Foraging efficiency and vigilance behaviour of impala: the influence of herd size and neighbour density

Group foraging can be beneficial for ungulates by decreasing the time required for vigilance, but it can also prove costly because of competition. To determine responses to gregarious behaviour, we studied foraging activity and vigilance of impala ( Aepyceros melampus ) near Kruger National Park, South Africa. We measured time spent foraging, vigilant, moving, grooming, engaging in social interactions and determined herd size and group distribution (i.e. density). We calculated accepted food abundance (AFA), food ingestion rate, steps per minute and percent vigilance for female, bachelor male and herd male impala. There was no relationship between herd size and vigilance, but vigilance decreased with increasing density ( t _1,311 = 4.91, P  <0.0001). Additionally, AFA decreased ( t _1,61 = 5.96, P  <0.0001) and steps per minute increased ( t _1,311 = 14.38, P  <   0.0001) as more individuals fed in close proximity to each other. Impala could be altering their behaviour to accommodate a perceived change in resources because of intraspecific competition and these adjustments might be related more to the distribution of herd members than to herd size. Further studies should examine the behaviour of gregarious animals in relation to the distribution of herd members in addition to group size.     

Male and female agonistic and affiliative relationships in a social group of farm cats (Felis catus L.)

A dominance hierarchy based on the outcome of agonistic encounters was found among male and female domestic cats. A female dominated over some males. The dominance concept is also discussed in terms of social bonding. The relationships among adult females were amicable, whereas adult males showed reciprocal tolerance. The flow of affiliative behaviour was directed mainly from females to one male of the group. The analysis of marking behaviour showed that this male sprayed urine and rubbed the perioral and cheek regions of the face on the objects of the environment at a higher rate than the other members of the group. Nevertheless, rubbing the perioral and cheek regions of the face on objects was not correlated to dominance rank, possibly because it has some function in social communication other than territorial defence against strangers. No relationships have been found between claw scratching, rolling on the ground and social rank, or between the former and other marking behaviour. It is concluded that claw scratching and rolling were not utilised to mark territory.     

Impala social behaviour: birth behaviour

Data collected from observations of eighteen parturient impala females are presented. Females attempt to isolate from the breeding herds 2 to 3 h before parturition. They have a characteristic appearance and behaviour at this stage and are having obvious labour contractions. Nearly all the females sought some cover and a vegetation type including many bushes was used frequently as a birth site. Only four of eleven females left the herd and found cover without harassment by either the territorial male or bachelor males. Many were also followed by impala females. The birth process is described; final expulsion of the foetus occurs in a recumbent position. All birth fluids and membranes are eaten by the mother. Much attention is paid to cleaning up the birth site, baby and herself. Time from birth to expulsion of the placenta is variable and the placenta is normally eaten as soon as it is expelled. Impalas can stand at about 15 min after birth and attempt to suckle soon after this. Only some mothers help the baby to find the udder. Unsteady running occurred at about 25 min after birth. Impalas have a preference for giving birth between 10·00 and 14·00 hours. The significance of this behaviour and the other behaviours shown as anti-predator devices are discussed. Individual differences in behaviour and early care of the young are noted.     

Activity and space patterns of Pyrenean desman (Galemys pyrenaicus) suggest non-aggressive and non-territorial behaviour

The Pyrenean desman is considered a flagship species for biodiversity and evolution. However, its scientific knowledge is still under development and currently on debate, particularly in relation to its behavioural ecology and social organization. Based on the previous hypothesis of individual desmans being solitary and territorial, activity and space patterns were considered to be arranged to avoid conspecifics. However, recent insights into the species’ social behaviour revealed non-interspecific avoidance. With this study, we provide novel insights into the activity and space patterns of the desman, and their relation to its social behaviour. A total of 30 individuals were trapped, 18 of which provided informative radiotracking data to study (1) activity behaviour, (2) the proportion of the home range utilized and the distances travelled, and (3) the dominant movement directionality. Activity and space use patterns were affected by daylight and seasonality, but not by sex, age or number of other conspecifics sharing the home range. In contrast to the previous observations, individuals did not show a pattern of directionality in their movements. Noticeably, we observed encounters between individuals without evidence of aggressive behaviour. Our results suggest that desmans do not alter their activity or their spatial behaviour to avoid encounters with conspecifics, as previously suggested. These novel findings provide more evidence of a social structure and organization with social interactions and non-aggressive behaviour. This is of relevance for management actions and for the conservation of this endemic mammal.     

Social behaviour and heart rate in tammar wallabies (Macropodidae: Macropus eugenii)

In an enclosed group of tammar wallabies the behaviour of all individuals and the heart rate (HR) of three males were recorded simultaneously. The social structure was characterized by the dominance relationships between the males and the sexual preference of all males for the oldest female. During non-oestrus, dominance was obvious when a male intervened terminating the sexual behaviour of a subdominant one. The relatively low amount of agonistic behaviour between the males is assumed to be due to the clear dominance relationships. Locomotion as well as sexual and agonistic behaviour were accompanied by an acceleration of HR. The HR pattern depended on the intensity of the behaviour, its duration and the animal's identity. Commonly performed sexual behaviour and approach/retreat encounters gave rise to HR values not different from HR values during locomotor activity without interactive components. Social events that occurred infrequently—like fights, and the mating of an oestrous female—were accompanied by strong accelerations of HR indicating high cardiac effort. The mild cardiac activity during common social encounters is consistent with a strategy of maximizing energy conservation. HR patterns during specific behaviour could not entirely be accounted for by the energetic costs of activity. Spreading the forelimbs in response to specific stimuli—like the fly-over of raptors—was indicative of a strong HR response even if the animal was motionless. During specific social encounters like the sexual behaviour of the alpha-male following an intervention—HR responses revealed that arousal might exceed motor activity in affecting HR. Individual differences of these HR responses are attributed to the age, experience and social status of a male.     

The contributions of genotype and sex to variation in mating behaviour between geographical subspecies of the bank vole (Clethrionomys glareolus scheber)

The sexual behaviour of bank voles from the mainland of Great Britain, and from the island of Skomer, differs in a number of respects. Voles from the mainland population show in general greater sexual activity. These variations may be understood as adaptations to ecological differences, particularly in population density. The reciprocal hybrids resemble the mainland population in a number of aspects of their mating behaviour. The possible evolutionary importance of this dominance is discussed. The behaviour of the bank vole is put into the perspective of generalizations made about other mammals. The separate influence of the male and of the female on quantitative variation between the two forms was investigated by studying mating between members of different populations. Some aspects of mating behaviour are shown to be due primarily to the male, others primarily to the female and others to the particular combinations involved. This situation is discussed in relation to the separate action of natural selection on members of each sex.     

Sexual Interference in the Sword-tailed Newt,Cynops ensicauda popei (Amphibia: Salamandridae)

This paper describes competitive interaction between males of the salamandrid Cynops ensicauda popei. The sexual behaviour of the male is influenced by the presence of a second, rival, male. Competition for access to the female and other male interactions continuously interrupt ongoing courtships. Spermatophore pick-up success is lower than in encounters involving single pairs. During triad encounters, fewer spermatophores were deposited than in dyad encounters, due to interference during the creep stage of courtship. Two basic forms of male sexual interference were distinguished: ‘lure away’ during the creeping stage, from a position that is different from the direction of creep; and ‘female mimicry’, during which the male presses his snout alternately against the courting male's tail and the female's snout. In both forms, a rival male may shove one of the courting individuals away. The repertoire of sexual behaviour patterns of Cynops ensicauda popei appears to be less complex and varied than that of most Triturus species. The courtship of Cynops places less emphasis on display behaviour with tail and body, and a stronger emphasis on the creeping stage, where the male carefully leads the responsive female over a series of spermatophores, but during which rival males may interfere in a variety of ways.     

Reciprocal Allogrooming in Dam-reared and Hand-reared Impala Fawns

Among adult females and males of African antelope impala are unique in their performance of reciprocal allogrooming. The occurrence of this behaviour in neonatal impala fawns was explored in a free-ranging impala herd at the San Diego Wild Animal Park where 5 dam-reared fawns were observed from birth through 10 weeks of age. One-way maternal grooming and reciprocal allogrooming with the dam and non dam partners emerged as distinct behavioural systems. Maternal grooming, directed mostly to the anogenital area, was typical of that seen in other ungulates, and sharply declined over the first two weeks. Reciprocal allogrooming, characterized by alternate exchanges of grooming bouts with a partner in the same manner as in adults, was seen as early as 3–8 d after birth. All fawns were grooming with unrelated adult females by the end of the second week. By week 2 virtually every measure of reciprocal allogrooming by fawns (grooming delivered per hour, reciprocity, and percent of encounters initiated) was as high as for adults. The appearance of this reciprocal allogrooming pattern, especially at such an early age, appears to be unique among ungulates, and possibly mammals in general. Three hand-reared impala fawns, deprived of the opportunity to interact with older herdmates, but having access to impala fawns and heterospecific fawns, were observed from 1–3 mo of age. The hand-reared impala showed no alteration in the occurrence of reciprocal allogrooming behaviour compared with the dam-reared control fawns, indicating that allogrooming experience with older animals was not required for the appearance of reciprocal allogrooming at an early age. Interestingly, hand-reared fawns persisted in grooming heterospecific fawns despite the fact that heterospecifics rarely reciprocated grooming. We postulate that the strong predisposition for impala young to groom others may be related to the threat of tick infestation in the impala's ecotone habitat.     

Wrestling in the red-spotted newt (Notophthalmus viridescens): resource value and contestant asymmetry determine contest duration and outcome

When an unpaired male red-spotted newt encounters a male and female engaged in amplexus courtship, he attempts to displace the amplectant male, or ‘owner’, by wrestling with him. In this study, it was found that the amount of time an intruding male invests in wrestling is determined by both the size of the female, which seems to be a reliable indicator of her fecundity, and the length of the intruder relative to the owner. In 90% of all wrestling encounters observed, the owner retained the female. Successful displacement was rare and occurred only in encounters in which the intruder was at least as long as the owner. Similar contest behaviour in other amphibians is briefly discussed.     

Influence of gonadal hormones and social housing conditions on agonistic,copulatory, and related socio-sexual behaviour in the Mongolian gerbil (Meriones unguiculatus)

The Mongolian gerbil is colonial, but its social organization within its colonies is not known. Pair formation has been suggested by breeders and has been observed under semi-natural conditions. Family units remain stable under such conditions. In the solitary-living hamster (Cricetus auratus), agonistic behaviour decreases in male-female encounters during female heat. This report concerns the influence of female and male gonadal hormones on agonistic behaviour in gerbils studied in cagemate and non-cagemate encounters. Further, the occurrence of mating, marking, and drumming was studied in the same situations. No effects were found of either the female or the male endocrine condition on the low amount of agonistic behaviour between cagemates. A reduction in the high amount of agonistic responses in encounters between non-cagemates was indicated. It was related to the castrated-male and induced-oestrous female conditions. Copulatory behaviour was not influenced by social maintenance conditions. Marking and drumming were most frequent in females in oestrous and intact males. The occurences in the females were influenced by the individual's endocrine condition, but not by the opponent's. Intact males in encounters with non-cagemates marked more frequently when the females were in anoestrous. Drumming in the intact males increased in tests with oestrous versus anoestrous females but occurred more frequently in encounters with non-cagemates than in those with cagemates. The behaviour of castrated males was not influenced by the endocrine or by social housing conditions. The limited variation in agonistic behaviour relative to the endocrine conditions may reflect a non-solitary social organization.     

The effects of luteinizing hormone, oestrogen and ovariectomy on the agonistic behaviour of female Quelea quelea

Previous work with male Quelea showed that agonistic behaviour in relation to individual distance is controlled by luteinizing hormone (LH), rather than by testosterone, and that males are more aggressive than females. Experiments with female groups are reported which show that: (a) LH injections increase aggressive encounter frequency; (b) ovariectomy in the breeding season (but not outside it) also increases encounter frequency; and (c) oestrogen injections decrease encounter frequency. The effects of LH were shown to be specific to agonistic responses rather than mediated through changes in activity. Correlations between changes in natural hormone levels and encounter frequency support the injection findings. These results are consistent with the hypothesis that LH controls aggressive encounters over individual distance in the female as in the male and that oestrogenic inhibition of this LH-mediated aggressiveness is a cause of female subordination and the lower encounter frequency found in female groups. The annual cycle of encounter frequency is described and the significance of different systems of hormonal control is discussed.     

The evolution of sociality in the spitting spider, Scytodes fusca (Araneae: Scytodidae)—evidence from observations of intraspecific interactions

The intraspecific behaviour of the communal-territorial spitting spider, Scytodes fusca , from North Queensland, Australia, is described from laboratory observations. This spider is tolerant of most conspecific intruders on the web but adults, particularly females, do display aggression to other adults by charging, chasing and spitting. During courtship, both the male and the female tap on the web. Courtship in S. fusca is similar to that of a non-social scytodid Scytodes thoracica. Males often stay with the female for a time after mating. Adults appear to be more cannibalistic toward third than toward first and second instar juveniles, This finding is discussed in relation to the mechanisms of intraspecific recognition. The social behaviour of S. fusca is compared with that of other scytodids and the possible evolution of sociality in S. fusca from the parent offspring association is discussed.     

Social Behaviour and Relationships in a Herd of Camargue Horses

Social relationships were studied in a herd of Camargue horses and are described in terms of certain aggressive and non-aggressive interactions and in terms of spatial patterning. Relationships between mares and their foal and yearling offspring were studied in detail. The effect of the activity of the herd on spatial patterning is described and changes in social behaviour due to the horsefly season are discussed.     

Birth and perinatal behaviour in family groups of the common marmoset (Callithrix jacchus jacchus), compared to other primates

Birth and perinatal behaviour in five family groups of the common marmoset are described. Information is given on 16 births, seven of which are described in detail.The birth process was divided into five stages; data are presented on the durations of the stages and the behaviours exhibited by the female in each stage.Infant marmosets cling unaided to the pelage of the female from birth, and the behaviour of neonates and the reactions of the female to infants are discussed.The interactions of other group members to the mother and neonates were analysed. The adult male's first interactions with the neonates appeared to occur much earlier (less than one hour after birth) than in other non-human primate species.Perinatal behaviour in marmosets is compared to that recorded in other non-human primate species and discussed in relation to the social organisation of C. j. jacchus.     

Agonistic and courtship behaviour patterns in the skink Chalcides viridanus (Fam. Scincidae) from Tenerife

There have been relatively few attempts to quantitatively describe behaviours in scincid lizards. Chalcides viridanus is a small body-sized skink endemic of Tenerife (Canary Islands). We describe and quantify 18 behaviour patterns (both social and agonistic) of this species, some of which have not been described before for other scincids. Video recordings of male–male, female–female, and male–female interactions were made under laboratory conditions, with controlled light–dark cycle and temperature. We describe several agonistic and courtship behaviour patterns. Within the first context, we detected a new agonistic behaviour for a scincid, “Snout to body”, that appeared at the beginning of agonistic sequences; it consisted of each animal placing its snout in contact with the other individual’s lateral side of the body. The amplitude of head movement during “Head bobbing” was lower than that described for many other lizard species. Agonistic behaviours were shown in intrasexual staged encounters both within males and females. The comparison of behaviour patterns of both types of intrasexual encounters showed that females were more active, exhibiting significantly higher frequencies of behaviour than males. Specifically, females showed the “Snout to body” pattern more frequently than males. In male–female encounters we detected courtship and copulation patterns only in April, when males performed “Bites” and “Snout to body” directed at females.     

Socially induced and rapid increases in aggression are inversely related to brain aromatase activity in a sex-changing fish, Lythrypnus dalli

Social interactions can generate rapid and dramatic changes in behaviour and neuroendocrine activity. We investigated the effects of a changing social environment on aggressive behaviour and brain aromatase activity (bAA) in a sex-changing fish, Lythrypnus dalli. Aromatase is responsible for the conversion of androgen into oestradiol. Male removal from a socially stable group resulted in rapid and dramatic (> or =200%) increases in aggression in the dominant female, which will become male usually 7-10 days later. These dominant females and recently sex-changed individuals had lower bAA but similar gonadal aromatase activity (gAA) compared to control females, while established males had lower bAA than all groups and lower gAA than all groups except dominant females. Within hours of male removal, dominant females' aggressive behaviour was inversely related to bAA but not gAA. These results are novel because they are the first to: (i) demonstrate socially induced decreases in bAA levels corresponding with increased aggression, (ii) identify this process as a possible neurochemical mechanism regulating the induction of behavioural, and subsequently gonadal, sex change and (iii) show differential regulation of bAA versus gAA resulting from social manipulations. Combined with other studies, this suggests that aromatase activity may modulate fast changes in vertebrate social behaviour.     

Daily activity of impala

The basic activity pattern of impala in the Serengeti is described quantitatively; seasonal variations and those related to social status are discussed. The adaptiveness of the basic pattern, especially in the preponderance of vulnerable and mobile activities in the daytime and of static activities at night, is discussed. The cost of various territorial activities is stressed; the presence of females in his territory disturbs the basic activity pattern of the territorial male, reducing his time spent on feeding and ruminating. The relationship between the occurrence of males in shade and some environmental factors is analysed.     

Etude des interactions sociales entre male et femelle chez la Genette (Genetta genetta L.): Relations entre marquage olfactif et agression

Study on Social Interactions between Male and Female Genets (Genetta genetta L.): Relations between Scent Marking and Aggression The aim of this work was to study the relations between marking behaviour and aggression in genets (Genetta genetta L.). Observations on social interactions were made on a socially stable pair and during encounters between unfamiliar pairs. In the first case, there appeared to be a relationship between marking activity in the male and aggression. During aggression periods, marking with the ano-urogenital region increased in the male and decreased in the female. Another striking variation concerned flank rubbing and hindleg rubbing. In both male and female these marking frequencies increased significantly during aggression periods and flank rubbing was modified by visual cues. Data recorded during encounters between unfamiliar pairs (one male being introduced in a female's cage) showed that agonistic behaviours were released through visual cues in females and olfactory cues in males. The same changes in marking frequencies were observed. These results also showed that ano-urogenital marking was inhibited in females during aggression periods. The adaptative significance of these phenomena is discussed.     

Social dominance in dairy cattle

The meaning of “social dominance” and the problems associated with its use are discussed. Most problems disappear if dominance is restricted to the phenomenon that in every pair of animals one member can inhibit the behaviour of the other. The dominance order of the group is the sum of all such inhibitory relationships. Dominant animals probably have been aggressive in the past to obtain their dominant positions, but a dominant animal need not be aggressive now. Measures of the dominance position of animals in a herd should be based on observations in the particular herd, contain sufficient observations to be reliable, reflect the actual magnitude of differences between animals, and be normally distributed. The method of Beilharz and Mylrea (1963) meets these requirements. The application of this method is discussed.The dominance structure of three dairy herds near Freiburg in South West Germany were studied. None of the herds had an individual cow who was dominant over all others in the herd. Parameters of social structure varied from herd to herd. The findings are discussed in relation to differences in space per animal, manner of recruitment of young stock to the herds, presence of horns and other features. It is concluded that the dominance relationship of any pair of animals is a result of learning, with many different factors, including trivial ones, being involved in the initial serious formation of the relationship. Once learnt, dominance relationships for each pair of animals persist for a long time. While bidirectional dominance relationships are found in young animals, where none has yet accepted dominance of others, mature animals of all species of domestic livestock generally have clear unidirectional dominance relationships. The dominance order of the group may be no more than the sum of the individual relationships. Dominant animals are freer than others in attending to stimuli from outside the herd.     

Sound production and reproductive behaviour of the armoured catfish Corydoras paleatus (Callichthyidae)

Sound production during reproductive behaviour, dyadic encounters and distress situations was investigated in the callichthyid catfish Corydoras paleatus. Sounds were broad-band, pulsed, acoustic signals produced during abduction of the pectoral spines. Only males emitted trains of sounds during courting and trains of sounds of shorter duration during dyadic encounters. Several males, which are usually smaller than females, courted one gravid female without obvious cooperation or competition between them. During mating, one previously vocalizing male clasped the female's barbels with one pectoral spine and inseminated the eggs. The number of successful spawnings, days until spawning, and number of eggs laid was not related to the number of males (one, two or three) combined with one female. Males did not behave aggressively towards each other during courting or in dyadic encounters. In distress situations, when fish were hand held, both sexes and juveniles produced single sounds. The dominant frequency was negatively correlated with body size and the sound duration was positively correlated with relative length of pectoral spines (standardized to body length). This acoustical behaviour in C. paleatus differs considerably from Hoplosternum thoracatum, a representative of the callichthyine subfamily, in which vocalization was observed during territorial behaviour in males and aggressive behaviour in both sexes. This is the first report of a major difference in vocalizing behaviour within one teleost family.