Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Sexual selection and reproductive careers in mandrills (Mandrillus sphinx)

In sexually dimorphic, polygynous species, where males provide little parental care and competition between males for access to fertile females is high, sexual selection theory predicts sex differences in age-specific reproductive output and mortality profiles, and greater variance in lifetime reproductive success in males than in females. We examined age-specific reproductive output, mortality patterns and the extent and causes of variation in reproductive success for a semi-free-ranging colony of mandrills (Mandrillus sphinx, Cercopithecidae) in Franceville, Gabon, using long-term (20 year) demographic records and microsatellite parentage analysis. Although differences in the demography and feeding ecology of this closed, provisioned colony, in comparison with wild mandrills, limit interpretation of our results, sex differences in reproductive output and mortality showed the patterns predicted by sexual selection theory. Mortality was higher in males than in females after sexual maturity, and lifespan was significantly shorter in males (mean 14 year) than in females (>22 year). Age at first reproduction was significantly earlier in females (mean 4.2 year) than in males (11.6 year), and male reproductive output declined earlier. All females of breeding age produced offspring; while only 17 of 53 sexually mature males (32%) sired. Males sired a maximum of 41 offspring, versus 17 in females, and variance in male reproductive output was significantly greater than in females at all ages. The most important influence on variation in lifetime reproductive output in both sexes was joint variation between length of the breeding period and reproductive rate, due to lower reproductive rates in younger animals. Finally, social rank significantly influenced reproductive output in both sexes: high-ranking females began their reproductive careers earlier and had a higher subsequent reproductive rate than low-ranking females, while males that achieved top rank during their career sired far more offspring than males that did not.Electronic Supplementary Material Supplementary material is available for this article at     

Sexual selection and spring arrival times of red-necked and Wilson's phalaropes

Summary Seasonal changes in sex ratios during a 4-year study of red-necked phalaropes and a 3-year study of Wilson's phalaropes showed that females preceded males to breeding and/or courtship areas. The degree to which females preceded males may have been constrained by harsh weather, as the arrival of red-necked phalaropes was synchronous in 1983, when spring was unusually late. Neither sex defended territories; females competed vigorously for direct access to males. These findings show that selection for sexually asynchronous arrival need not act through territoriality. We interpret early female arrival as an adaptation for obtaining mates and coclude that sexual selection may be an important determinant of arrival times in mate defense social systems.     

Hierarchical dominance structure in reintroduced California condors: correlates, consequences, and dynamics

Populations of reintroduced California condors (Gymnogyps californianus) develop complex social structures and dynamics to maintain stable group cohesion, and birds that do not successfully integrate into group hierarchies have highly impaired survivability. Consequently, improved understanding of condor socioecology is needed to inform conservation management strategies. We report on the dominance structure of free-ranging condors and identify the causes and consequences of rank in condor populations by matching social status with the behavioral and physical correlates of individual birds. We characterized the hierarchical social structure of wild condor populations as mildly linear, despotic, and dynamic. Condor social groups were not egalitarian and dominance hierarchies regulated competitive access to food resources. Absence of kin-based social groups also indicated that condor social structure is individualistic. Agonistic interactions among condors were strongly unidirectional, but the overall linearity and steepness of their hierarchies was low. Although one aggressive male maintained the highest dominance rank across the 3-year observation period, there was considerable fluidity in social status among condors within middle and lower rank orders. Older condors were more dominant than younger birds and younger males supplanted older females over time to achieve higher status. Dominance rank did not predict the amount of time that a bird spent feeding at a carcass or the frequency that a bird was interrupted while feeding. Thus, younger, less dominant birds are able to obtain sufficient nutrition in wild social populations.     

Achromatic color variation in black-capped chickadees,Poecile atricapilla: black and white signals of sex and rank

Sexual dichromatism and phenotypic variation in elaborate male traits are common products of sexual selection. The spectral properties of carotenoid and structurally-based plumage colors and the patch sizes of melanin-based plumage colors have received considerable attention as sexual signals in birds. However, the importance of variation in achromatic plumage colors (white, gray and black) remains virtually unexplored, despite their widespread occurrence. We investigated a potential signal function of the achromatic black and white plumage of black-capped chickadees (Poecile atricapilla). We captured and color-banded 178 free-living chickadees and assessed winter flock dominance hierarchies by tabulating pairwise interactions at feeders. We recaptured 73 of these birds and measured plumage coloration for six body regions using a reflectance spectrometer and the area of melanin-based plumage patches from standardized photographs. We found extensive individual variation in chickadee plumage traits and considerable sexual dichromatism. Male black-capped chickadees have significantly brighter white plumage than females, larger black patches, and greater plumage contrast between adjacent white and black plumage regions. We also found rank differences in the plumage reflectance of males; high-ranking males, who are preferred by females as both social and extra-pair partners, exhibit significantly darker black plumage and grow their feathers more rapidly than low-ranking males. This variation among individuals reveals a potential signal function for achromatic plumage coloration in birds.     

Imperfect female choice and male mating skew on leks of different sizes

We present a model of error-prone female choice on leks, and investigate the effects of different degrees of error on the distribution of mating success among males present at leks of different sizes. At higher levels of error, the best male is predicted to gain a smaller share of matings, while low-ranking males gain a larger share. Males who are of high rank but not the most desirable on the lek do best at intermediate levels of error, since the top-ranked male does not then claim all the matings, but assessment is still sufficiently accurate for females to discriminate between high-ranking and low-ranking competitors. The effects of error are shown to be more pronounced on larger leks, due to smaller expected differences in mating value between males of adjacent ranks. This interaction between lek size and error suggests that observed negative relationships between lek size and mating skew need not be attributed solely to intrasexual competition, as previously suggested, but could also be a result of imperfect choice. Received: 20 February 1998 / Accepted after revision: 25 October 1998     

Do males and females differ in the feeding of large and small siblings? An experiment with the bluethroat

Males and females have been reported to differ in their feeding of large and small siblings in several species of birds. According to recent hypotheses, this phenomenon may be related to a sexual conflict over avian hatching patterns. We designed an experiment to test for the existence of such a sex difference by manipulating nestling size hierarchies of the bluethroat (Luscinia s. svecica) in two directions; half the broods were “asynchronized” to yield large size-differences within broods and the other half were “synchronized” to yield small size-differences. In all broods, nestlings were categorized as being either large or small according to body mass. We recorded male and female food distribution by video early (day 4 after hatching) and late (day 8) in the nestling period. Males and females did not differ in their distribution of food among different-sized nestlings. With large size-differences, both males and females fed large nestlings nearly twice as often as small ones. In contrast, when the size-differences were small, food was more evenly distributed among nestlings. Early in the nestling period, males fed more nestlings during each feeding visit than did females. Our finding that male and female bluethroats do not differ in the feeding of large and small siblings is in contrast to most previous studies. Variation in costs and benefits to males and females from feeding different-sized nestlings, and restrictions to parental choice due to nestling interactions, may explain interspecific variation. Received: 27 June 1997 / Accepted after revision: 26 January 1998     

Sexual selection on mature male parr of masu salmon (Oncorhynchus masou): does sneaking behavior favor small body size and less-developed sexual characters?

In many salmonid species, males exhibit morphological dimorphism associated with alternative mating behaviors. ”Precocious males” have a small body size with little or no development of sexual characters and adopt sneaking to gain access to females, while ”migratory males” of large body size and well-developed secondary sexual characters fight. We quantified selection on precocious male parr of masu salmon (Oncorhynchus masou) under simulated natural conditions to examine the contribution of morphology to sneaking success. In contrast to the prediction that sneaking behavior favors small body size, we detected selection favoring relatively large body size for sneaking success. This selection pressure was caused by the dominance hierarchy within parr and may have been facilitated by indifference of dominant migratory males to parr. Unlike the secondary sexual characters exhibited by migratory male salmon, such as the hooked snout and humped back, no morphological characters other than body size contributed to the reproductive success of masu salmon parr. This non-contribution may have been responsible for the lack of development of sexual characters in precocious males. Received: 15 November 1999 / Accepted: 20 May 2000     

Behavioral correlates of non-random mortality among free-ranging female vervet monkeys

Summary Over a 3.5 year period, illness and predation operated in a non-random manner on free-ranging vervet monkeys in Amboseli National Park. As a result, there was no correlation among adult females between dominance rank and reproductive success. Deaths due to illness were concentrated among low-ranking individuals, and appeared to occur as a result of restricted access to food and water during the dry season. In contrast, deaths due to predation were concentrated among high-ranking individuals. The precise cause of such increased vulnerability could not be determined.High-ranking females alarm-called at higher frequencies than low-ranking females, and were also more aggressive than low-ranking females during intergroup encounters. In contrast, low-ranking females were more likely to initiate friendly interactions with the members of other groups. The non-random distribution of causes of mortality suggests that individuals living in the same social group may confront different selective pressures. Perhaps as a result, individuals appear to respond differently to similar social and environmental variables.     

Sexual maturity and hermaphroditism of the red porgy Pagrus pagrus (Teleostei: Sparidae)

Gonadal changes related to sexual maturity were examined by histology in reared populations of the red porgy Pagrus pagrus L., aged 0.5 to 6.5 years. More than 50% of red porgies were mature at the age of 4 years although some individuals were already mature at the age of 3, as indicated by changes in the gonadosomatic index, plasma levels of vitellogenin, 17β-estradiol and testosterone in females and males. Sex frequency distribution analysis in relation to the age and histological analysis of the gonads indicated the presence of males originated from the sex change of adult functional females. P. pagrus is a protogynous species. Some females had not changed sex up to the age of 6.5 years. Received: 16 July 1998 / Accepted: 29 April 1999     

Biology of the deep-sea octopus Bathypolypus sponsalis (Cephalopoda: Octopodidae) from the western Mediterranean Sea

Several aspects of the biology of Bathypolypus sponsalis were studied from 297 individuals (115 males, 180 females and 2 indeterminates) caught in a depth range of 200–800 m depth in the western Mediterranean Sea. The paper presents data on sizes (length-weight relationships, size-frequency distributions) and reproduction (sex ratio, maturation, condition), and also analyses of the diet of B. sponsalis from samples taken throughout the year. Length-weight relationships showed that females are heavier than males at the same mantle length. Although mature individuals were found all year round, the maximum number occurred in spring and summer. Sexual maturation data revealed that males mature at smaller sizes than females. The gonadosomatic index increased with maturity in both sexes; the increase was gradual in males, but abrupt in females. The digestive gland index was used as a condition index and showed a differential behaviour with maturity; it increased gradually in females, but decreased in males. Like other octopus species, B. sponsalis appears to be an opportunistic predator, feeding on a great variety of preys. Stomach content analysis yielded a total of 19 different prey items belonging to four major groups (Crustacea, Mollusca, Ophiuroidea and Osteichthya). The first three groups were the more frequent preys, since crustaceans, molluscs and ophiuroids appeared in 76%, 49% and 30% of the stomachs, respectively. Decapoda Reptantia (among crustaceans) and cephalopods and bivalves (among molluscs) constituted the more abundant prey items. While the Decapoda Reptantia group was significantly more abundant in stomach contents of females, gastropods were taken more frequently by males. These differences in diet could reveal females as a more active predators than males. Received: 5 March 2000 / Accepted: 7 November 2000     

Occurrence of large numbers of carcasses of the large,grazing copepod Calanus cristatus from the Japan Sea

Carcasses of Calanus cristatus were discovered in plankton samples collected from the Japan Sea throughout the year from 1970 to 1985. Many carcasses of copepodite Stages IV and V occurred in the layer between 15 and 300 m below a distinct thermocline. The number of copepodite Stage V carcasses also peaked in the layer between 1 500 and 2 000 m. The highest density of copepodite Stages IV and V carcasses was 169 individuals per 1 000 m³ and 1 573 individuals per 1 000 m³, respectively. Carcasses of adults occurred at depths below 500 m and numbers of males and females per 1 000 m³ were 1 to 16 and 1 to 42, respectively. Living males were larger in catch number than living females, but the relationship for carcasses was the opposite. Weight of carcasses was 15 to 25% of living C. cristatus. Carcasses contained about 51% carbon and 8% nitrogen by weight. Carcasses may have been drifting for more than one year in the epipelagic layer under the thermocline because of their slow decomposition rate.     

Sex differences in the movement patterns of free-ranging chimpanzees (<Emphasis Type="Italic">Pan troglodytes schweinfurthii</Emphasis>): foraging and border checking

Most social primates live in cohesive groups, so travel paths inevitably reflect compromise: decision processes of individuals are obscured. The fission–fusion social organisation of the chimpanzee, however, allows an individual's movements to be investigated independently. We followed 15 chimpanzees (eight male and seven female) through the relatively flat forest of Budongo, Uganda, plotting the path of each individual over periods of 1–3 days. Chimpanzee movement was parsed into phases ending with halts of more than 20 min, during which individuals fed, rested or engaged in social activities. Males, lactating or pregnant females and sexually receptive females all travelled similar average distances between halts, at similar speeds and along similarly direct beeline paths. Compared to lactating or pregnant females, males did travel for a significantly longer time each day and halted more often, but the most striking sex differences appeared in the organisation of movement phases into a day's path. After a halt, males tended to continue in the same direction as before. Lactating or pregnant females showed no such strategy and often retraced the preceding phase, returning to previously visited food patches. We suggest that female chimpanzee movements approximate an optimal solution to feeding requirements, whereas the paths of males allow integration of foraging with territorial defence. The ‘continually moving forwards’ strategy of males enables them to monitor their territory boundaries—border checking—whilst foraging, generally avoiding the explicit boundary patrols observed at other chimpanzee study sites.     

Vigilance in wild Thomas's langurs (Presbytis thomasi ): the importance of infanticide risk

This study examines vigilance as a behavioural indicator of the importance of infanticide risk by comparing the infanticide avoidance hypothesis with the predation avoidance and mate defence hypotheses for wild Thomas's langurs (Presbytis thomasi) in Sumatra. We found that all individuals were more vigilant in situations of high predation risk, i.e. lower in the trees and in the absence of neighbours. Females were also more vigilant on the periphery of the group. However, there were variations in vigilance levels that could not be accounted for by the predation avoidance hypothesis. Males without infants showed higher levels of vigilance in areas of home range overlap than in non-overlap areas during the early phase of their tenure, strongly suggesting mate defence. In these areas of home range overlap where Thomas's langur groups can interact, males may attack females and infants, and so the infanticide risk for males and females with infants is likely to be high in these areas. Only females with infants, but not males with infants or females without infants, showed higher vigilance levels in overlap areas than in non-overlap areas; in addition, in overlap areas, females with an infant were more vigilant than females without an infant, while this was not the case in non-overlap areas. Both females and males with infants were more vigilant high in the trees than at medium heights in overlap areas but not elsewhere. These findings can only be explained by the infanticide avoidance hypothesis. In contrast to predator attacks, infanticidal male attacks come from high in the canopy, and only occur in overlap areas. There was a significant sex difference in vigilance, but males were only more vigilant than females without an infant, and not more vigilant than females with an infant. We conclude that vigilance varied mainly in relation to the risk of predation and infanticide. Mate competition only played a role for males during the early phase of their tenure. Predation risk seems to offer the best explanation for vigilance for all individuals in the absence of infants. Both predation risk and infanticide risk played a role for females and males with infants. Received: 4 April 1998 / Accepted after revision: 6 September 1998     

Males are faster foragers than females: intersexual differences of foraging behaviour in the Apennine chamois

Availability of food resources and individual characteristics can influence foraging behaviour, which can differ between males and females, leading to different patterns of food/habitat selection. In dimorphic species, females are usually more selective in food choice, show greater bite rates and spend more time foraging than males. We evaluated sexual differences in foraging behaviour in Apennine chamois Rupicapra pyrenaica ornata, during the warm season, before the rut. Both sexes selected nutritious vegetation patches and spent a comparable amount of time feeding. However, males had a significantly greater feeding intensity (bite rate) and a lower search effort for feeding (step rate), as well as they spent more time lying down than females. Females selected foraging sites closer to refuge areas than males. In chamois, sexual size dimorphism is seasonal, being negligible in winter–spring, but increasing to 30–40 % in autumn. Our results suggest that males enhance their energy and mass gain by increasing their food intake rate during the warm season, to face the costs of the mating season (November). Conversely, females seem to prioritize a fine-scale selection of vegetation and the protection of offspring. A great food intake rate of males in the warm season could have developed as a behavioural adaptation leading herbivores to the evolutionary transition from year-round monomorphism to permanent dimorphism, through seasonal dimorphism.     

Effects of male dominance and courtship display on female choice in the ring-necked pheasant

Male traits and behaviours acting in mate choice and intrasexual competition are expected to be congruent. When studying their evolution, this often makes it difficult to differentiate between these two components of sexual selection. Studies are therefore needed on mate choice in conjunction with the role of displays and dominance. We present the results from two experiments conducted to investigate the effects of male dominance and courtship displays on female choice in the ring-necked pheasant, Phasianus colchicus, controlling for differences in morphological male traits. We found: (1) different courtship behaviours had different effects on female choice: females were mainly attracted by the feeding courtship behaviour, while another courtship display (the lateral display) was effective in producing the copulation-acceptance response by the females; (2) subordinate males performed the courtship behaviour before females less frequently than dominant males, and females reinforced intrasexual selection by choosing dominant males, and (3) subordinate males in visual contact with a dominant became less attractive to females. The results support the idea (armament-ornament model) that female pheasants may benefit from using traits selected in male-male competition as clues for mate choice. Received: 23 October 1997 / Accepted after revision: 7 October 1998     

Food or sex—males and females in a sex role reversed pipefish have different interests

In a sex role reversed pipefish, Syngnathus typhle, we found that basic life history allocations were directly influenced by sexual selection. We investigated time allocation to foraging and mating, respectively, in a choice experiment, giving males and females, of small or large body size, a choice between food and a potential partner. We found that males were more interested in foraging than mating, i.e., were more frequently observed in front of the food than in front of the partner, whereas females were more interested in the potential partner. This reflects sexual selection operating differently on the two sexes, as males and females are relatively similar in other life history traits, such as growth, mortality, age of maturity, dispersal, and parental expenditure. Moreover, large individuals allocated more time to mating activities, small to feeding. Individuals more interested in mating compared to food were subsequently more critical when given a choice between a large (high-quality) and a small (low-quality) partner, whereas individuals more interested in food were not selective. These findings are consistent with our predictions: sex-role reversed males can be relatively sure of achieving one or more matings, and should allocate more time to feeding and, hence, to parental investment, growth and/or future reproduction. Females, on the other hand, have more uncertain mating prospects and should allocate time to imminent reproductive activities, thereby foregoing other life history traits such as growth and future egg production. By this, they also sacrifice future fecundity and attractiveness.     

Activity pattern and spatial strategy in Pachygrapsus marmoratus (Decapoda: Grapsidae) from Mediterranean and Atlantic shores

The spatial strategies and activity patterns of two populations of Pachygrapsus marmoratus (Fabricius), from Italy and Portugal, were studied in the field. The Mediterranean site was a non-tidal splash zone about 30 cm wide, while the Atlantic site was a 30 m wide intertidal belt. Almost all P. marmoratus of both populations were active on the emerged rocks, rarely entering the sea. The Mediterranean crabs actively fed on the algal turf throughout the 24 h, while in Portugal active crabs were more numerous at night. In the Italian population, all crabs were packed together, although large males had larger activity areas than females and smaller males. In Portugal, the large males were more concentrated in the sublittoral fringe, while both small males and females were confined to the eulittoral and littoral fringe. The Italian P. marmoratus performed feeding loops within areas smaller than 9 m². Although most Portuguese crabs showed the same spatial strategy, some of them performed feeding migrations to the sublittoral fringe, covering distances of up to 20 m. Environmental factors (e.g. temperature), physical factors (e.g. refuge availability) and biotic factors (e.g. competitors and predators) of the different shores probably determine the behavioural plasticity of␣P.␣marmoratus. Received: 1 February 1998 / Accepted: 13 November 1998     

The cost of sexual coercion and heterospecific sexual harassment on the fecundity of a host-specific, seed-eating insect (Neacoryphus bicrucis)

Two species of seed-eating true bugs, Neacoryphus bicrucis (Heteroptera: Lygaeidae) and Margus obscurator (Heteroptera: Coreidae) co-occur on ragwort, Senecio tomentosus, in southern Georgia, USA. Males of both species sometimes engage in chases and protracted grappling with females that flee initial mountings. Sometimes genital coupling occurs while the wriggling female is restrained in the male's grasp. Chases, grappling, and mounting attempts are misdirected toward heterospecific females, heterospecific males, or conspecific males. In a laboratory study, confinement of mated N. bicrucis females with either conspecific or heterospecific males reduced fecundity by approximately one-half relative to mated females confined only with other females. Perhaps as a consequence of this, N. bicrucis females frequently leave areas of high host plant density, where they prefer to oviposit, when males are abundant. The abundance of each species is positively correlated with host plant density but the two species rarely occur together on the same plants. This may be an effect of heterospecific courtship which induces the flight of N. bicrucis more than the flight of M. obscurator. The laboratory results suggest that copulations following chases and grappling represent sexual harassment, not a mechanism of active female choice for a vigorous mate. As sexual harassment imposes high fitness costs that favor abandonment of host plants, it may, when misdirected, incidentally limit habitat use by ecologically similar species. Received: 3 October 1998 / Received in revised form: 21 January 1999 / Accepted: 14 February 1999     

Competition with flies promotes communal breeding in the burying beetle,Nicrophorus tomentosus

Communal breeding through nest-sharing may benefit cooperating individuals indirectly, in increased inclusive fitness, or directly, when environmental constraints reduce the fitness of solitary breeders. Burying beetles provide extensive parental care and can breed either in pairs or in larger groups of unrelated males and females. Parentage of communally-reared broods is usually shared but is skewed in favor of the individuals of each sex that provide longer care. Females provide care longer than males, and two females are more likely to remain together in the brood chamber than two males are. Flies and other burying beetles are the major competitors for carcasses and this study suggests that it is competition with flies that promotes communal breeding inNicrophorus tomentosus On medium-size carcasses (35–40 g) the presence or absence of oviposition by flies had a significant effect on the size of the brood reared, and on large carcasses (55–60 g) the number of beetles present, two or four, had a significant effect on brood size. On both medium and large carcasses, pairs rearing broods on flyblown carcasses had fewer young than pairs on clean carcasses or foursomes on flyblown carcasses. There was a strong trend for an interaction effect between number of beetles and competition with flies (Table 1). Duration of parental care was not affected by competition with flies except for that of the first male to depart, which provided care longer on flyblown carcasses (Table 2). Pairs and foursomes were equally able to defend the carcass and brood from conspecific intruders and from larger intrudingNicrophorus orbicollis (Table 3).