorhodion camtschaticum (Pall.) Small (Ericaceae) and elucidated the geographical distribution of genetic variation. The divergence history among populations from the Aleutian Islands, Kamchatka and northern Japan was inferred by analyses based on coalescent simulations. Results Populations were genetically distinct between northern Japan and Beringia. In spite of the genetic differentiation, the divergence between Beringia and northern Japan occurred during or after the last glacial period [maximum-likelihood estimates (MLEs) = 10.6–15.5 ka with 95% highest posterior densities (HPDs): 1.7–43.5 ka]. The latter region has a larger population size than the former region. In contrast, widespread genetic similarity was revealed among the geographically separated populations in the Aleutian Islands and southern Kamchatka. Among these populations, a model assuming population decline fits better than a model that assumes either constant population size or population growth. Main conclusions Our study suggests that insular refugia were unlikely for T. camtschaticum in the Aleutian Islands. Instead, T. camtschaticum may have had a wider range during LGM, encompassing both the Aleutian Islands and southern Kamchatka, or have colonized the Aleutian Islands from a LGM refugium that was the source of colonization into southern Kamchatka as well or encompassed the latter region. Our study provides novel insight into the biogeographical history of terrestrial biota in the Aleutian Islands.

Arctic–alpine plants have enormous ranges in the Northern Hemisphere. Phylogeographic studies have provided insights into their glacial survival as well as their postglacial colonization history. However, our understanding of the population dynamics of disjunct alpine populations in temperate regions remains limited. During Pleistocene cold periods, alpine populations of arctic–alpine species in East Asia were either connected to an ice-free Beringia refugium or they persisted with prolonged isolation after their establishment. To estimate which of these scenarios is more likely, we elucidated the genetic structure of Phyllodoce caerulea (Ericaceae) in Beringia and northern Japan, East Asia. Sequence variation in multiple nuclear loci revealed that P. caerulea can be distinguished into northern and southern groups. A demographic analysis demonstrated that the north–south divergence did not predate the last glacial period and detected introgression from Phyllodoce aleutica, relative widely distributed in East Asia, exclusively into the southern group. Therefore, although there has been genetic divergence between northern Japan and Beringia in P. caerulea, the divergence is unlikely to have resulted from their prolonged geographic separation throughout several cycles of glacial and interglacial periods. Instead, our study suggests that the introgression contributed to the genetic divergence of P. caerulea and that the range of P. caerulea was plausibly connected between northern Japan and Beringia during the last glacial period. Overall, our study not only provides a biogeographic insight into alpine populations of arctic–alpine plants in East Asia but also emphasizes the importance of careful interpretation of genetic structure for inferring phylogeographic history.

Given that East Asia is located south-west of Beringia and was less glaciated during the Pleistocene, species at higher latitudes were able to expand their range in this region during climate cooling. Although southward migration is an inevitable colonization process, the biogeographical history of the disjunct ranges of higher-latitude species in East Asia has been investigated less extensively. Here, we assess whether their disjunct distributions in the Japanese archipelago connected sufficiently with Beringia or persisted in isolation following their establishment. Sequences of nine nuclear loci were determined for Cassiope lycopodioides (Ericaceae) from the Japanese archipelago as well as its surrounding areas, Kamchatka and Alaska. According to the geographical pattern of genetic diversity, the northern populations from Kamchatka to the northern part of the Japanese archipelago were similar genetically and were differentiated from populations in central Japan. Our study suggested that the distribution of C. lycopodioides was connected between the northern part of the Japanese archipelago and south-western Beringia due to Pleistocene climate cooling. Conversely, central Japan harboured a disjunct range after its establishment. These inferences suggest that widespread range expansion in northern East Asia was plausible for species distributed in Beringia. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113, 497–509.

Disjunct geographic distributions of boreal plant species at the southern edges of their ranges are expected to result in low genetic diversity and high genetic differentiation in the disjunct populations. This prediction was tested in a riparian willow, Salix arbutifolia, distributed in the northeastern Eurasian continent and the Sakhalin, Hokkaido, and Honshu Islands, using chloroplast DNA haplotypes and nuclear microsatellite genotypes. Hokkaido and Honshu populations shared a chloroplast haplotype identical to a closely related species, S. cardiophylla. This haplotype was divergent from haplotypes in the Eurasian continent (Primorsky) and the Sakhalin Island. In the nuclear microsatellites, most Hokkaido populations were genetically closer to Primorsky populations than to Sakhalin populations in spite of the geographical vicinity between Sakhalin and Hokkaido. The unexpected genetic divergence between Sakhalin and Hokkaido implies a complicated history of migration and colonization. The most peripheral populations in Honshu had the lowest genetic diversity and were most differentiated from the others. Thus, low genetic diversity and high genetic differentiation at the range periphery suggest substantial effects of genetic drift on genetic structure in the disjunct populations of Salix arbutifolia at the southern edge of its range.

The circumarctic ranges of arctic-alpine plants are thought to have been established in the late Pliocene/early Pleistocene, when the modern arctic tundra was formed in response to climate cooling. Previous findings of range-wide genetic structure in arctic-alpine plants have been thought to support this hypothesis, but few studies have explicitly addressed the temporal framework of the genetic structure. Here, we estimated the demographic history of the genetic structure in the circumarctic Kalmia procumbens using sequences of multiple nuclear loci and examined whether its genetic structure reflects prolonged isolation throughout the Pleistocene. Both Bayesian clustering and phylogenetic analyses revealed genetic distinction between alpine and arctic regions, whereas detailed groupings were somewhat discordant between the analyses. By assuming a population grouping based on the phylogenetic analyses, which likely reflects a deeper intraspecific divergence, we conducted model-based analyses and demonstrated that the intraspecific genetic divergence in K. procumbens likely originated during the last glacial period. Thus, there is no need to postulate range separation throughout the Pleistocene to explain the current genetic structure in this species. This study demonstrates that range-wide genetic structure in arctic-alpine plants does not necessarily result from the late Pliocene/early Pleistocene origin of their circumarctic ranges, and emphasizes the importance of a temporal framework of the current genetic structure for understanding the biogeographic history of the arctic flora. This article is protected by copyright. All rights reserved.

Aim Based on seven consecutive seasons of biotic survey and inventory of the terrestrial and freshwater plants and animals of the 30 major islands of the Kuril Archipelago, a description of the biodiversity and an analysis of the biogeography of this previously little known part of the world are provided. Location The Kuril Archipelago, a natural laboratory for investigations into the origin, subsequent evolution, and long-term maintenance of insular populations, forms the eastern boundary of the Okhotsk Sea, extending 1200 km between Hokkaido, Japan, and the Kamchatka Peninsula of Russia. A chain of more than 56 islands, the system is only slightly smaller than the Hawaiian Islands, covering an area of 15,600 km 2 and

Summary Following climate cooling at the end of the Tertiary, arctic-alpine plants attained most of their extant species diversity. Because East Asia was not heavily glaciated, the importance of this region as a location for the long-term persistence of these species and their subsequent endemism during the Pleistocene was proposed in early discussions of phytogeography. However, this hypothesis remains to be fully tested. Here, we address this hypothesis by elucidating the phylogenetic history of Phyllodoce (Ericaceae). A phylogenetic tree based on multiple nuclear loci revealed that Phyllodoce nipponica was not derived from widespread species such as the arctic-alpine Phyllodoce caerulea, but rather represented an independent lineage sister to the clade of widespread relatives. Molecular dating indicated a mid-Pleistocene divergence of P. nipponica. These findings exclude the hypothesis that P. nipponica was derived from an arctic-alpine species that extended its range southwards during recent glacial periods. Instead, our results support the hypothesis that P. nipponica is an ancestral species which persisted in the Japanese archipelago during the mid- and late Pleistocene. Our findings demonstrate support for the early proposal and shed light on the importance of the Japanese archipelago for the evolution and persistence of arctic-alpine species.