The onset of seasonal quiescence in the female Bennett's wallaby (Macropus rufogriseus rufogriseus)

The breeding season of the non-lactating Bennett's wallaby terminates when animals enter the state of seasonal quiescence. To examine this transition, pouch young were removed from females at intervals which were 3, 4 or 8 weeks (6, 11 and 8 animals respectively) after the winter solstice. Within 48 days, 3, 1 and 1 females gave birth respectively, indicating that these animals were not in seasonal quiescence when pouch young were removed. Those animals which did not give birth were either in seasonal quiescence or had undergone a non-pregnant cycle. To differentiate between the 2 possibilities, techniques which would ensure the detection of pregnant and non-pregnant cycles were assessed in 8 females during the breeding season. As has been previously reported for the wallaby, changes in peripheral progesterone concentrations and the vaginal smear occurred during pregnant and non-pregnant reproductive cycles. In addition, mating was detected by marking the male with a mixture of coloured crayon and paraffin wax. It was concluded that reproductive cycles in female wallabies could be monitored by collecting blood samples 2 times each week for progesterone determination and daily examination of females for mating marks. These techniques were then used to study the onset of seasonal quiescence in 9 females. All animals continued to show reproductive cycles after the winter solstice and it was not until 10 weeks after the winter solstice that all animals were in seasonal quiescence. This represents an increase in the duration of the breeding season over that previously reported for this species.     

Plasma concentrations of progesterone and testosterone in captive woolly opossums (Caluromys philander)

Plasma testosterone and progesterone concentrations were measured in captive woolly opossums, a didelphid marsupial originating from neotropical forests in French Guyana. Although not exposed to cyclic environmental conditions as in the field, both sexes exhibited spontaneous circannual changes in sexual hormones. Males showed synchronous variations in plasma testosterone characterized by significant elevated concentrations during April and September (8.6 +/- 1 ng/ml, N = 5) and lower levels from May to July (3.6 +/- 0.4 ng/ml). In females, synchronous periods of 2-3 successive oestrous cycles occurred. Between these periods, females remained acyclic. The oestrous cycle, determined by urogenital smears, lasted 28-45 days (n = 14) and included a 20-day spontaneous luteal phase in which progesterone concentrations reached 30-40 ng/ml plasma. Even though testosterone concentrations in paired males increased significantly in response to oestrous periods of the paired females, spontaneous circannual rhythms of sexual activity were not well synchronized between the sexes in captivity. When compared to field data, sexual activity of captive animals followed a pattern similar to that in wild animals, without any changes in males but with a delay of 3 months in females.     

Seasonal changes in testosterone and corticosterone levels in four social classes of a desert dwelling sociable rodent

Animals have to adjust their physiology to seasonal changes, in response to variation in food availability, social tactics and reproduction. I compared basal corticosterone and testosterone levels in free ranging striped mouse from a desert habitat, comparing between the sexes, breeding and philopatric non-breeding individuals, and between the breeding and the non-breeding season. I expected differences between breeders and non-breeders and between seasons with high and low food availability. Basal serum corticosterone was measured from 132 different individuals and serum testosterone from 176 different individuals of free living striped mice. Corticosterone and testosterone levels were independent of age, body weight and not influenced by carrying a transmitter. The levels of corticosterone and testosterone declined by approximately 50% from the breeding to the non-breeding season in breeding females as well as non-breeding males and females. In contrast, breeding males showed much lower corticosterone levels during the breeding season than all other classes, and were the only class that showed an increase of corticosterone from the breeding to the non-breeding season. As a result, breeding males had similar corticosterone levels as other social classes during the non-breeding season. During the breeding season, breeding males had much higher testosterone levels than other classes, which decreased significantly from the breeding to the non-breeding season. My results support the prediction that corticosterone decreases during periods of low food abundance. Variation in the pattern of hormonal secretion in striped mice might assist them to cope with seasonal changes in energy demand in a desert habitat.     

Seasonal and sexual dimorphisms in the green anole forebrain

During the breeding season, male green anole lizards extend a throat fan (dewlap) in courtship. This behavior is facilitated by testosterone (T). Females extend a much smaller dewlap less often, even with the same dose of T. During the nonbreeding season when T is low, dewlap extension is reduced. To determine if parallels exist between structure and function, we investigated neuron soma size and density in the preoptic area (POA) and ventromedial nucleus of the amygdala (AMY), which are involved in the display behavior, in breeding and nonbreeding males and females. Cells from breeding animals were larger than cells from nonbreeding animals, but they were not sexually dimorphic. No significant effects existed in neuron density. This experiment indicates that portions of the anole forebrain important for sexual behavior are plastic and might be influenced by seasonal changes in steroid hormones. To investigate whether T can reverse the seasonal difference in soma size in both sexes, gonadectomized nonbreeding anoles were implanted with an empty or T propionate-filled capsule; animals were also tested for male-typical courtship behavior. Males and females treated with T had higher rates of dewlap extension, but across treatment groups these rates were greater in males. Neuron soma size in the POA and AMY was larger in males than females, but no effects of treatment were detected. Taken together, the results indicate that T can stimulate behavior in the nonbreeding season and suggest that a dissociation exists between the regulation of the courtship display and soma size of relevant brain regions.     

Olfactory communication and the importance of different odour sources in the ferret (Mustela putorius f. furo)

Olfactory communication has a major role in solitary carnivores, particularly during the breeding season. Scent-marking behaviour in ferret (Mustela putorius f. furo) has been well studied, but not the significance of the deposited odours for the animal. We compared the interest in urogenital, anal and body odours of opposite-sex conspecifics between male and female ferrets and investigated both sexes’ discrimination ability (familiar vs. unfamiliar) for these three opposite-sex odours and the use of its ability by females during the breeding season. Males are more interested than females in the urogenital odour of the opposite sex. Male body odour probably did not interest females and males are somewhat interested in the female one. However, anal odour of the opposite sex is as interesting for males as for females and both sexes are able to discriminate familiar anal odour from an unfamiliar one. Moreover, females could use this ability when they must choose to have some sensory contact between olfactory familiar and unfamiliar males since they prefer to be in contact with the former. These results are coherent with behavioural differences observed between male and female solitary mustelids in the field and in captivity, and the consortship success of resident males compared to roaming males.     

Reproductive biology of the Red-necked wallaby (Macropus rufogriseus banksianus) and Bennett's wallaby (M. r. rufogriseus) in captivity

Bennett's wallaby ( Macropus r. rufogriseus ) of Tasmania give birth from late January to early August in marked contrast to the Red-necked wallaby ( M. r. banksianus ) of mainland south-eastern Australia which produced young in all months. Within the breeding season however, the lengths of the oestrous cycle and gestation period are similar in the two forms and did not differ by more than 0.5 days. The gestation period of about 30 days extended to almost the length of the oestrous cycle of approximately 33 days. Birth was closely followed by mating which normally resulted in fertilization and subsequent embryonic diapause. Renewed blastocyst development was initiated by removal or loss of a pouch young and birth followed about 27 days later.
Unlike other macropodids with a similar breeding pattern, birth, as a result of renewed blastocyst development near the end of a large young's pouch life, did not occur within a day or two of the permanent emergence of the young, but followed 16 to 29 days later. In M. r. rufogriseus , young that left the pouch permanently in the non-breeding period were not replaced by new young until the beginning of the next breeding season two to four months later, and blastocysts resulting from mating of females without pouch young at the end of the breeding season remained quiescent until the next breeding season five to eight months later.
Females of both subspecies first mated at an age of about 14 months, and males were producing mature spermatozoa by about 19 months.
Young first left the pouch for short periods at about 230 days of age and permanently at about 280 days.
Observations are also given on reproductive behaviour, interpretation of vaginal smears, sex ratio of young, selection of teat by pouch young, and development of morphological features in known-age young that may be used as an aid in age determination.     

Male influence on oestrous cycles in female woolly opossum (Caluromys philander)

Plasma progesterone concentrations and the occurrence of oestrous cycles were studied in isolated woolly opossums subsequently subjected to male influences during a 40-day period. Pairing (N = 48) or exposure to male urine (N = 15) resulted in all females exhibiting oestrous during the stimulation phase, providing evidence that the activation of ovarian activity in the woolly opossum involves pheromonal cues from males. The latency of occurrence of oestrous in stimulated females depended upon their sexual state before male stimulation. In anoestrous females, the mean latency was 20.7 +/- 0.9 days (N = 35), a value which agrees with the duration of the follicular phase. In females which first entered oestrous before male stimulation, the latency of induced oestrous was inversely correlated to the date of occurrence of the previous oestrous. The inter-oestrous interval was normal (38.1 +/- 1 days, N = 5) when females were in oestrous at the beginning of male stimulation. In contrast, the inter-oestrous interval was significantly shortened (28.7 +/- 2 days, N = 7) or lengthened (51.1 +/- 1.7 days, N = 16) depending on whether females were in the luteal or follicular phases at the beginning of male stimulation. During pairing several females became pregnant and gave birth 24 +/- 0.9 days (N = 13) after copulation. In the woolly opossum, the response to male influences involves mechanisms similar to those observed in eutherians and results in enhancement and synchronization of oestrous cycles in females. Pheromonal interactions could play an important role in synchronizing oestrous cycles in wild females during the dry season, a period when animals regroup to feed on spatially localized food resources.     

Testosterone and year‐round territoriality in tropical and non‐tropical songbirds

Past studies have suggested a fundamental difference in testosterone concentrations between tropical and northern latitude male birds, with the convention being that males in the tropics express much lower levels of testosterone. However, recent comparative studies have shown that tropical males with a short and synchronous breeding season (i.e. a breeding season typical of northern species) express maximum testosterone levels similar to those of northern latitude birds. Here, we ask the converse: do northern latitude songbirds that express a defining life‐history characteristic typical of the tropics, i.e. year‐round territoriality, have an annual testosterone profile similar to that of tropical songbirds? For the few year‐round territorial species for which data are available, we found that seasonal testosterone profiles and seasonal maxima in plasma testosterone were similar between males of tropical and non‐tropical species. For example, males of both groups expressed seasonal maxima during the period when females were fertile, and testosterone levels at this time were similar. In contrast, this and other studies show that species with seasonal territories typically express maximum testosterone levels earlier in the breeding cycle, when territories are first being established. Taken together, we suggest that specific life‐history traits may play a more important role in determining testosterone profiles of tropical and non‐tropical birds than breeding latitude and encourage further studies to allow for more formal comparisons.     

Sex-specific patterns in body condition and testosterone level changes in a territorial migratory bird: the Bluethroat Luscinia svecica

The Bluethroat Luscinia svecica is a migratory passerine that exhibits a socially monogamous pair bond and a high level of parental care. Males are territorial both when wintering and breeding whereas females are territorial only in winter. We investigated changes in body condition and testosterone levels during successive life-history stages and determined their relationships. Sex-specific patterns were observed in the variation in body condition and testosterone level. Male body condition varied mainly during the winter. It peaked at the onset of the prenuptial moult and then decreased, whereas it remained stable throughout breeding. In contrast, female body condition varied mainly during the breeding season. It increased during the prelaying stage and then abruptly decreased until fledgling provisioning. As in other monogamous and territorial passerines, testosterone levels in Bluethroat males were low during winter, increased in late winter, peaked during the prelaying stage and then decreased when provisioning young. In wintering females, territorial competition caused testosterone levels to rise. These females were able to produce territorial vocalizations and exhibit aggressive postures. Females showed higher mean testosterone levels than males when wintering whereas the opposite was observed when breeding. Our data from wintering female Bluethroats support the 'challenge hypothesis' under which high testosterone levels are associated with periods of social instability, and testosterone can regulate female territorial behaviour during this period.     

The hormonal and behavioral response to group formation, seasonal changes, and restraint stress in the highly social Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus) (Chiroptera: Pteropodidae)

This study examined behavioral and physiological responses (changes in inter-animal spacing, total glucocorticoids, testosterone, and body mass) to the formation of breeding and same-sex groups in two bat species, the socially gregarious Malayan Flying Fox (Pteropus vampyrus) and the less social Little Golden-mantled Flying Fox (Pteropus pumilus). We hypothesized that social instability, especially in the breeding groups and especially in P. vampyrus, would result in elevated glucocorticoids and that social facilitation of breeding and/or male-male competition would result in persistently higher levels of testosterone in breeding males. Seasonal rhythms in all measures were also predicted, and the glucocorticoid stress response was expected to vary by sex, season, and group type. Nearly all animals responded to group formation with elevated glucocorticoids, but, for breeding animals (especially aggressive male P. vampyrus), these responses persisted over time. In both species, breeding group formation resulted in elevated testosterone in males. Glucocorticoids, testosterone, testes volume, and body mass generally peaked in the breeding season in males (late summer and early autumn), but the seasonal glucocorticoid peak in females occurred in late winter and early spring. All animals responded to restraint stress with elevations in glucocorticoids that largely did not differ by sex, time of year, reproductive condition, group type, or, in lactating females, the presence of her pup. Changes in both behavior and physiology were more evident in P. vampyrus than in P. pumilus, and we believe that their underlying social differences influenced their responses to group formation and to the changing seasonal environment.     

Effects of testosterone treatment and season on the frequency of dewlap extensions during male-male interactions in the lizard Anolis sagrei

Lizards of the genus Anolis extend and retract a large and often brightly colored throat fan called a dewlap. The dewlap in most anoles is a sexually dimorphic structure. It is larger in males than females and males use the dewlap more frequently and in more contexts than do females. In the present study we investigated whether plasma testosterone (T) levels and season affect the frequency of dewlap use in male--male interactions in the brown anole, Anolis sagrei. We manipulated plasma T levels by implanting adult, castrated males with pellets that delivered no T, a moderate dose of T, or a high dose of T. In tests with stimulus males, castrated males that were treated with a moderate or with a high dose of T had a significantly higher frequency of dewlap extensions than did castrated males that received no T. However, the frequency of dewlap extensions in castrated males that received the high dose of T did not differ significantly from castrated males that received the moderate dose of T or from non-castrated control males. Males captured during the breeding season and tested in the laboratory had a significantly higher frequency of dewlap extensions than did males captured in the postbreeding season and tested in the same manner. These results suggest that plasma T levels affect how frequently males of A. sagrei extend their dewlaps in male--male interactions and that seasonal changes in male dewlap use may be due to seasonal differences in plasma T levels.     

Seasonal changes in testosterone levels in wild Mexican cottontails Sylvilagus cunicularius

We studied serum testosterone levels in the endemic Mexican cottontail, Sylvilagus cunicularius, which has been reported to show seasonal breeding. Animals were trapped in the wild and in a field enclosure in the National Park La Malinche in central Mexico over a period of five years. Serum testosterone (T) levels were quantified by ELISA from blood samples. T levels of adult males were lowest around 4 months after the onset of the annual reproductive season and were already high prior to the onset of breeding. As expected, the T levels of adult females were consistently lower than in males, and there were no differences in T level with respect to female reproductive state. There were no detectable sex-specific differences in juveniles and subadults, but there was a marked increase in T levels between juvenile and adult males. Overall, our study clearly reflects and confirms the seasonal breeding strategy of this species, showing high similarities to the much better studied European rabbit.     

Virilization of the male pouch young of the tammar wallaby does not appear to be mediated by plasma testosterone or dihydrotestosterone.

Virilization of the male urogenital tract of all mammals, including marsupials, is mediated by androgenic hormones secreted by the testes. We have previously demonstrated profound sexual dimorphism in the concentrations of gonadal androgens in pouch young of the tammar wallaby Macropus eugenii during the interval when the urogenital sinus virilizes. To provide insight into the mechanisms by which androgens are transported from the testes to the target tissues, we measured testosterone and dihydrotestosterone in plasma pools from tammar pouch young from the day of birth to Day 150. Plasma testosterone levels were measurable (0.5-2 ng/ml) at all times studied, but there were no differences between males and females. These low concentrations of plasma testosterone appear to be derived from the adrenal glands and not the testes. Plasma dihydrotestosterone levels in plasma pools from these animals were also low and not sexually dimorphic. We conclude that virilization of the male urogenital tract cannot be explained by the usual transport of testosterone or dihydrotestosterone in plasma but may be mediated by the direct delivery of androgens to the urogenital tract via the Wolffian ducts. Alternatively, circulating prohormones may be converted to androgens in target tissues.     

Changes in female reproductive condition following male take-overs in a colony of hamadryas and hybrid baboons

This paper describes and discusses events observed in the Madrid colony of hamadryas and hybrid baboons, when a novel group of 3 adult males, 3 adult females and 1 unweaned infant was introduced to the resident colony comprising 12 adult females, 11 juveniles and 6 unweaned infants. Novel males took over resident adult females in any reproductive condition, and all acyclic females (i.e. lactating, pregnant and immature) exhibited a dramatic enhancement of sexual activity. Lactating females shortened their postpartum amenorrhoea periods and resumed oestrous cycles around day 14 following the introduction of the novel males, without infanticide occurring. Their return to breeding condition was not affected by the age of their current infant or the day they were taken over by the males. A female in an early stage of pregnancy aborted spontaneously and resumed oestrus on day 26. The other pregnant female significantly shortened her gestation time, delivered a viable infant on day 13, and resumed breeding activity 39 days post-partum (on day 52), while suckling her infant. A cycling female adopted and suckled a 74-day-old infant, continued showing oestrous cycles and conceived. Immature females reached menarche significantly earlier than expected and only then joined one of the newly established harem units. It is argued that the observed enhancement of sexual activity was not imposed by the males' aggressive behaviour but rather was a spontaneous female response to male novelty. This single causal factor was potent enough to override the role that nutrition and lactation normally play in the control of the females' reproductive activity. Differences in latency until the appearance of the response were probably due to the different constraints imposed by the female's current reproductive state. It was also hypothesized that when (a) sexual swellings are attractive to males, (b) novel males are especially active in the process of bonding with new females, and (c) males are important sources of coalitionary support, females might by developing sexual swellings compete more successfully against other females and attain a higher position in the female hierarchy of the newly established unit. This would have the ultimate effect of increasing their potential reproductive success. In several cases females did gain socially by coming into oestrus, but attained no immediate reproductive advantage.     

Differences in Fecal Androgen Patterns of Breeding and Nonbreeding Kori Bustards (Ardeotis kori)

To better understand breeding conditions to promote reproduction in captive kori bustards, fundamental endocrine studies measuring fecal androgen metabolites in male and female kori bustards were conducted. Feces collected weekly from males and females were analyzed for testosterone using enzyme‐linked immunoassay. Results from adult males (n = 5), adult females (n = 10), immature males (n = 10), and immature females (n = 10) revealed seasonally elevated testosterone concentrations in fertile, but not nonfertile adult males and females (P > 0.05). Adult females that were not maintained in a breeding group, or that did not produce eggs, did not demonstrate increases in testosterone compared to egg laying counterparts. In males, but not females, seasonal testosterone increases were accompanied by weight gain. Peaks in male fecal androgen metabolites ranged from 10‐ to 22‐fold higher than nonbreeding season (181.5 ± 19.1 vs. 17.0 ± 0.94 ng/g; P < 0.05). Mean breeding season values for adult males were 83.6 ± 6.1 ng/g vs. nonbreeding season values of 12.3 ± 0.73 ng/g (P < 0.05). In females, average breeding season testosterone concentrations were approximately 4‐fold higher than nonbreeding season (55.9 ± 6.0 vs. 14.5 ± 1.8 ng/g), with peaks 10‐ to 30‐fold higher. Results show that noninvasive fecal androgen metabolite analysis can provide a means of predicting fertility potential of male and female kori bustards and might be utilized to assess effects of modifying captive environments to promote reproduction in this species. Zoo Biol. 32:54‐62, 2013. © 2012 Wiley Periodicals, Inc.     

Annual reproductive activity of Caiman crocodilus fuscus in captivity

We describe the gonadal and abdominal fat storage cycles for a population of Caiman crocodilus fuscus kept in captivity at a breeding farm. The reproductive cycles of males and females are clearly seasonal. Vitellogenic follicles were observed in females year‐round; however, the follicular growth that produces the preovulatory follicles for the breeding season occurs in November–May (most of the dry season to the beginning of the rainy season). These months correspond with the presence of physiologically reproductive males. Oviposition takes place during the wet season (April–August) and the births occur at the end of this season (September–November). Fat storage was not related to reproductive activity, and did not vary by month or between sexes. Although environmental and hormonal conditions have been improved for these captive animals, they maintain a strongly seasonal reproductive cycle, historically fixed to oviposit in the most favorable period for nest construction and hatching during the rainy season. Zoo Biol 22:121–133, 2003. © 2003 Wiley‐Liss, Inc.     

Why are Male Columbian Ground Squirrels Territorial?

Male territorial defence is a component of many vertebrate mating systems and is often regarded as a tactic for acquiring mates. Traditionally considered within the context of overt site‐specific defence, territoriality actually may have several components which encompass a variety of behavioural tactics (e.g. post‐copulatory mate‐guarding, defence of resources that females need, defence of area around females) that underlie a mating system. The purpose of our study was to evaluate such influences on the territorial behaviour of male Columbian ground squirrels in southwestern Alberta, Canada. Males were dominant and territorial if they defended a minimum convex polygon activity range by chasing other males more within the activity range than they were chased. Subordinate males had no territory and were chased throughout their ranges, but they competed for mates by increasing chases in their activity range when nearby females were oestrous. Dominant males exhibited conditional breeding tactics, tending to chase other dominant males from their territory when nearby females were oestrous, but travelling outside their activity ranges to chase subordinate males when females were not oestrous. Although females mated first with a dominant male on whose territory they resided (and in order from oldest to youngest if several territories overlapped), mating pairs were not exclusive, as females usually mated with additional males. Males also guarded females after copulation and defended females directly just before oestrus, rather than defending territory per se during those times. Thus, males possess a repertoire of behaviours that complement site‐specific territoriality, and territory ownership serves to facilitate a first mating with females that live on the territory.     

Annual physiological changes in individually housed squirrel monkeys (Saimiri sciureus)

This study investigated whether annual changes in physiology occur in individually housed squirrel monkeys (Saimiri sciureus). Physiological measures were monitored for 20 months. Over the course of the study, all individually housed males and females exhibited clear annual changes in gonadal and adrenal hormone levels, and males exhibited species‐typical changes in body weight. Females exhibited a typical pattern of hormonal changes, with elevations in gonadal steroids occurring during the same months as elevations in cortisol. Males, however, exhibited an atypical pattern, as elevations in hormone levels were not synchronized with each other; rather, elevations in testosterone occurred out of phase with changes in cortisol and body weight. The timing of annual events in individually housed subjects was compared to that in nearby social groups, in which the timing of the breeding season from year to year was determined by social group formations and was outside the naturally occurring breeding season. Elevations of ovarian and adrenocortical hormones in individually housed females were synchronized with indices of breeding in heterosexual social groups. Similarly, weight gain in males was associated with elevations in cortisol and, as with socially housed males, tended to precede seasonal breeding in the social groups. In contrast, annual testosterone elevations for individually housed males were not synchronized with breeding in nearby social groups. We conclude that direct physical interaction is not required for the annual expression of breeding readiness. Synchrony of seasonality among squirrel monkeys may be accomplished by distant social cues in females, but males may require physical interaction for complete synchrony of annual physiological changes. Am. J. Primatol. 47:93–103, 1999. © 1999 Wiley‐Liss, Inc.     

The influence of social cues on the reproductive endocrinology of male brown-headed cowbirds: field and laboratory studies

Captive male brown-headed cowbirds exposed to long days exhibit gonadal growth and have elevated plasma testosterone (T) levels. This photoperiodic response is enhanced if males are housed with female cowbirds: Photostimulated males with females increase plasma testosterone levels sooner than do individually housed photostimulated males. Peak plasma T levels are similar in both groups, although peak levels are maintained longer in males housed with females. The gonadal cycle is similarly affected; males in the presence of females have earlier gonadal recrudescence and maintain mature gonads longer than do photostimulated males without females. Plasma corticosterone levels increase in the unpaired males, suggesting that removal of social cues is stressful for these birds. Free-living paired males have significantly higher plasma testosterone levels than do unpaired/unknown males early in the season, when social relationships are being established; the levels are similar thereafter. There is no difference between the two groups in testicular maturation rates; nor do they differ in plasma corticosterone levels at any time of the season. These results suggest that social stimuli are important in modulating the secretion of testosterone in males early in the season when pairing occurs, and possibly late in the season as well, probably to prevent termination of breeding prior to that of females.     

Monogamy and sex role reversal in the pipefish Corythoichthys haematopterus

We observed the mating pattern and social behaviour of the pipefish Corythoichthys haematopterus in temperate waters of Japan during three successive breeding seasons. Males cared for a clutch in their brood pouch for 9-19 days until hatching and had several broods in the season with nonbrooding intervals of only 1 or 2 days. The population sex ratio was female biased and some females were always excluded from reproduction. Although males were sometimes courted by unmated females together with their regular partners, they always mated with the latter. The pair bond was maintained until the next season if both members survived. When males lost their partners, they remated with neighbouring unmated females within a few days. In contrast, widowed females remained unmated for a long time. Females had larger home ranges and were more active in courtship displays than males. This pipefish provides the first example of sex role reversal among monogamous syngnathid fish. We suggest that mate guarding by females is a primary proximate factor for maintenance of monogamy in this fish. Copyright 2001 The Association for the Study of Animal Behaviour.