Analysis of genetic information from the antlers of Rangifer tarandus (reindeer) at the rapid growth stage.

Reindeer is the only deer species in which both males and females regularly grow antlers, providing an excellent model for studying the rapid growth and annual regeneration of antlers. The study of genetic information from reindeer is the basis for revealing the unique mechanism of antler growth. In the present study, we obtained 18.86 GB of clean reads, which were assembled to obtain 94,575 unigenes (average length: 704.69). Among these reads, 30,980 sequences were identified by searching a database of known proteins and then annotated with Gene Ontology (GO) terms, Clusters of Orthologous Groups (COG) classifications and Kyoto Encyclopedia of Genes and Genomes (KEGG) pathways. All 7,480 simple sequence repeats (SSRs) were detected. A total of 84,435 and 82,226 high-quality single-nucleotide polymorphisms (SNPs) were identified in male and female reindeer, respectively. We identified 31 genes that were highly expressed in reindeer antlers. These genes regulate cell activities that are closely associated with the process of rapid tissue growth. Our results provide a basis for studying reindeer antlers and for further studying the molecular genetics, population genetics, and functional genomics of reindeer.

Multi-pathogen serological survey of migratory caribou herds: A snapshot in time.

Pathogens can impact host survival, fecundity, and population dynamics even when no obvious disease is observed. Few baseline data on pathogen prevalence and diversity of caribou are available, which hampers our ability to track changes over time and evaluate impacts on caribou health. Archived blood samples collected from ten migratory caribou herds in Canada and two in Greenland were used to test for exposure to pathogens that have the potential to effect population productivity, are zoonotic or are emerging. Relationships between seroprevalence and individual, population, and other health parameters were also examined. For adult caribou, the highest overall seroprevalence was for alphaherpesvirus (49%, n = 722), pestivirus (49%, n = 572) and Neospora caninum (27%, n = 452). Lower seroprevalence was found for parainfluenza virus type 3 (9%, n = 708), Brucella suis (2%, n = 758), and Toxoplasma gondii (2%, n = 706). No animal tested positive for antibodies against West Nile virus (n = 418) or bovine respiratory syncytial virus (n = 417). This extensive multi-pathogen survey of migratory caribou herds provides evidence that caribou are exposed to pathogens that may have impacts on herd health and revealed potential interactions between pathogens as well as geographical differences in pathogen exposure that could be linked to the bio-geographical history of caribou. Caribou are a keystone species and the socio-economic cornerstone of many indigenous cultures across the North. The results from this study highlight the urgent need for a better understanding of pathogen diversity and the impact of pathogens on caribou health.

Toward the Restoration of Caribou Habitat: Understanding Factors Associated with Human Motorized Use of Legacy Seismic Lines.

Populations of boreal and southern mountain caribou in Alberta, Canada, are declining, and the ultimate cause of their decline is believed to be anthropogenic disturbance. Linear features are pervasive across the landscape, and of particular importance, seismic lines established in the 1900s (legacy seismic lines) are slow to regenerate. Off-highway vehicles are widely used on these seismic lines and can hamper vegetative re-growth because of ongoing physical damage, compaction, and active clearing. Restoration of seismic lines within caribou range is therefore a priority for the recovery of threatened populations in Alberta, but a triage-type approach is necessary to prioritize restoration and ensure conservation resources are wisely spent. To target restoration efforts, our objective was to determine factors that best explained levels of off-highway vehicles use on seismic lines intersecting roads. We investigated the relative importance of local topography, vegetation attributes of seismic lines, and broad-scale human factors such as the density of infrastructures and the proximity to recreation campsites and towns to explain the observed levels of off-highway vehicles use. We found that off-highway vehicles use was mainly associated with local topography and vegetation attributes of seismic lines that facilitated ease-of-travel. Broad-scale landscape attributes associated with industrial, recreation access, or hunting activities did not explain levels of off-highway vehicles use. Management actions aimed at promoting natural regeneration and reduce ease-of-travel on legacy seismic lines within caribou ranges can be beneficial to caribou recovery in Alberta, Canada, and we therefore recommend restrictions of off-highway vehicles use on low vegetation, dry seismic lines in caribou ranges.

"Generality of mis-fit"? The real-life difficulty of matching scales in an interconnected world.

A clear understanding of processes at multiple scales and levels is of special significance when conceiving strategies for human-environment interactions. However, understanding and application of the scale concept often differ between administrative-political and ecological disciplines. These mirror major differences in potential solutions whether and how scales can, at all, be made congruent. As a result, opportunities of seeking "goodness-of-fit" between different concepts of governance should perhaps be reconsidered in the light of a potential "generality of mis-fit." This article reviews the interdisciplinary considerations inherent in the concept of scale in its ecological, as well as administrative-political, significance and argues that issues of how to manage "mis-fit" should be awarded more emphasis in social-ecological research and management practices. These considerations are exemplified by the case of reindeer husbandry in Fennoscandia. Whilst an indigenous small-scale practice, reindeer husbandry involves multi-level ecological and administrative-political complexities-complexities that we argue may arise in any multi-level system.

How the timing of weather events influences early development in a large mammal.

Capturing components of the weather that drive environment-animal interactions is a perennial problem in ecology. Identifying biologically significant elements of weather conditions in sensible statistics suitable for analysis of life history variation and population dynamics is central. Meteorological variables such as temperature, precipitation, and wind modulate rates of heat loss in animals, but analysis of their effects on endothermic species is complicated by the fact that their influence on energy balance is not invariably linear, even across the thermoneutral range. Rather, the thermal load imposed by a given set of weather conditions is a function of organisms' metabolic requirement, which, crucially, may vary spontaneously both seasonally and across different life phases. We propose that the endogenous component of variation in metabolic demand introduces a temporal dimension and that, as a consequence, the specific effect of meteorological variables on energy balance and attendant life history parameters is a function of the timing of weather events with respect to the organism's metabolic rhythm(s). To test this, we examined how a spontaneous increase in metabolic demand influenced the effect of weather on early development in a large mammal. Specifically, we examined interaction between the exponential rise in the energy requirements of pregnancy and depth of snow, which restricts dams' access to forage, on the body mass of reindeer calves (Rangifer tarandus) at weaning. As expected, we detected a significant temporal component: the specific negative effect of snow on weaning mass was not constant, but increased across pregnancy. The life history response was therefore better predicted by interaction between the magnitude and the timing of weather events than by their magnitude alone. To our knowledge, this is the first demonstration of the influence of an endogenous metabolic dynamic on the impact of weather on a life history trait in a free-living mammal. Evaluating weather variables with respect to endogenous variation in metabolic demand adds biological realism and is likely to improve understanding of the influence of environmental variation on life history traits in many ecological contexts.

Population densities, vegetation green-up, and plant productivity: impacts on reproductive success and juvenile body mass in reindeer.

Global warming is expected to cause earlier springs and increased primary productivity in the Arctic. These changes may improve food availability for Arctic herbivores, but may also have negative effects by generating a mismatch between the surge of high quality food in the spring and the timing of reproduction. We analyzed a 10 year dataset of satellite derived measures of vegetation green-up, population densities, calf body masses and female reproductive success in 19 reindeer (Rangifer tarandus) populations in Northern Norway. An early onset of spring and high peak plant productivity had positive effects on calf autumn body masses and female reproductive success. In addition, body masses and reproductive success were both negatively related to population density. The quantity of food available, as determined by the onset of vegetation green-up and plant productivity over the summer were the main drivers of body mass growth and reproductive success. We found no evidence for an effect of the speed of spring green-up. Nor did we detect a negative mismatch between early springs and subsequent recruitment. Effects of global warming on plant productivity and onset of spring is likely to positively affect sub-Arctic reindeer.

Is there a cost of parasites to caribou?

SUMMARY: Macroparasites potentially play a significant but often ignored role in the ecology and dynamics of wild ruminant populations. In the Arctic, parasites may impact on host populations by exacerbating the effects of seasonal and limited forage availability on the condition, fecundity and survival of individuals. We studied the effects of abomasal nematode parasites and warble flies, Hypoderma tarandi, on condition and pregnancy of caribou Rangifer tarandus in the Dolphin-Union herd, Nunavut, Canada. By the end of winter, female caribou over 2 years old showed a significant decrease in body weight with increasing nematode burden, and a decrease in back fat depth with increasing warble abundance. These effects were exaggerated in the non-pregnant fraction of the population. High warble larvae burdens were also associated with significantly reduced probability of being pregnant. Our research demonstrates a negative relationship between parasites and caribou condition that may have consequences for their fitness. Additionally, we discuss the possibility that muskox Ovibos moschatus share some parasite species with the caribou and could lead to elevated burdens in the sympatric host. Parasites may have been a contributory factor in a previous winter range-shift of the caribou herd and this may reflect a form of apparent competition between the two ungulate species.

Warming, plant phenology and the spatial dimension of trophic mismatch for large herbivores.

Temporal advancement of resource availability by warming in seasonal environments can reduce reproductive success of vertebrates if their own reproductive phenology does not also advance with warming. Indirect evidence from large-scale analyses suggests, however, that migratory vertebrates might compensate for this by tracking phenological variation across landscapes. Results from our two-year warming experiment combined with seven years of observations of plant phenology and offspring production by caribou (Rangifer tarandus) in Greenland, however, contradict evidence from large-scale analyses. At spatial scales relevant to the foraging horizon of individual herbivores, spatial variability in plant phenology was reduced--not increased--by both experimental and observed warming. Concurrently, offspring production by female caribou declined with reductions in spatial variability in plant phenology. By highlighting the spatial dimension of trophic mismatch, these results reveal heretofore unexpected adverse consequences of climatic warming for herbivore population ecology.

Evidence for a trade-off between early growth and tooth wear in Svalbard reindeer.

Ruminants depend on efficient physical degradation of forage through chewing to increase the surface area of the food particles presented to the microflora. Fossil evidence suggests that increased molar height is an adaptation for wear tolerance in dry ecosystems with sparse vegetation, but no study has shown selection pressure for hypsodonty in contemporary ruminants. We explored the relationships between particle size in rumen, tooth wear (scanned molar occlusal topography), age and body mass of female Svalbard reindeer living in an arctic desert at 78 degrees latitude on Svalbard. We predicted that (H1) if the rumen particle size is determined mainly by constraints due to tooth wear, and if tooth wear is mainly a function of age, average particle size in rumen should increase with age. From allometric relations it is known that larger individuals can survive on a lower-quality diet, we therefore predicted (H2) larger particle sizes with increases in (ln) body mass, irrespective of age and wear. Lastly, if there is a trade-off between growth and tooth wear in dry ecosystems (a selection pressure for hypsodonty), we predicted (H3) that teeth of heavier animals should be more worn than those of lighter animals of the same age. The proportion of small particles (<1.0 mm) decreased rapidly with increasing age (consistent with H1). Heavier females within an age class had more worn teeth (consistent with H3) than lighter ones. A close-to-isometric relationship between particle size and body mass suggested that heavier animals partly compensated for reduced tooth efficiency by chewing more. We provide the first evidence of a trade-off between fast early growth and wear (a somatic cost) of a senescence-related trait--the structure and height of the molar--in a wild ruminant inhabiting an arctic desert where selection pressure for increased tooth height is expected. This suggests that foraging conditions are more extreme than the environment in which the species originally evolved.

Good reindeer mothers live longer and become better in raising offspring.

Longevity is the main factor influencing individual fitness of long-lived, iteroparous species. Theories of life history evolution suggest this is because increased longevity allows individuals to (i) have more breeding attempts (time component), (ii) accumulate experience so as to become better able to rear offspring (experience component) or (iii) because individuals reaching old age have above-average quality (quality component). We assess empirically the relative influences of time, experience and quality on the relationship between longevity and individual fitness among female reindeer. Fitness increased with longevity due to all three processes. All females increased in success with age up to their penultimate year of life (experience component), the success of the terminal-breeding occasion was strongly dependent on longevity. Long-lived females had more successful breeding attempts during their life (time component), and had higher reproductive success at all ages, especially during the last year of life (individual quality component) than short-lived females. Our study reveals a more complex relationship between longevity and fitness in large mammals than the simple increase of the number of reproductive attempts when living longer.

Temporal changes in concentrations of serum amyloid-A and haptoglobin and their associations with weight gain in neonatal reindeer calves.

Age-related changes in serum concentrations of two acute phase proteins (APPs), haptoglobin (Hp) and serum amyloid-A (SAA) were investigated in newborn reindeer calves. Repeated blood samples were obtained from 51 reindeer calves at ages 0-32 days (2-4 samples from each calf). An increase of SAA concentrations was observed during the first 2 weeks of life. However, by the end of the observation period, SAA concentrations had decreased to levels below those of the first week. Serum Hp concentrations increased throughout the observation period. SAA concentrations in the second week had a negative association with weight gain during the entire study period (4 months). These time-related changes in APP concentrations suggest that these proteins have a role in the defence and adaptation mechanisms of newborn reindeer calves. Possible reasons for these changes include the presence of APP mediators in the colostrum, exposure to environmental pathogens after birth and age-related changes in hepatic synthesis of APP.

(90)Sr, (210)Po and (210)Pb in lichen and reindeer in Norway.

Concentrations of (90)Sr, (210)Po and (210)Pb in lichen and reindeer were studied in central (Østre Namdal) and southern Norway (Vågå) during 2000-2003. The study focussed on potential differences in concentrations of these nuclides in reindeer of different ages. Concentrations of (90)Sr in bones of approximately 10 year old adult females were about 40% higher than those in calves' antlers ((90)Sr concentrations in antlers and bones of calves are similar), while the available data from Vågå suggest that (90)Sr concentrations in reindeer calves decreased with an effective ecological half-time of 9.03+/-0.06 years during 1988-2002. Furthermore, (90)Sr concentrations were 50-80% higher in bone of reindeer of a similar age from Vågå compared to those from Østre Namdal. Concentrations of (210)Po and (210)Pb in muscle and liver tissues were comparable to those reported for reindeer in other Nordic areas, with no significant difference in (210)Po and (210)Pb concentrations between adults and calves or between reindeer from the two different study areas.

Global climate change and reindeer: effects of winter weather on the autumn weight and growth of calves.

Reindeer/caribou (Rangifer tarandus), which constitute a biological resource of vital importance for the physical and cultural survival of Arctic residents, and inhabit extremely seasonal environments, have received little attention in the global change debate. We investigated how body weight and growth rate of reindeer calves were affected by large-scale climatic variability [measured by the North Atlantic Oscillation (NAO) winter index] and density in one population in central Norway. Body weights of calves in summer and early winter, as well as their growth rate (summer to early winter), were significantly influenced by density and the NAO index when cohorts were in utero. Males were heavier and had higher absolute growth than females, but there was no evidence that preweaning condition of male and female calves were influenced differently by the NAO winter index. Increasing NAO index had a negative effect on calves' body weight and growth rate. Increasing density significantly reduced body weight and growth rate of calves, and accentuated the effect of the NAO winter index. Winters with a higher NAO index are thus severe for reindeer calves in this area and their effects are associated with nutritional stress experienced by the dams during pregnancy or immediately after calving. Moreover, increased density may enhance intra-specific competition and limits food available at the individual level within cohorts. We conclude that if the current pattern of global warming continues, with greater change occurring in northern latitudes and during winter as is predicted, reduced body weight of reindeer calves may be a consequence in areas where winters with a high NAO index are severe. This will likely have an effect on the livelihood of many northern indigenous peoples, both economically and culturally.

Reproductive success in reindeer males in a herd with varying sex ratio.

In polygynous species, male reproductive success is often correlated with dominance status of individual males and sex ratio in the population. Reindeer, Rangifer tarandus, is a polygynous species, and here we compared the variation in male reproductive success and dominance status during two successive years in a herd with a male:female sex ratio of 1:7 and 1:3. Copulations were recorded, together with data on male dominance hierarchy and size of mating groups. Male reproductive success was estimated by paternity analysis of calves using microsatellite DNA markers. The distribution of reproductive success among the males was highly skewed for both years with the most dominant male also being the most successful. The largest mating group was established in the herd with the least skew in sex ratio. In this herd some of the adult males present were less reproductively successful than some of the more subordinate younger males. Estimates of the mating group size of males, correcting for dominance status when more than one male is present in the groups, gave good prediction of individual males' reproductive success.

Muscle fibre growth in undernourished reindeer calves (Rangifer tarandus tarandus L.) during winter.

To study whether moderate under-nutrition causes muscle wasting, reindeer (Rangifer tarandus tarandus L.) calves were fed either pelleted reindeer feed ad libitum (n=8) or restricted amounts of lichens (n=8). The restricted amount was 60% of ad libitum intake of lichens, and the feeding period was 6 weeks preceded by a 2-week adjustment period. Biopsy samples from the middle gluteal muscle (M. gluteus medius) for the analysis of fibre composition and area, as well as for the activity of cathepsin B were taken before the restriction period in November and January, and after the restriction period in April. In all calves the muscle fibre composition remained unchanged during the winter. In the lichen group, the fibre size also remained unchanged, whereas in control calves the cross sectional area of type I and type IIA fibres increased significantly from November to April. Cathepsin B activity decreased in all calves from November to January and remained at that low level for the rest of the study period, which suggests an attenuated rate of protein degradation. These results can be taken as an indication that moderate under-nutrition causes no muscle wasting in reindeer calves, and the decreased availability of nitrogen is partially compensated for by adaptive decrease in protein degradation. Interestingly the adaptive changes in protein metabolism are equally well seen in the well-fed controls as in the undernourished lichen-fed reindeer.

Parasites, biodiversity, and population dynamics in an ecosystem in the high arctic.

The abomasa of 163 Svalbard reindeers (Rangifer tarandus platyrhynchus) from Nordenskiöld Land, Spitsbergen (78 degrees N, 15 degrees E) were examined for adult and juvenile nematodes. Spitsbergen has midnight sun from late April to late August, arctic night from late October to mid-February, permafrost, and snow cover from October to June. Plant growth is restricted to 6-8 weeks, usually starting mid-June. In the reindeer calves, which are born in June prevalence and intensity of infection increased over the winter. All young (1 year) and adult (older than 1 year) reindeers were infected. Variations in worm burden, fraction of adult nematodes, male to female ratio in the nematodes, number of adult nematodes, number of juvenile nematodes, and distribution of juvenile nematodes on length groups were analysed for relationships with sex and age of host, and time of the year. The material was further analysed for relationships between the reindeer's content of stored fat and worm burden and its elements. It is concluded that transmission of nematodes to the reindeer continues over the winter and that arrested larval development has not been strongly selected for in this High Arctic locality. Infection continued to increase in adult males but not in adult females with age. This is explained as a balance between gain and loss of worms in the females connected to levelling off of food intake at end of somatic growth. In adult male reindeers there was a negative trend between stored fat and infection at all seasons. In reproducing females there were significant negative relationships in winter-spring, while in yeld females no significant relationship was found. The nematodes belonged to the following taxa: Ostertagia gruehneri m. gruehneri, O. gruehneri m. arcticus, Marshallagia marshalli m. marshalli, M. marshalli. m. occidentalis, Teladorsagia circumcincta m. circumcincta, T. circumcincta. m. trifurcata, and T. circumcincta. m. davtiani. There appears not to be any specially adapted species among the helminth parasites of reindeer (Rangifer tarandus) and muskoxen (Ovibos moschatus) in the High Arctic. In the Low Arctic Uminmakstrongylus pallikuukensis of muskoxen may be an example of such adaptation.

Annual serum PICP and ICTP and antler growth in female reindeer (Rangifer tarandus tarandus).

Annual cycle of type I collagen formation and degradation and antler growth was studied in six adult female reindeer, Rangifer tarandus tarandus. Blood samples were collected twice a week during 1 year. Antler length was measured weekly during the antler growth period. An assay for human PICP, the carboxyterminal propeptide of type I procollagen, was used as an indicator of type I collagen formation and an assay for bovine ICTP, the carboxyterminal telopeptide of type I collagen, as an indicator of type I collagen degradation. PICP was clearly increased during the antler growth period. Also ICTP was slightly elevated during antler growth, but the highest values were found in autumn and winter. Our statistical analysis revealed that changes in lagged values (from 3 to 6 weeks) of PICP could be linked to the subsequent changes in the growth rate of the antler, although the highest values of PICP were found during the final third of antler growth. ICTP had significant predictive power as well, but the connection with the growth rate seemed more immediate than that of PICP. In conclusion, antler collagen synthesis can be predicted by PICP, but also ICTP was related to the antler growth.

Predicting effects of naturally acquired abomasal nematode infections on growth rate and food intake in reindeer using serum pepsinogen levels.

An experiment was performed on semiwild reindeer with naturally acquired parasite infections to assess whether measures of serum pepsinogen concentration and fecal egg counts can be used to predict effects of abomasal nematodes on reindeer weight gain and food intake. Food intake and weight gains were lower in infected calves compared with calves where parasites had been removed by anthelmintic treatment. Among the infected animals, concentration of pepsinogen in serum was correlated with food intake and weight gains. Concentration of abomasal nematode eggs in feces was not. Our results suggest that measures of pepsinogen levels in serum, but not fecal egg counts, may be used to predict the effects of abomasal nematodes on food intake and weight gain of reindeer.

Ivermectin treatment did not increase slaughter weight of first-year reindeer calves.

To investigate if antiparasitic treatment of reindeer calves during the summer could increase their carcass weight during the slaughter period in autumn and winter, 529 reindeer calves were allocated to three groups, weighed, and marked with individually numbered plastic ear tags in early July, 1995. One of the groups was left untreated, another was treated with ivermectin injection at 200 micrograms/kg, and the third with pour-on ivermectin at 500 micrograms/kg. Following slaughter, carcass weights were received from 231 animals, and there was no difference between the treatment groups.

Seasonal levels of reproductive hormones and their relationship to the antler cycle of male and female reindeer (Rangifer tarandus).

Seasonal levels of LH, FSH, testosterone (T), estradiol, progesterone (P), and prolactin (PRL) were determined in the plasma of five adult bulls, and five barren and four pregnant cows of Alaskan reindeer (Rangifer tarandus), which were sampled every 3 weeks for 54 weeks. The male reproductive axis was sequentially activated; LH peaked in May-June (2 ng/ml), FSH in June (51 ng/ml), and T in September (11.8 ng/ml). LH levels in females reached a maximum in both groups at the end of August (the beginning of the rut). Seasonal variation in FSH was minimal in pregnant cows, but exhibited one elevation (41 ng/ml) in barren ones in November. T levels in cows remained at barely detectable levels. The decrease of T values observed in both groups in December and March was not significant. PRL peaked in May in cows (135 ng/ml pregnant, 140 ng/ml non-pregnant) and in June in bulls (92 ng/ml). Estradiol was highest in bulls in the rut (August), in non-pregnant cows in January and in pregnant cows in April, shortly before parturition. P levels in the pregnant cows rose from September and peaked (9 ng/ml) shortly before parturition in April. In the non-pregnant females P values increased and decreased several times before peaking (5 ng/ml) in March. In the males, the variation of T and estradiol levels correlated relatively well with the antler cycle but in the females the variation of neither estradiol, progesterone nor T appeared to be related to mineralization or casting of antlers.