Parental care and infant development in a family group of captive Sichuan golden monkeys (Rhinopithecus roxellanae): First 20 days

A male Sichuan golden monkey (Rhinopithecus roxellanae) was born on July 24, 1986 to an adult pair on loan from the Chinese government. Physical and behavioral development of the infant, as well as the behavior of the parents, were recorded for the first 20 days of the infant's life. (On Day 21 the trio was returned to the Chongqing Zoo in the People's Republic of China.) The infant pelage was dark gray at birth but soon began changing to the orange-gold color typical of the adults. Forty hours of behavioral data were collected using a combination of scan sampling and all occurrences of selected behaviors techniques. The infant pushed against his mother and pulled toward objects in the enclosure beginning at 4 days of age. Time spent in these behaviors increased steadily and on Day 15 the infant broke contact with his mother for the first time. Mother-infant contact appeared to be maintained by both individuals mutually up to the infant's 10th day of the life. The percentage of time that the infant spent actively embracing his mother began to decline after that while the female continued to spend the majority of her time embracing the infant. During the first few days following the birth, the male always initiated and maintained proximity to the female and the infant. This “vigilant” behavior had declined somewhat by the end of the study.     

Observations of a daytime birthing event in wild titi monkeys (Callicebus oenanthe): implications of the male parental role

Behaviors displayed during birth events, including prenatal, parturition, and postpartum periods, can give insight into caretaking roles in species in which the offspring is cared for by individuals other than the mother. Titi monkeys, genus Callicebus, exhibit a pair-bonded social system along with intense male care of offspring. Here, I report the first case of a birth seen in the wild in a group of Callicebus oenanthe, a little-known species in northern Peru, and describe infant care and development during the first few months after birth. Detailed behavior during the birth sequence as well as ad libitum data postpartum were recorded on nursing and infant care behaviors, including infant carrying, infant grooming, anogenital cleaning, protection, playing, and food sharing. In the 3 h preceding and during parturition, the male remained in contact with or in close proximity to the female (<1 m). The male licked and examined the newborn 3 min after parturition, carried the infant within 24 h after birth, was the main carrier of the infant, and was the predominant manual groomer of the infant during the first 4 months after birth. I argue that the male plays an integral role during the birth in order to establish his bond with the infant as well as reinforce infant care duties to the female and in predator vigilance. In light of various explanations for the exhibition of intense paternal care, I suggest that the male titi monkey provides infant care to release the female of these duties in order that she may spend more time foraging, thus potentially increasing the pair’s overall reproductive output.     

A method for encouraging maternal care in the giant panda

We developed a method to facilitate maternal care in a giant panda (Ailuropoda melanoleuca) female that rejected her cub immediately after parturition. After removal of the cub, the female was systematically exposed to a regime of infant‐related stimuli, including a surrogate toy panda, accompanied by infant vocalizations and urine, and the mother's own milk. The female displayed several measures of maternal behavior toward the surrogate, for example, spending 61% of her time holding the surrogate in positions typical for the species. There was some evidence that maternal proficiency increased across the 4 weeks of the experiment. Results also indicate that the female was responsive to both infant vocalizations and urine, but not milk. After a transitional period in which we assisted the female in her efforts to nurse and groom the infant, all maternal care‐giving responsibilities were returned to the mother. Over the next 3 months, the pattern of maternal care followed the species‐typical course of declining mother‐infant contact, grooming, overall interaction time, and responsiveness to the cub's vocalizations. This study marks the first successful reunification of a giant panda mother with an infant separated at birth and, it is hoped, will serve as a model for similar efforts elsewhere. Zoo Biol 19:53–63, 2000. © 2000 Wiley‐Liss, Inc.     

Adoption in free-ranging red howler monkeys,Alouatta seniculus of Venezuela

This report describes three cases of infant adoption in red howler monkeys (Alouatta seniculus) of Venezuela. In one case, a lactating female adopted her infant granddaughter after the infant's mother disappeared and the infant survived for the next 25 months. In the second case, an adult female adopted her infant granddaughter for nine days while the infant's mother was still a group member. In the third case, a nulliparous female temporarily adopted an infant from a neighboring group. The details and the possible reasons for these adoption are described.     

Within-group cannibalism by adult male chimpanzees

In the Mahale Mountains National Park, Tanzania, a young adult male chimpanzee was observed to feed on a 3-month-old male infant of the same unit-group. Four other adult males and an adult female shared the carcass. The mother of the victim had immigrated from a neighboring unit-group four years previously. Circumstantial evidence strongly suggests that the first-observed cannibal male also killed the infant. The adult male and the mother of the victim had been familiar socially and sexually with each other since the female immigrated. Since the mother of the victim had usually been ranging in the peripheral part of the unit-group's range, i.e., the overlapping area of the two unit-group's ranges during pregnancy and soon after birth, the infanticidal male might have had reason to suspect the paternity of her infant. Four such cases of within-group cannibalism by adult males suggest that the female range and association pattern before and after parturition are key factors allowing an infant to survive. The possibility of male-biased infanticide is also discussed.     

The birth of a wild mountain gorilla (Gorilla gorilla beringei)

The live birth of a wild mountain gorilla is described. Details of the birth process are given along with the mother’s periparturitional behaviour and the responses of others to the birth. The mother, a multiparous female, gave no observable indication that birth was imminent, and delivered her infant quickly and easily. During parturition, a subadult male showed interest in the proceedings. Following delivery, several animals approached the mother, who then left the birth site. The mother cleaned her infant and by two hours postpartum had established ventro-ventral contact and, possibly, nursing. These data are related to relevant observations from the literature and unpublished data on other animals from this study.     

Control of postpartum mating behavior in free-ranging rhesus monkeys

This paper explores the effect that a dependent rhesus macaque infant (Macaca mulatta) has on the timing of its mother's resumption of mating. Toward this end, behavioral data from the rhesus populations at Tughlaqabad, India, and Cayo Santiago, Puerto Rico, are compared. Specifically, the intervals between parturition and subsequent resumption of mating for individual parous females are examined in order to shed light on the relative importance of environmental and internal factors (such as lactational infertility) in the control of rhesus mating behvior. At both study sites it was found that the later a given female gave birth during a birth season the shorter was her parturition-mating interval and the younger was her infant when she resumed mating activity. It is suggested that this finding is the result of an interaction of environmental and internal factors; both act in concert to determine how soon after parturition a particular rhesus female will resume mating activity.     

Tactics of care for young infants by forest-living ruffed lemurs (Varecia variegata variegata): Ground nests,parking, and biparental guarding

Over a 4 month period, systematic and ad libitum observations were conducted on two adult female black-and-white ruffed lemurs (Varecia variegata variegata) and their infants in a 3.5 ha forest enclosure. The females were mother and daughter, members of a family group that had been semifree-ranging for 2 years and 3 months at the time of the births. One to two weeks before parturition, the females independently constructed nests, in which they kept their infants during the first few weeks following parturition. The older mother, cage-reared herself, prepared at least one nest. Her daughter, who has lived in the forest since late juvenescence, prepared at least four. Two to three weeks after parturition, the mothers moved their infants high into trees. During periods of maternal absence, the infants were often alone, and they rarely or never moved, vocalized, or groomed themselves. The mothers often kept their infants together after nests were no longer used. Each infant nursed freely from both lactating females. The infants were carried orally only by their mothers and were never transported by clinging to the pelage of any group members. Previously, researchers suggested that ruffed lemurs build nests for care of infants high up in trees. The present observations, however, suggest that two major modes of neonate care in Varecia exist: serial use of multiple ground nests and “parking” of infants high in trees. Advance preparation of several nest sites, relative lack of large predators, alternate maternal and paternal guarding of infants, infant immobility during absence of mother, and rapid infant development make this tactic of care for neonates plausible.     

Pregnancy,gestation and parturition in free-ranging tana river crested mangabeys (Cercocebus galeritus galeritus)

Mean gestation for seven free-ranging Tana River crested mangabeys (Cercocebus galeritus galeritus) is 180 days (SE = 4.49). All females showed postconception sexual swellings after the first 2 months of pregnancy, and two of the seven copulated with males at this time. One birth was observed; observations of a second infant began less than 1 hr after birth. Details of parturition are given, with responses of group members to the events. Adult and juvenile females showed more sustained interest in the new infants than adult or juvenile males.     

An attempted within-group infanticide in wild chimpanzees

Two high-ranking adult male chimpanzees (Pan troglodytes) of M group in the Mahale Mountains National Park, Tanzania, tried to get a newborn female infant of an adult female who was born in M group. The mother was not seen to mate with these adult males during the probable conception cycle of the infant, but she disappeared from the group during the later cycle, when she may have been inseminated by a male of one of the neighboring unit groups. The adult males failed to get the infant because the grandmother of the newborn and her female friend cooperated to protect the mother and infant from attacks by the males. The sexual selection hypothesis for infanticide by adult male chimpanzees holds for this observed case. The sudden disappearance of another infant, a healthy female, strongly suggests the killing of female infants too. Therefore, the asserted male-biased infanticide in chimpanzees appears to be less tenable.     

First occurrence of twins in provisioned free-ranging Tibetan macaques (Macaca thibetana) at Huangshan, China

A first case of newborn Tibetan macaque (Macaca thibetana) twins was observed in a free-ranging group at Huangshan, China. The female that gave birth to the twins was studied during their first 5 months post-partum, and her activity budget was compared to those of adult females with single or no offspring in order to assess her behavioral changes. Our report shows that female Tibetan macaques can produce twins, and that twins can successfully survive. The adult female with twins spent more time foraging and resting, but less time moving and engaged in other social behaviors than adult females with a single infant or no infant. Our report provides a case of successfully surviving twins in a wild environment and suggests that the mother modified her behavior patterns to adapt to the heavy burden. We conclude that both food provisioning and the mother’s behavioral strategies facilitated the survivorship of twins. This expands our understanding of the reproductive biology of Tibetan macaques.     

Observations of parturition among captive patas monkeys (Erythrocebus patas)

Six cases of labor and delivery by laboratory-housed patas monkeys (Erythrocebus patas) are described. Five of the births occurred during the daytime, and the sixth occurred just after the usual “lights out” time. Although it was impossible to determine the actual onset of labor, the final 1.5–0.5 hr before parturition were observed in all cases. Throughout labor, females tended to begin a contraction every 2–4 min and contractions usually lasted 35–60 sec. Females did not show common patterns of change in contraction frequency or duration as birth neared. It was not possible to describe patas monkey labor and delivery in terms of “stages.” Postpartum, females typically began cleaning their infants within 90 sec and picked them up within 5 min. Placental delivery times varied among females, as did the extent of placental consumption. One “mother-only” - reared female proved to be an inadequate mother.     

Allomaternal care: helper-assisted birth in the Rodrigues fruit bat, Pteropus rodricensis (Chiroptera: Pteropodidae)

We report observations of allomaternal care, or epimeletic (care-giving) behaviour, exhibited by one female toward another, before, during, and after giving birth, in the Rodrigues fruit bat, Pteropus rodricensis (Chiroptera: Pteropodidae). Interactions between the expectant mother and other roost mates, from the onset to completion of birth, were observed continuously over a period of approximately three hours. The attending female assisted the mother when she: (I) intermittently groomed her anovaginal region; (2) grasped her with partially outstretched wings; (3) 'tutored' her in a feet-down birthing posture; (4) groomed the emerging pup; and (5) physically assisted the mother by manoeuvring the pup into a suckling position. This is the first report of extensive assistance given by a female bat to another during parturition. We suggest that epimeletic behaviour may be common in the highly social Chiroptera, especially among primiparous or inexperienced females and/or when an expectant mother experiences difficulty during labour and parturition. Our observations support the hypothesis that roosting associations in bats may be as important as relatedness in explaining the occurrence of apparently altruistic acts.     

Age at first reproduction in philopatric female muriquis (<Emphasis Type="Italic">Brachyteles arachnoides hypoxanthus</Emphasis>)

Female northern muriquis (Brachyteles arachnoides hypoxanthus or B. hypoxanthus) at the Estação Biológica de Caratinga/RPPN-FMA, Minas Gerais, Brazil typically disperse from their natal groups at an average age of 6.1±0.6 years (median =6.0 years, range =5.3–7.8 years, n =22), prior to the onset of puberty and sexual activity. Immigrants do not conceive until at least their second mating season, and the minimum interval from immigration to first reproduction has been 2.0 years. Age at first reproduction in dispersing females, previously estimated at 8.9 years, has now been documented at 9.0 and 9.25 years for two females whose birth dates are known. This is older than the 7.5 years at which the only female previously known to have reproduced in her natal group gave birth. Here, we present new data from a second female that reproduced in her natal group. This female (BA) was first observed to copulate at 5.5 years, and gave birth to her first infant at 7.25 years of age. Her 1.75-year cycling-to-first conception delay was only slightly shorter than the minimum recorded for immigrant females, and thus was not responsible for her young age at first reproduction compared to dispersing females. Although our sample size is small, our findings suggest that early puberty may permit females to reproduce in their natal group, implying a possible link between life history trade-offs and dispersal patterns. Because the only two females that have reproduced in their natal group were maternal sisters, it is also possible that maternal effects on age at puberty can impact dispersal patterns, particularly in small populations of primates.     

Group responses to parturition inMacaca arctoides

The birth of an infant to the highest ranking female in a captive social group of stumptail macaques (Macaca arctoides) is described. Along with certain details of delivery behavior, the responses of group members to the birth are given. The alpha male and two juvenile males were sexually aroused just prior to and during parturition. Adult females paid no special attention to the birth. A subadult nulliparous female and a young adult primiparous female showed interest in the eating of the placenta by the female. The new mother's juvenile son was the monkey closest to her for about two hours after the birth; approaches by other monkeys, especially adult males were avoided. Time of delivery, signs of labor, and the behavior of the mother are also included.     

An observed birth in a free-living chimpanzee (Pan troglodytes schweinfurthii) in Gombe National Park,Tanzania

During 19 years of study of chimpanzees in the Gombe National Park only one birth has been observed; this is probably the first such observation for any chimpanzee in the natural habitat. The birth took place in a nest in a tall tree during the morning. Details were recorded by Tanzanian field staff from a neighbouring tree. Labour and parturition are described as well as the mother’s care of the infant immediately after birth. The mother consumed the placenta as she lay in another nest. Throughout the birth process the mother’s juvenile son remained close by and watched with apparent interest. Another mother and her offspring were present during the birth and an adult male approached the mother while she was feeding on the placenta. Their behaviour is described.     

Grouping-patterns and inter-group relationships of Japanese wild boars (Sus scrofa leucomystax) in the Rokko mountain area

Grouping-patterns and inter-group relationships of Japanese wild boars (Sus scrofa leucomystax) were studied at a provisioning site in the Rokko mountain area, Hyogo Prefecture, Japan, from 1982 to 1983. The observations were divided into four study periods. Almost all wild boars were individually identified and their kin-relationships were followed from the beginning of the study. To examine grouping-patterns of Japanese wild boars, the degree of association between any two individuals, based on coexisting time at the provisioning site, was analyzed for each study period. Mother and her litter group (MLG), female littermate group (FLG) and solitary individual were recognized grouping-patterns. Groups did not associate with each other. Male offspring separated from their mother before 1 year of age and become solitary. Separation of female offspring occurred before the subsequent parturition of their mother. The proximate cause of separation was increased aggression by the mother. After separation, female offspring formed a FLG. However, the FLG seemed to be generally a temporary one, maintaining its stability only until parturition of it's members. Thereafter adult females had no close association with any other individuals, except for their own infants. Male offspring disappeared from their natal area, while female offspring could remain there.     

Questionnaire-based survey of parturition in the queen

The lack of scientific data concerning whether parturition in the queen proceeds normally or not may prevent veterinarians and cat owners from recognizing parturition problems in time. A questionnaire-based study of parturition in 197 queens was performed to determine several parameters of parturition and their influence on its progress. The mean length of gestation was 65.3 days (range 57 to 72 days) and it decreased with increasing litter size (P = 0.02). The median litter size was 4.5 kittens (range 1 to 9), with more males (53%) than females (46%) (P = 0.05). Sixty-nine percent of the kittens were born in anterior presentation and 31% in posterior presentation, indicating that either can be considered normal in the cat. Males were born in posterior position (34%) more often than females (26%) (P = 0.03). The mean birth weight was 98 g (range of 35 to 167 g) and decreased with increasing litter size (P < 0.01). Mean birth weight was higher in males and kittens born in posterior presentation (P < 0.01). Forty-four (5%) of the 887 kittens were stillborn. This was not correlated with the presentation at expulsion but stillborn kittens were more often female (P = 0.02) and weighed less than those born alive (P = 0.04). The median interkitten time was 30 min (range 2 to 343 min) and 95% were born within 100 min after expulsion of the preceding kitten. The interkitten time as a measure of the progress of parturition was not influenced by the kitten's gender, presentation at expulsion, birth weight, or stillbirth, or by the parity of the queen. The results of this study can be used to develop reference values for parturition parameters in the queen, both to determine whether a given parturition is abnormal and as the basis for a parturition protocol.     

Sow responsiveness to human contacts and piglet vocalization during 24 h after onset of parturition

Sow responsiveness towards external disturbances and concurrent postural changes are proposed to be an important cause of early piglet crushing. The aim of the present study was to assess whether loose housed sows change their responsiveness over time within the first 24 h after birth of the first piglet upon exposure to different types of human contact and towards piglets’ scream. The responsiveness of the sows (n = 17) was scored during: (i) blood sampling of the sow during 24 h after the onset of parturition, (ii) human handling of a piglet at 0, 30 min, 1, 2, 4, 8 and 24 h after birth, (iii) screaming when a piglet was trapped underneath the sow and (iv) exposure to playback of piglets’ screams at 10 and 24 h after the onset of parturition. A sow was scored as responsive if she changed her posture in response to the stimuli. The behavioral scores were analyzed during three predefined periods: parturition (from birth of first piglet to birth of last piglet), phase 1 (from birth of last piglet to 12 h after birth of first piglet) and phase 2 (from 12 h after birth of first piglet to 24 h after birth of first piglet). The responsiveness of sows towards humans during blood sampling differed between the three periods (p < 0.01), whereas it did not differ between periods during human handling of piglets. During blood sampling, fewer sows were responsive during phase 1 (5%) compared to during parturition (11%) and the later phase 2 (17%). We did not detect any temporal changes in sow responsiveness towards natural incidences of screaming of own trapped piglet between the three periods (it remained high: 80%), whereas sows exposed to playback of piglet screams had a higher probability (p < 0.05) to react at 12 h (50%) than at 24 h (25%). In conclusion, the responsiveness of sows toward direct human contact was lower during the first 12 h postpartum. The careful handling of piglets in the home pen had a minimal effect on the probability of postural changes in sows. However, sows were highly reactive towards the screaming of own trapped piglet during the whole 24 h period pp. The relative lower responsiveness towards playbacks, decreasing from 12 to 24 h pp, cast doubt upon the piglet scream playback test as a useful approach to evaluate maternal responsiveness in sows.