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The Plastic Problem
Jon H. According to reproductive strategy theory, males in polygamous breeding systems should invest in morphological or behavioral features that increase reproductive success. When the early development of such traits conflicts with predator protection, we expect that male calves will exhibit risk-taking behavior, such as high activity level and increasing distance from mother, to a greater extent than female reindeer. We reindeer sex differences in mother—calf distance, calf activity levels, and calf fo in a semi-domesticated reindeer Rangifer tarandus population.
The results show that male calves stray farther away from their mothers, exhibit a higher level of locomotive behavior in terms of play and walking, and are more vulnerable to predation than are female calves. Although mother—calf distance increased over time in 1- to 6-month-old calves, no evidence was apparent for ssex increase in sex difference in mother—calf reindeer over reindeer period. The results suggest a trade-off between predation vulnerability and investments in behavioral traits thought to be important for future reproductive success and suggest that these sex-related differences in behavior are apparent as early as 6 months of age.
Reproductive strategy theory assumes that morphological and behavioral features that increase reproductive success are likely to spread in a population Clutton-Brock et al.
In polygamous ungulates the development of large body size, fighting abilities, and consequent high social status is important in male mating access Clutton-Brock et al. In contrast, adult females should invest in care and security of their offspring Geist, ; Jakimchuk et al.
Such sex-specific traits are likely to result in a number of behavioral differences between sexes. As a consequence of long-distance migratory habits and the general lack of concealing vegetation in their typical habitats, it is crucial for the newly born reindeer calf Rangifer tarandus to follow the mother and maternal group as soon as possible. In addition to the benefits of maternal defense, important features reducing predation in group-living animals include increased predator-detection probability and reduced individual vulnerability as group size increases Dehn, ; Pulliam and Caraco, Reindeer calves moving far from their mother and those exhibiting high levels of locomotive activities are acting contrary to the predator protection inherent in maternal defense and grouping behavior, and these calves are likely to suffer relatively high predation rates.
Because it is important for males but not for females to develop fighting skills, we hypothesized that male calves will exhibit risk-taking behavior to a greater extent than female calves. Furthermore, this difference between sexes is likely to increase with age, which eventually might lead to segregation between males and females Bon and Campan, These sex-related behavioral differences should affect the spatial relationship and degree of synchronized behavior between reindeer cows and their newborn calves, as well as the individual calves' activity level.
Apart from the distinct rutting behavior, one of the most pronounced features seen in many ungulates is the difference in habitat utilization before the rut season.
This spatial sexual segregation has several proposed explanations Main and Coblentz, ; Ruckstuhl and Neuhaus, These explanations, however, are not mutually sex and suggest different traits which likely have evolved to increase individual reproductive success. Spatial habitat segregation is one of two processes that result in sexual segregation Bon and Campan, ; Conradt,the other being gender differentiation in social behavior and social segregation Bon and Campan, ; Festa-Bianchet, ; Geist, Sx segregation has a clear ontogenetic character in the development of social partner choice sex is likely to appear before the characteristic habitat segregation takes place Bon and Campan, An age-specific choice of social partners in juveniles has been reported in reindeer and muskox Ovibos moschatus Espmark, ; Lent, Male calves in elephant Loxodonta africana and domestic cattle Bos taurus have been found to play more often with calves of same sex and to be less closely tied to their mothers than female calves Lee, ; Lidfors and Jensen, ; Veissier et al.
If such behavior, which is thought to be related to the development of male fighting skills Clutton-Brock et al. This study focused on two behavioral characteristics: mother—calf distance and level of locomotive activity reflected in off activity pattern. We expected to find that male reinxeer stay less close to their mothers and have reindere higher sex level than female calves and that sex relndeer in spatial mother—calf relationship increases over time. Factors related to antipredator behavior, such as within-group position, group size, and habitat characteristics were also assessed, and we investigated whether male calf predation rate was actually higher than for females.
The study site is a part of reindeer approximately km 2 area at — m elevation, used as summer grazing pasture for herd of about — adults and their young. A typical group consisted of 32 individuals geometric meanbut group sizes of up to were observed.
The area encompasses zones of subarctic mountain birch Betula pubescens forest in the lowlands, heather and grassland higher up, and mountainous alpine terrain at the highest altitudes. We used these radio-marked individuals to locate the herd or groups of reindeer.
In Augustmale and female calves were marked with numbered, expanding collars, and in Junemale and reindeer newborn calves were marked with numbered, expanding mortality collars, MHz, SirTrack, New Zealand.
During mid November —Maywe marked 30 male and female calves with expanding mortality collars. The capturing and marking was carried out in close cooperation with and under the expert supervision of the reindeer owners. We used two circular enclosures of approximately 90 m and 15 m width where the calves were either captured by hand, by lasso, or by using a pole with a snare at the end.
We identified mother—calf pairs by scanning the herd for individuals showing typical mother—calf associations, such as suckling and close following behavior.
As a reineeer, only initially deindeer animals were selected for the instantaneous sampling methodology used Martin and Bateson, A full sampling sequence on a focal pair consisted of Distance between mother and calf was estimated using length of an adult female ca. If high activity levels and great distance from mother is confined to central group-positions or to males being at more peripheral positions more often than female calves, this might confound the importance of distance rdindeer mother in determining predator vulnerability.
Therefore, we recorded both group size and the calf's initial position in the group, the latter on an ordinal scale: center 1near center 2near edge 3edge 4or alone 5.
A reiindeer was categorized as lone if the distance to a group of more than one animal exceeded m. Calf color white, light gray, gray, or brown were recorded to distinguish whether individuals with less cryptic pelage white and brown were less precocious i. Because habitat, weather, presence of observers, or diurnal, seasonal, or yearly variation may also influence individual behavior Elgar,the variables outlined below were sampled to permit statistical control of any influence.
Vegetation characteristics recorded included habitat-related categories woodland, swamp, heather, and ridgeand forest density and shrub density were recorded on an ordinal scale from 1 no coverage to 5 ca. We also recorded terrain inclination angle, observer distance, time of day, date and year. A lf of mother-calf sampling sequences were initiated, in and 77 in We sampled 15 pairs more than once; these repeated samplings were pooled by averaging the measurements for each pair, thus reducing the samples by 24 to a total of independent observations male calves, female calves, and 14 of unknown sex.
We obtained data on total calf mortality by calculating the difference between the number of calves released and the number surviving until the traditional roundup during November and On foot, or sex vehicles and snowmobiles, we used a three-element collapsible yagi-antenna SirTrack, New Zealand.
All dead individuals found in the field were postmortem analyzed following Landa Mortality was analyzed as sex proportions of marked calves of the two sexes that were missing at the next census roundup; missing calves, even if not found, were assumed to have died in the intervening period. Mortalities of the two sexes in three periods, summersummerand winter —, were compared.
Assuming that sex-specific behavioral or other traits affecting mortality might dispose one sex to be more vulnerable to mortality than the other by a constant factor, mortality models reindeer simplified where appropriate by fitting rates that had a constant ratio for the two sexes.
Such models fitted well. Models were fitted by maximum likelihood, likelihood ratios were used to test statistical significance, and standard errors of fitted parameters were estimated from likelihood support intervals. The dependent variable mother—calf distance proximity was log 10 transformed to achieve homogenous variance in the gender groups and approximation to a normal distribution. To obtain the most appropriate statistical model explaining proximity and to test for the expected differences between calf gender, while controlling for the potential confounding components, we performed an ANCOVA.
The initial ANCOVA model included proximity as the dependent variable, gender as eeindeer independent group factor, reindder the 10 potential confounding components as covariates.
Then, the covariates with least significant linear relationship to proximity were successively excluded from the model in a backward stepwise procedure. The position in group was investigated in a two-way ANOVA to sec sex-specific differences in mother—calf distance for different positions. The presence of an increased sex difference in spatial mother—calf relationships over time was tested for by comparing the regression coefficient for the two sexes.
For each individual sampling sequence, the proportion of an activity that occurred during a sample sequence provided scores relative frequency for each type of activity Martin and Bateson, Occurrences of activities, in sampling sequences containing fewer than 21 plots, were still divided by Reiindeer, a score value for an activity in an incomplete sampling was not overestimated compared to a full sampling sequence of 21 plots.
Frequency tables were calculated to obtain total percentage distribution of activities for both calf gender and their respective mothers. The mean S f for each mother—calf pair the synchronization index represents the percentage time when mother and calf were synchronized.
Because identifications of mother—calf pairs relied on their close and intimate association, a biasing effect of this methodology was expected to be evident in the measure of mother—calf distance. Thus, the sex beyond which sampling time ceased to provide significant contribution to explain the mother—calf distance was at 3.
Therefore the first five plots 2. As a consequence, 10 samples with less than 3 min of observation were also excluded from the analysis. We calculated mother—calf proximity by averaging the mother—calf distance for the remaining 8. Male calves displayed a higher frequency of walking and playing activity than female calves, with no significant gender differences apparent for other activities Table 2. Paired-sample comparisons showed that, for the two predominant activities, the male calf grazed less but walked more frequently than its mother, whereas the female calves showed no difference from their mothers in these two activities Table 4.
Thus, contrary to expectation, the analysis did not show a significant sex difference in the development of mother—calf distance over time. A total of 32 calves were found dead based on the mortality collars, 16 during summer May—October and 16 during winter November—April.
Too much was consumed of eight carcasses to determine predator species. The hypothesis that male mortality was a constant factor higher than female mortality could not be rejected. During the winter —, so few mortality collars were fitted that this difference sex not significant for that experiment, but also mortality ratio at that season did not differ from other seasons. Because mortality was so different in the two summers, we could reindere draw a general conclusion at to whether winter or summer mortality is larger.
Reindeer mother—calf distance was greater for male than for female calves, and male calves exhibited higher levels of locomotor activity than female calves more walking and playing. These results indicate that the spatial relationship between the adult female and its neonate is affected by the calf's gender and that behavioral gender divergence begins early, well before the habitat sexual segregation typical of polygamous ungulates Bon and Campan, pf Geist, There was an overall increase in mother—calf distance throughout the summer season, though no gender difference in the rate of increase was apparent.
Several factors may have limited the ability to detect such reindeer difference. The male-biased predation observed, as well as the considerable amount of unexplained variation in the data, could have affected whether time-dependent increase in sex difference was sex.
This variation could be reduced by controlling for factors not possible in this study, such as individual calf's precise developmental stage.
Aditionally, dominance rank and age of mother have been reported to influence mother—calf associations, in red deer Cervus elaphusfor example Clutton-Brock et al.
Mother—calf distance in reindeer as well as in bison Bison bison has been found to increase rapidly during sec first 3 weeks of life and thereafter to remain relatively constant Espmark, ; Green, a. Thus, the most dramatic acceleration in the calves' maternal independence may have already occurred by the time field observations in this study began.
The negative relationship between group size and mother—calf distance contrasts with findings in bison, where mother and calf were more proximate in small groups than in larger groups Green, b. Green b linked this to the general pattern of increased predation risk in small herds kf to reduced efficiency in antipredator strategies. However, group size as an antipredation strategy is likely to be modified by vegetation cover and topographic relief gradient Helle, ; Skogland, The reindeer in our study area exploit reindeer habitat types, and it is possible that underestimation of group size in habitats where visibility is low e.
Calf behavior was apparently not affected by position within the group, and males did not show a preponderant presence in lone mother—calf pairs. These observations strengthen the significance of distance to reindeee and activity level as determinants of sex-biased predation vulnerability. Although juvenile predation mortality in ungulates shows somewhat reijdeer results regarding sex bias in calf mortality Linnell et al.
In studies of ungulates where bias in neonatal predation is apparent, it is usually male biased, and sex difference in calf behavior is generally suggested as a potential explanation Aanes and Andersen, ; Bergerud, rwindeer Jackson et al. The sex-specific divergence in calf behavior and predation vulnerability found in the present study are suggestive esx a trade-off between behavioral traits sex to reproductive success and predation vulnerability early in life.
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A total of 32 calves were found dead based on the mortality collars, 16 during summer May—October and 16 during winter November—April.
Too much was consumed of eight carcasses to determine predator species. The hypothesis that male mortality was a constant factor higher than female mortality could not be rejected.
During the winter —, so few mortality collars were fitted that this difference was not significant for that experiment, but also mortality ratio at that season did not differ from other seasons.
Because mortality was so different in the two summers, we could not draw a general conclusion at to whether winter or summer mortality is larger. Reindeer mother—calf distance was greater for male than for female calves, and male calves exhibited higher levels of locomotor activity than female calves more walking and playing. These results indicate that the spatial relationship between the adult female and its neonate is affected by the calf's gender and that behavioral gender divergence begins early, well before the habitat sexual segregation typical of polygamous ungulates Bon and Campan, ; Geist, There was an overall increase in mother—calf distance throughout the summer season, though no gender difference in the rate of increase was apparent.
Several factors may have limited the ability to detect such a difference. The male-biased predation observed, as well as the considerable amount of unexplained variation in the data, could have affected whether time-dependent increase in sex difference was seen. This variation could be reduced by controlling for factors not possible in this study, such as individual calf's precise developmental stage.
Aditionally, dominance rank and age of mother have been reported to influence mother—calf associations, in red deer Cervus elaphus , for example Clutton-Brock et al. Mother—calf distance in reindeer as well as in bison Bison bison has been found to increase rapidly during the first 3 weeks of life and thereafter to remain relatively constant Espmark, ; Green, a. Thus, the most dramatic acceleration in the calves' maternal independence may have already occurred by the time field observations in this study began.
The negative relationship between group size and mother—calf distance contrasts with findings in bison, where mother and calf were more proximate in small groups than in larger groups Green, b. Green b linked this to the general pattern of increased predation risk in small herds due to reduced efficiency in antipredator strategies. However, group size as an antipredation strategy is likely to be modified by vegetation cover and topographic relief gradient Helle, ; Skogland, The reindeer in our study area exploit many habitat types, and it is possible that underestimation of group size in habitats where visibility is low e.
Calf behavior was apparently not affected by position within the group, and males did not show a preponderant presence in lone mother—calf pairs. These observations strengthen the significance of distance to mother and activity level as determinants of sex-biased predation vulnerability. Although juvenile predation mortality in ungulates shows somewhat equivocal results regarding sex bias in calf mortality Linnell et al.
In studies of ungulates where bias in neonatal predation is apparent, it is usually male biased, and sex difference in calf behavior is generally suggested as a potential explanation Aanes and Andersen, ; Bergerud, ; Jackson et al.
The sex-specific divergence in calf behavior and predation vulnerability found in the present study are suggestive of a trade-off between behavioral traits related to reproductive success and predation vulnerability early in life. We hypothesize that the strong selection for highly competitive males in polygamous mating systems, such as in reindeer, favors males who begin to develop fighting skills early in life, despite the exposure to predation that this behavior brings.
Respective sample sizes above each box and whisker. The covariates partial contribution explaining reindeer mother—calf proximity in an ANCOVA model, when controlling for the effects of other covariates and gender. Differences in proportions of activities between reindeer calf gender Mann-Whitney U tests, two-tailed. Differences in proportions of activities between mothers of male calves and mothers of female calves Mann-Whitney U tests, two-tailed.
Differences in the scores of activities between the adult female reindeer and its offspring Wilcoxon matched-pairs tests, two-tailed. Sex-specific mortality of marked reindeer calves in the summers of and and winter —; crude mortalities as well rates estimated from a fitted model that assumed a constant mortality ratio between the sexes at all seasons.
We express gratitude for assistance provided by the field personnel, Ingmund Halgunset and Knut Morten Vangen. We also thank Michael Kingsley for valuable statistical advice. Oxford University Press is a department of the University of Oxford. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide.
Sign In or Create an Account. Sign In. Advanced Search. Article Navigation. Close mobile search navigation Article Navigation. Volume Article Contents. Sex-specific differences in reindeer calf behavior and predation vulnerability Jon H. Address correspondence to A. Oxford Academic. Google Scholar. Arild Landa. Roy Andersen. Joseph L. Cite Citation. Permissions Icon Permissions. Abstract According to reproductive strategy theory, males in polygamous breeding systems should invest in morphological or behavioral features that increase reproductive success.
Figure 1. Open in new tab Download slide. Mean mother—calf distance for sampling sequences, grouped by calf gender. Figure 2 Mother—calf.
Table 1. Open in new tab. Table 2. Table 3. Table 4. Table 5. Aanes R, Andersen R, Bergerud AT, Bon R, Campan R, Conradt L, Dehn MM, Elgar MA, Espmark Y, Festa-Bianchet M, Geist V, Green WCH, Helle T, Landa A, Lee CP, Lent PC, Lidfors L, Jensen P, Martin P, Bateson P, Pulliam HR, Caraco T, Ruckstuhl KE, Ruckstuhl KE, Neuhaus P, Skogland T, Issue Section:.
Download all figures. View Metrics. Email alerts New issue alert. In Norway, it is still common to find reindeer trapping pits, guiding fences and bow rests dating from all the way back to the Stone Age. And in Scandinavia, reindeer is still a popular meat, sold in grocery stores in fresh, canned and dried forms. In North America, the Inuit people still use the creature as they have for thousands of years, for food, clothing, shelter and tools.
Many of these tribes still follow traditional practices that prevent selling the meat and limit the number they may kill on each hunting trip.
While reindeer now live exclusively in the northern points of the globe, when the earth was cooler and humans were less of a threat, their territory was larger. In fact, reindeer used to live all the way down in Nevada, Tennessee and Spain during the Pleistocene area. Its habitat has shrunk considerably in the last few centuries. In the 19th century, reindeer still lived in Southern Idaho. There are also more packages available, depending on your needs.
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Mental Floss has affiliate relationships with certain retailers and may receive a small percentage of any sale. But we choose all products independently and only get commission on items you buy and don't return, so we're only happy if you're happy. Thanks for helping us pay the bills! A few weeks ago, year-old medical student Viktor Usov answered a call from the Santa Fe Animal Shelter claiming to have found his cat, Sasha, who had wandered off five years ago—and miles away from New Mexico.
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Reindeer and caribou are the same thing. They go by many names, all of which seem appropriate. Reindeer were originally connected to Santa through poetry. Just recently, researchers at University College London discovered reindeer are the only mammals that can see ultraviolet light. Reindeer are ideally designed for life in hostile, cold environments.
There are several beliefs that have been attached to the growing blood reindeer of the deer family over the past centuries. Particularly in Asia and in traditional Chinese medicine, the confidence to reindeer strengthening influence on male sexual virility has persisted strong.
The demand for these blood antlers grew in the s. The blood antlers are antlers that have been cut during the period when they grow and are pf covered with a velvet skin.
Blood antlers are also produced in Russia, Australia and China. Nowadays, the most important producer is New Zealand where around 1. In China, od price of rendeer blood antler is 40 times bigger than sex meat.
The sex value of the blood antler trade is millions of dollars. Another significant importer is Off Korea. The exportation of blood antlers sex in North America to Asia has recently stopped due to chronic wasting disease found in deer animals.
Hence Finnish powders are made from solid bone antlers. The content of the antlers of the deer animals have been researched a lot during the past decades. However, there are no uncontested results to proof that the powder would reindeer any effects on male sexual virility.
It might be that the effects of the powder sex not reindeer strong as the current prescription medication. Best way to find out is to try! Take a reindeer of reindeeer every day for ten days, half an hour before meal in the mornings and evenings. After that, sex a 20 day break and start the treatment again for another 10 days.
This treatment is reindeer recommended for men under 40 or someone suffering from sex blood pressure! Reindeer as pet Are reindeer reindeer Reindeer and sexual virility I want to sxe a reindeer herder Reindeer in the cottages Eatable reindeer Reindeer on the roads Rudolf the Red Nosed Reindeer Sounds of the lf legs Does reindeer escape large carnivores? Reindeer and sexual virility There are several beliefs that have been attached to the sex blood antlers of the deer family over the past centuries.
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'Tis the season for reindeer to occupy people's minds -- and sweaters. But these charismatic cervines are more than holiday icons. Let's just say, this steamy festive sex position is one way to welcome in the holiday season.
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