Shoaling Decisions in Angelfish: The Roles of Social Status and Familiarity

Abstract Past research has shown that angelfish, Pterophyllum scalare , are capable of discriminating between shoals composed of familiar dominant and subordinate companions, whereas they show no preference for shoals of unfamiliar conspecifics. In this study, the relative importance of familiarity and social status (shoal factors) on the shoaling decision of juvenile angelfish, which also differed in social status (individual factor), was investigated as very little is known about such tradeoffs in fishes. Dominant and subordinate individuals were given the choice to shoal with a group of conspecifics composed of familiar dominants vs. unfamiliar dominants and composed of familiar subordinates vs. unfamiliar subordinates. The findings demonstrate that fish with different social status differed in their shoaling preference. Subordinate test fish showed a preferential association with familiar subordinates over unfamiliar subordinates, but preferred the unfamiliar shoal over the familiar one when both shoals were constituted by dominant individuals. The shoaling behaviour shown by dominant test fish, on the other hand, indicated no significant preference for any of the shoals regardless of their composition. A replicate preference test carried out 2 h 30 min after the first one indicated that the association pattern was relatively consistent. Results suggest that angelfish are able to differentiate between the stimulus shoals and demonstrate that the pervasive influence of familiarity on the shoaling decision may be restrained or overridden by the composition of the familiar shoals and the social status of the test fish.     

Rank and reproduction in cooperatively breeding African wild dogs: behavioral and endocrine correlates

African wild dogs (Lycaon pictus) live in cooperative packswith a clear-cut dominance hierarchy in each sex. Reproductionis largely monopolized by the dominant male and female. Alphafemales produced 76% of all litters in the Selous Game Reserveand 81% in Kruger National Park. Only 6-17% of subordinate femalesgave birth each year, compared to 82% of dominant females. Innonmating periods, subordinate females had higher estrogen levelsand higher estrogen/progestin ratios than alpha females, apparentlypreventing ovulation. During mating periods, subordinate femaleshad lower estrogen levels than dominants, mated less often,and were less aggressive. Subordinate males mated at low rates,wore less aggressive than dominants, and had lower testosteronelevels. Beta males were similar to alpha males behavioraDy andhormonaUy, suggesting that alpha males may share paternity withbeta males. If paternity is more evenly shared than maternity,then subordinate males have a larger incentive than subordinatefemales to remain in the pack. Following this expectation, dispersalin Selous was female biased (49% versus 24% dispersing annually).Perhaps as a result of mortality associated with dispersal,the adult sex ratio was male biased, although the pup sex ratiowas unbiased. In Kruger, neither dispersal nor the adult sex-ratiowas biased. Reproductive suppression is widely thought to becaused by social stress in subordinates, but basal cortkosteronelevels were higher in dominants than in subordinates     

Group composition affects male reproductive partitioning in a cooperatively breeding cichlid

Individuals within groups of cooperatively breeding species may partition reproduction, with the dominant pair often taking the largest share. The dominant's ability to reproductively control subordinates may depend on differences in competitive ability, due to, e.g. body size differences, but may also depend on the number of same‐sex competitors inside the group. We tested experimentally whether subordinates reproduce more when these subordinates are large or when a second subordinate of the same sex need to be controlled by the dominants, using the cooperatively breeding cichlid Neolamprologus pulcher. Dominant pairs were assisted by a large and a small unrelated subordinate; sexes of these fish were varied in a full‐factorial design (giving four treatments). Dominant males lost significantly more parentage to the large subordinate male when a small subordinate male was also present, compared to when a small subordinate female was present. However, subordinate paternity was generally low and did not significantly curb total dominant male reproductive output, which was more affected by the sizes and numbers of reproductive females present inside his group. Dominant female maternity, clutch sizes and total output did not depend on the treatments. Subordinate–subordinate reproduction was virtually absent (one out of 874 offspring). Female subordinates were more likely to provide care for their own broods. In contrast, male subordinates did not adjust their level of care to their parentage. Variability in female subordinate alloparental brood care was particularly high, with females showing more care than males in general. We also detected effects of growth rate and food ration on parentage independent of the treatments, most notably: (i) a trade‐off between dominant male growth rate and paternity; (ii) a decrease in dominant male paternity with increasing food ration; (iii) a positive effect of growth rate on paternity in small males. We conclude that dominant males should be sensitive to the number and sizes of subordinate males present in their group, particularly when these subordinates are not helpful or grow fast, and food is plentiful. Dominant females should be less sensitive, because female subordinates do not appear to impose reproductive costs and can be helpful through alloparental brood care.     

鳄蜥对熟悉和陌生个体气味的辨别

个体辨别对于减少同种争斗以及配偶选择具有重要意义。我们用棉棒粘取鳄蜥(Shinisaurus crocodilurus)尿液作为气味源,以香水作为对照,测定鳄蜥对熟悉个体气味、陌生个体气味以及香水的舔舌次数和舔舌潜伏期,来探讨鳄蜥通过化学信息辨别熟悉和陌生个体的能力。结果显示,不论是雌性还是雄性,对不同个体尿液的舔舌次数均显著高于对香水的,舔舌潜伏期显著短于香水的;尽管雄性对陌生同性个体气味与熟悉同性个体气味的舔舌次数无显著差异,但对前者的舔舌潜伏期显著短于后者;雄性对陌生雌性气味的舔舌次数显著多于熟悉雌性气味的,对前者的舔舌潜伏期显著短于后者;雌性对陌生雄性气味的舔舌潜伏期显著短于对熟悉雄性气味的;雄鳄蜥对陌生雌性气味的舔舌次数显著多于雌鳄蜥对陌生雄性的。结果表明,鳄蜥能辨别同种个体的化学信息,并能通过化学信息来辨别熟悉和陌生个体,推测鳄蜥的这种辨别能力对其领域分配以及繁殖交配有重要作用。     

Olfactory communication and the importance of different odour sources in the ferret (Mustela putorius f. furo)

Olfactory communication has a major role in solitary carnivores, particularly during the breeding season. Scent-marking behaviour in ferret (Mustela putorius f. furo) has been well studied, but not the significance of the deposited odours for the animal. We compared the interest in urogenital, anal and body odours of opposite-sex conspecifics between male and female ferrets and investigated both sexes’ discrimination ability (familiar vs. unfamiliar) for these three opposite-sex odours and the use of its ability by females during the breeding season. Males are more interested than females in the urogenital odour of the opposite sex. Male body odour probably did not interest females and males are somewhat interested in the female one. However, anal odour of the opposite sex is as interesting for males as for females and both sexes are able to discriminate familiar anal odour from an unfamiliar one. Moreover, females could use this ability when they must choose to have some sensory contact between olfactory familiar and unfamiliar males since they prefer to be in contact with the former. These results are coherent with behavioural differences observed between male and female solitary mustelids in the field and in captivity, and the consortship success of resident males compared to roaming males.     

Social Status and Feeding in Atlantic Salmon Salmo salar Parr: The Effect of Visual Exposure to a Dominant

Groups of four Atlantic salmon parr were established and the dominant individual identified. Feeding responses were then observed in the remaining (subordinate) fish housed in one of three conditions: in the same compartment as the dominant fish (full exposure), separated from it by a transparent partition (visual exposure) or behind an opaque partition (isolated). Subordinate parr in the full exposure condition responded to and ate fewer food pellets than did the dominant fish. However, subordinate parr in the visual exposure condition and in the isolation condition fed actively, indicating that neither visual intimidation nor long-term stress-induced reduction of appetite were responsible for the reduced food intake of subordinate parr in direct contact with dominants. Dominant parr and subordinates in the visual exposure condition, but not subordinates in the exposed condition, moved further from their feeding station when attacking food items that were contested by a competitor compared with the distance moved for uncontested items. The presence of a competitor therefore forced both dominant and visually exposed subordinates to pay higher costs while foraging.     

Social status, activity and preputial glands of wild and domestic house mice

Scent marking by deposition of urine, and the preputial glands, of adult, male, wild house mice, Mus mmmlus L., were studied and compared with those of an outbred domestic strain. The preputial glands of dominant wild mice were always heavier than those of subordinates. No dominance relationships could be established among the domestic mice. For study of scent marking each mouse was observed singly in a residential maze. Dominant wild mice marked more than the subordinates. The domestic mice scent-marked much less even than the subordinate wild mice. Subordinate wild mice spent less time than the dominant wild mice outside the nest; but the number of excursions outside the nest made by the subordinates resembled that of the dominants. Hence social status influenced the pattern of movements in a structured environment.     

Mating Resets Male Cricket Aggression

An animal’s motivational state can significantly impact its behavior. We examined the effects of mating on the aggression of male Acheta domesticus crickets. Pairs of males were allowed to establish dominance and subordinance and were then physically separated. Subordinate males were then allowed to either copulate with a female or to have chemo-tactile contact with, but to not copulate with, a female. Less than 15 min after separation, all male pairs engaged in a second agonistic encounter. Subordinate males that copulated with females were significantly more aggressive toward their dominant partners than un-mated subordinate males. Many mated subordinates became dominant. Allowing a subordinate male to contact, but not copulate with, a female had a similar effect, suggesting that chemo-tactile cues from the female are sufficient to elicit this change in aggression.     

雌性布氏田鼠对雄鼠气味的辨别

通过雌性布氏田导其配偶和陌生雄鼠气味的辨别实验,以及无交配经历的成年雌鼠对社会等级不同的陌生雄鼠气的辨别实验发现,有生育经历的雌鼠对陌生雄鼠气味的探究行为明显多于对配偶气味的控究,但雌鼠在配偶雄鼠气味周围的停留时间、自身修饰行为和何处行为的发生频次和持续地陌生雄鼠,没有交配经历的雌鼠对处于从鼠地位的雄性个体气味探究行为多于对优势雄性的控究,而其他行为表现没有明显差异。表明,雌性布氏田鼠可以利用气味     

The differential responses to the odor of unfamiliar male conspecifics in dominant,subordinate, and isolated mice

Male ddY mice were used to investigate the responses toward the odor of unfamiliar male mice. They were reared either in isolation from 3 weeks of age or in pairs. At 10 weeks of age, they went through a 4-day test period, in which the responses of the mice toward the odor of unfamiliar dominant and subordinate mice were investigated by differentiating the responses into the time to investigate the odor and the final decision of avoidance or preference. All the mice went through the encounter tests after this odor test in order to investigate the relationship between the responses toward the odor and the behavior toward the odor donor. The pair-reared dominants and the subordinates responded to the odor of the dominant and subordinate mice differentially. This indicated that they could discriminate the dominance status of unfamiliar mice by odor. Isolated mice did not respond to the odor of the dominant and subordinate mice differentially. Social experiences were concluded to be necessary to discriminate the dominance status of unfamiliar mice through odor. The length of the investigation time and the final decision of avoidance or preference did not correlate in pair-reared or isolated mice, and these 2 scales were concluded to be different. But the dominant mice that investigated the odor for longer periods tended to show aggression in the later encounter test.     

The Effects of Food Deprivation in Dyadic Encounters in Apodemus sylvaticus

The nature of dominance in relationships among male Apodemus sylvaticus was investigated using multivariate analysis and by experimental manipulation of the behaviour of dominants. The behaviour of dominant mice was typified by greater frequencies and durations of exploration, approach and pursuit of subordinates in short encounters in a small arena. The direction of encounters normally remained constant over 48 h. When the dominant mouse in a dyad was deprived of food there was a change in its behaviour and an immediate change in the behaviour of the subordinate. Subordinates showed greater interest in starving, though known, dominants, explored more freely and indulged more in pursuit than control subordinate mice i. e. those matched with dominants on an adequate diet. Results suggest that in A. sylvaticus, dominance is associated with the inhibition of the activity of a subordinate and the relationship is maintained by the behaviour of the dominant rather than the deference of the subordinate.     

Effects of social stress during early pregnancy on litter size and sex ratio in the golden hamster (Mesocricetus auratus)

Primiparous female hamsters were mated to proven breeders and stressed during early pregnancy. Females were housed singly throughout gestation except for Days 4, 5 and 6 when they were paired for 10-min intervals 3 times each day with another female matched for age, weight and day of pregnancy. Within each of the pairs, one female was consistently dominant to the other. Controls were exposed to a novel area instead of a conspecific. At parturition, all pups were counted, sexed and weighed. There were no significant differences between litter sizes or sex ratios (defined as % male) of control and dominant females. Litter sizes produced by control or dominant dams were significantly larger than those of subordinate dams, and litter sex ratios of dominants were significantly higher than those of subordinates. Subordinate dams produced fewer males than did dominant dams, but there was no difference in the number of females produced. Also, subordinate dams produced smaller pups than control dams. Examination of uterine implantation sites and fetal resorptions indicated that fetal loss occurred between Days 5 and 10 of pregnancy. These results suggest that subordinate dams produce smaller litters via selective resorption or spontaneous abortion of males in utero and that those males they do produce are smaller than those produced by dominant or control dams. We suggest that males are more susceptible in utero to effects of maternal stress in this species, and may require more maternal investment to survive to term.     

Effect of the Presence of Subordinates on Dominant Female Behaviour and Fitness in Hierarchies of the Dwarf Angelfish Centropyge bicolor

In many hierarchical animal societies, dominant individuals control group membership owing to their power to evict subordinates. In such groups, the presence of subordinates, and therefore group stability, is continually dependent on subordinates being tolerated by dominants. The dominant decision to tolerate or evict is, in turn, dependent on the costs and benefits to dominants of subordinate presence. We investigated the effect of subordinate presence on dominants in the female dominance hierarchy of the dwarf angelfish Centropyge bicolor, using both observations of natural groups and experimental removals of subordinates. We found that the presence of subordinates had no effect on dominant access to resources, as measured by dominant foraging rates and home range areas, nor on dominant fitness, as measured by growth rates and spawning frequencies. Our results suggest that the presence of subordinates has a neutral effect on the current fitness of dominants, so that dominants have no great incentive to evict subordinates. We discuss the possibility that tolerance of subordinates might be further explained by considering future fitness, as dominant females in these haremic protogynous angelfish stand to inherit the male position, whereupon subordinate females change from potential competition to useful mates.     

Subordinate superb fairy-wrens (Malurus cyaneus) parasitize the reproductive success of attractive dominant males

Explanations of cooperative breeding have largely focused on the indirect benefits philopatric offspring gain from investing in kin. However, recent molecular studies have revealed that in many species subordinates provision unrelated offspring. This has led to the re-evaluation of the direct and indirect benefits of helping behaviour. In this study, we used microsatellite genotyping to assess the extra-group reproductive success of subordinate superb fairy-wrens (Malurus cyaneus), a species with extremely high rates of extra-group paternity. Extra-group subordinate males sired 10.2% (193 out of 1895) of all offspring sampled between 1993 and 2000 and 21.4% (193 out of 901) of all illegitimate offspring sired by known males. The extra-group success of subordinates was greatly influenced by the attractiveness of their dominant male. Subordinates of attractive dominants sired more extra-group young than did average dominants. Evidence suggests that mate choice in superb fairy-wrens is error-prone and subordinates can gain direct reproductive benefits through parasitizing the reproductive success of attractive dominants.     

Gender Differences in the Costs that Subordinate Group Members Impose on Dominant Males in a Cooperative Breeder

In group-living species, a dominant male's ability to monopolize reproduction, and the cost of doing so, are expected to vary with a group's gender composition. We used spawning observations of a group-living cichlid, Neolamprologus pulcher , to test this expectation. We constructed groups that contained a dominant breeding pair and either two male subordinates, one male and one female subordinate or two female subordinates. Parasitic spawning by male subordinates was more common in groups with two male subordinates than in groups with one male and one female subordinate. Female subordinates were never observed laying eggs in dominant females' clutches, but three female subordinates laid independent clutches. During spawning, frequencies of dominant male aggression towards male and female subordinates were similar. Dominant males were less aggressive during non-reproductive periods. The declines were greater for female subordinates, such that, during non-reproductive periods, dominant males were more aggressive towards male subordinates. Aggression towards each subordinate was also affected by the second subordinate's gender; the direction of that effect differed for large and small subordinates. Male subordinates approached breeding shelters less often than female subordinates, and both male and female subordinates approached shelters more frequently when the second subordinate was male. Collectively, these patterns suggest: (1) that male subordinates impose higher costs on dominant males than female subordinates do and (2) that the presence of a second male subordinate imposes additional costs beyond those of the first male subordinate. We discuss the implications of these effects for dominant and subordinate group members.     

Recognition of interspecific familiar versus unfamiliar odours among bank voles and yellow-necked mice

Two experiments (laboratory and field-laboratory) were designed to determine whether individual bank volesClethrionomys glareolus (Schreber, 1780) and yellow--necked miceApodemus flavicollis (Melchior, 1834) could distinguish heterospecific odour cues from familiar and unfamiliar individuals. In the laboratory experiment each male bank vole was familiarized for 24 h with odour (cotton wool impregnated with urine and faeces) of male yellow-necked mice and yellow-necked mice were familiarized with odour of male bank voles. In the field-laboratory experiment the individual bank voles and yellow-necked mice captured at the same point were considered familiar and transfered to the laboratory. In laboratory, these individuals were tested in a box (for 5 h) affording them the choice between the odours of familiar and unfamiliar heterospecific males. Bank voles discriminated between familiar and unfamiliar yellow--necked mouse odours. Male yellow-necked mice seemed to have a similar ability to recognise odours of familiar bank voles. It is proposed that interactions between these two species occur not only on the species level, but also on the level of individual. This phenomenon (probably asymmetric) can play an important role in spatial orientation, and influence direct contacts between individuals of these species.     

The Effect of Social Hierarchy on Captive Coyote (Canis latrans) Foraging Behavior

Foraging efficiency of individuals in pack forming species may be influenced by social dynamics within a pack. The effects of social hierarchy in particular may influence individual foraging behavior in canids, such as coyotes (Canis latrans). To examine the impact of social hierarchy on foraging behavior, we tested 16 captive coyotes in eight naturally established dominant–subordinate pairs, using the guesser–knower paradigm. We measured the efficiency of subordinate coyotes to relocate a food resource when alone and then allowed pairs to forage together, such that subordinates had prior knowledge of food location but dominants did not. To determine whether (1) subordinates used a direct or discursive strategy to obtain food in the presence of a dominant and (2) dominants used an exploitative or independent strategy to obtain food in the presence of a subordinate with previous knowledge, we measured their search efficiency (e.g., correct choice of area, feeder, and latency to correct feeder). Results showed subordinates learned to relocate food and increase efficiency when alone. In a social context, however, subordinate efficiency decreased. That is, subordinates approached the correct area, but searched more feeders before finding the correct one. Dominants initially used an independent search strategy but then quickly displaced the subordinate and monopolized the resource, reducing subordinate efficiency further. Despite continual displacement and reduction in efficiency, subordinates did not alter their foraging strategy over time. Our results suggest prior information can improve individual foraging advantage, but that social status strongly impacts individual foraging efficiency in social species such as coyotes.     

Male Mating Success in the Presence of a Conspecific Opponent in a Madagascar Hissing Cockroach,Gromphadorhina portentosa (Dictyoptera: Blaberidae)

The influence of a conspecific competitor on male mating behavior was examined in a Madagascar hissing cockroach, Gromphadorhina portentosa. Previous studies have suggested that both male-male competition and female discrimination during courtship interactions may influence male mating success. Familiar pairs of males with a known social association were placed in an arena with a single virgin female and observed. As expected, subordinate males mated significantly less often than their dominant opponents. In pairs in which one male mated, dominant individuals limited the access of subordinates to females. Dominant males displayed an increased frequency and duration of interaction with the female. However, in pairs where both males remained unmated, the mating behavior of dominant and subordinate males did not differ significantly. As interactions progressed, as in the case of males that remained unmated, subordinate males gained increased access to the female. Mated males tended to be larger than their opponent although within a rank, males that mated were no larger than those that remained unmated. These results are discussed in light of the possible roles of male-male competition and female discrimination during courtship interactions.     

Dominance Interactions Regulate Worker Mating in the Polygynous Ponerine Ant Gnamptogenys menadensis

In queenless ants, workers compete aggressively to be reproductives instead of sterile helpers. To limit the number of reproductives, either mating or egg-laying by mated workers can be regulated. In Gnamptogenys menadensis , all mated workers reproduce, and field data presented here indicate that mating is not random. We removed gamergates experimentally (n=16 groups) to induce mating and gamergate replacement. Virgin workers engaged in agonistic interactions, allowing us to classify them into two groups: dominants performed antennal boxing and biting, while subordinates did not. However, a few subordinates cooperated to immobilize individual dominant workers. This effectively reduced the number of dominants because immobilized dominants became subordinates. Six to 12 wks after the removal of reproductives, between one and eight dominants per colony started to perform 'sexual calling' outside the nest entrance. Sixteen (out of 33) dominants mated with foreign males introduced in five experimental groups. None of the subordinate workers mated. The probability of mating is thus regulated by dominance interactions among workers and immobilizations initiated by infertile workers in this polygynous queenless ant.     

Differential response to the odor of familiar intruder mice in male mice (Mus musculus)

Responses to the odor of familiar intruder male mice according to their dominance were investigated. Responses were classified into 2 aspects: the investigation of the odor and the decision-making regarding avoiding it or not. The results varied according to the dominance of the respondents and the odor donor, and also according to the context of previous encounter situations. The dominants that had attacked an intruder dominant mouse responded randomly to its odor, whereas the dominants that had fought with it tended to avoid the odor. The subordinates that had observed an intruder dominant mouse being attacked by its dominant cagemate preferred the passage with the intruder's odor. The odor of a subordinate mouse was neither avoided nor preferred by either the dominants or subordinates. It was suggested that mice distinguished the dominance of the odor donor regardless of the context of the previous encounter situation, but they responded differently according to it and also according to their own dominance status.