Escaping parasitism in the selfish herd: age, size and density-dependent warble fly infestation in reindeer

It has been suggested that animals may escape attack from mobile parasites by aggregating in selfish herds. A selfish herd disperses the risk of being attacked among its members and the per individual risk of parasite infection should therefore decrease with increasing animal density through the encounter–dilution effect. Moreover, in a selfish herd, dominant and agile animals should occupy the best positions and thereby receive fewer attacks compared to lower ranked animals at the periphery. We tested these predictions on reindeer ( Rangifer tarandus tarandus ) parasitized by warble flies ( Hypoderma tarandi ). Warble flies oviposit their eggs on reindeer during summer and induce strong anti-parasitic behavioural responses in the herds. In this period, reindeer are sexually segregated; females and calves form large and dense herds while males are more solitary. After hatching, the warble fly larvae migrate under the skin of their host where they encyst. In the present study encysted larvae were counted on newly slaughtered hides of male calves and 1.5 year old males from 18 different reindeer herds in Finnmark, northern Norway with large contrasts in reindeer density. In reindeer, body mass is correlated with fitness and social status and we hypothesized that individual carcass mass reflected the animal's ability to occupy the best positions within the herd. Larval abundance was higher among the 1.5 year old males than among the calves. For calves we found in accordance with the selfish herd hypothesis a negative relationship between larval abundance and animal density and between larval abundance and body mass. These relationships were absent for the 1.5 year old males. We suggest that these differences were due to different grouping behaviour where calves and females, but not males, aggregated in selfish herds where they escaped parasitism.     

Behavioural Correlates of Female Social Status and Birth Mass of Male and Female Calves in Reindeer

In polygyny, male breeding success depends more than female success on body size, and male birth mass may therefore be more strongly correlated with mother's social dominance status. Among female reindeer, Rangifer tarandus, the rate at which a female took over winter feeding craters from conspecifics depended on rank and the mass loss decreased with increasing rank. Birth mass of male calves depended on mother's social rank, but that of female calves did not. Male foetuses appeared to be more effective than females to utilize resources associated with mother's phenotypical superiority.     

Chillingham Cattle: Dominance and Affinities and Access to Supplementary Food

A herd of cattle of natural sex ratio and age distribution, inhabiting a 134-ha park in northern England, was studied during supplementary feeding in 4 winters. Interactions could be summarised by conventional dominance hierarchies, more strictly linear and less stable among males than among females. Personal associations among individuals were not important, but affinities among social classes were, in determining the composition of feeding groups. Dominant males often fed in the same groups as dominant females. Dominant animals were less often seen to feed alone, implying that social dominance did not confer exclusive access to food. Cattle often fed in groups of two or three; certain combinations (notably those including two males, or one male and one female, or three calves) were stable, others unstable, notably combinations of females and calves, or of two or three females. This implies that females may defend resources more vigorously against other females than males do against other males.     

Adaptive adjustment of offspring sex ratio and maternal reproductive effort in an iteroparous mammal

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.     

Fecal hormones as a non-invasive population monitoring method for reindeer

Proper management of threatened species requires knowledge of population sizes and structures, however current techniques to gather this information are generally impractical and costly and can be stressful on the animals. Non-invasive methods that can produce high quality and accurate results are better alternatives. In winter 2010, we collected blood and fecal samples from 2 reindeer (Rangifer tarandus) populations (Kaamanen, Finland and Svalbard, Norway) to investigate the feasibility of using fecal progesterone metabolites to help estimate the reproductive status, the sex, and the age structures of the populations. We first examined the relationship between plasma progesterone and fecal progesterone metabolite concentrations. We further assessed whether fecal progesterone metabolite levels would clearly differ among calf, yearling, and adult and between pregnant and non-pregnant females. We quantified fecal progesterone metabolites (using enzyme immunoassay) and plasma progesterone (using radio immunoassay) of females and males of different ages from the 2 herds. We found in both populations that fecal progesterone metabolite levels reflected plasma progesterone concentrations. However, the range of fecal progesterone metabolite concentration was much wider in Finland than in Svalbard, possibly due to differences in diet or body condition. We determined a threshold value of 1.31 ng/ml plasma progesterone and 2025.93 ng/g dried fecal progesterone metabolites to identify pregnant reindeer from non-pregnant animals with 100% accuracy. We found a significant difference in fecal progesterone metabolite concentrations only between calves and yearlings/adults in Finland. We could not differentiate among males, non-pregnant adults, or calves of either sex; therefore identification of sex may have to rely on the use of DNA techniques. Our results suggest that hormone concentration, in combination with fecal DNA and pellet morphometry techniques, may provide important population parameters and is a valuable tool for the monitoring of reindeer and may have an application for threatened populations of woodland caribou throughout the winter and early spring. © 2011 The Wildlife Society.     

Evidence of alphaherpesvirus infections in Alaskan caribou and reindeer

Background  

The reindeer (Rangifer tarandus tarandus) industry in Alaska began with animals imported from Siberia (Russia) in the 1890's. Cervid herpes virus 2 (CvHV2) is endemic in reindeer in Scandinavia. We sought to determine if the same virus, or similar herpesviruses, were circulating in Alaskan reindeer and caribou (Rangifer tarandus granti). Serum samples from 292 reindeer were collected during annual reindeer handlings (1988-2005) near Nome, Alaska. In 2005, swab samples were collected from 40 calves from this herd, near Nome, Alaska. In 2007, ocular and nasal swab samples were collected from 30 apparently healthy reindeer calves near Wales, Alaska. Samples of plasma and white blood cells were collected from three Alaskan caribou herds, Mulchatna (n = 24), Teshekpuk (n = 34) and the Western Arctic (n = 87) in 2009.     

Microsatellite DNA evidence for genetic drift and philopatry in Svalbard reindeer

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

Induced orphaning reveals post-weaning maternal care in reindeer

A common by-product of human harvesting is orphaning of calves in autumn. Despite this, there are few studies in northern and temperate ungulates evaluating the fate of orphaned calves and the potential benefits to offspring and costs to mothers of post-weaning maternal care. We manipulated orphaning and forage distribution during winter for two herds of reindeer (Rangifer tarandus): one nonfed and the other supplementally fed to increase level of interference competition. Both herds consisted of females with and without calves at heel and orphaned calves. We measured survival and somatic losses during winter and distances between mother–calves and adult females–orphans within the herds. All females survived the winter, and there was no evidence of post-weaning maternal cost in terms of female’s mass loss. The winter mortality among calves was negligible and did not differ between orphans as compared to nonorphans. However, nonorphaned calves lost less mass and stayed closer to their mothers than orphans to adult females, suggesting that increased mortality might occur in harsher winters. This tended to be more marked in the fed group where interference competition was more likely due to feed being concentrated both in space and time. Reduced mass loss in nonorphans is therefore most likely due to mothers sharing and defending feeding resources and protecting their offspring from harassment by other herd members during their first winter. We conclude that hunting practise of northern and temperate ungulates where females having calf/calves at heel are intentionally or non-intentionally harvested, may have demographic side effects at least in harsh winters.     

Dispersal of yearling yellow-bellied marmots (Marmota flaviventris)

Field studies of the social organization of yellow-bellied marmots (Marmota flaviventris) in the vicinity of Gothic, Colorado, were initiated in 1962. Data collected during the subsequent 14 years are sufficient to analyse the behaviours that correlate with the time yearling male and yearling female marmots leave their natal home sites. Dispersal of yearling male yellow-bellied marmots was delayed when there were many yearling males in a harem, when yearling males were underweight, and when the rates of amicable interactions between yearling males and adults were high. Dispersal of yearling females was delayed when adults behaved amicably toward them and when the rates of adult aggression were low. Dispersal of yearling females was independent of the number of females in a harem. These observations are in accord with a model of dispersal that relates sexual differences in dispersal to maternal reproductive tactics, and to responses of yearlings that would maximize their own fitness.     

Mating Group Size and Stability in Reindeer Rangifer tarandus: The Effects of Male Characteristics,Sex Ratio and Male Age Structure

In polygynous mating systems, males compete intensely for mates and may mate several females during a single reproductive season. Accordingly, factors influencing the ability of males to control a larger number of females during the breeding season can provide information on the processes underlying sexual selection. In ungulates, age, body mass and social rank are considered good predictors of the reproductive success of males, but how male age structure and sex ratio in the population influence mating group (MG) dynamics has received little empirical testing. Between 1996 and 2005, we manipulated male age‐ and sex structure and monitored MG dynamics in a reindeer (Rangifer tarandus) population. We investigated the influence of male characteristics, percentage of males and male age structure on MG size and stability. We found that males with higher social rank (that were also older and heavier) controlled larger MGs (therefore had greater mating opportunities) and had more stable MGs (corresponding to a higher ability to maintain and control females) than males of lower social rank. Moreover, MG size and MG stability decreased as the percentage of males in the population increased, most likely resulting from greater male–male competition and increased female movements. Male age structure did not influence MG stability. Given the positive relationship between mating success and MG size (and likely MG stability), frequent female movements and intense competition among males to control females seem to be the principal components of reindeer MGs dynamic.     

Social behaviour of the calves of semi-wild Maremma cattle,Bos primigenius taurus

The social behaviour of semi-wild free-ranging Maremma calves, Bos primigenius taurus, was observed daily in their first 2 months from birth at the estate La Trappola (Ponticelli Reserve), 10 Km south-west of Grosseto, Tuscany, Italy. In the first 2–3 days of life the calves hid in the maquis, while the cows grazed in the surrounding area. The time spent at close range to the cows decreased with the increase in the age of the calves, but the time spent far from the mother was always greater than that spent at close or intermediate ranges. Lying down was the dominant activity at a distance over 15 m from the mother, while the time spent in standing still decreased daily. The daily mean of the minutes spent by the calves with their peers reached peaks between their 11th and 40th days of life. Sucking bouts reached a peak between 09.00 and 10.00 h, after the herd reached the grazing area from the night dormitory. Another peak occurred between 15.00 and 16.00 h. Calves solicited and terminated sucking bouts spontaneously more than cows. The mean duration of each sucking bout increased with calf age, but was negatively correlated with the mean daily number of sucking bouts. Immediately before or after a sucking bout, cows mainly sniffed their calves in the muzzle and tail area. Calves sniffed and licked the cows mainly in the area of the udder. Play activity reached a peak in the early morning, decreased strongly at mid-day and increased again in mid-afternoon. Grazing, and to a lesser extent moving to and from the pasture land, were the most frequent activities occurring before and after play. Frontal pushes were the most preferred kind of play by both females and males. Female calves soliciting play in males showed significantly more “frontal pushing” than “non-reproductive mounts”. The male calves were more often initiators of play than were females.     

Taxonomic Notes on the Swedish Domestic Reindeer, Genus Rangifer

In order to study the taxonomic relationship between domestic mountain and forest reindeer in Sweden nine skull characters on 126 and 62 animals, respectively, were measured and analysed. In addition antler width and shape of antler beam in cross-section were also measured. The differences indicate that the two groups of domestic reindeer are not entirely identical, although the differences are clearly not as plain as has been claimed. Comparing these results with earlier results it also appears that domestic forest reindeer are more in accordance with Rangifer tarandus tarandus than with R. t. jennicus , thus supporting the theory that possible influence of the latter subspecies on domestic reindeer has been considerably reduced as a result of long and intense cross breeding between the two groups of domestic reindeer.     

Phantoms of the forest: legacy risk effects of a regionally extinct large carnivore

The increased abundance of large carnivores in Europe is a conservation success, but the impact on the behavior and population dynamics of prey species is generally unknown. In Europe, the recolonization of large carnivores often occurs in areas where humans have greatly modified the landscape through forestry or agriculture. Currently, we poorly understand the effects of recolonizing large carnivores on extant prey species in anthropogenic landscapes. Here, we investigated if ungulate prey species showed innate responses to the scent of a regionally exterminated but native large carnivore, and whether the responses were affected by human‐induced habitat openness. We experimentally introduced brown bear Ursus arctos scent to artificial feeding sites and used camera traps to document the responses of three sympatric ungulate species. In addition to controls without scent, reindeer scent Rangifer tarandus was used as a noncarnivore, novel control scent. Fallow deer Dama dama strongly avoided areas with bear scent. In the presence of bear scent, all ungulate species generally used open sites more than closed sites, whereas the opposite was observed at sites with reindeer scent or without scent. The opening of forest habitat by human practices, such as forestry and agriculture, creates a larger gradient in habitat openness than available in relatively unaffected closed forest systems, which may create opportunities for prey to alter their habitat selection and reduce predation risk in human‐modified systems that do not exist in more natural forest systems. Increased knowledge about antipredator responses in areas subjected to anthropogenic change is important because these responses may affect prey population dynamics, lower trophic levels, and attitudes toward large carnivores. These aspects may be of particular relevance in the light of the increasing wildlife populations across much of Europe.     

Summer movements,predation and habitat use of wolves in human modified boreal forests

Grey wolves (Canis lupus), formerly extirpated in Finland, have recolonized a boreal forest environment that has been significantly altered by humans, becoming a patchwork of managed forests and clearcuts crisscrossed by roads, power lines, and railways. Little is known about how the wolves utilize this impacted ecosystem, especially during the pup-rearing summer months. We tracked two wolves instrumented with GPS collars transmitting at 30-min intervals during two summers in eastern Finland, visiting all locations in the field, identifying prey items and classifying movement behaviors. We analyzed preference and avoidance of habitat types, linear elements and habitat edges, and tested the generality of our results against lower resolution summer movements of 23 other collared wolves. Wolves tended to show a strong preference for transitional woodlands (mostly harvested clearcuts) and mixed forests over coniferous forests and to use forest roads and low use linear elements to facilitate movement. The high density of primary roads in one wolf’s territory led to more constrained use of the home territory compared to the wolf with fewer roads, suggesting avoidance of humans; however, there did not appear to be large differences on the hunting success or the success of pup rearing for the two packs. In total, 90 kills were identified, almost entirely moose (Alces alces) and reindeer (Rangifer tarandus sspp.) calves of which a large proportion were killed in transitional woodlands. Generally, wolves displayed a high level of adaptability, successfully exploiting direct and indirect human-derived modifications to the boreal forest environment.     

Courtship Disruption at the Lek in the Guianan Cock-of-the-Rock

During 4 seasons of study at a large (55 male) lek of Guianan cock-of-the-rock (Rupicola rupicola) in Suriname, we observed 3906 female courtship visits. Of these, 1171 (30.0%) were disrupted by adult males, 430 (11.0%) were disrupted by yearling males, and 289 (7.4%) were disrupted by females. This is one of the highest levels of courtship disruption reported among lek-breeding birds. Disruption was performed by approximately half of the territorial adult males each year, spanning the full range of mating success and territory location. Courtship disruption occurs in two contexts among adult male cock-of-the-rock. The most frequently performed disruptions were low-intensity supplantings and threats directed toward more successful territorial neighbors. These appear to reflect attempts to improve or maintain territorial status within the male social organization. Less commonly, individual males persistently directed high-intensity disruption toward specific females. Such males succeeded in redirecting the females' mate choice to themselves in 27% of instances. We found no support for the hypothesis that females choose males on the basis of their immunity to disruption. Reproductively successful males received disruption at proportionately equal or greater levels than unsuccessful males. Courtship disruption by yearling cock-of-the-rock occurred primarily in the context of practice courtship by these non-territorial immatures. Disruption by females was uncommon and did not appear to deny or even delay access to any male. We discuss these results in the light of recent proposals concerning the role of courtship disruption in the operation of mate choice and the determination of male dispersion on leks.     

The effects of twinning on mother—Offspring behavior in range beef cattle

Methods to increase the productivity of beef cattle include induced twinning and multiple fostering of calves on nurse cows, but little is known about the capacity of cows to rear more than one calf in a rangeland environment.Eight single and 14 twin calves and their mothers were studied in a 30.4-ha enclosure in a California rangeland environment from 2 to 20 weeks following parturition. Data were obtained on ingestive, grooming, agonistic, play and sexual behaviors, as well as spatial proximity of calves to mothers and peers.Initially, twins spent more time suckling their mothers than did single calves (presumably due to insufficient milk intake), but later adapted by utilizing natural forage and obtaining milk from alien cows. By 15–20 weeks of age, twins were suckling their dams for less than one-quarter of the time spent at 3–6 weeks, whereas the suckling time of single calves did not decline. Thus, twins may experience a natural early weaning from mother's milk.The hypothesis that mother—Offspring bonds are weaker with twins than with singles was supported by the fact that mothers bearing twins groomed their offspring less than mothers with singles, and twins were normally found at greater distances from their mothers than singles.Most long-distance cow—Calf separations resulted from the cow moving to feed or to obtain water. Reunion of mother and young depended on auditory communication; the calf normally traversed most of the intervening distance.     

Identification and breeding of yearling Piping Plovers

ABSTRACT Age of first breeding is an important life history trait. Many Piping Plovers (Charadrius melodus) do not breed as yearlings, but little information is available concerning the age of first breeding. From 2000 to 2006, we marked 991 chicks in three areas in Saskatchewan, Canada, and subsequently determined when 102 (49 females and 53 males) first bred. Females bred significantly earlier, on average, than males. More females (68%) bred as yearlings than did males (41%; P= 0.04), with most others first nesting when 2‐yr old (29% of females and 50% of males). As expected from differences between the sexes in age of first breeding, younger females were more likely to pair with older males than were younger males with older females. Chicks that hatched early in the breeding season did not breed at an earlier age than those that hatched late in the year. Unlike older birds, juvenile Piping Plovers do not replace flight feathers during their first winter. As a result, 18 of 27 yearlings (67%) had worn outer primaries, whereas only one of 123 (1%) older birds had worn primaries. In addition, whereas 20 of 24 yearlings (83%) retained a few buff‐tipped median coverts, none of 119 known older birds had such coverts. As a result, we were able to identify all yearlings by their worn primaries, buff‐tipped median wing coverts, or both. Wing lengths of yearling Piping Plovers were 3% shorter than those of older birds, presumably due to wear. Because there is no evidence of differences in adult survival rates between the sexes and breeding habitat is available, we speculate that fewer yearling males than females breed because primary wear may reduce the ability of yearling males to perform aerial breeding displays.     

Birth-season interactions of adult female Japanese Macaques(Macaca fuscata) without newborn infants

The affiliative interactions of 11 adult female Japanese macaques that did not deliver an infant during the 1981 birth season of the Arashiyama West troop were examined. Consideration was given to the effects of kinship as a structuring element in these birth-season interactions and to the degree of association with various categories of troop members based on age, sex, and (in the case of adult females) whether or not the females were new mothers. Females without infants interacted predominantly with their yearling off-spring, although it was the behavior of the offspring that precipitated the interaction. These females were active in soliciting affiliation with nonkin new mothers, whereas female matrilineal relatives with new infants approached and remained in proximity to them more than did nonrelated new mothers. Females without newborns groomed and approached nonkin infants more than infants within their own matriline, and these infants were predominantly those of females in the highest-ranking matriline of the troop. Adult males were responsible for 40% of all grooming received from nonkin by the females without newborns, and these males approached them significantly more than did other adult females without infants. These patterns demonstrate that the structure of social relationships is influenced by the particular dynamics of troop contexts such as birth seasons, as well as by enduring, broad-based affinities which are less affected by cyclic changes in troop context.     

Alternative Mating Behaviors of Young Male Bullfrogs

Yearling male bullfrogs often attempt to intercept and matewith females attracted to older, larger males (male parasitism)instead of defending a territory and attracting their own mates.Mating behaviors like male parasitism are thought to occur onlywhen individuals suffer some strong disadvantage in using thebehaviors that are more typical of their species. In this paper,I consider the relative benefits and costs of territorialityvs male parasitism for yearling males. Yearlings employing maleparasitism are only occasionally successful at mating, and inyears of reduced male-male competition, they produce significantlyfewer young than territorial males of their age. Yearling malesthat vary in the degree to which they employ territorial behaviorsinstead of male parasitism do not differ measurably in growthrate or chances of mortality; however, indirect informationsuggests that territoriality should entail a greater chanceof mortality. A computer simulation based solely on observeddemographic parameters predicts that yearling males should beterritorial whenever they can compete as effectively as two-year-oldmales. This condition appears to provide little restrictionon the breeding tactics of yearling males; however, data onmale-male aggressive encounters show that the size differencesbetween one-year-old and two-year-old males are usually sufficientto preclude yearling males from competing successfully. Thus,the observed behavioral plasticity of yearling males may bebest interpreted as a means of adjusting their behavioral tacticsto levels of male-male competition that can vary within andbetween breeding seasons.     

Use of climatic data to assess the effect of insect harassment on the autumn weight of reindeer (Rangifer tarandus) calves

Considerable behavioural evidence supports harassment by insects as the most important causal link between warm summer temperatures and low body condition of reindeer Rangifer tarandus , and that insect activity is influenced by weather condition. However, much less is known about the effect of insect harassment on individual performance, measured as reduced weight gain during summer, and the related consequences on both the reindeer pastoral economy and reindeer as a biological resource. Using climatic data, this paper develops a simple index for the analysis of insect harassment that takes into consideration weather variables known to significantly affect insect activity and/or the level of insect harassment. The insect harassment index, which is based on mid-day ambient temperature ≥13 °C, wind speed <6 m/s and cloud cover <40%, is further used to test the hypothesis that insect harassment has a negative effect on reindeer performance during summer in three Norwegian populations. Results show that harassment by insects negatively affects the autumn weight of reindeer calves, most probably through reduced grazing time and increased energy expenditure, but also indirectly by negatively influencing milk production of the dam. Moreover, female calves were more vulnerable to insect harassment than males. Insect harassment may have consequences on future reproductive performance, calving time, calf birth weight and hence neonatal mortality, and thus affect reindeer productivity. The presented index is easy to estimate and may be used to quantify and compare harassment levels on various reindeer summer grazing areas for management purposes. Our results also suggest that the expected temperature increase in the course of global warming may increase the insect-related stress on reindeer.