Genomic ancestry and social dynamics of the last hunter-gatherers of Atlantic France.

Since the early Holocene, western and central Europe was inhabited by a genetically distinct group of Western Hunter-Gatherers (WHGs). This group was eventually replaced and assimilated by the incoming Neolithic farmers. The western Atlantic façade was home to some of the last Mesolithic sites of mainland Europe, represented by the iconic open-air sites at Hoedic and Téviec in southern Brittany, France. These sites are known for the unusually well-preserved and rich burials. Genomic studies of Mesolithic European hunter-gatherers have been limited to single or a few individuals per site and our understanding of the social dynamics of the last Mesolithic hunter-gatherers of Europe and their interactions with incoming farmers is limited. We sequenced and analyzed the complete genomes of 10 individuals from the Late Mesolithic sites of Hoedic, Téviec, and Champigny, in France, four of which sequenced to between 23- and 8-times genome coverage. The analysis of genomic, chronological and dietary data revealed that the Late Mesolithic populations in Brittany maintained distinct social units within a network of exchanging mates. This resulted in low intra-group biological relatedness that prevented consanguineous mating, despite the small population size of the Late Mesolithic groups. We found no genetic ancestry from Neolithic farmers in the analyzed hunter-gatherers, even though some of them may have coexisted with the first farming groups in neighboring regions. Hence, contrary to previous conclusions based on stable isotope data from the same sites, the Late Mesolithic forager community was limited in mate-exchange to neighboring hunter-gatherer groups, to the exclusion of Neolithic farmers.

The ephemeral record: The role of opportunistic animal resources in the archaeology of Pampa and Patagonia.

The acquisition of opportunistic animal resources by hunter-gatherers-such as scavenged carcasses-is a well-known subsistence strategy. It is frequently mentioned in the context of the history of early human evolution, but not regularly considered among the strategies utilized by more recent foragers of the Southern Cone of South America. Historical and ethnographic information presented here suggests that the use of opportunistic animal resources was a strategy used under a number of conditions but only partially documented in the archaeological literature. We also present archaeological evidence from four sites-Guardia del Río, Paso Otero 1, Ponsonby, and Myren-in different Pampean and Patagonian settings where relevant guanaco (Lama guanicoe) bone assemblages were recovered. These sites record minimal anthropic involvement, basically cut marks on some guanaco bones accompanied by few stone tools, that we interpret as evidence for access and use of water logged immobilized or recently dead animals. Archaeological evidence for the use of these scavenging strategies is difficult to obtain at large archaeological sites that usually result from multiple occupations, since the distinction between the acquisition of actively pursued versus opportunistic animal resources is not straightforward. One conclusion from our review is that the best places to find and recognize this evidence are archaeological sites resulting from ephemeral occupations. The inclusion of these sites gives us access to crucial and rarely documented evidence for the long-term survival of hunter-gatherers.

Adipose development is consistent across hunter-gatherers and diverges from western references.

Despite agreement that humans have evolved to be unusually fat primates, adipose patterning among hunter-gatherers has received little empirical consideration. Here we consider the development of adiposity among four contemporary groups of hunter-gatherers, the Aka, Savanna Pumé, Ju'/Hoansi and Agta using multi-level generalized additive mixed modelling to characterize the growth of tricep skinfolds from early childhood through adolescence. In contrast to references, hunter-gatherers show several consistent patterns: (i) children are lean with little fat accumulation; (ii) no adiposity rebound at 5 years is evident; (iii) girls on average have built 90% of their body size, and reach menarche when adiposity is at its maximum velocity; and (iv) a metabolic trade-off is evident in young, but not older children, such that both boys and girls prioritize skeletal growth during middle childhood, a trade-off that diminishes during adolescence when height velocity increases in pace with fat accumulation. Consistent results across hunter-gatherers living in diverse environments suggest that these patterns reflect a general forager pattern of development. The findings provide a valuable baseline for adipose development not apparent from reference populations. We emphasize both generalized trends among hunter-gatherers, and that inter-populational differences point to the plasticity with which humans organize growth and development.

Hydrocarbons in Formicidae: influence of chemical footprints on ant behavioral strategies.

When an insect walks, it leaves chemical cues that derive from the arolium, a tarsal structure. These cues may contain important information about other species that occur in their community and can then mediate interactions of competition, predation, and information about resources with ants from their own colony. The compounds of these cues are released into the substrate in the form of chemical footprints. There are still few species studied, and little is known about the behavior of ants regarding these signals and how they use them in their interactions. Therefore, the aim of this study was to assess the behavioral strategy of different ant species when confronted with chemical footprints left by other ants, as well as identify their compounds and their relationship with the cuticular hydrocarbon profile. The experiments were performed using a Y-maze, where in one of the arms, there were chemical footprints of their own species or of other species, and the other Y arm was footprint-free. The chemical compounds of footprints and cuticle were analyzed by gas chromatography-mass spectrometry. The results show that foragers of all species detect and respond to the presence of chemical cues in the form of footprints left by other ants. Foragers of all species followed footprints of individuals of the same species both nestmates and non-nestmates; however, Neoponera villosa avoided the footprints of Cephalotes borgmeieri, and C. borgmeieri avoided the footprints of the other two species. The chemical compositions of the cuticle and footprints are related to each other and are specific to each species.

Human consumption of large herbivore digesta and its implications for foraging theory.

Vegetal matter undergoing digestion in herbivores' stomachs and intestines, digesta, can be an important source of dietary carbohydrates for human foragers. Digesta significantly increases large herbivores' total caloric yield and broadens their nutritional profile to include three key macronutrients (protein, fat, and carbohydrates) in amounts sufficient to sustain small foraging groups for multiple days without supplementation. Ethnographic reports of routine digesta consumption are limited to high latitudes, but the practice may have had a wider distribution prehistorically. Including this underappreciated resource in our foraging hypotheses and models can substantively change their predictions. Assessing the explanatory power of kilocalorie-centered models relative to ones that attend to humans' other nutritional requirements can help us better address major questions in evolutionary anthropology. This paper explores the foraging implications of digesta in two contexts-sex-divided subsistence labor and archaeologically observed increases in plant use and sedentism-using estimates of available protein and carbohydrates in the native tissues and digesta, respectively, of a large ruminant herbivore (Bison bison).

Complex mortuary dynamics in the Upper Paleolithic of the decorated Grotte de Cussac, France.

The Mid-Upper Paleolithic (Gravettian) karstic Grotte de Cussac (France) contains two areas of human remains in the context of abundant (and spectacular) parietal engravings. The first area (loci 1 and 2) includes the skeleton of a young adult male in a bear nest, rearranged by postdecomposition inundation, and the variably fragmentary remains of at least two individuals distributed across two bear nests, sorted anatomically and with most of the elements constrained to one side of one nest. The second area (locus 3) retains remains of two adults and an adolescent, in upper hollows and variably distributed down the slope, largely segregated into upper versus lower body groups. The only decoration associated with the human remains is red pigment on some of the bones or underlying sediment. The human remains indicate variable nonnatural deposition and manipulation of human bodies, body portions, and skeletal elements of at least six individuals. Moreover, Cussac is unusual in the association of these remains with exceptional parietal art. The complex Cussac mortuary pattern joins growing evidence from other Gravettian sites of variable treatment of individuals after death, within and across sites, in terms of formal deposition of the body versus postmortem manipulation versus surface abandonment. It provides a window onto the social diversity and the complex interactions of the living and the dead among these successful Late Pleistocene foragers.

Large-scale cooperation in small-scale foraging societies.

We present evidence that people in small-scale mobile hunter-gatherer societies cooperated in large numbers to produce collective goods. Foragers engaged in large-scale communal hunts and constructed shared capital facilities; they made shared investments in improving the local environment; and they participated in warfare, formed enduring alliances, and established trading networks. Large-scale collective action often played a crucial role in subsistence. The provision of public goods involved the cooperation of many individuals, so each person made only a small contribution. This evidence suggests that large-scale cooperation occurred in the Pleistocene societies that encompass most of human evolutionary history, and therefore it is unlikely that large-scale cooperation in Holocene food producing societies results from an evolved psychology shaped only in small-group interactions. Instead, large-scale human cooperation needs to be explained as an adaptation, likely rooted in distinctive features of human biology, grammatical language, increased cognitive ability, and cumulative cultural adaptation.

The emergence and intensification of early hunter-gatherer niche construction.

Hunter-gatherers, especially Pleistocene examples, are not well-represented in archeological studies of niche construction. However, as the role of humans in shaping environments over long time scales becomes increasingly apparent, it is critical to develop archeological proxies and testable hypotheses about early hunter-gatherer impacts. Modern foragers engage in niche constructive behaviors aimed at maintaining or increasing the productivity of their environments, and these may have had significant ecological consequences over later human evolution. In some cases, they may also represent behaviors unique to modern Homo sapiens. Archeological and paleoenvironmental data show that African hunter-gatherers were niche constructors in diverse environments, which have legacies in how ecosystems function today. These can be conceptualized as behaviorally mediated trophic cascades, and tested using archeological and paleoenvironmental proxies. Thus, large-scale niche construction behavior is possible to identify at deeper time scales, and may be key to understanding the emergence of modern humans.

The Role of Colony Temperature in the Entrainment of Circadian Rhythms of Honey Bee Foragers.

Honey bees utilize their circadian rhythms to accurately predict the time of day. This ability allows foragers to remember the specific timing of food availability and its location for several days. Previous studies have provided strong evidence toward light/dark cycles being the primary Zeitgeber for honey bees. Work in our laboratory described large individual variation in the endogenous period length of honey bee foragers from the same colony and differences in the endogenous rhythms under different constant temperatures. In this study, we further this work by examining the temperature inside the honey bee colony. By placing temperature and light data loggers at different locations inside the colony we measured temperature at various locations within the colony. We observed significant oscillations of the temperature inside the hive, that show seasonal patterns. We then simulated the observed temperature oscillations in the laboratory and found that using the temperature cycle as a Zeitgeber, foragers present large individual differences in the phase of locomotor rhythms for temperature. Moreover, foragers successfully synchronize their locomotor rhythms to these simulated temperature cycles. Advancing the cycle by six hours, resulting in changes in the phase of activity in some foragers in the assay. The results are shown in this study highlight the importance of temperature as a potential Zeitgeber in the field. Future studies will examine the possible functional and evolutionary role of the observed phase differences of circadian rhythms.

Sexual dimorphism in chimpanzee (Pan troglodytes schweinfurthii) and human age-specific fertility.

Across vertebrates, species with intense male mating competition and high levels of sexual dimorphism in body size generally exhibit dimorphism in age-specific fertility. Compared with females, males show later ages at first reproduction and earlier reproductive senescence because they take longer to attain adult body size and musculature, and maintain peak condition for a limited time. This normally yields a shorter male duration of effective breeding, but this reduction might be attenuated in species that frequently use coalitionary aggression. Here, we present comparative genetic and demographic data on chimpanzees from three long-term study communities (Kanyawara: Kibale National Park, Uganda; Mitumba and Kasekela: Gombe National Park, Tanzania), comprising 581 male risk years and 112 infants, to characterize male age-specific fertility. For comparison, we update estimates from female chimpanzees in the same sites and append a sample of human foragers (the Tanzanian Hadza). Consistent with the idea that aggressive mating competition favors youth, chimpanzee males attained a higher maximum fertility than females, followed by a steeper decline with age. Males did not show a delay in reproduction compared with females, however, as adolescents in both sites successfully reproduced by targeting young, subfecund females, who were less attractive to adults. Gombe males showed earlier reproductive senescence and a shorter duration of effective breeding than Gombe females. By contrast, older males in Kanyawara generally continued to reproduce, apparently by forming coalitions with the alpha. Hadza foragers showed a distinct pattern of sexual dimorphism in age-specific fertility as, compared with women, men gained conceptions later but continued reproducing longer. In sum, both humans and chimpanzees showed sexual dimorphism in age-specific fertility that deviated from predictions drawn from primates with more extreme body size dimorphism, suggesting altered dynamics of male-male competition in the two lineages. In both species, coalitions appear important for extending male reproductive careers.

Changes in human skull morphology across the agricultural transition are consistent with softer diets in preindustrial farming groups.

Agricultural foods and technologies are thought to have eased the mechanical demands of diet-how often or how hard one had to chew-in human populations worldwide. Some evidence suggests correspondingly worldwide changes in skull shape and form across the agricultural transition, although these changes have proved difficult to characterize at a global scale. Here, adapting a quantitative genetics mixed model for complex phenotypes, we quantify the influence of diet on global human skull shape and form. We detect modest directional differences between foragers and farmers. The effects are consistent with softer diets in preindustrial farming groups and are most pronounced and reliably directional when the farming class is limited to dairying populations. Diet effect magnitudes are relatively small, affirming the primary role of neutral evolutionary processes-genetic drift, mutation, and gene flow structured by population history and migrations-in shaping diversity in the human skull. The results also bring an additional perspective to the paradox of why Homo sapiens, particularly agriculturalists, appear to be relatively well suited to efficient (high-leverage) chewing.

Postpartum Maternal Mood Among Hadza Foragers of Tanzania: A Mixed Methods Approach.

Infant and maternal mortality rates are among the highest in the world in low and middle-income countries where postpartum depression impacts at least one in five women. Currently, there is a dearth of data on maternal mood and infant health outcomes in small-scale non-industrial populations from such countries, particularly during the postnatal period. Here, we present the first investigation of postpartum maternal mood among a foraging population, the Hadza of Tanzania. We administered the Edinburgh Postnatal Depression Scale (EPDS) to twenty-three women, all with infants under the age of 12 months. Semi-structured interviews on happiness and unhappiness during the post-partum period served as a validity cross-check for the EPDS. The combined results of the EPDS surveys and the interview responses suggest that a high proportion of Hadza women experience significant mood disturbances following birth and that postpartum unhappiness is associated with self-reports of pain, anxiety, and disturbed sleep patterns. These findings suggest that many of the mothers in our sample are experiencing post-partum unhappiness at levels similar to or higher than those reported for low to middle income countries in general, including Tanzania. These data are critical for improving our understanding of the etiologies of postpartum mood disturbances cross-culturally.

lncRNA profile of Apis mellifera and its possible role in behavioural transition from nurses to foragers.

BACKGROUND: The behavioural transition from nurses to foragers in honey bees is known to be affected by intrinsic and extrinsic factors, including colony demography, hormone levels, brain chemistry and structure, and gene expression in the brain. However, the molecular mechanism underlying this behavioural transition of honey bees is still obscure. RESULTS: Through RNA sequencing, we performed a comprehensive analysis of lncRNAs and mRNAs in honey bee nurses and foragers. Nurses and foragers from both typical colonies and single-cohort colonies were used to prepare six libraries to generate 49 to 100 million clear reads per sample. We obtained 6863 novel lncRNAs, 1480 differentially expressed lncRNAs between nurses and foragers, and 9308 mRNAs. Consistent with previous studies, lncRNAs showed features distinct from mRNAs, such as shorter lengths, lower exon numbers, and lower expression levels compared to mRNAs. Bioinformatic analysis showed that differentially expressed genes were mostly involved in the regulation of sensory-related events, such as olfactory receptor activity and odorant binding, and enriched Wnt and FoxO signaling pathways. Moreover, we found that lncRNAs TCONS_00356023, TCONS_00357367, TCONS_00159909 and mRNAs dop1, Kr-h1 and HR38 may play important roles in behavioural transition in honey bees. CONCLUSION: This study characterized the expression profile of lncRNAs in nurses and foragers and provided a framework for further study of the role of lncRNAs in honey bee behavioural transition.

The meta-group social network of early humans: A temporal-spatial assessment of group size at FLK Zinj (Olduvai Gorge, Tanzania).

Humans are the only primates that maintain regular inter-group relationships and meta-group social networks that enable the inter-group flow of individuals. This is the basis of the band/tribe concept in the anthropology of modern foragers. The present work is a theoretical approach to the development of analytical tools to understand group size and the temporal scale of site occupation in the archaeological record. We selected FLK Zinj as one of the oldest examples of a taphonomically-supported anthropogenic site in which both variables (group size and time) could be modelled using a combination of modern forager regression estimates from their camp sizes and estimates derived from the combined use of modern African foragers' meat consumption rates per day per capita during the dry season and minimum estimates of flesh yields provided by the carcass parts preserved at FLK Zinj. This approach provides the basis for a testable hypothesis which should be further tested in other Oldowan sites. An estimate of 18-28 individuals occupying FLK Zinj was made, which is similar to the estimated 16 individuals of one of the 1.5 Ma Ileret Homo erectus footprint trails. It also shows a similar proportional distribution to Dunbar's equations (group size to neocortex ratio) as documented in modern foragers, which suggest that most of the social network of H. erectus was in the meta-group level as is the case of modern foragers. Irrespective of the range of variation discussed for both variables (group size and length of time represented), it is argued that neither small estimates of time nor small group sizes can account for the formation of FLK Zinj.

Foraging mode, relative prey size and diet breadth: A phylogenetically explicit analysis of snake feeding ecology.

Foraging modes (ambush vs. active foraging) are often correlated with a suite of morphological, physiological, behavioural and ecological traits known as the "adaptive syndrome" or "syndrome hypothesis." In snakes, an ecological correlate often reported in the literature is that ambush-hunting snakes have a higher relative meal size compared to actively foraging snakes which feed on smaller prey items. This "large meal versus small meal" feeding hypothesis between ambush and active foragers has become a widely accepted paradigm of snake feeding ecology, despite the fact that no rigorous meta-analysis has been conducted to support this generalization. We conducted a phylogenetically explicit meta-analysis, which included ca. 100 species, to test this paradigm of snake feeding ecology. We gathered data on prey size by inducing regurgitation by palpation in free-ranging snakes and by examining the stomach contents of preserved museum specimens. When we found prey, we recorded both snake and prey mass to estimate relative prey mass (prey mass/snake mass). We also reviewed published studies of snake feeding ecology to gather similar information for other species. Ambush and active foragers did not differ in minimum or average meal size but the maximum meal sizes consumed by ambush-foraging snakes were larger than the maximum meal sizes eaten by active foragers. This results in ambush-foraging snakes consuming a significantly wider range of meal sizes, rather than being large meal specialists compared to active foragers. We argue that ambush foragers evolved to be more opportunistic predators because they encounter prey less frequently compared to active foragers. This hypothesis is further supported by the fact that ambush foragers also exhibited marginally wider diet breadths, consuming a broader range of prey types in comparison with active foragers. Our study challenges aspects of the foraging syndrome as it is currently conceived, and our results have important implications for our understanding of how foraging mode has shaped the behaviour and physiology of ambush-foraging snakes.

Age-related tooth wear in African rainforest hunter-gatherers.

OBJECTIVES: Central African small-scale foragers subsist primarily on hunting game activities and wild plant-food gathering. Starch-rich tubers are underground storage organs (USOs) and staple food resources in savanna and tropical rainforests. However, little is known about the effect of USO consumption on tooth wear development in living hunter-gatherers. We report age- and sex-dependent tooth wear rates in forest-dwelling Baka Pygmies with well-documented wild-yam-tuber-based diet to explore the long-term impact of USO mechanical hardness and abrasiveness on the wearing down of the teeth. MATERIALS AND METHODS: Percentages of dentine exposure (PDEs) of permanent left mandibular first molars (M1 ) were recorded using in vivo high-resolution replicas of Baka individuals (aged 8-33 years), inhabiting Le Bosquet district in Cameroon (Western Africa). Regression and covariance analyses were used to test the effect of individual aging by sex on PDE rates. RESULTS: We found a strong increase of PDE by age among Baka individuals. No evidence of sexual dimorphism in wear patterns suggests similar sex-related dietary and masticatory demands during growth. Overall, greatest dentine exposure values ≈4% denote unexpected slow wear down rates for foraging diets relying on USO consumption. DISCUSSION: The low molar wear rates with age found in Baka Pygmies contrast with extensive wear rates in savanna-dwelling foragers, reflecting differences in thermal processing techniques affecting fracture toughness and grittiness of mechanically challenging foods. Our findings reveal that culture-specific dietary proclivities influence tooth wear among foraging behaviors with important implications in hominin dietary versatility and abrasive stress on chewing surfaces.

Diet and adult age-at-death among mobile foragers: A synthesis of bioarcheological methods.

OBJECTIVES: The research explores whether the combined study of cortical bone histology, bone morphology, and dietary stable isotopes can expand insights into past human health and adaptations, particularly dietary sufficiency and life span. MATERIALS AND METHODS: Midthoracic rib cortices from 54 South African Late Holocene adult skeletons (28 M, 24 F, two sex undetermined) are assessed by transmitted-light microscopy for cross-sectional area measurements, osteon area (On.Ar), osteon population density, and presence/absence of secondary osteon variants. Values for δ13 Cbone collagen , δ15 Nbone collagen , 14 C dates, Southwestern and Southern Cape geographic regions, body size measures, estimated ages-at-death from both morphological and histological methods are integrated into analyses, which include Spearman correlations, χ2 tests and Kruskal-Wallis ANOVAs. RESULTS: There is reduced On.Ar variability with higher δ15 N (r = -.41, p = .005); rib %cortical area and δ15 N are negatively correlated in the Southern Cape group (r = -.60, p = .03). Osteon variants are more common in older adults; histological ages at death are significantly older than those determined from gross morphology. DISCUSSION: We found bone tissue relationships with measures of diet composition, but indicators of dietary adequacy remain elusive. Relationships of tissue quality and isotopes suggest that some Southern Cape adults lived long lives. Osteon variants are associated with age-at-death; some association with diet remains possible. Gross morphological methods appear to underestimate adult ages-at-death, at least among small-bodied adults.

Dental calculus reveals Mesolithic foragers in the Balkans consumed domesticated plant foods.

Researchers agree that domesticated plants were introduced into southeast Europe from southwest Asia as a part of a Neolithic "package," which included domesticated animals and artifacts typical of farming communities. It is commonly believed that this package reached inland areas of the Balkans by ∼6200 calibrated (cal.) BC or later. Our analysis of the starch record entrapped in dental calculus of Mesolithic human teeth at the site of Vlasac in the Danube Gorges of the central Balkans provides direct evidence that already by ∼6600 cal. BC, if not earlier, Late Mesolithic foragers of this region consumed domestic cereals, such as Triticum monococcum, Triticum dicoccum, and Hordeum distichon, which were also the main crops found among Early Neolithic communities of southeast Europe. We infer that "exotic" Neolithic domesticated plants were introduced to southern Europe independently almost half a millennium earlier than previously thought, through networks that enabled exchanges between inland Mesolithic foragers and early farming groups found along the Aegean coast of Turkey.

A Markov model for planning and permitting offshore wind energy: A case study of radio-tracked terns in the Gulf of Maine, USA.

Quantifying and managing the potential adverse wildlife impacts of offshore wind energy is critical for developing offshore wind energy in a sustainable and timely manner, but poses a significant challenge, particularly for small marine birds that are difficult to monitor. We developed a discrete-time Markov model of seabird movement around a colony site parameterized by automated radio telemetry data from common terns (Sterna hirundo) and Arctic terns (S. paradisaea), and derived impact functions that estimate the probability of collision fatality as a function of the distance and bearing of wind turbines from a colony. Our purpose was to develop and demonstrate a new, flexible tool that can be used for specific management and wind-energy planning applications when adequate data are available, rather than inform wind-energy development at this site. We demonstrate how the tool can be used 1) in marine spatial planning exercises to quantitatively identify setback distances under development scenarios given a risk threshold, 2) to examine the ecological and technical trade-offs of development alternatives to facilitate negotiation between objectives, and 3) in the U.S. National Environmental Policy Act (NEPA) process to estimate collision fatality under alternative scenarios. We discuss model limitations and data needs, and highlight opportunities for future model extension and development. We present a highly flexible tool for wind energy planning that can be easily extended to other central place foragers and data sources, and can be updated and improved as new monitoring data arises.

Skill learning and the evolution of social learning mechanisms.

BACKGROUND: Social learning is potentially advantageous, but evolutionary theory predicts that (i) its benefits may be self-limiting because social learning can lead to information parasitism, and (ii) these limitations can be mitigated via forms of selective copying. However, these findings arise from a functional approach in which learning mechanisms are not specified, and which assumes that social learning avoids the costs of asocial learning but does not produce information about the environment. Whether these findings generalize to all kinds of social learning remains to be established. Using a detailed multi-scale evolutionary model, we investigate the payoffs and information production processes of specific social learning mechanisms (including local enhancement, stimulus enhancement and observational learning) and their evolutionary consequences in the context of skill learning in foraging groups. RESULTS: We find that local enhancement does not benefit foraging success, but could evolve as a side-effect of grouping. In contrast, stimulus enhancement and observational learning can be beneficial across a wide range of environmental conditions because they generate opportunities for new learning outcomes. CONCLUSIONS: In contrast to much existing theory, we find that the functional outcomes of social learning are mechanism specific. Social learning nearly always produces information about the environment, and does not always avoid the costs of asocial learning or support information parasitism. Our study supports work emphasizing the value of incorporating mechanistic detail in functional analyses.