Reproductive biology and management of captive black-footed ferrets (Mustela nigripes)

A captive breeding program is being conducted with black-footed ferrets (Mustela nigripes), an endangered species. Results of 5 years of study are reported. Simple, but specialized, nontraumatic handling techniques allowed assessment of reproductive status with minimal stress, which was important in breeding management. Black-footed ferrets are sexually mature and may successfully reproduce in their 1st year. Proestrus lasts approximately 2-3 weeks. Duration of estrus in unbred females was 32–42 days; females usually bred within 20 days. Most breeding activity occurred during April. Mean gestation length was 42.7 days (±0.7, range 42–45 days), litter size averaged 3.0 kits (±1.4, range 1–6 kits), and weaned kits/litter averaged 2.4 (±1.7, range 1–6 kits). Weaning rate of kits was 80%. Sex ratio of kits was essentially 1:1. Productivity was greatest among females ?3 years of age. Rapid expansion of the captive population is possible and will be important for genetic management of the species and for achieving the primary goal of the recovery program, which is to return black-footed ferrets to the wild.     

Lifetime survival rates and senescence in northern elephant seals

The aim of this study was to extend 40 yr of prior demographic work on northern elephant seals (Mirounga angustirostris) at Año Nuevo, California, by including the oldest animals. We used a Bayesian mark‐recapture analysis to estimate lifelong survival and lifespan of a cohort of 372 weaned pups branded in 1985–1987 and resighted until 2008. Annual survival probability of females averaged 86.3%/yr at ages 5–16, then declined until age 21, the age of the oldest female. Male survival was lower, averaging 67.7%/yr from age 1 to age 15, the age of the oldest male. Northern elephant seal females in the expanding population at Año Nuevo live longer than southern elephant seal females (M. leonina) at colonies whose populations are declining. This comparison suggests that high survival of females is a key factor in population growth.     

The Effects of Birth Weight and Maternal Care on Survival of Juvenile Steller Sea Lions (Eumetopias jubatus)

Steller sea lions were listed as endangered following a collapse of the western distinct population beginning in the late 1970s. Low juvenile survival has been implicated as a factor in the decline. I conducted a multistate mark-recapture analysis to estimate juvenile survival in an area of the western population where sea lions are showing signs of recovery. Survival for males and females was 80% between 3 weeks and 1 year of age. Approximately 20% of juveniles continued to be nursed by their mothers between ages 1 and 2 and 10% between ages 2 and 3. Survival for juveniles that suckled beyond 1 year was 88.2% and 89.9% to ages 2 and 3, respectively. In contrast, survival for individuals weaned by age 1 was 40.6% for males and 64.2% for females between ages 1 and 2. Birth mass positively influenced survival for juveniles weaned at age 1 but had little effect on individuals continuing to suckle. Cumulative survival to age 4 was double that estimated during the population decline in this region. Evidence suggests that western Steller sea lions utilize a somewhat different maternal strategy than those in the eastern distinct population. Western adult females generally invest more in their pups during the first year but wean offspring by age 1 more often. This results in better survival to age 1, but greater mortality between ages 1 and 3 compared to the eastern population. Different maternal strategies may reflect density dependent pressures of populations at opposite levels of abundance.     

Reproductive biology and ecology of female polar bears (Ursus maritimus)

Data on age-specific natality rates, litter size, interbirth interval, age of first reproduction, reproductive senescence, age of weaning and cub survival were determined for a free-ranging population of polar bears inhabiting Hudson Bay, Canada, near the southern limit of the species range. Serum progesterone levels were also determined for females at different stages of their reproductive cycle to provide corroborative support for the reproductive parameters described. Animals were live captured using immobilizing drugs and each animal uniquely marked for future identification. First parturition occurred at four or five years of age and the age-specific natality rate increased with age until approximately 20 years, after which it dropped markedly. At least 40% of adult females displayed two-year interbirth intervals and 55% of cubs in their second year were independent of their mother. Mean size of cub litters in spring was 1.9 and 13% of litters had three or more cubs. The natality rate for 5–20-year-old females was estimated as 0.9, higher than that reported for any more northerly polar bear populations where two-year interbirth intervals are rare, fewer than 5% of yearling cubs are weaned and triplet litters occur with less than 1% frequency. Cub mortality was initially high and declined with age. Although cubs in western Hudson Bay were weaned at a younger age and a lighter weight than their counterparts in more northern populations, cub mortality rates were similar. The reason for the marked differences in reproductive parameters in the western Hudson Bay population is not known. We speculate that sea-ice conditions may be sufficiently different to allow weaned bears at a lighter body weight to hunt seals more successfully there than further north.     

Demography,female life history,and reproductive profiles among the chimpanzees of Mahale

Demography provides critical data to increase our understanding of the evolution, ecology, and conservation of primate populations. The chimpanzees of the Mahale Mountains National Park, Tanzania, have been studied for more than 34 yr on the basis of individual identification and standardized attendance records. From this long-term study, we derived the following demographic data: The major cause of death was disease (48%), followed by senescence (24%) and within-species aggression (16%). Fifty percent of Mahale chimpanzees died before weaning. The median ages of female life history variables were: first maximal swelling, 10.0 yr (n = 5); emigration, 11.0 yr (n = 11); and first birth, 13.1 yr (n = 5). The median period of adolescent infertility was 2.8 yr (n = 4) when calculated from the age at immigration to that at first birth. Female fecundity was highest between 20 and 35 yr of age, with an annual birth rate of 0.2. Twenty-six females that were observed from a young age (10-13 yr) to death at various ages (15-40 yr) gave birth to an average of 3.9 and weaned an average of 1.4 offspring. Twenty-five females that were observed from middle age (18-33 yr) to death in older age (31-48) gave birth to an average of 2.7 and weaned an average of 2.0 offspring. The post-reproductive lifespan for female chimpanzees was defined as the number of years that passed from the year when the last offspring was born to the year when the female died, minus 5. Twenty-five percent of old females had a post-reproductive lifespan. The interbirth interval after the birth of a son (x = 72 mo) tended to be longer than that after the birth of a daughter (x = 66 mo). The extent of female transfer, which is a rule in chimpanzees, is influenced by the size and composition of the unit group and size of the overall local community.     

Reproductive behaviour of female Siberian lemmings during the increase and peak phase of the lemming cycle

The reproduction of female Siberian lemmings in the increase and peak phases of the lemming cycle was investigated in connection with a ship-borne expedition along the Siberian arctic tundra. The cycle phase of each studied lemming population was determined using demographic analyses, i.e. current density indices (captured lemmings per 100 traps per 24 h), information on previous density (frequency of old lemming faeces and runways), and information from dendrochronological analyses revealing the most recent winters with a high intensity of willow-stem scarring caused by lemmings. The cycle phase determination was corroborated with data on the age profiles of the populations. The reproductive behaviour of female lemmings differed markedly in relation to cycle phase. In increase-phase populations, all captured females (including young and winter born) were reproducing (had embryos or were lactating), and females started to reproduce early in life, i.e. when <2 months old. By contrast, in peak-phase populations, only 6% of the young females and 63% of the winter-born ones were reproducing, and females did not start to reproduce until they were 5–6 months old. The average number of embryos per reproducing female was significantly higher in increase-phase populations than in peak-phase ones. It is concluded that the rapid population growth in lemmings during the increase phase can largely be explained by the early (young age) reproductive start and, consequently, the shorter generation time, the high proportion of females taking part in reproduction, and the large litters produced. Similarly, a delay in the start of reproduction, a lower proportion of reproducing females, and smaller litter sizes produced by peak-phase lemming populations can contribute substantially to the deceleration in the population increase and possibly lead to a decline. Received: 14 June 1999 / Accepted: 15 November 1999     

Reproductive performance of captive collared peccaries (Tayassu tajacu) in the eastern Amazon

This study analyses reproductive parameters resulting from 74 parturitions in a population of captive collared peccaries in the eastern Amazon (Belém, State of Pará, Brazil) during a 65-month survey. Parturitions were homogeneously distributed throughout the year. The average litter size was 1.85 newborns per parturition. Collared peccary females had a newborn sex ratio of 52.6% females and 47.4% males. The mean age at first parturition was 639, although, the earliest first parturition occurred at age of 381 days. Estimated parturition-conception interval was 58 days. Mean farrowing interval was 196 days and mean production was 1.03 litters and 1.86 newborns per year per female. Mortality rate in newborns less than 2 months old was 26.9% of the total newborn population. Most identified causes of death were abandonment by the mother (22.2%) and trauma due to aggression (13.9%). Newborn deaths occurred mainly during the first two days of life. This study suggests that the collared peccary possesses interesting reproductive parameters for its introduction in captive breeding programmes in the Amazon region.     

Demography of Black Bears in Hunted and Unhunted Areas of the Boreal Forest of Ontario

Abstract: We estimated relative density, survival, and reproduction of American black bears (Ursus americanus) from capture-recapture and telemetry data collected from 1989 to 1999 in the unhunted Chapleau Crown Game Preserve (CCGP) and nearby hunted areas in the boreal forest of Ontario, Canada. We tested for combinations of effects of age class, sex, year, years of food shortage, encumbrance status, and residency (on or off the Game Preserve) on vital rates. Results from live captures, remote captures, and bait-station hit rates indicated that density was highest inside CCGP. Total survival of adult females, subadults, and cubs were similar among residents and nonresidents of CCGP, but yearling survival was lower among CCGP residents. Adult females were approximately twice as likely to die and nearly 10 times as likely to be cannibalized (risk ratio [RR] = 9.62, 95% CI = 2.088–44.29) while encumbered with cubs of the year. Nonresidents of CCGP had greater risk of being harvested (RR = 4.00, 95% CI = 1.19–13.46) but similar risk of being cannibalized (RR = 0.875, 95% CI = 0.300–2.55) relative to CCGP residents, suggesting that harvest mortality was additive to other forms of mortality. Residents of CCGP had older ages at primiparity and lower litter-production rates than bears resident in hunted areas. Few litters were produced in years following food shortages, but litter size was unaffected. We recommend that managers allow for additive harvest mortality and reduced survival of bears encumbered with cubs of the year, and we caution that assuming density-compensatory increases in cub production could optimistically bias estimates of population growth.     

Population structure and spatial pattern in the dioecious shrub Ceratiola ericoides

Abstract. The dioecious shrub Ceratiola ericoides (Florida rosemary) dominates xeric, infrequently burned Florida scrub vegetation, often to the near-exclusion of other woody species. We studied the spatial pattern, age, sex and size structure of four populations in Florida, USA: two coastal scrub populations subject to recurrent local disturbances due to sand movement, and two inland scrub populations in sites periodically burned by stand-replacing fires. The age structure of individual genets was estimated from node counts and used to describe the age structure of the populations. The sex ratio of males to females was not significantly different from 1:1, except within a female-biased coastal population subject to frequent sand movement. Node counts indicated that the mean age for reproductive individuals was 15 - 16 yr for the inland populations and 13 - 16 yr for the coastal populations. In all sites, there was no difference in mean age between males and females. Vegetative reproduction was uncommon except for the least-disturbed coastal population where 72 % of the reproductive individuals originated through layering. Individuals were generally randomly dispersed at the coastal sites, whereas significant aggregation of males and females occurred in the inland sites where the populations were initiated following fire. Seedling recruitment was continuous in the disturbed coastal scrub site, where 35% of the individuals were juveniles. Most juveniles were dispersed from 0.5 to 0.75 m around females. At one of the inland sites, where juveniles comprised 11% of the population, juveniles were clustered at 0.25 to 5.75 m around females. Coastal populations were all-aged, while inland populations were uneven-aged. Recruitment appears to follow periods of disturbance; infrequent fire in the inland populations and continuous sand movement on the coast are factors initiating recruitment.     

Food consumption and growth of California sea lions (Zalophus californianus californianus)

The daily food consumption of 26 California sea lions at the Harderwijk Marine Mammal Park was recorded. Average annual food consumption of males increased with age to stabilize at approximately 4,000 kg/year by the age of 10 years. Females showed a rapid increase in average annual food consumption until they were 3 years old. Thereafter, females housed outdoors averaged 1,800 kg/year, whereas those housed indoors ate approximately 1,400 kg/year. Between the ages of 4 and 7 years, the food intake of males began to fluctuate seasonally, decreasing between May and August. The low food intake in summer was associated with an increase in aggressive behavior. Seasonal fluctuation in the food intake of non‐reproductive females was negligible. Between the ages of 6 and 8 years, many females began to reproduce successfully. Pups were born in May and June. The females' food intake decreased approximately 3 days before birth and ceased the next day. Feeding resumed the day after birth, and by 2 days after birth, it had usually returned to normal. On average, female intake increased in the year of conception, the year of birth, during which the pup was suckled for 6 months, and the following calendar year, during which the pup was weaned. Pups began to eat fish at approximately 11 months of age. When forcefed, they were fully weaned within 2 to 23 days. Male weight and body length increased until approximately 20 years of age. Females increased in body length until 6 years and in weight until approximately 13 years of age. The relationship between standard body length and body weight is given. The heavier an animal is, the lower is its food intake as a percentage of body weight. Zoo Biol 19:143–159, 2000. © 2000 Wiley‐Liss, Inc.     

Effect of age at first mating on primary sex-ratio of the two-spotted spider mite

This paper determines the influence of initial age at mating on the primary sex-ratio of the two-spotted spider miteTetranychus urticae Koch. The specific question asked is whether females that mate at ages beyond ecdysis, or are inseminated at ecdysis by virgin males of set ages, have daughter productions different from control females mated at ecdysis by randomly aged males.Age of initial female mating did not effect survival. Females mated at ecdysis produced more daughters than those mated at age 5 or 10 days. However, daughter production was equal for all female treatments in the post-mating period. All treatments stopped producing daughters at approximately the same age, regardless of when mated. Son production was lowest for females mated at ecdysis. However, son production by females mated when 5 or 10 days old was lower than that of females mated at ecdysis.Male age at mating did not effect female survival. Females exposed to virgin males either 1 or 7 days old produced more daughters than those exposed to males either 11 or 20 days of age. However, a substantial proportion of females in these latter cohorts were not mated. If only mated females are considered, daughter production is the same between all cohorts.Delayed age at first mating influenced sex-ratio only via the females, specifically by changes in daughter production (as opposed to changes in survivorship). Daughters that would have been produced between ecdysis and mating were never recovered by females that mated at ages beyond ecdysis. Thus, these daughters were lost from a sex-ratio viewpoint. Initial age of male mating did not change sex-ratio.     

Reproductive performance of otters Lutra lutra (Linnaeus, 1758) in Eastern Germany: low reproduction in a long-term strategy

The reproductive status of female otters ( Lutra lutra ) was determined from an examination of 518 carcasses collected from Eastern Germany between 1950 and 2001. In Germany otters mate throughout the year. Significant seasonal differences are evident with a distinct peak in summer. Females were in breeding condition between the ages of three and 15 years, with the majority between six and nine years. The mean litter size at birth was 2.36, which was the lowest when compared with other inland populations. There were no significant differences in litter size for regions, seasons or age. Prenatal losses account for 26.31%. The mortality between birth and first appearance of cubs following their mother is about 29%. An effective reproduction rate of 78.4% was deduced from the known age structure of the sample and the proportion of breeding females. Reproductively active females show a significantly higher body condition than non-reproductive females. Following these investigations into breeding status, suggestions on reproductive output and the life history of the otter are discussed.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 329–340.     

Reproductive responses to variation in temperature and food supply by house mice: II. Lactation

Lactating HS/Ibg house mice housed at 21 degrees C and 5 degrees C were assigned to 3 feeding regimes: ad libitum, daily rations of 80% of ad libitum, or 60% of ad libitum beginning on the day that they bore litters. Significant interaction between temperature and food restriction was found for litter survivorship, pup survivorship, litter size, female body weight, and cumulative biomass production. The interaction was due to a magnification of the effects of food restriction at the colder temperature: i.e. mice fed ad libitum were similar at the two temperatures, mice fed the 80% ration differed, and mice fed the 60% ration differed to a greater extent. The dominant response to food restriction was cannibalism by females, which might be associated with the rate of loss in body weight by the female on the days preceding cannibalism of one or more pups. Incidents of cannibalism tended to involve a limited number of pups and to be repeated until a sustainable litter size was reached. In 3 of the food-restricted treatments, females weaned relatively large litters of relatively small pups, but in the most severe treatment (in the group fed the 60% ration at 5 degrees C), the females weaned small litters of large pups. The patterns of cannibalism and variable relative investment in individual pups reflect the aggressive breeding strategy of this classic colonizing species.     

Life history characteristics of brown trout in lakes at different stages of acidification

In 66 Norwegian lakes with allopatric populations of brown trout Salmo trutta that ranged from highly acidified to non-acid (pH 4·7–6·6) there was a significant inverse correlation between mean age and relative abundance (CPUE). As mean annual survival rates were not significantly related to CPUE, this population response may indicate that recruitment was lower in low-density lakes. Age-specific masses were significantly higher in populations that had declined in abundance than in unaffected populations. Mean body mass at sexual maturity in both males and females was inversely correlated with CPUE. Between populations, high age at maturity was associated with high survival rates in both sexes. Age at maturity correlated positively with specific growth rate between the ages of 1 and 2 years in females only.     

Reproductive output, maternal age, and survivorship in captive common marmoset females (Callithrix jacchus)

Common marmosets (Callithrix jacchus) demonstrate significant variation in reproductive output on both a yearly and lifetime basis in comparison to other anthropoid primates. We explore the factors that may be most important in determining reproductive variation in captive common marmosets. Studies have found that maternal age, maternal condition, and dam survivorship are related to reproductive output; however, these reports are not in agreement and are far from conclusive. With the use of a large, multicolony, demographic database pooling data across five marmoset colonies, we examined litter information for 1,649 litters, and reproductive summaries for 400 dams to assess 1) how reproductive output variation (total production, total weaned production) is determined by litter size, interbirth interval (IBI), age at first birth, and dam survival age; 2) the relationship between maternal age and reproductive output variables; and 3) relationship between the reproductive output variables and survival. We used stepwise regression procedures to describe the amount of variation in lifetime reproductive output among dams, and found that mean litter size accounted for 18% of the variance in total production, survival age accounted for 10.6%, age at first birth accounted for 8.8%, and mean IBI accounted for 5%. For total (nonzero) weaned production, survival age accounted for 7.6% of variance, age at first birth accounted for 7.2%, mean IBI accounted for 2%, and mean litter size accounted for 1.6%. We identified significant effects (P<0.05) of maternal age on litter size and IBI length, but no effect of dam age on weaned litter size. Cox proportional hazards regression analyses revealed significant effects (P<0.01) of number of litters, age at first birth, and site on dam survivorship. Dams that produced more litters showed higher survivorship. Age at first birth showed a positive relationship with dam survivorship, i.e., dams that delayed first reproduction had higher survival. Our findings about reproductive variation in marmosets may have practical applications for the management of marmoset breeding colonies.     

Comparative studies of senescence in natural populations of guppies

Investigators have rarely sought evidence for senescence in natural populations because it is assumed that relatively few individuals will survive long enough in the wild to exhibit the intrinsic increase in mortality with age expected from senescent individuals. Nevertheless, senescence has been documented in some natural populations, mostly in birds and mammals. Here we report on a comparative study of senescence in two natural populations of guppies (Poecilia reticulata). We document senescence as an age-specific increase in mortality rate, with use of mark-recapture studies and implementation of program MARK for analysis of such observations. Extrinsic mortality was controlled for by choosing populations that experience low rates of predation because they coexist with only a single piscine predator (Rivulus hartii). These populations differ in their evolutionary history because one was native to such a site whereas the other was introduced to a site that previously contained no guppies. The source of the introduced guppies was a high-predation population downstream below a barrier waterfall. Theory predicts that the guppies derived from a high-predation locality should experience senescence at an earlier age than the native low-predation population; however, the historical differences among these populations are also confounded with everything else that differs among the two localities. We found that females from a natural low-predation population have delayed senescence compared with the recently established population and hence that the differences among localities in senescence conform to theoretical predictions. The males from natural low-predation environments also had lower overall mortality rates, but contrary to predictions, the pattern of senescence for males did not differ between populations. The difference between the sexes is potentially attributable to two factors that lower the statistical power for distinguishing differences in the age-specific acceleration of mortality in males. One factor is that males have higher mortality rates, so fewer survive to advanced ages. A second is that we had a greater ability to discriminate among older age classes in females. We also found that the introduced population sustained a higher rate of disease than the native low-predation population. Such disease may represent a confounding factor in our comparison, but it may also reflect one of the trade-offs inherent in the life-history differences of these populations.     

Reproductive lifespan and reproductive performance in SPF C3H mice: the onset of reproductive life and production efficiency (author's transl)]

To improve the production system, the onset and the termination of reproductive life of C3Hf/HeMsNrs mice mated immediately after weaning and reared for 400 days of life, were studied. From weaning females mated with a full grown male (group A), the first litter was obtained at a mean age of 47 days, suggesting the first copulation at 26 days of age. The age of males at the first copulation was estimated to be at 44 days of age from the age giving the first litters in weanling males mated with weanling (group B) and full grown (group C) females. The sex ratio of litters delivered by young dams tended to be excess in males. The reproductive performance of dams in later life was not affected by the parturition in earlier age. The production efficiency with weaned youngs per pair during the first 200 days after mating was the highest in group A. It was found from these results that the C3H females attained their sexual maturity at 5 to 6 days after weaning, being available for breeding without any deletion in reproductive performance.     

Age structure and reproduction in feral New Zealand populations of the house mouse (Mus musculus), in relation to seedfall of southern beech

Abstract

After good seedfalls by southern beech (Nothofagus spp.), density indices for mice increased, the breeding season for adult females was brought forward, and winter breeding was observed. The end of the breeding season of the following summer was brought forward only slightly if the maximum density achieved was not great; incomplete data on 2 very high-density populations suggested a much earlier end to breeding. In early winter following a good seedfall the recruitment of juveniles was temporarily increased. After spring, few young mice entered the population, though breeding continued. The distribution of age classes shifted steadily upwards as population density rose. The peak population of 1976–77 comprised mainly old mice, still breeding, but without significant recruitment. No effects of density on sex ratio, litter size, or body weight could be detected from these data. In all populations studied, density decreases were ultimately due to failure of recruitment and the disappearance of ageing, non-breeding mice. The proximate causes of this mortality were probably the onset of cold weather in 1976–77 and predation by stoats in 1979–80.     

Characterization of age and cuticular hydrocarbon variation in mating pairs of house fly,Musca domestica,collected in the field

House flies, Musca domestica L., were collected in copula over two summers from six dairies located in three climatically distinct regions in the U.S.A. southern California, Minnesota and Georgia. Ages of males and females from a total of 511 mating pairs were estimated using pterin analysis. Cuticular hydrocarbon profiles and gonotrophic ages of females also were evaluated. Mean age of mating males ranged from 54 to 102 degree‐days (DD) (4–10 days based on field air temperatures), depending on the farm. Very young males (< 10–20 DD) and old males (> 200 DD) were rare in mating pairs. Mean female age at mating ranged from 20 to 46 DD (2.5–4 days). All mating females had eggs in the early stages of vitellogenesis and 99.2% were nulliparous. However, some older and parous females were collected, demonstrating that re‐mating can occur in the field. Head width measurements of mating pairs suggested that assortative mating by size did not occur. The cuticular hydrocarbon profiles of females were determined, with emphasis on (Z)‐9‐tricosene (muscalure). Overall, only 55% of mating females had detectable amounts (> 4 ηg per fly) of (Z)‐9‐tricosene. Of the females that had detectable (Z)‐9‐tricosene, variation in amount per female was high in all fly populations, and thus was not statistically related to the size or age of the mating female. The proportion of mating females with detectable levels of (Z)‐9‐tricosene varied by geographic region. Seventy‐one, 63, and 27% of females from southern California, Minnesota and Georgia had detectable amounts of (Z)‐9‐tricosene. Principal components analysis of the eight most abundant hydrocarbons from mating females, by state, revealed state‐level distinctiveness of hydrocarbons in house fly populations, which may reflect genetic variation associated with environmental stresses in those geographical zones.     

Population size, female fecundity, and sex ratio variation in gynodioecious Plantago maritima

Theory predicts that the sex ratio of gynodioecious populations (in which hermaphrodites and females coexist) will be affected by the relative female fitness of females and hermaphrodites, and by founder events and genetic drift in small populations. We documented the sex ratio and size of 104 populations of the gynodioecious, perennial herb Plantago maritima in four archipelagos in eastern Sweden and western Finland (from latitude 53 to 64 degrees N). The sex ratio varied significantly both among and within archipelagos (range 0-70% females, median 6.3% females). The frequency of females was highest in the northernmost archipelago and lowest in the southernmost archipelago. As predicted, females were more frequently missing from small than from large populations, and the variance in sex ratio increased with decreasing population size. The relative fecundity of female plants (mean seed output per female/mean seed output per hermaphrodite) ranged from 0.43 to 2.16 (median 1.01, n = 12 populations). Among the 12 populations sampled for seed production (four in each of three archipelagos), the frequency of females was positively related to relative fecundity of females and negatively related to population size. The results suggest that the local sex ratio is influenced both by the relative fecundity of females and hermaphrodites and by stochastic processes in small populations.