S. O. MacDonald

Beringia: Intercontinental exchange and diversification of high latitude mammals and their parasites during the Pliocene and Quaternary

ABSTRACT Beringia is the region spanning eastern Asia and northwestern North America that remained ice-free during the full glacial events of the Pleistocene. Numerous questions persist regarding the importance of this region in the evolution of northern faunas. Beringia has been implicated as both a high latitude refugium and as the crossroads (Bering Land Bridge) of the northern continents for boreal mammals. The Beringian Coevolution Project (BCP) is an international collaboration that has provided material to assess the pattern and timing of faunal exchange across the crossroads of the northern continents and the potential impact of past climatic events on differentiation. Mammals and associated parasite specimens have been collected and preserved from more than 200 field sites in eastern Russia, Alaska and northwestern Canada since 1999. Previously, fossils and taxonomic comparisons between Asia and North America mammals have shed light on these events. Molecular phylogenetics based on BCP specimens is now being used to trace the history of faunal exchange and diversification. We have found substantial phylogeographic structure in the Arctic and in Beringia in mustelid carnivores, arvicoline rodents, arctic hares and soricine shrews, including spatially concordant clades and contact zones across taxa that correspond to the edges of Beringia. Among the tapeworms of these mammalian hosts, new perspectives on diversity have also been developed. Arostrilepis horrida (Hymenolepididae) was considered to represent a single widespread and morphologically variable species occurring in a diversity of voles and lemmings in eastern and western Beringia and more broadly across the Holarctic region. The BCP has demonstrated a complex of at least 10 species that are poorly differentiated morphologically. The diversity of Paranoplocephala spp. and Anolocephaloides spp. (Anoplocephalidae) in Beringia included relatively few widespread and morphologically variable species in arvicolines. BCP collections have changed this perspective, allowing the recognition of a series of highly endemic species of Paranoplocephala that demonstrate very narrow host specificity, and additional species complexes among arvicolines. Thus, extensive, previously unrecognized, diversity for tapeworms of 2 major families characterizes the Beringian fauna. By elucidating evolutionary relationships and phylogeographic variation among populations, species and assemblages, refined views of the sequence and timing of biotic expansion, geographic colonization and impact of episodic climate change have been developed for Beringia. Ultimately, Beringia was a determining factor in the structure and biogeography of terrestrial faunas across the Nearctic and Neotropical regions during the Pliocene and Quaternary.

High-latitude diversification within Eurasian least shrews and Alaska tiny shrews (Soricidae)

Abstract A novel shrew was discovered recently in Alaska and described based on morphological characters as Sorex yukonicus. This species is closely allied to Sorex minutissimus, a widespread shrew ranging through Eurasia. Together their distribution spans Beringia, a large Pleistocene nonglaciated area that connected Asia and North America. Beringia was repeatedly divided due to raised sea levels during Pleistocene interglacials and subsequently reconnected during glacials. We tested predictions related to the influence of large-scale geologic events on genetic variability through a phylogeographic analysis of both species of shrew using evidence from 3 independent genetic loci. We found low genetic divergence between S. minutissimus and S. yukonicus across continents. However, major phylogeographic breaks were found for Eurasian and Maritime Northeast Asia populations. Neither species is reciprocally monophyletic for any of the loci examined. Coalescence times for all pairwise population comparisons within both species fall within the Wisconsinan–Weichselian glacial (<130 thousand years ago), and significant population expansion estimates date to the Holocene suggesting that divergence between these taxa is minimal and may not warrant recognition of 2 distinct species. Phylogeographic relationships and sequence divergence estimates place populations of North American S. yukonicus and Siberian S. minutissimus as most closely related, and together they are sister to European S. minutissimus. We conclude that populations east and west of the Bering Strait represent a single Holarctic species, S. minutissimus. Temporal changes in range based on ecological niche predictions and a comparative assessment of other codistributed taxa provide a preliminary view of potential Last Glacial Maximum refugia in northern Asia.

Insular arctic ground squirrels (Spermophilus parryii) of the North Pacific: indigenous or exotic?

Abstract We lack critical information for oceanic archipelagos worldwide related to the origin and status of insular faunas. In southwestern Alaska, in particular, a need exists to tease apart whether specific insular populations are naturally occurring or are the result of exotic introductions by humans. We analyzed variation in mitochondrial sequences of the cytochrome-b gene and 8 nuclear microsatellite loci across 215 individuals representing 17 populations (12 insular) to refine our understanding of the history of the previously identified Southwest clade of arctic ground squirrels (Spermophilus parryii). We found significant geographic structure that suggests long-term isolation and diversification (Ushugat Island and Cold Bay), but we also documented closely related populations that are likely the result of human-mediated introductions. The latter instances (Kavalga and Unalaska islands) corroborate reports from early Alaska explorers.

Arctic Museum Collections--Special Issue The Beringian Coevolution Project: Holistic Collections of Mammals and Associated Parasites Reveal Novel Perspectives on Evolutionary and Environmental Change in the North

The Beringian Coevolution Project (BCP), a field program underway in the high northern latitudes since 1999, has focused on building key scientific infrastructure for integrated specimen-based studies on mammals and their associated parasites. BCP has contributed new insights across temporal and spatial scales into how ancient climate and environmental change have shaped faunas, emphasizing processes of assembly, persistence, and diversification across the vast Beringian region. BCP collections also represent baseline records of biotic diversity from across the northern high latitudes at a time of accelerated environmental change. These specimens and associated data form an unmatched resource for identifying hidden diversity, interpreting past responses to climate oscillations, documenting contemporary conditions, and anticipating outcomes for complex biological systems in a regime of ecological perturbation. Because of its dual focus on hosts and parasites, the BCP record also provides a foundation for c...