Mother-young Relationships in Captive Ungulates: Behavioral Changes over Time

Ungulates are frequently classified into two groups: the “followers”, in which young initially stay close to their mother and the “hiders”, in which young remain at some distance from their mother, often concealed in vegetation. To examine differences in mother-young behavior between the follower and hider groups and among species within these groups, we studied 63 mother-young pairs, representing 22 ungulate species, for 7 weeks after birth of the young. The follower group had consistently higher scores than the hider group on measures of mother-young association, and lower scores on the percentage of time the young spent lying; however, there was considerable variation within each group. In follower species, two measures of mother and young association decreased over time. In hiders, contact between mother and young did not increase over time although the distance between them, while both were lying, decreased. This reduction in the resting distance between mother and young did not always coincide with other changes possibly indicative of the end of the hiding period. The amount of time young hiders spent lying decreased over time.     

Maternal behavior and early development of pampas deer (Ozotoceros bezoarticus) fawns in a semi-captive environment

Maternal–infant interaction and behaviors are adapted to reduce the risk of predation and increase the probability of survival of the species. We determined how mother–young encounters and communication change by the development of the fawn (locomotion and feeding) throughout lactation in pampas deer (Ozotocerus bezoarticus). We registered the behavior of mother–fawn dyads for the first 60 postnatal days. Our results revealed three main developmental stages in the fawn, hiding (~0–19 days), active (~20–39 days), and adult-like (~40–60 days). In the hiding stage, the fawn stayed hidden, closer to the mother, and spent less time active, the mothers vocalized more often than the fawn, licked the fawns intensively while nursing, and were responsible for initiating nursing. In the active stage, licking behavior and vocalizations of the mother decreased while the fawns ate solid food, initiated nursing bouts, and displayed bursts of running activity. Finally, in the adult-like stage, the fawn showed many signs of nutritional weaning, and similar patterns of activity to their mothers. However, the duration and frequency of nursing bouts were unchanged during the study. We described mother–fawn interaction in pampas deer for the first time, and found that increased running activity in fawns anticipates nutritional weaning. Our results also suggest that to adapt to the development of the fawn, mothers only modify the dynamics of interaction and encounters rather than the time invested in nursing, an activity that also plays a role in maintaining mother–young affiliation.     

Stone handling behavior in rhesus macaques (Macaca mulatta), a behavioral propensity for solitary object play shared with Japanese macaques

Stone handling (SH) behavior was systematically studied in a captive troop of rhesus macaques housed at the Primate Research Institute of Kyoto University, and compared with the results of long-term studies of this behavior in Japanese macaques, to evaluate the similarities of SH behavior in these two closely related species. Similar to Japanese macaques, rhesus macaques showed age-related differences in SH. Young animals were more active and displayed more SH patterns and bouts than did adults. Furthermore, the young displayed SH at a higher frequency and their bouts were of a shorter duration, compared to adults. Young adults were more active and displayed more patterns than did older adults. On the other hand, older adults were more conservative and displayed fewer patterns, and engaged in them for longer durations. All individuals displayed SH more frequently in relaxed environmental and social conditions. While lacking an apparent immediate adaptive value, practice of the behavior has been proposed to have long-term functional value for neural and cognitive development in the young and for the maintenance or repair of neuro-pathways in aging macaques that habitually perform the behavior. The results presented here are consistent with what we know about Japanese macaque SH. Given the uniformity of SH behavioral parameters and these two macaque species’ close phylogenetic relatedness, we propose that a similar functional and adaptive value for SH can be inferred for rhesus macaques.     

Leaders benefit followers in the collective movement of a social sawfly

The challenges of maintaining cohesion while making collective decisions in social or aggregating insects can result in the emergence of a leader or leaders. Larval aggregations of the steel-blue sawfly Perga affinis forage nocturnally, and some larvae lead the aggregation on foraging trips more often than expected by chance. We investigated the relationship between these leader and follower roles by comparing the weight and growth of individual larvae with different roles. Our observations reveal no significant difference between the growth of leaders and followers, suggesting that the role of leadership may not provide direct foraging benefits. However, by experimentally manipulating the social structure of larval aggregations, we found that individuals within aggregations that comprise a mixture of leaders and followers enjoy higher growth rates than those in aggregations comprising a single behavioural type. These data demonstrate, for the first time, individual benefits to maintaining a balance of leader and follower roles within larval aggregations, and highlight the importance of considering the perspectives of both leaders and followers when investigating the evolutionary significance of this behavioural variation within animal groups.     

An evolutionary framework for understanding habitat partitioning in plants

Many plant species with overlapping geographic ranges segregate at smaller spatial scales. This spatial segregation—zonation when it follows an abiotic gradient and habitat partitioning when it does not—has been experimentally investigated for over a century often using distantly related taxa, such as different genera of algae or barnacles. In those foundational studies, trade-offs between stress tolerance and competitive ability were found to be the major driving factors of habitat partitioning for both animals and plants. Yet, the evolutionary relationships among segregating species are usually not taken into account. Since close relatives are hypothesized to compete more intensely and are more likely to interact during mating compared to distant relatives, the mechanisms underlying habitat partitioning may differ depending on the relatedness of the species in question. Here, I propose an integration of ecological and evolutionary factors contributing to habitat partitioning in plants, specifically how the relative contributions of factors predictably change with relatedness of taxa. Interspecific reproductive interactions in particular are understudied, yet important drivers of habitat partitioning. In spatially segregated species, interspecific mating can reduce the fitness of rare immigrants, preventing their establishment and maintaining patterns of spatial segregation. In this synthesis, I review the literature on mechanisms of habitat partitioning in plants within an evolutionary framework, identifying knowledge gaps and detailing future directions for this rapidly growing field of study.     

PATTERNS OF PARENTAL CARE AND PARENTAL INVESTMENT IN MARSUPIALS

I. Information on growth, development and care of young has been assembled for 62 species of marsupial. 2. During gestation, development of the marsupial embryo proceeds only so far as to allow the neonate to make its way from the urogenital opening to the mammary area on the abdomen of the female where it attaches to a teat. Specific structural adaptations keep the neonate firmly attached to the teat for at least the first month after birth. 3. Six types of pouch are distinguished ranging from lateral folds of skin, which do not cover the mammary area or enclose the developing young, to a fold of skin that covers the mammary area and forms a deep pouch, completely enclosing the developing young. 4. Although the young is very small at birth and birth is rapid, specific changes in the behaviour of females occur around the time of birth, and a specific birth position is adopted. 5. The time at which marsupial young leave the pouch cannot be equated with birth in eutherians, because of the considerable variations in the type of pouch and in patterns of parental care. From a consideration of the functional development of the young in the pouch, it is suggested that the nearest equivalent to eutherian birth is the time at which the marsupial young achieves homeostasis, when it is well furred and endothermic. 6. Maternal behaviour is influenced by the type of pouch. In all species, the mother keeps the young and the pouch clean by licking, especially when the young are wholly within the pouch or attached to the abdomen. In species with reduced pouches where young are left in a nest at an early stage of development, maternal behaviour includes nest building, defence, and retrieving and carrying the young. These functions are performed by the pouch itself in species with large deep pouches in which the young is carried for a much larger part of its development, and other specific maternal behaviours are infrequent. 7. The patterns of parental care are reviewed over all families of marsupial. Not all members of a family have the same pattern of parental care, which appears to be influenced by many factors including body size, type of pouch, diet, litter size and other aspects of life history strategy. 8. Three patterns of parental care are distinguished: (A) As soon as young begin to release the teat they are no longer carried by the mother, and are left in a nest when still barely furred, ectothermic and before the eyes open. This pattern is found in species with large litter size and a pouch reduced or absent, e.g. some Dasyuridae and some Didelphidae. (B) Young remain in the pouch after they begin to release the teat but are left in a nest, at a later stage of development than in A, when well furred, endothermic and with eyes open. After first pouch exit, there is generally a period when young return to the pouch from time to time. This pattern is found in species with well developed pouches and litters of I or > 1 e.g. Peramelidae, some Didelphidae. (C) Young remain in the pouch after they begin to release the teat. At first pouch exit, the young is well furred and endothermic, and leaves the pouch only for brief periods, gradually spending more time out until permanent pouch exit. It is not usually left in a nest. This pattern is found in species with well developed pouches and litters of one, e.g. Macropodidae. 9. Pattern A is seen particularly in the smaller species in any family, where large litter size means that by the time young release the teat, the litter is about 50% of maternal body weight and a considerable burden. In such species, young are left in a nest as soon as possible. In larger species with patterns B or C, litter size is smaller, and by the time they are no longer carried by the female, the litter is still only 20% of maternal weight. 10. Whatever the pattern of parental care, mortality from birth to permanent pouch exit is not unusually high in marsupials in comparison with eutherians. 11. I suggest that the presence of the pouch and the associated patterns of parental care are important determinants of social organization in marsupials. For much of the period of dependence, the young is small, attached to a teat or in a pouch. The male can make little contribution to parental care, and there is little room for improvement in the care of young in complex social groups. In most species, the female on her own is sufficient caretaker. The male is most likely to increase his own biological fitness by going off to mate again and leaving the female to raise his offspring. 12. Patterns of energy expenditure on offspring by female marsupials were assessed throughout the development of young. Investment before birth was assessed by weight of the neonate, during development by growth rate and the time for which the young was carried (pouch life), and total investment by weight of young at weaning and time from birth to weaning. Regression of measures of investment against maternal body weight allowed comparison of investment in animals of different size. 13. Investment in young before birth is very small. Neonatal marsupials range in size from 0·01 to 1 g, and the largest is less than 0·2 % of the size of the mother. Larger mothers produce larger young which are smaller relative to the mother than are the young of smaller species. Individual young in the family Dasyuridae are particularly small. 14. Growth rates in g/d were calculated over the period from permanent pouch exit to weaning. There is a very close correlation between growth rate and maternal body weight - that of litters increases as the 0·78 power of body weight. During this period the growth rate of individuals is comparable with that of eutherian young during lactation, and in litters it is higher still, suggesting that the difference in patterns of growth are not due to the lower metabolic rate of marsupials. As in eutherians there is considerable individual variation in growth rate; it is very high in litters of small dasyurids, which have individual rates comparable to those of larger species. Young of the family Peramelidae grow and develop rapidly; those of the arboreal folivore Phascolarctos do both slowly. I 5. Pouch life, the period for which the young is carried by the mother, increases with body size; as expected, species with pattern A parental care have shorter pouch lives than species of the same size with patterns B or C, reflecting the early stage of development at which young are left in the nest in pattern A. 16. Time from birth to weaning is also longer in larger species. There is a close relationship of age at weaning with maternal weight, with some significant exceptions. For their size, the family Peramelidae have a very short time from birth to weaning, and the time from pouch exit to weaning is particularly short. Many arboreal species have longer periods of dependence than expected from their size. 17. The weight at weaning of individual young is closely related to MBW^0·71, but the weight of one young relative to maternal body weight shows no trend with size, and ranges from 25–61 %, with a mean of 42 %. 18. Parental Investment, as measured by the function Wt. of litter at weaning × 100/MBW, decreases with increasing size of mother as MBW^0·28. The highest levels of investment are found in very small species. In many small species of the family Dasyuridae, a litter at weaning is > 300% MBW. By contrast, investment in the family Peramelidae is low - at weaning a litter of three is about 50% MBW, comparable with investment in a single young of the family Macropodidae. 19. The evolution of patterns of parental care and investment appears to follow three main lines: (1) Species with large litter size, high levels of investment in litters and in individual young. Investment is directed to growth and not to carrying the young in the pouch, since young are left in a nest at an early stage. Typical of this group is the family Dasyuridae, in which many species make few reproductive attempts per year. (2) Species with litters of more than one, low levels of investment in litters and in individuals, but rapid growth and development of young. Because of the small relative size of young they are carried in the pouch for a large part of the period from birth to weaning. This pattern is shown by the family Peramelidae, and seen as an adaptation to rapid and repeated reproduction in an environment with an extended favourable season. (3) Species with small litter size, lower total investment, but investment in individual young is not low, and investment in carrying young to an advanced stage of development is high. Patterns of this type are found in the Diprotodonta, with extreme development in the Macropodidae. 20. Many of the measures of investment have been expressed as a power function of maternal body weight. The exponents of body weight in these functions are such as to suggest that an important underlying variable is metabolic rate. 21. It has been suggested elsewhere that the marsupial mode of reproduction evolved as an adaptation to environmental uncertainty, in that it allows a reproductive attempt to be abandoned at any time much more readily than in eutherians, thereby increasing the likelihood that a female will survive to reproduce again. I consider this suggestion in the light of patterns of parental investment. For small, short-lived species, any reproductive attempt represents a substantial part of its lifetime reproductive output. Investment in any one reproductive attempt is high, and the cost of replacing an abandoned attempt is so high that it seems unlikely that the desertion of offspring would be an important reproductive strategy in small ancestral marsupials, although it may be an important response to environmental uncertainty in certain large modern macropodids.     

Summer and winter torpor use by a free-ranging marsupial

Torpor is usually associated with low ambient temperatures (T(a)) in winter, but in some species it is also used in summer, often in response to limited food availability. Since the seasonal expression of torpor of both placental and marsupial hibernators in the wild is poorly documented by quantitative data, we investigated torpor and activity patterns of the eastern pygmy-possum Cercartetus nanus (17.4 g) over two seasons. We used radio telemetry to track animals during winter (n=4) and summer (n=5) in a warm-temperate habitat and found that torpor was used in both seasons. In winter all animals entered periods of short-term hibernation (from 5 to 20 days) containing individual torpor bouts of up to 5.9 days. In summer, torpor bouts were always <1 day in duration, only used by males and were not related to daily mean T(a). Pygmy-possums entered torpor at night as T(a) cooled, and rewarmed during the afternoon as T(a) increased. Individuals interspersed torpor bouts with nocturnal activity and the percentage of the night animals were active was the same in summer and winter. Our study provides the first information on torpor patterns in free-ranging C. nanus, and shows that the use of torpor throughout the year is important for energy management in this species.     

Using proximity measures to describe mother-infant relationships

What are the relative roles of mothers and infants in maintaining proximity with one another? How do the relative roles of the mother and infant change over time? What do proximity measures tell us about the mother-infant relationship? Questions such as these have been addressed using proximity indices devised by Hinde and coworkers in the 1960s. In this paper, I review how these indices are derived and discuss what they do, and do not, infer about the nature of the mother-infant relationship. An additional index is proposed which indicates whether the mother or the infant is responsible for the majority of changes in proximity.     

Preference of 12-h-old kids for their mother goat is impaired by pre-partum-induced anosmia in the mother

We investigated whether kids were able to discriminate their own mother from an alien one in a two-choice test on the day of birth when they had access to acoustic, visual and olfactory cues from their mother, and whether this discrimination depended on the selective maternal behaviour of the mother (i.e. exclusive nursing of own kids). When given the choice between their own mother and an alien equivalent dam, 8-h-old kids did not show a significant preference for their dam, whereas 12- and 24-h-old kids did. When given the choice between their own and an alien mother that were both non-selective because they had been rendered peripherally anosmic by irrigation of the nostrils with zinc sulphate, 12-h-old kids did not show a significant preference for their mother. These results are similar to those reported in sheep and may suggest that the contrast of behaviour between their own and an alien mother existing in normosmic does is important for discrimination of dams by kids at this age. Finally, testing 8-h-old kids in a smaller enclosure resulted in some improvement of their performance, although they still failed to display a significant preference for their mother. On the whole, kids are able to discriminate between their own and an alien mother goat as early as previously reported in lambs. The impairment of this ability when mothers are anosmic and not selective suggests that acceptance behaviours displayed by the mother may serve as one of the cues orientating the choice of the kid when given the choice between intact mothers. Finally, the present results do not suggest the existence of fundamental differences in the establishment of a preference for the mother between lambs, which are followers, and kids, which are hiders.     

Assessing cow comfort: effects of two floor types and two tie stall designs on the behaviour of lactating dairy cows

Our objective was to test the effects of two types of stall flooring and two tie stall designs on the behaviour of dairy cows. Using two cross-over designs, 16 lactating Holstein cows were housed for 3-week periods in each of four tie stalls to compare the effects of concrete versus mattress flooring, and a narrow versus a wide stall front opening. When cows were housed on a mattress flooring they increased their total lying time by 1.8h per day (51.0 versus 43.4% time per day). The duration of individual lying bouts was longer on concrete flooring (78 versus 62min) and the same was true for the duration of individual bouts of standing (80 versus 48min). On mattress flooring cows stood up and lay down more often than on concrete (28 versus 20 per day, respectively). On concrete cows spent more time standing without eating (35.2 versus 28.1% of sampling observations). Cows consistently made use of the additional space provided with the wide stall front opening, placing their head in the manger area during 70.6% of lying observations compared to only 43.4% of observations in the narrow stall front. However, measures of general activity were not significantly different for the two stall designs. Cows spent similar amounts of time eating and consumed similar quantities of feed in all four tie stalls. Our results suggest a lack of comfort may be apparent in reduced time spent lying and a subsequent increase in time spent standing without eating. A lower frequency of standing and lying and an increase in the bout duration of lying suggests that cows on concrete flooring have difficulty standing up and lying down.     

Observations on the maternal behaviour of free-ranging domestic pigs

The maternal behaviour of four gilts and one sow having her second litter was studied in two enclosures, measuring 7 and 13 ha. The females left the flocks approximately 24 h before farrowing and moved between 2.5 and 6.5 km during a period of continuous wandering lasting 4–6 h. During this wandering, the pigs seemed to examine and discriminate between different possible farrowing sites. When a site had been chosen, the pigs constructed nests resembling those reported for the European wild boar. After farrowing, the mother and litter remained in or close to the nest for about 9 days. Thereafter, the nests were suddenly abandoned and the piglets changed from hiders to followers. The female and the young joined the flock, but the litter tended to remain an intact social unit throughout lactation and mixed with other piglets only occassionally during resting. No cross-suckling was observed, although some attempts to steal milk from alien mothers were recorded. The piglets were weaned at between 14 and 17 weeks of age. The maternal behaviour was similar to that reported for wild boars and feral pigs. There were individual differences in maternal behaviour among the animals studied, which may be interpreted either as genetically fixed inter-individual differences or as variable strategies, adopted to increase the lifetime reproductive success.     

Social object play among young Japanese macaques (Macaca fuscata) in Arashiyama, Japan

Social object play (SOP), i.e., social play using portable object(s), among young Japanese macaques (Macaca fuscata; 0-4 years old) in the Arashiyama E troop was studied using a modified sequence sampling method from July to October 2000. SOP was a relatively common activity for most of the young macaques and often continued for long periods. Participants used many kinds of object, including edible natural objects and artificial objects, such as plastic bottles, but they never used provisioned food or wild fruit in SOP bouts. An analysis of long bouts (>/=0.5 min) revealed the following interactive SOP features: (1) at any given time, participants used only one object, and only one participant held the object; (2) during SOP play-chasing, the object holder was likely to be chased by others; (3) during long bouts, the object changed hands frequently; and (4) agonistic competition for an object among young macaques was rare. Combinations of sexes, ages, relative ranks, or matrilines of the object holder and non-holder did not affect the tendency that the holder was chased by non-holder(s) during play-chasing. Even when there was a change in object holders, the repetitiveness of this interactive pattern, i.e., that the holder would be chased during SOP bouts, distinguished the SOP structure from that of other types of social play without object(s). General proximate social play mechanisms, such as self-handicapping or role taking, were associated with SOP. Other mechanisms that affected SOP included the following: (1) young macaques treated an object as a target in play competition, and (2) 'being the holder of a target object' was associated with the 'role of the chasee.'     

Social behavior and genital swelling in pregnant chimpanzees (Pan troglodytes)

Previous reports of increased sexual activity associated with genital swelling during chimpanzee pregnancy have suggested that the tumescent pregnant female may be regarded as a regularly cycling estrous female. Analysis of additional data from seven pregnant and eight cycling females, however, indicated that tumescent pregnant females differed from cycling females in some social interactions with males. As in earlier reports, pregnant females were more receptive to male-initiated copulatory bouts. In addition, pregnant females initiated grooming bouts with males much less frequently than did cycling females, and males spent less time grooming and less frequently inspected the genitals of pregnant than cycling cagemates. This tendency to decrease social interaction during pregnancy is consistent with field reports that pregnant chimpanzees prefer the company of their offspring and other noncycling females. In addition, the genital swelling patterns of 40 chimpanzee pregnancies were analyzed. Length of gestation averaged 231.5 days. Although a high degree of individual variation existed, females were swollen an average of 41% of the days observed, yet reached maximum swelling only 8.7% of the time. Genital swelling was less frequent during the third trimester, averaging 50.5, 47.9, and 22.6% tumescent days in each trimester. Young pregnant females showed maximum swelling more often than did older females. This finding is discussed with respect to field reports of intercommunity transfer of young, tumescent female chimpanzees.     

Validation of triaxial accelerometers to measure the lying behaviour of adult domestic horses

Examining the characteristics of an animal’s lying behaviour, such as frequency and duration of lying bouts, has become increasingly relevant for animal welfare research. Triaxial accelerometers have the advantage of being able to continuously monitor an animal’s standing and lying behaviour without relying on live observations or video recordings. Multiple models of accelerometers have been validated for use in monitoring dairy cattle; however, no units have been validated for use in equines. This study tested Onset Pendant G data loggers attached to the hind limb of each of two mature Standardbred horses for a period of 5 days. Data loggers were set to record their position every 20 s. Horses were monitored via live observations during the day and by video recordings during the night to compare activity against accelerometer data. All lying events occurred overnight (three to five lying bouts per horse per night). Data collected from the loggers was converted and edited using a macro program to calculate the number of bouts and the length of time each animal spent lying down by hour and by day. A paired t-test showed no significant difference between the video observations and the output from the data loggers (P=0.301). The data loggers did not distinguish standing hipshot from standing square. Predictability, sensitivity, and specificity were all >99%. This study has validated the use of Onset Pendant G data loggers to determine the frequency and duration of standing and lying bouts in adult horses when set to sample and register readings at 20 s intervals.     

Early locomotor behaviour in genetic stocks of chickens with different growth rates

Reduction in exercise increases the occurrence of lameness in meat-type chickens. Locomotor activity is dramatically reduced during the finishing period in chickens from fast-growing genetic types compared to slow-growing genetic types, but it is not known whether this difference is already present during the starting period and may be influenced by genetic factors. In order to define the effect of genetic origin on early locomotor behaviour, exercise was compared from 1 to 22 days of age in two meat-type chicken stocks differing in growth rate: male broilers (B) which grow fast and are often lame, and male "label rouge" chickens (L) which grow slowly and are rarely lame.Time budget (lying, standing, drinking, eating, walking) was measured by scanning in six repetitions of five birds (density=2.5 birds/m(2)) at 1, 8, 15 and 17 days of age. Standing bouts were analysed by focal sampling at 2-3, 6-7, 13-14 and 20-21 days of age.B chicks spent less time standing than L chicks at 15 days of age (B=13+/-2%, L=24+/-1%, P<0.01) and 17 days of age, and spent more time lying at 17 days of age (B=73+/-3%, L=60+/-4%, P<0.05).The major part (74%) of the total active time observed by focal sampling was linked to feeding activity. At 2 and 3 days, the activity of B chicks was half that of L chicks during standing bouts (duration of walking per bout: 19+/-4 s for B; 45+/-4 s for L, P<0.05). The activity observed by focal sampling during non-feeding bouts at 20-21 days was significantly correlated with the corresponding data recorded at 2-3 days in the same chicks in the B stock but not in the L stock.We concluded that (1) both B and L genetic stocks have the same overall activity during the first 3 days of age (scanning) but they exhibit different organisation and composition of standing bouts (focal sampling). (2) Genetic factors are probably involved in the expression of locomotor behaviour in very young chicks. (3) The correlations between the levels of activity at early and later ages suggest that selection of young mobile broiler chicks might increase activity at a later age and might therefore reduce the occurrence of leg abnormalities.     

Community Organization Among Neotropical Nectar-Feeding Birds

Assemblages of neotropical hummingbirds are organized accordingto parameters of available resources and morphological-behavioralattributes of particular hummingbird species. We distinguishfive types of flowers relative to hummingbird foraging, andwe define six community roles for hummingbirds in exploitationof various flower types. These roles are: high-reward trapliners,which visit but do not defend nectar-rich flowers with longcorollas; territorialists, which defend dense clumps of somewhatshorter flowers; lowreward trapliners, which forage among avariety of dispersed or nectar-poor flowers; territory-parasitesof two types (large marauders and small filchers); and generalists,which follow shifting foraging patterns among various resources.Simple communities on islands usually contain one species oflow-reward trapliner or generalist and one territorial species,and sometimes support one high-reward trapliner; often thesespecies are sexuallydimorphic. More complex mainland communitiessupport varying numbers of species in different roles, dependingon the relative importance and constancy of different flowertypes. High-reward trapliners are particularly important inforest under-stories, while forest canopies and open habitatssupport large numbers of shorter-billed, mobile birds fillingthe other five roles. We conclude by pointing out the many parallelsthat exist with other consumer groups.     

Mother-young relationships in Belgian Blue cattle after a Caesarean section: characterisation and effects of parity

The systematic use of Caesarean section in "double muscled" Belgian Blue cattle can induce ethical concerns. The aims of the following study was to characterise mother-young relationships in such a situation and to assess the effect of parity. Fifteen heifers and 15 cows of the Belgian Blue breed were observed using video recording when isolated with their calf during the 3 days following Caesarean, a rapid surgery with rare occurrence of aggressive behaviour and no sign of cows' discomfort or weakness. All calves were bottle-fed mother's colostrum once before first suckling occurred. Heifers' calves received a supplementary number of three such artificial meals, while cows' calves needed only one. The overall median time to first licking of the calf by the mother was 3.3min without any effect of parity. The mean licking frequency was 29.2+/-15.8 per 24h: heifers licked their calf less frequently than cows, respectively 23.4+/-15.3 per 24h versus 35.1+/-14.5 per 24h (P<0.05). The mean total licking duration was 42.2+/-25.9min per 24h: there was no significant difference between heifers and cows. The overall median time to first suckling was 6.1h, without any effect of parity. The mean suckling frequency was 8.4+/-4.8 per 24h: for half of these suckling bouts, the calf was situated on the left side of the mother, i.e. the side of the scar. The suckling frequency was negatively correlated with the number of artificial meals (r(s)=-0.45,P<0.05). Heifers suckled their calf less often than cows, respectively 6.1+/-3.1 per 24h versus 10.7+/-5.2 per 24h (P<0.01), but the left side proportion was not significantly different. The mean suckling total duration was 35.3+/-21.0min per 24h. The calf was on the left side of the mother during half of this time. Heifers suckled their calf during a shorter time than cows, respectively 26.9+/-20.0min per 24h versus 43.8+/-19.1min per 24h (P<0.05), and the left side proportion was not significantly different. Licking frequency and suckling duration were positively correlated (r=0.43,P<0.05). Only 10% of the mothers (one heifer and two cows) butted their calf and kicking was never observed. Despite the difficult comparison with results of the literature according to various methodologies, mother-young relationships were considered as similar to those reported after natural calving. In our study, cows can be considered as better mothers than heifers.     

Nocturnal behaviour of female Mythimna convecta

Nocturnal behaviour of Mythimna convecta (Walker) (Lepidoptera: Noctuidae) virgin females was studied in the laboratory under 20 °C and 16:8 LD conditions. The periodicity of activity, feeding, calling, pre-oviposition extrusion and oviposition varied with female age and hour of the scotophase. Females called for the first time between the 2nd and 11th scotophases with the peak in the 4th scotophase. Maximum calling occurred on the 7th hour of the scotophase. Young moths called more frequently with shorter bouts while old moths called less often but with longer bouts. In the presence of older females, moths spent significantly more time in pre-oviposition extrusion and resting and less in activity and feeding than they did when only females of the same age or younger were present. There were no significant differences for calling suggesting that pheromones of older calling females did not affect calling of younger females.     

Suckling behaviour in the brown long-eared bat (Plecotus auritus)

Thirty-two adult female brown long-eard bats were taken into captivity. Eight individuals gave birth to single young in captivity (known mother-young pairs), 10 were lactating when captured (putative mother-young pairs), and the remaining 14 bats were non-reproductive. Bats were maintained in five groups consisting of females from sigle(n=3) or mixed (n=2) wild roosts. All bats were housed in outdoor, free-flight enclosures and fed mainly on free-flying noctuid moths. Bats were individually were determined daily (n=152) for a single gruup of bats containing four known mother-young pairs and five non-reproductive bats. The probability of a being attached to the nipple declined from 100% of records at 1-5 days of age to 5% of records at 36-40 days of age. Females were always found suckling their own young. Suckling associations were determined using infra-red sensitive video-recordings of bat behaviour within the roost box. For both known (n=8) and putative mother-young pairs (n=10), there were no records of young attached to lactating females other than their own mothers (from the same or different wild roosts).