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1  Boundaries and definitions for the Arctic and Subarctic regions according to the Programme for the Conservation of Arctic Flora and Fauna (CAFF), shown in polar projection. Boundaries can be defined by the isotherms, habitat, latitude or geopolitical zones. Source map was developed by cartographer Philippe Rekacewicz (UNEP/GRID- Arendal) and is made available by CAFF at  _of_the_arctic. (For color version of this figure, the reader is referred to the web version of this book.) 

1 Boundaries and definitions for the Arctic and Subarctic regions according to the Programme for the Conservation of Arctic Flora and Fauna (CAFF), shown in polar projection. Boundaries can be defined by the isotherms, habitat, latitude or geopolitical zones. Source map was developed by cartographer Philippe Rekacewicz (UNEP/GRID- Arendal) and is made available by CAFF at _of_the_arctic. (For color version of this figure, the reader is referred to the web version of this book.) 

Context in source publication

Context 1
... Earth’s northern circumpolar regions present landscapes of outstand- ing beauty, incomparable fauna and flora and small, widely dispersed human settlements. These are integrated biological systems, where peo- ples of great resilience remain culturally close to the land and dependant on an array of natural resources, while living with the extremes of short summers and long cold winters (Rausch, 1951; Anisimov et al., 2007). The North encompasses those regions extending from the borders of the Sub- arctic (above 50°N) to beyond the Arctic Circle (north of 66°33 ′ N) ( Fig. 1.1). The latter marks the approximate limit for the current northern treeline and the circumpolar zone characterised seasonally by periods of constant polar night or midnight sun. Treeless tundra habitats dominate the Arctic where the average temperatures for the warmest month do not exceed 10°C; mean annual temperatures for the western subregions of the Arctic range from −20 to +12°C with minimal precipitation varying from 5 to 150mm (Callaghan et al., 2004e). The Arctic transitions into more south- erly Subarctic environments dominated primarily by taiga forests and a more complex mosaic of habitats defined by latitude and altitude. Northern ecosystems were formed by complex abiotic and biotic mechanisms in a crucible driven by episodic climatological processes and environmental perturbation extending across the late Pliocene and Quaternary during the past 3–5 million years (Myr) (e.g. Hopkins et al., 1982; Andersen and Borns, 1994 Dynesius and Jansson, 2000; Jansson and Dynesius, 2002; Callaghan et al., 2004a,c; Hewitt, 2004a,b). Contemporary patterns of faunal complexity reflect extinction events largely coincidental with the thermal maximum that signalled the termination of continental glaciation only 10 thousand years ago (Ka) (e.g. Barnosky et al., 2004). Consequently, northern biotas are typically characterised as relatively sim- ple, low-diversity assemblages with short trophic linkages, few pathogens and limited resilience or capacity for adaptation to environmental change (e.g. Callaghan et al., 2004a) (Table 1.1). A gradient of declining diversity with increasing latitude (from taiga forests to polar deserts) is also accom- panied by a shift or increase in dominance for some species, which may be manifested seasonally or annually (e.g. Callaghan et al., 2004b). High- latitude biotas are now at their minimal extent relative to patterns of diver- sity and geographic distribution that characterised faunas during the mid to late Pleistocene (e.g. Guthrie, 1984; Callaghan et al., 2004a). At high latitudes, vulnerable systems of low diversity continue to undergo significant and in some instances accelerating change due largely to human activity, both local and distant. These perturbations have the potential for broader impacts at a global scale (Callaghan et al., 2004a,d; Anisimov et al., 2007; Lawler et al., 2009; Post et al., 2009). Increasingly, substantial discussion has focused on the status and future of northern ecosystems, but these assessments have been limited to free-living organisms. Parasitism, pathogens and diseases have not yet been an integral component of this discourse (Kutz et al., 2009a). Parasites represent in excess of 40–50% of the organisms on Earth and complex assemblages of macroparasites (helminths and arthropods) and microparasites (viruses, bacteria and protozoans) shape ecosystems, food webs, host demographics and behaviour (e.g. Marcogliese, 2005; Hudson et al., 2006; Dobson et al., 2008). In some ecosystems, the biomass for macroparasites exceeds that for apex predators such as birds and fishes, suggesting a substantial role for otherwise obscure organisms at local to regional scales (Kuris et al., 2008). Parasites can cause disease and mortal- ity may influence the dynamics of wildlife populations and, in the worst- case scenarios, contribute to extinction events. Parasites are ubiquitous and diverse members of all biological commu- nities including those at high latitudes. All animals in circumpolar regions are susceptible to infection by characteristic assemblages of macroparasites and microparasites. Parasites can have subtle to severe effects on individual hosts or broader impacts on host populations that may cascade through ecosystems. Parasitic diseases have dual significance: (1) influencing sus- tainability for species and populations of diverse invertebrates, fishes, birds and mammals and (2) secondarily affecting food security, quality and avail- ability for people. Additionally, as zoonoses, some parasites can infect and cause disease in people and are a primary issue for food safety and human health (e.g. Kutz et al., 2009b; Jenkins et al., 2011). Sustainability, security and safety of ‘country foods’ are of concern at northern latitudes where people maintain a strong reliance on wildlife species. The potential signifi- cance of zoonoses is magnified in the North by the intimate linkage between wildlife and people. Subsistence food chains depend on the harvesting of free-ranging mammals, birds and fish for food, fibre and other animal prod- ucts. Understanding the role and influence of parasites in northern systems emerges from explorations of history, biogeography and the intricate eco- logical linkages among fishes, birds, mammals, domesticated species and people in the context of broader global connections (e.g. Hoberg, 2010). Despite the extreme environmental conditions in the North, the triad of host, parasite and environment remains ...

Citations

... With respect to parasitic infections, more than 100 parasite species have been reported to infect or infest reindeer [89][90][91][92][93][94], many of which are also shared with other ruminants. Protozoan parasites include Eimeria species [95], Cryptosporidium and Giardia [96], Entamoeba [97], Besnoitia [98,99] and Toxoplasma gondii [100][101][102][103]. Reindeer in Fennoscandia are intermediate hosts for Sarcocystis spp. ...
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The reindeer (caribou) Rangifer tarandus is a Cervidae in the order Artiodactyla. Reindeer are sedentary and migratory populations with circumpolar distribution in the Arctic, Northern Europe, Siberia and North America. Reindeer are an important wild and domesticated species, and have developed various adaptive strategies to extreme environments. Importantly, deer have also been identified to be putative zoonotic carriers, including for parasites, prions and coronavirus. Therefore, novel insights into immune-related markers are of considerable interest. Peptidylarginine deiminases (PADs) are a phylogenetically conserved enzyme family which causes post-translational protein deimination by converting arginine into citrulline in target proteins. This affects protein function in health and disease. Extracellular vesicles (EVs) participate in cellular communication, in physiological and pathological processes, via transfer of cargo material, and their release is partly regulated by PADs. This study assessed deiminated protein and EV profile signatures in plasma from sixteen healthy wild female reindeer, collected in Iceland during screening for parasites and chronic wasting disease. Reindeer plasma EV profiles showed a poly-dispersed distribution from 30 to 400 nm and were positive for phylogenetically conserved EV-specific markers. Deiminated proteins were isolated from whole plasma and plasma EVs, identified by proteomic analysis and protein interaction networks assessed by KEGG and GO analysis. This revealed a large number of deimination-enriched pathways for immunity and metabolism, with some differences between whole plasma and EVs. While shared KEGG pathways for whole plasma and plasma EVs included complement and coagulation pathways, KEGG pathways specific for EVs were for protein digestion and absorption, platelet activation, amoebiasis, the AGE–RAGE signaling pathway in diabetic complications, ECM receptor interaction, the relaxin signaling pathway and the estrogen signaling pathway. KEGG pathways specific for whole plasma were pertussis, ferroptosis, SLE, thyroid hormone synthesis, phagosome, Staphylococcus aureus infection, vitamin digestion and absorption, and prion disease. Further differences were also found between molecular function and biological processes GO pathways when comparing functional STRING networks for deiminated proteins in EVs, compared with deiminated proteins in whole plasma. This study highlights deiminated proteins and EVs as candidate biomarkers for reindeer health and may provide information on regulation of immune pathways in physiological and pathological processes, including neurodegenerative (prion) disease and zoonosis.
... Specimens of S. baturini were loaned from the Museum of Southwestern Biology, Division of Parasites, University of New Mexico in Albuquerque, USA. They originate from the intestines of three species of mustelids (Martes americana, Martes caurina, Mustela erminea), which were collected and dissected in the Beringia Coevolution Project (see Hoberg et al. 2003;Cook et al. 2005;Koehler et al. 2009b). Accession numbers and additional data on the samples are summarised in Table 1. ...
Article
Specimens of Soboliphyme baturini (Dioctophymatoida), collected from martens in Alaska, were investigated by scanning electron microscopy (SEM) and histological sections. Soboliphyme species are recognized by their large oral sucker, which is a muscular organ made of radial and longitudinal musculature. A ring of 6 papillae is present on the outer side of the oral sucker, as well as an individually variable pattern of additional papillae. Papillae are also present in a roughly longitudinal line along the body. We document the histological anatomy of the nervous system, the cervical sacs, pharynx and intestine, the musculature, the mesenteries and the reproductive organs. Several characters documented from S. baturini are proposed as apomorphies of Dioctophymatida: the anterior position of the brain ring, the presence of cervical sacs and the presence of two extracellular and four muscular mesenteries connecting the intestinal system to the body wall.
... Specimen-rich museum collections of tapeworms held at the Institute of Systematics and Ecology of Animals in Novosibirsk, Russia (ISEA) have been a key to understanding faunal structure in Eurasia, especially eastern Europe and northern Asia (e.g., . Detailed knowledge of the host and geographic distribution for species of Arostrilepis has emerged from the collections of the Beringian Coevolution Project and Integrated Inventories of Biomes of the Arctic as the source of extensive specimens, archives, and insights about biogeography and systematics of arvicolines and tapeworms across north-western North America, through Alaska and into eastern Siberia (Hoberg et al. 2003Cook et al. 2005Cook et al. , 2017. In contrast, potential records of cestodes that may be attributable to Arostrilepis from the central and eastern regions of the Nearctic have been unsubstantiated by archived specimens (e.g., Rausch 1952Rausch , 1957Schiller 1952;Voge 1952;Makarikov et al. 2013;Makarikov and Hoberg 2016). ...
... Our current understanding of the structure and assembly of the Arostrilepis-Arvicolinae rodent fauna has been consistent with origins in Eurasia and independent episodes of geographic expansion into North America during the Quaternary (Hoberg et al. 2003Cook et al. 2005Cook et al. , 2017Makarikov et al. 2013). Discovery of multiple species in eastern North America, and the resulting phylogenetic framework, however, requires modification of this narrative for expansion and establishment from Eurasia. ...
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Species of the genus Arostrilepis were discovered and definitively identified for the first time in rodents from geographically disparate localities along the Appalachian Mountain range of eastern North America (West Virginia, Virginia, and Maine). These are the first confirmed records for species of Arostrilepis occurring east of the Rocky Mountains and the Mississippi River in North America. Arostrilepis gardneri n. sp. is described on the basis of specimens obtained from two phylogenetically divergent rodent hosts: Southern Red-Backed Vole Myodes gapperi (Cricetidae: Arvicolinae) (from West Virginia) and the Woodland Jumping Mouse Napaeozapus insignis (Dipodidae: Zapodinae) (West Virginia, Virginia, and Maine). Additionally, in a mixed infection, specimens of Arostrilepis insperata n. sp. were also found in a Southern Red-Backed Vole from West Virginia. These previously unknown species are primarily distinguished from congeners based on shape, dimensions, and spination (pattern, shape, and size of spines) of the cirrus. Specimens of A. gardneri n. sp. are further characterized by the relative position and length of the cirrus-sac, arrangement of the testes, and relative size of the external seminal vesicle and seminal receptacle. Specimens of A. insperata n. sp. are structurally most similar to A. macrocirrosa from the western Nearctic and Palearctic but with consistently greater dimensions for the cirrus-sac, testes, and seminal receptacle. Phylogenetic analysis of Arostrilepis spp. using partial sequences of the mitochondrial cytochrome b gene and the nuclear second ribosomal internal transcribed spacer strongly supported the status of A. gardneri n. sp. and A. insperata n. sp. within an unresolved clade of congeners in Red-Backed Voles (Myodini and species of Myodes). Our observations extend the known geographic distribution for species of Arostrilepis to the Appalachian Mountains in either a disjunct or possibly continuous but patchy range across North America. Prior observations, summarizing field and museum collections, had suggested that geographic ranges for a diverse assemblage of Arostrilepis in North America were largely restricted to the north-western region of the continent, with historical connections to Beringia and Eurasia. Recognition of a more extensive distribution is consistent with a history of episodic biotic expansion and isolation under a dynamic of taxon pulses for arvicoline rodents and an associated parasite fauna in the Nearctic during the Quaternary. Occurrence in a dipodid rodent represents an event of host colonization from an arvicoline source.
... Fossil fuels, or hydrocarbons, due to their composition of carbon and hydrogen, have been a very attractive energy source mainly related to flexibility in transformation, efficiency, and high combustibility. Coal, oil, and gas have been used for more than two centuries (Hilyard 2012) and continue to be the base of the current global energy system despite the clear signals of depletion and decline in the main supply basins producing since the beginning of the 20th century. ...
... Parasites are an important component of Arctic ecosystems, influencing the health and sustainability of wildlife populations and the people who depend on them (Hoberg et al. 2003;Davidson et al. 2011). ...
... To use parasites as indicators of environmental health and to track or predict changes in parasitism and animal health, comprehensive data on parasite diversity, distributions, and life cycles are essential (Hoberg et al. 2003;Hoberg and Brooks 2008;. Although considerable progress has been made in defining the diversity and ecology of parasites found in/on Arctic vertebrates (Rausch 1974;Hoberg et al. 2012a), there remain substantial knowledge gaps. ...
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Parasites are key components of Arctic ecosystems. The current rate of climate and landscape changes in the Arctic is expected to alter host-parasite interactions, creating a significant concern for the sustainability of Arctic vertebrates. In addition to direct effects on host populations, changes in parasite loads on wildlife can have significant impacts on the people who depend on these organisms for food. Parasites play important roles in maintaining ecosystem stability through the regulation of host populations, and can provide unique insights into ecosystem structure. The present review examines the literature on the parasites of harvested wildlife in the Canadian North, including studies in the Yukon, Northwest Territories, Nunavut, northern Quebec, and Newfoundland and Labrador. For host species with higher mobility, we included records from other regions, such as Greenland, Russia, and the Canadian Subarctic, when no parasitological studies were available for the Canadian North. In addition, we searched databases of the Parasite Collection at the Canadian Museum of Nature and the United States Parasite Collection for records from the Canadian North. We found records for 248 species of macroparasites in vertebrate species of country food of animal origin in the Canadian North, including flatworms, roundworms, thorny-headed worms, ticks, lice, fleas, flies, and tongue worms. This review highlights the need to extend the study of the parasites that infect the primary species of harvested wildlife in the Canadian North. More detailed information on parasite communities is particularly important as climate and landscape change raises the possibility that new parasite species will colonize the region. Building a DNA barcode library for the parasites from country food in the area will facilitate their identification and monitoring.
... The summers are short, cool, and dry, providing a narrow developmental window for ectotherms, including parasites, and a short growing season for endotherms and plants [1][2][3]. Despite these adversities, a diverse group of flora and fauna, ranging from large mammals to microscopic parasites, constitutes Arctic biodiversity [4][5][6]. Studies have shown that these organisms develop unique physiological and behavioural strategies to cope the extremes [7,8]. ...
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Background: Umingmakstrongylus pallikuukensis and Varestrongylus eleguneniensis are two potentially pathogenic lungworms of caribou and muskoxen in the Canadian Arctic. These parasites are currently undergoing northward range expansion at differential rates. It is hypothesized that their invasion and spread to the Canadian Arctic Archipelago are in part driven by climate warming. However, very little is known regarding their physiological ecology, limiting our ability to parameterize ecological models to test these hypotheses and make meaningful predictions. In this study, the developmental parameters of V. eleguneniensis inside a gastropod intermediate host were determined and freezing survival of U. pallikuukensis and V. eleguneniensis were compared. Methods: Slug intermediate hosts, Deroceras laeve, were collected from their natural habitat and experimentally infected with first-stage larvae (L1) of V. eleguneniensis. Development of L1 to third-stage larvae (L3) in D. laeve was studied at constant temperature treatments from 8.5 to 24 °C. To determine freezing survival, freshly collected L1 of both parasite species were held in water at subzero temperatures from -10 to -80 °C, and the number of L1 surviving were counted at 2, 7, 30, 90 and 180 days. Results: The lower threshold temperature (T0) below which the larvae of V. eleguneniensis did not develop into L3 was 9.54 °C and the degree-days required for development (DD) was 171.25. Both U. pallikuukensis and V. eleguneniensis showed remarkable freeze tolerance: more than 80% of L1 survived across all temperatures and durations. Larval survival decreased with freezing duration but did not differ between the two species. Conclusion: Both U. pallikuukensis and V. eleguneniensis have high freezing survival that allows them to survive severe Arctic winters. The higher T0 and DD of V. eleguneniensis compared to U. pallikuukensis may contribute to the comparatively slower range expansion of the former. Our study advances knowledge of Arctic parasitology and provides ecological and physiological data that can be useful for parameterizing ecological models.
... Parasites are an important component of Arctic ecosystems, influencing the health and sustainability of wildlife populations and the people who depend on them (Hoberg et al. 2003;Davidson et al. 2011). Beginning in the 1940s, a succession of parasitologists and ecologists have explored parasitism in the Arctic, advancing understanding of the structure and function of host-parasite systems in this setting. ...
... The Simmons Collection not only represents a parasitological dataset with an intrinsic value to assess ecological questions, it also, and maybe even more importantly, highlights the relevance of proper archiving samples in accredited repositories for the study of wildlife diseases [77,78]. Archival collections are the window to study past geographic and genetic distribution of disease agents and also are a direct opportunity to maximize the impact of federal and private funds over time [14,[79][80][81]. ...
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Gastrointestinal helminths can have a detrimental effect on the fitness of wild ungulates. Arctic and Subarctic ecosystems are ideal for the study of host-parasite interactions due to the comparatively simple ecological interactions and limited confounding factors. We used a unique dataset assembled in the early seventies to study the diversity of gastrointestinal helminths and their effect on fitness indicators of Dall’s sheep, Ovis dalli dalli, in the Mackenzie Mountains, Northwest Territories, Canada. Parasite diversity included nine species, among which the abomasal nematode Marshallagia marshalli occurred with the highest prevalence and infection intensity. The intensity of M. marshalli increased with age and was negatively associated with body condition and pregnancy status in Dall’s sheep across all the analyses performed. The intensity of the intestinal whipworm, Trichuris schumakovitschi, decreased with age. No other parasites were significantly associated with age, body condition, or pregnancy. Our study suggests that M. marshalli might negatively influence fitness of adult female Dall’s sheep.
... The study site and its history of data acquisition Long-term research initiatives provide a key area where we might enhance the integration of specimen-based investigations within experimental systems ). Long-term experiments offer great potential for providing site-intensive temporal series of specimens to document and understand historical biotic responses to change and predict possible biodiversity scenarios in the Anthropocene era (Hoberg et al. 2003, Cook et al. 2005. The Sevilleta Long-Term Ecological Research (LTER) site in central New Mexico, United States, has been collecting community data on the regional small-mammal fauna at multiple sites since 1989. ...
... Furthermore, combined treatments of release and removal trapping can minimize "data deficiency" as a factor driving regulation of field investigations, particularly studies that incorporate specimen collection of inherently rare species (Winker et al. 2010). Ultimately, holistic sampling of hosts, parasites, and associated natural-history information can more accurately be extended to generalizable models of biodiversity responses across temporal and spatial scales (Hoberg et al. 2003, Winker 2004). ...
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Despite increasing use of specimens from natural-history collections, continued field sampling has met with growing resistance attributable to changing societal values. Widespread perception persists that the removal of individuals from wild populations will affect the integrity of natural communities. Ecological studies often document the resilience of wildlife to sustainable removal and the negligible contributions to mortality of scientific collecting compared with those of other natural or anthropogenic-induced causes. Nevertheless, few studies have directly assessed the consequences of specimen removal on populations or communities. We present long-term ecological research data that suggest removal trapping has negligible impacts on the species richness, diversity, or abundance of small mammals. The maintenance and future growth of natural-history archives for integrated biodiversity sciences may hinge on increased dedication to specimen vouchering across ecological and evolutionary disciplines and wider acceptance by regulatory authorities and funding agencies. The effects of low-intensity collecting should be investigated for other taxa and across biomes.
... A second grant from NSF was 21 awarded in 2004 to Cook (University of New Mexico) and Hoberg,supporting 22 additional collections in Beringia, and finally a third grant for fieldwork to Cook,23 Hoberg, Kurt Galbreath (Northern Michigan University) and Eric DeChaine 24 (Western Washington University) was awarded in 2012, allowing mammal/parasite 25 collections to be extended beyond the traditional boundaries of Beringia into central 26 Asia (Mongolia) and interior Canada. A series of collaborative efforts with state 27 (e.g., Alaska Department of Fish and Game) and federal agencies (e.g., US Forest 28 Service, US Geological Survey, National Park Service) and international partners 29 (e.g., Institute of Biological Problems of the North in Magadan, Siberia; Yukon 30 Department of Renewable Resources; National University of Mongolia) augmented 31 NSF-funded fieldwork and significantly expanded geographic and taxonomic 32 sampling of mammal and parasite populations radiating from the Bering Strait (e.g., 33 Hoberg et al. 2003Hoberg et al. , 2012aCook et al. 2005). These museum collections are site-34 ...
... It may differ from the final official version of record. ecological time, constitutes a cornerstone for systematic and historical biogeographic 120 studies (e.g., summarized in Hoberg et al. 2003Hoberg et al. , 2012aCook et al. 2005). Critically, 121 this foundation provides a trajectory for development and application of new 122 methods for exploring genomic diversity, molecular systematics, ecology, 123 phylogeography, conservation genetics, physiology, and epidemiology. ...
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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 comparative analyses that can document the effects of dynamic change on the geographic distribution, transmission dynamics, and emergence of pathogens. By using specific examples from carnivores, eulipotyphlans, lagomorphs, rodents, ungulates, and their associated parasites, we demonstrate how broad, integrated field collections provide permanent infrastructure that informs policy decisions regarding human impact and the effect of climate change on natural populations.