Ontogeny of Aggressive Behaviour in Schools of Yellowtail, Seriola Quinqueradiata

We studied ontogenetic changes in social interactions, especially in aggressive behaviour of the migratory marine yellowtail, Seriola quinqueradiata (Carangidae), and compared these to morphological and physiological changes. No agonistic interactions were observed during the larva period until 10 mm in total length (TL), at approximately 20 days after hatching. Typical shivering behaviour with J-posture was observed during metamorphosis, when fin rays and calcification of vertebra were completed and there was an increase of tissue thyroid hormone. The onset of aggressive behaviour was just after metamorphosis to the juvenile period, and coincided with a significant increase in tissue cortisol levels. The onset of schooling behaviour was at 12 mm TL, slightly after the onset of aggressive behaviour. From observations of individual aggressive behaviour within juvenile schools, we found three categories of social rank: dominants (10-20%), intermediates (10-20%), and subordinates (60-80%). There was an inverse relationship between social rank and cortisol concentration. Otoliths of dominant fish in 8 experimental groups were labeled and the fish were returned to their groups. Six labeled dominants appeared after 1 day and three after 1 week rearing, respectively, indicating that social rank was maintained for at least 1 week (binomial distribution, p < 0.05). Dominants were larger than subordinates after 1 day rearing, whereas dominants were smaller after 1 week rearing. From long-term rearing experiments using individual otolith marking, larvae that showed the J-posture more frequently tended to become dominants after metamorphosis, indicating a positive correlation between the J-posture and aggressive behaviour. Synthesizing all results from behavioural experiments, we generated a behavioural model for the triggering mechanism of aggressive behaviour and size selection of school members.     

Fighting Tactics of Fallow Bucks (Dama dama,Cervidae): Reducing the Risks of Serious Conflict

Competition between male fallow deer (Dama dama), during the breeding season was studied to determine if conflict strategies were consistent with the reduction of risk. Agonistic interactions between males were analysed in relation to age, dominance rank and availability of mating opportunities. The breeding season was divided into two main periods: the pre-rut began when all males had cleaned the velvet from their antlers and ended on the last day before matings were first observed, while the rut refers to the period between the first and last matings. Overall, socially mature males (≥4yr old) were involved in more interactions than immature (≤3yr old) males. Males established dominance rank largely by non-contact agonistic interactions during the pre-rut and there was substantial carry-over of rank to the rut, when it was correlated with mating success. The mating success of males was skewed; mature males achieved 99.4% of the matings and immature males accounted for 0.6%. A mature male was 13 times more likely to fight than an immature male; the mature males that fought most often did so between 0.4 and 0.5 times per hour. During the rut, the number of fights was positively correlated across days with the number of matings. The majority of agonistic interactions (79%) comprising dyads of immature males, involved antler contact. In contrast, mature males engaged antlers in only 42% of their interactions. Fights between mature males lasted more than twice as long as those between immature males and were more likely to occur between opponents with similar dominance ranks. However, towards the end of the rut formerly mismatched opponents were more likely to fight. Thus males operated conditional competitive strategies to decide when to interact and fight. The persistence of rank order from the pre-rut period to the rut and the tendency for mature males to resolve disputes without antler contact, served to reduce the frequency of fights and therefore the risk of serious injury.     

Deferred agonistic behavior in a long-lived scincid lizard Eumeces laticeps

Summary A laboratory experiment with the broad-headed skink (Eumeces laticeps) involving staged agonistic encounters demonstrates that larger males have an advantage over smaller ones in agonistic bouts. Field data on head wounds produced by intraspecific fighting during the breeding season show a much higher frequency of new wounds among males over 100 mm in snout-vent-length than in smaller males. The significant difference in new-wound frequency strongly suggests avoidance of fights by the small males, which is corroborated by laboratory and field observations. Access by males to reproductively active females depends on the ability to defeat other males in aggressive contests virtually always involving head biting if the males are of nearly equal size. Because the probability of winning agonistic encounters increases with size, young males avoid fights with older males. Aggressive contests with larger males and reproductive attempts other than courtship in the absence of larger males are deferred.Aggressive behavior in E. laticeps may be employed in direct defense of females, but might also be expressed in defense of specific sites and/or territories. In the laboratory, males in their home cages were significantly more likely to win encounters with males of similar size than were males fighting in the home cages of opponents. This suggests that encounter site could be important in determining encounter outcome and that field study of possible site defense or territoriality is needed.     

Effects of male call pitch on female behaviour and mate fidelity in little penguins

Playback experiments with two loudspeakers were conducted to examine how female little penguins (Eudyptula minor) respond to bray calls of males that varied in (1) dominant frequency (low vs high) and (2) familiarity (current partner vs stranger). In the first experiment, there was no statistically significant difference in the number of approaching females between low- and high-pitch calls. In the second experiment, using calls from the current partner and a stranger, females were more likely to approach closer to their own partners calls, regardless of the pitch of the strangers call. We conclude that female little penguins may recognize their partners calls (or at least familiar male calls) and appear to maintain mate fidelity even when larger, potentially more successful, strangers are on offer.     

Seasonal dynamics of agonistic displays in territorial and non-territorial males of goitered gazelle

Aggression serves a great variety of social functions, one of which is protection of individual territories from intruders. Territorial males of many antelope species show aggressive noncontact displays, and only rarely fight. It has been suggested that ungulate males tend to have more frequent and longer aggressive interactions with rivals of similar age or social status than with males of dissimilar status. In the present paper, we test whether territorial and non-territorial males behave in a similar manner and avoid fights, and whether or not they preferentially direct aggressive and longer agonistic interactions towards males of similar age or social status, rather than towards other classes of males. We found that territorial males usually avoided straight fights with peers, and instead mainly used noncontact displays in aggressive interactions. In contrast, non-territorial males used fights considerably more often, especially during the onset of territoriality in April to May, when these males had their most frequent aggressive interactions. Territorial bucks aggressively interacted most frequently with non-territorial males and significantly less often with other territorial males, but agonistic noncontact displays between territorial males lasted the longest. In contrast, non-territorial males addressed their aggressive noncontact displays and fights most often to peers and less frequently to sub-adults. Asymmetry in the social status of territorial vs. non-territorial males was likely responsible for the distinctively different agonistic behaviors shown by the two types of males, which in turn are likely due to the different costs and benefits each male can accrue from these aggressive interactions.     

Flexibility in the multi-modal courtship of a wolf spider, Schizocosa ocreata

Male Schizocosa ocreata wolf spiders court females with synchronous visual and seismic displays. We tested whether male S. ocreata modify their courtship in relation to light environment, and associated utility of the visual components. Males were generally more active and more likely to perform the major courtship element (jerky tapping) when in the light. One courtship element (arching) was only observed in the light while another (vertical leg-extend) was only observed in the dark. Courtship in the dark retained visual components, suggesting spiders cannot disengage these components of multi-modal display even when superfluous. Once initiated, there was no evidence that performance rate or time spent in each courtship element differed in the light and in the dark.     

Aggression among Lekking Male Fallow Deer (Dama dama): Territory Effects and Relationship with Copulatory Success

We studied the aggressive behavior of territorial male fallow deer (Dama dama) at two leks. Daily number of escalated fights was not correlated with number of matings. A dominance index including all agonistic interactions was weakly correlated with copulatory success; the correlation was stronger when each half of the rut was considered separately. Dominance likely changed over the rut due to fatigue. We ranked lek territories by the number of copulations seen in each. Males that won fights were likely to take over the loser's territory only if it ranked higher than the one they held. Winners were more likely to fight with a third male after defeating an opponent that held a higher-ranking territory than one from a lower-ranking territory. Exposure to attacks by third-party males may be a major cost of fighting. Males appear aware of the relative value of different lek territories, but the frequency of aggression was not scaled to potential fitness benefits, possibly because males seldom fought with opponents they were unlikely to beat. The outcome of interactions on the lek may not always reflect the relative dominance rank of the contestants. Females are unlikely to use the outcome of fights as a direct criterion for mate selection.     

On the distinction between niche and competitive ability: Implications for coexistence theory

The meaning of niche and competitive ability have long been surrounded by controversy. The reason for this stems from the obscure relationship that exists between these terms. This extends from the views of Darwin through Eltonian tradition to current views in which the meaning of competitive ability is implicitly infused into the paradigm of niche. Distinct operational definitions for niche and competitive ability are therefore established with special reference to plants. It is proposed that potential niche refer explicitly to a theoretical hyperspace of places where a species would leave descendents if all biotic interactions were precluded, and that competitive ability refer to the relative capacity to leave descendents in a particular place in the face of restrictions imposed by competitive interaction. This leads to a qualitative comprehensive theory for coexistence which may be extended to any type of biotic interaction. Niche and competitive ability are both determined by the biological attributes of a species and may be independently adjusted in a population by natural selection in contexts of competition. Species coexistence in nature may therefore be a consequence of alternative evolutionary mechanisms which may operate to various degrees in concert: (1) natural selection leading to niche differentiation; (2) an ongoing process of reciprocal selection (coevolution) which maintains an approximate balance in relative competitive abilities for contested resources.     

Evolution of delayed reproductive senescence in male fruit flies: Sperm competition

Populations ofDrosophila melanogaster that had been subjected to long-term selection favoring either delayed or rapid senescence were compared with respect to age-specific components of male reproductive success involving sperm competition. These components of reproductive success were divided into those related to sperm defense (protection of sperm from other males), and into those related to sperm offense (ability to mate with previously mated females and to displace the sperm of other males). Males were tested at four ages ranging from 1–2d to 5–6 wk after eclosion. Several aspects of sperm defense capability showed clear evidence of senescent decline. Furthermore, males from populations selected for delayed senescence were superior to males from control (rapid senescence) populations with regard to components of sperm defense. The superiority of males from populations with delayed senescence either increased as a function of male age, or was present at all ages tested. These results indicate that the rate of reproductive senescence in maleD. melanogaster can be altered in predictable directions by artificial selection. There were no differences between selection regimes with regard to sperm offense, and most components of sperm offense did not show clear evidence of senescence. The improved late-age reproductive success of males from populations selected for delayed senescence did not appear to entail any cost or trade-off at early ages with respect to the reproductive traits examined in these experiments.     

First copulation increases longevity and fecundity of <Emphasis Type="Italic">Histiostoma feroniarum</Emphasis> (Acari: Astigmata: Acaridida) females

I investigated the influence of insemination at different life stages on female fitness in the mite Histiostoma feroniarum. In this species, males guard immature females at the tritonymph stage to inseminate them immediately after the last moulting. Four groups of females were studied (1) females inseminated naturally, i.e. mating occurred immediately after guarding, and then the male was removed (IF/0M), (2) naturally inseminated females, where after insemination the male was replaced by two additional males (IF/2M), (3) virgin females reared without males (VF/0M) and (4) mature, virgin females to which two virgin males were added 3 days after last moulting (VF/2DM). The results show that females inseminated naturally (IF/0M) have higher longevity and fecundity than either virgin (VF/0M) or late-inseminated females (VF/2DM). Furthermore, longevity and fecundity of the former (IF/0M) was also greater than that of females naturally inseminated and subsequently exposed to males (IF/2M). One may suggest that seminal fluids have a positive effect on female fitness. When delayed insemination occurs, such positive effect may not be observed due to a change in features of the sperm access system. Harassment may explain decreased longevity and fecundity of females inseminated naturally compared to females that received additional males.This revised version was published online in May 2005 with a corrected cover date.     

Parallel swims as a means of intra- and interspecific assessment in stream fishes

Synopsis Several North American minnow species are known to engage in the action pattern parallel swimming, an activity that allows males competing for nests or nesting sites an opportunity to assess each other. These agonistic displays have been hypothesized to occur only between males of similar size. We provide evidence to support this hypothesis for two minnow species and a sucker species. Competition is strong both intra- and interspecifically for nest sites and we document the occurrence of interspecific parallel swims between three species.     

Aggressive habituation and rim circling in the social organization of bluegill sunfish (Lepomis macrochirus)

Synopsis The rim circling (RC), and aggressive habituation of nesting bluegills were studied in natural colonies during the six days of the reproductive period following spawning. To examine the stimulus specificity of aggressive habituation, selected test individuals and four neighbors were observed on four occasions during one day. After the first observation period established baseline data on aggressive interactions, the appearances of three of the neighbors were manipulated and further observations were made. Two more observation periods followed, separated by a sham manipulation to allow for possible disturbance effects. Fish lengths, inter-nest distances, and positions in the colony were also recorded.An analysis of the data demonstrated that aggressive interaction increased for the three manipulated neighbors. There were no significant differences among the data for the fourth (control) neighbors. The enhanced aggressive interactions between the test and manipulated males were initiated by the males at least partially in response to the changed appearance of the neighbor. Further, the level of aggressive interaction returned to normal after about 2 hours, indicating that the test male had habituated to the new appearances of his neighbors. Thus visual appearance plays an important role in the habituation of aggression between neighboring bluegills. It was not possible to determine if the behavior of the neighbor was also an important cue used in aggressive habituation, but this seems likely. Advantages of aggressive habituation are discussed. It is noteworthy that aggression decreased as internest distance increased.Levels of RC varied significantly over the six days, in a time course dissimilar from aggression. The RC rate is dependent on size of male, nest locality, and circling activity of neighbors. Observations on the synchrony of rim circling indicated social facilitation. It is argued that RC probably fulfills an advertisement function important in defending the brood. The motivation for RC is likely complex, and its performance reflects a mixed aggressive and reproductive strategy in the nesting population. Aggressive habituation functions in the social organization of bluegills to facilitate the occurrence of group displays such as RC.     

Female choice in Drosophila: evidence from Hawaii and implications for evolutionary biology

Details of female choice of mate in Drosophila silvestris of Hawaii strikingly parallels epigamic behavioral systems in many other animals and may be common in other species of Drosophilidae. Females respond selectively to male circling, wing displays, songs and tactile stimulation with foreleg cilia, a quantitative character that is highly variable in some populations. I hypothesize that the female can exert choice based on these cues from individual males that differ genetically by quantitative trait loci. Laboratory tests show that one third of courting males are repeatedly rejected in favor of a minority of alpha males. This result imposes non-random mating at the local population level. Past multiple-choice lab tests, widely used to measure isolation between pairs of populations or species of Drosophila may be flawed, since random mating has been assumed in the interpretation of results. Pre-mating sexual selection is clearly a powerful intrapopulation force in population biology. This view creates difficulties for discerning any proposed simultaneous interpopulation selective events in the presence of strong female choice. The long-held theory assuming that there is significant selection for pre-mating isolation between groups is questionable.     

Sound production through the substrate during reproduction in the mottled sculpin,Cottus bairdi (Cottidae)

Synopsis During reproduction maleCottus bairdi defend cavities beneath stones and perform defense and reproductive displays. Using a geophone to detect substrate vibrations under dark conditions (infrared viewing), we recorded three types of sounds. Knocks are produced during head nods and an acoustically similar sound is produced when the fish slaps the head to the substrate. A third sound, the drum roll appears to be a fast repetition of several knocks followed by a head slap. We argue that these signals traveling through the substrate are of greater importance than sounds traveling through the water because (1) the substrate vibration attenuates at a much lesser rate than the water vibration and, (2) even near riffles, which generate much water vibration, the background noise in the substrate is low enough for the fish to detect such sounds.     

No female mate choice in Mallee dragon lizards, Ctenophorus fordi

Aspects of sexual selection were studied in a sexually monomorphic Australian agamid lizard (Ctenophorus fordi), in particular with respect to the sensory exploitation hypothesis. In enclosure trials, females were offered the choice between large vs. small males and, in a different experiment, males with blue vs. normal head color. The rationale for these experiments was: firstly, to establish if females actively solicit copulations; secondly, if so, do females solicit copulations non-randomly with respect to male size (because large males may have access to food resources); thirdly, if male coloration is manipulated to match traits of congeneric, conspicuous and sexually dimorphic species, do females show preference for this novel trait (in accordance with the sensory exploitation hypothesis). The corresponding manipulations were also made in a free-living population where the distribution of females on the home ranges of color-manipulated males were monitored. Blue-headed males were accepted as mating partners both in the staged mating trials and in the natural population. Females appeared not to express any kind of active or passive mate choice (rejection); in only one out of 62 trials did a female approach a male herself rather than being approached by the male(s). There was no discrimination against any male category regardless of size or color within a female's receptive period and the manipulation of male head color in the natural population did not result in spatial re-distribution of females. Thus, a female appears to mate unselectively within her receptive period. Rejection behaviors were used only outside of the receptive period to communicate, to all males, that the female is not receptive.     

Tests to determine the responsiveness of free-ranging Japanese monkeys in food-getting situations

Experiments were performed on a free-ranging Japanese monkey troop (Arashiyama-B troop) in order to study the various levels of response caused by excitement and the characteristics of each monkey's action as observed under a series of stimulating situations carried out step by step through the giving of peanuts to the monkeys in various ways. The following results were obtained: (1) Three types of responses, that is, Threat response, Fear response, and Negative response were observed. (2) In response types, fairly evident individual differences exist along side sexual and age differences. (3) Similarity of response tendencies were not readily recognizable between a mother and her children. (4) Between two sisters with a slight disparity of age, the younger showed a stronger threat response. (5) Among females more than 4 years old, the lower the rank, the stronger the fear response.     

Maintenance of female mimicry as a reproductive strategy in bluegill sunfish (Lepomis macrochirus)

Synopsis In species where male reproductive success is dependent on male competition and aggression, alternative reproductive patterns, thought to represent a reduction in male reproductive effort, sometimes occur. Female mimicry in bluegill sunfish (Lepomis macrochirus) is an example of an obligate alternative male strategy. Female mimics are small, sexually mature males which mimic the details of female behavior, and gain access to functional females attracted to the nests of large, aggressive territorial males. The costs of female mimicry relative to nesting male behavior are discussed and two hypotheses, deception and mutual gain, are presented to explain the tolerance and courtship of female mimics by nesting males.     

The sexual function of wallowing in male wild boar (Sus scrofa)

The wild boar (Sus scrofa) is a medium-sized ungulate that produces a large variety of marks. Traditionally, wallowing is an activity that has fundamentally been related to several processes such as thermoregulation, reduction in the number of ectoparasites or disinfection of wounds in the skin of males resulting from fights to reach females. In this study, I analysed another possible function of wallowing in free-ranging wild boar populations when temperatures are low and parasitic numbers are not significant. The results indicate that during the sampling time, from October to February, the number of wild boars with a mud layer on their skin was 47, out of a total of 558 wild boar analysed. Of these 47 wild boars, 44 were males and within this group, 40 individuals were adult males. The wallowed males were bigger and older than non-wallowed males hunted at the same time. In addition, I found that males wallowed mainly during autumn (90.9% of all cases). In the total number of wild boars analysed, I did not notice individuals with wounds resulting from fights other than from the hunting day. On the other hand, the analysis of the females reproductive systems indicated that the rut period took place during the last days of October and the first days of November. From these results, the coincidence of the rut period with the maximum of wallowing in adult males could be indicative of a sexual function of this activity.     

Urinary testosterone-metabolite levels and dominance rank in male and female bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>)

The correlation between testosterone (T) and dominance rank may vary among species, and is expected to become stronger as the importance of aggressive competition for rank increases. However, it may also vary among social situations within a species, showing a stronger correlation during socially unstable periods. Knowledge on this topic in great apes, especially in females, is scant. This study presents the first data on the relationship between T and dominance rank in both sexes of the bonobo (Pan paniscus). For each period (four socially unstable and two stable ones), linear rank orders were determined and subsequently correlated with the accompanying mean urinary T-metabolite concentrations (measured as immunoreactive 5-androstan-17-ol-3-one). No correlation between these two variables was found for either sex among individuals during socially unstable or stable periods. Also, within an individual over the six periods, no relationship of T with rank could be demonstrated. These results suggest that either the outcomes of aggressions have no influence on T levels, or such clear outcomes appear insufficiently frequent to affect T levels over longer periods. Even during the unstable periods, the rate of aggressions was not higher than during stable periods, suggesting that frequencies of aggression have little effect on rank. Further analyses indeed demonstrated no correlation between frequencies of overall aggressions or any type of aggressive behavior separately, or rank. Perhaps factors other than the frequency of displayed aggressions alone have a marked influence on a bonobos rank, for example, coalition partners. Overall, in bonobos, T apparently does not form a physiological reflection of social status.     

A field survey of the bredding habits of Eretmodus cyanostictus, a biparental mouthbrooding cichlid in Lake Tanganyika

Oral incubation of young or mouthbrooding reduces the selective advantages of care by two parents and thus biparental care is rare among mouthbrooding fish. We surveyed the breeding biology of Eretmodus cyanostictus, a biparental mouthbrooder from Lake Tanganyika, to understand what factors maintain biparental care. We found larger males than females, a male-biased sex ration and indications that spawning is synchronized around the full moon. These preliminary findings suggest that the benefits of desertion for males are low; males may maximize their reproductive success by helping raise young while females regain reproductive condition.