Maternal effects on post-weaning physical and social development in juvenile mountain goats (Oreamnos americanus)

Little is known about maternal effects on post-weaning development, yet they may be important because maternal care could have long-term consequences only evident when offspring approach adulthood. We have assessed the effects of maternal age, current reproduction (presence of a kid of the year) and social rank on the body mass, horn length and social rank of 1- and 2-year-old mountain goats (Oreamnos americanus). Maternal reproductive status and social rank did not affect the mass or horn length of either yearlings or 2-year-olds. Maternal age was positively correlated with yearling body mass for males but not females. We could not detect any maternal age effects on body mass of 2-year-olds. Maternal age and spring forage quality were positively correlated with horn length of yearlings of both sexes, but not of 2-year-olds. Juvenile females showed compensatory growth in mass between 1 and 2 years of age, but males did not. Neither sex showed compensatory growth in horn length. None of the maternal characteristics we examined directly affected the social rank of juveniles, which increased with body mass. Social rank in female mountain goats seems to be established early in life and maintained to adulthood. By affecting yearling development, maternal age could affect the reproductive success of males.     

Reproductive effort in relation to maternal social rank in reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves body-mass change during their first winter). In the models, we included September females body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females social rank, whereas late preweaning daily growth rate of calves was not influenced by females rank. Calves preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.Communicated by R. Gibson     

Social transmission of food handling in the context of triadic interactions between adults and young canaries (<Emphasis Type="Italic">Serinus canaria</Emphasis>)

We studied the factors that enhance food recognition and consumption in young canaries when confronted with adults. In contrast to previous studies on canaries, in which social transmission of food habits was studied in the context of dyadic interactions (one juvenile–one adult), we proposed a more realistic framework in which young canaries were studied in the context of triadic interactions, free or not, with adults of both sexes. We found that during free interactions, the young bird only eats with a familiar male and that this association enhances the social transmission of seed handling. When the juvenile was separated from the adults by a transparent partition, it only learned to husk seed if it was present at the feeder at the same time as a familiar adult acting as a demonstrator. The presence of adults that are familiar but do not act as demonstrators does not facilitate social transmission of handling. However, the presence of a familiar, demonstrating female had also no effect on this transmission. Coordination of the actions of the experienced bird and of the naive subject is required for social transmission to occur. Action coordination does not depend solely on the level of familiarity between partners but also on the role played by the demonstrator (here, the adult male) that looks after the juvenile during its transition towards independence.     

Individual roosting strategies in a flock-living bird: movement and social cohesion of hooded crows (Corvus corone cornix) from pre-roost gatherings to roost sites

The function of the conspicuous pre-roost gathering in communally roosting birds is poorly known. We studied movement and social cohesion of radio-tagged hooded crows (Corvus corone cornix) from their daytime location via pre-roost to roost in autumn and winter. With increasing snowfall the previous 24 h, the crows attended pre-roosts farther from the territory, and moved longer from pre-roost to roost. The crows became more likely to roost communally as distance to their territory increased. Attending a pre-roost on average almost doubled the travel distance to a communal roost. Crows were much more likely to join the same roost when they had attended the same pre-roost than otherwise. Breeding mates were more likely to keep together from pre-roost to roost than were other assigned pairs of a territorial male and female. For assigned pairs of non-mates, cohesion from pre-roost to roost decreased with increasing distance to roost, and was higher when both crows roosted communally compared to when at least one of them roosted in its territory. When both roosted communally, cohesion decreased with increasing snowfall and increasing number of crows attending the pre-roost, increased with increasing snow depth, and became higher among two females, compared to other combinations of two crows, with increasing number of crows attending the pre-roost. These patterns may be interpreted as supporting several of the current hypotheses on the function of pre-roost gathering. Electronic Publication     

Weight comparisons and litter size manipulation in Columbian ground squirrels (Spermophilus columbianus) show evidence of costs of reproduction

I studied reproductive costs in the female Columbian ground squirrel (Spermophilus columbianus) using individually marked animals. I compared weight changes during the active season and over winter, and mortality for females that did and did not wean young. Females raising young were heavier at emergence in that spring than unsuccessful ones. Females that did not raise young gained more weight during summer, were heavier than successful females at the time of entry into hibernation, and were heavier emerging from hibernation the following spring. Over-winter mortality was higher for females that reared young compared to reproductively unsuccessful females. A food supplementation experiment showed that energy-rich food can accelerate individuals’ weight gain. Interactions between litter size, birth weight, weight at emergence from the natal burrow, survival of young to yearling age, and maternal fitness were also studied. Litter sizes were experimentally manipulated to evaluate how females cope with costs of rearing one additional young. Birth weight of juveniles was positively correlated with survival to emergence from the natal burrow and with survival to yearling age. Partial litter loss was higher in experimentally enlarged litters than in either experimentally decreased or control litters. Total litter loss, survival of adult females or the probability of weaning young the following year were not affected by the litter size manipulation. Females appear to adjust the size of their litter before birth, and to some extent during lactation, to their ability to wean young. Received: 20 January 2000 / Received in revised form: 12 March 2000 / Accepted: 18 March 2000     

Nestling size rank in the little egret (Egretta garzetta) influences subsequent breeding success of offspring

Few studies have investigated the long-term fitness consequences of nestling size hierarchies in altricial birds. In this study, we investigated whether or not the size rank order of siblings influences subsequent breeding success in the little egret, Egretta garzetta. From a marking program allowing individual recognition of wild birds, we obtained data on the breeding success of 56 pairs comprising individuals for which the size rank order was known. The breeding success in these pairs was positively influenced by the age of the marked bird but negatively affected by the laying date of the pair and the size rank order of the marked individual. There was also a significant difference between breeding colonies. We suggest two main hypotheses for a link between size rank order of individuals and their breeding success and we discuss our results in relation to current hypotheses on the adaptive value of hatching asynchrony. Received: 10 August 1998 / Accepted after revision: 13 December 1998     

Intraspecific parasitism and split sex ratios in a monogynous and monandrous ant (Leptothorax nylanderi)

Sex ratios were bimodally distributed in a population of the monogynous and monandrous ant Leptothorax nylanderi during each of 3 study years. The population-wide investment ratios suggested worker control of sex allocation. Nest-level variation in the proportional investment in virgin queens was not affected by the presence or absence of a queen and only slightly by collecting year, but was correlated with nest size, total sexual investment and, unexpectedly, with differences in nestmate relatedness: small, low-investment nests and nests with several worker lineages produced male-biased sex ratios. Colonies containing several worker lineages arise from usurpation of mature colonies by unrelated founding queens and the fusion of unrelated colonies under strong nest site limitation. In contrast to facultatively polygynous and polyandrous species of social insects, where workers can maximize their inclusive fitness by adjusting sex ratios according to the degree of relatedness asymmetry, workers in mixed colonies of L. nylanderi do not benefit from manipulating sex allocation, as here relatedness asymmetries appear to be the same as in homogeneous colonies. Received: 7 December 1999 / Received in revised form: 29 February 2000 / Accepted: 13 March 2000     

Parental care of replacement clutches in common terns (Sterna hirundo)

In long-lived seabirds with low annual reproductive output, the renesting decision after breeding failure is critical, and the parents have to weigh benefits of replacement clutches against possible future reproductive costs. In this study, we investigated factors influencing renesting decisions in common terns (Sterna hirundo) and compared aspects of breeding biology and body mass between two breeding attempts by the same pairs in each of 4 years of heavy losses due to predation. Renesting birds were characterized by early laying dates and by a high age. Among early breeders, high egg mass reduced the probability of renesting. A long relaying interval coincided with low mass of replacement eggs in one year, and short intervals with high egg mass in another. Further, egg mass decreased and relaying intervals increased the later the predation events occurred. Evidence of high levels of parental care of replacement clutches came from body mass data: female mass increase prior to egg laying was higher in the second attempt than in the first, whereas male mass was lower during the second courtship period than during the first. Male mass also affected relaying intervals and mass of replacement eggs. We conclude that common terns expend high levels of parental care of replacement clutches. Intrinsic factors related to individual quality (age, body condition) seemed most important for renesting decisions and for the degree of parental care, but foraging conditions seemed to have modifying effects. Received: 13 August 1999 / Received in revised form: 5 February 2000 / Accepted: 13 March 2000     

The sex in short supply for matings varies over small spatial scales in a katydid (Kawanaphilanartee, Orthoptera: Tettigoniidae)

In katydids such as Kawanaphilanartee, a female bias in the operational sex ratio (OSR) results in female competition for mates and male choice of mates. Previous work showed that the excess of sexually active females occurs when food availability is low, in part because less food increases the propensity of females to mate as they forage for the large edible spermatophores produced by males. In this study with K.nartee, a pollen-feeding species, we estimate natural variation in numbers of sexually active males and females by assessing male calling activity and the propensity of females to respond to experimental calling males. We found an excess of sexually active males at a site with many flowers and an excess of sexually active females at a site with few flowers about 900 m away. Between-site differences in gut masses of calling males were consistent with the hypothesis that pollen availability controls OSR. Finally, at a third site where flowers were at first scarce, we found that the initial excess in sexually active females changed to an excess of sexually active males after a clump of grass-trees flowered. The mean gut mass of all sampled males from this site increased after flowering. The large variation in OSR that we document for K. nartee highlights the importance of identifying the appropriate spatial and temporal scales over which OSRs are measured in studies of factors controlling sexual selection. Received: 13 May 1997 / Accepted after revision: 27 October 1997     

Carbon-based balanced trophic structure and flows in the offshore Lake Ontario food web before (1987-1991) and after (2001-2005) invasion-induced ecosystem change

Replicate mass-balanced solutions to Ecopath models describing carbon-based trophic structures and flows were developed for the Lake Ontario offshore food web before and after invasion-induced disruption. The food webs link two pathways of energy and matter flow: the grazing chain (phytoplankton-zooplankton-fish) and the microbial loop (bacteria-protozoans) and include 19 species-groups and three detrital groups. Mass-balance was achieved by using constrained optimization techniques to randomly vary initial estimates of biomass and diet composition. After the invasion, production declined for all trophic levels and species-groups except Chinook salmon. The trophic level (TL) increased for smelt, adult sculpin, adult alewife and Chinook salmon. Changes to ecotrophic efficiencies indicate a reduction in phytoplankton grazing, increased predation pressure on Mysis, adult smelt and alewife and decreased predation pressure on protozoans. Specific resource to consumer TTE changed; increasing for protozoans (8.0-11.5%), Mysis (0.6-1.0%), and Chinook salmon (1.0-2.3%) and other salmonines (0.4-0.5%) and decreasing for zooplankton (20.2-15.1%), prey-fish (9.7-8.8%), and benthos (1.7-0.6%). Direct trophic influences of recent invasive species were low. The synchrony of the decline in PP and species-group production indicates strong bottom-up influence. Mass balance required an increase of two to threefold in lower trophic level biomass and production, confirming a previously observed paradoxical deficit in lower trophic level production. Analysis of food web changes suggest hypotheses that may apply to other similar large pelagic systems including, (1) as pelagic primary productivity declines, overgrazing of zooplankton results in an increase in protozoan production and a loss of trophic transfer efficiency, (2) habitat and food web changes increased Mysis predation on Diporeia and contributed to their recent decline, and (3) production of Chinook salmon, the primary piscivore, was uncoupled from pelagic production processes. This study demonstrates the value of food web models to better understand the impact of invasive species and to develop novel hypotheses concerning trophic influences.     

Genetic population structure and sociogenetic organisation in Formica truncorum (Hymenoptera; Formicidae)

Summary The genetic population structure and the sociogenetic organization of the red wood ant Formica truncorum were compared in two populations with monogynous colonies and two populations with polygynous colonies. The genetic population structure was analysed by measuring allele frequency differences among local subsets of the main study populations. The analysis of sociogenetic organisation included estimates of nestmate queen and nestmate worker relatedness, effective number of queens, effective number of matings per queen, relatedness among male mates of nestmate queens and relatedness between queens and their male mates. The monogynous populations showed no differentiation between subpopulations, whereas there were significant allele frequency differences among the subpopulations in the polygynous population. Workers, queens and males showed the same genetical population structure. The relatedness among nestmate workers and among nestmate queens was identical in the polygynous societies. In three of the four populations there was a significant heterozygote excess among queens. The queens were related to their male mates in the polygynous population analysed, but not in the monogynous ones. The data suggest limited dispersal and partial intranidal mating in the populations with polygynous colonies and outbreeding in the populations having monogynous colonies. Polyandry was common in both population types; about 50% of the females had mated at least twice. The males contributed unequally to the progeny, one male fathering on average 75% of the offspring with double mating and 45–80% with three or more matings. Correspondence to: L. Sundström     

Fluctuating asymmetry and sperm transfer in male decorated field crickets (Gryllodes sigillatus)

Fluctuating asymmetry (FA) in limb size in female decorated field crickets (Gryllodes sigillatus) was associated with a reduction in the size of the spermatophore and the amount of sperm transferred by males and an increase in the time taken to transfer a spermatophore following introduction of a female. There was a weaker negative relationship between limb asymmetry in males and sperm number but no significant relationship between asymmetry in either sex and spermatophylax size. In line with a previous study, female size did not appear to influence spermatophore production or mating decisions by males. The results imply that developmental instability affects both gamete production and mating decisions among males, although the relationships between spermatophore size, sperm number and asymmetry in females are unlikely to be the result of males perceiving differences in female FA. Received: 24 July 1999 / Received in revised form: 22 November 1999 / Accepted: 31 December 1999     

Reproductive suppression and inbreeding avoidance in wild populations of co-operatively breeding meerkats (Suricata suricatta)

Meerkats live in co-operatively breeding familial groups in which reproduction is monopolised by a dominant pair of breeders. Offspring of the breeders are behaviourally subordinate, and typically remain in their natal group as sexually mature, non-breeding helpers. In this study, we investigated the proximate factors limiting subordinate reproduction. Evidence for reproductive suppression by dominants was investigated by comparing life history, behaviour and hormonal profiles of dominants and subordinates. Baseline levels of plasma luteinising hormone (LH) were significantly higher in dominant than in subordinate females. However, following an exogenous injection of gonadotrophin-releasing hormone (GnRH), both categories had comparable concentrations of circulating LH. There were no significant differences in pre- or post-GnRH challenge LH levels in dominant or subordinate males. Reproduction in both dominant and subordinate females rarely occurred in the absence of unrelated males. Given that groups typically comprise parents and offspring, lack of suitable mates emerged as the primary constraint on subordinate reproduction. When this constraint was removed, subordinates typically bred but at a lower rate than dominants. This difference in reproduction may be attributed to intrasexual competition manifested through direct interference by dominant females through subordinate evictions, infanticide and the abandoning of subordinate litters. We argue that differences in reproductive regulation within mammalian co-operative breeding systems may be explained by differences in the mating strategy (inbreeding versus outbreeding) and the probability that subordinates in obligate outbreeding species will encounter unrelated opposite-sex partners. Received: 19 April 2000 / Accepted: 17 July 2000     

Food abundance and parental care in yellow warblers (Dendroica petechia)

Emlen and Oring (1977) suggested that monogamy in birds is maintained because of the need for strict biparental care. A corollary of their suggestion is that paternal care should decrease under conditions of high food abundance. An alternative is that paternal care would increase if males take advantage of the higher food abundance by trying to reduce the length of the nestling feeding period. We tested these two ideas using yellow warblers (Dendroica petechia) by providing some pairs with supplemental food, thereby reducing the importance of biparental care. However, the extra food did not decrease paternal effort, nor did it increase it (Fig. 2). Early in the nestling period experimental females brooded more but visited their nestlings less than did control females, but later, when brooding times decreased, experimental females fed their nestlings more than did control females (Fig. 3). There were no significant differences in nestling survival (Fig. 5), but nestlings in the control treatment were larger and heavier up to 6 days old (Fig. 6). The main effect of supplemental food was on maternal, not paternal behaviour. Models of biparental care assume interdependence between the parental effort of both parents. In this species, however, males and females provide for their brood independently from each other.     

Body condition and parasite intensity correlates with escape capacity in Iberian hares (Lepus granatensis)

Preys require effective anti-predator traits to escape from predator attacks, whereas predators focus on individuals that have lower fitness. This fitness reduction is due to the effects of many regulatory forces such as parasites. We directly observed in the field the escape performance of the Iberian hare (Lepus granatensis) against coursing predators (measured as the time to be taken by greyhounds) and examined the relationships between parasite loads and duration of escape behaviours to test the hypothesis that predators would more easily catch substandard individuals. We found a negative relationship between the hare’s escape duration and parasite burdens in various taxa (Taenia pisiformis cisticercus, intestinal Coccidia) as well as with parasite diversity. Moreover, the escape duration of the hares is related positively to their condition. In the particular case of T. pisiformis cisticercus, we found a negative relationship with body condition, suggesting that its effect on anti-predation ability could be mediated by direct spoliation of host resources and/or by increasing requirements for anti-parasite defence. As parasite loads were not directly manipulated, experimental studies are needed to elucidate any causal links between parasite and anti-predatory capacity in Iberian hares.     

Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Dominance interactions affected patterns of non-reproductive division of labor (polyethism) in the eusocial wasp Mischocyttarus mastigophorus. Socially dominant individuals foraged for food (nectar and insect prey) at lower rates than subordinate individuals. In contrast, dominant wasps performed most of the foraging for the wood pulp used in nest construction. Social dominance also affected partitioning of materials collected by foragers when they returned to the nest. Wood pulp loads were never shared with nest mates, while food loads, especially insect prey, were often partitioned with other wasps. Dominant individuals on the nest were more likely to take food from arriving foragers than subordinate individuals. The role of dominance interactions in regulating polyethism has evolved in the eusocial paper wasps (Polistinae). Both specialization by foragers and task partitioning have increased from basal genera (independent-founding wasps, including Mischo-cyttarus spp.) to more derived genera (swarm-founding Epiponini). Dominance interactions do not regulate forager specialization or task partitioning in epiponines. I hypothesize that these changes in polyethism were enabled by the evolution of increased colony size in the Epiponini. Received: 8 December 1997 / Accepted after revision: 28 March 1998     

Ontogeny of provisioning in two populations of wild golden lion tamarins (Leontopithecus rosalia)

Provisioning weaned young is an important part of the cooperative rearing system of marmosets and tamarins (family: Callitrichidae). Juvenile callitrichids receive a substantial proportion of their diet from all adult group members, whereas juveniles of most other primate species only receive food from the mother infrequently via scrounging. We conducted a longitudinal study of provisioning to 21 young wild golden lion tamarins through their first year of life in two Brazilian reserves. Juveniles were predicted to experience higher provisioning rates at the site with mature forest (the presumed ancestral habitat) and in territories that contained a higher proportion of preferred habitat within the mature forest reserve. Key measures of provisioning did not differ substantially between habitat types. The rates at which juveniles begged for food were very similar across habitats, while the influence of habitat on the rates at which juveniles received food was small and varied across age groups. The most pronounced differences between the reserves were: adult resistance to food-transfer attempts was less frequent, food-offering calls before food transfer were more frequent, and the proportion of prey provisioned was higher in the reserve in which earlier successional-stage forest predominated. Within the mature forest reserve, begging success of young juveniles covaried positively with preferred habitat. Rather than overt caretaker–offspring conflict, an information–feedback loop between juveniles and adults appears to drive provisioning changes as juveniles mature. Direct measures of resource abundance are needed to help clarify the mechanisms by which forest successional stage influences provisioning.     

High relatedness and inbreeding in two species of haplodiploid eusocial thrips (Insecta: Thysanoptera) revealed by microsatellite analysis

Relatedness and inbreeding coefficients were estimated for Oncothrips tepperi and O. habrus, two species of eusocial gall-forming thrips, using data from two microsatellite loci. Relatedness between sets of individuals (sexes and castes) for O. tepperi ranged from 0.5 to 0.85 and for O. habrus estimates were centered around 0.5. Multiple mating in O. habrus was implicated in lowering relatedness within a gall. We estimated the inbreeding coefficient to be 0.38 for both species, the highest estimates known for eusocial insects. Our findings imply that high relatedness and inbreeding are important for the evolution and form of social behavior in Australian gall-forming Thysanoptera. Received: 14 November 1997 / Accepted after revision: 14 March 1998     

Shifting foraging strategies in colonies of the social wasp Polybia occidentalis (Hymenoptera, Vespidae)

Social insect colonies can be expected to forage at rates that maximize colony fitness. Foraging at higher rates would increase the rate of worker production, but decrease adult survival. This trade-off has particular significance during the founding stage, when adults lost are not replaced. Prior work has shown that independent-founding wasps rear the first workers rapidly by foraging at high rates. Foraging rates decrease after those individuals pupate, presumably reducing the risk of foundress death. In the swarm-founding wasps, colony-founding units have many workers, making colony death by forager attrition less likely. Do swarm-founding wasps show similar shifts in foraging rates during the founding stage? We measured foraging rates of the swarm-founding wasp, Polybia occidentalis at four stages of colony development. At each stage, foraging rates correlated with the number of larvae present, which, in the founding stages, correlated with the number of cells in the new nest. Thus, foraging rates appear to be demand-driven, with the level of demand in the founding stage set by the size of nest that is constructed. During the founding stage, foraging rates per larva were high initially, suggesting that colonies minimize the development times of larvae early in the founding stage. Later in the stage, foraging rates decreased, which would reduce worker mortality until new workers eclose. This pattern is similar to that shown for independent-founding wasps and likely results from conflicting pressures to maximize colony growth and minimize the risk of colony death by forager attrition.     

Mating strategies and mating success of fallow (Dama dama) bucks in a non-lekking population

The rutting behaviour of bucks in an enclosed population was investigated between 1988 and 1990. A substantial proportion of the matings were observed. After preliminary observations in the 1987 rut we categorised bucks into one of four rutting strategies based mainly on their degree of territoriality. We investigate the effects of age, dominance and mating strategy on mating success. Territories were aggregated in an area of oak woods and mating success was highly skewed. Bucks of between 5 and 7 years old achieved the majority (over 90%) of observed matings. Mating success was highly correlated with dominance but only weakly related to fighting success. The possession of a territory was crucial to achieving high reproductive success, with a 38-fold difference between the most and least successful strategies. Bucks pursuing the different strategies also differed in the time they commenced groaning, timing of matings, mating interference and the locations where they achieved their matings. Although high-ranking males devoted considerable effort to obtaining and defending a territory only 36% of each buck's matings were achieved on his territory and males tended to abandon these sites when the tendency of females to visit them decreased.