Structure, biodiversity, and historical biogeography of nematode faunas in holarctic ruminants: morphological and molecular diagnoses for Teladorsagia boreoarcticus n. sp. (Nematoda: Ostertagiinae), a dimorphic cryptic species in muskoxen (Ovibos moschatus)

Discovery of the ostertagiine nematode Teladorsagia boreoarcticus n. sp. in muskoxen, Ovibos moschatus, from the central Canadian Arctic highlights the paucity of knowledge about the genealogical and numerical diversity of nematode faunas characteristic of artiodactyls at high latitudes across the Holarctic. Teladorsagia boreoarcticus is a dimorphic cryptic species distinguished from Teladorsagia circumcincta/Teladorsagia trifurcata in domestic sheep by a 13% divergence in the ND4 region of mitochondrial DNA, constant differences in the synlophe, and significantly longer esophageal valve, spicules, gubernaculum, and bursa. Teladorsagia boreoarcticus represents an archaic component of the North American fauna and may have a Holarctic distribution in muskoxen and caribou. Recognition of T. boreoarcticus in muskoxen, in part, corroborates hypotheses for the existence of a cryptic species complex of Teladorsagia spp. among Caprinae and Cervidae at high latitudes and indicates the importance of climatological determinants during the late Tertiary and Pleistocene on diversification of the fauna. Also reinforced is the concept of the North American fauna as a mosaic of endemic and introduced species. Discovery of a previously unrecognized species of Teladorsagia has additional implications and clearly indicates that (1) our knowledge is incomplete relative to potentially pathogenic nematodes that could be exchanged among domestic and wild caprines; (2) we do not have sufficient knowledge of the fauna to understand the ecological control mechanisms (limitations) on dissemination and host range; and (3) an understanding of historical and geographical influences on the genealogical diversity and distribution of nematode faunas in domestic and wild ruminants is requisite to define the interface between agricultural and natural ecosystems across the Holarctic.     

Brucella suis biotype 4: a case of granulomatous nephritis in a barren ground caribou (Rangifer tarandus groenlandicus L.) with a review of the distribution of rangiferine brucellosis in Canada

Severe granulomatous nephritis caused by Brucella suis biotype 4 was found in a barren ground caribou (Rangifer tarandus groenlandicus) from Northwest Territories, Canada. A review of the distribution of human and animal cases of brucellosis in northern Canada indicated that B. suis biotype 4 is distributed widely and is probably enzootic in most Canadian caribou herds.     

Prevalence of Toxoplasma gondii antibodies in barren-ground caribou (Rangifer tarandus groenlandicus) from the Canadian Arctic

Prevalence of antibodies to Toxoplasma gondii was determined in 147 barren-ground caribou (Rangifer tarandus groenlandicus) from 5 herds in the Northwest Territories and Nunavut, northern Canada, by the modified agglutination test (MAT). In the mainland herds (Bluenose, Bathurst, and Beverly), antibodies were found in 43 (37%) of 117 caribou, and MAT titers were 1:25 in 10, 1:50 in 24, and 1:500 in 9. In the island herds, only 1 (4.3%) of 23 animals sampled from the North Baffin Island herd was positive (titer = 1:25) and no antibodies were detected in 7 caribou from the Dolphin and Union herd. The high prevalence of antibodies to T. gondii in the mainland caribou herds indicates that caribou meat may contain viable T. gondii.     

Prioritization of landscape connectivity for the conservation of Peary caribou

Adequate connectivity between discontinuous habitat patches is crucial for the persistence of metapopulations across space and time. Loss of landscape connectivity is often a direct result of fragmentation caused by human activities but also can be caused indirectly through anthropogenic climate change. Peary caribou (Rangifer tarandus pearyi) are widely dispersed across the islands of the Canadian Arctic Archipelago and rely on sea ice to move seasonally between island habitats throughout their range. Seasonal connectivity provided by sea ice is necessary to maintain genetic diversity and to facilitate dispersal and recolonization of areas from which caribou have been extirpated. We used least‐cost path analysis and circuit theory to model connectivity across Peary caribou range, and future climate projections to investigate how this connectivity might be affected by a warming climate. Further, we used measures of current flow centrality to estimate the role of High Arctic islands in maintaining connectivity between Peary caribou populations and to identify and prioritize those islands and linkages most important for conservation. Our results suggest that the Bathurst Island complex plays a critical role in facilitating connectivity between Peary caribou populations. Large islands, including Banks, Victoria, and Ellesmere have limited roles in connecting Peary caribou. Without rigorous greenhouse gas emission reductions our projections indicate that by 2100 all connectivity between the more southern Peary caribou populations will be lost for important spring and early‐winter movement periods. Continued connectivity across the Canadian Arctic Archipelago, and possibly Peary caribou persistence, ultimately hinges on global commitments to limit climate change. Our research highlights priority areas where, in addition to emission reductions, conservation efforts to maintain connectivity would be most effective.     

Mitochondrial DNA and microsatellite DNA variation in domestic reindeer (Rangifer tarandus tarandus) and relationships with wild caribou (Rangifer tarandus granti, Rangifer tarandus groenlandicus, and Rangifer tarandus caribou)

Reindeer (Rangifer tarandus tarandus) in Alaska are semidomestic livestock descended from 1280 animals introduced from Siberia, Russia, approximately 100 years ago. Genetic variation at 18 microsatellite DNA loci and the cytochrome b gene of mitochondrial DNA (mtDNA) was quantified in reindeer from Alaska, Siberia (Russia), and Scandinavia and compared with wild North American caribou. Mean sequence divergence among 15 mtDNA haplotypes in reindeer was 0.007 substitutions per nucleotide site, and reindeer mtDNA is polyphyletic with caribou mtDNA. Microsatellite allele and mtDNA haplotype frequencies are similar between Alaskan and Russian reindeer and differentiated between these and Scandinavian reindeer. The frequencies of microsatellite alleles and mtDNA haplotypes are different in reindeer and wild caribou (Rangifer tarandus granti, Rangifer tarandus groenlandicus, and Rangifer tarandus caribou). Alaskan reindeer have maintained a genetic variation comparable to that in Russia and differentiated from that of wild caribou, >100 years after their introduction to Alaska.     

An epizootic of besnoitiosis in captive caribou (Rangifer tarandus caribou), reindeer (Rangifer tarandus tarandus) and mule deer (Odocoileus hemionus hemionus)

Besnoitia sp. was diagnosed in two caribou (Rangifer tarandus caribou) which died of pneumonia at the Assiniboine Park Zoo (Winnipeg, Manitoba, Canada) in 1983. During the following 3 yr besnoitiosis spread to an isolated herd of caribou, to mule deer (Odocoileus hemionus hemionus) and to reindeer (Rangifer tarandus tarandus). Reduction of exposure to biting insects appears to have reduced the transmission of besnoitiosis within the reindeer herd. The morbidity rate was approximately 82% in caribou and 67% in mule deer over the age of 2 mo. Most animals with clinical signs were euthanized; this precluded an estimation of the disease-related mortality rate. Twenty-eight caribou, 10 mule deer and three reindeer have been euthanized or died as a result of this epidemic. Attempts to artificially transmit the disease to potentially susceptible intermediate and definitive hosts were unsuccessful.     

Invasion,establishment, and range expansion of two parasitic nematodes in the Canadian Arctic

Climate warming is occurring at an unprecedented rate in the Arctic and is having profound effects on host‐parasite interactions, including range expansion. Recently, two species of protostrongylid nematodes have emerged for the first time in muskoxen and caribou on Victoria Island in the western Canadian Arctic Archipelago. Umingmakstrongylus pallikuukensis, the muskox lungworm, was detected for the first time in 2008 in muskoxen at a community hunt on the southwest corner of the island and by 2012, it was found several hundred kilometers east in commercially harvested muskoxen near the town of Ikaluktutiak. In 2010, Varestrongylus sp., a recently discovered lungworm of caribou and muskoxen was found in muskoxen near Ikaluktutiak and has been found annually in this area since then. Whereas invasion of the island by U. pallikuukensis appears to have been mediated by stochastic movement of muskoxen from the mainland to the southwest corner of the island, Varestrongylus has likely been introduced at several times and locations by the seasonal migration of caribou between the island and the mainland. A newly permissive climate, now suitable for completion of the parasite life cycles in a single summer, likely facilitated the initial establishment and now drives range expansion for both parasites.     

Yersiniosis in free-ranging muskoxen on Banks Island, Northwest Territories, Canada.

Sixty-seven muskox (Ovibos moschatus) carcasses, 53 skeletal remains and two sick muskoxen were seen during an aerial survey of the Thomsen River region, northern Banks Island, Northwest Territories, Canada in late July 1986. Complete necropsies were performed on 29 muskoxen estimated to have died within the previous 3 to 5 days. Twenty were diagnosed with acute yersiniosis due to Yersinia pseudotuberculosis serotype 1B and were in excellent body condition. A diagnosis could not be made on seven animals due to marked autolysis; however, these muskoxen also were in excellent body condition. The remaining two were aged, emaciated muskoxen. This report describes the first occurrence of yersiniosis in free-ranging muskoxen and the first documentation of large scale mortality due to this disease in a free-ranging population of wild ungulates.     

Serum biochemistry, serology, and parasitology of boreal caribou (Rangifer tarandus caribou) in the Northwest Territories, Canada

Boreal caribou (Rangifer tarandus caribou) are an ecologically and culturally important wildlife species and now range almost exclusively in the boreal forests of Canada, including the Northwest Territories, northern Alberta, and British Columbia. Boreal caribou are threatened throughout their Canadian range because of direct and indirect natural and anthropogenic factors. In the Northwest Territories, however, they have a continuous range that overall has not yet been subjected to the same degree of anthropogenic habitat fragmentation and degradation that has occurred elsewhere in Canada. To monitor the health of boreal caribou populations and individuals, we collected blood from 104 adult, female boreal caribou captured between March 2003 and February 2006 and measured serum biochemical parameters. Serum creatinine was higher in pregnant than in nonpregnant caribou. Several biochemical parameters differed among years, but they tended to be similar to those reported for reindeer. Serum antibodies were found to an alphaherpesvirus, Toxoplasma gondii, and to the Mycobacterium avium subspecies paratuberculosis in 37.5, 2.9, and 1.3% of boreal caribou, respectively. Fecal samples were collected from 149 boreal caribou, and Cryptosporidium sp. oocysts, Giardia sp. cysts, trichostrongyle ova, dorsal-spined nematode larvae, cestode ova, and Eimeria sp. were found. Trypanosoma sp. was detected in the blood of 72.1% of boreal caribou. Eimeria sp., Cryptosporidium sp., and Giardia sp. have not been previously reported in boreal caribou.     

Herpesvirus infection in woodland caribou in Alberta, Canada

Sera and genital swabs collected from 121 adult woodland caribou (Rangifer tarandus caribou) in five subpopulations in northern Alberta, Canada, between December 1997 and October 1999, were examined for evidence of infection with herpesviruses or pestiviruses. No virus was isolated from sera or swabs, and no antibodies against bovine viral diarrhea virus were detected. However, 63 (52%) of the 121 animals had neutralizing antibody titers against bovine herpesvirus 1. There was sufficient serum from 37 of the 121 caribou to allow parallel testing for antibodies against a new alphaherpesvirus isolated from an elk (Cervus elaphus nelsoni), and 20 animals had antibodies against this virus. Paired sera collected 11 mo apart from 14 caribou showed seroconversion in seven animals, indicating that an active herpesvirus infection was present. Virus neutralization data suggest that these caribou are infected with a distinct alphaherpesvirus.     

Modern and ancient DNA reveal recent partial replacement of caribou in the southwest Yukon

The long‐term persistence of forest‐dwelling caribou (Rangifer tarandus caribou) will probably be determined by management and conservation decisions. Understanding the evolutionary relationships between modern caribou herds, and how these relationships have changed through time will provide key information for the design of appropriate management strategies. To explore these relationships, we amplified microsatellite and mitochondrial markers from modern caribou from across the Southern Yukon, Canada, as well as mitochondrial DNA from Holocene specimens recovered from alpine ice patches in the same region. Our analyses identify a genetically distinct group of caribou composed of herds from the Southern Lakes region that may warrant special management consideration. We also identify a partial genetic replacement event occurring 1000 years before present, coincident with the deposition of the White River tephra and the Medieval Warm Period. These results suggest that, in the face of increasing anthropogenic pressures and climate variability, maintaining the ability of caribou herds to expand in numbers and range may be more important than protecting the survival of any individual, isolated sedentary forest‐dwelling herd.     

Antler stiffness in caribou (Rangifer tarandus): testing variation in bone material properties between males and females

Vertebrate bones, including deer antler, often exhibit variation in their mechanical properties that corresponds to differences in the functional demands they encounter. Among deer, antlers are found in both males and females only in caribou (Rangifer tarandus). Several differences between male and female R. tarandus in behavior and physiology might subject their antlers to differing demands, potentially making divergence of antler material properties between the sexes advantageous. Alternatively, antler material properties might not differ between male and female R. tarandus because both sexes are members of the same species, and the properties of their antlers could, therefore, have emerged under similar pressures and constraints through evolution. To test for sexual dimorphism in antler material properties, we compared the stiffness (Young's modulus of elasticity) of antler specimens from male and female caribou using three-point bending tests. Despite behavioral and physiological differences between males and females, stiffness values did not differ significantly between the sexes in caribou, with a mean (+/-S.E.) stiffness of 5.8+/-0.4GPa across all specimens. This value differed by less than 10% from the values published for R. tarandus specimens of unknown sex, verifying the comparability of bone material property data collected across multiple studies, and lending confidence to recent analyses of the evolution of antler stiffness in deer that have drawn on literature data.     

Evidence of herpesvirus infection in Woodland Caribou in Saskatchewan

Sera were collected from 40 female and two male woodland caribou (Rangifer tarandus caribou) in Saskatchewan (Canada) from March 1992 to January 1995, inclusive. The samples were examined for antibodies against smooth Brucella spp., five serovars of Leptospira interrogans, bovine viral diarrhea virus, and bovine herpesvirus 1 (BHV-1). Twenty-two (52%) of 42 sera exhibited positive reactions to BHV-1 by a modified serum neutralization test, and the prevalence correlated positively with the age of the animals. No antibodies were detected against the other pathogens. This is the first reported evidence of herpesvirus infection in isolated populations of woodland caribou in western Canada.     

Survival in the Rockies of an endangered hybrid swarm from diverged caribou (Rangifer tarandus) lineages

In North America, caribou ( Rangifer tarandus ) experienced diversification in separate refugia before the last glacial maximum. Geographical isolation produced the barren-ground caribou ( Rangifer tarandus groenlandicus ) with its distinctive migratory habits, and the woodland caribou ( Rangifer tarandus caribou ), which has sedentary behaviour and is now in danger of extinction. Herein we report on the phylogenetics, population structure, and migratory habits of caribou in the Canadian Rockies, utilizing molecular and spatial data for 223 individuals. Mitochondrial DNA analyses show the occurrence of two highly diverged lineages; the Beringian–Eurasian and North American lineages, while microsatellite data reveal that present-day Rockies' caribou populations have resulted from interbreeding between these diverged lineages. An ice-free corridor at the end of the last glaciation likely allowed, for the first time, for barren-ground caribou to migrate from the North and overlap with woodland caribou expanding from the South. The lack of correlation between nuclear and mitochondrial data may indicate that different environmental forces, which might also include human-caused habitat loss and fragmentation, are currently reshaping the population structure of this postglacial hybrid swarm. Furthermore, spatial ecological data show evidence of pronounced migratory behaviour within the study area, and suggest that the probability of being migratory may be higher in individual caribou carrying a Beringian–Eurasian haplotype which is mainly associated with the barren-ground subspecies. Overall, our analyses reveal an intriguing example of postglacial mixing of diverged lineages. In a landscape that is changing due to climatic and human-mediated factors, an understanding of these dynamics, both past and present, is essential for management and conservation of these populations.     

Geographic pattern of serum antibody prevalence for Brucella spp. in caribou, grizzly bears, and wolves from Alaska, 1975-1998

Blood samples were collected from 2,635 caribou (Rangifer tarandus), 1,238 grizzly bears (Ursus arctos), and 930 wolves (Canis lupus) from throughout mainland Alaska during 1975-98. Sera were tested for evidence of exposure to Brucella spp. Serum antibody prevalences were highest in the northwestern region of the state. In any specific area, antibody prevalences for caribou and wolves were of a similar magnitude, whereas antibody prevalence for bears in these same areas were two to three times higher.     

Connecting the dots in databases of endangered species: a Bayesian hierarchical imputation strategy for missing Peary caribou (Rangifer tarandus pearyi) population data

Peary caribou (Rangifer tarandus pearyi) is listed as Endangered under Canada's Species at Risk Act. Studying rare and endangered species can be particularly challenging due to the constraints posed by incomplete datasets owing to poor weather conditions, lack of technology, organizational deficiencies, and high survey costs in remote areas. A defensible way to remedy data gaps and thus improve the robustness of any given modelling exercise is through imputation schemes, which are typically selected on the basis of missing data patterns. In this study, we develop a regression-based imputation method aiming to identify the linkages among available population records in time and space. One of the fundamental assumptions of our imputation model is that there is a subset of primary islands that act as the core areas from where the Peary caribou populations migrate to secondary or satellite islands. Specifically, we delineated six distinct geographic clusters (Banks, Axel Heiberg, Bathurst, Boothia, Melville and Mackenzie King) across the Canadian Arctic Archipelago, and within each cluster Peary caribou seasonal inter-island movements take place in any given year. Parameterized with Bayesian inference techniques, our imputation modelling strategy has a flexible structure that can accommodate non-monotonic spatiotemporal patterns, and was thus able to capture more than 65% of the variability in our dataset. The two islands (Banks and Northwest Victoria) of the Banks island complex exhibited an average decline rate of 6% per year over the past four decades, which collectively reflects the dramatic population decrease from the early 1970s until the late 1990s, as well as the distinct recovery after the early 2000s. Similar “wax-and-wane” cycles characterize the Peary caribou population patterns on Melville and Bathurst island complexes. Our analysis provides evidence of positive rate of change of the population trends on Axel Heiberg and Ellesmere Islands, which likely stems from pockets on those islands, where favorable climatic/geomorphological conditions, and rich vegetation prevail. In stark contrast, overharvesting, higher predation, adverse climatic conditions, and human-induced forage shortage have led to a dramatic decline, nearing their extirpation, on the Boothia island complex.     

Modeling density dependence and climatic disturbances in caribou: a case study from the Bathurst Island complex, Canadian High Arctic

Peary caribou Rangifer tarandus pearyi is the northernmost subspecies of Rangifer in North America and endemic to the Canadian High Arctic. Because of severe population declines following years of unfavorable winter weather with ice coating on the ground or thicker snow cover, it is believed that density-independent disturbance events are the primary driver for Peary caribou population dynamics. However, it is unclear to what extent density dependence may affect population dynamics of this species. Here, we test for different levels of density dependence in a stochastic, single-stage population model, based on available empirical information for the Bathurst Island complex (BIC) population in the Canadian High Arctic. We compare predicted densities with observed densities during 1961–2001 under various assumptions of the strength of density dependence. On the basis of our model, we found that scenarios with no or very low density dependence led to population densities far above observed densities. For average observed disturbance regimes, a carrying capacity of 0.1 caribou km⁻² generated an average caribou density similar to that estimated for the BIC population over the past four decades. With our model we also tested the potential effects of climate change-related increases in the probability and severity of disturbance years, that is unusually poor winter conditions. On the basis of our simulation results, we found that, in particular, potential increases in disturbance severity (as opposed to disturbance frequency) may pose a considerable threat to the persistence of this species.     

Wolves,white‐tailed deer,and beaver: implications of seasonal prey switching for woodland caribou declines

Population increases of primary prey can negatively impact alternate prey populations via demographic and behavioural responses of a shared predator through apparent competition. Seasonal variation in prey selection patterns by predators also can affect secondary and incidental prey by reducing spatial separation. Global warming and landscape changes in Alberta's bitumen sands have resulted in prey enrichment, which is changing the large mammal predator–prey system and causing declines in woodland caribou Rangifer tarandus caribou populations. We assessed seasonal patterns of prey use and spatial selection by wolves Canis lupus in two woodland caribou ranges in northeastern Alberta, Canada, that have undergone prey enrichment following recent white‐tailed deer Odocoileus virginianus invasion. We determined whether risk of predation for caribou (incidental prey) and the proportion of wolf‐caused‐caribou mortalities varied with season. We found that wolves showed seasonal variation in primary prey use, with deer and beaver Castor canadensis being the most common prey items in wolf diet in winter and summer, respectively. These seasonal dietary patterns were reflected in seasonal wolf spatial resource selection and resulted in contrasting spatial relationships between wolves and caribou. During winter, wolf selection for areas used by deer maintained strong spatial separation between wolves and caribou, whereas wolf selection for areas used by beaver in summer increased the overlap with caribou. Changing patterns in wolf resource selection were reflected by caribou mortality patterns, with 76.2% of 42 adult female caribou mortalities occurring in summer. Understanding seasonal patterns of predation following prey enrichment in a multiprey system is essential when assessing the effect of predation on an incidental prey species. Our results support the conclusion that wolves are proximately responsible for woodland caribou population declines throughout much of their range.     

An investigation of arterial disease in Alaskan reindeer and Caribou.

No significant lesions of atherosclerosis or other vascular diseases were found in the aorta and coronary arteries of 34 reindeer and 15 caribou (Rangifer tarandus). Serum lipid, phospholipid, cholesterol and triglyceride levels were similar in caribou and reindeer and did not differ greatly from those reported in other ruminants.