Suckling behavior of eland antelopes (Taurotragus spp.) under semi-captive and farm conditions

Mother–offspring interactions soon after parturition play a key role in the survival of mammals. We investigated the suckling behavior of semi-captive Western Derby eland (Taurotragus derbianus derbianus) in a 60-ha enclosure covered by dense savanna vegetation in Senegal and farmed Common eland (T. oryx) on an open 2-ha pasture in the Czech Republic. We hypothesized that the basic pattern of suckling bout duration and mother–offspring interactions would be similar between species, but would vary in response to the environmental conditions and breeding system. During three calving periods, we observed the suckling of 27 and 23 calves of Derby and Common elands, respectively, between the ages of 1–5 months, and the interactions between mother and calf before and during suckling. Suckling bout duration increased with the age of the calves for both elands. However, in Derby elands we recorded longer suckling bouts in male than female calves and shorter suckling bouts in primiparous mothers than multiparous ones; no differences were found in farmed Common elands. The animals’ active approach to mother–offspring contact, for example naso–anal contact, and initiation and termination of suckling, resulted in longer suckling bouts in Derby elands. The results suggest that Derby elands that range over a large enclosure with dense vegetation cover adjust their maternal behavior in compliance with potential predator risk, facing a trade-off between nursing and vigilant behavior in the wild. The suckling behavior of farmed elands, on the other hand, reflects the conditions of captivity without predators and with the small available area enabling permanent visual contact of animals.     

Playful interactions in family groups of the common marmoset (Callithrix jacchus jacchus)

Play behaviour in six family groups of common marmosets (Callithrix jacchus jacchus) is described. Playful behaviour patterns comprised a distinct repertoire of behaviour and the patterns formed ordered sequences. The differences in the play of adults and offspring are described and analysed. Adult play was infrequent and most took the form of adult-offspring interactions; it contained fewer bouts of chasing and was less active, and adults elicited fewer bouts. No differences attributable to sex could be determined in the play of offspring and the only pattern that decreased with age was bouncing gait. Possible partner preferences were analysed, and it was found that marmoset twins showed an overall preference for playing with the twin. The vocalizations heard during bouts of play were recorded and analysed and their relationship to non-playful vocalization discussed; during bouts of play younger animals were found to vocalize to older animals. Marmoset play wrestling differed in form from aggressive wrestling, and patterns occurred during aggressive interactions which were not observed in play. The play of marmosets is discussed with regard to the socioecology of the species and compared to the play of other mammalian species.     

Food choice causes interrupted feeding in the generalist grasshopper Schistocerca americana: further evidence for inefficient decision-making

Individual grasshoppers were observed foraging on either single food plant species or binary mixtures of them. Individuals ate slightly less on the mixtures, but grew faster on them. In the mixtures, individuals fed regularly from both plant species, typically eating both within a single meal. Feeding bouts were longer on the preferred plant species in the mixture. The pattern of foraging behavior was markedly different in mixtures and single food treatments. In mixtures, although the total length of time engaged in ingestion over the day was slightly less than in single food treatments, there were relatively more short feeding bouts and more gaps between feeding bouts within meals, so that the time spent in activities associated with the food was markedly longer on mixtures. This is discussed in terms of efficiency and the problem of choice that faces individual generalists. The efficiency factor and its impact on vigilance is added to the other costs of being a polyphagous herbivore.     

Decision making in grooming by Japanese macaques (Macaca fuscata)

I analyzed the temporal organization of individual Japanese macaques’ (Macaca fuscata) grooming sequences in 14 mothers and 13 offspring of different age/sex classes and 4 nonkin females. I hypothesized that preceding grooming affects subsequent grooming by the same individual. Grooming bouts were likely to be terminated as the bouts became longer when females groomed nonrelatives. Moreover, the duration of first bouts was longer than that of following bouts. These effects were also seen in grooming of mothers by their offspring > 1 year old and that of adult and adolescent female offspring by their mothers. In contrast, neither the duration of first bouts nor the number of preceding bouts had much effect on the occurrence or duration of subsequent bouts in any subject.     

Maternal attendance behaviour of sympatrically breeding Antarctic and subantarctic fur seals,Arctocephalus spp., at Macquarie Island

Maternal attendance behaviour was studied in Antarctic (Arctocephalus gazella) and subantarctic fur seals (Arctocephalus tropicalis) which breed sympatrically at subantarctic Macquarie Island. Data on attendance were obtained using telemetric methods. Both species undertook two types of foraging trips: overnight foraging trips which were of less than 1 day duration and occurred exclusively overnight, and extended foraging trips which lasted longer than 1 day. The mean duration of overnight foraging trips was 0.43 and 0.39 days, while the duration of extended foraging trips was 3.6 and 3.8 days in A. gazella and A. tropicalis, respectively. The duration of overnight and extended foraging trips did not differ significantly between species. Two types of shore attendance bouts that differed in duration were also observed in these species. Short attendance bouts lasted less than 0.9 days, while long attendance bouts lasted longer than 0.9 days. Short attendance bouts lasted 0.4 and 0.5 days, while long attendance bouts lasted 1.6 and 1.7 days in A. gazella and A. tropicalis, respectively, and did not differ significantly between species. The most significant differences between the attendance behaviour of both species was in the percentage of foraging time allocated to overnight foraging trips (15% and 25% in A. gazella and A. tropicalis, respectively), and the percentage of time spent ashore (30% and 38% in A. gazella and A. tropicalis, respectively). The nearness of pelagic waters to Macquarie Island is considered to be the main reason that lactating females are able to undertake overnight foraging trips. These trips may be used by females as a means of optimising the costs of fasting and nursing ashore. Females may be able to save energy by only nursing pups when milk transfer efficiencies are high, and reduce the time and energy costs of fasting ashore when milk transfer efficiency is low. Of the female A. gazella that still carried transmitters at the end of lactation, 83% continued regular attendance for between 21 and 150 days post-lactation (when data collection ceased). Overwintering of A. gazella females at breeding sites has not been previously reported in other populations. Accepted: 10 November 1998     

Increased Anxiety in Offspring Reared by Circadian Clock Mutant Mice

The maternal care that offspring receive from their mothers early in life influences the offspring’s development of emotional behavior in adulthood. Here we found that offspring reared by circadian clock-impaired mice show elevated anxiety-related behavior. Clock mutant mice harboring a mutation in Clock, a key component of the molecular circadian clock, display altered daily patterns of nursing behavior that is fragmented during the light period, instead of long bouts of nursing behavior in wild-type mice. Adult wild-type offspring fostered by Clock mutant mice exhibit increased anxiety-related behavior. This is coupled with reduced levels of brain serotonin at postnatal day 14, whose homeostasis during the early postnatal period is critical for normal emotional behavior in adulthood. Together, disruption of the circadian clock in mothers has an adverse impact on establishing normal anxiety levels in offspring, which may increase their risk of developing anxiety disorders.     

Evolutionary genetics of the suiformes as reconstructed using mtDNA sequencing

We have amplified and sequnced the entire mitochondrial DNA cytochromeb gene from four species of Suidae: babirusa, warthog, bearded pig, and some specimens belonging to different subspecies and populations of wild and domestic pigs (Sus scrofa). These sequences were aligned with additional mammalian sequences retrieved from the literature and were used to obtain phylogenetic trees of the Suiformes (Artiodactyla). Several species of Carnivora, Perissodactyla. Cetacea, and other Artiodactyla were used as outgroups. Molecular phylogenetic relationships among the Suiformes reflect their current taxonomy: Hippopotamidae, Tayassuidae, and Suidae are separated by deep genetic gaps, and the division of the Suidae into the subfamilies Babyrousinae., Phacochoerinae, and Suinae has strong genetic correlates. Cytochromeb sequences show differences among Asian and Western populations ofSus scrofa, agreeing with other genetic information (karyotypes blood groups, and protein variability). The two Italian subspecies of wild boar have unique mtDNA cytochromeb haplotypes. The evolutionary rates of cytochromeb sequences are different at transitions versus transversions as well as at first, second, and third positions of codons. Therefore, these classes of substitutions reached different levels of mutational saturation. Only transversions and the conservative first and second position substitutions are linearly related to genetic distances among the Suiformes. Therefore, divergence times were computed using unsaturated conserved nucleotide substitutions and calibrated using paleontological divergence times between some Artiodactyla. Transversions apparently evolve at remarkably regular rates in ungulate taxa which have accumulated less than 20% estimated sequence divergence, corresponding to about 40–45 million years of independent evolution. Molecular, information suggests that Hippopotamidae and Tayassuidae are not closely related (as stated by Pickford, 1986, 1989, 1993) and that the origin of babirusa and warthog (about 10–19 and 5–15 million years ago, respectively) is more recent than supported by current evolutionary reconstructions. The inferred origin of bearded pig is about 2.1 million years old, and genetic divergence among differentSus scrofa populations is probably a Pleistocene event. The addition of new sequences of Suiformes does not help in resolving the phylogenetic position ofHippopotamus amphibius, which shows weak but recurrent linkages with the cetacean evolutionary lineage.To whom correspondence should be addressed.     

Anatomy and histology of the babirusa (Babyrousa babyrussa) stomach

Stomachs from six adult and one 12-month-old babirusa (Babyrousa babyrussa) were collected from zoological gardens. The babirusa stomach was larger than that of the domestic pig (Sus scrofa), and possessed a large diverticulum ventriculi. Its gastric glands were confined to a small, easily identifiable unit at the end of the corpus ventriculi, and the connections between the different stomach parts were wide and unrestricted, with the exception of the well-defined connection between the diverticulum ventriculi and the fundus ventriculi. Microscopically, the structure of the cardiac, gastric and pyloric glands was similar to that of Sus scrofa. However, the mucusproducing cardiac glands of babirusa occupied a larger area within the stomach (> 70% versus 33%). The pH in the lumen of the cardiac gland area lay between 5.3 and 6.4 and micro-organisms were found here. It was hypothesised that the babirusa is a nonruminant foregut-fermenting frugivore/concentrate selector.     

Social organization, reproduction and rearing strategies of Callimico goeldii: new clues from the wild

The callitrichines are a specialized radiation of primates that are characterized in part by variable social systems and cooperative infant care. Callimico goeldii, unlike the other callitrichines, have single rather than twin offspring, reducing the need for allocare and permitting synchronous breeding within groups. Low mortality rates among offspring and unstable social groups are suggested to be possible factors that have led to single births among C. goeldii. Single offspring may benefit from greater maternal investment and more frequent food sharing than twin offspring, factors that may help to explain why C. goeldii reaches sexual maturity more rapidly than other callitrichines. In addition, increased breeding opportunities for young C. goeldii females may have selected for rapid maturation rates among this species. Postpartum ovulation and aseasonal resource availability appear to permit females to have biannual birth seasons, further increasing the potential reproductive output.     

Faecal steroid metabolites for non-invasive assessment of reproduction in common warthogs (Phacochoerus africanus), red river hogs (Potamochoerus porcus) and babirusa (Babyrousa babyrussa)

The objectives of this study were to analyse faecal steroid metabolites in African and South East Asian pig species kept in European zoos. Species studied were the warthog (Phacochoerus africanus), the red river hog (Potamochoerus porcus) and the babirusa (Babyrousa babyrussa). Faecal samples were collected 1-3 times per week from non-pregnant and pregnant captive female warthogs (n = 9), red river hogs (n = 7) and babirusas (n = 5). Enzyme-immunoassays for faecal progesterone, androgen, and oestrogen metabolites, were tested for their ability to determine follicular and luteal phases. In all three species, oestrous cycles could be monitored with 20alpha-OH- and 20-oxo-pregnane assays. In contrast, oestrogens and androgens were not useful in characterising follicular activity during the oestrous cycle in any species. Faecal 20alpha-OH- and 20-oxo-pregnane values were significantly correlated. Faecal pregnane concentrations revealed species-specific differences. Luteal phase values of 20alpha-OH-pregnanes were considerably higher than 20-oxo-pregnanes; 20alpha-OH-pregnanes were in the range of 3-10 microg/g in warthogs and red river hogs, whereas concentrations were 30-200 microg/g faeces in the babirusa. Regular oestrus cycles had a length of about 35 days in all three species studied. Results indicated a seasonal influence on the occurrence of reproductive cycles in the warthog with anoestrous periods in the European summer. The red river hog was found to be a seasonal and poly oestrous breeder; oestrus cycles started by January and continued until summer. In contrast, the babirusa showed non-seasonal ovarian cyclicity. In pregnant red river hogs, progesterone metabolites were comparable to luteal phase values of the oestrous cycle during the first 3 months of gestation, but did further increase during the last month of pregnancy. Oestrogens and 17-oxo-androstanes were significantly elevated during the second half of gestation. In summary, the reproductive biology of three exotic pig species was studied using non-invasive faecal steroid analysis and these methods were used for comparative investigations of oestrous cycles, pregnancy and seasonality.     

Behavioural development of twin siamangs (Hylobates syndactylus)

Multiple births are very rare among gibbons. The birth of siamang twins at the Zürich Zoo in 1992 therefore presented a valuable opportunity to observe the development of the twins and to contrast it with a survey of previous reports on the development of single offspring of siamang and gibbons of thelar group. Furthermore, the hypothesis that the presence of twins among siamang may facilitate the occurrence of helping behaviour (defined as the care of offspring by individuals other than their parents) was re-examined (Dielentheis et al., 1991). The Zürich twins (one male and one female) were observed for a total of 74 hr during their first year of life. The results show that: (1) The twins exhibited more rapid behavioural development than that reported for single offspring. (2) A clear difference between the twins was observed: the female twin developed more rapidly than the male. (3) Neither the twins' father nor their older sisterRama was ever observed carrying the twins. The hypothesis ofDielentheis et al. (1991) is hence not supported by the present study, although it is possible that the older sisterRama did not carry the twins because she was younger than the juvenile in that study. (4) Siamangs may have a longer maturation period than gibbons of thelar group.     

Mother-calf relationships and maternal investment in European bison<Emphasis Type="Italic">Bison bonasus</Emphasis>

Mother-young relationships, up to the third month of the calves’ life, were investigated in European bisonBison bonasus (Linnaeus, 1758) from Białowieża National Park breeding centres (Poland). During the first week, calves spent most of their time close to their mothers and were rarely seen in the vicinity of other group members. Highest suckling rate, most frequent vocal contact and agonistic behaviour of mothers towards intruders, were other characteristics of this period. Although being a forest species, European bison exhibit a following type of strategy for offspring protection, typical for ungulates living in open areas; no hiding phase was observed. Suckling rates were similar in calves of both sexes, however, the pattern of nursing was different: male calves suckled longer, but in rarer bouts than female calves. The mothers’ condition, reflected by their social rank, did not influence the suckling rate. No sex differences were recorded in other maternal activities or mother-calf distance. The presumption of a lack of sex-biased maternal investment in European bison is confirmed by the results of other research.     

Intragroup agonistic behavior in captive groups of the common marmosetCallithrix jacchus jacchus

Agonistic behavior was observed in five captive groups of the monogamous monkey Callithrix jacchus jacchus,which normally produces twin offspring every 5 months. Fighting is first recorded when twins are aged 5–10 months. These “twin fights” determine differences in status between combatants that can be detected 5 months later. From what is known of the social behavior of the species, it is postulated that an individual’s twin is its most serious rival and that early fighting is advantageous because injury is minimal while the animal retains its milk dentition. In contrast to twin fights, aggression between same-sex juveniles and adults does not involve long fights and probably serves to reinforce preexisting differences in status. It often results in forcing a group member to the periphery, and therefore, its main function may be to drive mature, same-sexed individuals out of the group. Apart from the relationship between twins, status within the family group is age related and not normally determined by aggression.     

Local variation in weather conditions influences incubation behavior and temperature in a passerine bird

Incubation is an important component of avian parental care and slight changes in incubation temperature can affect offspring phenotype. Although many extrinsic and intrinsic factors may generate variation in incubation temperature, they remain underexplored under natural conditions. Using a robust data set encompassing 55 nests, 22 816 behavioral observations, and > 1 million paired ambient and egg temperatures, we describe the relationships among abiotic factors, female incubation behavior, incubation temperature, and incubation period for tree swallows Tachycineta bicolor. We report a large amount of individual variation in incubation behaviors and average incubation temperatures for our study population. The average on‐bout incubation temperature was 34.1°C, with daily egg temperatures ranging from 18.0–39.2°C. Females modulated the number of times they left the nest and the amount of time they stayed off the nest according to interactions between precipitation and temperature patterns. Models generated from our observations predicted that the number of female off‐bouts was the lowest under warm and dry conditions while more off‐bouts were taken under cold and dry or warm and wet conditions. During cold and dry conditions, females stayed off their nest ～4 times longer than under warm and dry conditions. However, this pattern was reversed under periods of rainfall; females tended to take shorter off‐bouts when it was rainy and cold compared to longer off‐bouts during warmer rain events. Furthermore, variation in female behavior was associated with differences in overall incubation temperature such that females that maintained greater incubation constancy produced higher incubation temperatures at a given ambient temperature than those that displayed lower incubation constancy. Our results provide perspective on the timing of breeding, as some of the advantages of breeding early may be countered by cooler, early season temperatures and precipitation that cause reproducing females to favor self‐maintenance at a potential cost to optimal incubation temperatures for offspring development.     

Maternal care and infant development in <Emphasis Type="Italic">Callimico goeldii</Emphasis> and <Emphasis Type="Italic">Callithrix jacchus</Emphasis>

Callimico goeldii gives birth to single offspring, whereas other callitrichids, including Callithrix jacchus, twin. This study compares maternal effort and infant development in C. goeldii and C. jacchus; it is the first study to look at nursing frequency. Infants were observed from birth for 7 weeks in two captive groups each of C. goeldii and C. jacchus. C. goeldii mothers physiologically invested the same or less than C. jacchus mothers. C. goeldii mothers gained the same amount of weight during pregnancy in absolute terms as did the smaller C. jacchus. This results in a smaller gain in proportion to maternal weight but an equivalent proportional gain on a per fetus basis. C. goeldii mothers nursed their infants less based on duration of nursing bouts compared with C. jacchus mothers. C. goeldii mothers transported their infants exclusively through the first 2 weeks of life, which is longer than C. jacchus mothers, who exclusively transported infants only during the first week of life. As maternal infant carriage declined, other group members transported offspring in both species. C. goeldii infants engaged in independent locomotive sequences later in development and tasted solid foods less frequently than C. jacchus infants when compared at equivalent ages. A single, opportunistic milk sample obtained from a C. goeldii mother when her infant was 48 days old indicates that C. goeldii milk contains gross energy from crude protein within the range of variation observed in Callithrix milk. Despite the similarities in milk quality and prenatal effort in individual fetuses, C. goeldii infants gain weight faster from 0 to 18 months than do C. jacchus infants. A reduction in litter size allows C. goeldii mothers to spend more time carrying their infant and to delay weaning, thereby allowing accelerated infant and juvenile growth rates compared with C. jacchus.     

Feeding behavior ofLemur catta andLemur fulvus

The feeding behavior of two sympatric species of lemurs, Lemur cattaand Lemur fulvus,was studied in an enclosure simulating a natural habitat at the Duke University Primate Center. L. fulvusspent less time feeding during the day than L. catta.But the former species ate more fruit and had longer feeding bouts on preferred food items than L. catta.They also had a shorter food passage time than L. cattaand their choice of resting places was more influenced by food distribution. Furthermore, the two lemur species ate parts of different plant species and showed different reactions to chemical plant components. According to these results, L. fulvusis a more conservative feeder than L. catta.These interspecific differences in feeding behavior may be one of a number of differences that allow the two species to coexist. In allopatry, however, L. fulvusmay also adopt feeding patterns similar to those of L. catta.But L. cattawas never found to change its feeding strategies in different areas. It may be this option of L. fulvusto adopt different feeding strategies in different situations that allows this species to have the widest range of all Malagasy lemurs. Duke University Primate Center Publication No. 259.     

Pupal Period Affects Calling Behavior of the Wheat Moth, Pseudaletia sequax (Lepidoptera: Noctuidae)

The effects of pupal period and age on calling behavior of virgin females of Pseudaletia sequax were determined. Calling behavior of groups of females of similar age of pupation was observed every 10 min for six calling days. Females were considered calling when they assumed a characteristic position: wings and abdomen elevated, displaying the ovipositor and presumably releasing pheromone. Calling behavior was discontinuous. Females with a shorter pupal period took more time to start calling than females with a longer pupal period and called for a longer period. On the first day of calling, females took more time to initiate calling, on average during the seventh hour of the scotophase, whereas on the second day of calling, most females initiated calling during the fifth hour of the scotophase. On subsequent days of calling, the average time to initiate calling changed to the fourth hour of the scotophase. The mean time of calling per day decreased significantly with the number of days since first calling, and the mean number of calling bouts increased with days since first calling.     

Sexual,Seasonal and Group Size Differences in the Allocation of Time between Vigilance and Feeding in the Greater Rhea,Rhea americana

We studied the effect of sex and group size on the proportion of time a greater rhea, Rhea americana, allocates to vigilance and feeding during the breeding and the non-breeding seasons. We analysed 175 records of focal animals that were feeding alone or in groups of 2 to 26 birds. In both seasons, males spent more time in vigilance and less time in feeding than females. Both sexes spent more time in vigilance and less time in feeding during the breeding season. Sexual and seasonal differences in vigilance were the result of different mechanisms. Males had shorter feeding bouts than females but there were no sexual differences in the length of the vigilance bouts. On the contrary, seasonal differences were the result of males and females having longer vigilance bouts during the breeding season but there were no seasonal differences in the length of the feeding bouts. During the non-breeding season, individual vigilance was higher in rheas foraging alone than in groups. In this case, solitary birds had longer vigilance and shorter feeding bouts than birds foraging in groups. We discuss the possible effect of intragroup competition and food availability on the allocation of time between feeding and vigilance in this species.     

Are species differences in maternal effects arising from maternal care adaptive?

Parental care benefits offspring through maternal effects influencing their development, growth and survival. However, although parental care in general is likely the result of adaptive evolution, it does not follow that specific differences in the maternal effects that arise from care are also adaptive. Here, we used an interspecific cross‐fostering design in the burying beetle species Nicrophorus orbicollis and N. vespilloides, both of which have elaborate parental care involving direct feeding of regurgitated food to offspring, to test whether maternal effects are optimized within a species and therefore adaptive. Using a full‐factorial design, we first demonstrated that N. orbicollis care for offspring longer regardless of recipient species. We then examined offspring development and mass in offspring reared by hetero‐ or conspecific parents. As expected, there were species‐specific direct effects independent of the maternal effects, as N. orbicollis larvae were larger and took longer to develop than N. vespilloides regardless of caregiver. We also found significant differences in maternal effects: N. vespilloides maternal care caused more rapid development of offspring of either species. Contrary to expectations if maternal effects were species‐specific, there were no significant interactions between caretaker and recipient species for either development time or mass, suggesting that these maternal effects are general rather than optimized within species. We suggest that rather than coadaptation between parents and offspring performance, the species differences in maternal effects may be correlated with direct effects, and that their evolution is driven by selection on those direct effects.