Sexually selected females in the monogamous Western Australian seahorse

Studies of sexual selection in monogamous species have hitherto focused on sexual selection among males. Here, we provide empirical documentation that sexual selection can also act strongly on females in a natural population with a monogamous mating system. In our field-based genetic study of the monogamous Western Australian seahorse, Hippocampus subelongatus, sexual selection differentials and gradients show that females are under stronger sexual selection than males: mated females are larger than unmated ones, whereas mated and unmated males do not differ in size. In addition, the opportunity for sexual selection (variance in mating success divided by its mean squared) for females is almost three times that for males. These results, which seem to be generated by a combination of a male preference for larger females and a female-biased adult sex ratio, indicate that substantial sexual selection on females is a potentially important but under-appreciated evolutionary phenomenon in monogamous species.     

Sexual selection in the monogamous barn swallow (Hirundo rustica). II. Mechanisms of sexual selection

Mechanisms of sexual selection in the monogamous, sexually dimorphic barn swallow (Hirundo rustica) were studied during a seven-year period. First, the sex ratio of reproducing adults was male-biased, and mated males had significantly longer tail ornaments than unmated males. Secondly, some of the unmated individuals later committed infanticide and became mated with the mother of the killed brood. Fathers of killed broods had significantly shorter tails than other males, and there was a tendency for infanticidal males to have longer tail ornaments than other unmated males. Thirdly, long-tailed male barn swallows were more successful in acquiring extra-pair copulations than other males, and females involved in extra-pair copulations, as compared to females not involved in such copulations, had mates with shorter tail ornaments. Fourthly, male barn swallows having long tails as compared to short-tailed males acquired mates in better body condition. Females mated to long-tailed males reproduced earlier, laid more eggs and were more likely to have two clutches than were females mated to short-tailed males. Finally, females mated to long-tailed males put more effort into reproduction than did other females, as evidenced by their relatively larger contribution to feeding of offspring. Thus, at least five different components of sexual selection affected male reproductive success. Selection arising from differential success during extra-pair copulations, differential reproductive success and differential male reproductive effort thus accounted for most of the selection on tail ornaments in male barn swallows.     

Male mate choice in the chameleon grasshopper (Kosciuscola tristis)

In many species, males can increase their fitness by mating with the highest quality females. Female quality can be indicated by cues, such as body size, age and mating status. In the alpine grasshopper Kosciuscola tristis, males can be found riding on subadult females early in the season, and as the season progresses, males engage in fights over ovipositing females. These observations suggest that males may be competing for females that are either unmated (early season) or sperm‐depleted (late season). We thus hypothesised that male K. tristis may be choosy in relation to female mating status, and specifically, we predicted that males prefer females that are unmated. We conducted behavioural experiments in which males were given the choice of two females, one mated and one unmated. Contrary to our prediction, males did not mate preferentially with unmated females. However, copulation duration with unmated females was, on average, 24 times the length of copulation with mated females. While female K. tristis can reject mates, we did not observe any evidence of overt female choice during our trials. Females may gain additional benefits from mating multiply and may therefore not readily reject males. While our experiment cannot definitively disentangle female from male control over copulation duration, we suggest that males choose to invest more time in copula with unmated females, perhaps for paternity assurance, and that male mate assessment occurs during copulation rather than beforehand.     

Infanticide and effectiveness of pup protection in bank voles: does the mother recognise a killer?

Infanticide, the killing of conspecific young, has been documented in numerous species of mammals, especially rodents. In that infanticide is costly to the victim mother, natural selection should favour counter-strategies by females to protect their pups. We studied the frequency of infanticide by male and female bank voles (Clethrionomys glareolus) and the effectiveness of dams in recognising and deterring infanticide. In trials in which unprotected pups were exposed to voles of both sexes, one third of male and female intruders killed pups. When mothers were present at the nest site, not a single female and only 2 of 25 males were able to commit infanticide. Females acted aggressively towards all intruders and hence did not discern between infanticidal and non-infanticidal males and females. Aggression of dams against any intruder indicates that all strange individuals near the nest site of a territorial species form an equal threat to pups. However, the presence of the aggressive dam is sufficient to deter most intruders from harming offspring by keeping them at a distance from the nest site. Heat run before mating, in a species with postpartum oestrus and vulnerable pups in the nest, also might lure potential infanticidal males away from the nest site and obscure the nest location. Received in revised form: 17 October 2001 Electronic Publication     

Monogamy and sex role reversal in the pipefish Corythoichthys haematopterus

We observed the mating pattern and social behaviour of the pipefish Corythoichthys haematopterus in temperate waters of Japan during three successive breeding seasons. Males cared for a clutch in their brood pouch for 9-19 days until hatching and had several broods in the season with nonbrooding intervals of only 1 or 2 days. The population sex ratio was female biased and some females were always excluded from reproduction. Although males were sometimes courted by unmated females together with their regular partners, they always mated with the latter. The pair bond was maintained until the next season if both members survived. When males lost their partners, they remated with neighbouring unmated females within a few days. In contrast, widowed females remained unmated for a long time. Females had larger home ranges and were more active in courtship displays than males. This pipefish provides the first example of sex role reversal among monogamous syngnathid fish. We suggest that mate guarding by females is a primary proximate factor for maintenance of monogamy in this fish. Copyright 2001 The Association for the Study of Animal Behaviour.     

Proximate and ultimate factors explaining floating behavior in long-lived birds

Avian populations are composed of reproductive individuals coexisting in space and time with young, non-breeding conspecifics or “floaters”. Despite the fact that first breeding can be delayed for years and can exert profound effects on fitness, many aspects of the behavior, ecology and physiology of young floaters remain poorly understood. By means of combining endocrine, behavioral and life history information from a population of black kites (Milvus migrans) monitored long-term, we tested several hypotheses suggesting endocrine function as a determinant of floating status. Sexual function in non-breeding males, estimated through determination of systemic testosterone and progesterone levels, was similar to that in reproductively active conspecifics. Floating females, on the contrary, displayed an endocrine pattern of circulating estrogens and progesterone that was parallel in timing but reduced in magnitude as compared to breeders. Our results suggest that floaters are not physiologically constrained to reproduce, but the cost-benefit balance of attaining complete gonadal function is sexually dependent. While young, unmated males could increase their breeding prospects by attaining sexual maturity regardless of their social environment, natural selection would favor females relying on social cues to mature. Consistent with the sexual roles of socially monogamous species, gonadal recrudescence and testosterone production would allow unmated males to access breeding resources (e.g. through male-male competition and extra-pair fertilizations). Unmated females, on the contrary, would reduce physiological costs by means of delaying ovarian maturation until establishing pair bonds with a male providing access to breeding resources.     

Infant Mortality Risk and Paternity Certainty Are Associated with Postnatal Maternal Behavior toward Adult Male Mountain Gorillas (Gorilla beringei beringei)

Sexually selected infanticide is an important source of infant mortality in many mammalian species. In species with long-term male-female associations, females may benefit from male protection against infanticidal outsiders. We tested whether mountain gorilla (Gorilla beringei beringei) mothers in single and multi-male groups monitored by the Dian Fossey Gorilla Fund’s Karisoke Research Center actively facilitated interactions between their infants and a potentially protective male. We also evaluated the criteria mothers in multi-male groups used to choose a preferred male social partner. In single male groups, where infanticide risk and paternity certainty are high, females with infants <1 year old spent more time near and affiliated more with males than females without young infants. In multi-male groups, where infanticide rates and paternity certainty are lower, mothers with new infants exhibited few behavioral changes toward males. The sole notable change was that females with young infants proportionally increased their time near males they previously spent little time near when compared to males they had previously preferred, perhaps to encourage paternity uncertainty and deter aggression. Rank was a much better predictor of females’ social partner choice than paternity. Older infants (2–3 years) in multi-male groups mirrored their mothers’ preferences for individual male social partners; 89% spent the most time in close proximity to the male their mother had spent the most time near when they were <1 year old. Observed discrepancies between female behavior in single and multi-male groups likely reflect different levels of postpartum intersexual conflict; in groups where paternity certainty and infanticide risk are both high, male-female interests align and females behave accordingly. This highlights the importance of considering individual and group-level variation when evaluating intersexual conflict across the reproductive cycle.     

Infanticide in great reed warblers: secondary females destroy eggs of primary females

In 1994–1995 artificial nests with attached model eggs were put into territories that were known to have been occupied by male great reed warblers,Acrocephalus arundinaceusin previous years. Because the eggs were made of soft plasticine, predators left peckmarks in them and this enabled us to identify predators by comparing peckmarks with reference marks made by various species. Previous field data had suggested that infanticidal behaviour existed in our study population, as nests of primary females suffered a three times higher rate of nest loss during the egg-laying period than nests of secondary and monogamous females. The presence of infanticide was supported by the experiment. Small peckmarks resembling those of a great reed warbler occurred almost exclusively in territories occupied by great reed warblers, in particular when a new female settled in the territory. The newly settled females built nests closer to depredated than non-depredated nests. That small peckmarks occurred when new females settled strongly suggests that it is secondary female great reed warblers that commit infanticide on eggs of primary females. Females of low harem rank are expected to gain from infanticidal behaviour because a low ranked female gets a higher proportion of male parental investment when the nest of the primary female fails.     

Mating systems and territory in Lapwings Vanellus vanellus

In a population of Lapwings Vanellus vanellus studied for 5 years near Bergen, southwester Norway, 23–41% of the males were polygynous, having two and sometimes three mates, whereas 50–77% were monogamous and 0–21% remained unmated. Bachelors held territories in or immediately adjacent to the study area each of the years. As predicted from the Polygyny Threshold model, primary females generally laid eggs earlier than monogamous females, but the overlap in laying dates was substantial between these two classes. The number of females and start of breeding on the territories were analysed in relation to nine variables pertaining to safety from predation and to food. Of these, territory size correlated most consistently with number of females (positively) and the laying of first egg (negatively). Numbers of females observed were not significantly different from numbers expected from territory size in 4 out of 5 years; however, earlier start of breeding on large territories indicated that females did not settle according to the Neutral Mate Choice model. The Female-biased Sex Ratio hypothesis was refuted, unmated territorial males being available throughout the breeding seasons.     

Rapid temporal change in frequency of infanticide in a passerine bird associated with change in population density and body condition

Sexually selected infanticide, whereby unmated males obtaina mate by killing the dependent offspring of an already matedfemale, is a common alternative reproductive strategy in manyanimals. I estimated the frequency of infanticide in a populationof barn swallows, Hirundo rustica, during the period 1977–2002.Population size decreased by more than a factor 10 during thisperiod, and this decrease was associated with an increase inmortality, selecting for adults with better body condition.Density-dependent effects acted on infanticide through the relationshipbetween the relative number of unmated males and populationsize. Because males in prime condition are better able to defendtheir nests against infanticidal males, the frequency of infanticidedecreased as mean tail length and mean body condition of malesincreased during the study period. Therefore, a rapid decreasein population size, a decrease in the abundance of unmated males,and a concomitant increase in body condition have changed theimportance of infanticide from being a major cause of mortalityaccounting for more than 25% of all nestling mortality to beingalmost completely absent during a period of 25 years.     

Territory and monogamy among Kloss' gibbons (Hylobates klossii) in Siberut Island, Indonesia.

Behavior of Kloss' gibbons was studied from July 1 to October 7, 1972 in Siberut Island, off the coast of western Sumatra, Indonesia. Reproductive groups are monogamous families with a mean family size of 3.4 individuals (n = 11 families). Such families occupy territories averaging 6.7 ha. Adults defend their territories only against members of the same sex. This intrasexual defense of territory maintains the monogamous mating system. Other social units were unmated resident females, unmated resident males, floating males, and a courting pair. Males establish territories before mating, perhaps with help from their parents. Males guard their families against predators. Females lead progression through the territory. Subadults remain peripheral to their families but other family members tend to remain within 10 m of one another.     

Evolutionary significance of promiscuity in an aphidophagous ladybird, Propylea dissecta (Coleoptera: Coccinellidae)

The short- and long-term consequences of monogamy and promiscuity in an aphidophagous ladybird, Propylea dissecta (Mulsant) in terms of reproductive performance and offspring fitness were investigated. Promiscuous females were more fecund and laid more viable eggs than monogamous ones. Amongst promiscuous females, those mated with multiple males (freedom for mate choice) had significantly higher reproductive output than those mated daily with a new unmated male. The increased reproductive output of females mated with an unmated male each day in comparison to monogamous ones might possibly be a result of their increased ejaculate size. The further increase in reproductive output, when a female was allowed to choose a mate, may be because of repeated preferential mating with the fitter males. Results on long-term consequences of promiscuity revealed faster development and increased survival in offspring of promiscuous females at 25, 27 and 30 degrees C. The percent adult emergence was more strongly influenced than development by the mating regime. There were no statistically significant differences in survival of offspring of promiscuous females (mate choice) at the three temperatures, while the percent survival of offspring of monogamous females differed significantly. Offspring of promiscuous females (mate choice) were fittest in terms of development and survival, which indicates their probable better genetic constitution and subsequently that of chosen mates. Thus, promiscuity has both short- and long-term benefits and is advantageous to ladybirds. The presence of long-term benefits in terms of fitter offspring probably indicates the evolutionary rationale behind the prevalence of promiscuity in ladybirds.     

Sexual signalling by females: do unmated females increase their signalling effort?

Theory predicts that females should invest least in mate searching when young, but increase their effort with age if they remain unmated. Few studies have examined variation in female sexual signalling. Female Dawson''s burrowing bees (Amegilla dawsoni) search for males by signalling their receptivity on emergence, but many leave the emergence site unmated and must attract males at feeding sites. Female bees prevented from mating on emergence had more extreme versions of cuticular hydrocarbon profiles that make them attractive to males, lending empirical evidence of adaptive shifts in female mating effort.     

Mating is associated with immediate changes of the hydrocarbon profile of Leptothorax gredleri ant queens

While sexual communication is often characterized by attempted manipulation, both sexes agree about females reliably signalling their receptivity. Female sexuals of the ant Leptothorax gredleri quickly became unattractive to males after their first copulation. This loss of attractiveness coincided with almost immediate changes in their cuticular hydrocarbon (CHC) profiles. Already 30 min after mating, the CHC profiles of female sexuals had significantly lower relative amounts of branched alkanes and higher amounts of linear alkanes than those of unmated and freshly mated female sexuals. Discriminant analysis did not distinguish between the profiles of freshly mated and unmated female sexuals, suggesting that the extremely rapid modification of CHC profiles is not caused by males marking females with anti-aphrodisiac CHCs. Instead, the new profile is produced by the female sexuals themselves. In addition to making them unattractive to males, this change may also help mated female sexuals when seeking adoption into established colonies.     

New Case of Intragroup Infanticide in the Chimpanzees of Gombe National Park

Researchers have reported a total of 31 infanticides in 4 different chimpanzee (Pan troglodytes) populations. Though infanticide is infrequent, low reproductive rates of females likely make it a strong selective pressure in the species. We report a new incident of intragroup infanticide in Gombe National Park, Tanzania, in which a community male attacked a 3.5-yr-old male. We then consider the infanticide in terms of adaptive and nonadaptive explanations for infanticide including the social pathology, by-product of male aggression, nutritive benefits, resource competition, and sexual selection hypotheses. The incident reported here is not well explained by any of them. While the infanticide is puzzling in terms of ultimate explanations for infanticide, it provides a good context in which to consider proximate mechanisms for offspring recognition. The incident provides some evidence that males may use their mating history with the mother to assess paternity likelihood.     

Polygyny in the pied flycatcher,Ficedula hypoleuca: a test of the deception hypothesis

The deception hypothesis has been proposed as an explanation for polygyny in the pied flycatcher. According to this hypothesis, already-mated males hide their mating status with polyterritorial behaviour and thereby increase their chances of obtaining a second mate. In a study area at Oslo, Norway, secondary females raised 84% as many fledglings as did concurrent monogamous and primary females. The unmated males sang most of the time near their nest site, whereas the already-mated males frequently disrupted singing for longer periods in their secondary territories to visit their primary nest; such visits to the primary nest occurred both before and after the time of their second mating. The behaviour of the males suggests that deception of females is not an evolutionarily stable strategy, as an observant female would soon discern the male's status. Another difficulty with the deception hypothesis is that secondary females laid larger clutches than primary females. The number of young fledged from secondary nests was not dependent on the distance to the primary nest. The deception hypothesis was not supported by the data, and the reduced reproductive success of secondary females may be explained by the cost of searching for a mate.     

Monogamy does not last long in Pontonia mexicana, a symbiotic shrimp of the amber pen-shell Pinna carnea from the southeastern Caribbean Sea

Theory suggests that symbiotic species should be monogamous and form long-lasting heterosexual pairs in/on hosts when inhabiting scarce and small hosts in environments where predation risk away from hosts is high (e.g., tropical subtidal). This prediction was tested with Pontonia mexicana which inhabits the relatively small and scarce amber pen-shell Pinna carnea in the tropical Caribbean. In agreement with theory, P. mexicana were found dwelling as heterosexual pairs in the mantle cavity of pen-shell individuals with more frequency than expected by chance alone. However, additional observations suggested that male-female pairing of P. mexicana does not last long. First, males paired with females that were close to molt and become sexually receptive more frequently than expected by chance alone. In monogamous species in which pairing appears to be long lasting, males occur with females in the same host, independent of the reproductive condition of the female. Second, the body size of paired shrimps was poorly correlated and the relationship between host and shrimp body size was weak. If males and females of P. mexicana were staying within host individuals for long periods of time, a tight correlation between host and shrimp size and between males and females in a pair would have been found. Lastly, sexual dimorphism in terms of cheliped size was evident; males invested considerably more resources to this body structure compared to females. In monogamous species in which pairing appears to be long-term, sexual dimorphism is low or absent given that sexual selection is weak in this mating system. Overall, our data suggests that monogamy in P. mexicana does not last long and that males switch among host individuals in search of receptive females. Manipulative experiments are necessary to understand the conditions favoring short- and long-term monogamy in symbiotic crustaceans.     

Potentially infanticidal behavior in the Amazon river dolphin (<Emphasis Type="Italic">Inia geoffrensis</Emphasis>)

Infanticide by males is a phenomenon common in species in which the reproductive output of large numbers of females can be monopolized by a small number of males. It is thought to increase a male’s fitness, at the expense of the fitness of the infant’s parents, by bringing females into season more quickly. Infanticide by males has been recorded in just three cetacean species. We report aggressive behavior suggestive of infanticide in a fourth, the Amazon river dolphin (Inia geoffrensis). We observed and photographed a series of attacks on a neonate Amazon river dolphin by a large male, with apparent protective behavior by the mother. Although infanticide was not confirmed, the forceful, aggressive behaviors were highly suggestive of infanticidal behavior and represent another important data point for comparative studies of infanticide in mammals. Amazon river dolphins may have a polygynous, polyandrous, or promiscuous mating system, the latter two of which are not the norm in species in which the reproductive output of large numbers of females are monopolized by a small number of males. However, sexual dimorphism, high rates of aggression by males, socio-sexual object-carrying displays by males, and a long interbirth interval suggest that successful male Amazon river dolphins may be able to monopolize a large proportion of mating opportunities, and it is plausible that male dolphins can improve their reproductive success by bringing females into estrous sooner by killing the offspring of other males.     

The Role of Female Behavior in Maintaining Monogamy of a Coral-Reef Filefish

The corallivorous filefish Oxymonacanthus longirostris is a predominantly monogamous species without parental care. Polygyny reduces female fitness because monogamous females benefit from male assistance with feeding–territory defense that secures both a feeding area and time to feed. I studied the strategy of females to prevent polygyny, with the help of field observations and manipulative field experiments. Monogamous females always behaved aggressively towards intruding floater females. When a caged male or female was placed within the territory of a monogamous pair, the resident female was more likely to pay attention to the caged female than to the caged male. Experimentally transplanted floater females succeeded in settling as secondary females in the monogamous territory more frequently when the resident female was caged. These results suggest that monogamous females behave aggressively towards floater females to defend their mating status; such aggressive behavior is successful in preventing polygyny. Furthermore, the removal experiment showed that polygynous (especially secondary) females moved to territories of widowed males located near them more frequently than monogamous females did. This outcome indicates that polygynous females can change their mating status and escape polygyny by moving to another territory if the opportunity occurs. Thus, female behavior can play an important role in the maintenance of monogamy and in lowering the level of polygyny within populations of this species.     

Polyandry enhances offspring survival in an infanticidal species

The adaptive significance of polyandry is an intensely debated subject in sexual selection. For species with male infanticidal behaviour, it has been hypothesized that polyandry evolved as female counterstrategy to offspring loss: by mating with multiple males, females may conceal paternity and so prevent males from killing putative offspring. Here we present, to our knowledge, the first empirical test of this hypothesis in a combined laboratory and field study, and show that multiple mating seems to reduce the risk of infanticide in female bank voles Myodes glareolus. Our findings thus indicate that females of species with non-resource based mating systems, in which males provide nothing but sperm, but commit infanticide, can gain non-genetic fitness benefits from polyandry.