Do human activity and infrastructure disturb domesticated reindeer? The need for the reindeer’s perspective

In recent decades, human–Rangifer (reindeer and caribou) interactions have increasingly been studied from a scientific perspective. Many of the studies have examined Norwegian wild reindeer or caribou in North America. It is often questioned whether results from these studies can be applied to reindeer in managed herds, as these animals have been exposed to domestication and are also more used to humans. In order to examine the domesticated reindeer’s reactions to various disturbance sources, we reviewed 18 studies of the effects of human activity and infrastructure on 12 populations of domesticated reindeer and compared these to studies on wild reindeer and caribou; based on this, we discuss the effects of domestication and tameness on reindeer responses to anthropogenic disturbance. We also consider the relevance of spatial and temporal scales and data collection methods when evaluating the results of these studies. The reviewed studies showed that domesticated reindeer exhibit avoidance behaviours up to 12 km away from infrastructure and sites of human activity and that the area they avoid may shift between seasons and years. Despite a long domestication process, reindeer within Sami reindeer-herding systems exhibit similar patterns of large-scale avoidance of anthropogenic disturbance as wild Rangifer, although the strength of their response may sometimes differ. This is not surprising since current Sami reindeer husbandry represents an extensive form of pastoralism, and the reindeer are not particularly tame. To obtain a true picture of how reindeer use their ranges, it is of fundamental importance to study the response pattern at a spatial and temporal scale that is relevant to the reindeer, whether domesticated or wild.     

Polymorphism of the mtDNA control region in wild reindeer Rangifer tarandus (Mammalia: Artioodactyla) from the European part of Russia

Genetic diversity of wild reindeer (Rangifer tarandus) inhabiting the European part of Russia, including Komi Republic, Arkhangelsk oblast, Murmansk oblast, and the Republic of Karelia was characterized using sequence polymorphism of the mtDNA control region. Despite of currently low population number of wild reindeer, they were characterized by a high level of genetic diversity (?? = 0.018; H = 0.872 to 0.914). Phylogenetic analysis showed close relationships between European reindeer and wild reindeer of Siberia. In reindeer from Murmansk oblast a haplotype in common with the wild reindeer form Southwestern Norway was described. The reindeer sample examined contained no haplotypes earlier described for the reindeer of Central Norway. It is suggested that in recent past wild reindeer from the European north of Russia formed one population with the reindeer from the north of the Asian part of Eurasia.     

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.     

Why don't Svalbard reindeer migrate?

Reindeer and caribou are best known as migratory, seasonally nomadic animals; many continental populations, for example, travel between distinct summer and winter ranges which may lie hundreds of km apart. Much less is known about the movements of animals belonging to island populations. This paper describes seasonal and annual movements of wild reindeer Rangifer tarandus platyrhynchus on the high arctic archipelago of Svalbard, based on observations of nine animals captured and individually marked in Adventdalen, Spitsbergen, between 1977 and 1982. Four ear-tagged reindeer (one male and three females) were followed extensively for between four and seven years. Five radio-collared females were followed intensively for seven months in 1982. Svalbard reindeer seem neither to undertake long migrations nor to be nomadic within seasons like mountain reindeer or barren-ground caribou. They appear instead to use small, traditional, seasonal home ranges more, for example, like red deer or wild sheep. This atypical behaviour is discussed in relation to the dispersion of reindeers' resources in Svalbard.     

Evidence of alphaherpesvirus infections in Alaskan caribou and reindeer

Background  

The reindeer (Rangifer tarandus tarandus) industry in Alaska began with animals imported from Siberia (Russia) in the 1890's. Cervid herpes virus 2 (CvHV2) is endemic in reindeer in Scandinavia. We sought to determine if the same virus, or similar herpesviruses, were circulating in Alaskan reindeer and caribou (Rangifer tarandus granti). Serum samples from 292 reindeer were collected during annual reindeer handlings (1988-2005) near Nome, Alaska. In 2005, swab samples were collected from 40 calves from this herd, near Nome, Alaska. In 2007, ocular and nasal swab samples were collected from 30 apparently healthy reindeer calves near Wales, Alaska. Samples of plasma and white blood cells were collected from three Alaskan caribou herds, Mulchatna (n = 24), Teshekpuk (n = 34) and the Western Arctic (n = 87) in 2009.     

Foraging conditions, tooth wear and herbivore body reserves: a study of female reindeer

Several aspects of the life history of reindeer (Rangifer tarandus) are related to the nutritional condition of the animals. Moreover, compensatory growth and fattening in summer decreases with age. The interaction of tooth wear and the standing crop of lichens on age-related variation in body size and tissue reserves was examined to evaluate the proximate causes of density-dependent food limitation on life history parameters in female reindeer. Studies in nine semi-domesticated free-ranging reindeer herds showed that molar height depended on the mean standing crop of terrestrial lichens in winter habitats. The extent of tooth wear had the strongest effect on body reserves among the oldest females (11–14 years). This indicates that severe tooth wear limits the animals′ ability to process food efficiently and, hence, to maintain their body reserves. Both tooth wear and the biomass of lichens influenced body mass in old females, probably because on heavily exploited winter ranges reindeer had to use higher proportions of lower-ranking coarser foods, especially dwarf shrubs. Received: 1 December 1997 / Accepted: 13 July 1998     

Taxonomic Notes on the Swedish Domestic Reindeer, Genus Rangifer

In order to study the taxonomic relationship between domestic mountain and forest reindeer in Sweden nine skull characters on 126 and 62 animals, respectively, were measured and analysed. In addition antler width and shape of antler beam in cross-section were also measured. The differences indicate that the two groups of domestic reindeer are not entirely identical, although the differences are clearly not as plain as has been claimed. Comparing these results with earlier results it also appears that domestic forest reindeer are more in accordance with Rangifer tarandus tarandus than with R. t. jennicus , thus supporting the theory that possible influence of the latter subspecies on domestic reindeer has been considerably reduced as a result of long and intense cross breeding between the two groups of domestic reindeer.     

Nutritive value of terricolous lichens for reindeer in winter

Abstract:In contrast to virtually all other species of ruminants, reindeer (Rangifer tarandus) eat substantial amounts of lichens in winter. Several different species of lichens are eaten and most are highly palatable. The chemical composition and the in vitro digestibility of six species of terricolous lichens commonly eaten by reindeer were measured. Ruminal inoculum was obtained in winter from six free-ranging reindeer that had been grazing on natural pastures in northern Norway. Analysis of the plant parts recovered from their rumens confirmed that the animals had been eating a mixed diet that included both lichens and vascular plants. The chemical composition and the in vitro digestibility of the different species of lichens varied considerably between species and genera. Cetraria islandica, C. nivalis andCladonia arbuscula were highly digestible [69–77% dry matter (DM)], Stereocaulon paschale was poorly digestible (43% DM), whereas Cladonia stellaris and C. gracilis had an intermediate digestibility (56–57% DM). Mixing Cladonia stellaris with vascular plants (50:50) had no effect on the in vitro digestibility of the combined substrates. The in vitro digestibility of Cladonia stellaris in inoculum from two captive reindeer which had had no access to lichens was very low (10% DM). This shows that the source of inoculum used for digestibility trials has a major effect on the apparent digestibility of the substrates. Evidently, the extent to which reindeer are able to utilize lichens depends on the species that are selected and on what the animals have been eating recently.     

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.     

Strontium isotope evidence for migration in late Pleistocene Rangifer: implications for Neanderthal hunting strategies at the Middle Palaeolithic site of Jonzac, France

In order to understand the behaviours and subsistence choices of Palaeolithic hunter-gatherers, it is essential to understand the behavioural ecology of their prey. Here, we present strontium isotope data from sequentially-sampled enamel from three reindeer (Rangifer tarandus ssp.) and a single bison (Bison cf. priscus) from the late Middle Palaeolithic site of Jonzac (Chez-Pinaud), France. The results are used to investigate the ranging and migratory behaviours of these important prey species. We found that the bison had isotope values most consistent with a local range, while the three reindeer had values indicating a seasonal migration pattern. Due to the similarity of the patterning of two of the three reindeer and in conjunction with zooarchaeological results, we suggest that they may have been from the same herd, were likely killed around the same point during their seasonal round and may therefore be the product of a single hunting event or a small number of successive hunting events. The isotope analyses complement the zooarchaeological data and have allowed greater insight into the palaeoecology of these species, the palaeoenvironment, and Neanderthal site use and hunting strategies.     

Aggression and Nearest Neighbour Distances in Female Reindeer during the Rut

Data on nearest neighbours and aggressive behaviour were collected within rutting groups of semi-domesticated reindeer (Rangifer t. tarandus L.) in a 1300-ha pasture in Finnish Lapland, and wild forest reindeer (Rangifer t. fennicus Lönnb.) in a 15-ha enclosure in Central Finland. Adult cows of forest reindeer charged yearling males, yearling females and calves in that order of decreasing frequency. Semi-domesticated reindeer cows rushed yearling females more often than they did one another. One suggested influence of antagonism by cows were the observed intragroup associations of yearling females. Cows were tolerant toward their own yearling females, but not toward their own yearling males. Although no maternal aggression toward calves was found, male calves moved away more often from their mothers than did female calves during the post-rut period.     

From high voltage (300 kV) to higher voltage (420 kV) power lines: reindeer avoid construction activities

Demands for increased energy production have initiated several new high-voltage power line projects, of which hundreds of km will traverse reindeer (Rangifer tarandus tarandus) habitat in central and northern parts of Scandinavia. We investigated area use of semidomesticated reindeer in the Essand reindeer district’s summer range (Norway) in connection with a new 420 kV power line built in 2008/2009 to replace an existing 300 kV line. We used 6 years (2008–2013) of GPS telemetry data from 5 to 22 female reindeer per season. During the construction period compared to the period before and after construction, predicted probability of use decreased on average 10 % within areas 6 km from the central infrastructure for the calving period, about 12 % within 3.5 km in summer and close to 13 % within 3.5 km in autumn. In the calving period prior to construction, as well as the calving period, summer and autumn for the years after construction, use of areas close to the infrastructure did not deviate from random. Resource selection functions showed significant effects of construction work, habitat quality, elevation and aspect on the area use of reindeer. We found no support for the hypothesis that power lines have negative effects on reindeer area use, independent of associated human activity during construction. Mitigation measures should focus on both the construction period of power lines, minimizing construction work when adjacent areas are utilized by reindeer, as well as keeping human activity to a minimum during operative years.     

High genetic connectivity and introgression from domestic reindeer characterize northern Alaska caribou herds

Defining genetic populations and detecting hybridization with introduced or domestic taxa are two major concerns for the conservation of population-level diversity. We studied the genetic population structure of large, migratory caribou herds (Rangifer tarandus granti) on Alaska’s North Slope and their potential hybridization with introduced domestic reindeer (R. t. tarandus). Using a population genetics approach, we determined: (1) whether the four caribou herds could be differentiated; (2) how distance and population size appear to drive genetic population structure; and (3) how contact with reindeer has affected the genetic identity of herds. Samples from four caribou herds (n = 245) and reindeer (n = 67) were analyzed at 19 microsatellite loci. We found that North Slope caribou are isolated by distance, with no differentiation among herd pairs except for the most geographically distant herds (F _st  = 0.003, Jost’s D = 0.023; P-values < 0.001). We detected reindeer-caribou admixture in all populations except Kodiak Island, including 8 % of individuals in caribou herds and 14 % of individuals in Seward Peninsula reindeer herds. However, considering the stable or increasing trend in North Slope herds, reindeer introgression has had no apparent deleterious effect on herd demographics. Our findings indicate long-term genetic exchange among North Slope caribou herds when their ranges overlap, and suggest that herd size may influence susceptibility to reindeer introgression. As North Slope herd ranges are increasingly altered by industrial development, this study can provide a baseline for detecting potential future impacts to what are currently large, diverse, and naturally evolving herds.     

Reindeer parasites, weather and warming of the Arctic

The protostrongylid nematode Elaphostrongylus rangiferi parasitizes reindeer (Rangifer tarandus) and may cause severe disease and death. A relationship between particularly warm summers and the occurrence of the parasite has been suggested. The hypotheses of a positive relationship between the abundance of the parasite and summer warmth were tested by yearly examination of the density of larvae of E. rangiferi in faeces of reindeer in northern Norway. In addition to summer temperature, summer precipitation and reindeer density were included. The abundance of the parasite increased with increasing summer temperature. It is predicted that warming of the Arctic will increase the abundance of E. rangiferi and cause more disease and death in reindeer.     

Population trends,distribution and conservation status of semi-domesticated reindeer (Rangifer tarandus) in China

The reindeer (Rangifer tarandus tarandus) is listed as a species of “least concern” on the Red List of Threatened Species of the World Conservation Union (IUCN). It is distributed in the northern part of China and semi-domesticated in the region. Here we present a comparison of the historical and current distribution and numbers of reindeer in China, as well as an assessment of the factors limiting population growth. Based on this information we explore potential protective measures for reindeer in China.The reindeer has been herded extensively by the Ewenki people, the only people to do so in China. Reindeer in China originated from the Wenduosenlin tundra highlands of Siberia. Capable of migrating more than 3000 km/year, the reindeer moved from the highlands to the Weitimu river during the 16th century and on to the Northeast area of China in 1654. Since 1957, 600 reindeer have settled in Qiqian in Inner Mongolia, and from 1965 more than 900 reindeer migrated from there to Mangui. In response to frequent flooding and a fluctuating reindeer population in the Mangui region, in 2003 the Ewenki people and more than 260 reindeer translocated from there to Aoluguya.Currently, eight meta-populations of reindeer comprising 773 individuals inhabit in the Arctic region of China. The reindeer population peaked in the 1970s at more than 1080 animals, but declined markedly from 1998 onwards mainly as a result of inbreeding, poaching, predation and tourism. The Chinese government's interest in wildlife protection is increasing, as evidenced by the introduction of wildlife protection laws and regulations, the importation of new reindeer varieties from Russia and the joining of the Association of World Reindeer Herders. However, natural reserves and national parks have still not been established within the reindeer's range in China to conserve this fragile population.     

Summer distribution of semi-domesticated reindeer relative to a new wind-power plant

Wind-power plants (WPs) within reindeer (Rangifer tarandus tarandus) habitat may have negative effects on reindeer habitat use. Avoidance effects towards a WP were tested by comparing reindeer distributions on a peninsula where a WP was built in 2006 with a control peninsula without a WP. Distributions were measured by direct observations during construction period, and in four subsequent years, and limited faecal pellet group counts along transects before, during and after the WP construction (2005–2010). We predicted higher reindeer density in the control than the WP peninsula and at increasing distances from the WP when controlling for habitat quality. We found no avoidance effects from the WP, with significantly more reindeer in the WP than the control peninsula. Faecal pellet group data supported a lack of negative effects towards the WP after construction compared to before, while area within 100 m from the access road to the WP was avoided during the construction period and for 3 years afterwards. Reindeer avoided low-quality habitat both in the control and WP peninsulas. Our study indicates that WP development might have minor effects on habitat use if built in poor habitats, at least for semi-domestic reindeer. Our results cannot be used to infer effects of a WP built in higher-quality habitats or where large-scale movements are less restrictive than on a peninsula. Disturbance effects of human infrastructure likely are context-dependent, and management should thus be careful in planning of WPs to minimize adverse effects.     

Neandertal mobility and large-game hunting: The exploitation of reindeer during the Quina Mousterian at Chez-Pinaud Jonzac (Charente-Maritime, France)

Neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges. Equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the relationships between hunting and subsistence strategies with other components of the Neandertals’ niche, such as mobility, site use, and lithic technology. The Quina Mousterian deposits from the rockshelter site of Chez Pinaud Jonzac (Charente-Maritime, France; hereafter Jonzac) offer an excellent opportunity to pursue these issues. This paper focuses on the extensive and well-preserved skeletal remains of reindeer (Rangifer tarandus) recovered from recent excavations of the site, representing at least 18 individuals that were hunted by Neandertals during the fall through winter. Our zooarchaeological results indicate that all ages of reindeer were hunted but adult individuals predominate. No bias is evident in the comparable frequencies of males and females. These prey were butchered on-site, with abundant evidence of meat filleting and marrow exploitation. In the excavated sample, the absence of hearths and the almost complete lack of burned bones or stones suggest that Neandertals were not using fire to assist with processing the reindeer carcasses. The zooarchaeological results presented here indicate that reindeer were hunted during a restricted window of time when they were seasonally abundant in the local area near Jonzac. Taken together with the lithic industry based on bifacial elements, the evidence is consistent with a pattern of site use by highly mobile hunter-gatherers making frequent, short-term visits. Ongoing research at Jonzac and other Quina Mousterian localities will contribute to a better understanding of Neandertal behavior during cold climate phases.     

Experimental oral transmission of chronic wasting disease to reindeer (Rangifer tarandus tarandus)

Chronic wasting disease (CWD), a transmissible spongiform encephalopathy of cervids, remains prevalent in North American elk, white-tailed deer and mule deer. A natural case of CWD in reindeer (Rangifer tarandus tarandus) has not been reported despite potential habitat overlap with CWD-infected deer or elk herds. This study investigates the experimental transmission of CWD from elk or white-tailed deer to reindeer by the oral route of inoculation. Ante-mortem testing of the three reindeer exposed to CWD from white-tailed deer identified the accumulation of pathological PrP (PrP(CWD)) in the recto-anal mucosa associated lymphoid tissue (RAMALT) of two reindeer at 13.4 months post-inoculation. Terminal CWD occurred in the two RAMALT-positive reindeer at 18.5 and 20 months post-inoculation while one other reindeer in the white-tailed deer CWD inoculum group and none of the 3 reindeer exposed to elk CWD developed disease. Tissue distribution analysis of PrP(CWD) in CWD-affected reindeer revealed widespread deposition in central and peripheral nervous systems, lymphoreticular tissues, the gastrointestinal tract, neuroendocrine tissues and cardiac muscle. Analysis of prion protein gene (PRNP) sequences in the 6 reindeer identified polymorphisms at residues 2 (V/M), 129 (G/S), 138 (S/N) and 169 (V/M). These findings demonstrate that (i) a sub-population of reindeer are susceptible to CWD by oral inoculation implicating the potential for transmission to other Rangifer species, and (ii) certain reindeer PRNP polymorphisms may be protective against CWD infection.     

Dental microwear textural analyses to track feeding ecology of reindeer: a comparison of two contrasting populations in Norway

Using dental microwear textural analysis, we aim to explore variations in feeding habits within and between wild reindeer populations (Rangifer tarandus) in Norway. Specimens from the two wild reindeer areas of Knutshø and Hardangervidda were hunted between late August and the end of September 2014, and found to share very similar topological and ecological characteristics (continental alpine tundra). Despite the homogenisation of plant resources at the end of the snow-free period, dental microwear textural analysis on cheek teeth indicates differences in feeding behaviors between the two populations, in that reindeer from Knutshø might display a more “grazing” signal than the ones from Hardangervidda. Such differences in dental microwear textures reflect the differences in feeding habits that could be linked with differences in population density of reindeer. The Knutshø reindeer density is lower than at Hardangervidda, where primary food resources like grasses, sedges, and lichens are less abundant. Analyses also show the differences in gender for the population of Knutshø, but more studies are necessary for the clearest interpretation of data. Our results emphasize the need to explore local variations at a monthly time frame in order to better assess the versatility of dental microwear textures before applying this kind of analysis to either extant or extinct multi-population scales.     

Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.