An analysis of inter‐ and intraspecific aggregations in two sympatric species of hermit crab (Decapoda,Anomura, Paguridae)

A series of experiments was conducted on the relationship between population density and spacing out in two sympatric species of crab, Pagurus bernhardus (L.) and Pagurus prideauxi Leach. A second series of experiments investigated the two species’ aggressiveness and activity by observing the behaviour of pairs of animals.

In the first series of experiments, animals became more aggregated (Clarke and Evans’ R) as the population density (p) was increased. The rate of aggregation (dR/dp) was greater when the species were mixed than when they were separate.

In the second series of experiments, activity, total encounters, and resolved encounters, were highest in paired P. bernhardus, lower in mixed pairs of the two species, and lowest in pairs of P. prideauxi. Larger individuals were more active than smaller ones in P. bernhardus and P. prideauxi pairs. In the mixed pairs, P. bernhardus were more active. Paired P. bernhardus were slightly more active than isolated P. bernhardus but this difference was not statistically significant. The activities of paired and isolated P. prideauxi were not significantly different. The distances between individuals in pairs were similar for all combinations.

The results are discussed in relation to the distribution of sympatric species.     

Temporal changes in food resources,parental feeding and breeding success of Heron Island silvereyes,Zosterops lateralis chlorocephala

The influences of the temporal change in food supply on the parental feeding effort and breeding success of silvereyes,Zosterops lateralis chlorocephala, was investigated on Heron Island, Australia. Food supply (arthropods and figs) declined as the breeding season progressed. The parental feeding rate and growth of nestlings were lower when food supply was poor. When available, dominant pairs fed their young more figs and fewer arthropods than lower ranking pairs. Dominant pairs raised heavier young than lower ranking pairs when food supply was poor, while there were no significant differences between them when food supply was rich. When food supply was rich, pairs delivering greater amounts of arthropods reared nestlings better, whereas feeding more figs did not improve growth of nestlings. When food supply was poor, pairs spending a longer time at the nest reared nestlings better.     

Female response to predation risk alters conspecific male behaviour during pre‐copulatory mate guarding

Mating behaviour often increases predation risk, but the vulnerability within mating pairs differs between the sexes. Such a sex difference is expected to lead to differences in responses to predation risk between the sexes. In the two‐spotted spider mite Tetranychus urticae, males engage in pre‐copulatory mate guarding because only the first mating results in fertilisation. We investigated (i) whether pre‐copulatory pairs are more conspicuous to the predatory mite Phytoseiulus persimilis than solitary females, (ii) whether the vulnerability to the predator differs between sexes within the pre‐copulatory pair, (iii) whether each sex of T. urticae responds to predation risk during pre‐copulatory mate guarding and (iv) whether T. urticae's response to predation risk affects predator behaviour. Because T. urticae females are immobile during pre‐copulatory mate guarding, we observed male behaviour to evaluate effects of predation risk. We found that the predators detect more pre‐copulatory pairs than solitary females and that more females than males of the pre‐copulatory pairs are preyed upon by the predators. The preference of spider mite males for pre‐copulatory pairs versus solitary females was affected by whether or not the female had been exposed to predators during development. Male T. urticae exposed to predation risk did not alter their behaviour. These results suggest that only the most vulnerable sex, that is the female, responds to predation risk, which modifies male behaviour. Regardless of T. urticae females’ experience, however, P. persimilis detected more T. urticae pre‐copulatory pairs than solitary females, suggesting that pre‐copulatory mate guarding itself is dangerous for T. urticae females when these predators are present. We discuss our results in the context of sex‐dependent differences in predation risk.     

The Effect of Social Environment on Male–Male Competition in Guppies (Poecilia reticulata)

We examined male–male competition in guppies (Poecilia reticulata) to test for evidence of hierarchy formation and any subsequent effects on male mating success by comparing the interactions of pairs of males that were siblings and life‐long tank mates with those of unrelated pairs that had never met. These pairs of males were first observed in the absence of a female; then a female was added to gauge the effects of the initial male–male interactions on male sexual behaviour. The unfamiliar/unrelated pairs engaged in significantly more aggressive interactions such as physical contacts, nipping and chasing than the familiar/related pairs. Based on several previous studies, we suggest that familiarity played a greater role than relatedness in the differences in behaviour that we observed. Our results suggest that, in some circumstances, more aggressive males may have more mating opportunities than less aggressive males. Our results also indicate that males adjust their aggressive and courtship behaviours to the perceived intensity of competition for mates, based on the number of mature males in their rearing tanks. We suggest that male–male competition for mating opportunities may play a more important role in the guppy mating system than previously thought.     

Body Size and the Maintenance of Reproductive Isolation in Stickleback,Genus Gasterosteus

The blackspotted stickleback Gasterosteus wheatlandi and the widely studied threespine stickleback G. aculeatus are sympatric throughout the former’s range and share many aspects of life history and reproductive behaviour. These two species differ significantly in size, with G. wheatlandi of both sexes measured at approximately 60% of the standard length of their G. aculeatus counterparts. This study concentrated on G. wheatlandi courtship behaviour and investigated its role in the maintenance of reproductive isolation with G. aculeatus. Specifically, the roles that (1) female body size plays in influencing male courtship preferences and (2) male body size and behaviour play in female courtship preferences were investigated through dummy and live conspecific and heterospecific stimulus presentations. Male G. wheatlandi courtship preferences are consistent with previously described patterns for G. aculeatus. Males of both species preferentially approach and court the larger of two simultaneously presented live or dummy females. Thus, the smaller G. wheatlandi males are indiscriminate with respect to assortative mate choice; not only preferring to approach and court more fecund conspecific females but, more significantly, G. aculeatus‐sized females. In contrast, females of both species demonstrate strong assortative courtship preferences. When presented with pairs of flask‐enclosed males, females of both species preferentially orient and court the conspecific male over the heterospecific. Similarly, when presented with a conspecific male and a heterospecific male presented singly, females prefer to enter the nest of the conspecific. Systematic analysis of the interactions between these pairs of fish (one male, one female) demonstrates that the breakdown of courtship in heterospecific courtship occurs late in the courtship sequence when the widely differing forms of male leading behaviour results in drastically differing female responses. I suggest that, as previously described in G. aculeatus, the supernormality effect plays a significant role in mediating adaptive mate choice behaviour in G. wheatlandi. However, the added element of a larger sympatric species introduces a possible cost in time and energy devoted to courting heterospecific, and sympatric, females that the larger G. aculeatus do not likely incur. There is substantial evidence from many sympatric G. aculeatus species pairs that there is assortative mate choice based on size and/or courtship behaviour. Courtship trials suggest a more pervasive role for females in assortative mate choice. Whether it is male body size per se, or in combination with behaviour, morphology or other cues, is unresolved in the present study.     

The genetic and environmental basis of adaptive differences in shoaling behaviour among populations of Trinidadian guppies,Poecilia reticulata

The degree of plasticity an individual expresses when moving into a new environment is likely to influence the probability of colonization and potential for subsequent evolution. Yet few empirical examples exist where the ancestral and derived conditions suggest a role for plasticity in adaptive genetic divergence of populations. Here we explore the genetic and plastic components of shoaling behaviour in two pairs of populations of Poecilia reticulata (Trinidadian guppies). We contrast shoaling behaviour of guppies derived from high‐ and low‐predation populations from two separate drainages by measuring the shoaling response of second generation laboratory‐reared individuals in the presence and absence of predator induced alarm pheromones. We find persistent differences in mean shoaling cohesion that suggest a genetic basis; when measured under the same conditions high‐predation guppies form more cohesive shoals than low‐predation guppies. Both high and low‐predation guppies also exhibit plasticity in the response to alarm pheromones, by forming tighter, more cohesive shoals. These patterns suggest a conserved capacity for adaptive behavioural plasticity when moving between variable predation communities that are consistent with models of genetic accommodation.     

Behaviour of cyclic bank voles under risk of mustelid predation: do females avoid copulations?

Mustelid odours have been shown to suppress breeding in captive bank voles (Clethrionomys glareolus) from cyclic populations (Ylönen 1989; Ylönen and Ronkainen 1994). The mechanism behind the suppression is unknown. Based on a series of behavioural trials and breeding experiments with pairs of bank voles in breeding condition, we suggest that the primary cause for breeding suppression is a change in female mating behaviour. Experimental female-male pairs (n=34) exposed to mustelid odour decreased their general activity compared to control pairs (n=34). When encountering males in behavioural trials, females exposed to stoat odour were more aggressive and actively avoided precopulatory behaviours of males. No copulations were observed in experimental pairs compared to five in control pairs during the behavioural trials. Males actively approached females in general but male behaviour did not change under exposure to mustelid odours. We suggest that females are more vulnerable to mustelid predators than males and therefore actively avoid copulations in the (indirect) presence of mustelids. As well as this behavioural response, internal abortive mechanisms (cf. Bruce 1959) could play a role in the observed breeding suppression.     

Cobreeding in the Burying Beetle,Nicrophorus vespilloides: Tolerance Rather Than Cooperation

Under intra‐ and interspecific competition, cooperative behaviour can provide direct fitness benefits if individuals work together to expel intruders. In the burying beetle Nicrophorus vespilloides, a relatively small species, multiple unrelated pairs can breed together, and individuals are weak competitors in interactions when competing with larger individuals of the same species and with larger species of the same genus. In field and laboratory studies, we found that two pairs together attracted significantly fewer congeneric and non‐congeneric competitors compared with single pairs. No other benefits were found. Communal breeding had large negative effects on fitness, as there were fewer offspring per pair and a higher chance of injuries. The higher chance of injuries reflected pairs fighting among themselves and not against competitors. These costs are much greater than the small benefit of fewer intruders. Why should a N. vespilloides breeding pair eventually allow another pair to join? A potential partial explanation is that these are not cooperative pairs in the traditional sense, but rather pairs have a higher tolerance for each other. From the resident's perspective, joining pairs are not expelled because the chance of injury makes the cost of fighting high. Joining others may have unusually low costs in this species because reproductive opportunities are rare, dependent on a resource that is unpredictable in time and space, and residents should be inclined to tolerating new pairs because of the cost of fighting. Thus, for N. vespilloides, communal breeding appears to be ‘making the best of a bad job’, providing some reproduction rather than none.     

On the rutting behaviour of the Himalayan goralNemorhaedus goral (Hardwicke, 1825)

The rutting behaviour of the Himalayan goralNemorhaedus goral was studied in an area of the Himalayan foothills, Himachal Pradesh, Northern India, from October 13 to 25, 1989. Twenty seven qualitatively different behaviour patterns were observed. Males showed a more extensive repertoire (25 patterns, 14 performed only by this sex) than did females (12 patterns, 1 of them-Reactive urination-unique). The Head butt and the Intentional head butt were the most frequently shown behaviour patterns of both sexes, and Naso-nasal contact followed. Males interacted mainly with females and usually seemed to ignore each other. Courtship activities peaked on October 22–25, when mounts were also recorded. “Following” was the only mating system observed. Pairs sometimes courted at short distances from each other, without eliciting any apparent aggressive reaction. No evidence of territoriality was found throughout our study period. The goral behaviour appears to be more closely related to that ofOreamnos (in quality and quantity of behaviour patterns, as well as the mating system) than to that ofCapricornis.     

The Effect of Inbreeding on Mating Behaviour of West Indian Sweet Potato Weevil Euscepes postfasciatus

When there is an inbreeding depression, mating with a kin individual is generally considered maladaptive behaviour. However, in some conditions, the inclusive fitness benefits from inbreeding may outweigh the costs of inbreeding depression, and thus, inbreeding tolerance is often adaptive. Inbreeding depression and the effect of relatedness on mating behaviour in the West Indian sweet potato weevil Euscepes postfasciatus were examined. No significant inbreeding depression was detected as indicated by body weight and number of progeny emerging from sweet potato roots. Male mating performance (i.e. number of mating occurrences per night) was adversely affected by inbreeding depression, but the effect was low (fitness loss was 6.3%). Although there were no significant differences in latency to mounting, pre‐copulatory guarding, copulation and post‐copulatory guarding duration between full‐sib and non‐kin pairs, the copulation rate of full‐sib pairs was significantly higher than that of non‐kin pairs. These results support the theoretical prediction that when inbreeding depression is weak, copulation with close relative individuals is favoured.     

Chromosomal interchange inEleutherine plicata Herb

Chromosome behaviour at metaphase I and anaphase I of meiosis inEleutherine plicata Herb. (2n=14) is studied. Cells with chromosome associations comprising an association of four long chromosomes, in addition to five bivalents were observed more frequently than those with seven bivalents. it is concluded that the ring of four is due to a segmental interchange between the two long non-homologous chromosome pairs. The ring of four at anaphase I showed delayed disjunction, bridge formation and irregular separation of chromosomes in a number of cells while the behaviour of the other bivalents was normal.     

Significance of Agonistic Vocalization in the Croaking Gourami (Trichopsis vittatus,Teleostei)

Trichopsis vittatus emits high amplitude sounds during agonistic encounters with conspecifics. The sound producing organ is derived from the structural components of the pectoral fins. The study involved muting a sample of subjects by removing two pectoral fin tendons without any further restriction in movements and behaviour. Mute and unaltered males were then placed together in pairs and the following agonistic behavioural elements were determined: attacks, lateral displays, sound production and frontal displays. Soniferous males had a higher probability of winning contests when size differences were small. In pairs with big size ratios, the larger males were more often successful. Besides visual and acoustical signals, lateral line stimuli seem to play no role in threatening displays. These results demonstrate that vocalization during agonistic encounters is important for becoming dominant in specific circumstances. Because of correlation between main frequencies and size, sound emission might be a very effective method of assessing the physical strength of an opponent.     

Diet influences male-female interactions in the bushcricketRequena verticalis (Orthoptera: Tettigoniidae)

The influence of diet on the courtship roles of male and female Requena verticaliswas investigated in the laboratory. The protein content of available food was found to affect the frequency of mating attempts. Pairs which were fed on a low-protein diet were involved in fewer mating attempts than pairs which were fed on a high-protein diet. Diet also influenced the relative frequencies of male and female rejections. Males rejected their virgin female partners more often than females rejected their male partners when the pairs were kept on a low-protein diet. The opposite was found when the female had mated once before. No difference in the frequency of male and female rejections was found when the pair was kept on a high-protein diet irrespective of the mating status of the female.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.     

A note on the effect of isolation during testing and length of previous confinement on locomotor behaviour during open-field test in dairy calves

This study investigated firstly if confined calves perform more locomotor behaviour when open-field tested in pairs than when tested individually, and secondly if length of confinement affects the build-up of motivation to perform locomotor behaviour. In the first experiment,14 calves were open-field tested on two successive days either individually or as a pair. Calves walked more and performed more locomotor play when tested in pairs, suggesting that it may be appropriate to avoid isolation when aiming to measure the effects of confinement on locomotor behaviour. In the second experiment, in each of three successive weeks, 24 calves had access to an exercise arena for 45 min daily on three successive days either: (i) the first 3 days, (ii) the third, fourth and fifth day, or (iii) the fourth, fifth and sixth day. On the seventh day the calves were released into the arena for 10 min (open-field test). All calves received all three treatments in a Latin square design. Calves performed more locomotor play, and they trotted more after 3 days without access to the arena than after 1 or 0 days, suggesting that in calves the motivation to perform locomotor play and trotting increases with length of confinement.     

Dominance hierarchy among workers changes with colony development in Polistes japonicus (Hymenoptera, Vespidae) paper wasp colonies with a small number of workers

Dominance hierarchy in the primitively eusocial wasp Polistes japonicus was analysed in four colonies for two periods: (1) the first-brood period, when only early emerging workers are present on the nest, and (2) the mixed-brood period, when the first and second (last) broods are present on the nest. The rank in the dominance hierarchy was determined based on a sociogram showing a dominance–subordinance relationship for all pairs of workers. During the first-brood period, older workers were likely to be more dominant (older dominance hierarchy), while the rank of workers was reversed during the mixed-brood period, with younger workers being likely to be more dominant (younger dominance hierarchy). However, the oldest and youngest workers were not always the top-ranked workers in the dominance hierarchy during the first- and mixed-brood periods, respectively, and during the mixed-brood period no younger dominance hierarchy was evident when the first or second brood was analysed separately. Higher ranked workers displayed dominance behaviour more frequently, and the lowest ranked worker hardly displayed dominance behaviour. Most workers displayed dominance behaviours primarily towards the worker ranked immediately below in the dominance hierarchy during the mixed-brood period but not during the first-brood period. The bodies of younger workers were larger for the mixed brood, but not for the first brood in some colonies or the second brood in all colonies. The association between body size and rank in the dominance hierarchy was negative during the first-brood period and positive during the mixed-brood period, with a nearly significant trend also seen even when the analysis was limited to the second brood. To explain the above temporal change from an older dominance hierarchy to a younger dominance hierarchy, we propose the hypothesis that the probability of a worker inheriting the colony increases rapidly with colony development, and consequently younger larger workers attempt to move up the dominance hierarchy in order to produce their own offspring by becoming the superseder late in colony development, rather than working harmoniously so as to boost the overall production of reproductive progeny for a colony, which is the strategy adopted early in colony development.     

Pair formation in the herbivorous rabbitfish Siganus doliatus

This study investigated the basis of pair formation in the abundant herbivorous rabbitfish Siganus doliatus on Orpheus Island, Great Barrier Reef. Pair formation was the most common social system in S. doliatus, with 67·4% of all individuals occurring in pairs. Pairs were stable (i.e. individuals remained with the same partner throughout the study) and pair members were found within 5 m of each other 82·9% of the time. Of the examined pairs, 25% were homosexual resulting in a proportion of heterosexual pairs (75%) that was significantly lower than expected if pairs were formed solely for reproductive reasons. Therefore, although reproduction appears to be the main driver of pair formation in S. doliatus, other factors are likely to influence this behaviour. The high density of individuals on the reef crest (5·7 ± 0·9 individuals 200 m^?2; mean ± s.e .) and extensively overlapping home ranges of pairs indicated that the defence of territories plays no role in pair formation. Instead, it appears that pair formation in S. doliatus is driven, in part, by other, non‐reproductive, ecological factors. It is suggested that pair formation allows for increased vigilance against predation and enables S. doliatus to execute a novel feeding behaviour.     

Variation in stimulus,seasonal context,and response to urine marks by captive Iberian wolves (<Emphasis Type="Italic">Canis lupus signatus</Emphasis>)

Urine marking behaviour was observed over 14 months in two captive pairs of Iberian wolves (Canis lupus signatus) by means of behaviour sampling. The study focused on the relative attractiveness of the different stimuli toward which urinations are directed, the seasonal context of urine marking, and the response by mates to sexually dimorphic action patterns. Kinds of marked substrates varied according to the posture used by wolves to urinate [raised-leg urination (RLU), flexed-leg urination (FLU), standing urination (STU), or squat urination (SQU)]. In RLUs and FLUs the wolves mainly used conspicuous substrates, whereas STUs and SQUs were carried out directly onto inconspicuous substrates. Wolves urinated on plants more than on other substrates, mostly selecting trees. A selection of trees according to their trunk diameter was also observed. The posts were chosen or avoided seemingly because of their specific characteristics such as size. In short, wolves marked with urine (RLUs and FLUs) those substrates that secured a greater effectiveness of marking. The mean duration of RLUs was lower than that of FLUs and SQUs. The rate of RLUs in males increased in winter (November–December) and in summer (July–August), while the rate of FLUs and SQUs in females and STUs in males did not increase during these periods. The wolves investigated RLUs more frequently than SQUs. The RLU display appears to be more ancestral than derived, similar in all species of canids and even in the Iberian wolf.     

Effects of Pair-Bond and Social Context on Male–Female Interactions in Captive Titi Monkeys (Callicebus moloch, Primates: Cebidae)

In monogamous species, an abiding relationship between a specific adult male and a specific adult female is a defining feature of the social system. The interactions between these individuals are influenced by many factors, including not only the history of their relationship (for example, development of a mutual bond), but also the immediate effects of the prevailing social context (for example, presence and sex of extra‐pair conspecifics). In this study we examined the effects of an existing bond and of social context on interactions between adult heterosexual pairs of the monogamous titi monkeys (Callicebus moloch). Twelve adult males and 12 adult females were tested with their cagemates and with an unfamiliar partner of the opposite sex in five social contexts: (1) mated male–female pair; (2) unfamiliar pair; (3) single female; (4) single male; and (5) empty stimulus cage. Results show that mated pairs were more affiliative than unfamiliar pairs and differentiated social contexts more sharply. Males were more responsive to context than females. Distance between mates was less and physical contact was more frequent in the presence of male–female pairs or a single male, than in the presence of a single female or an empty cage. These findings suggest that the presence and sex of strangers have a stronger influence on male–female interactions when the pair has an existing relationship.     

Comparison of affiliative behaviors between old and recently established pairs of golden lion tamarin,Leontopithecus rosalia

To evaluate the changes in the relationship between male and femaleLeontopithecus rosalia over time, I compared the interactions in four well established pairs at the National Zoological Park, Washington, D.C., to those in four newly established pairs at the Conservation and Research Center of the National Zoological Park, Front Royal, Virginia, U.S.A. I recorded the frequency of approaches, withdrawals, food offerings, food takings, and resisting food transfers. Male and female adults were the focal animals and the observations totaled 206.5 hr. Males of newly established pairs interacted more with females than males of old established pairs. While males generally began and maintained the interactions significantly more than females, females of recently formed pairs were initiators of affiliative behaviors significantly more than females of old pairs. These results demonstrate a qualitative difference in the relationship between males and females when old and new pairs are compared.