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.     

Farmers without borders—genetic structuring in century old barley (Hordeum vulgare)

The geographic distribution of genetic diversity can reveal the evolutionary history of a species. For crop plants, phylogeographic patterns also indicate how seed has been exchanged and spread in agrarian communities. Such patterns are, however, easily blurred by the intense seed trade, plant improvement and even genebank conservation during the twentieth century, and discerning fine-scale phylogeographic patterns is thus particularly challenging. Using historical crop specimens, these problems are circumvented and we show here how high-throughput genotyping of historical nineteenth century crop specimens can reveal detailed geographic population structure. Thirty-one historical and nine extant accessions of North European landrace barley (Hordeum vulgare L.), in total 231 individuals, were genotyped on a 384 single nucleotide polymorphism assay. The historical material shows constant high levels of within-accession diversity, whereas the extant accessions show more varying levels of diversity and a higher degree of total genotype sharing. Structure, discriminant analysis of principal components and principal component analysis cluster the accessions in latitudinal groups across country borders in Finland, Norway and Sweden. F_ST statistics indicate strong differentiation between accessions from southern Fennoscandia and accessions from central or northern Fennoscandia, and less differentiation between central and northern accessions. These findings are discussed in the context of contrasting historical records on intense within-country south to north seed movement. Our results suggest that although seeds were traded long distances, long-term cultivation has instead been of locally available, possibly better adapted, genotypes.     

High genomic diversity in the bank vole at the northern apex of a range expansion: The role of multiple colonizations and end‐glacial refugia

The history of repeated northern glacial cycling and southern climatic stability has long dominated explanations for how genetic diversity is distributed within temperate species in Eurasia and North America. However, growing evidence indicates the importance of cryptic refugia for northern colonization dynamics. An important geographic region to assess this is Fennoscandia, where recolonization at the end of the last glaciation was restricted to specific routes and temporal windows. We used genomic data to analyse genetic diversity and colonization history of the bank vole (Myodes glareolus) throughout Europe (>800 samples) with Fennoscandia as the northern apex. We inferred that bank voles colonized Fennoscandia multiple times by two different routes; with three separate colonizations via a southern land‐bridge route deriving from a “Carpathian” glacial refugium and one via a north‐eastern route from an “Eastern” glacial refugium near the Ural Mountains. Clustering of genome‐wide SNPs revealed high diversity in Fennoscandia, with eight genomic clusters: three of Carpathian origin and five Eastern. Time estimates revealed that the first of the Carpathian colonizations occurred before the Younger Dryas (YD), meaning that the first colonists survived the YD in Fennoscandia. Results also indicated that introgression between bank and northern red‐backed voles (Myodes rutilus) took place in Fennoscandia just after end‐glacial colonization. Therefore, multiple colonizations from the same and different cryptic refugia, temporal and spatial separations and interspecific introgression have shaped bank vole genetic variability in Fennoscandia. Together, these processes drive high genetic diversity at the apex of the northern expansion in this emerging model species.     

Cutting of lichen trees: a survival strategy used before the 20th century in northern Sweden

Sami reindeer herders in northern Fennoscandia have historically cut trees with high quantities of arboreal lichens to feed their animals during harsh winters. In this study we analysed temporal and spatial patterns of lichen-tree cuttings of Scots pine in a remote forest landscape in northern Sweden. We wanted to test the hypothesis that repeated cutting of such trees may have had long-lasting effects on the forest ecosystem that are still discernable today. Our analyses included an inventory of remaining lichen-tree stumps, and a field survey of forest variables to relate structural variation within the forest to these cuttings. We used dendrochronological methods to date stumps and reconstruct forest structure. Our results show that more than 7,000 trees were cut for this purpose over a period of almost 300 years within an area covering 6 km². Lichen-tree cutting was most intense during the late 18th century when ca. 600–1,000 trees were cut during each decade. We did not detect any significant relationship between past lichen-tree cutting and present small-scale forest heterogeneity, and suggest that this might be due to the small size of the cut trees and the long time elapsed since most of the cuttings. We conclude that vestiges such as stumps from lichen-tree cutting are highly valuable for our understanding of past land use. Furthermore, since they are located in an unlogged and very old forest, this highlights the need to acknowledge the interconnectedness between ecological and cultural values of such forests and the need to identify and protect them.     

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.     

Diet of the golden eagle Aquila chrysaetos during the breeding season in Sweden

During a five-year period, 1975–1979, a total of 2881 prey individuals of 65 prey species were collected at 162 golden eagle nests from northern Sweden and from the island of Gotland. In northern Sweden birds are taken in higher numbers than mammals but calculated as weight the two categories are of equal importance, The main prey during the breeding season are capercaillie, black grouse, willow grouse, ptarmigan, mountain hare and reindeer fawns which together form 91% of the total food biomass. The capercaillie and the black grouse are taken more in the southern part of the coniferous region than in the northern. In contrast, in northern areas, reindeer fawns are more preyed upon than in the South, Ptarmigan and willow grouse are the most commonly captured prey species in mountain areas. The total number of reindeer fawns taken (dead and/or alive) by the Swedish golden eagle population during one summer is estimated at 600 individuals. On Gotland the golden eagles take mammals more often than in its northern distribution area. Rabbit and hedgehog arc the most important species.     

Sami Resource Utilization and Site Selection: Historical Harvesting of Inner Bark in Northern Sweden

Native Sami people have lived for thousands of years in northern Fennoscandia, subsisting on reindeer herding, hunting, fishing and gathering of wild plants. Harvesting inner bark from Scots pine (Pinus Sylvestris L.) was a common practice and traces of this resource use are still evident in old forests. In this study we analyzed trees with bark-peelings in a nature reserve in northernmost Sweden to assess the magnitude and spatiotemporal patterns of inner-bark harvesting. We found that the same inner-bark harvest locale was used repeatedly, in nearly every decade from the late 16th to the late 19th century and that harvest levels varied greatly over time. The harvest levels we observed were surprisingly low, in comparison to levels suggested in other studies, and indicate that inner-bark was used as a supplementary health food during early spring when game meat was naturally lean, rather than being used as bulk food year-round.     

Synchronous short-term population fluctuations of some birds and mammals in Fennoscandia - occurrence and distribution

We examined population fluctuations of the voles Microtus agrestis L. and Clethrionomys glareolus Schreb. by biannual trapping; black grouse Tetrao tetrix L., mountain hare Lepus timidus L., and red fox Vulpes vulpes L. by using bag records from all over Sweden, questionnaires from south-central Sweden, and population indices from Grimsö Wildlife Research Area in south-central Sweden. Synchronous population fluctuations between voles as a group and the other species conformed to a 3–4 year periodic pattern in both autumn and spring populations only in northern and central Sweden (the boreal forest region). Spring populations of boreal forest grouse, hare, and probably also fox, lagged one year behind the voles. The northerly areas also formed geographical units of co-fluctuation within each game species. Using our own data as well as reviewing previous studies in Fennoscandia, we conclude that synchronized 3–4 year population fluctuations of voles and medium-sized herbivores are confined to the central and northern part of Fennoscandia, although voles may exhibit short-term population fluctuations further south. The synchronizing link between the herbivores could be (1) food plant quality and/or quantity and/or (2) predation. We could not reject either of these two plausible mechanisms as a cause of interspecifically synchronous fluctuations.     

High-resolution palynology reveals the land use history of a Sami <Emphasis Type="Italic">renvall</Emphasis> in northern Sweden

The limited availability of historical and archaeological evidence means that much is still unknown about the development of Sami reindeer herding in Fennoscandia in both the recent and more distant past. To address this problem, high-resolution palynological analyses, ¹⁴C and ²¹⁰Pb dating were undertaken on two adjacent (<25 m apart) peat profiles collected at a recently abandoned reindeer gathering pen (renvall) near Jokkmokk (~66.6°N, 19.8°E) in the boreal forest of northern Sweden. The aim was to assess the impact of Sami reindeer herding on the local environment through a study of pollen, coprophilous fungal spores, microscopic charcoal and sedimentology. The samples collected from within an annex to the renvall indicate cycles of use and abandonment of the pen on a multi-decadal timescale between ~ad 1800–2008, most obviously in the coprophilous fungal spore archive. The pattern and timing of these cycles confirm events previously known only from oral histories. Although the local pollen assemblage zones associated with the phasing of activity were reproducible in a second peat core beyond the boundary of the renvall, the coprophilous fungal spore signal in this paired profile was much less distinctive, possibly due to the typically shorter dispersal distances for these microfossils in comparison to pollen grains.     

Owl winter irruptions as an indicator of small mammal population cycles in the boreal forest of eastern North America

Contrary to what is observed in Fennoscandia, it seems to be widely accepted that small mammals do not exhibit multi-annual population cycles in the boreal forest of North America. However, in the last thirty years, irruptions of vole predators such as owls have been reported by ornithologists south of the North American boreal forest. While such southerly irruptions have been associated in Fennoscandia with periods of low abundance of small mammals within their usual distribution range, their possible cyclic nature and their relationships to fluctuations in vole densities at northern latitudes has not yet been demonstrated in North America. With information collected from existing data-bases, we examined the presence of cycles in small mammals and their main avian predators by using temporal autocorrelation analyses. Winter invasions of boreal owls ( Aegolius funereus ) were periodic, with a 4-yr cycle in Québec. Populations of one species of small mammal, the red-backed vole ( Clethrionomys gapperi ), fluctuated periodically in boreal forests of Québec (north to 48°N). Boreal owls show invasion cycles which correspond to years of low density of red-backed voles, the main food item for this owl species. In addition, winter observations of northern hawk owls ( Surnia ulula ) and great gray owls ( Strix nebulosa ) south of their usual range increased in years of low density of red-backed voles. Our results suggest that a 4-yr population cycle exists in the eastern boreal forest of North America for voles and owls, which is very similar to the one observed in Fennoscandia.     

Identification of semi-domesticated reindeer (Rangifer tarandus tarandus,Linnaeus 1758) and wild forest reindeer (R.t. fennicus,Lönnberg 1909) from postcranial skeletal measurements

Reindeer bones are common finds from archaeological sites from prehistoric and historic Fennoscandia. The interpretation of the reindeer bone finds, however, is often hindered by the difficulty to separate the different subspecies of reindeer using the postcranial skeletal morphology. In this paper, skeletal measurements of modern semi-domesticated reindeer and wild forest reindeer are explored with multivariate statistical methods in order to find suitable methods for subspecies identification. The results are then applied to archaeological reindeer bone finds from Northern Finland and archaeological implications of the results are discussed.     

High degree of population subdivision in a widespread amphibian

In general, amphibians are known to exhibit a higher degree of population subdivision than any other major animal taxa, but large-scale population genetic surveys of widely distributed species are still scarce, especially in the Eurasian continent. Using microsatellite markers and mitochondrial DNA sequences, we investigated the large-scale population genetic structure of the common frog (Rana temporaria)--one of the most widespread amphibians of the Palearctic region. Analyses of cytochrome b sequences revealed evidence for two distinct lineages inhabiting western and eastern parts of Europe. The separation of these lineages c. 700,000 years ago may have been induced by the onset of the Middle Pleistocene continental glaciations. Analyses of the variability of microsatellite loci within each of the clades revealed evidence for evolution of a high degree of population subdivision (FST approximately 0.23) even in northern Fennoscandia, colonized less than 10,000 years ago. The high level of substructuring is puzzling in the face of an apparently high dispersal capacity, as evidenced by the rather rapid recolonization of northern Europe. This suggests that processes other than restricted dispersal capacity need to be explored as explanations for the high degree of population subdivision in amphibians. The colonization of northern Europe has been accompanied by loss of genetic variability as evidenced by decreasing levels of intrapopulational genetic variability in microsatellite loci from south to north across Europe.     

The arctic fox Alopex lagopus in Fennoscandia: a victim of human-induced changes in interspecific competition and predation?

After a marked decline at the beginning of the 1900s, the arctic fox Alopex lagopus population in Fennoscandia has remained at a very low level. We suggest that the main cause for the population crash was winter starvation caused by (1) over-hunting of reindeer Rangifer tarandus populations, and thus reduced carcass availability in the mountains, and (2) increased interspecific competition for these carcasses because of increased invasion of red foxes Vulpes vulpes from lower altitudes. The failure of arctic fox populations to recover, despite increasing reindeer populations in the mid 1900s, can be explained by a concurrent strong increase in red fox numbers. Analyses of countywide hunting statistics from Norway 1891–1920 suggest that there actually was an increase in red fox numbers in the period of arctic fox decline, and that the increase in reindeer populations from the 1920s to the 1950s was accompanied by a new increase in red fox numbers. We conclude that restoring arctic fox populations most likely will require a substantial and lasting reduction of red fox populations.     

Landscape Agency and Evenki-Iakut Reindeer Husbandry Along the Zhuia River,Eastern Siberia

This interdisciplinary study applies a series of environmental tests developed in Fennoscandia to elaborate the patterns created by contemporary and past reindeer husbandry along the Zhuia River, Bodaibo district, Irkutsk oblast’, Russian Federation. We successfully used pollen and fungal spore analysis to document the long-term use of one site by ungulates–although it remains unclear if these animals were ‘domestic’ reindeer or not. The date of occupation could go as far back as the fourteenth century. The on-site phosphate analysis, attempted for the first time in Eastern Siberia, proved a useful tool for locating the sites of animal action although failed to specify the boundaries of that action. It did emphasise the importance of accounting for the agency of wind in ordering reindeer behaviour. Finally, the combined phosphate, botanical, and pollen work documented a history of succession of types of land-use from the hunting of Rangifer, to holding Rangifer, to the maintenance of meadows for horses or cattle, to the formation of cereal crops and vegetable patches. The combined use of these methods and a discussion of the ambiguities they produced suggests that they are best employed to find distinctive sites in the landscape which attract both people and animals and are less effective in documenting a Euro-American vision of trust or domination in human-animal relations.     

Population cycles in voles and lemmings: state of the science and future directions

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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.     

Diet overlap among ruminants in Fennoscandia

Information on overlap in resource use is central to understanding of interspecific exploitation competition and resource partitioning. Despite this, measures of diet overlap among northern ruminants in Fennoscandia is limited to one earlier study (reindeer and sheep). Diet overlap between sympatric moose and roe deer calculated with Schoener’s index was 20.7% and 33.6% during summer (data from one area) and winter (data from two areas), respectively, whereas average diet overlap between moose and red deer was 32.0% during winter (data from four areas). Diet overlap between a coastal island population of red deer and sheep was 59.3% during summer and 63.9% during winter. Summer diet overlap between a sheep and a goat population and a sheep and a reindeer population calculated with data on main types of forage plants was 77.0% and 55.1%, respectively. However, overlap calculated with main plant groups was sometimes considerably higher than when calculated for individual forage species. Neither difference in feeding type nor body mass successfully predicted diet overlap between species pairs (n=9), although there tended to be negative correlation (r _p =–0.586, P=0.098) between diet overlap of main plant groups (calculated across studies) and difference in feeding type. Received: 19 October 1999 / Accepted: 31 January 2000     

Climate Change Promotes the Emergence of Serious Disease Outbreaks of Filarioid Nematodes

Filarioid parasites represent major health hazards with important medical, veterinary, and economic implications, and considerable potential to affect the everyday lives of tens of millions of people globally (World Health Organization, 2007). Scenarios for climate change vary latitudinally and regionally and involve direct and indirect linkages for increasing temperature and the dissemination, amplification, and invasiveness of vector-borne parasites. High latitude regions are especially influenced by global climate change and thus may be prone to altered associations and dynamics for complex host-pathogen assemblages and emergence of disease with cascading effects on ecosystem structure. Although the potential for substantial ecological perturbation has been identified, few empirical observations have emanated from systems across the Holarctic. Coincidental with decades of warming, and anomalies of high temperature and humidity in the sub-Arctic region of Fennoscandia, the mosquito-borne filarioid nematode Setaria tundra is now associated with emerging epidemic disease resulting in substantial morbidity and mortality for reindeer and moose. We describe a host-parasite system that involves reindeer, arthropods, and nematodes, which may contribute as a factor to ongoing declines documented for this ungulate species across northern ecosystems. We demonstrate that mean summer temperatures exceeding 14°C drive the emergence of disease due to S. tundra. An association between climate and emergence of filarioid parasites is a challenge to ecosystem services with direct effects on public health, sustainability of free-ranging and domestic ungulates, and ultimately food security for subsistence cultures at high latitudes.     

Herbivores inhibit climate-driven shrub expansion on the tundra

Recent Pan-Arctic shrub expansion has been interpreted as a response to a warmer climate. However, herbivores can also influence the abundance of shrubs in arctic ecosystems. We addressed these alternative explanations by following the changes in plant community composition during the last 10 years in permanent plots inside and outside exclosures with different mesh sizes that exclude either only reindeer or all mammalian herbivores including voles and lemmings. The exclosures were replicated at three forest and tundra sites at four different locations along a climatic gradient (oceanic to continental) in northern Fennoscandia. Since the last 10 years have been exceptionally warm, we could study how warming has influenced the vegetation in different grazing treatments. Our results show that the abundance of the dominant shrub, Betula nana , has increased during the last decade, but that the increase was more pronounced when herbivores were excluded. Reindeer have the largest effect on shrubs in tundra, while voles and lemmings have a larger effect in the forest. The positive relationship between annual mean temperature and shrub growth in the absence of herbivores and the lack of relationships in grazed controls is another indication that shrub abundance is controlled by an interaction between herbivores and climate. In addition to their effects on taller shrubs (>0.3 m), reindeer reduced the abundance of lichens, whereas microtine rodents reduced the abundance of dwarf shrubs (<0.3 m) and mosses. In contrast to short-term responses, competitive interactions between dwarf shrubs and lichens were evident in the long term. These results show that herbivores have to be considered in order to understand how a changing climate will influence tundra ecosystems.