Stewardship of global collective behavior.

Collective behavior provides a framework for understanding how the actions and properties of groups emerge from the way individuals generate and share information. In humans, information flows were initially shaped by natural selection yet are increasingly structured by emerging communication technologies. Our larger, more complex social networks now transfer high-fidelity information over vast distances at low cost. The digital age and the rise of social media have accelerated changes to our social systems, with poorly understood functional consequences. This gap in our knowledge represents a principal challenge to scientific progress, democracy, and actions to address global crises. We argue that the study of collective behavior must rise to a "crisis discipline" just as medicine, conservation, and climate science have, with a focus on providing actionable insight to policymakers and regulators for the stewardship of social systems.

Large-herbivore nemabiomes: patterns of parasite diversity and sharing.

Amidst global shifts in the distribution and abundance of wildlife and livestock, we have only a rudimentary understanding of ungulate parasite communities and parasite-sharing patterns. We used qPCR and DNA metabarcoding of fecal samples to characterize gastrointestinal nematode (Strongylida) community composition and sharing among 17 sympatric species of wild and domestic large mammalian herbivore in central Kenya. We tested a suite of hypothesis-driven predictions about the role of host traits and phylogenetic relatedness in describing parasite infections. Host species identity explained 27-53% of individual variation in parasite prevalence, richness, community composition and phylogenetic diversity. Host and parasite phylogenies were congruent, host gut morphology predicted parasite community composition and prevalence, and hosts with low evolutionary distinctiveness were centrally positioned in the parasite-sharing network. We found no evidence that host body size, social-group size or feeding height were correlated with parasite composition. Our results highlight the interwoven evolutionary and ecological histories of large herbivores and their gastrointestinal nematodes and suggest that host identity, phylogeny and gut architecture-a phylogenetically conserved trait related to parasite habitat-are the overriding influences on parasite communities. These findings have implications for wildlife management and conservation as wild herbivores are increasingly replaced by livestock.

Macronutrient balancing affects patch departure by guerezas (Colobus guereza).

Foraging strategies are central in shaping social structure and grouping patterns in primates. We address Colobus guereza foraging strategies by investigating their patch departure decisions in relation to diet composition and nutrition. We examine whether guerezas are constrained in their intake of food in patches and thereby forage according to a fixed amount strategy that dictates patch departure. Additionally, we assess whether guereza employ a fixed time strategy or attempt to balance nutrients when foraging. We measured food patch occupancy time, intake rates, and analyzed foods for macronutrients, fiber, and condensed tannins. We determined that guerezas do not employ a fixed time foraging strategy; patch residence time varied widely between 1 and 290 min. They also did not depart patches or stop eating when they reached a specific intake of dry mass, macronutrients, or condensed tannins. However, guerezas maintained a macronutrient balance when feeding across patches, and the balance of protein to non-protein energy (fats and carbohydrates) in patches is the best indicator of time adult guerezas spent feeding in patches. Previous studies have shown that the protein-to-fiber ratio is important in predicting food selection for folivores and their biomass; however, we found that guerezas did not maximize protein and minimize fiber intake while foraging in patches, nor did they stay longer in patches with the highest ratio of protein to fiber concentrations. This study raises questions about the nutritional and social implications of patch depletion as a foraging strategy in folivorous monkeys where food limitation predicts competitive and social regimes. Am. J. Primatol. 79:e22495, 2017. © 2015 Wiley Periodicals, Inc.

New estimates indicate that males are not larger than females in most mammal species.

Sexual size dimorphism has motivated a large body of research on mammalian mating strategies and sexual selection. Despite some contrary evidence, the narrative that larger males are the norm in mammals-upheld since Darwin's Descent of Man-still dominates today, supported by meta-analyses that use coarse measures of dimorphism and taxonomically-biased sampling. With newly-available datasets and primary sources reporting sex-segregated means and variances in adult body mass, we estimate statistically-determined rates of sexual size dimorphism in mammals, sampling taxa by their species richness at the family level. Our analyses of wild, non-provisioned populations representing >400 species indicate that although males tend to be larger than females when dimorphism occurs, males are not larger in most mammal species, suggesting a need to revisit other assumptions in sexual selection research.

Zebras of all stripes repel biting flies at close range.

The best-supported hypothesis for why zebras have stripes is that stripes repel biting flies. While this effect is well-established, the mechanism behind it remains elusive. Myriad hypotheses have been suggested, but few experiments have helped narrow the field of possible explanations. In addition, the complex visual features of real zebra pelage and the natural range of stripe widths have been largely left out of experimental designs. In paired-choice field experiments in a Kenyan savannah, we found that hungry Stomoxys flies released in an enclosure strongly preferred to land on uniform tan impala pelts over striped zebra pelts but exhibited no preference between the pelts of the zebra species with the widest stripes and the narrowest stripes. Our findings confirm that zebra stripes repel biting flies under naturalistic conditions and do so at close range (suggesting that several of the mechanisms hypothesized to operate at a distance are unnecessary for the fly-repulsion effect) but indicate that interspecific variation in stripe width is associated with selection pressures other than biting flies.

Divergent water requirements partition exposure risk to parasites in wild equids.

For grazing herbivores, dung density in feeding areas is an important determinant of exposure risk to fecal-orally transmitted parasites. When host species share the same parasite species, a nonrandom distribution of their cumulative dung density and/or nonrandom ranging and feeding behavior may skew exposure risk and the relative selection pressure parasites impose on each host. The arid-adapted Grevy's zebra (Equus grevyi) can range more widely than the water-dependent plains zebra (Equus quagga), with which it shares the same species of gastrointestinal nematodes. We studied how the spatial distribution of zebra dung relates to ranging and feeding behavior to assess parasite exposure risk in Grevy's and plains zebras at a site inhabited by both zebra species. We found that zebra dung density declined with distance from water, Grevy's zebra home ranges (excluding those of territorial males) were farther from water than those of plains zebras, and plains zebra grazing areas had higher dung density than random points while Grevy's zebra grazing areas did not, suggesting a greater exposure risk in plains zebras associated with their water dependence. Fecal egg counts increased with home range proximity to water for both species, but the response was stronger in plains zebras, indicating that this host species may be particularly vulnerable to the elevated exposure risk close to water. We further ran experiments on microclimatic effects on dung infectivity and showed that fewer nematode eggs embryonated in dung in the sun than in the shade. However, only 5% of the zebra dung on the landscape was in shade, indicating that the microclimatic effects of shade on the density of infective larvae is not a major influence on exposure risk dynamics. Ranging constraints based on water requirements appear to be key mediators of nematode parasite exposure in free-ranging equids.

The gastrointestinal nematodes of plains and Grevy's zebras: Phylogenetic relationships and host specificity.

Equids are chronically infected with parasitic strongyle nematodes. There is a rich literature on horse strongyles, but they are difficult to identify morphologically and genetic studies on strongyles infecting other equid species are few, hampering studies of host specificity. We sequenced expelled worms from two sympatric zebra species in central Kenya to expand the strongyle phylogeny and used DNA metabarcoding on faecal samples to genetically characterize zebra nemabiomes for the first time. We generated sequences for several species new to public genetic reference databases, all of which are typical strongyles in wild zebras (i.e., the three species of Cylindropharynx and Cyathostomum montgomeryi), and identified their closest relatives. We also discovered an apparent fungus infecting a quarter of the expelled Crossocephalus viviparus worms, a hyperabundant nematode species in the family Atractidae, hinting at the possibility that zebra host-parasite dynamics may involve a zebra-fungus mutualism. The two zebra species had similar nemabiomes; we found a complete overlap in the list of nematode species they carry and very similar prevalence (i.e., proportion of hosts infected) for the different nematode species. Our study suggests limited host-specificity in zebra strongyles and high potential for transmission between the plains zebra and the endangered Grevy's zebra.

The non-invasive measurement of faecal immunoglobulin in African equids.

Eco-immunological research is encumbered by a lack of basic research in a wild context and by the availability of few non-invasive tools to measure the internal state of wild animals. The recent development of an enzyme-linked immunosorbent assay for measuring immunoglobulins in faecal samples from Soay sheep prompted us to optimize such an assay to measure immunoglobulin A (IgA: an antibody associated with parasitic nematode fecundity) in faecal samples from equids. We measured total IgA in domestic donkeys, wild plains zebras, and wild Grevy's zebras sharing the same landscape in central Kenya over two field seasons. Attempts to measure anti-nematode IgA more specifically, using a homogenized extract from a mixture of excreted nematodes, failed to clear background. However, we found that total IgA positively correlated with strongyle nematode faecal egg counts (FECs) in donkeys sampled during the wetter field season - a time when the donkeys were in good condition. Further, this relationship appeared among donkeys with high body condition but not among those with low body condition. Time lags of 1-4 days introduced between IgA and FEC measurements in repeatedly sampled donkeys did not yield correlations, suggesting that IgA and FEC roughly tracked one another without much delay in the wet field season. Such a direct IgA-FEC relationship did not appear for zebras in either the wet or dry field season, possibly due to higher interindividual variation in body condition among the free-roaming zebras than in the donkeys. However, Grevy's zebras had higher overall levels of IgA than either plains zebras or donkeys, potentially associated with their reportedly lower FECs at the population level. Our results suggest that equids may mount an IgA response to nematode egg production when the host is in good condition and that equid species may differ in baseline levels of mucosal IgA.

Patch depletion behavior differs between sympatric folivorous primates.

Food competition in group-living animals is commonly accepted as a critical determinant of foraging strategies and social organization. Here we examine food patch depletion behavior in a leaf-eating (folivorous) primate, the guereza (Colobus guereza). Snaith and Chapman (2005) studied the sympatric folivorous red colobus (Procolobus rufomitratus), which shares many food resources with the guereza. They determined that red colobus deplete the patches (feeding trees) they use, while we found contrary evidence for guerezas using the same methods. We found that the time guerezas spent feeding in a patch was affected by neither tree size, an indicator of food abundance, nor the size of the feeding group, an indicator of feeding competition. For their principal food item (young leaves), intake rate remained constant and coincided with a decrease in the distance moved to find food within a patch, implying that guerezas do not deplete patches. This points to a fundamental difference in the use of food by guerezas and red colobus, which may be linked to the large difference in their group sizes and/or to a disparity in their digestive physiologies. However, further analyses revealed that the number of feeders within a patch did not affect patch depletion patterns in either species, leaving the potential for a physiological basis as the most plausible explanation. Our research highlights the need for a more critical examination of folivorous primate feeding ecology and social behavior, as all folivorous primates are typically lumped into a single category in socioecological models, which may account for conflicting evidence in the literature.