Post-settlement emigration affects mortality estimates for two Bahamian wrasses

Understanding the dynamics of open marine populations is inherently complex, and this complexity has led to decades of debate regarding the relative importance of pre- versus post-settlement processes in structuring these populations. Movement between patches may be an important modifier of patterns established at settlement, yet local immigration and emigration have received less attention than other demographic rates. I examined loss rates from tagged populations of juvenile wrasses (yellowhead wrasse Halichoeres garnoti and bluehead wrasse Thalassoma bifasciatum) at two sites in the Bahamas. Assuming that all losses were due solely to mortality would have significantly underestimated survivorship of yellowhead wrasse by 29% and bluehead wrasse by 14%. On average, per capita mortality and emigration rates were higher for yellowhead than bluehead wrasse, but neither demographic rate differed between sites for either species. With respect to within-species density, bluehead wrasse mortality was density-dependent at the patch reef site, but mortality rates of yellowhead wrasse were consistently density-independent. Evaluating the effects of between-species density, yellowhead wrasse mortality increased with a decrease in bluehead wrasse density, but this effect was limited to the patch reef site. Emigration rates were not a function of either within-species or between-species density, but instead varied inversely with isolation distance. Numerous previous studies of coral-reef fish, conducted on patch reefs separated by only a few meters of sand and often using untagged fish, may have confounded losses due to emigration with those due to mortality. A better understanding of the factors affecting emigration in marine fishes is important to their effective management using spatial tools such as marine protected areas.     

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.     

Reversed Sex-Change in the Protogynous Reef Fish Labroides dimidiatus

Protogynous hermaphroditism, or female-to-male sex change, is known for many reef fishes including wrasses (family Labridae) in which large males monopolize mating. When the dominant male disappears from a polygynous group, the largest female may change sex within a few weeks. Such social control of sex change was first documented in harems of the cleaner wrasse Labroides dimidiatus almost 30 yr ago. To examine whether change of social status would induce males of L. dimidiatus to perform reversed sex-change, we conducted experiments: (i) releasing single males near lone males whose mates have been removed in the field; and (ii) keeping two males in a tank. Smaller males changed back to females when they became subordinate: it took 53–77 d (n=3) for them to complete gonadal sex change and release eggs in the aquarium. The male–male pairs performed spawning behavior, with the smaller male in the female role already 5–58 d before completion of gonadal sex change. This is the first report of reversed sex-change among protogynous wrasses. Moreover, we conducted another experiment, keeping a pair of a male and a larger female in a tank (n=1). We found sex change by both mates, which has not been reported from any fishes. Thus, the sex of L. dimidiatus is strictly determined by social status whenever it changes after mate loss.     

Courtship displays and coloration as indicators of safety rather than of male quality : the safety assurance hyposthesis

Male courtship displays and bright coloration are usually assumedto provide information to females about some aspect of themale's value as a mate. However, in some species, courtshipmay serve another function—namely, indicating the currentpredation risk at the mating site and assuring the female thatit is safe to mate there at this time. We developed this safetyassurance hypothesis (SAH) and tested its predictions in thebluehead wrasse (Thalassoma bifasciatum), a Caribbean reef fish. Females in this species come to males' territories to spawn,and males court each arriving female. Males with larger whiteflank patches court less intensely than less bright males.We show that such males are probably more visible to predatorsand thus need not court so intensely to provide the same degreeof safety assurance to a female. When model lizardfish predatorsare presented at spawning sites, males habituate to them quickly,but newly arriving females who see the predator are expectedto demand more assurance of site safety. Accordingly, and consistentwith the SAH, males court females more intensely (longer averagecourtship bout length) under such circumstances, but maleswith bright flank patches do not increase their courtship asmuch as duller males do. Despite this relatively low intensityof courtship, the spawning rate of bright males does not declinerelative to that of duller males in the presence of a predator,suggesting that bright coloration conveys a differential benefit.Females of species like the bluehead wrasse, who spawn repeatedlyover the course of their life, are expected to be more concernedwith their own risk of mortality during each spawning boutthan with the quality of a particular male. It is in such speciesthat we expect the SAH to be most applicable.     

Behaviour of the female beaugregory damselfish (Stegastes leucostictus)

Much less is known about the behaviour of female beaugregory damselfish than about males of the species. This study was initiated to determine behavioural patterns and interactions of female beaugregories on the back reef of Discovery Bay Marine Laboratory, North shore of Jamaica. Females fed, patrolled, and chased intruders in a significantly larger area and ate significantly more per 15-min observation period than did males. Males spent significantly more time patrolling their territories and chased significantly more total intruders than did females. However, this difference in total chases comes from the fact that males chased the bluehead wrasse, an egg predator, significantly more often than did females while all other species of intruder were chased the same by both sexes. The distance that females travelled from their homesite to court was significantly positively correlated to female body length. This increased distance travelled may relate to mate assessment. Female beaugregory damselfish may be able to be more selective in their choice of mate with increased size and distance travelled.     

Early gonadal development and primary males in the protogynous epinepheline, Cephalopholis boenak

Early gonadal development of the protogynous epinepheline, Cephalopholis boenak, was examined histologically in 289 specimens with standard length (L_S) of 42–130 mm, collected from May 2000 to April 2002 in Hong Kong waters, to determine male developmental pathways and establish its sexual pattern. All juvenile gonads developed an ovarian lumen with primary‐growth stage oocytes and scattered spermatogenic tissue prior to sexual differentiation and first sexual maturation. From this bisexual phase containing both female and male tissues, some gonads differentiated as ovaries with further oocyte growth to cortical‐alveolus and vitellogenic stages, the rest differentiated as testes with the proliferation of spermatogenic tissue and the formation of a sperm sinus. All testes retained the lumen and primary‐growth stage oocytes, and sperm sinuses ran within the gonad wall. Unlike most protogynous species, among functional males it was impossible to distinguish those resulting from juveniles through sexual differentiation (i.e. primary male) from those resulting from functional females through sex change (i.e. secondary male) based solely on testicular morphology. A proportion‐spermatogenic‐tissue index (I_ST) was, therefore, developed and determined to be a reliable quantitative indicator for distinguishing differentiating, primary males before a sperm sinus was evident, from differentiating females during sexual differentiation. Since sexually transitional specimens with the concominant appearance of degenerating vitellogenic, or later, stage oocytes and spermatogenic tissue in the gonads were previously noted from Hong Kong, diandric, protogynous hermaphroditism is confirmed in C. boenak. For species, such as this and other epinephelines, in which all males have the same testicular morphology, a complete analysis of a wide range of body sizes from juveniles to adults is necessary for understanding male developmental pathways, and determining sexual pattern.     

Behavioral and energetic costs of group membership in a coral reef fish

Animals in social aggregations often spend more time foraging than solitary conspecifics. This may be a product of the relative safety afforded by aggregations: group members can devote more time to foraging and less time to antipredator behaviors than solitary animals (the “risk reduction” effect). All else being equal, risk reduction should result in higher food intake for grouped animals. However, intragroup competition may force group members to spend more time foraging in order to obtain the same food ration as solitary individuals (the “resource competition” effect). We compared these opposing explanations of foraging time allocation in a coral reef fish, bluehead wrasse (Thalassoma bifasciatum). Aggregations of juvenile bluehead wrasse experience safety-in-numbers, and preliminary observations suggested that juveniles in aggregations spent more time foraging for copepods in the water column than solitary juveniles. However, the risk reduction and resource competition hypotheses are indistinguishable on the basis of behavioral observations alone. Therefore, we collected behavioral, dietary, and growth data (using otolith growth rings) for bluehead wrasse at multiple reefs around a Caribbean island. Despite spending more time foraging in the water column, grouped fish did not capture more prey items and had slower growth rates than solitary fish. Thus, the increased foraging time of grouped fish appears to reflect resource competition, not risk reduction. This competition may limit the size and frequency of aggregations among juvenile bluehead wrasse, which have been shown to experience reduced mortality rates in larger groups. Bluehead wrasse recruits also spent less time foraging but grew faster at sites where planktonic copepod prey were more abundant. This suggests the possibility that large-scale spatiotemporal variability in the abundance of planktonic copepods over coral reefs may produce corresponding variability in the dynamics of reef fish populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Queen-worker conflict over sex ratio: A comparison of primary and secondary sex ratios in the Argentine ant,Iridomyrmex humilis

We compare the primary sex ratio (proportion of haploid eggs laid by queens) and the secondary sex ratio (proportion of male pupae produced) in the Argentine ant Iridomyrmex humilis with the aim of investigating whether workers control the secondary sex ratio by selectively eliminating male brood. The proportion of haploid eggs produced by queens was close to 0.5 in late winter, decreased to less than 0.3 in spring and summer, and increased again to a value close to 0.5 in fall. Laboratory experiments indicate that temperture is a proximate factor influencing the primary sex ratio with a higher proportion of haploid eggs being laid at colder temperatures. Production of queen pupae ceased in mid-June, about three weeks before that of male pupae. After this time only worker pupae were produced. During the period of production of sexuals, the proportion of male pupae ranged from 0.30 to 0.38. Outside this period no males were reared although haploid eggs were produced all the year round by queens. Workers thus exert a control on the secondary sex ratio by eliminating a proportion of the male brood during the period of sexual production and eliminating all the males during the remainder of the cycle. These data are consistent with workers preferring a more female-biased sex ratio than queens. The evolutionary significance of the production of male eggs by queens all the year round is as yet unclear. It may be a mechanism allowing queen replacement in the case of the death of the queens in the colony.     

The effect of local stock density on social behavior and sex change in the Mediterranean labrid Coris julis

Synopsis Two stocks of Coris julis, a protogynous hermaphrodite labrid fish, were studied in the field in Corsica (France) to observe the effect of stock density variations on the timing of sex change. A higher density was associated with lower terminal phase male's territory size and an earlier sex change whilst the proportion of terminal phase males in the stock increased. As well, the average number of females courted per terminal phase male and the percentage of spawning with non-territorial male interferences remained similar in the two situations. These results indicate that territory availability should be considered as an important cue for wrasse sex change.     

Contribution to the biology of the redbanded perch, Ellerkeldia huntii (Hector), with a discussion on hermaphroditism

The presence of oocytes in the testes of several males indicates that the redbanded perch Ellerkeldia huntii exhibits protogynous hermaphroditism. All males appear to be secondary in origin; however the structure of the testes differ from most other protogynous serranids. In samples collected from the Poor Knights Is., New Zealand, males outnumbered females 2.7: 1 ( n = 52). The numbers in each sex were equal until a length of 100–110 mm, after which the male proportion increased to 100%. The oldest female was aged 2+ years, whereas males were found as old as 5+ years. There were no colour differences associated with sex. The mean testes weight was only 7% of the mean ovary weight. Both sexes grew at the same rate. Observations during December indicated that although many individuals were solitary, some were paired. The larger individuals of pairs were males, the smaller females. Individuals foraged in the algal canopy on gammarid amphipods and mysid shrimps. The average range of movements was only 2·2 m².     

Bi-directional sex change in coral reef fishes from the family Pseudochromidae: an experimental evaluation

Coral reef fishes exhibit a diversity of hermaphroditic strategies and comparisons among species with different ecological characteristics will help identify the underlying basis of this complexity. We used manipulative experiments to test the potential for bi-directional sex change in three species of Pseudochromis (Pseudochromidae): P. flavivertex, P. aldabraensis and P. cyanotaenia. The first two species are sexually monochromatic, whereas, P. cyanotaenia is sexually dichromatic. For each species, where two functional females were kept together, one individual in the pair changed sex to male. Where two functional males were kept together, one individual in the pair changed sex to female. In all three species, functional sex change from male to female (52-93 days) took longer than sex change from female to male (18-56 days). In the sexually dichromatic species, P. cyanotaenia, colour change accompanied adult sex change. Females that changed sex to male took on the bright colouration of males and males that changed sex to female took on the drab colouration of females. These results indicate that bi-directional sex change is probably widespread in the family Pseudochromidae and cannot be predicted by the presence or absence of secondary sexual characteristics.     

Social cues influence growth and sexual maturation of the male musk shrew (Suncus murinus)

Musk shrews were maintained from weaning (20 days of age) for 20 or 40 days in one of several social conditions. In Exp. 1, young males housed with adult females gained more weight and had heavier sex accessory organs than did young males housed with an adult male or reared alone. In Exp. 2 this same pattern of accelerated growth and sexual maturation was found when males were reared directly with an adult female or in a split cage where a wire barrier served to separate the male and his adult female cagemate. In Exp. 3, males were reared in cages containing clean or soiled bedding: soiled bedding was taken once every 5 days from the cage of an adult male, or a female. Under these conditions differences in the weights of reproductive tissues showed minimal variation with housing condition after 20 days of treatment. At that time males reared in soiled bedding taken from the cage of an adult female had accelerated development compared with control males. In Exp. 4, males were housed alone or in a split cage with an adult female which was separated by a wire mesh or a solid, opaque barrier. Males separated by a solid barrier from their female cagemates for 40 days had reproductive tissue weights equivalent to those measured in males reared alone. Taken together these results suggest that the presence of an adult female has dramatic effects on body growth and development of reproductive target tissues in young male musk shrews. Male-female social interactions could play an important role in the timing of puberty in this opportunistically breeding tropical mammal.     

A simple model for adaptive variation in the sex ratios of mammalian offspring

We present a simple mathematical model that describes how primary and secondary sex ratios of offspring may vary adaptively in order to maintain equal numbers of the sexes at the age of reproductive maturity. The model postulates that the sex of an offspring depends probabilistically on a weighted linear combination of maternal testosterone and male vulnerability. The model operates at population level, and is based on three physiological phenomena: first that maternal testosterone in follicular fluid is normally distributed, with levels above the mean more likely to be associated with the conception of males; secondly, that males are more vulnerable than females from conception onwards; and thirdly that under conditions of chronic stress, increased secretion of female testosterone coincides with increased male vulnerability. Thus during times of chronic stress, more males are conceived, but their number of live births is moderated by increased male loss. Variations in secondary sex ratios should therefore be related not only to the stressfulness of environmental conditions, but also to the timing of changes in stressfulness.     

Development of the testes in female domestic fowls submitted to an experimental sex reversal during embryonic life

The sex differentiation of the female chick embryo can be totally inverted toward the male sex by an early extraembryonic testis grafting. This sex reversal remains permanent, as shown by three adult fowls described in this paper. They possess two testes associated with normally differentiated male excretory ducts and their Müllerian ducts have regressed. The development of male sex characteristics such as external features, behavior and complete spermatogenesis is evidence that these cocks have endocrine and exocrine capabilities similar to those of normal cocks. Although these cocks were able to mate with female fowls, they were sterile. A mechanism is discussed by which grafted testes induce such modifications in females. Hypotheses considering the heterogametic sex (female in birds) as exerting a dominant influence on the phenotypic sexual differentiation can be discarded in light of our results because a homogametic testis provokes the definitive sex inversion of a female.     

Phenotypic variation between dimorphic males in corkwing wrasse

The age-frequency distributions for sneaker males (individuals with female secondary sexual characters) and territorial males (individuals with typical male sexual characters) of corkwing wrasse Symphodus melops in a Swedish population did not differ. Otolith analysis revealed that sneaker males were smaller than their territorial male counterparts as juveniles, but there was no evidence of a clear-cut threshold to determine the subsequent reproductive patterns. Thus, the expression of male morph in corkwing wrasse most likely is related to early growth pattern and the different morphs could be fixed for life. Adult sneaker males in general were smaller than territorial males of the same age. The relative gonad size and the abdomen in sneaker males were substantially enlarged compared with territorial males, but there was no difference in gonad water content between the two types of males. Sneaker males weighed less in relation to their length than territorial males. These results support theoretical sperm competition models which predict that the male morph with reduced attractiveness to females (i.e. sneakers) should invest more into spermatogenesis in order to achieve fertilization than the attractive male morph (i.e. territorial males).     

The Humphead Wrasse,Cheilinus Undulatus: Synopsis of a Threatened and Poorly Known Giant Coral Reef Fish

The humphead wrasse, Cheilinus undulatus, is the largest living member of the family Labridae, with a maximum size exceeding 2 m and 190 kg. Its geographic range covers much of the Indo-Pacific. The species is not common, recorded maximum adult densities rarely exceeding 20 fish/10,000 m². Small individuals are typically associated with high coral cover; larger fish are found mainly on outer or deep reefs, steep slopes and passes, singly or in small groups and seagrasses. However, for reproduction the species forms small spawning aggregations of tens to more than one hundred fish. The diet of the humphead wrasse includes large invertebrates and small fishes. The species attains at least 30 years and reaches sexual maturation at about 35–50 cm total length and <5 years of age. Most small adults are female while mainly males exceed 1 m and there is evidence of female to male sex change. The humphead wrasse is of considerable cultural value in some Pacific countries and is among the most prized in the live reef food fish export trade, for which it is often taken in its juvenile size range, either directly for sale or, increasingly, for grow-out to market size. It is also marketed chilled. The species is particularly sensitive to fishing pressure. In most fished areas, density and body size have dropped substantially. It appears to be particularly heavily targeted and depleted in SE Asia and in some places faces extirpation. The humphead wrasse is often taken by night spearfishing and by cyanide, with protection typically weak or non-existent, despite regulations and by a vulnerable assessment on the 1996 IUCN Red List. The humphead wrasse has not been reared successfully in hatcheries. Other giant reef fish share many similar problems and detailed study of the humphead wrasse contributes to a better understanding and conservation of all such species. This review examines and evaluates published and gray literature, original unpublished research and correspondence with almost 50 knowledgeable workers. It examines the value of such sources for quickly, but adequately, assessing the conservation and management status and key data gaps in species that are little known, vulnerable, difficult and expensive to study and may require urgent management or conservation action.     

Sex allocation and egg distribution of an autoparasitoid, Encarsia pergandiella (Hymenoptera: Aphelinidae)

Abstract. 1. Encarsia pergandiella Howard females develop as primary parasitoids on immature whiteflies, and males develop as secondary parasitoids on females of their own or a related species. The hypothesis that the sex ratio reflects the relative abundance of the two host types was tested in the laboratory using petri dish arenas with varying proportions of early fourth instar greenhouse whitefly (Trialeurodes vaporariorum (West.)) (primary hosts) and pupal female E.pergandiella (secondary hosts). Egg distribution was analysed with respect to sex ratio, super-parasitism and host discrimination.
2. The proportion of primary and secondary hosts parasitized in each treatment reflected the relative availability of each host type. Thus females presented with 75% primary hosts laid more female eggs than male. However, in all treatments, a greater proportion of secondary hosts were parasitized than would be expected from the proportion of secondary hosts available. This indicates that more male eggs were laid than expected.
3. More secondary hosts than primary hosts were superparasitized.
4. Host discrimination analysis using a new test statistic showed that females generally laid eggs at random with regard to previous parasitism of primary or secondary hosts. However, females in one treatment with 50% of each host type appeared to preferentially oviposit in secondary hosts which did not contain any eggs.