Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears

Despite significant sexual dimorphism and differing reproductive strategies in carnivores, sexual segregation is rarely studied and is often overlooked in the management of wild populations. Potential nutritional constraints imposed by sexual dimorphism and differing reproductive strategies between the sexes have important implications, particularly when combined with differential effects of human activities on sex and age classes. We examined the effects of sexual dimorphism, reproductive strategies, and human activities (bear-viewing and hunting) on resource use by different sex and age classes of brown bears (Ursus arctos). Sexual segregation of habitat use and effects of experimental bear-viewing were quantified at a single site in south-central Alaska, U.S.A., by capturing, collaring, and observing brown bears at a salt marsh and salmon stream. Effects of salmon capture rate, availability of alternative salmon runs, harvest pressure, and numbers of annual visitors on sex and age class use were examined from data collected or previously published from 13 other sites. Bear-viewing sites on salmon streams where salmon capture rates were low (<4 salmon/hour) resulted in low use by adult males (<10% of all bears), except for sites with falls. However, maximum male use of viewing areas also depended on the availability of alternative salmon streams and harvest pressure. Use of habitats by females with dependent young was significantly related to the prevalence of adult males at the site. Thus, both sexual dimorphism and differing reproductive strategies led to sexual segregation in habitat use by bears. As a result of infanticide, females with young appear to prioritize avoidance of male bears over avoidance of humans when choosing habitats, in contrast to responses documented in herbivores. Because carnivores often exhibit both sexual dimorphism and infanticide, selection for sexual segregation is likely to be high. In these cases, the nutritional demands of large adult males, balanced with responses to human activity, drive dynamic temporal and spatial distributions of individuals in the population.     

Males in the shade: habitat use and sexual segregation in the Galápagos sea lion (Zalophus californianus wollebaeki)

In many sexually dimorphic species adult sexes tend to segregate socially, spatially, or in habitat use. Several hypotheses have been formulated regarding underlying mechanisms. We investigated terrestrial habitat use and sexual segregation in a tropical otariid, the Galápagos sea lion (Zalophus californianus wollebaeki), where most of the hypotheses can be ruled out a priori. Factors relating to thermoregulation and costs of locomotion were of prime importance for habitat use. Habitats directly adjacent to the sea, with simple structured flat surfaces, shade, and tide pools were most frequented, but sexes and age classes differed in their usage patterns. Sexual segregation, both spatial and by habitat was pronounced in the reproductive period (RP), but remained high during the nonreproductive period (NRP). A GLM model of habitat use showed that in both seasons adult males frequented habitat types that adult females and other age classes used much less. Males were most abundant in suboptimal inland habitats, which offered only shade for cooling. Females with newborns differed in habitat use from females with older offspring and lone females. Spatial and habitat segregation are explained most parsimoniously as by-products of social processes, primarily intrasexual competition and female avoidance of male harassment, linked to the polygynous mating system.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Active swimming in meiobenthic copepods of seagrass beds: geographic comparisons of abundances and reproductive characteristics

The entry of meiobenthic copepods from sediments or seagrass blades into the water column and reproductive characteristics of actively migrating fauna were investigated from 1981–1986 in a temperate intertidal Zostera capricorni seagrass bed in Pautahanui Inlet, New Zealand and in a subtidal Thalassia testudinum bed in Tampa Bay, Florida, USA. Emergence of copepods in New Zealand varied over a tidalcycle, while in Florida a distinct diel periodicity was displayed. Selected copepod species in New Zealand had similar numbers emerging from sediments and/or blades over a 6 h period as the common copepods actively migrating from sediments in Florida. Daily abundances of emerging copepods (24 h) in Tampa Bay, Florida, were substantially greater than those in New Zealand, where migration is linked to tidal cover. In Z. capricorni meadows in New Zealand, sex ratios of copepods in sediments and on blades were dominated by females; males dominated water-column samples. In T. testudinum meadows in Tampa Bay, sex ratios of males to females, although of a lowermagnitude than in Z. capricorni beds, were higher in trap than in sediment samples. Differences in sex ratios, the availability in emergence traps of females of appropriate stage for mating, and observations on clasping in live samples from traps suggest that swimming behavior in copepods may be partly linked to prenuptial courtship. Meiobenthic copepods may use the water column as an important habitat for reproductive behavior.     

Male influence on sex allocation in the parasitoid wasp Nasonia vitripennis

Sex allocation is an important reproductive decision for parents. However, it is often assumed that females have substantial control over sex allocation decisions, and this is particularly true in haplodiploid insects, in which females apparently determine sex by deciding whether to fertilise an egg (and produce a diploid daughter) or not (and produce a haploid son). Mechanisms by which males may influence sex allocation are not so straightforward, and their potential influence on sex ratios has been somewhat neglected. Here, we test whether males influence offspring sex ratios in the parasitoid wasp Nasonia vitripennis. We show that some of the variation in observed sex ratios can be attributed to males when comparing the affect of male strain on sex ratio. We did not find among-male variation in sex ratio with a less powerful experiment using males from only one strain or an effect of male mating environment. Our data suggest that males can influence female sex ratios and contribute to the variation around the sex ratios optimal for females. However, the influence is not large, suggesting that females have more influence on sex allocation than do males. We conclude by considering whether male influences on sex ratio represent differences in male reproductive competence or deliberate attempts by males to increase their fitness by influencing daughter production.     

Mating system,optimal number of matings,and sperm transfer in the Argentine ant Iridomyrmex humilis

Summary In ants, because males have a finite sperm supply and females mate only at the beginning of their reproductive lives, it is possible to infer which is the limiting sex from a few parameters: the amount of sperm produced by males, the amount of sperm stored by females, and the numerical sex ratio. In the Argentine ant Iridomyrmex humilis mating takes place in the nest. Laboratory experiments and field data showed that the numerical sex ratio is heavily male-biased (10.1:1) and that the maximum number of sperm a female can store is similar to the number of sperm a male possesses. Thus females are the limiting sex in this species. In a set of mating experiments, one queen was presented with 1–20 males. The highest proportion of successful matings occurred when females were presented with two males. There was a significant negative correlation between the amount of sperm queens stored and the number of males present in the mating arena. This relationship most likely resulted from male interference during the copulation process. When several males were present in the arena, the mating pair was frequently disturbed by other males trying to copulate. Newly mated queens collected from the field stored 172,000 ± 76,000 sperm, a quantity most similar to that measured in laboratory mating experiments with a ratio of 5 or 10 males per queen. Because the operational sex ratio in I. humilis is highly male-biased, male interference may also decrease the amount of sperm queens store in the field. In many ants, fewer sperm stored by queens should decrease their reproductive success because they would run out of sperm earlier in their reproductive lifetimes. However, comparison of the amount of sperm present in young and old I. humilis queens collected in the field suggests that most use only a small proportion of their sperm supply during their lifetimes. Males mate once and discharge all their sperm during a single mating. Females may mate with several males but dissection of these males indicated that in most cases only one of them had empty seminal vesicles thus suggesting that a single male is responsible for most of the sperm transfer. Thus caution should be exercised in inferring multiple inseminations, as is frequently done in eusocial insects, from the observation of multiple copulations. Correspondence to: L. Keller     

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.     

Selective males and ardent females in pipefishes

Summary In the pipefishes Syngnathus typhle and Nerophis ophidion, males have been shown to limit female reproductive rate, and females to compete for access to males. Hence, these species fit the criteria for sex-role reversal. Males brood the eggs and provide the offspring with nutrients, oxygen and an osmoregulated environment. Moreover, in S. typhle both sexes prefer a larger mate when given a choice. Sexual selection theory predicts that males should be more choosy than females, and that was experimentally demonstrated in this study. We predicted that S. typhle males should be less eager to copulate than S. typhle females with an unattractive (i.e. small) mate. We measured eagerness as the time from the start of the experiment until copulation occurred. Males with unattractive partners took significantly longer to copulate than females with unattractive partners. Moreover, females invariably initiated the courtship dance, and resumed it quicker after copulation than did the males, again suggesting reproductive hesitation in males. Neither male nor female size per se was correlated with time until copulation. In N. ophidion, where we have previously shown that males prefer larger to smaller females, we found that females did not select males with regard to size. Our results are consistent both with earlier findings (males limit female reproduction and females compete for males) and with operational sex ratios in nature: in seven annual field samples in June, the numbers of S. typhle females with ripe eggs always significantly exceeded numbers of receptive males. Hence, the potential cost of being choosy in terms of lost matings is much higher in females than in males. In conclusion, S. typhle females were somewhat choosy, but less so than males, whereas N. ophidion females were not choosy at all. Correspondence to: A. Berglund     

Reconciling competing ecological explanations for sexual segregation in ungulates

Sexual segregation in ungulates has important conservation and theoretical implications, but despite numerous studies, the impetus for this behavioral pattern remains a topic of debate. Sexual segregation hypotheses can be broadly grouped into social and ecological explanations, but only ecological explanations can adequately describe why sexes use different areas and habitats. The reproductive strategy hypothesis (RSH) and forage selection hypothesis (FSH) are leading ecological explanations, and although both have received support in the literature, neither the collective basis for that support nor overlap between these hypotheses has been adequately evaluated. This review analyzed seasonal sex comparisons of habitat forage quantity (n=66), quality (n=67), and diet (n=63) from peer-reviewed studies of north temperate ruminants to test predictions of each hypothesis. Empirical data supported predictions of the RSH, but did not support two of three key predictions of the FSH. Males used habitats with greater forage quantity significantly more than females. But, contrary to predictions of the FSH, females did not use habitats with superior forage quality nor consume higher-quality diets more than males. Sexes typically used habitats and consumed diets of similar quality, and when differences were reported, males used higher-quality habitats significantly more than females. Results refute FSH arguments that differences in dietary requirements associated with sexual dimorphism are a universal explanation for sexual segregation in ungulates, but the physiological mechanism on which the FSH is predicated may explain why males consume poorer-quality diets when high-quality forage is scarce. The FSH, therefore, operates at a proximate level because it explains diet and habitat selection by males under certain environmental conditions, but multiple environmental factors may influence sexual segregation, and no single proximate explanation adequately describes this behavioral pattern. The RSH explains sexual segregation as the evolutionary response to differences in reproductive strategies: males choose habitats to maximize energy gains in preparation for rut, and females select habitats with combinations of resources that contribute to offspring survival. Consequently, the RSH provides an ultimate explanation that can be used to explain and interpret studies of sexual segregation in ungulates.     

Bigamy and mate choice in the biparental cichlid fish Cichlasoma nigrofasciatum

Summary Sixty percent of C. nigrofasciatum males showed bigamous behaviour when held in small, outdoor ponds at an adult sex ratio of 5 males to 10 females. No male held more than two pair bonds simultaneously. Bigamy had reproductive benefits; bigamous males, on average, fathered more broods that survived well into the free-swimming fry stage than did monogamous males. In ponds where adult size varied, no positive correlations were found, for either sex, between adult size and frequency of spawning or success of broodrearing, despite predictions that larger size would confer reproductive benefits on both sexes. A significant positive correlation was found between the sizes of mated individuals in the pond where both sexes varied in size. On theoretical grounds this relationship is likely to be more strongly influenced by female than by male choice of spawning partner.     

Operational sex ratios and behavioural sex differences in a pipefish population

In the pipefish Syngnathus typhle, only males brood embryos in specially developed brood pouches, supplying oxygen and nutrients. Laboratory studies have shown that this elaborate paternal care has led to sex-role reversal in this species: males limit female reproductive rate, females are the primary competitors for mates and males exercise greater selectivity in accepting mates. In the first field study of this pipefish, we describe mating behaviour in the wild and test the hypothesis that temporal variations in the operational sex ratio (OSR) determine sex differences in mating behaviour. Our study comprised two reproductive seasons of two sequential mating periods each, the latter separated by a lengthy interval of male brooding. During mating periods, females displayed to all males without wandering and males moved about searching for females, without reacting to all females. The OSR was least female-biased (or even male-biased) at the onset of the breeding season, when most pipefish were simultaneously available to mate, but became strikingly female-biased as males' pouches were filled. The OSR remained substantially female-biased during the second mating period, because few males became available to remate at any one time. As hypothesised, female-biased OSRs resulted in more female-female meetings. As well, females were above the eelgrass more often than brooding males, thus exposing themselves to conspecifics and/ or predators. In the second year, males arrived earlier than females on the breeding site and male pregnancies were shorter, because of higher water temperatures, so rematings occurred earlier. Males met more often during that year than the previous one, but male competitive interactions were still not observed. The field results support laboratory studies and demonstrate that behaviours associated with female-female competition are more prominent when the OSR is more female-biased. Correspondence to: A. Vincent     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

Genetic reconstruction of breeding patterns in gundis (Rodentia: Ctenodactylidae)

Current mating system theory predicts that the number of females breeding in a group will depend on the number of females in the group and the accessibility of unrelated males, whereas the number of males breeding in a group will depend on the ability of males to control access to reproductive females. By combining information on group composition with genetic data, I determined whether breeding patterns in a rock-dwelling rodent, Ctenodactylus gundi, were concordant with these expectations. C. gundi breeding patterns varied from facultative monogamy to uni-male polygyny to multi-male polygyny. The number of reproductive individuals of each sex in a group increased with group size. Whereas communal breeding among related females tended to increase female reproductive success, males breeding in the same group were unrelated to other group members and seemed to compete over access to matings. While some males were assigned offspring from neighboring social groups, most group-living males successfully monopolized the reproduction of females within their group. There was no evidence that females had multiply sired litters, although some bred with alternative males in separate breeding attempts. Although numerous individuals were not assigned as parents or offspring, genetic information enabled me to determine that most unassigned individuals were philopatric group members, whereas only a few were unrelated immigrants into their current social group. Together, these results provide evidence that C. gundi social groups represent fairly distinct reproductive units whose breeding patterns are dependent on group size and composition in accordance with theoretical predictions. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Possible causes and consequences of philopatry and breeding dispersal in red-backed shrikes <Emphasis Type="BoldItalic">Lanius collurio</Emphasis>

Studies of animal breeding dispersal have often focused on possible causes, whereas its adaptive significance has received less attention. Using an information-theoretic approach, we assessed predictions of four hypotheses relating to causes and consequences of breeding dispersal in a migratory passerine, the red-backed shrike Lanius collurio. As predicted by the reproductive performance hypothesis, probability of breeding dispersal in females (though not in males) decreased with increasing annual average number of fledglings produced in the past year, but there was no association with conspecific reproductive performance in either sex. The site choice hypothesis, stating that individuals disperse to improve breeding site quality, received support in males only, as dispersal probability was positively associated to a measure indicating low territory quality. The social constraints hypothesis, referring to dispersal in relation to intraspecific interactions, received little support in either sex. The predation risk hypothesis was hardly supported either. Consequences of dispersal were marginal in both sexes because neither fledgling production in females, nor territory quality in males improved after dispersal. In addition, males settled on territories closer to the forest edge than those occupied predispersal, which is opposite to the prediction of the predation risk hypothesis. We conclude that own reproductive success was the major factor determining dispersal behavior in females, whereas territory quality and possibly predation risk were most important in males. Overall, breeding dispersal appeared not to be adaptive in this dense population inhabiting an optimal habitat.     

Mate preferences in <Emphasis Type="Italic">Drosophila</Emphasis> infected with <Emphasis Type="Italic">Wolbachia</Emphasis>?

The maternally inherited bacterium Wolbachia pipientis generates strong reproductive incompatibilities between uninfected females and infected males (cytoplasmic incompatibility), significantly reducing both female and male reproductive success. Such fitness costs are thought to place selective pressure on hosts to evolve pre-copulatory preferences for mating with compatible mates, thereby enabling them to avoid the reproductive incompatibilities associated with Wolbachia. Therefore, uninfected females are predicted to prefer mating with uninfected males, whereas infected males are predicted to prefer mating with infected females. Despite these predictions, previous investigations of pre-copulatory mate preferences in Wolbachia-manipulated Drosophila have not found evidence of female preference for uninfected or compatible males. However, none of these studies utilised a design where focal individuals are provided with a simple choice in a relatively non-competitive situation. We examined both female and male pre-copulatory mate preference based on mate infection status in Drosophila simulans and D. melanogaster using simple choice assays involving between 30–50 replicates per treatment. Although we found no evidence of female pre-copulatory mate preferences in either species, male D. simulans exhibited some preference for mating with females of the same infection status. However, this preference was not evident when we repeated the experiment to confirm this finding. Consequently, we conclude that neither male nor female D. melanogaster and D. simulans exhibit significant Wolbachia-associated pre-copulatory mate preferences.     

Population differentiation in female sex pheromone and male preferences in a solitary bee

Population differentiation in female mating signals and associated male preferences can drive reproductive isolation among segregated populations. We tested this assumption by investigating intraspecific variation in female sex pheromone and associated male odour preferences among distant populations in the solitary bee Colletes cunicularius (L.) by using quantitative gas chromatography and by performing field bioassays with synthetic blends of key sex pheromone compounds. We found significant differences in sex pheromone blends among the bee populations, and the divergence in odour blends correlated positively with geographic distance, suggesting that genetic divergence among distant populations can affect sex pheromone chemistry. Our behavioural experiments, however, demonstrate that synthetic copies of allopatric female sex pheromones were cross-attractive to patrolling males from distant populations, making reproductive isolation by non-recognition of mating signals among populations unlikely. Our data also show that patrolling male bees from different populations preferred odour types from allopatric populations at the two sites of bioassays. These male preferences are not expected to select for changes in the female sex pheromone, but may influence the evolution of floral odour in sexually deceptive orchids of the genus Ophrys that are pollinated by C. cunicularius males.     

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.     

Spatial organization and mating system of Microtus townsendii

Summary Space use by individual Townsend's voles, Microtus townsendii, was investigated in spring and summer by means of radiotelemetry and intensive live trapping in undisturbed grasslands near Vancouver, British Columbia. Home ranges of males were larger than those of females; females had significantly larger ranges in spring than in summer. Most males and females maintained territories free of individuals of the same sex in spring. Male-female pairs had their exclusive territories closely overlapping each other. The 1:1 operational sex ratio and the spatial association of pairs of males and females suggest that the voles were monogamous in the spring of 1988 and that 50% of the males were monogamous in the spring of 1989. In summer, there was more intrasexual overlap between home ranges of males and females and female ranges were considerably smaller than those of males. Females were more philopatric than males and females thought to be members of the same family group lived adjacent to each other or had overlapping home ranges. Males overlapped with more than one female in summer, but most females still overlapped with only one male, which suggests that the mating system is polygynous in summer. Thirty-five percent of the philopatric females became pregnant for the first time when the male spatially associated with their mother in the spring was still alive and thus could potentially have mated with their fathers. Male and female territoriality in spring is the proximate mechanism for the limitation of breeding density by spacing behaviour.[/p] Offprint requests to: C.J. Krebs     

Loss of mating opportunities influences refuge use in the Iberian rock lizard,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>

Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.Communicated by A. Mathis     

Body size preferences in the pot-bellied seahorse <Emphasis Type="Italic">Hippocampus abdominalis:</Emphasis> choosy males and indiscriminate females

Male seahorses (genus Hippocampus) provide all post-fertilization parental care, yet despite high levels of paternal investment, these species have long been thought to have conventional sex roles, with female mate choice and male–male competition. Recent studies of the pot-bellied seahorse (Hippocampus abdominalis) have shown that sex-role reversal occurs in high-density female-biased populations, indicating that male mating preferences may lead to sexual selection on females in this species. Egg size, egg number, and offspring size all correlate positively with female body size in Hippocampus, and by choosing large mating partners, male seahorses may increase their reproductive success. While male brood size is also positively correlated with body size, small H. abdominalis males can carry exceptionally large broods, suggesting that the fecundity benefits of female preference for large partners may be limited. We investigated the importance of body size in reproductive decisions of H. abdominalis, presenting focal individuals of both sexes with potential mating partners of different sizes. Mating preferences were quantified in terms of time spent courting each potential partner. Male seahorses were highly active throughout the mate-choice trials and showed a clear behavioral preference for large partners, while females showed significantly lower levels of activity and equivocal mating preferences. The strong male preferences for large females demonstrated here suggest that sexual selection may act strongly on female body size in wild populations of H. abdominalis, consistent with predictions on the importance of female body size for reproductive output in this species. An erratum to this article can be found at     

Sperm economy and limitation in spiny lobsters

Sperm limitation, when female fertilisation success is constrained by the supply of sperm, is generally perceived to be an uncommon feature of reproduction in species which directly transfer gametes during copulation. Male size, previous copulations, and the balance of expected reproductive return and future mating opportunity may, however, limit the amount of sperm males transfer to females. We used laboratory experiments where mate size could be manipulated and its consequences on spermatophore size and clutch size determined, to show that in two genera of spiny lobsters (Crustacea: Palinuridae) male reproductive output limits the size of clutches brooded by females. In Panulirus argus from the Florida Keys, we show that while male size affects spermatophore area, males also vary the amount of ejaculate positively with female size. Furthermore, the area of the spermatophore has a greater influence than female size on subsequent clutch weight. In Jasus edwardsii from New Zealand, female size, male size and mate order all affect clutch weight. In both species, clutches fertilised by small males in the laboratory are significantly smaller than clutches fertilised by large males. These results suggest that to ensure they receive sufficient sperm, females should either mate several times prior to oviposition, mate as early as possible in the reproductive season, or choose large, preferably unmated males as partners and thus compete with other females for preferred males. Sperm-limited female fecundity has the potential to limit the egg production of fished populations where large males are typically rare. Received: 18 May 1998 / Received in revised form: 20 November 1998 / Accepted: 30 November 1998