The influence of age, sex and player position on skill demands in junior Australian football.

This study aimed to investigate how ecological constraints influence individual player skill demands in community-level junior Australian Football (AF). Footage from 68 junior AF games, incorporating 1874 unique players, were analysed for individual skill demands such as disposals, kicks, handballs, tackles, marks, and shots at goal per player per game. Multiple linear mixed effect models were created for each skill, using player as a random effect. The impact of age, relative area per player (RAPP), number of players on the team, quarters in midfield, and weather on skill demands was assessed in U9-U14 boys. Finally, the influence of sex on the same skills was explored in U11-U14 boys and girls. Younger players were found to have fewer disposals and handballs, but more tackles, per player than older players (all p < 0.05). Further, more quarters in the midfield increased all skill demands (all p < 0.001). Finally, girls had fewer disposals and marks, but more tackles, per player than boys (all p < 0.05). Overall, age, sex, and position significantly impacted players' exposure to skills in junior AF. Game designers should consider methods to improve skill involvement and reduce tackling exposure early in the junior pathway, with further emphasis on the girls' game.

Determinants of macaques' space use: A test for the ecological constraints model using GPS collars.

As a central topic in Behavioral Ecology, animal space use involves dynamic responses to social and ecological factors. We collared 22 rhesus macaques (Macaca mulatta) from six groups on Neilingding Island, China, and collected 80,625 hourly fixes over a year. Using this high-resolution location data set, we quantified the macaques' space use at the individual level and tested the ecological constraints model while considering various environmental and human interfering factors. As predicted by the ecological constraints model, macaques in larger groups had longer daily path lengths (DPLs) and larger home ranges. We found an inverted U-shape relationship between mean daily temperatures and DPLs, indicating that macaques traveled farther on mild temperature days, while they decreased DPLs when temperatures were too high or too low. Anthropogenic food subsidies were positively correlated to DPLs, while the effect of rainfall was negative. Macaques decreased their DPLs and core areas when more flowers and less leaves were available, suggesting that macaques shifted their space use patterns to adapt to the seasonal differences in food resources. By applying GPS collars on a large number of individuals living on a small island, we gained valuable insights into within-group exploitation competition in wild rhesus macaques.

Biogeographical |distributions of trickster animals.

Human language encompasses almost endless potential for meaning, and folklore can theoretically incorporate themes beyond time and space. However, actual distributions of the themes are not always universal and their constraints remain unclear. Here, we specifically focused on zoological folklore and aimed to reveal what restricts the distribution of trickster animals in folklore. We applied the biogeographical methodology to 16 taxonomic categories of trickster (455 data) and real (93 090 848 data) animals obtained from large databases. Our analysis revealed that the distribution of trickster animals was restricted by their presence in the vicinity and, more importantly, the presence of their corresponding real animals. Given that the distributions of real animals are restricted by the annual mean temperature and annual precipitation, these climatic conditions indirectly affect the distribution of trickster animals. Our study, applying biogeographical methods to culture, paves the way to a deeper understanding of the interactions between ecology and culture.

Ecogeography of group size suggests differences in drivers of sociality among cooperatively breeding fairywrens.

Cooperatively breeding species exhibit a range of social behaviours associated with different costs and benefits to group living, often in association with different environmental conditions. For example, recent phylogenetic studies have collectively shown that the evolution and distribution of cooperative breeding behaviour is related to the environment. However, little is known about how environmental variation may drive differences in social systems across populations within species, and how the relationship between environmental conditions and sociality may differ across species. Here, we examine variation in social group size along a steep environmental gradient for two congeneric cooperatively breeding species of fairywrens (Maluridae) and show that they exhibit opposing ecogeographic patterns. Purple-backed fairywrens, a species in which helpers increase group productivity, have larger groups in hot, dry environments and smaller groups in cool, wet environments. By contrast, superb fairywrens, a species with helpers that do not increase group productivity despite the presence of alloparental care, exhibit the opposite trend. We suggest differences in the costs and benefits of sociality contribute to these opposing ecogeographical patterns and demonstrate that comparisons of intraspecific patterns of social variation across species can provide insight into how ecology shapes social systems.

Mutual mother-pup vocal recognition in the highly colonial Cape fur seal: evidence of discrimination of calls with a high acoustic similarity.

The Cape fur seal (Arctocephalus pusillus pusillus) is one of the most colonial mammal species in the world. Females exclusively nurse their pups for 9 to 11 months, during which they alternate frequent foraging trips at sea with suckling periods ashore. The survival of the pup thus depends on the ability of the mother-pup pair to relocate each other among thousands of individuals. Previous work has demonstrated identity information encoded in pup-attraction (PAC) and female-attraction (FAC) calls. Here, we investigated vocal recognition between mother and pup using playbacks of PAC and FAC performed during the breeding season at Pelican Point, Namibia. Both females and pups were able to specifically discriminate the voice of their pup or their mother from non-affiliated pup or mother. Females were able to memorize previous versions of their pup's calls (evidence of recognition up to 73 days after pup's calls recording). Vocal recognition was demonstrated in pups from 1- to 13-weeks old age. Females and pups did not respond differently to the non-filial or non-mother (for pups) stimulus even if it had a strong acoustic similarity with the filial or mother stimulus. This suggested that Cape fur seal mother-pup pairs have high perceptual and cognitive abilities, allowing individuals to identify kin's vocalizations in a very noisy and confusing environment.

Patterns of sex-biased dispersal are consistent with social and ecological constraints in a group-living cichlid fish.

BACKGROUND: Sex-biased dispersal is a common and widespread phenomenon that can fundamentally shape the genetic structure of the social environments in which animals live. For animals that live in and move between social groups, sex-biased dispersal can result in an asymmetry in the degree of relatedness among cohabiting males and females, which can have strong implications for their social evolution. In this study, we measured the relatedness structure within and across groups of a wild population of Neolamprologus multifasciatus, a highly-social, shell-dwelling cichlid fish endemic to Lake Tanganyika, East Africa. In total, we genotyped 812 fish from 128 social groups at 20 microsatellite loci. Neolamprologus multifasciatus live at high densities, and also experience strong ecological constraints on free movement throughout their habitat. At the same time, they exhibit sex differences in the degree of reproductive competition within their groups and this makes them an excellent model system for studying the factors associated with sex-biased dispersal. RESULTS: Social groups of N. multifasciatus consist of multiple males and females living together. We found that cohabiting females were unrelated to one another (Lynch-Ritland estimates of relatedness = 0.045 ± 0.15, average ± SD), while males shared much higher, albeit variable, levels of relatedness to other males in their groups (0.23 ± 0.27). We uncovered a pronounced decline in relatedness between males living in separate groups as the spatial separation between them increased, a pattern that was not evident in females. Female dispersal was also markedly constrained by the distribution and availability of nearby territories to which they could emigrate. CONCLUSIONS: Our results indicate female-biased dispersal in N. multifasciatus. Our study also highlights how the spatial distribution of suitable dispersal destinations can influence the movement decisions of animals. We also emphasize how sex-biased dispersal can influence the relatedness structure of the social environment in which individuals interact and compete with one another.

Parentage analysis across age cohorts reveals sex differences in reproductive skew in a group-living cichlid fish, Neolamprologus multifasciatus.

Group-living animals are often faced with complex reproductive decisions, namely how to partition within-group reproduction, how to obtain extra-group reproduction and how these two means of reproduction should be balanced. The solutions to these questions can be difficult to predict because ecological conditions can affect the scopes for within-group and extra-group reproduction in complex ways. For example, individuals that are restricted from moving freely around their habitats may have limited extra-group reproductive opportunities, but at the same time, groups may live in close proximity to one another, which could potentially have the opposite effect. The group-living cichlid fish Neolamprologus multifasciatus experiences such ecological conditions, and we conducted an intensive genetic parentage analysis to investigate how reproduction is distributed within and among groups for both males and females. We found that cohabiting males live in "high-skew" societies, where dominant males monopolize the majority of within-group reproduction, while females live in "low-skew" societies, where multiple females can produce offspring concurrently. Despite extremely short distances separating groups, we inferred only very low levels of extra-group reproduction, suggesting that subordinate males have very limited reproductive opportunities. A strength of our parentage analysis lies in its inclusion of individuals that spanned a wide age range, from young fry to adults. We outline the logistical circumstances when very young offspring may not always be accessible to parentage researchers, and present strategies to overcome the challenges of inferring mating patterns from a wide age range of offspring.

Spatial population genetic structure and colony dynamics in Damaraland mole-rats (Fukomys damarensis) from the southern Kalahari.

BACKGROUND: Non-random associations within and among groups of social animals can provide valuable insight into the function of group living and the evolution of social behaviour. Damaraland mole-rats (Fukomys damarensis) demonstrate extremely high levels of reproductive skew, and dispersal is considered to be male-biased in onset and frequency, although asymmetry in dispersal distance is yet to be investigated. Dispersal may be positively correlated with increasing favourable environmental conditions, such as rainfall, however, the effects of ecological constraints on dispersal and colony fission-fusion dynamics have not previously been demonstrated on a spatial scale. Here we provide the first spatial population genetic study for this species. We investigated genetic structure in a population of Damaraland mole-rats from the southern Kalahari in South Africa over 3 years, combining observational dispersal data from mark-recapture with population genetic data to evaluate (1) sex-bias in frequency and distance of dispersal in this species, and (2) the effect of rainfall on fission-fusion dynamics of colonies. RESULTS: Our results demonstrate (1) that both males and females favour local dispersal but on rare occasions may disperse over distances greater than 400 m, (2) that males may disperse over greater distances than females, and (3) that males more frequently immigrate into established neighbouring colonies than females, who predominantly disperse by colony fission, i.e. multiple individuals "budding" from their native colony into a neighbouring territory, thereby establishing new colonies. Furthermore, our results demonstrate (4) elevated dispersal and colony fission in association with increased rainfall, supporting the hypothesis that rainfall may play a significant role in the maintenance and/or disruption of reproductive skew in Damaraland mole-rat populations. CONCLUSION: This study represents the first fine-scale spatial population genetic study in Damaraland mole-rats, and provides relevant insights into colony fission-fusion dynamics in a social and cooperatively breeding species.

Chimpanzee ranging responses to fruit availability in a high-elevation environment.

Most primates experience seasonal fluctuations in the availability of food resources and face the challenge of balancing energy expenditure with energy gain during periods of resource scarcity. This is likely to be particularly challenging in rugged, montane environments, where available energy is relatively low and travel costs are high. Chimpanzees (Pan troglodytes) show extensive behavioral diversity across study sites. Yet, as most research has focused on low- and mid-elevation sites, little is known on how chimpanzees respond to periods of low fruit availability in harsh montane environments. We use focal follow and phenology data to investigate how fruit availability influences daily path length and monthly home range in chimpanzees living in Nyungwe National Park, a montane forest in Rwanda. Nyungwe chimpanzees decreased their daily travel distances during periods of fruit scarcity. However, this decrease in travel effort did not correspond with a decrease in foraging area. Instead, monthly homes ranges shifted location across the study period. Nyungwe chimpanzees occupy a relatively wide altitudinal range and the shifts in monthly home range location may reflect differences in the altitudinal distribution of food resources. Chimpanzee monthly diet was often dominated by one or two species and each of these species were confined to different elevation zones. One important species, Podocarpus latifolius, grew only at high elevations (2,600-2,950 m) and chimpanzees ranged at the altitudinal peak of their range for 2 consecutive months while feeding on this species. Thus, while high elevations are often thought to be harsh environments for primates, they can be an important part of a species' home range when they provide a refugium for densely distributed, important food species.

Experimental vacancies do not induce settlement despite habitat saturation in a cooperative breeder.

The paradox of cooperative breeding, whereby individuals assist others instead of reproducing independently, is generally explained through ecological constraints, but experimental evidence is scant. Here we performed the crucial test of the role of habitat saturation through the experimental creation of vacancies and found that, despite abundant presence of potential mates, subordinates are reluctant to disperse into suitable vacant habitat where conspecifics are absent. We argue that sudden disappearance of multiple group members might indicate a heightened risk of predation. Thereby the results of this study are consistent with the 'perceptual trap' hypothesis: the avoidance of habitats because cues do not accurately reflect their quality. Interestingly, this hypothesis can also explain previous findings, which were widely interpreted as evidence for ecological constraints as a driver of cooperative breeding. Our results can have considerable implications for conservation as they mean that opportunities for colonization might go unexploited.

Winter is coming: harsh environments limit independent reproduction of cooperative-breeding queens in a socially polymorphic ant.

Cooperative breeding animals frequently inhabit harsh environments. It is widely accepted that harsh environments hinder independent reproduction, and this constraint maintains individuals in family groups. Yet the assumption that harsh ecological conditions reduce the success of members of cooperative breeding groups when breeding independently has not been experimentally tested. We addressed this shortcoming using the socially polymorphic Alpine silver ant, Formica selysi. This species has single-queen (independent breeders) and multiple-queen (cooperative breeders) colonies coexisting within populations. We placed newly mated queens emerging from each type of colony to breed alone in either a harsh or mild winter condition and recorded their brood production and survival. Queens emerging from single-queen colonies were unaffected by the winter condition and were more successful at founding a nest independently than queens from multiple-queen colonies. By contrast, queens from multiple-queen colonies had higher mortality after a harsh than after a mild winter. These results support the long-held assumption that harsh environments constrain independent reproduction of members of cooperative breeding groups.

Effect of Group Size on Time Budgets and Ranging Behavior of White-Headed Langurs in Limestone Forest, Southwest China.

The ecological constraints model is well supported by data from most frugivorous primates; however, the prediction power of the model is weak for folivorous primates. From September 2016 to August 2017, we collected comparative data on time budgets, daily path lengths and diets of four groups of white-headed langurs (Trachypithecus leucocephalus), including two large groups (G-DS and G-ZWY) and two small groups (G-LZ and G-NN) in Chongzuo White-Headed Langur National Nature Reserve, Guangxi Province, Southwest China. The aim was to obtain evidence of foraging competition and to test the ecological constraints model on this highly folivorous primate in its karst habitat. The results showed that langurs in the larger groups spent more time traveling, less time resting, and had a longer average daily path length than those in the small groups. Diet composition and dietary diversity were not significantly different between the large and small groups. Our study demonstrates that langurs from large groups suffer scramble competition in limestone forests and supports the validity of the ecological constraints model for folivores.

Resolving the Paradox of Environmental Quality and Sociality: The Ecological Causes and Consequences of Cooperative Breeding in Two Lineages of Birds.

Cooperatively breeding animals occur in virtually every ecosystem on earth. Comparative and biogeographic studies suggest that both benign and harsh-as well as stable and fluctuating-environments can favor the evolution of cooperative breeding behavior. The fact that cooperative societies occur in environments of such contrasting quality creates a paradox of environmental quality and sociality. The dual benefits framework-which leads to the prediction that the ecological consequences of sociality (e.g., range size) vary depending on the benefits that individuals of each species receive by forming social groups-offers a potential resolution to this paradox. Here we use a case study of two avian lineages, starlings (Sturnidae) and hornbills (Bucerotidae), in which environmental unpredictability appears to have opposite effects on the evolution of cooperation to test the dual benefits framework. Consistent with previous work, harsh and unpredictable environments promote cooperative breeding behavior in starlings, which in turn leads to larger geographic ranges. However, cooperatively breeding hornbills occur in benign and stable environments, but sociality does not influence range size. Our study suggests that the paradox of environmental quality and sociality arises largely because cooperative breeding is an umbrella term encompassing social species that form groups for different reasons. We demonstrate that differentiating among the functional causes of social group formation is critical for developing a predictive framework for understanding the evolution of cooperative breeding behavior.

Successional change of testate amoeba assemblages along a space-for-time sequence of peatland development.

It is well established that in ombrotrophic bogs, water-table depth (WTD) is the primary environmental control on testate amoeba distribution. However, the environmental controls on testate amoebae in minerotrophic fens are less well known and successional change in their assemblages associated with fen-bog peatland development has been scarcely investigated. Here we investigate a peatland space-for-time sequence resulting from postglacial rebound on the west coast of Finland, to assess successional patterns in testate amoeba communities and their relationships with environmental variables during peatland development. Sample sites along a 10-km transect from coast to inland ranged from a recently emerged wet meadow to a mature bog. Environmental variables (e.g., peat thickness, carbon and nitrogen content, pH, WTD and vegetation) were measured alongside testate amoeba samples. Results showed that even though the distribution of testate amoebae was to some extent determined by the succession stage, many taxa had wide WTD and pH ranges. The primary environmental control for many taxa changed along the succession. In conclusion, the ecological constraints on testate amoebae in minerotrophic systems are more complex than in bogs. The detected patterns also complicate the use of testate amoebae as a primary proxy in palaeoecological reconstructions where fen-to-bog shifts occur.

Why do some animals mate with one partner rather than many? A review of causes and consequences of monogamy.

Why do some animals mate with one partner rather than many? Here, I investigate factors related to (i) spatial constraints (habitat limitation, mate availability), (ii) time constraints (breeding synchrony, length of breeding season), (iii) need for parental care, and (iv) genetic compatibility, to see what support can be found in different taxa regarding the importance of these factors in explaining the occurrence of monogamy, whether shown by one sex (monogyny or monandry) or by both sexes (mutual monogamy). Focusing on reproductive rather than social monogamy whenever possible, I review the empirical literature for birds, mammals and fishes, with occasional examples from other taxa. Each of these factors can explain mating patterns in some taxa, but not in all. In general, there is mixed support for how well the factors listed above predict monogamy. The factor that shows greatest support across taxa is habitat limitation. By contrast, while a need for parental care might explain monogamy in freshwater fishes and birds, there is clear evidence that this is not the case in marine fishes and mammals. Hence, reproductive monogamy does not appear to have a single overriding explanation, but is more taxon specific. Genetic compatibility is a promising avenue for future work likely to improve our understanding of monogamy and other mating patterns. I also discuss eight important consequences of reproductive monogamy: (i) parentage, (ii) parental care, (iii) eusociality and altruism, (iv) infanticide, (v) effective population size, (vi) mate choice before mating, (vii) sexual selection, and (viii) sexual conflict. Of these, eusociality and infanticide have been subject to debate, briefly summarised herein. A common expectation is that monogamy leads to little sexual conflict and no or little sexual selection. However, as reviewed here, sexual selection can be substantial under mutual monogamy, and both sexes can be subject to such selection. Under long-term mutual monogamy, mate quality is obviously more important than mate numbers, which in turn affects the need for pre-mating mate choice. Overall, I conclude that, despite much research on genetic mating patterns, reproductive monogamy is still surprisingly poorly understood and further experimental and comparative work is needed. This review identifies several areas in need of more data and also proposes new hypotheses to test.

Individual dispersal delays in a cooperative breeder: Ecological constraints, the benefits of philopatry and the social queue for dominance.

Delayed dispersal is a key step in the evolution of familial animal societies and cooperative breeding. However, no consensus has been reached on the ecological and social circumstances driving delayed dispersal. Here, we test predictions from the ecological constraints and benefits of philopatry hypotheses as well as the recently proposed dual benefits hypothesis to better understand the evolution of group-living and cooperative breeding. Furthermore, we consider how individual social circumstances within groups affect dispersal decisions. We examine 11 years of life-history information on a wild population of cooperatively breeding southern pied babblers Turdoides bicolor. We investigate the effects of ecological conditions, natal-group membership and individual social context on male and female dispersal delays, disperser survival and acquisition of dominance. Female dispersal decisions are generally unconstrained by ecological or social circumstances. In contrast, males disperse in response to relaxed ecological constraints, decreases in nepotistic tolerance or when low social rank in the queue for dominance decreases their likelihood of gaining a dominant breeding position. Early dispersal by end-of-queue males often leads to a head-of-queue subordinate position in a non-natal group, thereby increasing access to dominant breeding positions. However, males and females remaining in natal groups gain benefits of philopatry via increased survival and, for head-of-queue males, very high likelihood of acquisition of a breeding position. Overall, predictions from the dual benefits hypothesis best describe these results, while some predictions from each of the ecological constraints and benefits of philopatry hypotheses were supported. The benefits of living and working together (collective action benefits) in large stable groups are of central importance in shaping dispersal delays in southern pied babbler societies. In addition, position in the subordinate social queue for dominance is the key in determining access to reproduction, particularly for males. This research highlights the importance of considering the costs and benefits of individual social circumstances in dispersal decisions and illustrates how the dual benefits hypothesis offers new perspectives in understanding delayed dispersal.

Delayed dispersal and prolonged brood care in a family-living beetle.

Delayed juvenile dispersal is an important prerequisite for the evolution of family-based social systems, such as cooperative breeding and eusociality. In general, young adults forego dispersal if there are substantial benefits to remaining in the natal nest and/or the likelihood of dispersing and breeding successfully is low. We investigate some general factors thought to drive delayed juvenile dispersal in the horned passalus beetle, a family-living beetle in which young adults remain with their families in their natal nest for several months before dispersing. Fine-scale population genetic structure indicated high gene flow between nest sites, suggesting that constraints on mobility are unlikely to explain philopatry. Young adults do not breed in their natal log and likely disperse before reaching breeding age, suggesting that they do not gain direct reproductive benefits from delayed dispersal. We also examined several ways in which parents might incentivize delayed dispersal by providing prolonged care to adult offspring. Although adult beetles inhibit fungal growth in the colony by manipulating both the nest site and deceased conspecifics, this is unlikely to be a major explanation for family living as both parents and adult offspring seem capable of controlling fungal growth. Adult offspring that stayed with their family groups also neither gained more mass nor experienced faster exoskeleton development than those experimentally removed from their families. The results of these experiments suggest that our current understanding of the factors underlying prolonged family living may be insufficient to explain delayed dispersal in at least some taxa, particularly insects.

Costs and benefits of group living in primates: an energetic perspective.

Group size is a fundamental component of sociality, and has important consequences for an individual's fitness as well as the collective and cooperative behaviours of the group as a whole. This review focuses on how the costs and benefits of group living vary in female primates as a function of group size, with a particular emphasis on how competition within and between groups affects an individual's energetic balance. Because the repercussions of chronic energetic stress can lower an animal's fitness, identifying the predictors of energetic stress has important implications for understanding variation in survivorship and reproductive success within and between populations. Notably, we extend previous literature on this topic by discussing three physiological measures of energetic balance-glucocorticoids, c-peptides and thyroid hormones. Because these hormones can provide clear signals of metabolic states and processes, they present an important complement to field studies of spatial and temporal changes in food availability. We anticipate that their further application will play a crucial role in elucidating the adaptive significance of group size in different social and ecological contexts.This article is part of the themed issue 'Physiological determinants of social behaviour in animals'.

Mother-child emotion dialogues: the disrupting effect of maternal history of communal sleeping.

The study examined associations between mothers' upbringing background (kibbutz vs. home) and maternal-sensitive guidance of emotional conversations with their preschool children; children's cooperation and exploration; and the coherence of their conversations. Using a quasi-experimental design, 112 children and their mothers (72 kibbutz raised, 40 home) completed the Autobiographical Emotional Event Dialogue. We hypothesized that maternal kibbutz upbringing would be associated to lower levels of mothers' sensitive guidance of the conversations, children's lower cooperation and exploration, and lower overall coherence. Results showed no upbringing-related differences for the mothers, but significant differences were found for the children, with children of kibbutz-raised mothers showing less cooperation and exploration, and lower levels of coherence for these dyads. The role of maternal background in shaping the interaction with their child is discussed.

Climate predicts which sex acts as helpers among cooperatively breeding bird species.

Among avian cooperative breeders, help in raising offspring is usually provided by males or by both sexes. Sex bias in helping should evolve in response to sex-specific ecological constraints on independent reproduction, with mate shortage for males and breeding vacancy shortage for each sex. Given that male-biased adult sex ratios are prevalent among birds, we predict that male-only helping mainly occurs in temperate species where fast population turnovers deriving from low adult annual survival allow all adult females to hold breeding vacancies, whereas some males overflow as helpers, and both-sex helping in tropical species where saturated habitats prevent not only males, but also females from breeding themselves. As expected, we found that across species, adult survival increased towards tropical zones and warmer climates, and higher adult survival tended to be associated with both-sex helping. Furthermore, sex bias in helping was predicted by latitude and ambient temperature. Our findings of demographic response of species to climate as a potential determinant of bias in helper sex uncover how ecological constraints operate to limit independent reproduction in sex-specific ways.