Sperm Limitation Research Articles

Environmental and fishing effects on the abundance, size and sex ratio of the blue‐throated wrasse, Notolabrus tetricus, on South Australian coastal reefs

Abstract The blue‐throated wrasse, Notolabrus tetricus (Richardson), is a site‐attached protogynous hermaphroditic species, with a small home range of 1000–2000 m2, and the most abundant fish on inshore coastal reefs of South Australia. Long‐term studies over 18–24 years were conducted at an island reserve site and a fished mainland reference site. At the reserve site the retention rate of fish (reduced by mortality + emigration) was directly correlated with female mean size; the female:male sex ratio ranged from 10 to 20:1, consistent with an induction‐inhibition model of sex‐change. At the fished reference site, female mean size declined under intense fishing, and the sex ratio became strongly skewed towards females. Additional spatial surveys over >2000 km of coast showed that: (1) juvenile and adult abundance was variously determined by habitat features, such as substratum‐type, depth, bottom relief, wave exposure and algal canopy cover, and by geographical factors, such as distance off‐shore and within gulfs; and (2) that female mean size and the sex ratio were strongly influenced by recreational fishing, and to a small extent by bottom relief and other factors. The results suggest that female mean size and the sex ratio together can be used as an indicator of recreational fishing intensity at local scales. As populations of sequential hermaphroditic species are sensitive to size‐selective harvesting, management measures should aim to prevent sperm limitation, reproductive failure and skewed sex ratios in this species.

Courtship rate signals fertility in an externally fertilizing fish

Sperm limitation is widespread across many animal species. Several mechanisms of sperm allocation have been proposed, including optimal allocation according to clutch size and equal allocation across females. However, considerably less effort has been directed at investigating the behavioural signals associated with sperm limitation in males, which may include mating rate and the intensity of courtship. We investigated whether multiple successive spawnings affect individual male fertilization success, mating rates and courtship rates in Japanese medaka (Oryzias latipes). Across an average of 17 spawning events per male, fertilization success decreased from 83.7 per cent for the first spawning to 40 per cent for the last spawning while courtship rate decreased from 3.4 to 1.5 min⁻¹. Females appeared to respond to male sperm depletion by reducing clutch size. Our results suggest that male Japanese medaka are sperm-limited, and that courtship rate may be an honest indication of fertilization ability.

Sea urchin fertilization in a warm, acidified and high pCO 2 ocean across a range of sperm densities

Marine invertebrate gametes are being spawned into an ocean simultaneously warming, acidifying and increasing in pCO 2. Decreased pH/increased pCO 2 narcotizes sperm indicating that acidification may impair fertilization, exacerbating problems of sperm limitation, with dire implications for marine life. In contrast, increased temperature may have a stimulatory effect, enhancing fertilization. We investigated effects of ocean change on sea urchin fertilization across a range of sperm densities. We address two predictions: (1) low pH/increased pCO 2 reduces fertilization at low sperm density and (2) increased temperature enhances fertilization, buffering negative effects of acidification and increased pCO 2. Neither prediction was supported. Fertilization was only affected by sperm density. Increased acidification and pCO 2 did not reduce fertilization even at low sperm density and increased temperature did not enhance fertilization. It is important to identify where vulnerabilities lie across life histories and our results indicate that sea urchin fertilization is robust to climate change stressors. However, developmental stages may be vulnerable to ocean change.

Spawning behaviors of two congeneric brittle stars

Broadcasting gametes for external fertilization is a common reproductive strategy of many marine invertebrates; however, little is known about adult behaviors because of the brevity and unpredictability of spawning. We studied the spawning of 2 shallow-water brittle stars — the subtidal Ophiocoma dentata and the intertidal O. scolopendrina — in southern Taiwan by inducing males to spawn after exposure to conspecific ovary suspensions. Laboratory results indicated that male O. dentata preferred to cohabit with females, but that male-male antagonism did not occur. In the field, males with female companions were more inducible to spawn than other males. In contrast, O. scolopendrina did not pair; instead, males rapidly approached released eggs before spawning. Despite this, 53% of simulated egg releases in tide pools did not induce any male to spawn. Male spawning capacities in both species were relatively limited; spawning inducibility rates and disc- pumping frequencies decreased with repeated inductions. Evidence of sexual competition was observed during the spawning induction of O. scolopendrina in tide pools. Only males that arrived first were likely to spawn, whereas late individuals had low rates of spawning. The fitness of female O. scolopendrina may be limited by the shortage of spawning males; however, at the same time, males face sexual competition in addition to sperm limitation. This is due to the female-biased sex ratio, and the limited spawning capacity of unevenly distributed males.

Variable reproductive success in fragmented populations

Marine habitats are naturally patchy and anthropogenic disturbance can further fragment them. Many marine animals are sessile as adults or obligate inhabitants of particular habitats, so populations living in isolated patches of habitat are linked largely by dispersal of planktonic larvae. Theoretically, larvae are more likely to find and settle into large patches of habitat than small patches, thus small habitat patches may experience a more discontinuous supply of recruits resulting in small populations with unusual size- or age-structures or odd sex ratios — conditions where Allee effects on reproductive success are likely. We tested this hypothesis for the Caribbean spotted spiny lobster ( Panulirus guttatus), an obligate inhabitant of coral patch reefs whose mating dynamics are size-dependent. We found that P. guttatus were less abundant on small reefs where their size structure and per capita reproductive success were significantly more variable, particularly among large females that are susceptible to sperm limitation that diminishes fertilization rates. These results are indicative of Allee effects and provide a mechanistic understanding of how size-dependent mating dynamics influence reproductive success in ways that alter population dynamics in patchy habitats.

Quantity matters: male sex pheromone signals mate quality in the parasitic wasp Nasonia vitripennis

Sexual selection theory asserts that females are well adapted to sense signals indicating the quality of potential mates. One crucial male quality parameter is functional fertility (i.e. the success of ejaculates in fertilizing eggs). The phenotype-linked fertility hypothesis (PLFH) predicts that functional fertility of males is reflected by phenotypic traits that influence female mate choice. Here, we show for Nasonia vitripennis, a parasitic wasp with haplodiploid sex determination and female-biased sex ratios, that females use olfactory cues to discriminate against sperm-limited males. We found sperm limitation in newly emerged and multiply mated males (seven or more previous matings) as indicated by a higher proportion of sons in the offspring fathered by these males. Sperm limitation correlated with clearly reduced pheromone titres. In behavioural bioassays, females oriented towards higher doses of the synthetic pheromone and were attracted more often to scent marks of males with a full sperm load than to those of sperm-limited males. Our data support the PLFH and suggest that N. vitripennis females are able to decrease the risk of getting constrained to produce suboptimal offspring sex ratios by orienting towards gradients of the male sex pheromone.

Dangerously few liaisons: a review of mate‐finding Allee effects

AbstractIn this paper, we review mate‐finding Allee effects from ecological and evolutionary points of view. We define ‘mate‐finding’ as mate searching in mobile animals, and also as the meeting of gametes for sessile animals and plants (pollination). We consider related issues such as mate quality and choice, sperm limitation and physiological stimulation of reproduction by conspecifics, as well as discussing the role of demographic stochasticity in generating mate‐finding Allee effects. We consider the role of component Allee effects due to mate‐finding in generating demographic Allee effects (at the population level). Compelling evidence for demographic Allee effects due to mate‐finding (as well as via other mechanisms) is still limited, due to difficulties in censusing rare populations or a failure to identify underlying mechanisms, but also because of fitness trade‐offs, population spatial structure and metapopulation dynamics, and because the strength of component Allee effects may vary in time and space. Mate‐finding Allee effects act on individual fitness and are thus susceptible to change via natural selection. We believe it is useful to distinguish two routes by which evolution can act to mitigate mate‐finding Allee effects. The first is evolution of characteristics such as calls, pheromones, hermaphroditism, etc. which make mate‐finding more efficient at low density, thus eliminating the Allee effect. Such adaptations are very abundant in the natural world, and may have arisen to avoid Allee effects, although other hypotheses are also possible. The second route is to avoid low density via adaptations such as permanent or periodic aggregation. In this case, the Allee effect is still present, but its effects are avoided. These two strategies may have different consequences in a world where many populations are being artificially reduced to low density: in the first case, population growth rate can be maintained, while in the second case, the mechanism to avoid Allee effects has been destroyed. It is therefore in these latter populations that we predict the greatest evidence for mate‐finding Allee effects and associated demographic consequences. This idea is supported by the existing empirical evidence for demographic Allee effects. Given a strong effect that mate‐finding appears to have on individual fitness, we support the continuing quest to find connections between component mate‐finding Allee effects (individual reproductive fitness) and the demographic consequences. There are many reasons why such studies are difficult, but it is important, particularly given the increasing number of populations and species of conservation concern, that the ecological community understands more about how widespread demographic Allee effects really are, and why.

In vitro fertilization experiments using sockeye salmon reveal that bigger eggs are more fertilizable under sperm limitation.

Although theory and widespread evidence show that the evolution of egg size is driven primarily by offspring and maternal fitness demands, an additional explanation invokes sperm limitation as a selective force that could also influence egg size optima. Levitan proposed that constraints from gamete encounter in external fertilization environments could select for enlargement of ova to increase the physical size of the fertilization target. We test this theory using in vitro fertilization experiments in an externally fertilizing fish. Sockeye salmon (Onchorhyncus nerka) females show considerable between-individual variation in ovum size, and we explored the consequences of this natural variation for the fertilization success of individual eggs under conditions of sperm limitation. By engineering consistent conditions where in vitro fertilization rate was always intermediate, we were able to compare the sizes of fertilized and unfertilized eggs across 20 fertilization replicates. After controlling for any changes in volume through incubation, results showed that successfully fertilized eggs were significantly larger than the eggs that failed to achieve fertilization. Under conditions without sperm limitation, fertility was unaffected by egg size. Our findings therefore support Levitan's theory, demonstrating empirically that some element of egg size variation could be selected by fertilization demands under sperm limitation. However, further research on sperm limitation in natural spawnings is required to assess the selective importance of these results.

How to go extinct by mating too much: population consequences of male mate choice and efficiency in a sexual–asexual species complex

Selection acting on individuals is not predicted to maximize population persistence, yet examples that explicitly quantify conflicts between individual and population level benefits are scarce. One such conflict occurs over sexual reproduction because of the cost of sex: sexual populations that suffer the cost of producing males have only half the growth rate compared to asexuals. Male behaviour can additionally impact population dynamics in a variety of ways, and here we study an example where the impact is unusually clear: the riddle of persistence of sperm‐dependent sexual–asexual species complexes. Here, a sexually reproducing host species coexists with an ameiotically reproducing all‐female sperm parasite. Sexual–asexual coexistence should not be stable because the proportion of asexually reproducing females will rapidly increase and the relative abundance of the sexually reproducing host species will decline. A severe shortage of males will lead to sperm limitation for sexual and asexual females and the system collapses. Male mate choice could reduce the reproductive potential of the asexual species and thus potentially prevent the collapse. In the gynogenetic (sperm‐dependent parthenogenetic) Amazon molly Poecilia formosa and its host (P. latipinna or P. mexicana), males discriminate against asexual females to some extent. Using a population‐dynamical model, we examine the population dynamics of this species complex with varying strengths of male discrimination ability and efficiency with which they locate females and produce sperm. The sexual species would benefit from stronger discrimination, thus preventing being displaced by the asexual females. However, males would be required to evolve preferences that are probably too strong to be purely based upon selection acting on individuals. We conclude that male behaviour does not fully prevent but delays extinction, yet this is highly relevant because low local extinction rates strongly promote coexistence as a metapopulation.

Sperm depletion as a consequence of increased sperm competition risk in the European bitterling, Rhodeus amarus

Sperm depletion occurs when males are unable to maintain ejaculate size over successive ejaculations and appears to be widespread across multiple taxa. We used the European bitterling, a fish that lays its eggs in the gills of freshwater mussels, in experiments to investigate sperm depletion over different temporal scales at different levels of sperm competition risk. The spermatozoa density of territorial male ejaculates declined over five consecutive ejaculations, but mean ejaculate size was significantly larger if the male was exposed to a rival, suggesting males were sensitive to sperm competition risk and modulated ejaculate size in accordance with theoretical models. However, following successive matings over the course of a day, males exposed to rivals produced significantly smaller ejaculates than males with a low risk of sperm competition. Results are discussed with reference to a possible intersexual conflict over the size and distribution of ejaculates in the context of male sperm depletion and female sperm limitation.

Reproductive biology and implications for management of the orange‐spotted grouper Epinephelus coioides in the southern Arabian Gulf

The reproductive biology of Epinephelus coioides was determined from the examination of 1455 individuals collected between July 2005 and June 2007 in the southern Arabian Gulf. Histological preparations of gonads indicated that males were either derived from a juvenile phase or the transition of postspawning females, confirming a diandric protogynous sexual pattern. The spawning season was well defined, occurring once a year during April and early May. Peaks in spawning occurred after the full and new moons and was completed within a single lunar cycle. The presence of mature males over the entire size and age range and the absence of inactive mature females during the spawning season suggested that the population was not constrained by sperm limitation. While specimens undergoing sexual transition were only observed in size and age ranges of 335-685 mm total length (L(T)) and 5-6 years, patterns in the proportion of males in size and age classes suggested that sex change occurred at a relatively constant rate after female maturation up to the maximum size (1002 mm L(T)) and age (11 years). Relationships between reproductive output and capacity with size and age indicated that conventional regulations that equate the mean size at first capture to sexual maturation are unsuitable for the management of E. coioides. The maximum age, small size and young age at sexual maturation (L(min)= 320 mm L(T), 2 years, for females and 242 mm L(T), 1 year, for males) conflict with the general pattern for large epinepheline groupers and may be a direct result of the intensive demersal fishery in the southern Arabian Gulf.

Multiple mating in the ant Cataglyphis cursor: testing the sperm limitation and the diploid male load hypotheses

Multiple mating (i.e., polyandry) by queens in social Hymenoptera is expected to weaken social cohesion since it lowers within-colony relatedness, and hence, indirect fitness benefits from kin selection. Yet, there are many species where queens mate multiply. Several hypotheses have been put forward to explain the evolution and maintenance of polyandry. Here,we investigated the ‘sperm limitation’ and the ‘diploid male load’ hypotheses in the ant Cataglyphis cursor. Genetic analyses of mother-offspring combinations showed that queens mate with up to 8 males, with an effective mating frequency of 3.79. Significant paternity skew (unequal contribution of the fathers) was detected in 1 out of 5 colonies. The amount of sperm stored in the spermatheca was not correlated with the queen mating frequency, and males carry on average enough sperm in their seminal vesicles to fill one queen’s spermatheca. Analyses of the nuclear DNA-content of males also revealed that all were haploid. These results suggest that the ‘sperm limitation’ and the ‘diploid male load’ hypotheses are unlikely to account for the queen mating frequency reported in this ant. In light of our results and the life-history traits of C. cursor, we discuss alternative hypotheses to account for the adaptive significance of multiple mating by queens in this species.

Male reproductive success during spawning events of the octocoral Pseudopterogorgia elisabethae

Many sessile organisms rely on the transport of one or both types of gametes for repro- ductive success. While sperm limitation has been reported in some species, high fertilization success has been observed in others including the branching octocoral Pseudopterogorgia elisabethae. In such cases, males may compete for eggs to fertilize and the local dispersion of sperm will be an important component of male success. Individual reproductive success of 91 male P. elisabethae colonies found within a 400 m 2 area on San Salvador Island, Bahamas was examined based on microsatellite analysis of planulae collected from female colonies. Larvae which were brooded on the surface of 4 different females were collected during 2 separate spawning events. Males in the study area were sampled, and parentage was assessed using 5 microsatellite loci and the paternity assign- ment program CERVUS 3.0. Twenty-five percent of the larvae were assigned sires with ≥80% likeli- hood and 87% of the planulae could have been sired by males within the study area. The success of males was not related to either distance to the female or to male colony size. On average, successful sperm only travelled 5.2 m and the success of individual males within the 400 m 2 study site was not significantly different from random expectations. Male P. elisabethae may follow spawning strategies that increase the likelihood of siring some planulae, spread success across multiple females and reduce variance in reproductive success.

Gamete plasticity in a broadcast spawning marine invertebrate

Sperm competition has classically been thought to maintain anisogamy (large eggs and smaller sperm) because males are thought to maximize their chance of winning fertilizations by trading sperm size for number. More recently it has been recognized that sperm quality (e.g., size, velocity) can also influence sperm competition, although studies have yielded conflicting results. Because sex evolved in the sea, debate has continued over the role of sperm competition and sperm environment in determining both sperm and egg size in externally fertilizing broadcast spawners. Remarkably, however, there have been no direct tests of whether broadcast spawners change the traits of their gametes depending on the likelihood of sperm competition. We manipulated the density (and thus, sperm environment) of a broadcast spawning ascidian (Styela plicata) in the field and then determined whether the phenotype of eggs and sperm changed. We found that sperm from adults kept at high density were larger and more motile than sperm from low-density adults. In vitro fertilizations revealed that sperm from high-density adults also lived longer and induced less polyspermy. Adult density also affected egg traits: eggs from high-density adults were smaller targets for sperm overall but produced larger ovicells than eggs from low-density adults. This suggests that broadcast spawning mothers balance (potentially conflicting) pre- and postzygotic selection pressures on egg size. Overall, our results suggest that sperm competition does not represent a strong force maintaining anisogamy in broadcast spawners. Instead, sperm limitation seems to select for large eggs and smaller, more numerous sperm.

Male Mating Constraints Affect Mutual Mate Choice: Prudent Male Courting and Sperm‐Limited Females

Costs of sperm production may lead to prudence in male sperm allocation and also to male mate choice. Here, we develop a life history-based mutual mate choice model that takes into account the lost-opportunity costs for males from time out in sperm recovery and lets mate competition be determined by the prevailing mate choice strategies. We assume that high mating rate may potentially lead to sperm depletion in males, and that as a result, female reproduction may be limited by the availability of sperm. Increasing variation in male quality leads, in general, to increased selective mate choice by females, and vice versa. Lower-quality males may, however, gain access to more fecund higher-quality females by lowering their courting rate, thus increasing their sperm reserves. When faced with strong male competition for mates, low-quality males become less choosy, which leads to assortative mating for quality and an increased mating rate across all males. With assortative mating, the frequency of antagonistic interactions (sexual conflict) is reduced, allowing males to lower the time spent replenishing sperm reserves in order to increase mating rate. This in turn leads to lower sperm levels at mating and therefore could lead to negative effects on female fitness via sperm limitation.

The significance of intercolonial variation of cuticular hydrocarbons for inbreeding avoidance in ant sexuals

The reproductive biology of social Hymenoptera is characterized by lifelong sperm storage by queens and sperm limitation in males. Both sexes are therefore expected to be especially choosy about their mating partners. In particular, sexuals should avoid sibmating because of the resulting risk of producing sterile diploid male offspring. Colonies of the ant Leptothorax gredleri are specialists of patchy habitats and often live in subpopulations of fewer than a few dozen colonies with restricted gene flow between patches. Sexuals therefore have a high probability of mating with a related partner. Using gas chromatography and mass spectrometry, we found that the blend of cuticular hydrocarbons, which is generally thought to be important in nestmate recognition in social insects, was colony specific in both sexes. In principle, this might provide sexuals with a chemical cue for the avoidance of sibmating. When allowed to choose among equal numbers of mating partners from their own and another nest in flight cages, sexuals had only a weak although significant preference for mating with non-nestmates. Other characters might therefore be more important than relatedness in mate choice.

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (Poecilia mexicana, P. formosa)

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.

GAMETE TRAITS INFLUENCE THE VARIANCE IN REPRODUCTIVE SUCCESS, THE INTENSITY OF SEXUAL SELECTION, AND THE OUTCOME OF SEXUAL CONFLICT AMONG CONGENERIC SEA URCHINS

Sea-urchin species differ in susceptibility to sperm limitation and polyspermy, but the influences of gamete traits on reproductive variance, sexual selection, and sexual conflict are unknown. I compared male and female reproductive success of two congeners at natural densities in the sea. The eggs of the species occurring at higher densities, Strongylocentrotus purpuratus, require higher sperm concentrations for fertilization but are more resistant to polyspermy compared to S. franciscanus. Both species show high variance in male fertilization success at all densities and high variance in female success at low densities, but they differ in female variance at high densities, where only S. franciscanus shows high female variance. The intensity of sexual selection based on Bateman gradients is high in males of both species, variable in S. franciscanus females, and low in S. purpuratus females. Strongylocentrotus franciscanus females experience sexual selection at low densities and sexual conflict at high densities. Strongylocentrotus purpuratus may rarely experience sperm limitation and may have evolved to ameliorate sexual conflict. This reduces the variance in female fertilization, providing females with more control over fertilization. Sperm availability influences sexual selection directly by determining sperm-egg encounter probabilities and indirectly through selection on gamete traits that alter reproductive variances.

Consequences of inbreeding depression due to sex-linked loci for the maintenance of males and outcrossing in branchiopod crustaceans

Androdioecy, where males co-occur with hermaphrodites, is a rare sexual system in plants and animals. It has a scattered phylogenetic distribution, but it is common and has persisted for long periods of evolutionary time in branchiopod crustaceans. An earlier model of the maintenance of males with hermaphrodites in this group, by Otto et al. (1993), considered the importance of male-hermaphrodite encounter rates, sperm limitation, male versus hermaphrodite viability and inbreeding depression suffered by selfed progeny. Here I advance this model in two ways: (1) by exploring the conditions that would allow the invasion of hermaphrodites into a dioecious population and that of females into an androdioecious population; and (2) by incorporating a term that accounts for the potential effects of genetic load linked to a dominant hermaphrodite-determining allele in androdioecious populations. The new model makes plausible sense of observations made in populations of the species Eulimnadia texana, one of a number of related species whose common ancestor evolved hermaphroditism (and androdioecy) from dioecy. In particular, it offers an explanation for the long evolutionary persistence of androdioecy in branchiopods and suggests reasons for why dioecy has not re-evolved in the clade. Finally, it provides a rather unusual illustration of the implications of the degeneration of loci linked to a sex-determining locus.

SIZE‐SELECTIVE HARVESTING ALTERS LIFE HISTORIES OF A TEMPERATE SEX‐CHANGING FISH

Selective mortality, whether caused naturally by predation or through the influence of harvest practices, initiates changes within populations when individuals possessing certain heritable traits have increased fitness. Theory predicts that increased mortality rates will select for changes in a number of different life history characteristics. For example, fishing often targets larger individuals and has been shown repeatedly to alter population size structure and growth rates, and the timing of maturation. For sex-changing species, selective fishing practices can affect additional traits such as the mature population sex ratio and the timing of sexual transformation. Using historical comparisons, we examined the effects of exploitation on life history characteristics of California sheephead, Semicossyphus pulcher, a temperate protogynous (female-male sex changer) labrid that inhabits nearshore rocky environments from central California, USA, to southern Baja California, Mexico. Recreational fishing intensified and an unregulated commercial live-fish fishery developed rapidly in southern California between the historical and current studies. Collections of S. pulcher from three locations (Bahia Tortugas, Catalina Island, and San Nicolas Island) in 1998 were compared with data collected 20-30 years previously to ascertain fishery-induced changes in life history traits. At Bahia Tortugas, where fishing by the artisanal community remained light and annual survivorship stayed high, we observed no changes in size structure or shifts in the timing of maturation or the timing of sex change. In contrast, where recreational (Catalina) and commercial (San Nicolas) fishing intensified and annual survivorship correspondingly declined, males and females shifted significantly to smaller body sizes, females matured earlier and changed sex into males at both smaller sizes and younger ages and appeared to have a reduced maximum lifespan. Mature sex ratios (female:male) increased at San Nicolas, despite a twofold reduction in the mean time spent as a mature female. Proper fisheries management requires measures to prevent sex ratio skew, sperm limitation, and reproductive failure because populations of sequential hermaphrodites are more sensitive to size-selective harvest than separate-sex species. This is especially true for S. pulcher, where different segments of the fishery (commercial vs. recreational) selectively target distinct sizes and therefore sexes in different locations.