Spawning behavior and haremic mating system in the corallivorous butterflyfish,Chaetodon trifascialis, at Kuroshima Island, Okinawa

Territorial and spawning behavior ofChaetodon trifascialis were investigated on a small patch of reef at Kuroshima Island, Okinawa, Japan. Three males and 8 females inhabited the reef, each individual defending a territory against conspecifics of the same sex. Each male territory included 2 or 3 female territories. In the daytime, each male frequently visited the females living in its territory. At dusk in the full or new moon periods, courtship began within the female territories, pair spawning subsequently occurring within or near those territories. When a male actively courted a female in the territory of a second male, the latter male immediately chased off the intruder. Thus, mating occurred only between a male and females living in former's territory. This is the first report of a haremic mating system among butterflyfishes (Chaetodontidae).     

The Haremic Mating System and Mate Choice in the Wide-Eyed Flounder, Bothus podas

The social structure and reproductive behaviour of the wide-eyed flounder, Bothus podas, was studied in the coastal waters around the Azorean Islands. Both sexes are territorial throughout the year. Adult males defend large territories, which include several smaller female territories. Intraspecific agonistic behaviour was frequent and differed between sexes: males were more aggressive towards other males, while females were only aggressive towards each other and juveniles. During the reproductive season and only at dawn, territorial males court and mate successively with females in their territories, and females seem to show mating fidelity to their dominant male. Such territoriality and mating patterns indicate a haremic social system in the wide-eyed flounder. In order to identify potential factors influencing female mate choice acting on this haremic system, we examined male mating success and some of its potential correlates. We found no evidence for female preference for any of the males' physical or territory characteristics. However, courtship effort was strongly correlated with the total number of attempted and successful spawnings, indicating that females seem to mate preferentially with males that court them more vigorously. Thus, our data suggest that courtship plays an important role in determining male mating success in the wide-eyed flounder and, that it may possibly serve as an honest indicator of male `quality' for female choice.     

Reversed Sex Change in The Haremic Protogynous Hawkfish Cirrhitichthys falco in Natural Conditions

Bi‐directional sex change has recently been reported in a range of reef fishes, including haremic species that were earlier thought to be protogynous (female to male). However, the occurrence of this phenomenon and the social conditions driving the reversion of males to females (reversed sex change) have been poorly documented under natural conditions. Reversed sex change is predicted to occur in low‐density populations where facultative monogamy is common. However, few studies have evaluated this over a long period in such populations. We documented the occurrence of bi‐directional sex change during a 3‐yr demographic survey of a population characterised by small harem sizes in haremic hawkfish Cirrhitichthys falco. New males were derived following a change in sex of functional females (secondary males; n = 3) and juveniles always matured first as females (n = 3). Thus, C. falco exhibited a typical protogynous sexual pattern, consistent with a range of haremic fish species. We observed reversed sex change in two males. In both cases, all the females disappeared from their harems and the neighbouring males expanded their territories to encompass the territories of the sex changers. However, bachelor males did not always revert to females. A dominant male experienced bachelor status twice but regained mating opportunities following the immigration of a female into his territory or by taking a female from a neighbouring harem. Thus, we conclude that bachelor males use reversed sex change as a facultative tactic to regain reproductive status in a haremic mating system. In addition, we discuss the influence of harem size upon occurrence of reversed sex change.     

Two alternative female tactics in the polygynous mating system of the threadsail filefish,Stephanolepis cirrhifer (Monacanthidae)

Reproductive behavior of the threadsail filefishStephanolepis cirrhifer was studied at Kashiwajima, southern Shikoku, Japan. This species spawned in pairs on the sandy bottom, the eggs being scattered over an area of about 15 cm in diameter and attached to sand particles. After spawning, males departed immediately, while the females remained at the site to guard the eggs for a few minutes. Thereafter the eggs were left unguarded for three days until hatching. Females spawned only once daily, whereas males mated with multiple females in succession. The reproductive males established territories, in which 1–4 resident females defended smaller territories from each other. The harem size changed according to some ecological conditions, such as population density. Moreover, the males also mated with visiting non-resident females. Thus, the two alternative tactics of females resulted in two mating patterns, haremic polygyny and female visiting of male territories, in a single population ofS. cirrhifer.     

Harem structure and female territoriality in the dwarf hawkfish <Emphasis Type="Italic">Cirrhitichthys falco</Emphasis> (Cirrhitidae)

We investigated the territoriality and the spatial and mating relationships of the haremic hawkfish, Cirrhitichthys falco, on a reef off Kuchierabu-jima Island in southern Japan. Each individual maintained a territorial home range which was defended against same-sex conspecifics at the boundary of the home range. The territory of each male encompassed the territories of 2–3 females, allowing the male to completely monopolize mating opportunities with those females. Based on our observations, we classified the harem type of C. falco as a territorial female type. Large juveniles maintained independent home ranges outside the female territories. In contrast, small juveniles were allowed to cohabit within the territory of an adult female. Stomach contents analysis revealed that the smallest size class of C. falco fed primarily on copepods. In contrast, all other size classes fed primarily on decapods. Together, these results suggest that female territoriality plays an important role in defending food resources.     

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.     

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.     

Spawning and biparental egg-care in a temperate filefish,Paramonacanthus japonicus (Monacanthidae)

Synopsis Reproductive habits of a temperate filefish, Paramonacanthus japonicus, were studied on a rocky reef at Tsuyazaki, Fukuoka, Japan, from 1989 through 1990. Males had territories of 30–70m² and defended them from conspecific males and potential egg predators such as another filefish, Stephanolepis cirrhifer. Egg masses were found on the sandy bottom in male territories. Individual discrimination of males and females occurring in three male territories revealed that males and females stayed in stable pairs during one month of observation in 1989. In these stable pairs, males fed only within their territories, but females occasionally foraged outside. The occurrence of egg masses within male territories and biparental egg care showed that fish were reproducing as monogamous pairs. Contrary to this, males tagged in 1990 changed their territories after the disappearance of females, and males and females mated polygamously. Spawning was observed only four times during the study period, between 1633 and 1754h. Prior to spawning, the female prepared a spawning bed on the sandy bottom. The male nuzzled the female and the pair spawned, touching their gonopores on the spawning bed. Spawning was very quick and took only 1–3 seconds. The adhesive eggs were spherical with a diameter of 0.56 mm. They were mixed with sand particles and formed a doughnut-shaped mass of about 4 cm in diameter. One egg mass contained 3300–3800 embryos of similar developmental stage, which hatched 2–3 days later. P. japonicus appears to be monogamous but may also practice polygamy when pair-bonds are unstable.     

Mating system of the lefteye flounder,Engyprosopon grandisquama

Underwater observations were conducted on the reproductive behavior and mating system of the lefteye flounder,Engyprosopon grandisquama, off Nagashima, southwest of Kyushu Island, Japan. Two types of males were found: large males, which defended territories against other large males, and small males, which did likewise but only against smaller males. Large males established territories which encompassed or ovelapped the home ranges of 1 or 2 cohabitant females. Territories of the small males, in which a smaller female often maintained a home range, overlapped those of large males. Pair spawning occurred around sunset. Mating of large males with cohabitant females was observed 36 times and that of a small male with a smaller female once. Pair formation was assorted by body size, paired males being larger than females in most cases. Thus, inE. grandisquama, gigamous large males were common, small males occurring within the former's erritorial boundaries mating with smaller females.     

Female defense polygyny in the territorial triggerfish <Emphasis Type="Italic">Sufflamen chrysopterum</Emphasis>

The occurrence of polygyny requires specific environmental conditions such as female aggregation or patchy resource distribution. However, it is difficult to determine the factors responsible for polygyny in species in which the territories of both sexes overlap. To overcome this, we performed female removal experiments in the polygynous triggerfish Sufflamen chrysopterum (Balistidae) in the Okinawa coral reef. Both sexes defended their territories exclusively against consexuals of the same species, and female aggregation was absent. Each male territory included 1–3 female territories, and nonterritorial males were significantly smaller than territorial males. Further, the body size of territorial males was positively correlated with that of the largest female in their territories, and larger males tended to mate with more females. The results of the female removal experiments (n = 10 females) indicated that females competed for better territories rather than larger mates. In contrast, males abandoned the territories once the females emigrated. These results strongly suggest that males defend females rather than sites and compete for larger and a greater numbers of females. Thus, in S. chrysopterum, female defense polygyny occurs in the absence of female aggregation.     

Bigamy or monogamy with maternal egg care in the triggerfish,Sufflamen chrysopterus

Reproductive behavior and mating systems of the triggerfish,Sufflamen chrysopterus (Balistidae), were studied on the fringing reef of Sesoko Island, Okinawa. Both males and females maintained territories against consexual adults, feeding on benthic animals within their own territories. Each male territory overlapped one or two female territories, with mating occurring between the cohabitants. The monogamous males were smaller and foraged more frequently than the bigamous ones, suggesting that the former allocated more energy to growth rather than to improving reproductive success. Pair spawning occurred around sunrise, females only taking care of the demersal eggs until hatching, which occurred around sunset of the same day. On spawning days females foraged less frequently than usual, but as frequently as males. Females spawned at intervals of 5–7 days, usually shifting sites within their territories. Thus both feeding and spawning sites were available for females within their territories, providing males with the opportunity to monopolize females by defending their territories.     

Conflicting Territory Use in Males and Females of a Monogamous Ungulate,the Kirk's Dikdik

In monogamous mammals it is often unclear why males do not defend larger territories to attract more than one female. I investigated the territoriality of the monogamous Kirk's dikdik, Madoqua kirki, a dwarf antelope, in which food resources increase with territory size and some males defend enough resources for more than one female. Yet, all males are paired monogamously. When males were removed from small territories, their female partners spent more time outside of their territories than females in large ones. When females were removed, their male partners almost never left. Pairs in small territories spent more time together than pairs in large ones. Paired males left mostly together with their females, apparently not on their own initiative. Presumably because females in small territories left more often, their males spent more time outside in the female's company than males in large territories. I argue that males in smaller territories can keep better track of their females and that they can effectively reduce their females' time outside. Male intrusion pressure was unrelated to territory size, but it increased in the presence of unguarded females. If large territories decrease the ability to mate guard, and if unguarded females attract competing males, then defending large territories may be uneconomical, even it they could attract more than one female. On the other hand, territories must be large enough to satisfy the requirements of a single female.     

Breeding site fidelity in willow ptarmigan: the influence of previous reproductive success and familiarity with partner and territory

Summary Breeding site fidelity is high in willow ptarmigan: only 9% of males and 31% of females switched territories between years. Unpaired males were more likely to switch territories than paired males. For paired males, survival of their previous partner and reproductive success in year x did not influence probability of switching in year x+1. A female was more likely to switch territories if her previous partner disappeared. If her partner returned, she had a higher probability of switching if she did not produce chicks the previous year. Most hens moved to the territories of older males, although hens paired with unfamiliar older males did not have higher reproductive success than those paired with yearlings. Individuals that paired with their previous partner laid earlier and produced heavier chicks than those paired with unfamiliar partners. Excluding birds paired with familiar partners, survival and reproductive success in year x+1 was similar for males and females that did or did not switch territories. Males had a higher probability of producing chicks after switching than before, but females were more likely to lose their clutch after switching. For both sexes, birds that switched territories were as successful as the birds that replaced them on their former territories. We conclude that high site fidelity in willow ptarmigan is maintained because of the benefits of pairing with a familiar partner.     

Mating patterns of female Leon Springs pupfish Cyprinodon bovinus

In a field study of Leon Springs pupfish Cyprinodon bovinus, two questions about female promiscuity were investigated. First, were females selective in the males with whom they spawned or were they unselective, spawning randomly among males? Second, how promiscuous were the females, i.e. with how many males did they spawn? If simply spawning with many males maximized a female's reproductive success, then females might be expected to spawn randomly with as many males as possible. Alternatively, if females were selective but engaged in multiple mating, they would limit their spawning to preferred males. In the only wild population of this endangered fish, breeding males defend closely associated territories in the shallow margins of a single desert pool. No territories were observed elsewhere in the pool. Therefore, all territorial males were present simultaneously and females could survey all of them, depositing any number of eggs with one, a few or many males. Rather than spawning randomly, females surveyed many males first, visited relatively few males and ultimately spawned with a small fraction of those available males. With increasing numbers of spawns, however, females increased the number of different mates with whom they spawned. Thus, females showed a bet‐hedging tactic of having a narrow mate preference while also laying eggs in the territories of other males, possibly to reduce egg predation and to avoid inbreeding.     

Female Mate Preference for Males Having Long and Symmetric Fins in the Bower-holding Cichlid Cyathopharynx furcifer

Female mate preference in a bower-holding cichlid, Cyathopharynx furcifer, was studied in Lake Tanganyika. Most males held territories with crater-shaped bowers in sand, but some males held territories without bowers. Territories were distributed adjacently and females visited them to spawn. After engaging in circling behaviour with the male, a female deposited eggs in the bower. Soon after spawning, the female picked the eggs up into her mouth and brooded them in places away from male territories. Female mate choice appeared to follow three steps: 1) females visited only bower-holding male territories, and more frequently visited territories of males that performed courtship displays at a higher frequency and had longer pelvic fins; 2) females preferred to start circling with males having longer and more symmetrical pelvic fins; 3) females chose males with more symmetrical pelvic fins as their mates. Less than 7% of females that visited male territories spawned eggs in the bowers. In contrast to other bower-holding species, bower size did not correlate with male reproductive success in C. furcifer. Bowers may therefore be essential as spawning sites or may function as a species recognition character for females. Female choice may be dependent instead on males having long and symmetrical pelvic fins apparent during the circling behaviour carried out in the bowers.     

Timing of mate-locating by males in relation to female activity in the carpenter beeXylocopa varipuncta (Hymenoptera: Apidae)

Males of the carpenter beeXylocopa varipuncta Patton wait for females to visit them as they hover at landmark territories along ridgelines on some spring afternoons. While hovering, males advertize their presence by releasing a pheromone that attracts passing females. If males have limited time to invest in territorial hovering and signaling, then they should engage in these activities more often at times when mate-searching females are most likely to visit landmark territories. The number of females flying near male territories varies greatly over the course of afternoons and from day to day. Measures of female activity and male territorial activity were highly correlated at one study site, both in terms of changes within afternoons and in terms of day-to-day fluctuations. This result supports the hypothesis that males ofX. varipuncta time their mateattracting behavior to maximize contacts with receptive females.     

Territorial and mating behavior ofXylocopa fimbriata F. andXylocopa gualanensis Cockerell from Costa Rica

The territorial and mating behavior of two Xylocopaspecies X. fimbriataF. and X. gualanensisCockerell from Costa Rica are described. Male territorial activity was common during February through April in the higher areas of a dry forest savanna. Both species maintained territories at the same location within hollows in the foliage of Ardesia revolutaH.B.K., a small evergreen tree which occurs commonly in the dry forest. Xylocopa fimbriatamaintained territories during early morning hours, whereas the territorial period for X. gualanensisoccurred during late afternoon. Males of both species infrequently marked territories. It is suggested that a secretion released from their mesosomal glands is wiped onto their legs during frequent leg-body and leg rubbing which occurs while males hover. The secretion is occasionally applied to marking points on vegetation by their secretion-contaminated legs. Males repeatedly marked select locations when females were near their territories regardless of whether females were searching nearby branches for nesting sites, gathering provisions, or passing through their areas. Some females oriented to and entered certain male territories. An example of female mate choice in selection of male territories is provided. Females flew into male territories from 2–4 cm down-wind of male scent-marked locations. As females approached these locations, males hovered downwind in close proximity to the females' approach path. As females hovered a few centimeters downwind of the scented focal point, males approached from downwind and mounted. The males behavior and the role of scent-marking in the reproductive behavior of lek polygyny is discussed.     

Female preference for males that have exclusively marked or invaded territories depends on male presence and its identity in the subterranean rodent Ctenomys talarum

Territorial scent‐marking provides chemical records of male competitive interactions that are available to females, who gain valuable information to assess and identify best quality partners. In this context, the solitary subterranean rodent tuco‐tuco (Ctenomys talarum) offers excellent possibilities to evaluate the effects of male exclusive scent‐marking of territories on female assessment. For evaluation, we used wild caught individuals of C. talarum, manipulated their scent marks within the territories in captive conditions and staged preference tests where females were able to choose between exclusive and invaded territories. The evaluation was performed in two scenarios considering the identity of the intruder scent mark: territories invaded by a strange male and territories invaded by a neighbour male. Females investigated the chemical cues deposited on the substrate of the exclusively marked territory more frequently. Next, females displayed equal interest to scent samples of both males presented in a Y‐maze. Finally, when females could gain access to both individually isolated males and their scent‐marked territories, they spent more time within invaded territories despite they visited them with the same frequency. Moreover, females tried to get in contact by scratching the mesh of the owner of the invaded territory more frequently. We found that females of C. talarum evaluate the homogeneity (exclusiveness) of scent marks within a male territory and then show preferences in relation to the identity of the intruder's scent –whether strange or neighbour.     

Evolution of Female Egg Care in Haremic Triggerfish,Rhinecanthus aculeatus

Paternal care is predominant among telcost fishes with external fertilization. This study describes maternal care in a haremic coral-reef fish and discusses the possible factors leading to its evolution. Both sexes of the triggerfish Rhinecanthus aculeatus (Balistidae) maintained territories; some individuals for more than 8 years. Each male's territory overlapped 2-3 female territories. Pair-spawning occurred around sunrise. Only females cared for the demersal eggs until hatching, which occurred just after sunset on the day of the spawning. No predation was observed on eggs under the maternal care, but experimental removal of parental females decreased the hatching rate to nearly zero. Egg-guarding females foraged as frequently as males, but less than half of non-spawning days. Spawning occurred only in the periods of about 1 wk around the new or full moon, and individual females spawned up to three times in each period. Thus, the maternal care did not significantly affect the duration of the females' spawning intervals, while males would suffer mate loss if they performed parental care. In this situation, maternal care should be the evolutionarily stable strategy. Evolutionary transition from no care to maternal care and then to biparental care is suggested in the Balistidae.     

Landmark Territoriality in the Neotropical Paper Wasps Polistes canadensis (L.) and P. carnifex (F.) (Hymenoptera: Vespidae)

Male Polistes canadensis and P. carnifex aggregate along crests of prominent ridges in Santa Rosa National Park, Costa Rica. At these sites males of both species defend territories (trees and shrubs) by chasing conspecific rivals. Territories do not contain nests or resources that are collected by females. Chasing by territorial males reduces the amount of time spent by intruders in a territory. I describe and contrast male territorial behavior of both species. Some male P. canadensis are territorial while others in the same area exhibit patrolling behavior, flying from one occupied territory to another. Males of P. carnifex exhibit territoriality only. Patrolling in P. canadensis is an outcome of relatively high male density along the ridge, rendering territories in short supply, as shown by the observation that experimentally vacated territories are seized rapidly by formerly patrolling males. Due to a high intraspecific intrusion rate, territorial male P. canadensis spend less time perching and more time flying and chasing intruders from their territories than do male P. carnifex. Males of these two species also differ in the placement of their territories along the ridgeline; P. canadensis occupy territories in saddles while P. carnifex occupy those at peaktops. I show that this divergent spatial pattern is not maintained by competitive exclusion of either species by the other, and I discuss alternative explanations for their separate spatial distributions. Comparative data suggest that males are territorial because females restrict matings to within territories, and I discuss alternative hypotheses to explain this bias in female behavior.