Abstract: A fundamental goal of sociobiology is to explain how complex social behaviour evolves, especially in social insects, the exemplars of social living. Although still the subject of much controversy, recent theoretical explanations have focused on the evolutionary origins of worker behaviour (assistance from daughters that remain in the nest and help their mother to reproduce) through expression of maternal care behaviour towards siblings. A key prediction of this evolutionary model is that traits involved in maternal care have been co-opted through heterochronous expression of maternal genes to result in sib-care, the hallmark of highly evolved social life in insects. A coupling of maternal behaviour to reproductive status evolved in solitary insects, and was a ready substrate for the evolution of worker-containing societies. Here we show that division of foraging labour among worker honey bees (Apis mellifera) is linked to the reproductive status of facultatively sterile females. We thereby identify the evolutionary origin of a widely expressed social-insect behavioural syndrome, and provide a direct demonstration of how variation in maternal reproductive traits gives rise to complex social behaviour in non-reproductive helpers.

Abstract: Abstract Analysis of salivary cortisol is increasingly used to assess stress responses in horses. Since spontaneous or experimentally induced increases in cortisol concentrations are often relatively small for stress studies proper controls are needed. This requires an understanding of factors affecting salivary cortisol over longer times. In this study, we have analysed salivary cortisol concentration over 6 mo in horses (n = 94) differing in age, sex, reproductive state and housing. Salivary cortisol followed a diurnal rhythm with highest concentrations in the morning and a decrease throughout the day (P < 0.001). This rhythm was disrupted in individual groups on individual days; however, alterations remained within the range of diurnal changes. Comparison between months showed highest cortisol concentrations in December (P < 0.001). Cortisol concentrations increased in breeding stallions during the breeding season (P < 0.001). No differences in salivary cortisol concentrations between non-pregnant mares with and without a corpus luteum existed. In stallions, mean daily salivary cortisol and plasma testosterone concentration were weakly correlated (r = 0.251, P < 0.01). No differences in salivary cortisol between female and male young horses and no consistent differences between horses of different age existed. Group housing and individual stabling did not affect salivary cortisol. In conclusion, salivary cortisol concentrations in horses follow a diurnal rhythm and are increased in active breeding sires. Time of the day and reproductive state of the horses are thus important for experiments that include analysis of cortisol in saliva.

Abstract: Feeding habits of free-ranging wildebeest and zebra were monitored in a semi-arid nature reserve, bordering the southwestern part of Kruger National Park, South Africa. The purpose of study was to distinguish and define the feeding niches of two roughage grazers that occur in similar habitat types. The monthly compositions of diets were evaluated by direct observations of feeding bouts over a period of two years when rainfall patterns were average and animal populations were stable. Other analyses evaluated the standing biomass of grass species in the reserve during the wet summer and dry winter seasons.
A considerable overlap of grass species composition was found in the diets of wildebeest and zebra. Ordination of bi-monthly records of the diet composition showed greater variations in scores of grasses in zebra diet in comparison to wildebeest. Seasonal patterns were more apparent in the wildebeest diet. Preference ranking of grass species indicated that zebra diet remained constant in winter and summer. Wildebeest diet however, alternated with seasons, showing high preferences during the winter months for grass species which were rejected during summer.
The combined assessment of results from three separate statistical methods analysing temporal patterns and preferences in diet composition revealed contradictory trends. The solution, however, relied on the initial assumptions posed. Hence, wildebeest and zebra are essentially generalist feeders which show a limited amount of preference in their choice of diet.

Abstract: This experiment investigated the effects of positive and negative tactile handling on the stress physiology and behaviour of dairy heifers. Forty-eight 5-14-month-old nonlactating Holstein-Friesian heifers were allocated to one of two handling treatments, either positive or negative tactile handling, over four time replicates. Handling was imposed twice daily, 2-5 min per session and involved moving animals individually along a 64 m outdoor route. The negatively handled heifers took longer to approach within 1 and 2 m of a stimulus person in a standard test, than their positively handled counterparts (P<0.001) and had a greater flight distance to an approaching stimulus (P<0.001). The time taken by the heifers to approach within 1 and 2 m of a familiar person was similar to that taken to approach within 1 and 2 m of an unfamiliar person in the standard test (P<0.05). There was a tendency for heifers to have a greater flight distance from the approaching unfamiliar person than from the approaching familiar person (P=0.06). The negatively handled heifers had greater (P<0.05) increases in total cortisol concentrations 5, 10 and 15 min after exposure to a human and had higher (P<0.05) free cortisol concentrations in the afternoon than the positively handled heifers. It is concluded that the nature of the human contact affects the subsequent behavioural response of heifers to humans. This behavioural response may extend to other humans through the process of stimulus generalisation, although there was some evidence of moderate discrimination. Negative handling results in an acute stress response in the presence of humans and also leads to a chronic stress response. Further research into the effect of these stress responses on milk production and welfare in fearful cows in a commercial situation is suggested.

Abstract: Mortality is high in zoo-housed black rhinoceros (Diceros bicornis), and the reproductive rates of captive white rhinoceros (Ceratotherium simum) are unsustainably low. To determine the possible role of stress in the causation of these problems, we analyzed weekly fecal samples collected for 1 year from black (10 males and 16 females) and white (six males and 13 females) rhinoceroses at 16 zoos for corticoid metabolite concentrations. Fecal corticoid profiles were examined in relation to behavior as rated by keepers in a questionnaire, luteal phase ovarian cycles of females (Brown et al., 2001), and socioenvironmental factors. We compared individual fecal corticoid profiles by examining hormone means and variability (i.e., standard deviation (SD) and coefficient of variation (CV)). For the black rhinos, higher mean corticoid concentrations were found at zoos where rhinos were maintained in enclosures that were exposed to the public around a greater portion of the perimeter. Higher variability in corticoid excretion was correlated with higher rates of fighting between breeding partners and higher institutional mortality rates. Black rhino pairs that were kept separated exhibited lower corticoid variability and less fighting activity when they were introduced during female estrous periods compared to pairs that were kept together every day. For white rhinos, significantly lower mean corticoids were found for individuals that rated higher on “friendliness to keeper.” Higher corticoid variability was found in noncycling as compared to cycling white rhino females. Noncycling females exhibited higher rates of stereotypic pacing and lower frequencies of olfactory behaviors. Interindividual differences in mean corticoids in both species appeared to be related to responsiveness to humans, whereas corticoid variability was related to intraspecific social relationships. More importantly, high corticoid variability appeared to be an indicator of chronic or “bad” stress, because of its association with potentially deleterious consequences in each species (i.e., fighting and mortality (black rhino), and reproductive acyclicity (white rhino)). Our results provide evidence that social stressors may cause chronic stress in black and white rhinos, and that this contributes to the captive-population sustainability problems observed in each species. Zoo Biol 0:1–18, 2005. © 2005 Wiley-Liss, Inc.