Mating behavior of the marine copepodOithona davisae

Oithona davisae Ferrari and Orsi were collected from the innermost region of Tokyo Bay, Japan, in 1980 and 1981. The mating behavior of this marine cyclopoid copepod consists of several steps, starting with the paddling of the male in random search of a mate. This behavior is followed by a spiraling movement in pursuance of a mate. Next is the copulatory grasp during which the male grasps the fourth swimming legs of the female partner by means of his first antennae. Whilst in this position, the male's urosome vibrates to allow the spermatophores to extrude from his genital openings. The mating behavior then culminates in the spermatophore transfer. Males do not grasp the uro-some or caudal setae of the mating partner before proper copulatory grasp. The specialized setae of the female's fourth legs may help the male to grasp her legs firmly. Spiraling occurs when the male approaches or traverses the trail of a female that is ready to copulate and that presumably emits a sex-attractant pheromone. The turning radius reduces gradually from more than 1 mm to ca. 0.25 mm as the male approaches the mate female. Females may register spiraling as a mate (male)-approaching signal. Spiraling may lead the male to locate a pheromone source more accurately, and to promote diffusion of the pheromone to prevent other males from pursuing the source. This swimming strategy can increase the copulatory chance of mature virgin females.     

Density affects female and male mate searching in the fiddler crab, <Emphasis Type="Italic">Uca beebei</Emphasis>

In most species, only one sex searches for mates while the other waits. Models of sex-specific mate-searching behavior predict single-sex searching, but the factors that determine which sex searches are not understood. In this study, we examine the effects of density and predation risk on mate-searching behavior in the fiddler crab Uca beebei. U. beebei is one of the few fiddler-crab species in which both sexes search for mates. In a field experiment conducted in Panama, we manipulated crab density and perceived predation risk in replicate plots. Females searched more and males searched less at high densities. At high levels of perceived predation risk, both sexes similarly reduced their search rates. Observations of plots that naturally varied in crab density show that females were more likely to search for mates in areas of higher density, where there were more males. Females may preferentially search for mates in high-density areas because the abundance of nearby burrows, into which they can run to escape predators, decreases their costs of searching and because the abundance of males and male burrows facilitates comparisons and thus may increase their benefits from searching. Males at high densities decrease their mate-searching rate perhaps in response to the increase in female searching and to the corresponding increase in the intensity of their competitors' mate-attraction signals.     

Parental investment in two solitary wasps Ancistrocerus adiabatus and Euodynerus foraminatus (Eumenidae: Hymenoptera)

Summary Females of the solitary eumenid wasps Ancistrocerus adiabatus and Euodynerus foraminatus control the adult size of their offspring by the amount of food provded to the larvae. For both species, larger females provision more offspring and collect more food than smaller females. Males of E. foraminatus, upon emergence as adults, fight for control of the nest area. The winning brother remains at the nest and mates with his sisters as they emerge several days later. Males of A. adiabatus are not as agonistic as males of E. foraminatus so that a number of males may remain near the natal nest. When females nest near each other, male-male interactions increase, and the likelihood of sibling mating decreases. Under conditions of clumped nesting, females make larger offspring and invest more in males than when nesting in isolation.     

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

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

Seasonal variation in male-female competition,cooperation and selfish hoarding in a monogamous songbird

Both cooperation and conflict between the sexes are commonplace in monogamous mating systems. However, little is known about how cooperation and competition varies seasonally in monogamous species that maintain permanent territories. We presented territorial pairs of male and female New Zealand robins (Petroica australis) with a large supply of insect prey at monthly intervals for 2 years. Behavioural observations after food presentation were then made to quantify seasonal and sexual differences in aggressive interactions over prey, prey acquisition rates, mate provisioning, offspring provisioning, selfish food hoarding and cache retrieval. Data were used to evaluate sex-specific behavioural strategies of mediating competition for food. Results showed that males aggressively excluded females from experimental food sources year-round. Females only accessed food sources when males left them unattended. Consequently, females acquired fewer prey than males. After controlling for differences in prey acquisition, both sexes consumed similar amounts of prey in the non-breeding season. Even though males aggressively excluded females from accessing food sources directly, males fed large amounts of prey to females during the breeding season. Both sexes provisioned young at similar rates. Males cached less prey than females in the breeding season but more prey than females in the non-breeding season. Females showed similar caching intensities year-round. Although males tried to defend their hoards, females frequently retrieved male-made caches. Overall, results showed that although New Zealand robins cooperate to raise offspring during the breeding season, conflict between the sexes occurs year-round. Males and females display different behavioural strategies to gain access to experimental food sources, which appear to lessen male–female competition for food and evenly distribute food resources between the sexes.     

An experimental study of paternal behavior in red-winged blackbirds

Summary The effect of brood size and female nesting status on male parental behavior was investigated in red-winged blackbirds Agelaius phoeniceus using brood size manipulation experiments. Male redwings allocated parental effort on the basis of brood size and nestling age. Males began assisting females only at nests with at least three offspring older than three days. Female nesting status had no singificant influence on male parental care. When females were unable to meet a brood's demand for food, males assisted females with nestling feeding. Females did not reduce the amount of food delivered to nestlings when males assisted. The amount of food brought to nestlings by the male was additional to the amount of food provided by the female. Male assistance increased fledgling success. When female provisioning was sufficient to meet a brood's demand for food males did not assist. The value of male parental care varied inversely with the ability of the female to meet nestling food demands. The ability of unassisted females to provide sufficient food and to raise a brood of nestlings successfully appeared to be influenced by resource abundance.     

Dynamic female preference for multiple signals in <Emphasis Type="Italic">Rhinogobius brunneus</Emphasis>

Many species base their choice of mates on multiple signals which provide them with different kinds of information. Choosers may assess the signals together to evaluate the overall quality of potential mates, but individuals often pay attention to different signals in different contexts. In Rhinogobius brunneus, a fish displaying exclusive male parental care, females generally prefer males showing larger first dorsal fins (FDF) and more active courtship displays as mates. Females choosing a mate usually initially assess the FDF and later utilize courtship for the final decision. In our experiments, females with different hunger states used different signals when selecting mates. Females in both hunger states preferred males with larger FDF in the first stage. In the second stage, well-fed females showed highly repeatable choice, whereas poorly fed females responded only to variation in the courtship activity of males. The males preferred by poorly fed females exhibited significantly higher offspring survival than nonpreferred males. Under conditions of food shortage, males allocate more energy to future reproduction at the expense of the present brood, and females may prioritize signals predictive of offspring survivorship over signals reflecting other aspects in male quality to minimize the losses in direct benefits. We conclude that R. brunneus females may employ information from both signals but dynamically adjust their prioritization of each signal to current conditions to ensure the choice that is currently most adaptive.     

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

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

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

How western meadowlarks respond to simulated intrusions by unmated females

Summary Western meadowlarks (Sturnella neglecta) were played back female calls at various stages of their breeding cycles, to examine how residents might respond to unmated females. Females reacted to intruders with aggressive displays and other responses that varied in intensity (Table 1). Males reacted not only to playback, but also to their mates' responses, often intervening in them. The responses of both males and females, however, were strongly affected by the stage of the breeding cycle at which an intrusion was simulated (Table 3). Females responded to playback most strongly before they began incubating eggs; thereafter, the strength of their responses declined rapidly. Males' also responded less strongly at later stages of their mate's nesting cycle, but the strength of their responses declined less rapidly than did the females'. When the reactions evoked from mated individuals were compared, females responded at least as strongly as their mates, until after their young had fledged (Fig. 3). These seasonal changes in the intensity of females' responses suggest that decisions about how strongly to respond take account of the costs of responding to an intruder, as well as its benefits. Our results suggest that interactions among mated and unmated females may affect the timing of bigamy and, in some cases, its incidence as well.     

Mating behaviour of the tiger blue butterfly (Tarucus theophrastus): competitive mate-searching when not all females are captured

Summary Females of the tiger blue butterfly (Tarucus theophrastus) fly upwind in search of the larval hostplant. Males perch or patrol the downwind edges, searching for incoming females or those already on the bush. A model of competitive mate-searching is developed for the case where not all receptive females are contacted by searching males: the model differs from the earlier ones of Parker (1970, 1974), particularly when few males are involved. Observed distributions of males upon bushes of different sizes agree better with the predictions of the game theory model than with a random distribution of males. The described model may be widely applicable to populations where females remate, but frequently evade detection by searching males.     

Association patterns of sailfin mollies (Poecilia latipinna): alternative hypotheses

Individuals may associate with each other due to a variety of selective forces, such as intra- and intersexual selection, and conspecific recognition. Previous studies have concluded that mate choice governs association behavior in polygynous species of fish. I examined whether mate choice underlies the preference for larger individuals by examining preference for association (time spent in proximity to a fish) not only between opposite-sex individuals but also between same-sex individuals of the live-bearing sailfin molly (Poecilia latipinna). Males and females from three size classes were tested with a large and a small object fish of the same and opposite sex. Females preferred to associate with larger over smaller males. Males also preferred to associate with larger over smaller females, as expected. The same female and male test fish also preferred to associate with larger over smaller fish of the same sex. Moreover, females demonstrated no significant difference in their strength of preference (large–small) when offered males or females. The same held true for males. When males and females were subsequently tested with one large male and one large female, females tended to prefer large males while males showed no significant preference for association based on sex. In another experiment, females were tested with a large female and a small male, and significantly preferred the former. These findings suggest that association patterns may have arisen under a variety of conditions, such as predation pressures, shoaling behavior, and associative preference behavior. The assumption that association behavior is a uniformly sufficient predictor of mate choice in fish needs to be re-examined for P. latipinna and other species. Received: 6 November 1998 / Received in revised form: 12 May 1999 / Accepted: 12 May 1999     

Feeding and reproductive behaviour and their effect on catchability of the spanner crab Ranina ranina

The behaviour of 16 adult (8 male and 8 female) spanner crabs (Ranina ranina), collected off southern Queensland, Australia, was monitored continuously in captivity by closed-circuit television for fifteen months from September 1982. Spanner crabs spent most of the time buried in the substrate, emerging mainly only when food became available. They remained emerged for twice as long on feeding days as on days without food. Females responded significantly faster than males to the presence of food. Response was slowest in October and November. There was no correlation between temperature and response time. The average feeding time was 2.0 min (SE=0.12), with no significant difference between males and females. Crabs without food were aggressive towards crabs with food, which sometimes led to fighting and wounding. These interactions also could result in food being transferred from one individual to another. When males interacted, food was transferred more often than when either females, or males and females interacted. Around moulting, male crabs did not feed for 52 d (SE=9.0) and females for 22 d (SE=2.2). This habit would reduce the frequency with which newly moulted crabs are caught in the baited tangle nets used by commercial fishermen. In mating interactions, copulation was always initiated by the males. Males dug up other crabs but, apparently unable to distinguish the sex of these individuals, attempted copulation with either sex. The majority of copulations occurred between midday and midnight and in the period August to December. The frequency of copulations with a female increased 10 d before she extruded eggs, after which it dropped to zero for the following 41 to 50 d. Eggs were carried for 39 to 44 d in the period September to November. The females remained emerged for long periods before extruding their eggs, but the period shortened immediately afterwards. This behaviour would lead to low catchability of ovigerous females. Because females respond more rapidly than males to a food stimulus, they may be more catchable in baited nets than are males. It is concluded that seasonal changes in the behaviour of spanner crabs could affect the number caught by baited tangle nets and may influence the sex ratio in catches.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Host plant density and territorial behavior of the seed bug,Neacoryphus bicrucis (Hemiptera: Lygaeidae)

Summary Adults of the seed bug, Neacoryphus bicrucis, are most numerous in large host plant patches and where host plants are most dense within patches. Males patrol small territories composed of host plants and expel other males and unreceptive females. Larger males aggressively exclude smaller males from high quality territories where both host plants and females are most dense. Consequently, large males mate more frequently than small males. Males remain for longer time in areas where host plants are more dense. However, mating experience influences tenure within a host plant patch such that males tend to remain where they have recently mated. Courtship is aggressive and male territorial behavior is similar to courtship behavior. Preference for territories in areas of high host plant density appears to have been selected to increase the rate of encounter with females which preferentially oviposit in areas of high host plant density.     

Daring females,devoted males,and reversed sexual size dimorphism in the sand-dwelling spider <Emphasis Type="Italic">Allocosa brasiliensis</Emphasis> (Araneae,Lycosidae)

Sexual selection theory predicts that a higher investment in offspring will turn females into the selective sex, while males will compete for accessing and courting them. However, there are exceptions to the rule. When males present a high reproductive investment, sex roles can reverse from typical patterns, turning males into the choosy sex, while females locate males and initiate courtship. In many spiders, males are smaller than females, wandering in search of sedentary females and maximizing the number of copulations. In the present study, we present findings on the sand-dwelling wolf spider, Allocosa brasiliensis, evidencing a reversal in typical courtship roles reported for the first time in spiders. Males were bigger than females. Females located males and initiated courtship. Copulation always occurred in male burrows and took place mainly in long burrows. Males donated their burrows to the females after copulation, closing the entrance before leaving with female cooperation from inside. Males would provide females with a secure place for ovipositing, being exposed to predation and diminishing their future mating possibilities until constructing a new burrow. The cost of vacating the burrow and losing the refuge in an unpredictable habitat, such as sand dunes, would explain the courtship roles reversal in this spider species. Results turn A. brasiliensis as a promising model for discussing the determinants of sex roles and the pressures that drive their evolution and maintenance. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

The reproductive behavior of Anthidium manicatum (Hymenoptera: Megachilidae) and the significance of size for territorial males

Summary Males of the wool-carder bee, Anthidium manicatum, patrol clumps of garden plants. Females of this species visit these plants for pollen, nectar, and pubescence; they also mate there. Females are polyandrous, with intervals between copulations as short as 35 s. Patrolling males defend their territories (0.1–1.3 m²) against other males and against other species of flower-visiting insects. Honey bees may be rendered unable to fly by the attacks of A. manicatum.Territory owners perform exploratory flights to other males' territories, changing territories often (median ownership 4–7 days; maximum 30 days) and flying up to 450 m to establish new territories. Territorial usurpations are nearly always by larger males.Female visitation rate is significantly correlated with number of flowers on a territory. The head size of territory-owner males shows significant correlation with territorial quality (measured by number of flowers, not area) and thus with number of female visits and copulatory opportunities. Some males fail to maintain territories and instead attempt to forage and copulate in other males' territories while the owners are otherwise occupied. Nonowner males are significantly smaller than owners, forage less often and from fewer flowers, and achieve significantly fewer copulations than owners. Females, however, do not reject smaller, nonowner males at a higher rate than they do larger, owner males; their choice for male size appears to be indirect, based instead on choice of food resource.The interval between a copulation and the male's next attempt with a different female is not shorter than that involving the same female. Males do not escort just-mated females about their teritories, as observed in Anthidium maculosum. Territorial behavior in this species most likely evolved through intrasexual competition for reproductive success which led to sexual dimorphism. The defense of a resourcebased territory is the mechanism used by a male to maximize his reproductive potential.     

Philopatry and nest founding in a primitively social bee,Halictus rubicundus

Summary Observations of inter- and intraseasonal dispersal patterns in the primitively social sweat bee Halictus rubicundus in New York reveal considerable philopatry in both sexes. Females overwinter away from the nest aggregation, in diapause for 8–11 months, and return to dig new nests close to the site of their natal nests (typically within 50 cm). Nests are all founded by siggle females (haplometrosis). Clumping of nests may reflect patterns of soil vegetation. Workers rarely enter or take up residence in non-natal nests. Males commonly return to both natal and non-natal nests, and actively patrol vegetation near the natal aggregation for females. These philopatric tendencies, in combination with the haplometrotic mode of nest founding, should have important effects on population structure, particularly in terms of population subdivision, mate choice, and intracolony interactions. The resulting population structure may be conducive to the evolution and maintenance of social behavior.     

Biology of Octopus vulgaris off the east coast of South Africa

The biology of Octopus vulgaris Cuvier inhabiting subtropical littoral reefs off the east coast of South Africa was investigated. Analyses of stomach contents and lair middens revealed that the mussel Perna perna was the dominant food organism. Growth rate of captive individuals was higher than has previously been recorded but food conversion was lower. Females became sexually mature at 900 g which is estimated to be attained in 3.6 months. Males became sexually mature at 400 g, which is estimated to be attained after 3 months. Results indicate that females live for 9–12 months and have the potential to reach 4 000 g in 240 d while males live for about 12–15 months and have the potential to achieve 4 000 g in 290 d. Mating and breeding occurred throughout the year although evidence for the seasonal migration of females is presented and discussed in relation to breeding and feeding behaviour.     

Experimental increase in food supply influences the outcome of within-family conflicts in Tengmalm’s owl

Within a family there are conflicts of interest between parents and offspring, and between male and female parents, over the supply of parental care. The observed pattern of parental care is the outcome of negotiations within the family, and may be influenced by environmental factors such as food abundance. We experimentally increased food supply to ten Tengmalm’s owl (Aegolius funereus) nests from hatching to fledging, mimicking natural cached prey. Ten un-supplemented nests served as controls. Parents and offspring were fitted with radio-tags. Food provisioning by parents was measured both in the (1) mid- and (2) late nestling stage and in the (3) early and (4) late post-fledging stage. In response to food supplementation, both males and females reduced food provisioning, but the effect was more pronounced in females. Females generally contributed much less to food provisioning than males, and food supplementation increased the difference between the sexes. Mass loss during the brooding stage was substantially lower for supplemented than for control females. Food supplementation did not improve offspring survival, and had no effect on body measurements of nestlings. In conclusion, parents of both sexes used the increased food supply to reduce the costs of caring for their current offspring, but females responded more strongly than males.