Effects of Passive Social Support in Juvenile Greylag Geese (Anser anser): A Study from Fledging to Adulthood

In general, support by social allies may reduce stress, increase an individual's status and facilitate access to resources. In Greylag geese (Anser anser), offspring stay with their parents for an entire year or even longer and thereby enjoy social support in encounters with other flock members. We investigated the influence of spatial distance to one's allies on the outcome of agonistic encounters in a natural flock situation for a total of 12 sibling groups after fledging. In addition, we tested two groups of hand‐raised juvenile geese over a time span of 11/2 yr. Passive (i.e. not interfering) human supporters of different familiarity were placed at a standard distance during food provisionings, which produced a tight flock situation. Success in agonistic interactions increased with decreasing distance to members of their social unit. The hand‐raised juveniles were more successful in agonistic interactions and showed increased feeding rates when accompanied by a familiar human than when alone or with a non‐familiar human. The effect of the presence of familiar humans on success in agonistic encounters significantly decreased with increasing age, while feeding rates remained elevated. The positive effects of social support were particularly evident in females. We conclude that social support has similar effects and functions in the highly social greylag geese as reported for social mammals.     

Spatial proximity among adult siblings in greylag geese (Anser anser): evidence for female bonding?

By staying close to allies, individuals may enjoy benefits through social support. In the socially monogamous greylag goose (Anser anser), pair-partners, parents, and even human foster parents may provide social support, facilitating access to resources or reducing agonistic pressure. In the present work, we analysed the spatial distribution of individuals within a semi-feral flock of 120 greylag geese, which contained 23 adult sibling groups of 2–4 individuals from 2 to 12 years old. During resting periods we scored dyadic distances between 28 focal individuals of different social categories, their siblings and unrelated control individuals of the same age. Adult female siblings (i.e. those hatched in the same year and raised together) rested significantly closer to each other than to either their brothers or unrelated control individuals. We attribute this to social attraction rather than to just a common preference for the same resting site. Thus, kinship bonds as expressed by cohesion might persist into adulthood, at least in the females. We discuss the potential benefits of proximity between related individuals with regard to reduced social stress via social support. Received: 21 February 2000 / Received in revised form: 16 May 2000 / Accepted: 17 May 2000     

Physiological implications of pair-bond status in greylag geese

In group-living vertebrates, reliable social allies play a decisive role in dealing with stressors. For example, support by social allies is known to dampen glucocorticoid responses. It remains unknown, however, how social embedding affects the sympatho-adrenergic axis as indicated by heart rate (HR) in non-human animals. We studied the relationships between HR, pair-bond status and distance from the pair-partner in twenty-five free-ranging greylag geese (Anser anser) in a natural social environment. In three individuals, we investigated HR responses following partner loss. Overall, we found a context- and sex-dependent difference in HR between paired and unpaired individuals, paired males having a lower HR during agonistic encounters, and unpaired females having a lower HR during resting. Also, in paired females HR increased with increasing distance from the partner. Our data suggest that HR is modulated by pair-bond status in greylag geese in a context- and sex-dependent manner, which may be representative for social vertebrates in general. Despite the low sample size, the present study indicates that proper social embedding may optimize an individual's physiological investment in the social domain. This reduces individual energy expenditure and may benefit health and reproductive success.     

Serial agonistic attacks by greylag goose families, Anser anser, against the same opponent

It is known from primates that alliance partners may support each other's interests in competition with others, for example, through repeated agonistic attacks against a particular individual. We examined serial aggressive interactions between greylag goose families and other flock members. We found that repeated attacks towards the same individual were common and that up to five serial attacks by family members followed an initial attack. Family size did not affect the frequency of such serial attacks. Juvenile geese evidently benefited most from active social support through serial attacks. About 60% of the juveniles' lost primary interactions were subsequently reversed by another family member. This may be one of the reasons why juveniles rank higher in the social hierarchy than would be expected from their age and size alone. Losses in serial attacks predominantly occurred against other, presumably higher-ranking, family geese and ganders. We propose three major functions/consequences of serial attacks. Analogous to primates, serial attacks in greylag geese may serve to reinforce a losing experience of an opponent defeated in a preceding attack. On the side of the winning family, serial attacks may reinforce the experience of winning. Both winning and losing experiences are linked with physiological consequences in higher vertebrates, affecting the future social performance of winners or losers. Finally, serial attacks may signal the agonistic potential of a family to other flock members. This is supported by heart rate data, which indicate that greylags are competent to interpret third-party relationships.     

Interspecific Mate Choice Following Cross-fostering in a Mixed Colony of Greylag Geese (Anser anser) and Canada Geese (Branta canadensis). A Study on Development and Persistence of Species Preferences1

The role of early experience in mate choice and species preference of geese was studied in an experiment where young greylag geese (Anser anser) were cross-fostered by Canada geese (Branta canadensis). In two successive years, all eggs from 9 nests of wild Canada geese were removed and replaced by greylag goose eggs. The young greylag geese then followed their Canada goose foster parents on their southward migration in autumn, to return with them in spring. Of 35 returning birds, all 16 females paired with greylag goose males (100%) whereas of 19 males 5 paired with Canada goose females (14%) and the remaining 14 with greylag goose females (74%). The pair bonds generally persisted as long as both birds were present, but after loss of a partner, the remaining bird usually re-mated. Even when this happened several times during the lifetime of a male, the new mate was always a Canada goose female, showing that the males were sexually imprinted to this species. The Canada goose females which had mated with the greylag ganders also remated when widowed, but their new mate could be either a Canada goose or a greylag goose.     

Ontogenie und Funktion der Rangordnung innerhalb einer Geschwisterschar von Graugänsen (Anser anser L.)

Between the siblings of a family of greylag geese (Anser anser L.) a rank order is established by fighting during the first days after hatching, and also by other, more complex interactions. The position of each goose in the rank order can be defined by three groups of behaviour patterns (aggressive behaviour, “greetings”, escape responses). A clear-cut ranking relation between two individuals is a prerequisite for their close cohesion. The survival function of rank order lies in diminishing the intensity and frequency of agonistic interactions, in diminishing social stress situations and in guaranteeing the close family cohesion.     

Heart rate modulation in bystanding geese watching social and non-social events

Simply observing other individuals interacting has been shown to affect subsequent behaviour and also hormones in 'bystander' individuals. However, immediate physiological responses of an observer have been hardly investigated. Here we present results on individuals' heart rate (HR) responses during various situations, which occur regularly in a flock of greylag geese (Anser anser, e.g. agonistic encounters, vehicles passing by). We recorded simultaneously HR and behaviour of 21 semi-tame free-roaming geese, equipped with fully implanted transmitters. We considered 304 social and 81 non-social events during which the focal individuals did not respond behaviourally. Independent of the spatial distance to the event, these HR responses were significantly greater in social contexts (e.g. departing or landing geese, agonistic interactions) than in non-social situations (e.g. vehicles passing by, thunder). Focal individuals showed a significantly higher maximum HR as well as a greater HR increase in response to agonistic interactions, in which the pair partner or a family member was involved, as compared with a non-affiliated goose. Also, HR was significantly higher when the bystander watched non-affiliated geese interacting, which were higher ranking than the focal. We conclude that these differences are due to different relevance of the recorded events for the focal individual, depending on the individuals involved in the observed interaction.     

Relationship between wild greylag and European domestic geese based on mitochondrial DNA

The origins of the European domestic goose are uncertain. The available information comes from archaeological findings and historical literature, but genetic evidence has hitherto been scarce. The domestic goose in Europe is derived from the greylag goose (Anser anser), but it is not known where the initial domestication took place and which of the two subspecies of greylag goose was ancestral. We aimed to determine the amount and geographical distribution of genetic diversity in modern populations of greylag geese as well as in different breeds of the domestic goose to make inferences about goose domestication. We studied DNA sequence variation in the mitochondrial control region of greylag geese from multiple populations across Europe and western Asia as well as specimens of domestic geese representing 18 modern breeds and individuals not belonging to any recognised breed. Our results show notable differences in genetic diversity between different greylag goose populations and the presence of six mitochondrial haplogroups which show a degree of geographical partitioning. The genetic diversity of the domestic goose is low, with 84% of sampled individuals having one of two major closely related haplotypes, suggesting that modern European domestic geese may derive from a narrow genetic base. The site of domestication remains unresolved, but domestic geese in Turkey were unusually diverse, indicating the importance of further sampling in the vicinity of the eastern Mediterranean and the Near East. There appears to be past or ongoing hybridisation between greylags and domestic geese in particular areas, consistent with field observations.     

The cytogenetics of domestic geese

Hybrids were produced between an African male and several Pilgrim female domestic geese. Partial karyotypes revealed a difference in the fourth largest pair of autosomal chromosomes. This chromosome pair was metacentric in the African, submetacentric in the Pilgrim, and heteromorphic in the hybrids. A similar difference between the putative wild ancestors of the African and Pilgrim breeds has been reported by others. These findings provide cytological evidence to support the traditional opinion that the African breed was derived from the Asiatic swan goose (Anser cygnoides) and the Pilgrim breed was derived from the European greylag goose (Anser anser).     

Heart rate during conflicts predicts post-conflict stress-related behavior in greylag geese

Background

Social stressors are known to be among the most potent stressors in group-living animals. This is not only manifested in individual physiology (heart rate, glucocorticoids), but also in how individuals behave directly after a conflict. Certain ‘stress-related behaviors’ such as autopreening, body shaking, scratching and vigilance have been suggested to indicate an individual''s emotional state. Such behaviors may also alleviate stress, but the behavioral context and physiological basis of those behaviors is still poorly understood.

Methodology/Principal Findings

We recorded beat-to-beat heart rates (HR) of 22 greylag geese in response to agonistic encounters using fully implanted sensor-transmitter packages. Additionally, for 143 major events we analyzed the behavior shown by our focal animals in the first two minutes after an interaction. Our results show that the HR during encounters and characteristics of the interaction predicted the frequency and duration of behaviors shown after a conflict.

Conclusions/Significance

To our knowledge this is the first study to quantify the physiological and behavioral responses to single agonistic encounters and to link this to post conflict behavior. Our results demonstrate that ‘stress-related behaviors’ are flexibly modulated by the characteristics of the preceding aggressive interaction and reflect the individual''s emotional strain, which is linked to autonomic arousal. We found no support for the stress-alleviating hypothesis, but we propose that stress-related behaviors may play a role in communication with other group members, particularly with pair-partners.     

Reproductive behaviour of female Chorthippus biguttulus grasshoppers

Female grasshoppers of acoustically communicating species assume series of reproductive states that are associated with particular behaviours. Studies on laboratory populations of Chorthippus biguttulus (L.) revealed that females of this species lack the period of ‘passive copulatory readiness’, increase their attractiveness to males by sound production and mate multiple times before their first oviposition. In particular, female Ch. biguttulus display a period of ‘primary rejection’ after their imaginal moult during which they reject male mating attempts followed by a period of ‘active copulatory readiness’ in which they produce acoustic signals and may copulate with courting males. Female stridulation generally stimulated male mating activity and stridulating females attracted more male mating attempts than mute females in the same cage, indicating that males preferentially court females that signal ‘active copulatory readiness’. After receipt of a spermatophore, Ch. biguttulus females displayed periods of ‘secondary rejection’ followed by re-establishment of ‘active copulatory readiness’. Acoustic responses of females to male songs, an indicator of reproductive readiness, were significantly reduced until 2 days after mating and remained slightly reduced in comparison to pre-mating levels. Some females mated multiple times before their first oviposition and cycled between ‘secondary rejection’ and ‘active copulatory readiness’.     

Assessing hunters’ ability to identify shot geese: implications for hunting bag accuracy

Reliable hunting bag statistics are a prerequisite for sustainable harvest management. Recently, Internet-based hunting bag reporting systems have been introduced in some European countries, e.g. Denmark, which may enable faster and more detailed reporting. However, reporting of waterfowl bags on a species-specific level may be biased from the individual hunters’ ability to correctly identify species, particularly because juvenile birds can only be identified from subtle differences. We assessed hunters’ ability to identify the five goose species huntable in Denmark. Identifications were made from a line-up of ten full-bodied geese including adults and juveniles. From a total of 2160 identifications made by active hunters, 85.5% were correct while 14.5% were assigned to a wrong species. Active hunters had on average an identification accuracy of 76.0%, highest for Canada goose (99.1%) and lowest for white-fronted goose (74.6%) and bean goose (73.7%). Identification accuracy was significantly lower for juvenile than for adult individuals of white-fronted and bean geese. Correcting the official Danish Bag Record (2013/2014) for identification accuracy, the bags of white-fronted and bean geese increase by 56.5 and 104.4%, respectively, while the bags of greylag and pink-footed geese decrease by 6.7 and 9.0%; the bag for Canada goose remains unchanged. Although identification accuracy is probably higher under field conditions, the study documents that inaccurate species identification is a source of bias in national bag statistics. Hence, improving identification skills by hunters is important to improve bag data accuracy when based on Internet reporting.     

Developmental stress predicts social network position

The quantity and quality of social relationships, as captured by social network analysis, can have major fitness consequences. Various studies have shown that individual differences in social behaviour can be due to variation in exposure to developmental stress. However, whether these developmental differences translate to consistent differences in social network position is not known. We experimentally increased levels of the avian stress hormone corticosterone (CORT) in nestling zebra finches in a fully balanced design. Upon reaching nutritional independence, we released chicks and their families into two free-flying rooms, where we measured daily social networks over five weeks using passive integrated transponder tags. Developmental stress had a significant effect on social behaviour: despite having similar foraging patterns, CORT chicks had weaker associations to their parents than control chicks. Instead, CORT chicks foraged with a greater number of flock mates and were less choosy with whom they foraged, resulting in more central network positions. These findings highlight the importance of taking developmental history into account to understand the drivers of social organization in gregarious species.     

Repeated grazing of a salt marsh grass by moulting greylag geese Anser anser– does sequential harvesting optimise biomass or protein gain?

The effects of simulated goose grazing on common saltmarsh-grass Puccinellia maritima plants were tested on a Danish salt marsh during the flightless moulting period of greylag geese Anser anser (3–21 June 1998). Plants in an area exclosed from the influence of grazing and the nutrient effects of goose faeces were subject to removal of youngest lamina at 3-, 6-, 9- and 18-day intervals during this period. Average biomass and protein accumulation between harvests was highest at defoliation intervals of 9 days or more. Field observations from two separate study areas demonstrated geese returned to regraze the Puccinellia sward after 6–8 days and oesophageal contents from feeding geese showed selection for lamina lengths consistent with the results of clipping every 6 days. Geese therefore regrazed Puccinellia patches at shorter intervals than expected were they to maximise their intake of biomass or protein at each visit. However, total cumulative lamina elongation, equivalent to the long term gain during the entire moult period, showed no significant difference between the three most intensive defoliation treatments, which were significantly greater than those of plants defoliated at 18 day intervals. Highest overall lamina protein levels were maintained at 6- and 9-day defoliation intervals. This suggests geese regrazed Puccinellia patches at a rate that maximised their number of harvests during the flightless period, but maintained highest protein levels and overall biomass in the sward. This suggests, in line with earlier studies, that moulting greylag geese combine dietary selection, reduced nitrogen excretion and regrazing patterns to meet protein demands during regrowth of flight feathers.     

Isolation and characterization of microsatellite marker loci in the greylag goose (Anser anser)

Ten novel polymorphic microsatellite loci were isolated and characterized from the greylag goose, Anser anser, a long-term monogamous and biparental bird. Additionally, five new primers pairs were designed based on previously published microsatellite locus sequences from closely related species. Multiplex polymerase chain reactions conditions were optimized for all 15 primer pairs. The number of alleles ranged from two to 12 per locus with an observed heterozygosity ranging from 0.07 to 0.85. This marker set will be used to determine rates and origins of extra-pair and parasitic young in a population of individually banded greylag geese with known life histories.     

Latitudinal-Related Variation in Wintering Population Trends of Greylag Geese (Anser Anser) along the Atlantic Flyway: A Response to Climate Change?

The unusually high quality of census data for large waterbirds in Europe facilitates the study of how population change varies across a broad geographical range and relates to global change. The wintering population of the greylag goose Anser anser in the Atlantic flyway spanning between Sweden and Spain has increased from 120 000 to 610 000 individuals over the past three decades, and expanded its wintering range northwards. Although population sizes recorded in January have increased in all seven countries in the wintering range, we found a pronounced northwards latitudinal effect in which the rate of increase is higher at greater latitudes, causing a constant shift in the centre of gravity for the spatial distribution of wintering geese. Local winter temperatures have a strong influence on goose numbers but in a manner that is also dependent on latitude, with the partial effect of temperature (while controlling for the increasing population trend between years) being negative at the south end and positive at the north end of the flyway. Contrary to assumptions in the literature, the expansion of crops exploited by greylag geese has made little contribution to the increases in population size. Only in one case (expansion of winter cereals in Denmark) did we find evidence of an effect of changing land use. The expanding and shifting greylag population is likely to have increasing impacts on habitats in northern Europe during the course of this century.     

Feeding by geese on the Filsø Farmland, Denmark, and the effect of grazing on yield structure of spring barley

From the end of August to early May on average 506400 goose days were spent on the farmland of Filsø, 92% by pink-footed geese, 7% by greylag geese and 1% by bean geese. 36% of the goose days were spent on stubbles (autumn), 38% on stubbles with undersown seed grass (autumn-spring), and 21% on new-sown barley fields (spring).
Geese feeding on spring barley consume kernels left on the surface or in the upper surface layers. In 1982 and 1983 trials were set up to quantify the effect of grazing. The effect on yield and plant structure was measured by comparing ungrazed (exclosures) and grazed plots. Grazing by geese significantly reduced the grain yield (7-20%), but the reduction was not proportional to goose usage. The maximum reduction in sprout density was observed in 1982 in an area with heavy goose usage; sprout density was reduced by 72%, but the subsequent yield in the same area was only reduced by 8%. Plants on grazed plots compensated for the reduced sprout density by having more tillers with ears and grains of higher weight. The effect of grazing on stubble, seed grass, and winter cereals on Filsø is discussed.     

布氏田鼠社会行为对光周期的适应格局

对实验室内不同光周期条件下的布氏田鼠陌生成体—亚成体间社会行为模式的初步研究显示:不同光周期下成体对亚成体（特别是对雄性亚成体）的行为反应具性二型特征,长光照周期下（light:dark,14L:10D)成体与亚成体同性组合间的争斗较异性组合间的明显,容忍性低,其中成体雄对亚成体雄尤甚。短光照周期下（light:dark,10L:14D）成体雄对亚成体雄、成体雌对亚成体雌及亚成体雄的相容性均较成体雄对亚成体雌的容忍性低,尤成体雌对亚成体雄的排斥表现更为突出,其容忍性也较长光照条件下对应组的低。结果表明,不同光周期条件下布氏田鼠成体与亚成体间的社会行为模式不同。在自然条件下,这些行为模式是其对环境季节变化的一种有效适应,它影响种群中未成年个体存活、补充、扩散重组以及社群结构、种群空间格局季节变化,对该鼠的生存和繁殖有利。     

Why do goose parents adopt unrelated goslings? A review of hypotheses and empirical evidence,and new research questions

Adoption of unrelated offspring by successful breeders is one form of brood mixing and alloparental care that is widespread among geese and other waterfowl. Biparental care and long-lasting family bonds in geese are likely to affect the costs and benefits of adoption. Most hypotheses that have been proposed to explain this behaviour assume that the separation of the gosling from its original family is accidental, and that adoption forms the 'best of a bad job' solution. For the gosling, adoption is therefore thought to be the only, and thus adaptive, option. For parents, some hypotheses assume that there are costs of adoption (intergeneration conflict), while others assume that there are benefits (mutually beneficial). The few studies of adoption in wild goose populations indicate cost-neutrality or that there are small benefits to parents of adopting young. This agrees with studies of brood size, which suggest that large families provide benefits for goslings and parents alike. By contrast, most observed adoption attempts involve parental aggression against the lone gosling. However, incidental observations are likely to be biased towards adoptions that involve conspicuous behaviour, such as aggression, and might overlook inconspicuous adoptions. Studies of individually marked goslings are needed to identify the background of the adoption goslings in order to identify whether in geese, as in some larids and altricial species, adoption might be an active strategy of offspring to improve their fitness prospects. In addition, more experimental studies are needed to test predictions about the costs and benefits of large families in geese.