Persistence of vigilance and flight response behaviour in wild reindeer with varying domestic ancestry

Knowledge about changes in behavioural traits related to wildness and tameness is for most mammals lacking, despite the increased trend of using domestic stock to re‐establish wild populations into historical ranges. To test for persistence of behavioural traits of wild reindeer (Rangifer tarandus L.) exposed to hunting, we sampled DNA, vigilance and flight responses in wild reindeer herds with varying domestic ancestry. Analyses of 14 DNA microsatellite loci revealed a dichotomous main genetic structure reflecting their native origin, with the Rondane reindeer genetically different from the others and with least differentiation towards the Hardangervidda reindeer. The genetic clustering of the reindeer in Norefjell‐Reinsjøfjell, Ottadalen and Forollhogna, together with domestic reindeer, supports a predominant domestic origin of these herds. Despite extensive hunting in all herds, the behavioural measures indicate increasing vigilance, alert and flight responses with increasing genetic dissimilarity with domestic herds. Vigilance frequency and time spent vigilant were higher in Rondane compared to Hardangervidda, which again were higher than herds with a domestic origin. We conclude that previous domestication has preserved a hard wired behavioural trait in some reindeer herds exhibiting less fright responses towards humans that extensive hunting has, but only slightly, altered. This brings novel and relevant knowledge to discussions about genetic diversity of wildlife in general and wild reindeer herds in Norway in specific.     

Adults only. Reindeer hunting at the middle palaeolithic site salzgitter lebenstedt, northern Germany

The Middle Palaeolithic site Salzgitter Lebenstedt (northern Germany), excavated in 1952, is well known because of its well-preserved faunal remains, dominated by adult reindeer (Rangifer tarandus). The archaeological assemblage accumulated in an arctic setting in an earlier part of the last (Weichsel) glacial (OIS5-3). The site is remarkable because of the presence of unique Middle Palaeolithic bone tools and the occurrence of the northernmost Neanderthal remains, but this paper focuses on an analysis of its reindeer assemblage. The results indicate autumn hunting of reindeer by Middle Palaeolithic hominids. After the hunt, carcasses were butchered and in subsequent marrow processing of the bones a selection against young and sub-adult animals occurred. Adults were clearly preferred, and from their bones, again, poorer marrow bones were neglected. This focus on primeness of resources has been documented in other domains of Neanderthal behaviour, but Salzgitter Lebenstedt is the best example yet known in terms of systematic and routinized processing of game. The Salzgitter Lebenstedt assemblage displays some remarkable similarities to the Late Glacial reindeer assemblages from the Ahrensburg tunnel valley sites. The subsequent review of the evidence on subsistence strategies from earlier periods of the European Palaeolithic shows that hunting of large mammals may have been a part of the behavioural repertoire of the Middle Pleistocene occupants of Europe from the earliest occupation onwards. At the same time, it is suggested that these early hunting strategies were incorporated in ways of moving through landscapes ("settlement systems") which were different from what we know from the middle parts of the Upper Palaeolithic onwards.     

Effects of Hunting on Response Behaviors of Wild Reindeer

ABSTRACT Because wild reindeer (Rangifer tarandus) are hunted in southern Norway, reindeer may perceive all recreationists as threats. Potential adverse effects of hunting on reindeer behavior may be exacerbated by other forms of recreation because the number of skiers and hikers in areas inhabited by reindeer has also increased. The Norefjell-Reinsjøfjell wild reindeer area is used extensively for recreation and tourism. Reindeer hunting was introduced in the area in 1992, and harvest rate has been stable at about 38% of winter herd size. We recorded behavioral responses of reindeer to a person approaching directly on foot or skis during 1992 and again in 2002–2006. Compared to 1992, flight-initiation distance increased and fewer groups assessed the observer before taking flight during 2002–2006. In winter, when reindeer are usually comparably more vigilant than in other seasons, flight-initiation distance increased from only 60 m to 115 m and escape distance decreased from 400 m to 210 m. Neither alert distance, calf carcass weights (23.6 ± 0.7 [SE] kg to 22.4 ± 0.2 kg), nor reindeer herd size (661 ± 73 to 579 ± 15) changed during the 15 years of our study. Reindeer appeared to habituate to the observer because they initiated flight at shorter distances as the number of approaches on the same day increased. In Norefjell-Reinsjøfjell, encounters with a person on foot or skis did not result in behavioral responses likely to entail substantial energy costs for reindeer; therefore, hunting at current levels appears compatible with other recreational activities.     

Heaps of fascinating fragments: reconstructing a medieval reindeer population based on bone middens from the Hardangervidda high mountain plateau,Norway

The results of a detailed morphological and pathological study on reindeer bones (Rangifer tarandus) from four medieval hunting stations on Hardangervidda are presented. As intensive marrow collecting left almost no bones intact, traditional sexing methods could only sparsely be applied. Alternative methods had to be explored to successfully assign the fragments to a sex. Employing linear discriminant analysis (LDA) on early‐ and non‐fusing skeletal elements, I have shown that (incomplete) calcanei, metapodia and phalanges I and II can be used successfully to assign a specimen to a sex and should no longer be excluded from osteometric analyses. Differences in the demographic compositions of the taphocoenoses lead to the assumption that hunters in the 11th century AD targeted large reindeer bucks, while at the 13th‐century sites, the complete biocoenose is represented, albeit in a different ratio. There seems to have been a shift in hunting technique: from selective hunting to mass hunting. Size wise, the reindeer from Hardangervidda were smaller than reindeer from contemporary assemblages from the Dovre area (central Norway), a population that is genetically different. Few pathologically affected bones were encountered in the material, but some cases of infections, bone lesions and a progressed osteosarcoma are described.     

Genetic analyses reveal independent domestication origins of Eurasian reindeer

Although there is little doubt that the domestication of mammals was instrumental for the modernization of human societies, even basic features of the path towards domestication remain largely unresolved for many species. Reindeer are considered to be in the early phase of domestication with wild and domestic herds still coexisting widely across Eurasia. This provides a unique model system for understanding how the early domestication process may have taken place. We analysed mitochondrial sequences and nuclear microsatellites in domestic and wild herds throughout Eurasia to address the origin of reindeer herding and domestication history. Our data demonstrate independent origins of domestic reindeer in Russia and Fennoscandia. This implies that the Saami people of Fennoscandia domesticated their own reindeer independently of the indigenous cultures in western Russia. We also found that augmentation of local reindeer herds by crossing with wild animals has been common. However, some wild reindeer populations have not contributed to the domestic gene pool, suggesting variation in domestication potential among populations. These differences may explain why geographically isolated indigenous groups have been able to make the technological shift from mobile hunting to large-scale reindeer pastoralism independently.     

Effects of weather and hunting on wild reindeer population dynamics in Hardangervidda National Park

Wild reindeer have a range that extends across the circumpolar region. In the last few decades, however, populations of wild reindeer have been on the decline. The reasons for these declines are poorly understood, but are suggested to be linked to both local and global climatic factors, disease, and human interference. Hardangervidda plateau in Norway is home to the largest wild reindeer population in Europe, and is at the southern end of its European range. This population is therefore of particular importance, particularly in the light of climate change. We investigated how weather and hunting have affected the wild reindeer population in Hardangervidda over the last two decades. Our findings suggest that the wild reindeer population in Hardangervidda is most affected by winter temperature and hunting, where colder temperatures and lower harvest rates typically result in higher growth rates. We did not find significant evidence for linear density dependence. Our results show trends across Hardangervidda, and give an indication of how region-wide weather and hunting pressure can affect the wild reindeer population. As new data emerge, future investigations should look into the existence and nature of density dependence and the influence of other weather and human disturbance related factors.     

Museum specimens reveal changes in the population structure of northern Fennoscandian domestic reindeer in the past one hundred years

Traditional reindeer herding of northern Fennoscandia has been based on seasonal movements independent of national borders. At the beginning of the 19th century, these yearly movements of reindeer were excessive, but during that century the borders between the Fennoscandian countries were closed. By analysing a 190‐base pair fragment of the mitochondrial DNA control region in 79 museum samples, we show that the reindeer of northern Fennoscandia were one homogenous population shortly after the national borders were closed. However, anthropogenic activity has effectively ended genetic exchange within northern Fennoscandia and has made the reindeer population within this region heterogeneous. Genetic input of eastern origin is also suggested within the extant Russian reindeer of the Kola Peninsula.     

Adaptation and niche construction in human prehistory: a case study from the southern Scandinavian Late Glacial

The niche construction model postulates that human bio-social evolution is composed of three inheritance domains, genetic, cultural and ecological, linked by feedback selection. This paper argues that many kinds of archaeological data can serve as proxies for human niche construction processes, and presents a method for investigating specific niche construction hypotheses. To illustrate this method, the repeated emergence of specialized reindeer (Rangifer tarandus) hunting/herding economies during the Late Palaeolithic (ca 14.7-11.5 kyr BP) in southern Scandinavia is analysed from a niche construction/triple-inheritance perspective. This economic relationship resulted in the eventual domestication of Rangifer. The hypothesis of whether domestication was achieved as early as the Late Palaeolithic, and whether this required the use of domesticated dogs (Canis familiaris) as hunting, herding or transport aids, is tested via a comparative analysis using material culture-based phylogenies and ecological datasets in relation to demographic/genetic proxies. Only weak evidence for sustained niche construction behaviours by prehistoric hunter-gatherer in southern Scandinavia is found, but this study nonetheless provides interesting insights into the likely processes of dog and reindeer domestication, and into processes of adaptation in Late Glacial foragers.     

Genetic diversity of wild reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>) of Taimyr: Analysis of polymorphism of the control region of mitochondrial DNA

Based on an analysis of the polymorphism of nucleotide sequences (n = 111) of the mtDNA control region (left domain), the genetic diversity of the largest population of wild reindeer Rangifer tarandus in Eurasia, which inhabits Taimyr peninsula, was studied. High levels of haplotype (H) and nucleotide (π) diversity (0.987 and 0.018, respectively) were revealed, which indicate the long existence and the sufficiently stable growth of this population. The absence of long periods of abrupt decrease in the number of the Taimyr wild reindeer population and/or facts of formation of its genetic diversity as a result of mixing of genetically distant conspecific populations is supported by the data on the pattern of mismatch distributions and the topology of the phylogenetic tree. The low level of genetic differences between reindeer from the western, central, and eastern groups reflects their common origin and close relationship.     

Sacrifice as the ideal hunt: a cosmological explanation for the origin of reindeer domestication

The Siberian Northeast shows striking parallels between the cosmologies of hunters and reindeer herders. What may this tell us about the transformation from hunting to pastoralism? This article argues for a structural identity between hunting and sacrifice, and for the domestication of the reindeer as the result of hunters' efforts to use sacrifice to control the accidental variables of the hunt. Hunters can practise their ethos of ‘trust’ with prey only through highly controlled ritual enactments. We describe two: the famous bear festival of the Amur Gulf region and the consecrated reindeer of the Eveny. Both express the same overall logic by which sacrifice functions as an ideal hunt. The animal is involved in a relation not of domination but of trust, while also undergoing a process of taming. We therefore suggest that the reindeer's domestication may be based not only on ecological or economic adaptations, but also on cosmology.     

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.     

Genetic diversity of wild reindeer <Emphasis Type="Italic">Rangifer tarandus</Emphasis> L. from the eastern part of the Kola Peninsula: Polymorphism of the mtDNA control region

Based on the results of the analysis of nucleotide sequence polymorphism in the hypervariable fragment (left domain) of the mtDNA control region (D loop), the genetic diversity of wild reindeer from the eastern part of the Kola Peninsula (Murmansk oblast, Terskii region) was studied. Low values of the genetic diversity indices for Eurasian wild reindeer were detected. The effect of domestic reindeer breeding on the development of genetic diversity of the wild reindeer population in the studied region (indicating a low degree of hybridization of domesticated and wild reindeer) was estimated.     

Foraging strategy and predator avoidance behaviour: an intraspecific approach

Relationships between predator avoidance behaviour (scanning and flocking) and foraging were studied in Calidris alpina, to test predictions regarding the effect of foraging techniques on such behaviours. The scanning hypothesis predicts that individuals with a tactile hunting technique and individuals with a visual hunting technique (both continuous searchers) do not differ in any variable related to scanning behaviour. The flocking hypothesis predicts that visually hunting individuals witl tend to form smaller flocks than tactile-foraging individuals. The two continuous feeding strategies did not differ among individuals in vigilance rate, nor in vigilance time or mean scan duration. However, with respect to flocking behaviour, visual foragers differed from tactile foragers in foraging flock size. The relationships between flocking behaviour and foraging strategy are discussed. The pattern found at the intraspecific level are the same as those found at interspecific level.     

Polymorphism of the mtDNA Control Region in Reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>) from the Mainland of the Northeastern Part of European Russia

Based on an analysis of the polymorphism of nucleotide sequences (n = 103) of the hypervariable fragment of the mtDNA control region (D-loop), the genetic diversity of wild and domestic reindeer from the mainland of the European Northeast of Russia (eastern parts of Arkhangelsk oblast, Komi Republic, and the Nenets Autonomous Okrug) was studied. Relatively high values of genetic diversity indices were detected. Phylogenetic analysis showed close relationships between reindeer of the study region and those of the tundra zone of Siberia. The influence of domestic reindeer on the formation of the genetic diversity of the wild reindeer was considered as negligible. Genetic lines belonging to an extinct group of forest reindeer from the Nizhny Novgorod region were found among the modern groups of reindeer from the European Northeast of Russia.     

Human Disturbances,Habitat Characteristics and Social Environment Generate Sex-Specific Responses in Vigilance of Mediterranean Mouflon

In prey species, vigilance is an important part of the decision making process related to predation risk effects. Therefore, understanding the mechanisms shaping vigilance behavior provides relevant insights on factors influencing individual fitness. We investigated the role of extrinsic and intrinsic factors on vigilance behavior in Mediterranean mouflon (Ovis gmelini musimon×Ovis sp.) in a study site spatially and temporally contrasted in human pressures. Both sexes were less vigilant in the wildlife reserve compared to surrounding unprotected areas, except for males during the hunting period. During this period, males tended to be less strictly restricted to the reserve than females what might lead to a pervasive effect of hunting within the protected area, resulting in an increase in male vigilance. It might also be a rutting effect that did not occur in unprotected areas because males vigilance was already maximal in response to human disturbances. In both sexes, yearlings were less vigilant than adults, probably because they traded off vigilance for learning and energy acquisition and/or because they relied on adult experience present in the group. Similarly, non-reproductive females benefited of the vigilance effort provided by reproductive females when belonging to the same group. However, in the absence of reproductive females, non-reproductive females were as vigilant as reproductive females. Increasing group size was only found to reduce vigilance in females (up to 17.5%), not in males. We also showed sex-specific responses to habitat characteristics. Females increased their vigilance when habitat visibility decreased (up to 13.8%) whereas males increased their vigilance when feeding on low quality sites, i.e., when concomitant increase in chewing time can be devoted to vigilance with limited costs. Our global approach was able to disentangle the sex-specific sources of variation in mouflon vigilance and stressed the importance of reserves in managing and conserving wild sheep populations.     

Diet shift of a facultative scavenger, the wolverine, following recolonization of wolves

1. Wolves Canis lupus L. recolonized the boreal forests in the southern part of the Scandinavian peninsula during the late 1990s, but so far there has been little attention to its effect on ecosystem functioning. Wolf predation increases the availability of carcasses of large prey, especially moose Alces alces L., which may lead in turn to a diet switch in facultative scavengers such as the wolverine Gulo gulo L. 2. Using 459 wolverine scats collected during winter-spring 2001-04 for DNA identity and dietary contents, we compared diet inside and outside wolf territories while controlling for potential confounding factors, such as prey density. We tested the hypothesis that wolverine diet shifted towards moose in the presence of wolves, while taking into account possible sexual segregation between the sexes. Occurrence of reindeer, moose and small prey was modelled against explanatory covariates using logistic mixed-effects models. Furthermore, we compared diet composition and breadth among habitats and sexes. 3. Occurrence of reindeer, moose and small prey in the diet varied with prey availability and habitat. As expected, diet contained more moose and less reindeer and small prey in the presence of wolves. Their diet in tundra consisted of 40% reindeer Rangifer tarandus L., 39% moose and 9% rodents. In forest with wolf, their diet shifted to 76% moose, 18% reindeer and 5% rodents; compared to 42% moose, 32% reindeer and 15% rodents in forest without wolf. This diet switch could not be explained by higher moose density in wolf territories. Female diet consisted of more small prey than for males, but there was a tendency for females to use the highly available moose carrion opportunistically and to hunt less on small prey within wolf territories. 4. Our study highlights how wolves increase scavenging opportunities for wolverines, and how sexual differences in diet may also apply to large scavengers. Due to their more restricted home range, female wolverines are forced to rely more on hunting small prey. The relatively high occurrence of wolf kills, however, forms an important food source to wolverines in this area. The recolonization of wolves may therefore have contributed to the consequent recolonization of wolverines into the same area.     

Behavioural Responses of European Roe Deer to Temporal Variation in Predation Risk

In natural environments, predation risk varies over time. The risk allocation hypothesis predicts that prey is expected to adjust key anti‐predator behaviours such as vigilance to temporal variation in risk. We tested the predictions of the risk allocation hypothesis in a natural environment where both a species‐rich natural predator community and human hunters are abundant and where the differences in seasonal and circadian activity between natural and anthropogenic predators provided a unique opportunity to quantify the contributions of different predator classes to anti‐predator behaviour. Whereas natural predators were expected to show similar levels of activity throughout the seasons, hunter activity was high during the daytime during a clearly defined hunting season. According to the risk allocation hypothesis, vigilance should then be higher during the hunting season and during daytime hours than during the non‐hunting season and night‐time hours. Roe deer (Capreolus capreolus) on the edge of Bia?owie?a Primeval Forest in Eastern Poland displayed vigilance behaviour consistent with these predictions. The behavioural response of roe deer to temporarily varying predation risks emphasises the behavioural plasticity of this species and suggests that future studies of anti‐predator behaviour need to incorporate circadian variation in predation pressure as well as risk gradients of both natural and anthropogenic predators.     

Population genetics of the native caribou (Rangifer tarandus groenlandicus) and the semi-domestic reindeer (Rangifer tarandus tarandus) in Southwestern Greenland: Evidence of introgression

Over the past centuries the native caribou ofWest Greenland has gone through extensive population size fluctuations, with reductionsas great as 90% in less than 20 years.Norwegian semi-domestic reindeer wereintroduced to the Nuuk area in 1952 because ofthe small number of caribou in Greenland.Although the reindeer and caribou wereinitially kept separated, mixing has occurredsince the 1970's. We investigated the genotypicstructure of caribou and reindeer in South-westGreenland, using five polymorphicmicrosatellite markers isolated from cattle,sheep, goat and red deer. A total of ninetysamples were collected, which included samplesfrom caribou of four different regions andsamples from two different reindeer herds.Based on the genetic variation of the fivemarkers, our results shows that the caribou andthe reindeer populations in the six regionssampled are genetically differentiated withineach group and the two subspecies aredifferentiated from each other. A likelyexplanation for the genetic isolation of thepopulations investigated is that naturalbarriers (glaciers and wide fjords) exists inthe area. Furthermore we found that introducedNorwegian domestic reindeer hybridized with thenative Greenlandic caribou in two areasneighbouring Nuuk.     

Microsatellite DNA evidence for genetic drift and philopatry in Svalbard reindeer

Mainland populations of Arctic reindeer and caribou Rangifer tarandus often undergo extensive movements, whereas populations on islands tend to be isolated and sedentary. To characterize the genetic consequences of this difference, levels of genetic diversity and subdivision of Svalbard reindeer (R. t. platyrhynchus) from two adjacent areas on Nordenskjiöldland, Spitsbergen were estimated using data from up to 14 microsatellites. The mean number of alleles per locus in Svalbard reindeer was 2.4 and mean expected heterozygosity per locus was 0.36. The latter value was significantly lower than in Canadian caribou and Norwegian reindeer but higher than in some other cervid species. Large samples of females (n = 743) and small samples of males (n = 38) from two sites ≈ 45 km apart showed genetic subdivision, which could be due to local population fluctuations or limited gene flow. To our knowledge, this is the first study to report significant differentiation at microsatellite loci in Rangifer at such short geographical distances. Neither population showed genetic evidence for recent population bottlenecks when loci unbiased with respect to heterozygosity were analysed. In contrast, false signals of a recent bottleneck were detected when loci upwardly biased with respect to heterozygosity were analysed. Thus, Svalbard reindeer appeared to conform to the paradigm of island populations made genetically depauperate by genetic drift.     

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.