Female choice on male quantitative traits in lizards — why is it so rare?

Female choice on the basis of male traits has been described in an array of taxa but has rarely been demonstrated in reptiles. In the sand lizard (Lacerta agilis), and possibly in other non-territorial reptiles, a male's contribution to a female's fitness is restricted to his genes. In order to choose males of high genetic quality, females have to trade the fitness gain against the costs of active choice. In a Swedish population of sand lizards, long-lived males sired offspring with higher embryonic survival compared to offspring sired by short-lived males. In spite of this female sand lizards did not mate selectively with older and/or larger males. There appeared to be mo reliable cues to male longevity; age-specific male body size was highly variable. Furthermore, estimates of male nuptial coloration did not covary with ectoparasite load and, hence, females cannot use male coloration as a cue to heritable resistance to pathogenic parasite effects. When cues to male genetic quality are poor, or inaccurate, and males make no parental investment, we predict that female choice will be rare. Sand lizard females mating with many partners lay clutches with higher hatching success. Thus, females may obtain good genes for their young by multiple mating, thereby avoiding costs associated with mate choice.     

Sequential female choice and the previous male effect in sticklebacks

Summary Female choice, identified as a major force in sexual selection theory, has recently been demonstrated in a number of species. These tests concentrated on simultaneous choice situations although females have to compare males sequentially in most territorial species, which is the more demanding task. Here it is shown that female three-spined sticklebacks, Gasterosteus aculeatus L., rate sequentially presented males according to their brightness. With increasing costs of sampling the females become less choosy. Furthermore, a male's attractiveness has a significant effect on the female's rating of the next male; a given male is rated higher when preceded by a duller male than by a brighter one and vice versa. Female sticklebacks use a stochastic decision rule in sequential mate choice that is attuned to the attractiveness of the present and previously encountered male. This demonstration of a previous male effect not only indicates an efficient mechanism for finding the best of a number of males but also extends the applicability of sexual selection theory.Offprint requests to: T.C.M. Bakker     

Nestmate recognition and the ontogeny of acceptability in the ant,Leptothorax curvispinosus

Summary In social insects, there is often a brief period following eclosion when workers are highly acceptable in alien nests of their own or other species. This study tested for such an acceptance period in the facultatively polygynous ant, Leptothorax curvispinosus, and compared the duration and effectiveness of this period for conspecific and heterospecific introductions. Workers that eclosed and aged for 1–70 h or 30 days in isolation were introduced into either their parental nests (n=24), alien conspecific nests (n=265), or nests of the closely related and biologically similar species, L. longispinosus (n=341). In alien conspecific nests, acceptance was maximal for workers aged 1–12 h at introduction (67.7% not attacked, 75.8% adopted) and gradually decreased until the level of nonaggression (after 60 h) and adoption (after 36 h) were not significantly different from 30-day-old workers (5.9% not attacked, 17.6% adopted). In heterospecific nests, acceptance was maximal for workers aged 1–4 h at introduction (34.8% not attacked, 37.0% adopted) but thereafter was not significantly different from 30-day-old workers (5.6% not attacked, 8.3% adopted). In their parental nests, workers were generally accepted regardless of age (4–56 h posteclosion, 95.8% not attacked, 100% adopted); a result that is consistent with previous research on older workers (38–157 days posteclosion). This study demonstrates an acceptance period that is more effective and of longer duration within than between these species but that, under uniform laboratory conditions, is often not necessary for the integration of workers into their parental colonies. Within colonies, acceptance periods might only be important during relatively brief periods in a colony's life history when eclosing workers produce genetically based nestmate recongition cues that are not already represented in the colony and must be learned by colony members (e.g., during early colony growth or following adoption of queens), or when young workers must acquire environmentally based nestmate recognition cues to achieve and maintain acceptability.     

A mechanism for female choice of large males in the treefrog Hyla chrysoscelis

Summary In this study, we combine both field and laboratory experiments to address the effects of female preference for certain call characteristics on a large-male mating advantage in the treefrog H. chrysoscelis. In laboratory-choice experiments, females always chose the call with the lower fundamental frequency when call rate and call intensity were held constant and the difference in frequency between the two calls was 15%. The lower frequency call was preferred by 8 out of 12 females when the difference in fundamental frequency was 7.4%. These results are consistent with field comparisons of the size of unmated males calling within 2 m of a mated male: Male body size was negatively correlated with fundamental frequency and the greater the size difference, the more likely that the larger male mated. In field choice experiments, females preferred males with higher call rates. Since size differences between males used in this experiment averaged only 2.3 mm, we would not expect the fundamental frequency of a male's call to be the best predictor of mating success. Laboratory results demonstrated that call rate could override female preference for the low frequency call over the high frequency call, while intensity could at least dilute this preference. However, individual males in the field varied both call rate and the call intensity as perceived by the female. We suggest that the interaction between call rate, male size and mating success should be studied further through the use of field-choice experiments.     

Underwater oviposition in a damselfly (Odonata: Coenagrionidae) favors male vigilance,and multiple mating by females

Summary Female Enallagma hageni oviposit underwater where they are inaccessible to males. I demonstrate that males guard submerged females rather than perch sites, and are behaviorally distinct from lone males at the water. In contrast to lone males, which always attempt to copulate with females presented to them, guarding males exhibit a conditional latency to remating which corresponds closely to the time required by females to oviposit a complete clutch of eggs. By ovipositing underwater, females decrease the risk that their eggs become exposed. Risks associated with submerged oviposition favor both mate guarding, and multiple, within-clutch matings by females. Both guarding mates and lone males rescue females that float on the water surface as the result of improper resurfacing. Such behavior reduces the mortality risk to females from 0.06 to 0.02 per oviposition bout. By remating between bouts, females benefit from the additional vigilance of lone males, who rescue floating females 1.4 times as often as original mates. A second consequence of multiple mating is an increase in the selective advantage of vigilance by mates. Because receptive females become scarce by early afternoon, whereas male density remains high, a male has little (3%) chance of encountering a second receptive female that day. However, he incurs a 42% risk of losing fertilizations if he abandons a mate. For male E. hageni mate guarding functions in the context of both natural and sexual selection. It insures that a mate lives to lay a complete egg clutch in addition to protecting a male's sperm investment.     

Does the “Teer” vocalization deter prospecting female red-winged blackbirds?

Summary A speaker-occupation experiment was conducted to determine whether female red-winged blackbirds (Agelaius phoeniceus) use the aggressive Teer vocalization to deter conspecific females that are prospecting for opportunities to settle. In each of 14 trials conducted during the breeding seasons of 1987, 1988, and 1989, tape recordings and a pair of loud speakers were used to defend suitable habitat. Observations of female red-winged blackbirds settling on these speaker territories and on adjacent control areas showed that speaker territories were settled significantly earlier than silent control territories. Each of the 14 first settlers on speaker territories included at least one speaker location within the bird's display area, and 9 of the 14 included both speaker locations. Additional observations showed that the rate at which prospecting females visited control and speaker territories, and the number of females eventually settling on them, were not significantly different. Finally, females that settled on speaker territories sang Teers at significantly higher rates than those on control territories. These results demonstrate that the Teer alone was not sufficient to deter prospecting female red-winged blackbirds. On the contrary, Teers given in the absence of an actual resident may have attracted settlers. It is suggested that the aggressive Teer vocalization might function to establish dominance relationships among the females sharing a polygynous male's territory.     

Comparisons of the feeding activity and diets of male and female cunners Tautogolabrus adspersus (Pisces: Labridae)

The diel distribution of feeding activity during the spawning season was compared for territorial male and female cunners Tautogolabrus adspersus in Conception Bay, Newfoundland, Canada between June and August, 1978. Territorial males feed significantly less often than females and concentrate their feeding activities in the morning, whereas females feed as frequently in the afternoon as in the morning. Based on the contents of alimentary tracts, territorial males, non-territorial males, and females have different diets. Since these three groups share the same habitat, the observed dietary differences probably reflect differences in their foraging behaviour. The relevance of these findings to the concept of time minimizer and energy maximizer is discussed.     

Female aggression and male peace-keeping in a cichlid fish harem: conflict between and within the sexes in Lamprologus ocellatus

Summary Conflicts of interest within and between the sexes are important processes leading to variability in mating systems. The behavioral interactions mediating conflict are little documented. We studied pairs and harems of the snail-shell inhabiting cichlid fish Lamprologus ocellatus in the laboratory. Due to their larger size, males controlled the resource that limited breeding: snail shells. Males were able to choose among females ready to spawn. Females were only accepted if they produced a clutch within a few days of settling. When several females attempted to settle simultaneously the larger female settled first. Females were least aggressive when guarding eggs. Secondary females were more likely to settle when the primary female was guarding eggs. In established harems females continued to be aggressive against each other. The male intervened in about 80% of female aggressive interactions. Male intervention activity correlated with the frequency of aggression among the females in his harem. The male usually attacked the aggressor and chased her back to her own snail shell. When a male was removed from his harem, aggression between females increased immediately and usually the secondary female was expelled by the primary female within a few days. Time to harem break-up was shorter the more mobile the primary females' young were and did not correlate with the size difference between harem females. Male L. ocellatus interfere actively in female conflict and keep the harem together against female interests. Female conflict presumably relates to the cost of sharing male parental investment and to the potential of predation by another female's large juveniles on a female's own small juveniles.Correspondence to: F. Trillmich     

Replacement males in the western bluebird: opportunity for paternity, chick-feeding rules, and fitness consequences of male parental care

In western bluebirds (Sialia mexicana), most pairs remain together for life and share equally in post-hatching parental care. We removed resident males of socially monogamous pairs during laying and after clutch completion to examine chick-feeding rules used by replacement males and current and future fitness consequences of paternal care. Replacement males were not infanticidal and 7 (47%) fed nestlings. Feeding replacement males and the females they joined fed at rates similar to controls. Females without a feeding replacement male compensated by feeding more themselves so that overall feeding rates were not compromised, but they reduced their brooding time. Unlike assisted females, unassisted females exhibited reduced nesting success and their 14-day-old chicks weighed less than controls. Field metabolic rates of unassisted females were 17% higher than those of control females, but the difference was not statistically significant. Older females were better able to raise young without the male's help than were yearling females. Female condition was not affected by male provisioning and unassisted females were as likely to survive to the next breeding season as assisted females. We found no future benefits of provisioning by replacement males; those that fed were no more likely to breed with the female on her subsequent attempt than were males that did not feed, and subsequent clutch sizes were not reduced for females rearing young without the male's help. These experiments suggest that male parental care increases nesting success in western bluebirds and that replacement males use an all-or-none rule to determine whether or not to feed chicks: if they are present during the fertile period they feed at typical rates; if they are not, they usually do not feed at all. Because chick-feeding by males is tied to opportunity for paternity, influences success in the current nest, and does not affect the male's future breeding success, it appears to be parental rather than mating effort. Received: 8 May 1998 / Accepted after revision: 23 July 1998     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Unattractiveness of mated females to males in the parasitoid wasp Spalangia endius

Despite common stereotypes, males are not always indiscriminate and eager when it comes to mating. In the parasitoid wasp Spalangia endius, the initial response of males to females was almost always one of apparent excitement; however, this was followed by a clear preference for virgin females over mated females in both no-choice and choice situations. The no-choice data were collected from videotapes of male-female pairs of all possible combinations of mated and virgin individuals. Neither female nor male mating status had a significant effect on likelihood of, or time until, contact or male courtship fanning. However, a males first retreat was sooner when the female was mated than when she was virgin; mated males exhibited their first retreat sooner than did virgin males; and mated females were less likely to be mounted than were virgin females. In addition to the videotapes, male-choice experiments were performed. When given a choice of a virgin and a mated female, both virgin and mated males were more likely to mount and copulate with the virgin. The difference in response to virgin versus mated females seemed to be less in virgin males than in mated males, perhaps due to virgin males greater eagerness to mate: when a virgin male and a mated male were presented with a dead virgin female, the virgin male was usually the first to respond to the female. That males preferentially retreated from and avoided mounting mated females appears to be adaptive given that mated females rarely copulated.Communicated by N. Wedell     

Sex ratio,monogamy and breeding success in the Midas cichlid (Cichlasoma citrinellum)

Summary The Midas cichlid (Cichlasoma citrinellum) is a large, aggressive monogamous fish from the Great Lakes of Nicaragua. It lives in an ecosystem where breeding sites are at a premium and where breeding success can be low. I tested the strength of the pair bond by manipulating the sex ratios in captive groups, using ratios of 1:1, 2:1, 4:1, 1:2 and 1:4 (:). No polygamy was seen at any of the ratios. Departure from a ratio of 1:1 decreased breeding success for those pairs that formed. Both sexes appeared to enforce the pair bond but did so in different ways: females attacked males and females equally but males attacked other males significantly more than they did females. These results, coupled with field observations, indicate the Midas cichlid is obligated to monogamy by the demands of competition for breeding sites, the need for two parents to protect the fry and by energetic constraints.     

Ammonium enhancement of dark carbon fixation and nitrogen limitation in zooxanthellae symbiotic with the reef coralsMadracis mirabilis andMontastrea annularis

The nutrient status (limitation vs sufficiency) of dinoflagellates (zooxanthellae) symbiotic with reef corals in Bermuda was assessed in 1989 and 1990 by measuring the enhancement of dark carbon fixation with 20 M ammonium by isolated symbionts. A colony ofMadracis mirabilis was kept in the laboratory and fed daily or starved for one month. Symbionts from fed portions of the colony had ammonium-enhancement ratios (NH _4dark ⁺ ; SW_dark;SW=seawater without added ammonium) similar to those of the original field population (1.2 to 1.3). Ammonium-enhancement ratios increased with starvation of the host (x1.7) as did values forV _D:V _L [(ammonium dark rate-seawater dark rate): light rate in seawater]. Both parameters indicated decreasing nitrogen sufficiency of the algae when the host was not fed, but starvation appeared to affect these algae less than symbionts of sea anemones. Field samples of zooxanthellae fromM. mirabilis (Three Hill Shoals and Bailey's Bay Flats) yielded results similar to those for fed corals, but those taken from Bailey's Bay Flats in May 1990 yielded exceptionally high values for enhancement (>3) andV _D:V _L indicating pronounced nitrogen limitation at the time of sampling. We sampled zooxanthellae from populations ofMontastrea annularis at 8 m (Three Hill Shoals) and 24 m (Soldier's Point) depths. Enhancement andV _D:V _L values for zooxanthellae from the 8 m corals were density-dependent: symbionts from corals with normal symbiont densities displayed the most nitrogen limitation (enhancement values=1.4 to 2.0), while those from bleached corals with lower density exhibited enhancement andV _D:V _L values typical of nitrogen-sufficient algae. Symbionts isolated from the 25 m corals yielded the highest values, and appeared to exhibit the least nitrogen-sufficiency for this species.     

Effect of intraspecific density on life history traits and population growth rate of Neanthes arenaceodentata (Polychaeta: Nereidae) in the laboratory

The effects of intraspecific density on life history traits and population dynamics of the nereid polychaete Neanthes arenaceodentata Moore were assessed in a laboratory experiment. Survival, growth, and fecundity were measured for one generation of worms at densities of 40, 80 and 160 worms per 840 cm² (1x, 2x, and 4x treatments, respectively). Density did not affect size (prior to pairing), percentage of worms paired, time to pairing, or size of mature paired males. Density did have a significant negative effect on survival, size of mature paired females, time to spawning, percentage of females that reproduced, and number of eggs per reproducing female. As density increased, mean survival was 90.0, 80.8 and 74.0%; mean size of mature females was 52.2, 49.2 and 48.1 segments; mean time to spawning was 100.6, 102.4 and 109.4 d; and mean fecundity was 881, 622, and 598 eggs per female, for 1x, 2x and 4x treatments, respectively. Increased density reduced the potential population growth rate, ; for a given rate of larval survivorship, was lower in the 2x and 4x treatments than the 1x treatment. Analysis of sensitivity of to changes in survivorship indicated that population growth rate at the highest density was sensitive to both changes in larval survivorship and the probability of producing a successful brood, although at low densities, was sensitive only to changes in larval survivorship. We attribute these density effects to aggressive bahavioral interactions between the worms, primarily the adults. This experiment identifies key life history traits that could be measured in future experiments to test population level responses of N. arenaceodentata populations to pollutants, both in the laboratory and in the field.Contribution No. 820 of the U.S. Environmental Protection Agency, Environmental Research Laboratory and Contribution No. 189 of the Marine Sciences Institute, University of Connecticut     

Symmetric fights as a measure of escalation potential in a symbiotic,territorial snapping shrimp

Summary The shrimp Alpheus armatus territorially defends the sea anemone it occupies, using as a weapon its large, specially modified snapping claw. This behavior was studied in experimental contests which were symmetric (matched individuals) with respect to sex, size, and residency. Characteristics of these contests were compared for two size-classes of male and female shrimp.There were no significant differences between these groups of shrimp in the number of bouts required to establish dominance or in the number of snaps exchanged. Large females had shorter contests than either small or large males, and losers of contests between large females were injured more frequently and more severely.This distinctiveness of large female contests could be interpreted as evidence that (i) controlling anemones is more important for large females, (ii) injuries are less important for large females, or (iii) large females lose the ability to assess one another because their contests are less frequent.If injuries are an accurate measure of the most important costs associated with fighting, then these results indicate that short contests are not necessarily the least costly, and that females can be more aggressive than males, as measured by escalation potential, in sexually selected species.     

Lack of innate preference for morph and species identity in mate-searching <Emphasis Type="Italic">Enallagma</Emphasis> damselflies

Insect mate recognition is often viewed as stereotypic, innate, and species-specific. However, male damselflies can learn to identify female-specific color morphs as potential mates. A suite of male mimicry hypotheses assume that heteromorphic females, which differ from males in color pattern, are more easily recognized as “female” and thus lack the inherent, anti-harassment advantage that the more male-like signal provides for andromorphs. Using two measures of male preference, we investigated whether naïve males have a preexisting sensory bias for a given morph color in Enallagma civile, a species that appeared to exhibit extreme plasticity in morph expression across generations within a breeding season. E. civile males raised in the absence of females exhibited no preference for either morph, whereas males raised with one female type exhibited a learned sensory bias for that morph. Male Enallagma also lacked a bias toward conspecific females over a congeneric sister species. In a naturally naïve population of Enallagma ebrium, males reacted sexually to both morphs of Enallagma hageni as often as they did to conspecific females, whose thoracic spectra were nearly identical with those of E. hageni. Moreover, despite the similar thoracic spectra of males and andromorphs, both of which reflected UV, males rarely reacted sexually to other males. Our results falsified implicit assumptions of male mimicry hypotheses, supported learned mate recognition, and suggested a scenario for speciation via sexual conflict.     

Does promiscuous mating preclude female choice? Female sexual strategies in chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d’Ivoire

According to sexual selection theory, females should selectively mate with high-quality males to enhance offspring survival and maximize reproductive success. Yet, chimpanzee (Pan troglodytes) females are known to mate promiscuously. Although there is substantial rationale for a promiscuous mating strategy, there is also a strong expectation that females should be selective, and the question arises as to whether promiscuity precludes female choice. The aims of this study are to: (1) compare wild female chimpanzee sexual strategies throughout estrus, and (2) determine whether females exhibit mate preferences for particular males. Over 2,600 h of data were collected on two habituated chimpanzee (Pan troglodytes verus) communities in the Taï National Park, Côte dIvoire. Female mate preferences were measured by quantifying proceptivity and receptivity toward males. Results indicate that all females exhibited proceptivity and resistance to male solicitations, but that there was substantial variation in their magnitudes within and among females. Female proceptivity rates were lower and resistance rates were higher in the periovulatory period (POP) when conception is most likely. Females were more selective during POP, and more promiscuous outside of POP, suggesting that females may follow a mixed reproductive strategy, being selective when conception is likely and more promiscuous when conception is unlikely. Results from this study emphasize the importance of considering the fertility window when determining female mate preferences, and of examining female behavior in POP and non-POP phases separately when evaluating hypotheses for multi-male mating.Communicated by C. Nunn     

Functional aspects of the pair bond in winter in Bewick's swans (Cygnus columbianus bewickii)

Summary The benefits to partners of monogamous pairs of maintaining continual spatial proximity in the non-breeding season were studied in Bewick's swans wintering in Norfolk, UK.When separated from their mates, females were less successful in aggressive encounters, were threatened more frequently and spent less time feeding than when close to them. Males also suffered reduced success in encounters and a higher frequency of threats by other flock members, though the effect was less pronounced than for females. This sex difference in effect of separation may be associated with the greater weight of males and the fact that success in encounters is related to weight.Partners appeared to assist each other by joining in aggressive encounters, as well as by inhibiting other birds from threatening their mates. However, the precise manner in which the female assisted the male is still obscure, since the highest intensity aggressive encounters — physical fights — involved only male partners. It is suggested that the male may fight harder in his mate's presence.Proximity of partners varied with situation and between different pairs. Partners maintained greater proximity in dense flocks than in dispersed flocks and showed a tendency to stay closer when feeding on winter wheat than on waste potatoes. Partners where the male was high-ranking spent more time together than those where the male was low-ranking.     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.     

Reproductive consequences of male body mass and aggressiveness depend on females’ behavioral types

Relatively few investigations explicitly test for concordant versus conflicting selection pressures from intrasexual versus intersexual selection. Here, we examine the effects of male body mass and behavioral type (BT) on reproductive success in the spider Anelosimus studiosus, with emphasis placed on the potential interaction between intrasexual and intersexual selection influences. Female A. studiosus exhibit either an aggressive-active or docile-passive BT, both of which co-occur in multifemale colonies. Males, in contrast, exhibit a more continuous distribution of behavioral tendencies. We investigated the male traits favored by females in five trial types: one docile female, one aggressive female, four docile females, four aggressive females, and two docile and two aggressive females. Male reproductive success was estimated by the number eggs produced by females following staged mating trials. In previous work, it was established that large aggressive males are favored in male–male contests, an intrasexual effect. However, large aggressive males were not universally favored here. We failed to detect an effect of male body mass or aggressiveness on reproductive success in trials with all docile females; however, in situations involving aggressive females, large aggressive males experienced diminished reproductive success relative to small docile males. Large, aggressive males were also more likely to be attacked and killed by aggressive females in the first 20 min of staged encounters and were more likely to be found dead after 72 h of unobserved interactions. Taken together, our data suggest that the reproductive consequences of male traits differ based on (1) the aspect of sexual selection being considered (intrasexual versus intersexual) and (2) the BT of their prospective mates: large aggressive males enjoy advantages in intrasexual selection and when courting docile females and small docile males experience reduced risk of cannibalism and increased reproductive success with aggressive females.