Intraspecific divergence in sperm morphology of the green sea urchin, <Emphasis Type="Italic">Strongylocentrotus droebachiensis</Emphasis>: implications for selection in broadcast spawners

Background  

Sperm morphology can be highly variable among species, but less is known about patterns of population differentiation within species. Most studies of sperm morphometric variation are done in species with internal fertilization, where sexual selection can be mediated by complex mating behavior and the environment of the female reproductive tract. Far less is known about patterns of sperm evolution in broadcast spawners, where reproductive dynamics are largely carried out at the gametic level. We investigated variation in sperm morphology of a broadcast spawner, the green sea urchin (Strongylocentrotus droebachiensis), within and among spawnings of an individual, among individuals within a population, and among populations. We also examined population-level variation between two reproductive seasons for one population. We then compared among-population quantitative genetic divergence (Q _ST) for sperm characters to divergence at neutral microsatellite markers (F _ST).     

The effect of alternative mating tactics on the fertilization success of a hermaphroditic seabass

In the simultaneously hermaphroditic marine fish, Serranus subligarius, male role individuals are known to pair spawn, group spawn and streak spawn. While the effects of these common mating tactics on mating success in the male role have been well studied, their consequences for the reproductive success of the individuals taking the female role have received little attention. To investigate those consequences, I observed mating behaviors and quantified fertilization success in natural and experimental settings during the summers of 2005-2008 at three sites with different local population densities. I observed focal individuals in 15-min increments and recorded the total number of spawns, number of streak spawns, size of participating spawners, and fertilization rate. The occurrence of small-sized individuals in the local population is associated with higher frequencies of streaking behavior; these small fish are most often first-year individuals reaching sexual maturity late in the spawning season (August/September). Spawns that included one or more streak spawners had a significantly lower average fertilization rate (89?%) than pair spawns without a streak spawner (97?%). This pattern was confirmed with a field manipulation experiment in which spawning events that included streakers again showed lower fertilization rates (93?%) than spawning events that did not include streakers (98?%). These lower fertilization rates occurred despite the fact that spawns that included multiple males produced, on average, 20?% more sperm than produced in spawns with only a single male. These results indicate that females incur a significant fitness cost when streakers invade a spawning event, a cost not attributable to sperm limitation or any direct effects on the female.     

Fertilization in the sea: are the hazards of broadcast spawning avoided when free-spawned sperm fertilize retained eggs?

In broadcast-spawning marine animals, rapid dilution and short lifespan of sperm following release may impose severely localized patterns of mating. Partial or total failure of external fertilization due to sperm limitation appears commonplace. However, it is not clear to what extent the restrictive kinetics of fertilization in water also constrain mating in animals that release sperm but retain their eggs for fertilization.<BR>The compound ascidian Diplosoma listerianum liberates sperm that are dispersed to other colonies and taken in prior to internal cross-fertilization. The fertile lifespan of sperm was found to be long (half-life ca. 8 hours), and a substantial number of fertilizations occurred with 24-hour-old sperm. Fertilizations were obtained from sperm concentrations that would typically produce little or no external fertilization. In a separate experiment, a very small piece of D. listerianum (dry weight less than 2 mg) sired abundant progeny throughout a 3840 l tank. Paternity of progeny in these experiments was confirmed by molecular markers. The same markers were used to extend, to over seven weeks, the known maximum period of storage of exogenous sperm prior to fertilization in this species.<BR>The production of only a few thousand sperm at a time by each zooid, poor synchronization of release between zooids, and the existence of many well-spaced exhalant openings in large colonies suggest that D. listerianum is incapable of generating a dense plume of sperm, even close to the source. It is suggested that, unlike external fertilization, successful internal cross-fertilization in D. listerianum is not dependent upon the interception of a dense cloud of gametes just released by a near neighbour. It seems instead that dilute, long-lived sperm can be extracted efficiently from seawater by this suspension feeder, potentially over a period of time. This capability, and other features of the life history, make it unlikely that sperm limitation is an acute problem in this species and comparable taxa, a conclusion with potential significance for expected patterns of mating, sex allocation and gamete attributes in sessile aquatic invertebrates. Variance in reproductive success between individuals due to differences in fertilization rate may be much lower than in broadcast spawners exhibiting external fertilization.     

Releasing small ejaculates slowly increases per‐gamete fertilization success in an external fertilizer: Galeolaria caespitosa (Polychaeta: Serpulidae)

The idea that male reproductive strategies evolve primarily in response to sperm competition is almost axiomatic in evolutionary biology. However, externally fertilizing species, especially broadcast spawners, represent a large and taxonomically diverse group that have long challenged predictions from sperm competition theory—broadcast spawning males often release sperm slowly, with weak resource‐dependent allocation to ejaculates despite massive investment in gonads. One possible explanation for these counter‐intuitive patterns is that male broadcast spawners experience strong natural selection from the external environment during sperm dispersal. Using a manipulative experiment, we examine how male reproductive success in the absence of sperm competition varies with ejaculate size and rate of sperm release, in the broadcast spawning marine invertebrate Galeolaria caespitosa (Polychaeta: Serpulidae). We find that the benefits of Fast or Slow sperm release depend strongly on ejaculate size, but also that the per‐gamete fertilization rate decreases precipitously with ejaculate size. Overall, these results suggest that, if males can facultatively adjust ejaculate size, they should slowly release small amounts of sperm. Recent theory for broadcast spawners predicts that sperm competition can also select for Slow release rates. Taken together, our results and theory suggest that selection often favours Slow ejaculate release rates whether males experience sperm competition or not.     

Density-dependent sexual selection in external fertilizers: variances in male and female fertilization success along the continuum from sperm limitation to sexual conflict in the sea urchin Strongylocentrotus franciscanus

Sperm competition and female choice are fundamentally driven by gender differences in investment per offspring and are often manifested as differences in variance in reproductive success: males compete and have high variance; most females are mated and have low variance. In marine organisms that broadcast spawn, however, females may encounter either sperm limitation or sperm competition. I measured the fertilization success of male and female Strongylocentrotus franciscanus over a range of population densities using microsatellite markers. Female fertilization success first increased and then decreased with mate density, limited at low density by sperm limitation and at high density by polyspermy. Mate density affected variance in fertilization success in both males and females. In males, the variance in fertilization success increased with mate density. In females, the pattern was more complex. The variance in female success increased similarly to males with increased mate density but then decreased to low levels at intermediate densities, where almost all eggs were fertilized. As density increased further, the female variances again increased as polyspermy lowered average fertilization success. Male and female variances differed only at intermediate densities. At low densities, both sexes may be under selection to increase fertilization success; at intermediate densities, males may compete; and at high densities, both sexes may be under selection to increase success by increasing (males) or decreasing (females) likelihood of fertilization during sexual conflict. Only within a narrow range of densities do patterns of sexual selection mirror those typically noted in internally fertilizing taxa.     

A theoretical investigation of sympatric evolution of temporal reproductive isolation as illustrated by marine broadcast spawners

Recent theory suggests that frequency-dependent disruptive selection in combination with assortative mating can lead to the establishment of reproductive isolation in sympatry. Here we explore how temporal variation in reproduction might simultaneously generate both disruptive selection and assortative mating, and result in sympatric speciation. The conceptual framework of the model may be applicable to biological systems with negative frequency-dependent selection, such as marine broadcast spawners or systems with pollinator limitation. We present a model that is motivated by recent findings in marine broadcast spawners and is parameterized with data from the Montastraea annularis species complex. Broadcast spawners reproduce via external fertilization and synchronous spawning is required to increase the probability of successful fertilization, but empirical evidence shows that as density increases, so does the risk of polyspermy. Polyspermy is the fusion of multiple sperm with an egg at fertilization, a process that makes the embryo unviable. Synchrony can therefore also act as a source of negative density-dependent disruptive selection. Model analysis shows that the interaction between polyspermy and spawning synchrony can lead to temporal reproductive isolation in sympatry and that, more generally, increased density promotes maintenance of genetic variation.     

The evolution of gonad expenditure and gonadosomatic index (GSI) in male and female broadcast‐spawning invertebrates

Sedentary broadcast‐spawning marine invertebrates, which release both eggs and sperm into the water for fertilization, are of special interest for sexual selection studies. They provide unique insight into the early stages of the evolutionary succession leading to the often‐intense operation of both pre‐ and post‐mating sexual selection in mobile gonochorists. Since they are sessile or only weakly mobile, adults can interact only to a limited extent with other adults and with their own fertilized offspring. They are consequently subject mainly to selection on gamete production and gamete success, and so high gonad expenditure is expected in both sexes. We review literature on gonadosomatic index (GSI; the proportion of body tissue devoted to gamete production) of gonochoristic broadcast spawners, which we use as a proxy for gonad expenditure. We show that such taxa most often have a high GSI that is approximately equal in both sexes. When GSI is asymmetric, female GSI usually exceeds male GSI, at least in echinoderms (the majority of species recorded). Intriguingly, though, higher male GSI also occurs in some species and appears more common than female‐biased GSI in certain orders of gastropod molluscs. Our limited data also suggest that higher male GSI may be the prevalent pattern in sperm casters (where only males release gametes). We explore how selection might have shaped these patterns using game theoretic models for gonad expenditure that consider possible trade‐offs with (i) somatic maintenance or (ii) growth, while also considering sperm competition, sperm limitation, and polyspermy. Our models of the trade‐off between somatic tissue (which increases survival) and gonad (which increases reproductive success) predict that GSI should be equal for the two sexes when sperm competition is intense, as is probably common in broadcast spawners due to synchronous spawning in aggregations. Higher female GSI occurs under low sperm competition. Sperm limitation appears unlikely to alter these conclusions qualitatively, but can also act as a force to keep male GSI high, and close to that of females. Polyspermy can act to reduce male GSI. Higher male than female GSI is predicted to be less common (as observed in the data), but can occur when ova/ovaries are sufficiently more resource‐intensive to produce than sperm/testes, for which some evidence exists. We also show that sex‐specific trade‐offs between gonads and growth can generate different life‐history strategies for males and females, with males beginning reproduction earlier. This could lead to apparently higher male GSI in empirical studies if immature females are included in calculations of mean GSI. The existence of higher male GSI nonetheless remains somewhat problematic and requires further investigation. When sperm limitation is low, we suggest that the natural logarithm of the male/female GSI ratio may be a suitable index for sperm competition level in broadcast spawners, and that this may also be considered as an index for internally fertilizing taxa.     

THE QUICK AND THE DEAD? SPERM COMPETITION AND SEXUAL CONFLICT IN SEA

Our view of sperm competition is largely shaped by game-theoretic models based on external fertilizers. External fertilization is of particular interest as it is the ancestral mode of reproduction and as such, relevant to the evolution and maintenance of anisogamy (i.e., large eggs and tiny, numerous sperm). Current game-theoretic models have been invaluable in generating predictions of male responses to sperm competition in a range of internal fertilizers but these models are less relevant to marine broadcast spawners, the most common and archetypal external fertilizers. Broadcast spawners typically have incomplete fertilization due to sperm limitation and/or polyspermy (too many sperm), but the effects of incomplete (<100% fertilization rates) fertilization on game-theoretic predictions are unclear particular with regards to polyspermy. We show that incorporating the effects of sperm concentration on fertilization success changes the predictions of a classic game-theoretic model, dramatically reversing the relationship between sperm competition and the evolutionarily stable sperm release strategy. Furthermore, our results suggest that male and female broadcast spawners are likely to be in conflict at both ends of the sperm environment continuum rather than only in conditions of excess sperm as previously thought. Across the majority of the parameter space we explored, males release either too little to too much sperm for females to achieve complete fertilization. This conflict could result in a coevolutionary race that may have led to the evolution of internal fertilization in marine organisms.     

Components of fertilization success in the bluehead wrasse, Thalassoma bifasciatum

In many species of marine organisms, males and females releasegametes directly into the water column. Although free-spawningmarine invertebrates appear to have highly variable fertilizationsuccess, in tropical reef fishes the average fertilizationsuccess is quite high, typically over 90%; nevertheless, substantialvariation has been reported, and fertilization has a directeffect on fitness. We investigated the factors affecting fertilization success in natural spawnings of the bluehead wrasse, Thalassoma bifasciatum. During a two-year study at a site in St. Croix,we found extensive and predictable variation in fertilizationsuccess in pair spawns of this reef fish. Fertilization successaveraged 95%, but was affected by the amount of sperm released,the water velocity at a site, the mating success of the male,and the size of the female. As sperm released in a spawn increases, and as water velocity at a site decreases, sperm concentrationsshould remain higher in the vicinity of eggs for a longer periodof time, and both of these factors are correlated with increasingfertilization success. The recent history of individuals withpartners or sites did not affect the fertilization successof their spawn. In an evolutionary context, the real and predictable variance in fertilization success in this species may influencethe mating choices of males and females. However, there iscurrently no evidence that females use differences in fertilizationsuccess among males or sites in their reproductive decisions.     

Female control of sperm ejection and retention in the cornsilk fly Euxesta eluta (Diptera: Ulidiidae)

Promiscuous mating systems provide the opportunity for females to bias fertilization toward particular males. However, distinguishing between male sperm competition and active female sperm choice is difficult for species with internal fertilization. Nevertheless, species that store and use sperm of different males in different storing structures and species where females are able to expel all or part of the ejaculates after copulation may be able to bias fertilization. We report a series of experiments aimed at providing evidence of female sperm choice in Euxesta eluta (Hendel), a species of ulidiid fly that expels and consumes ejaculates after copulation. We found no evidence of greater reproductive success for females mated singly, multiply with the same male, or mated multiply with different males. Female E. eluta possesses two spherical spermathecae and a bursa copulatrix for sperm storage, with a ventral receptacle. There was no significant difference in storing more sperm in spermathecae 24 h after copulation than immediately after copulation. Females mated with protein-fed males had greater reproductive success than similar females mated to protein-deprived males. Protein-fed females prevented to consume the ejaculate, retained more sperm when mated to protein-fed males than when mated to protein-deprived males. Our results suggest that female E. eluta can exert control of sperm retention of higher quality males through ejaculate ejection.     

Quantitative analysis of sperm plane circular movement in the blue mussels Mytilus edulis, M. trossulus and their hybrids

Fertilization success of free spawning organisms such as Mytilus species depends on gamete interactions. Therefore, gamete traits such as sperm movement are important for determining fertilization success in free spawning organisms. Since little is known about sperm movement pattern in Mytilus species, the purpose of this study was to investigate sperm movement pattern of blue mussel M. edulis, M. trossulus and their hybrids using computer-assisted sperm movement video analysis. Sperm of all genotypes were found to conduct circular movement in a two-dimensional plane. Furthermore, new sperm movement parameters, real time radius (R), angle change rate (θ) and the center of circular track (O(t)) were developed to verify and quantitatively describe the plane circular movement pattern using software (Image-J) that may be widely applied to sperm movement study in other organisms. Angle change rate was positively correlated to fertilization success. However, no correlation between fertilization and real time radius was detected. Although no interspecific differences were found in the radius, the F1 (first generation) hybrid sperm had a lower angle change rate than M. edulis and M. trossulus. Published studies have shown that sperm circular movement is more prevalent in aquatic broadcast spawning species than in species with mating behavior or internal fertilization. Therefore, a two-dimensional circular movement pattern in sperm may represent a trait that increases fertilization success for broadcast spawning species by either increasing gamete interaction rate at a small scale and/or avoiding swimming further away from the eggs before sperm detects the chemoattractant gradient.     

Germ-cell warfare in ascidians: sperm from one species can interfere with the fertilization of a second species

Ascidians (invertebrate chordates) are very abundant in many marine subtidal areas. They often live in dense multispecies clumps; thus, interspecific competition for space may be intense. Although most noncolonial species are broadcast spawners, their eggs can be fertilized only by sperm of the same species (1). Multiple fertilization is lethal and all animals have evolved blocks to polyspermy. Ascidian eggs block polyspermy by enzymatic (2) and electrical mechanisms (3). Sperm bind to N-acetylglucosamine groups on the vitelline coat (4, 5, 6, 7). Follice cells surrounding the vitelline coat release N-acetylglucosaminidase during egg activation (8), preventing the binding of all sperm but a few (2). I show here that this interaction is not species-specific; sperm from one species can cause glycosidase release from follicle cells of a second species. Furthermore, once glycosidase release has been induced, the subsequent addition of sperm from the egg-producing species fails to fertilize a substantial proportion of these eggs. This leads to the hypothesis that sperm from one species of ascidian can interfere with fertilization of a second species. While intraspecific sperm competition has been well documented in several taxa (9, 10), this is the first record of sperm competition between species, or interspecific sperm competition.     

Introgression despite substantial divergence in a broadcast spawning marine invertebrate

Understanding the relationship between reproductive isolation and time since divergence is critical to our understanding of speciation. One group for which we know little about the relationship between hybridization/introgression and time since divergence is the marine broadcast spawners. Here, we investigate the distribution of closely related cryptic species of marine broadcast spawners (Type A and B Ciona intestinalis) in areas of potential sympatry to determine whether these two types occur together and if so, whether they show evidence of hybridization and introgression. Then we combine our data with other studies to investigate general patterns of reproductive isolation versus divergence in marine broadcast spawners. We found that Type A and B C. intestinalis occurred sympatrically in 2007, and that 21 individuals show evidence of introgression in sympatry (out of approximately 500). Type A and B C. intestinalis are 12.4% divergent at mitochondrial COI (mtCOI), and in comparison with other marine broadcast spawning species at mtCOI, these two types may be near the upper limit of the range of divergence values in which introgression is still possible. However, introgression at divergence levels similar to those found in Ciona does exist, prompting questions about the strength of postmating prezygotic reproductive barriers in marine broadcast spawners.     

Sperm competition affects male behaviour and sperm output in the rainbow darter

Rainbow darters, Etheostoma caeruleum, are promiscuous fish with moderate rates of group spawning (between one and five males may simultaneously mate with one female). In this study, I examined male sperm output and male willingness to spawn under different levels of sperm competition intensity. One male and one female were allowed to spawn in an aquarium where they had visual and olfactory access to one of four treatments: four males, one male, zero males, or one female. Theory predicts that males should reduce sperm output when there are more than the average number of males at a group spawning (four-male treatment) and should increase sperm output when there are fewer than average males at a group spawning (one-male treatment). Mean sperm output did not differ among treatments. However, males released more sperm when spawning in the presence of competing males (four-male and one-male treatments pooled) than when spawning in the absence of competing males (zero-male and one-female treatments pooled). Males were also most likely to forego spawning opportunities when sperm competition intensity was high. Furthermore, male willingness to spawn was size dependent. Large males were more likely to forego spawning opportunities under high sperm competition intensity. Large males may be better off waiting for future spawning opportunities when there is a lower potential for sperm competition intensity.     

An advantage for young sperm in the house cricket Acheta domesticus

We show that males of the house cricket Acheta domesticus regularly expel sperm packages (spermatophores) independently of copulation and at a rate that is not affected by the presence of females. We then show for the first time that the age of sperm affects their likelihood of being stored by females after copulation; younger sperm were overrepresented in the female sperm storage organ and therefore in the sperm population used for fertilization. Our results suggest that the reproductive success of males may increase if they deliver ejaculates with young sperm, and the results may explain why the males of several species are regularly observed to discard ejaculates. Our results also suggest that phenomena such as female multiple mating, paternity bias, and/or exaggerated ejaculate sizes may be related to the advantage both genders gain by using young sperm.     

INTRASPECIFIC VARIATION IN SEX ALLOCATION IN A SIMULTANEOUS HERMAPHRODITE: THE EFFECT OF INDIVIDUAL SIZE

Within a population of simultaneous hermaphrodites, individuals may vary in both their current reproductive investment (biomass invested in gonads) and in how they allocate that investment between male and female function. In the chalk bass, Serranus tortugarum, estimates of both reproductive allocation and reproductive success as a male and a female can be made for individuals of different sizes. As individuals increase in size, their investment in gamete production increases, and there is a shift in allocation to a stronger female bias. Spawning frequency as a female in pair spawnings and as a male in both pair spawning and streaking (an alternative mating tactic) does not vary with individual size. As a result, larger individuals should release more sperm or eggs per spawn. Size-assortative pair spawning in this species leads to larger individuals having higher potential returns in total male reproductive success than smaller individuals, which should lead to increases in absolute levels of sperm production in larger individuals when individuals compete for fertilizations through sperm competition. However, smaller individuals contribute a smaller proportion of the sperm released in spawns with multiple spawners and thus are under more intense sperm competition than larger individuals, which should select for increases in male allocation in smaller individuals, all else equal. A local-mate-competition (LMC) model predicts that these factors select for increasing absolute male and female investment with individual size but a relative shift to more female-biased allocation as individual size increases. These predictions are supported by gonadal data. The predictions of average male allocation from the quantitative LMC model were 21.6% and 25.7%, whereas the collections averaged 21.3%. This close agreement of both the mean male allocation and its relative shift with individual size between model and data support the hypothesis that size-specific shifts in sex allocation in this species represent an adaptive response to patterns of mating success and sperm competition.     

Elevated pCO2 increases sperm limitation and risk of polyspermy in the red sea urchin Strongylocentrotus franciscanus

Anthropogenic carbon dioxide (CO₂) emissions and the resultant acidification of surface ocean waters are predicted to have far‐reaching consequences for biological processes in the marine environment. For example, because changes in pH and pCO₂ can alter sperm performance, ocean acidification may be accompanied by reductions in the success of fertilization in marine broadcast spawners. Several studies have attempted to determine the effects of elevated pCO₂ on marine invertebrate fertilization success, albeit with differing results. These conflicts may stem from the use of inappropriate sperm–egg contact times and, in several cases, the lack of measurements over a range of sperm concentrations extending from sperm‐limited conditions to polyspermy scenarios. In our study, we used biologically realistic sperm–egg contact times and a full range of sperm concentrations to assess the effect of elevated pCO₂ on fertilization in the broadcast spawning sea urchin, Strongylocentrotus franciscanus. Fertilization experiments were carried out in seawater bubbled with CO₂ to 400 (control), 800, and 1800 ppm. Using a fertilization kinetics model, we estimate that elevated pCO₂ levels both increased sperm limitation and reduced the efficiency of fast blocks to polyspermy. Thus, elevated pCO₂ decreased the range of sperm concentrations over which high fertilization success was likely. Given the inherent difficulties in achieving high fertilization success in broadcast spawners, raised pCO₂ levels are likely to exacerbate low fertilization success in low‐density populations or in areas with high water turbulence.     

Asymmetric conspecific sperm precedence in relation to spawning times in the Montastraea annularis species complex (Cnidaria: Scleractinia)

In broadcast spawners, prezygotic reproductive isolation depends on differences in the spatial and temporal patterns of gamete release and gametic incompatibility. Typically, gametic incompatibility is measured in no‐choice crosses, but conspecific sperm precedence (CSP) can prevent hybridization in gametes that are compatible in the absence of sperm competition. Broadcast spawning corals in the Montastraea annularis species complex spawn annually on the same few evenings. Montastraea franksi spawns an average of 110 min before M. annularis, with a minimum gap of approximately 40 min. Gametes are compatible in no‐choice heterospecific assays, but it is unknown whether eggs exhibit choice when in competition. Hybridization depends on either M. franksi eggs remaining unfertilized and in proximity to M. annularis when the latter species spawns or M. franksi sperm remaining in sufficient viable concentrations when M. annularis spawns. We found that the eggs of the early spawning M. franksi demonstrate strong CSP, whereas CSP appears to be lacking for M. annularis eggs. This study provides evidence of diverging gamete affinities between these recently separated species and suggests for the first time that selection may favour CSP in earlier spawning species when conspecific sperm is diluted and aged and is otherwise at a numeric and viability disadvantage with heterospecific sperm.     

Mechanisms of reproductive isolation among sympatric broadcast-spawning corals of the Montastraea annularis species complex

Many coral species spawn simultaneously and have compatible gametes, leading to controversy over the nature of species boundaries and the frequency with which hybridization occurs. Three western Atlantic corals, Montastraea annularis, M. faveolata, and M. franksi, typify this controversy; they all spawn sympatrically on the same evenings after the fall full moons. Here we show, in both Panama and the Bahamas for multiple years, how a variety of mechanisms may act in concert to reproductively isolate all three species. Field studies indicate that M. franksi spawns two hours earlier than the other two species, and the eggs released during this earlier period disperse an average of 500 m by the time the other species spawn. Field measures of fertilization indicate that peak fertilization occurs when spawning synchrony is high and that corals that spawn at the tails of the spawning distributions have greatly reduced fertilization success. Laboratory studies indicate that there is a gametic incompatibility between M. faveolata and the other two species. There are regional differences in the gametic compatibility of M. franksi and M. annularis. In Panama, the two species are completely compatible, whereas in the Bahamas, M. franksi sperm can fertilize M. annularis eggs but the reciprocal cross often fails. Gamete age influences patterns of fertilization, such that very young eggs seem resistant to fertilization and old sperm lose viability after two hours. In sum, the combination of temporal differences in spawning, sperm aging, gamete dispersal and dilution, and gametic incompatibility act in various combinations among the three species, making it unlikely that hybrid fertilization would occur.     

Scallop Size does not Predict Amount or Rate of Induced Sperm Release

Within populations of broadcast spawning marine invertebrates such as scallops, larger animals typically have larger gonads. Presumably, this means those larger males have more sperm to release than small males. However, there has never been a direct test of whether larger males actually release more sperm, at a higher rate, during spawning. To address this, we compared the allometry of induced sperm release with that of reproductive investment (gonad weight) in ripe males of 2 species of scallops, Chlamys bifrons and Chlamys asperrima. We did not find that larger scallops released more sperm or released it faster than small scallops, and were able to reject the hypothesis that instantaneous sperm release was related to body size in the same way as gonad weight. Consequently, we speculate that if larger broadcast spawning males do release more sperm, they may do so by spawning on more occasions within a reproductive season.