Both-ways sex change in monogamous coral gobies, Gobiodon spp.

Synopsis We confirmed both-ways sex change in the coral-dwelling gobies Gobiodon micropus, G. oculolineatus, G. quinquestrigatus and G. rivulatus rivulatus by mate-removal experiment in the field and by the aquarium experiment of keeping two consexual fish in a coral. Eight species of Gobiodon were found in Acropora corals on the reef flat of Sesoko Island, Okinawa, southern Japan. The 4 species mentioned above bred in monogamous pairs composed of a male and a female matched by size, and the male took care of eggs deposited on the coral branch. In G. quinquestrigatus and G. rivulatus rivulatus males were larger than females in newly formed pairs, and females grew faster than their mates until breeding. The growth-rate advantage in females seems to be the major factor in the evolution of female to male sex change. The gobies strongly depended on host corals, but they moved between the corals after mate loss or coral death to form new pairs. This provides opportunities for the evolution of male to female sex change; the ability to change sex in both directions reduces the frequency of risky movement between host corals to form new pairs. These conditions are very similar to those reported in the both-ways sex change of another coral-dwelling goby Paragobiodon echinocephalus.     

Sexual-Selection Models for Exaggerated Traits are Useful but Constraining

SYNOPSIS. Science is driven by productive hypotheses and technology,but these may sometimes limit the questions posed. For instance,Fisherian runaway sexual selection and related hypotheses havehelped us understand the evolution of exaggerated visual sexualdimorphism. Species with indistinguishable sexes, however, mayuse different behavioral mechanisms when pairing and thus possessdifferent adaptations. In the monomorphic Midas cichlid (Amphilophuscitrinellum), females chose large aggressive males in a restrainedsituation, as sexual selection predicts, but males did not choose.The nuchal hump of males swells coincidently with pair formation.However, overly large humps were shunned by females while thenormal— size hump facilitated sex recognition. This speciesis polychromatic, and pairs mate assortatively by color in Nicaragua.Some have suggested the Midas cichlid might therefore show howsexual selection produces explosive speciation of cichlids inAfrica. All females, however, are biased toward normal—colormales. The color of gold morphs modulates aggressive responsesof the other fish. All else equal, the benefit to gold in afight equals 15% more weight than the opponent. Pair formationsucceeds best when the typically smaller female of a pair isrelatively more aggressive than the male. The pair combination,gold male with normal female, is difficult to produce; makingthe female the same size as the male removes the disability.Pair formation is a negotiated process in which the male teststhe aggressiveness of the female relative to self. That putsthe behavioral mechanisms of the male and female in conflict.     

Sexual selection for large male size in a polyandrous butterfly: the effect of body size on male versus female reproductive success in Pieris napi

Sexual selection theory predicts that the larger sex shouldbe that for which fitness increases at the faster rate withsize. In butterflies, as in most invertebrates, females areusually the larger sex, but previous comparative analysis hasshown that relative male size increases with female polyandryamong butterflies. In agreement with this pattern, males arelarger than females in the strongly polyandrous green-veinedwhite butterfly, Pieris napi L., and in this article we assessthe size dependence of reproductive success in both sexes. Inan experiment where virgin males and females were released inthe field, we found no strong association between size and malemating success. However, laboratory experiments showed thatthere was a strong correlation between size and the ejaculatethat the male delivered to the female at mating and that largeejaculates delayed female remating for a longer time comparedto small ejaculates. Moreover, female P. napi utilize male-derivednutrients received at mating to increase their fecundity. Hence,large males sire more offspring both by way of donating morenutrients to female egg production and by way of delaying femaleremating (given that the last male to mate with the female willfather most of the offspring). Laboratory experiments showedthat the association between size and fecundity was low, ornonexistent, among P. napi females allowed to mate only once.However, weak size dependence was found for polyandrous females.We hypothesize that size dependence of female fecundity maybe especially weak among polyandrous butterflies because a fundamentalsource of variation in fecundity relates to their ability tofind nutrient giving males, an ability which may be unrelatedto female size. According to this hypothesis there is a causalassociation between weak size dependence of female fecundityand polyandry, and a strong size dependence of male reproductivesuccess that may underlie the comparative pattern of positivecorrelation between relative male size and polyandry.     

Social Reversal of Sex‐Biased Aggression and Dominance in a Biparental Cichlid Fish (Julidochromis marlieri)

In biparental species, aggression, dominance, and parental care are typically sexually dimorphic. While behavioral dimorphism is often strongly linked to gonadal sex, the environment—either social or ecological—may also influence sex‐biased behavior. In the biparental cichlid fish Julidochromis marlieri, the typical social environment for breeding pairs consists of large females paired with smaller males. While both sexes are capable of providing territory defense and parental care, the larger female provides the majority of defense for the pair, while the smaller male remains in the nest guarding their offspring. We examine the contributions of sex and relative mate size to these sex‐biased behaviors in monogamous J. marlieri pairs. Both female‐larger and male‐larger pairs were formed in the laboratory and were observed for territorial aggression (against conspecifics and heterospecifics), dominance, and parental care. In female‐larger pairs, territorial aggression and intra‐pair dominance were female‐biased, while in male‐larger pairs this bias was reversed. For both pairing types, the presence of an intruder amplified sex differences in territorial aggression, with the larger fish always attacking with greater frequency than its mate. Though less robust, there was evidence for plasticity of sex‐bias for some egg care related behaviors in the inverse direction. Our study suggests that relative mate size strongly influences the sex bias of aggression and dominance in J. marlieri and that this aspect of the social environment can override the influence of gonadal sex on an individual's behavior. The remarkable plasticity of this species makes Julidochromis an exciting model that could be used to address the relationship between proximate and ultimate mechanisms of behavioral plasticity.     

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.     

Bidirectional sex-change behavior and physiological aspects in the Gorgeous goby Lythrypnus pulchellus (Gobiidae)

The Gorgeous goby Lythrypnus pulchellus shows extreme sexual plasticity with the bidirectional sex-change ability socially controlled in adults. Therefore, this study describes how the hierarchical status affects hormone synthesis through newborn hormone waste products in water and tests the influence of body size and social dominance establishment in sex reversal duration and direction. The associated changes in behavior and hormone levels are described under laboratory conditions in male–male and female–female pairs of similar and different body sizes, recording the changes until spawning. The status establishment occurred in a relatively shorter time period in male and female pairs of different sizes (1–3 days) compared to those of similar size (3–5 days), but the earlier one did not significantly affect the overall time of sex change (verified by pair spawning). The changes in gonads, hormones, and papilla occurred in sex-changer individuals, but the first one was observed in behavior. Courtship started at 3–5 days in male pairs and from 2 h to 1 day in female pairs of both groups of different and similar sizes. Hormones did not gradually move in the new sexual phenotype direction during the sex-change time course. Nonetheless, estradiol regulated sex change and 11-ketotestosterone enabled bidirectional sex change and was modulated by agonistic interactions. Cortisol is associated with status and gonadal sex change. In general, similar mechanisms underlie sex change in both directions with a temporal change sequence in phases. These results shed new light on sex-change mechanisms. Further studies should be performed to determine whether these localized changes exist in the steroid hormone synthesis along the brain–pituitary gonad axis during social and bidirectional sex changes in L. pulchellus.     

Sex reversal in pairs of Lythrypnus dalli: behavioral and morphological changes

In Lythrypnus dalli, the bluebanded goby, reproductive success is primarily determined by functional sex, and functional sex is determined largely by rank in the dominance hierarchy. In most natural social groups of L. dalli, one male is at the apex of the hierarchy, and 1 to 7 females are lower in rank. When a male exits the group, a female ascends to the top of the hierarchy and becomes a male. We have examined this process in a simplified environment--a pair of females--that allows us to identify behavior associated with the formation of a dominance relationship and any other phenotypic changes associated with dominance, sex change or both. We found that pairs of L. dalli females quickly and readily form stable dominance relationships, with the dominant fish changing sex into a male. This dominant animal also rapidly increases in body size and length of its dorsal fin. In summary, dominant L. dalli females change sex in this simplified environment, providing excellent opportunities to examine the early behavioral and morphological changes associated with dominance and sex change.     

Genetic and environmental control of temporal and size-dependent sex allocation in a wind-pollinated plant

Sex allocation in hermaphrodites can be affected by spatial and temporal variation in resources, especially in plants where size-dependent gender modification is commonplace. The evolution of sex allocation will depend on the relative importance of genetic and environmental factors governing patterns of investment in female and male function. In wind-pollinated plants, theoretical models predict a positive relation between size and male investment because of the fitness advantages associated with more effective pollen dispersal. Theory also predicts that the timing and allocation to each sex function should depend on available resources. We grew maternal half-sibling families of annual, wind-pollinated, Ambrosia artemisiifolia in sun and shade treatments to investigate these predictions. There was significant genetic variation for female and male flower production in both sun and shade treatments. Size-dependent sex allocation occurred in the direction predicted by theory, with male flower production increasing more rapidly in larger plants. The timing of sex function also varied, with significant genetic variation for dichogamy within environments and plasticity of this trait between environments. Protandry was expressed more commonly in the sun and protogyny in the shade. The occurrence of dynamic sex allocation with changing size and experimental treatment indicates the potential for adaptive responses under different ecological conditions.     

Size, operational sex ratio, and mate-guarding success of the carrion beetle, Necrophila americana

When male insects guard females until oviposition, the benefitsfrom last-male sperm precedence must outweigh the costs of relinquishingadditional fertilizations. The profitability of guarding isincreased when males guard large, fecund females and when femalesare scarce because fewer fertilizations are sacrificed. However,the male reproductive success is not only determined by theprofitability of guarding but also by his ability to maintainguarding. In this study, we used male carrion beetles (Necrophilaamericana) to examine the effects of sex ratio, male relativesize, and female quality on the ability to guard. First, wepresent a model of mate guarding that explores factors, suchas sperm precedence, sex ratio, male size, and female quality,that influence the profitability of postcopulatory riding. Ourmodel predicts that large N. americana males should preferentiallyguard the largest female only when the sex ratio is male biasedand sperm precedence is above 80%. In contrast, small malesgain little from guarding because they are not likely to maintainit and be the last male to mate. Then, we tested these predictionsby manipulating sex ratio, relative male size, and female quality.All males in equal sex ratio and large males in male-biasedsex ratio guarded females significantly longer than did malesin female-biased sex ratio. In male-biased sex ratio, largemales guarded significantly longer and achieved more takeoversthan small males. Large females were guarded longer. The successof guarding males in this beetle depends on their size relativeto other males and the operational sex ratio.     

Climbing to reach females: Romeo should be small

The race for reaching mates by the time they are receptive, or sexual selection by scramble competition, has received little attention. We argue that smaller males are favored in species in which the male must climb to reach females located in high habitat patches. This new explanation we term the "gravity hypothesis" of sexual size dimorphism (SSD). We show that a simple biomechanical model of animal movement predicts that: (1) selection should favor a comparatively smaller size in the searching sex when searching involves climbing; and (2) this effect should be stronger in larger species than in smaller species. In reaching high habitats, smaller, faster searchers will be favored either through sexual selection by scramble competition and/or by escaping predation easier by running faster on vertical surfaces. Different spider species are found at a wide range of heights. We compiled a dataset of spider taxa and arranged their habitats according to four height categories, ranked from soil surface to trees. We show that, after controlling for phylogeny, both predictions of the gravity hypothesis of SSD are met. Thus, it appears that the constraint imposed by gravity on climbing males is a selective factor in determining male dwarfism.     

Further observations relating sex size ratios to mating success in calanoid copepods

Mating experiments using 153 pairs of Diaptomus leptopus Forbeswere video-taped in the laboratory. The following were measuredand scored: attempted capture of the female by the male, timeto successful capture and mounting, duration of copulation,spermatophore placement, time to clutch extrusion, prosome lengthsof all individuals and sex size ratios (female:male lengths)of all pairs. Mating success was not a function of sex sizeratio for D.leptopus at ratios commonly observed in nature inthe populations tested. However, this lack of relationship maynot be true of all populations. Photographic analysis of D.leptopusas well as D.birgei Marsh showed that males always held ontofemale genital segments in the vicinity of the spines with theirright fifth legs. Pearson correlations were calculated comparingprosome lengths to a variety of genital parts. The strongestrelationships were found with female genital segment width atthe level of the spines for both species, and male right fifthleg claw length for D.birgei. These data support the hypothesisthat the relationship between sex size ratio and mating successmay be species specific.     

Does gonad structure reflect sexual pattern in all gobiid fishes?

Synopsis In immature and adult females of protogynous gobies, small distinctive masses of cells associated with the ovarian wall develop into testis-associated glandular structures during sex change. These precursive accessory gonadal structures, or pAGS, have been found in females of known protogynous goby species, but not among gonochoric goby species, suggesting that their presence can be used as a species-specific indicator of protogyny within the family. However, a detailed examination of a developmental series of ovaries in two gonochoric species,Gobiosoma illecebrosum andG. saucrum, revealed the presence of a gonadal feature previously thought to be restricted to protogynous gobies. Among immature females of both species, pAGS-like structures having a similar appearance and placement as functional pAGS of protogynous gobies were found. In femaleG. illecebrosum, the size of these structures among immatures progressively decreased with maturation and were absent in all but the smallest adult females. A similar pattern was evident in a small sample ofG. saucrum. Population demography based on field collections showed thatG. illecebrosum exhibits sex ratios and male and female size-frequency distributions typical of gonochores and laboratory experiments indicated that final sexual identity was unaffected by social environment during the juvenile period. Thus, the presence of pAGS in juvenile femaleG. illecebrosum is not related to an ability to change sex at that ontogenic interval. Whether the transient pAGS observed here are vestiges of an ancestral protogynous condition is unknown. Based on their presence among immatures in two gonochore gobies, however, only the presence of pAGS in adult females should be used to predict protogyny among gobies.     

Sex allocation and nestling survival in a dimorphic raptor: does size matter?

Fisher's theory predicts equal sex ratios at the end of parentalcare if the costs and benefits associated with raising eachsex of offspring are equal. In raptors, which display variousdegrees of reversed sexual size dimorphism (RSD; females thelarger sex), sex ratios biased in favor of smaller males areonly infrequently reported. This suggests that offspring ofeach sex may confer different fitness advantages to parents.We examined the relative returns associated with raising eachsex of offspring of the brown falcon Falco berigora, a medium-sizedfalcon exhibiting RSD (males approximately 75% of female bodymass) and subsequent sex ratios. Female nestlings hatched eitherfirst or second did not receive more food nor did they hatchfrom larger eggs or remain dependent on parents for longer periodsthan male offspring from these hatch orders. Together with previousstudies this result indicates that even in markedly dimorphicspecies, the required investment to raise the larger sex islikely to be less than that predicted by body size differencesalone. Moreover, among last-hatched nestlings, both sexes faceda reduced food allocation and suffered a slower growth rateand thus final body size, with a concurrent increased probabilityof mortality. For last-hatched females the reduction in foodallocation was more marked, with complete mortality of all last-hatchedfemale nestlings monitored in this study. Once independent,males of any size but only larger females are likely to be recruitedinto the breeding population. The sex-biased food allocationamong last-hatched offspring favoring males thus reflects therelative returns to parents in raising a small member of eachsex.     

Male load-lifting capacity and mating success in the swarming caddis fly Athripsodes cinereus

Abstract. Males of the caddis fly Athripsodes cinereus (Curtis) (Trichoptera: Leptoceridae) swarm above the water surface of lakes and streams. Females enter swarms and are pursued until grasped by a male. The pair couple their genitalia in the air, and then the male alone flies the pair to the shore where they settle and complete the copulation. About 8% of the pairs (total n = 384 pairs) dipped in the water soon after the coupling manoeuvre and about 25% of those then separated. Males in dipping pairs ( n = 13) were on average smaller and relatively older than the males that successfully carried their mate to the shore ( n = 54). No differences were found for flight muscle ratio (weight of flight muscles/total body weight) or relative load (total load/flight muscle weight). Males were larger than females (wing length), though typically female Trichoptera are the larger sex. Large male body size in A. cinereus may be an adaptation for flight during pairing; i.e. larger males are more likely to be able to carry larger loads.     

Cortisone and the growth of populations of Mesocestoides tetrathyridia in mice.

Volumes and weights of populations of Mesocestoides tetrathyridia increased faster in SEC mice treated with cortisone, than in the controls. Cortisone did not alter the sex resistance of the host and the populations always grew faster in male than in female hosts. Larvae of the populations from cortisone treated mice were smaller than those from control mice, and those from males smaller than those from females.The average size of the larvae decreased as the population size increaed. Fast growing populations contained a higher percentage of larvae with 2 suckers. These forms may serve as indicators of accelerated division of larvae.     

Mating Patterns of Red-Eyed Treefrogs, Agalychnis callidryas and A. moreletii

Deviations from random mating in frogs are often explained by two different size‐based patterns. The large‐male mating advantage predicts that males found in amplexus with females will be larger on average than non‐amplectant males, whereas size‐assortative mating predicts that males and females found in amplexus will maintain an optimal size ratio. Both these pairing patterns are consistent with a female mating preference for larger males, or for males of a given size relative to the choosy female. I examined pairing patterns of two species of Neotropical hylids, Agalychnis callidryas and A. moreletii for three consecutive breeding seasons in Belize, Central America to evaluate whether mating behavior was influenced by either a large‐male mating advantage or size‐assortative mating. For each species, I compared size traits between amplectant and non‐amplectant males, and within amplectant pairs, and I quantified fertilization success for each amplectant pair. For both species I found evidence of deviations from random mating by size, but the nature of the deviations varied between species and among years. The proportion of eggs fertilized was consistently high among years for both species and there was no relationship between fertilization success and the size ratio of amplectant pairs. These data are consistent with female mate preference, but a role for male–male competition cannot be excluded. My findings suggest that mating patterns may be density‐dependent and that the nature and intensity of sexual selection may be increased by extreme environmental conditions.     

Sex Reversal and Fruit Formation on the Male Plants of Morus nigra L. by 2-Chloroethylphosphonic Acid

Ethrel treatment at 960, 1920, and 3840 parts 10–6 tomale plants of M. nigra produced intersex and female flowers.The intersexual flowers showed various degrees of transformationof male sex organs into female. The complete female flowersproduced were similar to female controls. Fruit setting occurredin inflorescences having both types of flowers which were similarin appearance but smaller in size than the untreated control.     

Adaptiveness of sex ratio control by the pupal parasitoid Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size

Adaptiveness of sex ratio control by the solitary parasitoid wasp Itoplectis naranyae (Hymenoptera: Ichneumonidae) in response to host size was studied, by examining whether differential effects of host size on the fitness of resulting wasps are to be found between males and females. The offspring sex ratio (male ratio) decreased with increasing host size. Larger hosts yielded larger wasps. Male larvae were less efficient in consuming larger hosts than female larvae. No significant interaction in development time was found between parasitoid sex and host size. Larger female wasps lived longer than smaller females, while longevity of male wasps did not increase with increasing wasp size. Smaller males were able to mate either with small or with large females, while larger males failed to mate with small females. Larger female wasps had a greater number of ovarioles and mature eggs at any one time than smaller females, although the number of eggs produced per host-feeding was not influenced by female wasps. Thus, the differential effect of host size on the fitness of males and females exists in I. naranyae. The basic assumption of the host-size model was therefore satisfied, demonstrating that sex ratio control by I. naranyae in response to host size is adaptive.     

Sperm competition and sex change: a comparative analysis across fishes

Current theory to explain the adaptive significance of sex change over gonochorism predicts that female-first sex change could be adaptive when relative reproductive success increases at a faster rate with body size for males than for females. A faster rate of reproductive gain with body size can occur if larger males are more effective in controlling females and excluding competitors from fertilizations. The most simple consequence of this theoretical scenario, based on sexual allocation theory, is that natural breeding sex ratios are expected to be female biased in female-first sex changers, because average male fecundity will exceed that of females. A second prediction is that the intensity of sperm competition is expected to be lower in female-first sex-changing species because larger males should be able to more completely monopolize females and therefore reduce male-male competition during spawning. Relative testis size has been shown to be an indicator of the level of sperm competition, so we use this metric to examine evolutionary responses to selection from postcopulatory male-male competition. We used data from 116 comparable female-first sex-changing and nonhermaphroditic (gonochoristic) fish species to test these two predictions. In addition to cross-species analyses we also controlled for potential phylogenetic nonindependence by analyzing independent contrasts. As expected, breeding sex ratios were significantly more female biased in female-first sex-changing than nonhermaphroditic taxa. In addition, males in female-first sex changers had significantly smaller relative testis sizes that were one-fifth the size of those of nonhermaphroditic species, revealing a new evolutionary correlate of female-first sex change. These results, which are based on data from a wide range of taxa and across the same body-size range for either mode of reproduction, provide direct empirical support for current evolutionary theories regarding the benefits of female-first sex change.     

Inter-group movement of females of the polygynous gobiid fish <Emphasis Type="Italic">Trimma okinawae</Emphasis> in relation to timing of protogynous sex change

Social conditions and function of inter-group movement of females of the polygynous goby, Trimma okinawae, have been studied at Akamizu Beach, Kagoshima, Japan. Some females moved from their original groups, where the male was still present, to other groups. Before the movement females sometimes temporarily visited the group into which they subsequently moved, suggesting they were able to assess social conditions during the visit. By moving, the females increased in size rank or escaped from similar-sized female competitors in their previous groups. Although the social ranks of the moving females in their original groups were lower than those of the resident females, the ratio of the number of females that changed sex to the number of females surviving at the end of the study did not differ for the two types of female. Inter-group movement of females may increase the probability of their changing sex to become a dominant male.