Socially controlled sex-change in the half-moon grouper,Epinephelus rivulatus,at Ningaloo Reef,Western Australia

The cues controlling sex-change have been elucidated for various species of hermaphroditic fishes that inhabit coral reefs, but not for the epinepheline serranids. A male removal experiment conducted on an assemblage of the half-moon grouper, Epinephelus rivulatus, demonstrated that protogynous sex-change in this species is socially controlled, possibly by the suppressive dominance of males and a threshold sex ratio. The experiment showed that a reproductively ripe female can change sex and become a male with ripening testis within 3 weeks. However, this process can be delayed, slowed, or stopped by the presence of other males in the area.     

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.     

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.     

Preservation of winter social dominance status in Brent Geese Branta bernicla bernicla within and across winters

Dominant and subordinate individuals in a group may benefit from the stability of the social dominance organisation, avoiding excessive waste of time and energy in aggressive interactions and reducing injury risks. Nevertheless, the likely evolutionary incentive for individuals to become, and furthermore to stay, dominant may destabilise such dominance hierarchies. In this context, the relative importance of fixed (e.g. sex, morphological size) and fluctuating (e.g. body condition, mating status, reproductive success, social unit size) traits influencing the establishment and preservation of dominance relationships could play a key role in group structure. We investigated the relative role of fixed and fluctuating traits on social status in Dark-bellied Brent Geese Branta bernicla bernicla which form large fairly unstable groups both within and across winters. We compared individual dominance scores of ringed Brent Geese during four consecutive winters. Brent Geese conserved their dominance score within a given winter irrespective of their age but were generally unable to conserve it across consecutive winters. As winter dominance scores correlated best with social unit size, dominance status thus appeared to be mostly a by-product of a fluctuating trait: breeding success in the previous summer. When we considered only adults that had the same social unit size during two consecutive winters, we observed a significant preservation of dominance scores. This result suggests that a fixed trait such as sex or morphological size may still play a role in setting dominance status.     

Social control of sex change in the Red Sea razorfish Xyrichtys pentadactylus (Teleostei,Labridae)

Synopsis The Red Sea razorfish, Xyrichtys pentadactylus, a territorial haremic labrid with dominance hierarchies within the harems. Theory predicts that primary males (fish developing initially as males) should be rare or nonexistent in haremic territorial species because the larger secondary males (males which have undergone sex and/or color change) limit access to females. Histological examination of gonads of 95 specimens showed that all males are derived from females by sex change (i.e. they are secondary males). During five months of field studies 100% of more than 200 observed matings were pair spawnings — the usual mating practice for monandric (having one type of male) species. Sex change in females was induced by male removal in nature. Isolation of four groups of females in aquaria showed that the largest female in the social group changes sex in the absence of a male, demonstrating that sex change is socially-controlled in this species.     

New aspects of sex change among reef fishes: recent studies in Japan

New aspects of sex change in reef fishes are reviewed with special emphasis on recent studies in Japan. For protogyny, studies on both monandric and diandric species have been conducted, but the distinction of primary males from prematurational secondary males seems to need further examination. For protandry, detailed field studies on anemonefishes have revealed alternative life-history styles associated with movements between hosts before or after maturation. The most interesting new aspect has been the discovery of 2-way sex change within a species. Conditions for evolution of 2-way sex change are examined in relation to the size-advantage model and social control mechanisms. A fish may change sex when it becomes dominant in a mating group, but a dominant fish may also change sex in the reverse direction when its social status changes to subordinate through inter-group movement. Two-way sex change has hitherto been reported only from basically protogynous fishes (e.g., Gobiidae, Pomacanthidae, Cirrhitidae, Epinephelinae). Possibilities of the reverse sex change in the protandrous anemonefishes are discussed with data from some unpublished studies.     

Spawning dynamics of orange roughy,Hoplostethus atlanticus,in mid-slope waters of New Zealand

Synopsis The social and reproductive biology of the sand tilefish,Malacanthus plumieri (Malacanthidae), was studied at Glover's Reef, Belize, where this species occurs in colonies over sand-rubble flats. Individuals each occupy a home burrow refuge and a surrounding home range. Home range overlap among adjacent fish of the same sex is low, and individuals defend exclusive use of much of their home range against all conspecifics except mates (i.e., territoriality). Areas defended by males overlap the territories of up to 6 females; and male territory area is positively related to the number of female residents. Males maintain dominance over females within their territories by aggression, including intervention into some female disputes. Females spawn pelagically-dispersed eggs as frequently as every day. Each female spawns near her burrow, almost exclusively with the male whose defended area encompasses her territory (harem polygyny). Tilefish colonies therefore consist of a mosaic of female territories over which adjacent male territories are superimposed. Histological evidence and observation of behavioral sex change in one female revealed thatM. plumieri is capable of protogynous sex reversal. Females did not change sex in response to removal of one male. Occurrence of small transitional fish indicates that the onset of sex change is controlled by factors other than size-related social hierarchies within harems or colonies.     

Social system of an inshore stock of the red hind grouper,Epinephelus guttatus (Pisces: Serranidae)

Synopsis According to sex allocation theory, the decision by a female in protogynous fish species to change sex or not should be influenced by, among other things, the mating sex ratio during spawning periods and/or by factors that vary directly with the spawning sex ratio, such as relative rates of behavioral interaction with males and females outside of spawning periods. In groupers that only spawn during a few weeks of the year in large aggregations, individuals must assess the relative value of changing sex or not entirely within the aggregation unless the social system during the remainder of the year provides a behavioral equivalent of the mating sex ratio. Fifty-five individuals of the red hind,Epinephelus guttatus, were tagged and repeatedly located during a 152-day period within a 100 × 100 m grid on a shallow forereef off southwestern Puerto Rico. The home ranges of 22 tagged individuals sighted 10 or more times were 112–5636 m² in area. Individual home ranges overlapped with the home ranges of 1–18 other individuals. Home ranges of small fish were not clustered within the borders of the home range of larger fish, i.e. fish did not form spatially defined social units. At the end of the study, 31 tagged individuals remained on the grid together with five newly sighted fish. All 36 individuals proved on histological examination to be females similar in size to females in the spawning aggregation of the following year. The sex ratio of this all-female inshore stock differed significantly from the sex ratio of that spawning aggregation. Hence, information predicting the reproductive value of a sex change is not available to females in the inshore stock during nonspawning months.     

Physical interactions facilitate sex change in the protogynous orange-spotted grouper,Epinephelus coioides

Sex change in teleost fishes is commonly regulated by social factors. In species that exhibit protogynous sex change, such as the orange-spotted grouper Epinephelus coioides, when the dominant males are removed from the social group, the most dominant female initiates sex change. The aim of this study was to determine the regulatory mechanisms of socially controlled sex change in E. coioides. We investigated the seasonal variation in social behaviours and sex change throughout the reproductive cycle of E. coioides, and defined the behaviour pattern of this fish during the establishment of a dominance hierarchy. The social behaviours and sex change in this fish were affected by season, and only occurred during the prebreeding season and breeding season. Therefore, a series of sensory isolation experiments was conducted during the breeding season to determine the role of physical, visual and olfactory cues in mediating socially controlled sex change. The results demonstrated that physical interactions between individuals in the social groups were crucial for the initiation and completion of sex change, whereas visual and olfactory cues alone were insufficient in stimulating sex change in dominant females. In addition, we propose that the steroid hormones 11-ketotestosterone and cortisol are involved in regulating the initiation of socially controlled sex change.