Agonistic relationships between female mountain gorillas (Gorilla gorilla beringei)

Females in some mountain gorilla groups can be ranked on the basis of decided, non-aggressive approach-retreat interactions. However, data on four gorilla groups show that females do not form clear agonistic dominance hierarchies. Most aggressive interactions between females were undecided, and most dyads had undecided dominance relationships (Fig. 2). Females ignored aggression or responded to it aggressively more often than they responded submissively (Fig. 3). Moreover, directional inconsistency in aggression (including escalated contests) was high, and in some groups females showed significant bidirectionality in initiation of aggression (i.e., the more often they received aggression from partners, the more often they directed aggression at those partners; Fig. 4). Assigning ranks on the basis of aggressive interactions has limited power to explain variation in rates of initiating aggression and in responses and outcomes to aggression. Aggression was most common during feeding, but usually did not interrupt feeding bouts. It was proportionately more common in contexts that seem to involve competition for access to males (Figs 5, 6). This is consistent with the argument that females depend crucially on male services. As predicted by current models of female primate social relationships, agonistic relationships between gorilla females are unlike those typical in cercopithecine primates. They resemble those of some other primates and some equids in which contest competition for food is relatively unimportant and ecological costs of female transfer are low.     

Female-enhanced male competition determines the first mate and principal sire in the spider Linyphia litigiosa (Linyphiidae)

Summary Two to five days before sexual maturation, female sierra dome spiders (Linyphia litigiosa: Linyphiidae) undergo a transformation in their behavior toward males that visit their webs. During this latter part of their penultimate instar, females change from consistently positioning themselves far away from males to actively maintaining close proximity, reactions I call avoidant and associative behavior, respectively. Consistent associative behavior ceases after the female's first mating and thus is limited to soon-to-mature penultimate females. When a mate-seeking male fords an associative female, he attempts to guard her until she matures; this is often a multi-day affair. In contrast, males guard immature avoidant and mature mated females for only a single day. This dichotomy in male guarding times can be understood by the fact that associative behavior signals that the female will soon develop peak reproductive value. Upon completion of their final molt, 98% of females immediately mate with the current guarding male. Secondary suitors are not as likely to achieve mating. Moreover, first mates father 1.8 times more offspring, on average, than secondary mates. Whenever they meet on any female's web, males fight until one of the contestants withdraws. Fights typically are intensive, occasionally deadly, and often result in usurpation of the web by the newly arriving male. Larger males win more fights, but other qualities (e.g., vigor and persistence) appear to be important when contestants differ by less than 10–20% in body weight. Prolonged (i.e., multi-day) guarding of associative females enhances the intrasexual selection process by ensuring that every male that arrives at the web finds it already guarded. Therefore every male that finds the web becomes a participant in a series of male-male conflicts and web usurpations which span the period between the resident female's commencement of associative behavior and her sexual maturation. Since unforced male departures from the webs of associative females are rare, victors are retained on the web until they themselves lose a fight. This facilitates a steady increase in the fighting ability of sequential guards throughout the associative period, up until female maturation and mating. On my study site, first mates represented the final winners in a combative sorting process based on a minimum average of 2 fights; they were heavier and larger than secondary mates and randomly sampled males. The combination of (1) associative behavior by nearly mature females, (2) high mating propensity of newly mature females, and (3) first male sperm priority, constitutes a system whereby females enhance male-male competition and boost the expected fighting prowess of the principal sire of their progeny. Since males appear to make no material contribution toward progeny, the female's behavior probably functions to improve the genetic constitution of the offspring. In addition, the timing of associative behavior may limit prolonged guarding by food-stealing males to a period (1) encompassing the female's pre-molt fast and (2) before the heavy yolking of eggs, thereby ameliorating the nutritional costs of intrasexual selection.     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Trade-off between pre- and postcopulatory sexual cannibalism in a wolf spider (Araneae,Lycosidae)

Sexual cannibalism can occur before, during or after mating. Relatively few experimental studies have examined why there is variation in the timing of sexual cannibalism. We examined the latency and number of attacks required for female spiders to capture male spiders pre- vs. postcopulation. We also examined the effects of female mating status and hunger level on the occurrence of pre- and postcopulatory cannibalism, which reflects the contribution of both relative capture success and female motivation to cannibalize males. Precopulatory cannibalism occurred after a shorter interval and required fewer chases and physical interactions for the female to successfully capture the male than was the case for postcopulatory cannibalism. Virgin females were more likely to engage in postcopulatory rather than precopulatory cannibalism and mated females vice versa. Those virgin females that did engage in precopulatory cannibalism had significantly lower body condition than virgin females engaging in postcopulatory cannibalism. While precopulatory cannibalism occurred more quickly and required fewer attacks by females, it comes at a potential cost of not mating with males. Hence, females are more likely to engage in precopulatory cannibalism if they have already mated or, if virgins, if they have low body condition. These results indicate that the decision of when to cannibalize males is dynamic and depends upon the relative value of a male as a mate versus a meal.     

Prior encounters with the opposite sex affect male and female mating behavior in a wolf spider (Araneae,Lycosidae)

Mate availability can vary widely in nature depending upon population density and sex ratio and can affect the ability of individuals to be selective in mate choice. We tested the effects of prior encounters with the opposite sex (i.e., exposure to the opposite sex either with or without mating) on subsequent mating behavior in two experiments that manipulated mate availability for both males and females in the wolf spider, Hogna helluo. The probability of mating in the experimental trial depended upon whether the prior encounter involved mating or not, and males and females responded in opposite directions. Exposure without mating resulted in a higher subsequent frequency of mating for females and a lower subsequent frequency of mating for males, while prior mating experience resulted in a lower frequency of female remating and a higher frequency of male remating. Prior exposure without mating did not affect female aggression. However, mated females engaged in precopulatory cannibalism more frequently than virgins. Mated males escaped postcopulatory cannibalism more frequently than virgins. Our results show that males respond to exposure without mating in the expected manner. However, prior mating (1 week earlier) had unexpected effects on males, which may be due to mated males being of higher quality. There were little or no effects of the size of the prior exposure individual or mate on subsequent mating behaviors. Further research is needed to determine why different species use different degrees of prior information in mate choice.     

Agonistic interactions in siskin flocks: Why are dominants sometimes subordinate?

Summary Dominance interactions among captive siskins were examined to see if the behavior of dominants reduced the risk of subordinates leaving the flock. The outcome of aggressive encounters was related to the possession status of the two birds (i.e., which bird was first to arrive at the contested resource) and the type of aggression used (i.e., display or attack). More dominant birds were successful whether they were possessors or intruders, and whether they attacked or displayed. When possessors, they tended to display, presumably because of the greater cost of attack and the lack of substantial benefits associated with it. When intruding, they tended to attack, possibly because attack is slightly more successful than display. When initiating encounters against dominants, subordinates were more successful if they were possessors than if they were intruders. Subordinates tended to use displays whether they were possessors or intruders, even though when the birds were intruding, displays were less successful than attacks. Subordinates may use display when intruding because attack holds a higher risk of retaliation. The fact that siskins can repel more dominant intruders merely by using displays suggests that dominants, by respecting possession and allowing reversals, are able to reduce the likelihood that subordinates will leave the flock. This may be to the dominants' long-term advantage, since they gain benefits from being in stable flocks.     

Partitioning behavior and intra- and interspecific interactions: a comparison between male and female groupers,Cephalopholis spiloparaea (Pisces: Serranidae: Epinephelinae)

Behavior of male and female groupers, Cephalopholis spiloparaea, observed, between 1990 and 1991 in Sasanhaya Bay, Rota, Northern Mariana Islands, was compared to detect differences in proportional use between sexes, and between daylight and pre-courtship time periods. This species has male-dominated haremic groups. Time partitioning could not be measured exactly. Instead, the numbers of intra-and interspecific interactions, foraging, resting, swimming, and maintenance behaviors of each sex were measured during the observation periods. The proportion of acts in each category out of the total number of acts, termed effort, was used in comparisons. Males were predicted to devote more effort toward intra-and interspecific interactions compared to females and to maximize reproductive success. Females were predicted to devote more effort towards foraging, compared to males. This behavior maximizes reproductive effort. Both were predicted to engage in greater intra-and interspecific interactions during the pre-courtship period, corresponding both to the approaching time of courtship and to increases in the numbers of potential spatial and trophic competitors. Intra-and interspecific interactions of both males and females were greater during the pre-courtship period. Differences in all classes of behavior between sexes were only weakly significant for two, however. Males had greater effort in intra-and interspecific interactions. Females had greater effort only in maintenance behavior. Both differences were evident only during the pre-courtship period. Foraging behavior by both sexes was virtually absent during daylight and pre-courtship periods. Fish sought shelter and were not incidentally observed foraging after dusk, suggesting that this species actively forages later at night or during early morning hours, just prior to and during sunrise.     

Factors influencing cannibalism in the wolf spider Pardosa agrestis (Araneae, Lycosidae)

Cannibalistic tendencies are well known in spiders and may be a significant factor influencing population size. The wolf spider, Pardosa agrestis, is the dominant non-web-building spider in a wide range of central European agricultural habitats. Preliminary field observations indicated an extended reproductive period, which results in a very wide size distribution of juvenile instars. We hypothesised that if cannibalism is enhanced by differences in size, especially during periods when prey is scarce, these populations might be susceptible to cannibalism in an ecologically significant way. Laboratory studies were conducted on juvenile P. agrestis in arenas. We analysed the following specific aspects of cannibalism: (1) the effect of the weight ratio between the opponents; (2) the effect of weight per se, and (3) the role of hunger level in determining cannibalistic tendencies of spiders. The role of weight and hunger were analysed in separate experiments, in both cases by controlling for the other variable. The results showed that cannibalism was strongly positively correlated with both weight ratio and hunger, but absolute size/age of an individual could not predict the occurrence of a cannibalistic event. These experiments generated the plausible hypothesis that cannibalism might be an important phenomenon in the regulation of real populations, which should be tested specifically in future field experiments. Received: 15 December 1997 / Accepted after revision: 10 October 1998     

Mate with the young, kill the old: reversed sexual cannibalism and male mate choice in the spider Micaria sociabilis (Araneae: Gnaphosidae)

Female mate choice is regarded as a strong selective force that significantly affects male mating success. In extreme cases, mate rejection can result in sexual cannibalism. However, males may choose between their partners as well. The killing of potential female mates, i.e. reversed form of sexual cannibalism, may be related to male mate choice. We examined male mate choice in the spider Micaria sociabilis, focusing on the roles of female mating status (virgin/mated), size and age. Reversed cannibalism reached its highest frequency in the period of generation overlap, i.e. when young males from the summer generation met old(er) females from the spring generation. These results suggest discrimination against old(er) females. The frequency of cannibalism was not affected by female mating status or female size. However, larger males from the summer generation were more cannibalistic than smaller males from the spring generation. We conclude that reversed sexual cannibalism might be an adaptive mate choice mechanism and can be explained in the context of the aggressive spillover hypothesis.     

Determinants of paternity success in the spider Pholcus phalangioides (Pholcidae: Araneae): the role of male and female mating behaviour

In double mating experiments, we examined whether and to what extent various male and female behavioural traits influence the course of mating and fertilization success in the cellar spider. In males, we focussed on pre-copulatory behaviour and on the rhythmic twisting movements that the male performs with his pedipalps during copulation. In females, we investigated remating decisions and the effect of female termination of copulation. Second males fertilized a high proportion of the eggs (P ₂: median 89%) despite much shorter second matings, with high variation in relative paternity success. The number of pedipalp movements (PPMs) of either male was a better predictor of paternity than copulation duration. Our results suggest that in second matings, PPMs help to remove sperm from previous males, whereas in first matings a high number of PPMs enhances fertilization success, either due to numerical sperm competition or cryptic female choice. Furthermore, we found a negative male age effect on paternity in second matings, implying that age-related deterioration of spermatozoa may promote variation in fertilization success. Female receptivity decreased significantly in second matings; only 70% of the females remated. Females that accepted a second copulation were found to terminate these much earlier and with higher probability than first matings. This suggests that the intensity of conflict between the sexes is higher in second matings. Increased intensity of sexual conflict may be responsible for stronger selection on male traits, as pre-copulatory behaviour and age only affected male copulatory performance and paternity in second matings. Electronic Publication     

Sexual cohabitation as mate-guarding in the leaf-curling spider Phonognatha graeffei Keyserling (Araneoidea, Araneae)

The leaf-curling spider Phonognatha graeffei incorporates a twisted leaf into the central hub of its orb-web that is used as a retreat. This species is unusual among orb-weaving spiders because males cohabit in the leaf retreat with both immature and mature females, mating with the former shortly after the female molts. Cohabitation appears to be a form of mate-guarding because cohabiting males respond agonistically to rival males that venture onto the web, and their behaviour depends upon the reproductive status of the female; males defending immature females are more aggressive than those defending virgin, adult females. Males copulate with previously mated females for significantly longer than with virgin females. Females may cannibalise cohabiting males, which occurs independently of whether the female has been deprived of food. Females that cannibalise a single male do not have a higher fecundity than non-cannibalistic females. Received: 2 February 1996 / Accepted after revision: 27 October 1996     

Chemically mediated interactions between the red algaPlocamium hamatum (Rhodophyta) and the octocoralSinularia cruciata (Alcyonacea)

Interactions between the red algaPlocamium hamatum J. Agardh (Rhodophyta) and other benthic organisms including the alcyonacean soft coralSinularia cruciata (Tixier-Durivault) were investigated on an inshore fringing reef environment in whichP. hamatum was the dominant large fleshy alga. Field observations of sessile reef organisms including octocorals and sponges living in close proximity toP. hamatum revealed that varying degrees of tissue necrosis were suffered by the invertebrates when in physical contact with the alga. In order to establish whether the chemical constituents of the alga, especially chloromertensene, played a role in this necrosis, manipulative field experiments were carried out in the Pelorus Channel, Palm Island group (18°34S; 146°29E), North Queensland, Australia, in November and December 1988. The first experiment involved the relocation of healthy plants and soft corals into contact and non-contact situations on a mesh grid. In all cases of contact betweenP. hamatum andS. cruciata, the soft coral suffered tissue necrosis (n=6,p=0.0022). The second experiment had the same design, but involved the use of artificial plants both uncoated and coated with natural levels of chloromertensene, in contact withS. cruciata. In all cases of contact with coated treatments, necrosis was observed inS. cruciata (n=4,p=0.025). In cases where uncoated artificial fronds were placed in contact with soft corals,S. cruciata showed minor abrasion effects, but no appreciable necrosis. Coated treatments were not fouled by epiphytes during the experiment and were not consumed by predators. Uncoated treatments were rapidly reduced in size by predation and any remaining material was biofouled. These experiments thus demonstrated that the deleterious effects observed in soft corals in the field were caused by contact with the algaP. hamatum, that these effects were indeed chemically mediated by chloromertensene, and that physical contact without chemical intervention caused no such deleterious effects. This is the first experimental evidence which conclusively demonstrates allelopathy between an alga and other marine organisms and identifies the compound responsible for the observed allelopathic effects.     

Reproductive success of solitary and communal Philoponella oweni (Araneae: Uloboridae)

Summary Philoponella oweni (Uloboridae) is a facultatively communal orb-web spider found in arid regions in the southwestern United States. Within one habitat solitary adult females and communal groups of adult females coexist.This study compared the reproductive output (egg production) and reproductive success (production of live offspring) of solitary and communal females. There was no significant difference in the mean number of egg-cases produced by solitary and communal females, but communal females produced significantly more eggs per egg-case. However, communal females suffered significantly higher rates of egg-case parasitism from the pteromalid wasp Arachnopteromalus dasys. As a result, the mean number of liver offspring produced by solitary and communal females did not differ. Thus, both strategies led to roughly equal reproductive success. The advantages of aggregation of adults may be significant in the evolution of social behavior in spiders.     

Kleptoparasites: a twofold cost of group living for the colonial spider, Metepeira incrassata (Araneae, Araneidae)

Several species of kleptoparasitic and araneophagic spiders (Araneae: Family Theridiidae, Subfamily Argyrodinae) are found in colonial webs of the orb-weaving spider Metepeira incrassata (Araneae, Araneidae) from Mexico, where they steal food and/or prey upon their spider hosts. Census data from natural M. incrassata colonies reveal that the incidence of these species increases with colony size. This pattern may reflect the presence of several other orb-weaving spiders, each with their own kleptoparasitic species, invading larger M. incrassata colonies. As the number of these associated spiders increases, so does the density and number of Argyrodinae species in M. incrassata colonies, suggesting that associated spiders might reduce their own kleptoparasite load by building their webs within M. incrassata colonies. This represents a twofold cost to M. incrassata, as a field enclosure experiment revealed that a primarily kleptoparasitic species (Argyrodes elevatus) may reduce prey available to their hosts, but a kleptoparasitic/araneophagic species (Neospintharus concisus) inflicts high mortality upon M. incrassata. However, the cost of kleptoparasitism and predation by these species may be offset in part for M. incrassata individuals in large colonies by certain defensive mechanisms inherent in groups, i.e., “attack-abatement” and “selfish herd” effects. We conclude that increased occurrence of kleptoparasitic and/or predatory Argyrodinae spiders is a consequence of colonial web building and is an important potential cost of group living for colonial web-building spiders.