A positive Link Between Male Testosterone and Spacing Behavior in Pair‐Bonded California Mice

The effects of testosterone (T) on male sexual behavior are well established, but its roles in several associated social behaviors such as pair‐bonding and paternal behavior are diverse. Recently, we reported that male T in response to pairing with a female predicts future paternal behavior in the monogamous and biparental California mouse (Peromyscus californicus). Here, we examined whether T in response to pairing is also associated with spacing behavior between mates under laboratory conditions and if these behavioral patterns are continued during pup care. Pairs of California mice were observed throughout their pair bond before and again after pups were born. We found that males with higher T post‐pairing remained closer to their mates in the weeks before pups were born, and pairs with higher proximity measures during the pair bond spent more time caring for pups together. Overall, these results suggest a similar mechanism for spacing behavior across distinct phases of the pair bond.     

Inter-sexual differences in Antarctic fur seal pup growth rates: evidence of environmental regulation?

We investigated the variation of Antarctic fur seal (Arctocephalus gazella) pup growth rates in response to sex, breeding season and duration of both maternal foraging trip and attendance bouts. Data were collected during five consecutive rearing seasons at Cape Shirreff, the most important breeding colony in the South Shetland Islands, Antarctica. Our results showed significant interannual and sexual variations in pup growth rates. Male pups grew significantly faster than female pups during 2000, 2001, 2002 and 2004 seasons, whilst during 2003 no difference was found. The interannual variation in pup growth rates was correlated with the interannual fluctuation in maternal foraging trip and attendance bouts. There was a significant effect of pup sex and maternal foraging trip duration on pup growth rates, which varied between years having foraging trip duration a major effect during 2003, when females spent more time at sea and interestingly on that year there were no sexual differences in pup growth rates. The effect of attendance bout on pup growth rates was not significant. Diet analysis showed that Antarctic krill (Euphausia superba) was the most frequent prey item during the study period. Analysis of krill size distribution showed a significant difference in krill length, during 2003, when A. gazella preyed upon the smallest sizes of krill. In this study, sex was the most important factor on pup growth rates, but when prey availability seemed more limited, there are longer foraging trips and shorter attendance bouts the sex factor became less significant.     

Pup development and maternal behavior in captive Key Largo woodrats (Neotoma floridana smalli)

This study describes the first systematic observations of maternal behavior and pup development of captive Key Largo woodrats (Neotoma floridana smalli) during the first 30 days of life. Data were collected on six litters of pups born to four dams between December 2006 and July 2007. Gestations for the six litters averaged 38 days and all dams exhibited adequate maternal care postpartum. Key Largo woodrat maternal and pup behavior was generally consistent with behavior observed in other woodrat species. We observed greater pup independence from the dam and a marked change in social interactions between days 13–22. No sex differences in pup development or maternal care were observed. Activity budgets were consistent across dams and across days within the observation period. Although dams spent much of their time inactive with pups attached to their teats, the average percent of intervals with at least one pup observed attached decreased steadily during the 30‐day observation period. Attachment of pups to the dams' teats did not interfere with dams' ability to forage. Feeding with pups attached and feeding following active detachment of pups were both common. Dams were observed to actively detach pups by performing a circular turning motion. This information has application for the future management of this endangered species in captivity and in the wild. Zoo Biol 27:394–405, 2008. © 2008 Wiley‐Liss, Inc.     

BEHAVIORAL AND PHYSIOLOGICAL MEASUREMENTS OF MATERNAL INVESTMENT IN THE STELLER SEA LION, EUMETOPIAS JUBATUS

The onshore and at-sea cycles of females, suckling behavior of pups and their milk intake were studied in Steller sea lions ( Eumetopias jubatus ) during 1983 at Año Nuevo Island, California. Females averaged approximately 21 h ashore and 36 h at sea. The trips to sea lengthened as pups aged, resulting in an overall decline in female time ashore to 30% by the sixth week following parturition. Activity budgets of pups showed no significant differences among suckling time, age and sex. Milk intake, estimated using labeled water studies, revealed that heavier pups consumed more milk than lighter ones (milk ingestion in ml/d = 4.26 + 0.0687 [Pup Mass in kg]). Mean milk intake was 1.78 ± 0.33 liters/d. Mean pup growth rate was 0.38 ± 0.1 kg/d. The results suggest that female attendance patterns are shaped by the increasing nutritional demands of growing pups and their increasing efficiency at suckling.     

Individual,Group, and Environmental Influences on Helping Behavior in a Social Carnivore

Variation in group composition and environment can affect helping behavior in cooperative breeders. Understanding of how group size, traits of individuals within groups, food abundance, and predation risk simultaneously influence helping behavior is limited. We evaluated pup‐guarding behavior in gray wolves (Canis lupus) to assess how differences in individuals, groups, and environment affect helping behavior. We used data from 92 GPS‐collared wolves in North America (2001–2012) to estimate individual pup‐guarding rates. Individuals in groups with low helper‐to‐pup ratios spent more time guarding young than those in groups with more helpers, an indication of load‐lightening. Female helpers guarded more than male helpers, but this relationship weakened as pups grew. Subset analyses including data on helper age and wolf and prey density showed such factors did not significantly influence pup‐guarding rates. We show that characteristics of individuals and groups have strong influences on pup‐guarding behavior in gray wolves, but environmental factors such as food abundance and predation risk from conspecifics were not influential.     

Ontogeny of Time Allocation in Black-tailed Prairie Dogs

The ontogeny of time allocation was studied in a field population of black-tailed prairie dogs, Cynomys ludovicianus. Pups first emerged from their natal burrows in late May. All individuals in the population, including all pups, were observed for 12 weeks following this emergence. Both cross-sectional and longitudinal analyses of ontogeny revealed considerable age differences. Upon first emergence, pups were extremely wary and spent most of their time vigilant and little time feeding. As pups aged, they increased time spent feeding and decreased time spent vigilant. Male and female pups behaved similarly. Pups differed from adults in their time allocation patterns and these differences changed as pups aged. Pups were initially more vigilant and fed less than adults, but became less vigilant than adults as the summer progressed. Yearlings behaved similarly to adults. Possible intrinsic (e.g., pup age and weight) and extrinsic (e.g., weather conditions, microhabitat location and coterie composition) influences on time allocation by pups were also examined. In general, extrinsic influences appeared to have more impact on pup behavior than intrinsic ones, suggesting that time allocation by pups may be largely context-dependent. Overall, the considerable number of age differences argues for more consideration of ontogeny in models of antipredator behavior.     

Factors affecting weaning weight in Antarctic fur seals Arctocephalus gazella at South Georgia

Summary Relationships between size, body condition, age and feeding-attendance patterns during pup rearing of female Antacrtic fur seals Arctocephalus gazella and their effects on the timing of birth and weaning, pup weight, growth and condition were studied at South Georgia in 1981–1982. Twenty-seven (6 male, 21 female) mother-pup pairs were followed from birth to weaning. The analysis of maternal effects was limited to female pups because of the small sample size of male weaners. High weaning weight was associated with those female pups whose mothers spent more time ashore attending their offspring. Weaning weight showed no relationship with perinatal duration, number of feeding trips to sea, days at sea or date of weaning. A further 63 mother-pup pairs were analysed for the effects of maternal body condition (weight/length), age and timing of birth on offspring body weight and condition. Pup weight and condition were weakly correlated with maternal age in female pups. Male pups born earlier in the season were heavier and in better condition. Maternal and offspring body weight and condition were unrelated. For the Antarctic fur seal population at South Georgia where the food supply was apparently not limiting in summer, maternal condition and foraging time were subordinate to maternal care on land (as expressed by attendance duration) in determining offspring weight at weaning.     

Aromatase inhibition during adolescence reduces adult sexual and paternal behavior in the biparental dwarf hamster Phodopus campbelli

Previous studies have failed to identify an activational role for estradiol in the paternal behavior of Phodopus campbelli fathers. However, none of these studies addressed a developmental role that estradiol might play in establishing paternal behavior in this species. Males were orally administered the aromatase inhibitor letrozole (1 mg/kg/day) for three days at 18, 34, or 90 days of age. As adults, males were tested for paternal and sexual behavior. Letrozole treatment at 18 days resulted in males that spent less time huddling over pups during the birth, and had higher pup losses and male-biased pup survival for the first litter. Letrozole treatment at 34 days resulted in males that had altered sexual behavior; males had a longer interval between mounts and between intromissions, and took longer to achieve ejaculations over the first three ejaculatory series. Furthermore, these males sired smaller first litters and produced second litters with a male-biased sex ratio. Males treated with letrozole as adults showed a modest increase in paternal care during the birth, but pup development and survival were not altered. There was no effect of treatment on attack or retrieval behavior either as sexually naive adults or as new fathers. Thus, the results of the present study suggest that estradiol acts during adolescence to establish the normal expression of midwifery behavior and sexual behavior during adulthood.     

Daily and seasonal variation in the haul-out behavior of the South American sea lion

In the South American sea lion (Otaria flavescens), daily fluctuations in abundance have been analyzed based on sequential counts of the number of animals hauled-out. However, no studies have analyzed haul-out activity in relation to an annual cycle or according to different age/sex classes. The objective of this study was to determine the daily and seasonal haul-out patterns of each age/sex class of South American sea lions as a function of the time of the day. A total of 222 days were analyzed in a breeding colony of Chile, from May 2008 to December 2010. During the non-breeding season (March to December) males, females, and juveniles showed a unimodal pattern, in which few sea lions are hauled-out in the morning and maximum numbers are found in the rookery during the early afternoon (1330–1630). In contrast, during the breeding season (austral summer) the proportion of individuals hauled-out shifted from a unimodal to a bimodal pattern, especially in the case of juveniles. Our results indicate that there are fine scale differences in haul-out behavior among age/sex classes, as well as larger scale seasonal differences in the proportion of sea lions ashore. These differences appear to be related to reproductive activities, food availability and thermoregulatory requirements. These patterns of seasonal variability of South American sea lion haul-out should be taken into consideration when planning surveys to estimate population abundance.     

ATTENDANCE PATTERNS OF CALIFORNIA SEA LION (ZALOPHUS CALIFORNIANUS) FEMALES AND PUPS DURING THE NON-BREEDING SEASON AT SAN MIGUEL ISLAND

The attendance patterns of California sea lions were studied during the non-breeding seasons from 1991 to 1994. Lactating females frequented the rookery to nurse their pups until weaning; most non-lactating females left the rookery for the season. Females spent over 70% of their time at sea except in 1993 when they spent 59% of their time at sea. The mean foraging trip length in the winter and spring ranged from 3.3 to 4.6 d; the mean nursing visit ranged from 1.2 to 1.4 d. The duration of foraging trips and nursing visits was variable over the season for individuals but no pattern of change was detected. Interannual and seasonal differences were not significant for time at sea, visits ashore, or foraging-trip duration before, during, or after the 1992-1993 El Niño event. Pups spent an average of 66.6% of their time ashore and up to three days away from the rookery during their mother's absence. Most females and pups stayed associated until April or May. The results suggest that seasonal movement of prey is more important in determining attendance patterns late in the lactation period than increasing energy demands of the pup.     

Mechanisms of Change in the Ontogeny of Black-tailed Prairie Dog Time Budgets

This study examines the behavioral mechanisms underlying changes in the time allocation patterns of black-tailed prairie dog (Sciuridae: Cynomys ludovicianus) pups. Previous analyses of time budgets had revealed substantial differences between pups (young of the year) and adults, and that pup time budgets changed significantly over time. This paper shows that these differences were due to two mechanisms: pups generally differed from adults in the frequency of performance of different behaviors; however, changes in pup behavior over time were due to changes in both bout frequencies and durations. Other influences on bout durations were also examined. Time of day dramatically affected the duration of most behaviors, with vigilance bouts longest in the morning and feeding bouts longest in the evening. The presence of a pup's mother above ground and the height of the vegetation a pup was in had much less influence on bout durations. Multiple regression analyses were used to identify other potential influences (e.g., weather and social conditions) on bout lengths. Of these, the distance of a pup from the nearest burrow produced the strongest relationships.     

Postbranding Survival of Steller Sea Lion Pups at Lowrie Island in Southeast Alaska

ABSTRACT Steller sea lion (Eumetopias jubatus) pups (n = 366) were hot-branded at Lowrie Island, Southeast Alaska, USA, in June 2001 and 2002 for vital-rates studies. To assess potential mortality following branding, we estimated weekly survival to 12 weeks postbranding using mark-recapture models. Survival estimates ranged from 0.984/week to 0.988/week, or 0.868 over the 12-week period; varied little with sex, year, and capture area; and were higher for larger than smaller male pups and unexpectedly lower for larger than smaller female pups. Inclusion of resights at 1–3 years of age prevented a −4.5% bias in cumulative survival to 12 weeks postbranding by accounting for pups that survived but permanently emigrated from Lowrie Island during the 12-week survey. Data from double-marked pups (i.e., branded and flipper-tagged) indicated the low brand-misreading probability of 3.1% did not bias survival estimates. Assuming survival differences between the first 2 weeks postbranding and later weeks were due entirely to the branding event, potential postbranding mortality of branded pups attributable to the branding event was 0.5–0.7%, or one pup for every 200 marked. Weekly survival of branded pups was nearly identical to estimates from a control group of undisturbed, unbranded pups born to 10–11-year-old branded adult females in 2005 (0.987–0.988/week) and similar to pup survival estimates from other otariid studies. Available data did not indicate substantial mortality to 12 weeks postbranding resulting from the branding disturbance, suggesting branding of Steller sea lion pups can be used effectively for investigations of population declines without significantly affecting population health or study goals.     

SEXUAL DIMORPHISM IN NEWBORN SOUTHERN SEA LIONS

Male southern sea lion, Otaria byronia , pups born at Peninsula Valdés, Argentina, were significantly heavier (13.7 ± 1.6 kg, n = 37 vs 12.3 ± 1.4 kg, n = 38) and longer (0.82 ± 0.04 m, n = 29 vs . 0.79 ± 0.03 m, n = 29) than females. The rate of mass and length gain during the first month of life did not differ between the sexes (males: 0.21 kg/d, 0.003 m/d; females: 0.20 kg/d, 0.004 m/d). Sex differences in pup body size were not reflected in female nursing behavior, attendance patterns, and duration of feeding trips. These results suggest that southern sea lion mothers invest more energy in sons than in daughters during gestation, a difference consistent with the importance of large body size to the reproductive success of males in a polygynous species.     

DISTRIBUTION AND POPULATION SIZE OF THE SOUTHERN SEA LION (OTARIA FLAVESCENS) IN CENTRAL AND SOUTHERN CHUBUT,PATAGONIA, ARGENTINA

Population size and distribution of Otaria flavescens in central and southern Chubut, Patagonia were determined by aerial censuses conducted during the breeding season in 1989 and 1995 and outside the season. Pup numbers were corrected by means of a linear regression between pup and adult male (AM) numbers obtained from terrestrial censuses carried out in 1994 and 1996. During the 1989 breeding season, 10,557 sea lions occurred at 29 locations; 2,800 (26.3%) were pups. During the 1995 breeding season, 14,887 animals were censused at 32 sites; 3,311 (22.2%) were pups. The corrected pup number increased to 4,852, indicating that around 32% is lost when counting from photographs. The corrected total number increased to 16,483; this fell to less than one half between breeding seasons (November 1990 and 1995). The 1995 breeding season census represents the highest number since 1972 (8,800) and the widest historical dispersion (37 sites year round contrasting with 13 in the late 1940s and 1972). However, the present number is about one half of the 33,000 censused about 50 yr ago. Using a correction factor developed for the north Paragonian population (1.8 × censused number) the present population size in the area can be estimated at around 29,669 individuals.     

Growth and survival of New Zealand sea lions, <Emphasis Type="Italic">Phocarctos hookeri:</Emphasis> birth to 3 months

Offspring birth mass and growth rate represent important life history traits, which influence many vital population and individual characteristics, while offspring survival is a key factor in variation in female reproductive success. For a threatened population of pinnipeds, such as New Zealand sea lions, Phocarctos hookeri, (Grey, 1844, NZ sea lions), understanding individual life history parameters and population dynamics is vital for their management and conservation. This is the first study of the behaviour of females during parturition, pup birth mass and growth, and pre-weaning survival of NZ sea lions, Enderby Island, Auckland Islands during austral summer breeding seasons, 2001/2002 to 2003/2004. Pregnant females arrived ashore 2.1 ± 0.16 days prior to giving birth. After parturition, mothers suckled their pups for 8.6 ± 0.16 days before leaving on their first foraging trip. Male pups were born significantly heavier than female (males 10.6 ± 1.4 kg, females 9.7 ± 0.9 kg). Pups lost on average 48 ± 0.14 g per day mass during the early postpartum period (between birth and mothers first foraging trip). Pup mortality did not vary by pup sex, birth mass, date of birth or any maternal characteristics however it varied significantly between years due to a bacterial infection epidemic (Pup mortality at 60 days: 2001 32%; 2002 21%; 2003 12%). The absolute growth rate per day for pups was 151 g/day over all years. Pup growth rate measured as the slope of linear line fitted to pup mass by age was consistently higher for pups with heavier birth mass, male pups and during the 2002 season. High offspring mortality and slow growth rates coupled with maternal foraging behaviour at their physiological limits may reflect a threatened species which has limited ability for population growth in an environment which is at the extreme of their historical range and impacted upon by fisheries.     

Behavioral budgets of captive sea otter mother–pup pairs during pup development

Instantaneous sampling of eight behavior categories for a captive mother sea otter (Enhydra lutris) and her four pups was conducted continuously during the first three months following each pup's birth. Initially, pup-grooming and rest occupied the largest proportion of the mother's time. Despite a steady decline in the amount of time the mother spent pup-grooming, the mother continued to groom the pup a substantial amount of time three months after the pup's birth. The proportion of time the mother spent resting, self-grooming, or swimming did not change significantly over the study period, suggesting that these behaviors are important to the mother's self-maintenance. The newborn pups' behaviors were initially limited to rest, active, and nursing, with large proportions of time spent resting and nursing. A significant increase in self-grooming occurred regularly by week 2, followed by increases in swimming (week 3), diving (week 6), and feeding (week 7). Three months after birth, the pups' behavior budgets were similar to their mother, with the exception that pups self-groomed substantially less than their mother self-groomed. The lower proportion of pup self-grooming may be due to the continued close proximity to their mother in captivity, which afforded extended association, and allowed the mother to continue to perform this complex behavior for the pup. © 1993 Wiley-Liss, Inc.     

Aromatase inhibition during adolescence reduces adult sexual and paternal behavior in the biparental dwarf hamster Phodopus campbelli

Previous studies have failed to identify an activational role for estradiol in the paternal behavior of Phodopus campbelli fathers. However, none of these studies addressed a developmental role that estradiol might play in establishing paternal behavior in this species. Males were orally administered the aromatase inhibitor letrozole (1 mg/kg/day) for three days at 18, 34, or 90 days of age. As adults, males were tested for paternal and sexual behavior. Letrozole treatment at 18 days resulted in males that spent less time huddling over pups during the birth, and had higher pup losses and male-biased pup survival for the first litter. Letrozole treatment at 34 days resulted in males that had altered sexual behavior; males had a longer interval between mounts and between intromissions, and took longer to achieve ejaculations over the first three ejaculatory series. Furthermore, these males sired smaller first litters and produced second litters with a male-biased sex ratio. Males treated with letrozole as adults showed a modest increase in paternal care during the birth, but pup development and survival were not altered. There was no effect of treatment on attack or retrieval behavior either as sexually naive adults or as new fathers. Thus, the results of the present study suggest that estradiol acts during adolescence to establish the normal expression of midwifery behavior and sexual behavior during adulthood.     

Endogenous 24-hour cycle of core temperature and oxygen consumption in week-old Zucker rat pups

Summary Experiments were designed to test whether or not the 24-h core temperature fluctuations in week-old rat pups are of endogenous origin. Lean (Fa/-) Zucker rat pups born on the same day to mothers maintained in two different colonies with light/dark cycles 12 h out of phase with each other were mother-reared through the first 3–4 days of life and then artificially reared simultaneously in constant dim light. Continuous, automatic measurement of core temperature and oxygen consumption during artificial rearing showed clear 24-h rhythms in 5- to 8-day-old pups. Each rhythm reached a daily minimum at a time corresponding to the beginning of the light period in the colony of origin. The amplitude of these rhythms did not diminish during artificial rearing, nor did the phase difference between the rhythms of pups originating in the two colonies systematically change. The persistent 12-h phase differences between these two groups of pups prove that the observed rhythms are not caused by exogenous stimuli. We conclude that the rat pup possesses an endogenous time-keeping mechanism that permits the expression of overt rhythmicity at the age of 1 week.Abbreviations Tc core temperature - LD-pups born to a mother entrained to a 7:00 to 19:00 light cycle, then artificially reared in constant dim light - DL-pups born to a mother entrained to a 19:00 to 7:00 light cycle, then artificially reared in constant dim light - SCN suprachiasmatic nucleus     

Hormonal Responses of Male Gerbils to Stimuli from Their Mate and Pups

Following copulation and cohabitation with a pregnant female, male gerbils show high levels of parental behavior toward their pups. The initiation of male parental behavior may be the result of neuroendocrine changes induced by cohabiting with the pregnant female or by pup stimuli. Experiment 1 examines the changes in androgen and prolactin levels in male gerbils cohabiting with females over the reproductive cycle. Gerbils were mated and blood samples taken from males for hormone analysis 1, 10, and 20 days after pairing and 3, 10, and 20 days after pups were born. A group of unmated male gerbils served as controls. Plasma prolactin levels of males were elevated throughout the female's pregnancy and lactation periods, but were only statistically significantly higher than those of unmated males 20 days after pups were born. Androgen levels rose during pregnancy and dropped significantly after the birth of the pups. These hormonal changes are similar to those found in males of monogamous birds and differ from those found in males of polygynous rodents such as the rat. Experiment 2 examined the hormonal responses of male and female gerbils to pup replacement after 4 hr of parent–pup separation. Female gerbils showed a significant elevation of prolactin levels 1 hr after pup replacement, but males did not. Males with pups returned showed no difference in androgen levels from males who did not have pups returned. Thus, male gerbils show neuroendocrine changes following long-term cohabitation with their mate and pups, but do not show acute hormone responses to pup removal and replacement. These results indicate that parental males have neuroendocrine changes associated with parental behavior and these differ from the neuroendocrine changes underlying female parental behavior.     

Reproductive patterns in the domestic dog--a retrospective study of the Drever breed

The aim of this study was to examine the differences, between seasons of the year, in the distribution of matings and whelpings, litter size, pup deaths, and sex ratio in domestic dogs. Furthermore, we wanted to examine the effects of age and parity of the bitch at the time of whelping on litter size, as well as the effect of litter size on gestational length. A final aim was to investigate the fertility and frequency of whelping problems in a private kennel of Drever dogs. Data from the Swedish Kennel Club (SKK) registry for the Drever breed during 1995-2006, comprising a total of 2717 litters, were analyzed together with more detailed data from a private, professional kennel of Drevers, with a total of 285 matings and 224 whelpings, during the same time period. The most matings took place during winter, and the fewest during summer; consequently, most whelpings occurred during the winter and spring seasons. Of the 285 mated bitches, 78.6% whelped, 6.25% experienced dystocia, and 5.36% underwent Cesarean section. The pup death rate was 7.6%. The largest litters were born during spring. Litter size was negatively correlated with duration of pregnancy (r=-0.18). Each pup more than average caused a shortening of the gestation by 0.25 days, and each pup less a corresponding lengthening. Bitches giving birth to their first litter after 4 years of age produced a smaller litter than younger bitches. Litter size decreased after 5 years in all bitches. The number of born pups at the private kennel increased from the first to the third parity, then decreased. The number of registered pups increased from the first to the second parity in the SKK data and from the second to the third parity in the data from the private kennel, then decreased. Mating a bitch only once resulted in a smaller litter size. None of the studied factors had any effect on the sex ratio of the pups. There were significant differences between males in whelping rate among the mated bitches, but no difference in mean litter size, which indicates a female problem rather than a male one. Available data suggest that the domestic dog is still under considerable seasonal influence, although modified by ambient and management factors.