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.

Reproductive allocation at one age is predicted to reduce the probability of surviving to the next year or to lead to a decrease in future reproduction. This prediction assumes that reproduction involves fitness costs. However, few empirical studies have assessed whether such costs may vary with the age at primiparity or might be overridden by heterogeneities in individual quality. We used data from 35 years' monitoring of individually marked semi-domestic reindeer females to investigate fitness costs of reproduction. Using multi-state statistical models, we compared age-specific survival and reproduction among four reproductive states (never reproduced, experienced non-breeders, reproduced but did not wean offspring, and reproduced and weaned offspring) and among contrasted age at primiparity. We assessed whether reproductive costs occurred, resulting in a trade-off between current reproduction and future reproduction or survival, and whether early maturation was costly or rather reflected differences in individual quality of survival and reproduction capabilities. We did not find any evidence for fitness costs of reproduction in female reindeer. We found no cost of gestation and lactation in terms of future reproduction and survival. Conversely, successful breeders had higher survival and subsequent reproductive success than experienced non-breeders and unsuccessful breeders, independently of the age at primiparity. Moreover, it was beneficial to mature earlier, especially for females that successfully weaned their first offspring. Successful females at early primiparity remained successful throughout their life, clearly supporting the existence of marked among-female differences in quality. The weaning success peaked for multiparous females and was lower for first-time breeders, indicating a positive effect of experience on reproductive performance. Our findings emphasize an overwhelming importance of individual quality and experience to account for observed variation in survival and reproductive patterns of female reindeer that override trade-offs between current reproduction and future performance, at least in the absence of harsh winters.

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. Longlived 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.

Calving and calf production were studied in an experimental reindeer (Rarigifcr tarandus L) herd in Inari, Kaamanen (69°10'N) in northern Finland. Most of the calving took place between May 10 and 29; 50% of the calves were born up to 22.5. and 90% up to 29.5. The percentage of calves (calves/hinds proceeding year) was on average 79.2%. The newborn sex ratio was 1:1. There was a large range (1.8 to 8.5 kg) in the birth-weight of newborn calves; male calves weighed about 0.3 kg more than female. The total loss of calves during summer and autumn was about 34.5% (range 6.3 to 100.0%). During the calving period 111 calves (12.2%) died. They were usually younger than one day, and on average weighed 1.3 kg less at birth than calves who lived until autumn. The hinds of calves dying very early were usually young and calving for the first time. The autumn-weight of hinds correlated significantly (r = 0.49) with the birth-weight of their calves. Over-5-year-old and over 80 kg hinds bore the heaviest calves (mean weight 6.3 kg) and the survival of these calves was very good. The first hinds to calve in the spring (calving up to 22.5.) are 5-year-old and older, which arc also the heaviest and produce the most vital calves.

Two different processes can lead to a change in individual reproductive output with age in long-lived iteroparous vertebrates. The senescence hypothesis predicts a decline of performance in old age, whereas the terminal allocation hypothesis predicts an increase. Using long-term (>30 years) individually based data of female reindeer, we first assessed age-specific variation in body mass and different components of reproductive output. Then we investigated the contribution of senescence and terminal allocation (the increase in components of reproductive output) processes for shaping observed patterns. We found that female reindeer body mass increased up to about 11.5 years of age, and decreased afterwards, supporting the senescence hypothesis. Calf birth mass, both in absolute terms or for a given female mass, first increased and then declined with female age, also supporting the senescence hypothesis. The female mass gain (June-September) decreased with increasing age, and female change in mass between 2 consecutive years decreased with female age, all patterns again supporting the senescence hypothesis. However, the autumn calf mass did not change with age. Calf body mass in autumn tended to be positively related to female mass gain, supporting a quality effect. Raising a calf had a marked negative effect on female mass gain, indicating energetic reproductive costs of raising a calf. Calf body mass in autumn did not influence yearly female mass change. Overall, our results provided consistent evidence for general effects of senescence on most components of reproductive output and highlighted that both individual heterogeneity and reproductive costs shape female reindeer reproductive tactics.

ABSTRACT. The focus of the work was to investigate dependency between reindeer density and lichen (Cladonia spp.) ranges in the Finnish semidomesticated reindeer management area. Secondly, we formed a model on the recovery rate of ungrazed woodland lichen ranges (29 sites) after forest fires to evaluate the potential productivity and time needed for Finnish lichen ranges to recover at optimal production. During 1974 -95, 59% of the variation in mean semidomesticated reindeer density (range: 0.7-3.0 reindeer/km 2 of the total land area) among the reindeer herding districts in Finland was explained by the proportion of land area covered by lichen ranges in these districts. Reindeer densities were highest in the districts where lichen ranges covered 20% to 30% of the area. Reindeer density on the total land area did not explain the condition of lichen ranges, but 58% of the condition was explained by the reindeer density on the lichen ranges. A condition level for lichen ranges of 1000 kg dry matter per hectare (d.m./ha) of lichen biomass can be considered adequate to ensure survival of reindeer and continued production of lichen. To maintain this level within the sedentary Finnish grazing system, winter reindeer densities on lichen ranges must remain below 5-7 reindeer/km 2 . According to our model, the maximum amount of living lichen in the woodland lichen stand at the climax stage is on average about 7000 kg d.m./ha. The maximum annual yield of lichen (175 kg d.m./ha) is produced by lichen stands that contain 2600-2800 kg d.m./ha of living lichen. Using our model and our 1995 -96 data, we calculated that the average lichen biomass on lichen ranges in the Finnish reindeer management districts was 13.0% of this optimum, and the average lichen production was 36% of the possible maximum annual yield. Our model indicates that the Finnish lichen ranges would have to remain ungrazed for an average of 18 years to recover to maximum production levels. However, the average time needed for the lichen ranges to recover to the level of 1000 kg lichen d.m./ha, would be only about 7 years.Key words: Rangifer, reindeer, Cladonia, lichen, pastures, condition, recovery, productivity RÉSUMÉ. Ces travaux portent tout d'abord sur l'étude de la dépendance existant entre la densité du renne et les grands pâturages de lichen (Cladonia spp.) dans le secteur finlandais de gestion du renne semi-domestique. Ensuite, on a conçu un modèle en se fondant sur le taux de récupération, après des incendies de forêt, d'étendues naturelles (29 emplacements) de lichen forestier non pâturées, en vue d'évaluer la productivité potentielle et le temps nécessaire aux grands pâturages de lichen finlandais pour récupérer à une production optimale. De 1974 à 1995, 59 p. cent de la variation dans la densité moyenne du renne semi-domestique (fourchette de 0,7 à 3,0 renne/km 2 de superficie totale des terres), parmi les districts où se trouvent les troupeaux de rennes en Finlande, s'expliquaient par la proportion du territoire qui était couvert par des pâturage...

In sexually dimorphic ungulates, male reproductive success depends on fighting with other males for access to females during a brief rutting season. Large body size is necessary for success in intrasexual competition, and a few large-sized males are often able to monopolize access to female groups. Earlier studies have reported that reproductive effort increases with age until prime-age is reached, and one study that population density lowered effort in (older) males. No study has directly assessed whether there is within-age-class variation in effort resulting from varying levels of intra-male competition. It is reported here the weight loss during the rutting season of 54 individual male reindeer Rangifer tarandus coming from eight herds with varying density (3.3-6.0 deer/km 2 ) and sex ratio (4-28% males). In agreement with earlier studies, reproductive effort was lower for young (1-to 2-year-old) than for prime-aged (3-to 5-year-old) males both on an absolute and relative scale. Among 1-year-old males (n = 33), effort was lower as sex ratio became closer to even, but density during the rutting season had no effect. This suggests that yearling males take a more active role when prime-aged males are absent. In addition to the insight into male ungulate life history, understanding male rutting behaviour may also have implications for population dynamics.

Large mammals in seasonal environments have a pattern of high-reproductive synchrony in spring, but how the timing of reproduction affects resource allocation decisions at different stages of the reproductive cycle remains largely unexplored. By manipulating the timing of conception in reindeer (Rangifer tarandus), we tested how the timing of conception affected sex ratio, gestation length and weight development of mother and offspring. Females that conceived at their first ovulation within the rut had a 60.5% probability of producing a male; in contrast, females that conceived a cycle later had a 31.3% probability of producing a male. Late conceiving females had gestation times that were 10 days shorter and the calves were 0.6 kg (9.2%) lighter at birth and 7.4 kg (14.7%) lighter in autumn. Over the year, female weight changes was similar between the groups suggesting reindeer follow a bet-hedging strategy; reducing the quality of this year's offspring to ensure their own future reproduction and survival. Harvesting is often selective leading to skewed sex ratios and age structure, which may influence the timing of reproduction due to females hesitation to mate with young males. Whenever this hesitation is strong enough to increase the frequency of recycling, harvesting is likely to have profound life history consequences.