One or several factors could explain sexual segregation, in which males and females of polygynous, sexually dimorphic species form separate herds during most of the year. Bighorn sheep (Ovis canadensis) are polygynous ungulates that exhibit extreme sexual dimorphism and segregate into ram and ewe herds outside of the rutting season. Four major hypotheses for sexual segregation were tested in a population of desert bighorn (O. c. mexicana) at the Red Rock Wildlife Area, New Mexico, from 1999-2001. We collected data on the size, composition, and location of ram and ewe groups during the summer period of segregation. Activity budgets were recorded for males in ram herds and females in ewe herds, and foraging selectivity was measured for males and females in mixed groups during early rut. Habitat was evaluated by measuring forage availability, ruggedness, and visibility at sites utilized by ram and ewe groups. Ram herds utilized areas with more available forage compared with ewe sites, while ewe groups preferred more rugged terrain than that used by ram groups. Ewe groups occurred much closer to free water sources than did ram groups. Bighorns in ram and ewe groups did not differ in foraging time or selectivity, nor did time spent moving, reclining, or ruminating differ between the sexes as predicted by the 'activity budget hypothesis'. The results support the predictions of the 'reproductive strategy-predation risk hypothesis', which proposes that males seek more abundant forage in order to build up body condition needed to maximize mating success (even if exposing themselves to greater predation risk), while females choose rugged terrain that minimizes predation risk to themselves and their offspring (even if sacrificing forage abundance). Female bighorns chose sites that provided access to water, also predicted by the 'reproductive strategy-predation risk hypothesis', indicating that lactation-related water requirements may constrain the movements of ewe groups and contribute to patterns of sexual segregation in desert bighorn.

Abstract Biting insects impose costs on hosts, including decreased feeding or resting time as the result of disturbance, blood loss, and disease transmission. Insect-repelling behaviors, such as ear-flicking, head-shaking, stamping, and grouping, have evolved in many ungulate species to minimize these costs. We studied female desert bighorn sheep (Ovis canadensis mexicana) at Red Rock Wildlife Area, New Mexico, during the summers of 1999 and 2000. We tested the predictions that: 1) bighorn sheep will increase insect-defense behavior when biting insects are more abundant, and 2) close aggregation of sheep will decrease the per capita insect harassment by means of a dilution effect. Numbers of midges and other biting insects increased in association with rising temperature and decreased with increasing wind speed. Ewes performed between 0 and 78 ear-flicks/min, and >5,000 ear-flicks over the course of a 12-h day. As predicted, the rate of ear-flicking was positively correlated with counts of biting insects, indicating that ear-flicking was a direct response to the irritation of attacking insects. We also found a negative correlation between the number of sheep clustered together within 1 body length and ear-flicking rate, suggesting that insect harassment is diluted when bighorn sheep bunch together. Bighorn sheep generally bedded on upper slopes and rocky outcrops exposed to gusts of wind. These results indicate that ear-flicking, grouping, and microhabitat choice might be important strategies for reducing the costs of biting insects in desert bighorn sheep.

Tick removal grooming may be centrally regulated by an internal timing mechanism operating to remove ticks before they attach and engorge (programmed grooming model) and/or evoked by cutaneous stimulation from tick bites (stimulus-driven model). The programmed grooming model predicts that organismic and environmental factors that impact the cost-benefit ratio of grooming (e.g. body size and habitat) will influence the rate of tick removal grooming. The body size principle predicts that smaller-sized animals, because of their greater surface-to-mass ratio, should engage in more frequent tick removal grooming than larger-bodied animals in order to compensate for higher costs of tick infestation. The body size principle may be tested intraspecifically between young and adult animals, or interspecifically among species of contrasting body sizes. To rigorously test the interspecific body size prediction, we observed the programmed grooming (oral and scratch grooming) of 25 species (or subspecies) of bovids at a tick-free zoological park in which stimulus-driven grooming was ruled out. Multiple correlation analysis revealed highly significant negative correlations between species-typical mass and mean species grooming rates when habitat was controlled for in the model. Species-typical habitat type (classified along a gradient from most open to most closed) was positively correlated with mean oral grooming rate, indicating that species tended to groom at a higher rate in woodland and forest habitats (where typical tick density would be high) compared with more open environments. Species mass accounted for up to two-thirds of the variation in grooming rate across species, whereas habitat accounted for ca. 20% of variation in oral grooming. Similar results were obtained when the analysis was expanded to include 36 species/subspecies of six different families. The body size principle can therefore account for a large proportion of species-typical differences in programmed grooming rate among ungulates. However, to understand the tick defence adaptations of very large mammals that rarely or never engage in oral or scratch grooming (e.g. elephants, giraffes, rhinoceros), alternative tick defence strategies must be considered, such as thick skin, wallowing, rubbing and tolerance of oxpeckers and other tick-eating birds. Copyright 2000 The Association for the Study of Animal Behaviour.