Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

Insights on the female reproductive system in Hippolyte inermis (Decapoda,Caridea): is this species really hermaphroditic?

Abstract. The majority of the published data about the reproductive biology of the decapod shrimp, Hippolyte inermis support the idea that this species is a protandric hermaphrodite, as is reported to be the case for certain other caridean species. However, our studies, based on the relative growth of the male reproductive appendage and histological examinations of the ovary, testes, oviducts, and deferentia vasa, indicate that there is no evidence supporting the occurrence of protandry. The first report of anomalies in the process of sexual inversion is credited to Reverberi (1950) ; however, we have obtained no evidence to support this phenomenon in H. inermis . We have not found either ovotestes or testes transforming into ovotestes. Therefore, we propose that H. inermis is a gonochoric species.     

Testing generalizations about latitudinal variation in reproduction and recruitment patterns with sicyoniid and caridean shrimp species

Summary

Study of latitudinal variation in seasonality of reproduction and recruitment of benthic marine invertebrates is useful in generating and testing hypotheses about causal factors acting on reproduction such as temperature and larval food supply that might be altered by changes in world climate. Analysis of latitudinal variation in reproductive patterns might be made with comparisons (a) among species with a common phylogenetic history from different latitudes and habitats and (b) among phylogenetically different taxa from the same location. Hypotheses on variation of reproductive seasonality with latitude are tested here with results of a study on nine species of caridean and two species of sicyoniid shrimp sampled from a tropical seagrass meadow in Puerto Rico. Breeding condition was determined by the presence or absence of incubated embryos (carideans) and the state of ovarian development in both carideans and sicyoniids. Recruitment was estimated from the percentage of individuals of monthly population samples in the juvenile size classes. Comparison of reproductive patterns among tropical, subtropical, and cool temperate Sicyonia spp. supports the paradigm of continuous reproduction in the tropics with increased restriction of breeding season with an increase in latitude. A greater intensity of breeding effort appears to accompany the shorter breeding period associated with an increase in latitude. At the tropical site most females of all caridean species carried embryos during all months of the year. With the onset of sexual maturity, caridean females produced consecutive broods for the rest of their relatively short (< 6 month) life span. In both sicyoniid and caridean species, recruitment occurred throughout the year but was highly variable, i.e., episodic rather than truly continuous or seasonal. Patterns of recruitment were highly concordant among but not between sicyoniid and caridean species, indicating that different sets of environmental factors controlled recruitment in the two groups. It is suggested that simultaneous study of adult reproduction and larval ecology is necessary to understand patterns of reproduction and recruitment. Coordinated effort on a global scale in studying latitudinal variation in reproduction and recruitment is suggested in order to predict the consequences of climate change on commercially and ecologically important marine invertebrate species.     

Proximate developmental mediators of sexual dimorphism in size: case studies from squamate reptiles

Sexual dimorphism in size (sexual size dimorphism; SSD) is nearlyubiquitous, but the relative importance of genetic versus environmentalcontrol of SSD is not known for most species. We investigatedproximate determinants of SSD in several species of squamatereptiles, including three species of Sceloporus lizards andthe diamond-backed rattlesnake (Crotalus atrox). In naturalpopulations of these species, SSD is caused by sexual differencesin age-specific growth. Males and females, however, may oftenshare similar potentials for growth: growth is strongly responsiveto the availability of food, and sexual differences in growthcan be greatly suppressed or completely absent under commonenvironmental conditions in the laboratory. Sexually divergentgrowth is expressed in natural environments because of inherentecological differences between males and females and becauseof potential epigenetic effects of sex-specific growth regulators.In field-active Sceloporus, sexual differences in growth rateare associated with sexual divergence in plasma testosterone.Experiments confirm that testosterone inhibits growth in speciesin which females are larger (for example, S. undulatus and S.virgatus) and stimulates growth in those in which males arelarger (for example, S. jarrovii). Interestingly, however, sexualdivergence in plasma testosterone is not accompanied by divergencein growth in S. jarrovii or in male-larger C. atrox in the laboratory.Furthermore, experimental effects of castration and testosteronereplacement on growth are not evident in captive S. jarrovii,possibly because growth effects of testosterone are supersededby an abundant, high-quality diet. In female-larger S. undulatus,growth may be traded-off against testosterone-induced reproductivecosts of activity. In male-larger species, costs of reproductionin terms of growth are suggested by supplemental feeding ofreproductive female C. atrox in their natural environment andby experimental manipulation of reproductive cost in femaleS. jarrovii. Growth costs of reproduction, however, do not contributesubstantially to the development of SSD in male-larger S. jarrovii.We conclude that the energetic costs of testosterone-induced,male reproductive behavior may contribute substantially to thedevelopment of SSD in some female-larger species. However, despitestrong evidence that reproductive investment exacts a substantialcost in growth, we do not support the reproductive cost hypothesisas a general explanation of SSD in male-larger species.     

Protandry models and their application to salmon

Mating systems characterized by restricted breeding seasons,male polygamy, and female monogamy are common among animals.In such systems (e.g., butterflies), the earlier emergenceof males than females to breeding areas (protandry) is a typicalphenological pattern. Protandry likely results from a timingstrategy that maximizes mating opportunities by males. In Pacific salmon (Oncorhynchus spp.), males typically arrive at the spawning grounds in advance of females. Using arrival-timing models,I found that under the mate-opportunity hypothesis, the matingsystem of salmon favors protandry. Protandry is predicted undera range of competitive scenarios, and the degree of protandryis especially sensitive to the duration of male spawning activity.Greater protandry is expected with increasing population sexratio (i.e., more males) when there is mate guarding, but lowerprotandry is expected with increasing population sex ratiowhen interference competition among males reduces male longevity.The timing of unequal competitors is expected to be similar,but among years, protandry may be less variable in the bettercompetitor.     

Testing three models on the adaptive significance of protandric simultaneous hermaphroditism in a marine shrimp

Protandric simultaneous hermaphroditism, as reported for shrimps in the genus Lysmata, is a sexual system in which individuals invariably reproduce as males first and later in life as simultaneous hermaphrodites. I tested three models (i.e., sex-dependent energetic costs, sex-dependent mortality rates and sex-dependent time commitments) in an attempt to explain the adaptive value of protandric simultaneous hermaphroditism in the shrimp L. wurdemanni. Specific assumptions and predictions of each model were evaluated using manipulative experiments. In the laboratory, males grew faster than simultaneous hermaphrodites of the same size and age, an indication that the female function incurs higher energetic costs of reproduction than the male function. Also, large SHPs were more successful in monopolizing food than small males. The sex-dependent growth rate and size-dependent resource holding power agree with predictions of the sex-dependent energetic cost model. The time that simultaneous hermaphrodites required for replenishing their sperm reservoirs after mating as males was much shorter (2 days) than the time required to brood one clutch of embryos (11 days). Also, small simultaneous hermaphrodites experienced heavier mortality due to predatory fishes than large ones. The sex-dependent reproductive time commitment and size-dependent mortality agree with predictions of the sex-dependent time commitment model. Conversely, I found no evidence that the sex-dependent mortality model explains protandric simultaneous hermaphroditism in the studied species. In contrast to model predictions, mortality due to predatory fishes suffered by simultaneous hermaphrodites was not greater than that suffered by males of the same body size. In L. wurdemanni, the relationship between sex-specific investment and reproductive success seems to change during ontogeny in a way that is consistent with an adaptive adjustment of sex allocation to improve age-specific reproductive success.     

Tests of the mate-guarding hypothesis for social monogamy: male snapping shrimp prefer to associate with high-value females

Social monogamy without biparental care has evolved in manytaxa, and a number of hypotheses have been developed to explainthis phenomenon. Several authors have suggested the importanceof male mate-guarding behavior in the evolution of social monogamy,although empirical support for this hypothesis is lacking. Inthe caridean shrimp genus Alpheus, social monogamy may resultfrom selection on males for long-term guarding of females becausemating is temporally restricted to a short time after the female'smolt. I used Alpheus angulatus to test two predictions of theextended mate-guarding hypothesis: Males should (1) be physiologicallycapable of predicting the timing of female sexual receptivity,and (2) prefer to associate with (guard) females that are closerto sexual receptivity. Data from a Y-maze experiment testingfor distance chemical communication showed that males of A.angulatus were attracted to water treated by exposure to premoltfemales, repulsed by water treated by exposure to intermoltmales and females, and did not appear to respond in either directionto water treated by exposure to premolt males. In mate choiceexperiments, significantly more males paired with premolt femalesthan with postmolt females. These data suggest that males ofA. angulatus engage in precopulatory mate-guarding behavior.Other factors (population density, sex ratio) may have playeda role in the temporal extension of mate guarding to socialmonogamy.     

Sex change in either direction by growth-rate advantage in the monogamous coral goby, Paragobiodon echinocephalus

The size-advantage model predicts the evolution of sex changeif the relative reproductive success of the sexes changes withsize or age. In the goby (Paragobiodon echinocephalus) the largesttwo fish, a male and a female matched by size, breed monogamouslyin each host coral. Because the female fecundity and male abilityof egg care increase with body size in a similar way, no size-fecundityadvantage exists. However, we found both protogyny and infrequentprotandry in a natural population of this species in Okinawa.New pairs were often formed after movement between host coralsand also sex change or sex differentiation of one or both members.In most new pairs males were larger than females, and femalesgrew much faster than their mates until breeding (growth-rateadvantage). The smaller member of a new pair should be the femalethat grows faster, because the smaller limits the reproductivesuccess of the pair. To form such a pair, the goby changed sexaccording to the sex and relative size of a new mate, as a status-dependentconditional strategy. The growth-rate advantage predicts predominanceof protogyny, but movement between host corals provides opportunitiesalso for protandry.     

Molecular systematics of peppermint and cleaner shrimps: phylogeny and taxonomy of the genera Lysmata and Exhippolysmata (Crustacea: Caridea: Hippolytidae)

Shrimps from the ecologically diverse genera Lysmata and Exhippolysmata are rare among marine invertebrates because they are protandric simultaneous hermaphrodites: shrimps initially mature and reproduce solely as males, and later in life become functional simultaneous hermaphrodites. Considerable progress on the reproductive ecology of members from these two genera has been achieved during the last decade. However, several outstanding issues of systematic nature remain to be addressed. Here, a molecular phylogeny of these two genera was used to examine the overall evolutionary relationship within and between species and genera, and to answer various questions related to the systematic status of several species. The present phylogenetic analysis, including 53 sequences and 26 species of Lysmata and Exhippolysmata, indicates that semiterrestrial shrimps from the genus Merguia represent the sister group to a second natural clade composed by shrimps from the genera Lysmata and Exhippolysmata. Also, the phylogenetic analysis confirmed that the genus Lysmata is paraphyletic, and includes the genus Exhippolysmata, as noted in a preliminary study. The tree partially supports the separation of species with or without a developed accessory branch into two different genera or subgenera (i.e. Lysmata and Hippolysmata having a well‐developed accessory branch, or not, respectively). The genetic distance between the cleaner shrimps Lysmata amboinensis and Lysmata grabhami was smaller than has been observed between other sister species. On the other hand, the topology of the tree indicates that these two entities are reciprocally monophyletic. Thus, this latter result, together with minor but constant differences in the colour pattern reported for these two entities, indicates that there is no reason to stop treating them as different valid species. This study enabled the long overdue resolution of standing taxonomic questions in shrimps from the genera Lysmata and Exhippolysmata. In the future, this phylogeny will help to reveal the conditions favouring the origins of several behavioural and morphological novelties in these unique shrimps. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 254–265.     

Protandry, sexual selection and climate change

Protandry refers to the earlier appearance of males before females at sites of reproduction. Sexual selection has been hypothesized to give rise to sex differences in benefits and costs of early arrival, thereby selecting for earlier appearance by the sex subject to more intense sexual selection. If sexual selection is more intense, there is a greater premium on early arrival among individuals of the chosen sex because of direct selection for earlier arrival. This hypothesis leads to the prediction that changes in the costs and benefits of early arrival related to changes in environmental conditions should particularly affect the sex that arrives first and hence the degree of protandry. I tested this hypothesis using the Barn Swallow Hirundo rustica. During 1971–2003, the degree of protandry increased significantly in a Danish population because males advanced arrival date while females did not. This earlier arrival by males compared with females was correlated with a significant increase by over 1.2 standard deviations in the length of the outermost tail feathers of males, a secondary sexual character, suggesting direct selection on both protandry and the secondary sexual character. Environmental conditions during spring migration in Northern Africa, as reflected by the normalized difference vegetation index, have deteriorated since 1984, resulting in increased mortality among males during spring migration, but not among females, and this deterioration of climatic conditions was positively correlated with an increasing degree of protandry. Likewise, an increase in April temperatures at the breeding grounds during recent decades is positively correlated with increased protandry, apparently because males can arrive earlier without increasing the fitness cost of early arrival. Local population size did not predict changes in arrival date. These findings suggest that rapid changes in climate can cause a change in degree of protandry and secondary sexual characters.     

Protandry and sexual dimorphism in trans-Saharan migratory birds

Earlier arrival to reproductive sites of males relative to females(protandry) is widespread among migratory organisms. Diversemechanisms have been proposed that may select for protandry,including competition for limiting resources (e.g., territories)or mates. In species with large variation in male reproductivesuccess, such as polygamous species and those with intense spermcompetition, early arriving males may accrue a fitness advantagebecause they acquire more mates or have larger chances of paternity.Comparative studies of birds have shown that sexual size dimorphism(SSD) is positively associated with the level of polygyny, whereasintense sperm competition is associated with sexual dichromatism(SD). Positive correlations between protandry and SSD or SDcan therefore be expected to exist across avian species. Becauselarge males are predicted to be better able to cope with adverseecological conditions early in the breeding season, selectionfor protandry, in turn, may have a correlated response on SSDamong migratory species breeding in boreal latitudes. Althoughprevious studies of birds have analyzed the association betweenSSD and protandry, none has analyzed SD in relation to protandry.Here we analyze the association between protandry during springmigration, SSD, and SD in 21 trans-Saharan monogamous migratorybird species. The difference in median migration dates betweenfemales and males, reflecting protandry, was positively associatedwith SD but not with SSD. Because dichromatism is positivelyrelated to sperm competition across species, present resultsare consistent with predictions derived from sexual selectionhypotheses for the evolution of protandry mediated by spermcompetition.     

Reproductive skew among males in a female-dominated mammalian society

The purpose of this study was to document patterns of reproductiveskew among male spotted hyenas (Crocuta crocuta), a speciesin which many normal mammalian sex roles are reversed. We usedpaternity determined from 12 microsatellite markers togetherwith demographic and behavioral data collected over 10 yearsfrom a free-living population to document relationships among reproductive success (RS), social rank, and dispersal statusof male hyenas. Our data suggest that dispersal status andlength of residence are the strongest determinants of RS. Natalmales comprise over 20% of the adult male population, yet theysire only 3% of cubs, whereas immigrants sire 97%. This reproductiveadvantage to immigrants accrues despite the fact that immigrants are socially subordinate to all adult natal males, and it providesa compelling ultimate explanation for primary dispersal inthis species. High-ranking immigrants do not monopolize reproduction,and tenure accounts for more of the variance in male reproductivesuccess than does social rank. Immigrant male hyenas rarelyfight among themselves, so combat between rivals may be a relativelyineffectual mode of sexual selection in this species. Instead,female choice of mates appears to play an important role in determining patterns of paternity in Crocuta. Our data supporta "limited control" model of reproductive skew in this species,in which female choice may play a more important role in limitingcontrol by dominant males than do power struggles among males.     

Sexual selection and mating patterns in a mammal with female-biased sexual size dimorphism

In mammals, species with highly male-biased sexual size dimorphismtend to have high variance in male reproductive success. However,little information is available on patterns of sexual selection,variation in male and female reproductive success, and bodysize and mating success in species with female-biased size dimorphism.We used parentage data from microsatellite DNA loci to examinethese issues in the yellow-pine chipmunk (Tamias amoenus), asmall ground squirrel with female-biased sexual size dimorphism.Chipmunks were monitored over 3 years in the Kananaskis Valley,Alberta, Canada. We found evidence of high levels of multiplepaternity within litters. Variation in male and female reproductivesuccess was equal, and the opportunity for sexual selectionwas only marginally higher in males than females. Male and femalereproductive success both depended on mating success. We foundno evidence that the number of genetic mates a male had dependedon body size. Our results are consistent with a promiscuousmating system in which males and female mate with multiple partners.Low variation in male reproductive success may be a generalfeature of mammalian species in which females are larger thanmales.     

Reproductive effort and costs of reproduction do not explain female-biased sex ratios in the moss Pseudocalliergon trifarium (Amblystegiaceae)

A fundamental assumption in life-history theory is that reproduction is costly. Higher reproductive investment for fruits than for flowers may result in larger costs of reproduction in females than in males, which is often used to explain male-skewed sex ratios in unisexual seed plants. In contrast, bryophytes have predominantly female-biased sex ratios, suggested to be a product of a higher average cost of sexual reproduction in males. Empirical evidence to support this notion is largely lacking. We investigated sex-specific reproductive effort and costs in the unisexual moss Pseudocalliergon trifarium that has a female-dominated expressed sex ratio and rarely produces sporophytes. Annual vegetative segment mass did not differ among male, female, and non-expressing individuals, indicating that there was no threshold-size for sex expression. Mean and annual mass of sexual branches were higher in females than in males, but branch number per segment did not differ between sexes. Prefertilization reproductive effort for females was significantly greater (11.2%) than for males (8.6%). No cost for sexual branch production in terms of reduced relative vegetative growth or decreased investment in reproductive structures in consecutive years was detected. A higher realized reproductive cost in males cannot explain the unbalanced sex ratio in the study species.     

Sex change in the Mollusca

Sex change in the Mollusca is almost exclusively protandric (male to female), and has only been reported among gastropods and bivalves. The adaptive value of protandry in these two classes most likely relates to the limited availability of females, and the consequent size-independent nature of male reproductive success (versus the size-dependent nature of reproductive success in females). In two well studied distantly related prosobranch gastropod superfamilies, Calyptracea and Patellacea, individuals of some species respond to local ecological changes by altering the age at which they change sex, although the critical ecological changes appear to be different. The physiological switch that activates sex change also appears to be different: it is found in the cemtral nervous system of the calyptracean Crepidula fornicata, and actually within the gonads of the patellacean Patella vulgata. Although the taxonomic breadth of studies on sex change is necessarily limited, and many questions remain to be answered, research on the Mollusca has produced a remarkable range of perspectives on sex change - from evolutionary to proximal; further research will benefit greatly from this breadth of knowledge.     

The cost of males in Daphnia pulex

Sex is paradoxical, because asexuals should replace their sexual ancestors by avoiding the demographic cost of producing males (hereafter referred to as the cost‐of‐males). Despite the large body of theoretical and empirical work dealing with the paradox of sex, the cost‐of‐males assumption has been rarely tested. In the present study, we tested the cost‐of‐males assumption in the cladoceran Daphnia pulex. Populations of this species consist of both cyclically parthenogenetic (i.e. sexuals) and obligately parthenogenetic (i.e. asexuals) lineages. In addition, some of the asexual lineages produce only female offspring, whereas others produce functional males, which can mate with sexual females. We compared the reproductive investment of sexuals, male‐producing asexuals, and non‐male‐producing asexuals when raised separately under various environmental conditions. We also determined the outcome of competition between pair‐wise combinations of these reproductive modes. The cost of males was evident when sexual and asexual females were raised separately: sexuals produced fewer female offspring. However, there was no cost of males when reproductive modes were raised in pairs, as sexuals won the competition with asexuals. Our results directly relate to the field conditions experienced by D. pulex. Sexuals might suffer the cost of males at the beginning of the season, when resource competition is low; but when conditions deteriorate as the population approaches carrying capacity, sexuals seem to be better competitors in spite of male production.     

Sexual selection and maintenance of sex: evidence from comparisons of rates of genomic accumulation of mutations and divergence of sex-related genes in sexual and hermaphroditic species of Caenorhabditis

Several hypotheses have been proposed to explain the persistenceof dioecy despite the reproductive advantages conferred to hermaphrodites,including greater efficiency at purging deleterious mutationsin the former. Dioecy can benefit from both mutation purgingand accelerated evolution by bringing together beneficial mutationsin the same individual via recombination and shuffling of genotypes.In addition, mathematical treatment has shown that sexual selectionis also capable of mitigating the cost of maintaining separatesexes by increasing the overall fitness of sexual populations,and genomic comparisons have shown that sexual selection canlead to accelerated evolution. Here, we examine the advantagesof dioecy versus hermaphroditism by comparing the rate of evolutionin sex-related genes and the rate of accumulation of deleteriousmutations using a large number of orthologs (11,493) in thedioecious Caenorhabditis remanei and the hermaphroditic Caenorhabditisbriggsae. We have used this data set to estimate the deleteriousmutation rate per generation, U, in both species and find thatalthough it is significantly higher in hermaphrodites, bothspecies are at least 2 orders of magnitude lower than the valuerequired to explain the persistence of sex by efficiency atpurging deleterious mutations alone. We also find that genesexpressed in sperm are evolving rapidly in both species; however,they show a greater increase in their rate of evolution relativeto genes expressed in other tissues in C. remanei, suggestingstronger sexual selection pressure acting on these genes indioecious species. Interestingly, the persistence of a signalof rapid evolution of sperm genes in C. briggsae suggests arecent evolutionary origin of hermaphrodism in this lineage.Our results provide empirical evidence of increased sexual selectionpressure in dioecious animals, supporting the possibility thatsexual selection may play an important role in the maintenanceof sexual reproduction.     

Sex‐related variation in migration phenology in relation to sexual dimorphism: a test of competing hypotheses for the evolution of protandry

Timing of arrival/emergence to the breeding grounds is under contrasting natural and sexual selection pressures. Because of differences in sex roles and physiology, the balance between these pressures on either sex may differ, leading to earlier male (protandry) or female (protogyny) arrival. We test several competing hypotheses for the evolution of protandry using migration data for 22 bird species, including for the first time several monochromatic ones where sexual selection is supposedly less intense. Across species, protandry positively covaried with sexual size dimorphism but not with dichromatism. Within species, there was weak evidence that males migrate earlier because, being larger, they are less susceptible to adverse conditions. Our results do not support the ‘rank advantage’ and the ‘differential susceptibility’ hypotheses, nor the ‘mate opportunity’ hypothesis, which predicts covariation of protandry with dichromatism. Conversely, they are compatible with ‘mate choice’ arguments, whereby females use condition‐dependent arrival date to assess mate quality.     

Temperature affects operational sex ratio and intensity of male-male competition: an experimental study of sand gobies, Pomatoschistus minutus

The operational sex ratio is intimately related to the intensityof sexual selection, but factors governing variation in theoperational sex ratio and their effects on mating competitionare still poorly understood. In this study, temperature wasfound to affect both the operational sex ratio and the intensityof male-male competitive interactions in the sand goby [Pomatoschistusminutus (Pallas)]. In an experiment with two different temperaturetreatments, the operational sex ratio became male biased inthe warm treatment (15°C) and males in that treatment interactedmore frequendy than in the cold treatment (8.5°C). Theseresults were as predicted since the potential reproductive rateof males increases faster with temperature than does the potentialreproductive rate of females. Thus, an environmental factor,water temperature, affects not only the reproductive rates ofthe sexes, but also the operational sex ratio and mating competition,and thereby the intensity of sexual selection. Operational sexratio was not found to be correlated with male behavior. Thismay suggest a direct effect of temperature or potential reproductiverates on mate competition. The mechanism behind the evolutionof such a direct relationship would, however, probably be theimpact of potential reproductive rates on operational sex ratio,which in turn direcdy affects sexual selection.     

<Emphasis Type="Italic">Lysmata rafa</Emphasis>, a new species of peppermint shrimp (Crustacea,Caridea, Hippolytidae) from the subtropical western Atlantic

Lysmata rafa n. sp. is described from freshly collected specimens from the Keys West Lakes, Florida Keys, and from a museum specimen collected at Bear Cut, Biscayne Bay, Florida. The new species is morphologically most similar to the western Atlantic Lysmata rathbunae Chace, 1970 and the eastern Pacific Lysmata gracilirostris Wicksten, 2000, but can be distinguished from them by the number of carpal segments in the second pereiopod; the length and dentition of the rostrum; the shape and number of spines on the dactylus of the third to fifth pereiopods; and the absence of a tooth on the pterygostomial margin of the carapace. Despite being a shallow-water species, L. rafa n. sp. has extremely elongate walking legs and third maxilliped that are more typical to deep-water or cave dwelling carideans.