Bold coloration and the evolution of aposematism in terrestrial carnivores

Several species of terrestrial carnivores (Mammalia: Carnivora) have bold contrasting color patterns that, in some species, apparently signal possession of noxious anal gland secretions, or even physical strength and great ferocity; yet the evolutionary drivers of both placement and patterning of these contrasting pelage colors on the body, and the ecological selection pressures underlying them, have yet to be systematically examined. Here we explore these issues and find not only that both boldly colored and dichromatic species do indeed often use anal gland secretions for defense, but also that such species are stockier, and live in more exposed habitats where other forms of antipredator defense are limited. We also show that white dorsa are found in sprayers that are primarily nocturnal; that horizontal stripes are found in species that have an ability to spray anal secretions accurately; and that facial stripes are found in burrowing species that typically leave only their heads exposed to attack. Our phylogenetic reconstructions suggest that aposematic coloration has evolved more than once in terrestrial carnivores. We finish by outlining five evolutionary routes for patterns of pelage coloration in this taxon.     

Roosting ecology and the evolution of pelage markings in bats

Multiple lineages of bats have evolved striking facial and body pelage makings, including spots, stripes and countershading. Although researchers have hypothesized that these markings mainly evolved for crypsis, this idea has never been tested in a quantitative and comparative context. We present the first comparative study integrating data on roosting ecology (roost type and colony size) and pelage coloration patterns across bats, and explore the hypothesis that the evolution of bat pelage markings is associated with roosting ecologies that benefit from crypsis. We find that lineages that roost in the vegetation have evolved pelage markings, especially stripes and neck collars, which may function in crypsis through disruptive coloration and a type of countershading that might be unique to bats. We also demonstrate that lineages that live in larger colonies and are larger in size tend not to have pelage markings, possibly because of reduced predation pressures due to the predator dilution effect and a lower number of potential predators. Although social functions for pelage color patterns are also possible, our work provides strong support for the idea that roosting ecology has driven the evolution of pelage markings in bats.     

The effect of competing carnivores on the feeding behaviour of leopards (Panthera pardus) in an African savanna

Knowledge of competition dynamics among Africa’s large carnivores is important for conservation. However, investigating carnivore behaviour in the field can be challenging especially for species that are difficult to access. Methods that enable remote collection of data provide a means of recording natural behaviour and are therefore useful for studying elusive species such as leopards (Panthera pardus). Camera traps and Global Positioning System (GPS) collars are powerful tools often used independently to study animal behaviour but where their data are combined, the interpretation of a species’ behaviours is improved. In this study we used data from baited camera trap stations to investigate the feeding habits of leopards at Malilangwe Wildlife Reserve, Zimbabwe. We investigated the influence of spotted hyenas, lions and other competing leopards on the feeding duration of leopards using Generalized Linear Mixed Effects Modelling. To test the influence of competing predators on resting distances from bait sites, eight leopards were fitted with GPS collars. Results showed that leopards spent the shortest time feeding on the baits in the presence of competing male leopards compared to other predators while lion presence caused animals to rest farthest from bait sites. Interaction analysis indicated that small‐bodied leopards spent significantly shorter durations feeding when spotted hyenas were present. Our findings demonstrate that competition from guild carnivores has negative impacts on the food intake of leopards, which may have implications for fitness and survival. This study provides a snapshot of the competition dynamics at bait sites which may give insight to ecosystem level interactions among large carnivores in savanna ecosystems.     

Mark-Recapture and Mark-Resight Methods for Estimating Abundance with Remote Cameras: A Carnivore Case Study

Abundance estimation of carnivore populations is difficult and has prompted the use of non-invasive detection methods, such as remotely-triggered cameras, to collect data. To analyze photo data, studies focusing on carnivores with unique pelage patterns have utilized a mark-recapture framework and studies of carnivores without unique pelage patterns have used a mark-resight framework. We compared mark-resight and mark-recapture estimation methods to estimate bobcat (Lynx rufus) population sizes, which motivated the development of a new "hybrid" mark-resight model as an alternative to traditional methods. We deployed a sampling grid of 30 cameras throughout the urban southern California study area. Additionally, we physically captured and marked a subset of the bobcat population with GPS telemetry collars. Since we could identify individual bobcats with photos of unique pelage patterns and a subset of the population was physically marked, we were able to use traditional mark-recapture and mark-resight methods, as well as the new “hybrid” mark-resight model we developed to estimate bobcat abundance. We recorded 109 bobcat photos during 4,669 camera nights and physically marked 27 bobcats with GPS telemetry collars. Abundance estimates produced by the traditional mark-recapture, traditional mark-resight, and “hybrid” mark-resight methods were similar, however precision differed depending on the models used. Traditional mark-recapture and mark-resight estimates were relatively imprecise with percent confidence interval lengths exceeding 100% of point estimates. Hybrid mark-resight models produced better precision with percent confidence intervals not exceeding 57%. The increased precision of the hybrid mark-resight method stems from utilizing the complete encounter histories of physically marked individuals (including those never detected by a camera trap) and the encounter histories of naturally marked individuals detected at camera traps. This new estimator may be particularly useful for estimating abundance of uniquely identifiable species that are difficult to sample using camera traps alone.     

Effects of Single‐ and Mixed‐Species Group Composition on the Flight Initiation Distances of Plains and Grevy's Zebras

Zebras, as prey species, attend to the behavior of nearby conspecifics and heterospecifics when making decisions to flee from predators. Plains zebras (Equus quagga) and Grevy's zebras (E. grevyi) frequently form mixed‐species groups in zones where their ranges overlap in Kenya. Although anecdotal observations suggest that Plains zebras are more flighty around humans than Grevy's zebras are, this has not been empirically confirmed, and relatively little is known about how they may influence each other's flight behavior. We addressed these questions by examining the flight initiation distances (FIDs) of Plains and Grevy's zebras in single‐species and mixed‐species groups from an approaching human. One target individual per group was approached steadily on foot, with start distance, alert distance, and FID recorded from this target. Using start distance and alert distance separately as covariates, 22 Plains zebras in single‐species groups exhibited a significantly longer mean FID than 15 Grevy's zebras in single‐species groups. The FIDs of 7 Plains zebras and 5 Grevy's zebras tested in mixed‐species groups were virtually equivalent and intermediate to those of Plains and Grevy's zebras in single‐species groups, suggesting a bidirectional moderating influence of heterospecifics on risk assessment. This effect was most pronounced for Plains zebras in mixed‐species groups that exhibited an FID that was significantly shorter than that of Plains zebras in single‐species groups. Our findings underscore the importance of recognizing that related equids may be differently impacted by anthropogenic stress.     

Similar but Different: Dynamic Social Network Analysis Highlights Fundamental Differences between the Fission-Fusion Societies of Two Equid Species,the Onager and Grevy’s Zebra

Understanding why animal societies take on the form that they do has benefited from insights gained by applying social network analysis to patterns of individual associations. Such analyses typically aggregate data over long time periods even though most selective forces that shape sociality have strong temporal elements. By explicitly incorporating the temporal signal in social interaction data we re-examine the network dynamics of the social systems of the evolutionarily closely-related Grevy’s zebras and wild asses that show broadly similar social organizations. By identifying dynamic communities, previously hidden differences emerge: Grevy’s zebras show more modularity than wild asses and in wild asses most communities consist of solitary individuals; and in Grevy’s zebras, lactating females show a greater propensity to switch communities than non-lactating females and males. Both patterns were missed by static network analyses and in general, adding a temporal dimension provides insights into differences associated with the size and persistence of communities as well as the frequency and synchrony of their formation. Dynamic network analysis provides insights into the functional significance of these social differences and highlights the way dynamic community analysis can be applied to other species.     

Motion camouflage induced by zebra stripes

The functional significance of the zebra coat stripe pattern is one of the oldest questions in evolutionary biology, having troubled scientists ever since Charles Darwin and Alfred Russel Wallace first disagreed on the subject. While different theories have been put forward to address this question, the idea that the stripes act to confuse or ‘dazzle’ observers remains one of the most plausible. However, the specific mechanisms by which this may operate have not been investigated in detail. In this paper, we investigate how motion of the zebra's high contrast stripes creates visual effects that may act as a form of motion camouflage. We simulated a biologically motivated motion detection algorithm to analyse motion signals generated by different areas on a zebra's body during displacements of their retinal images. Our simulations demonstrate that the motion signals that these coat patterns generate could be a highly misleading source of information. We suggest that the observer's visual system is flooded with erroneous motion signals that correspond to two well-known visual illusions: (i) the wagon-wheel effect (perceived motion inversion due to spatiotemporal aliasing); and (ii) the barber-pole illusion (misperceived direction of motion due to the aperture effect), and predict that these two illusory effects act together to confuse biting insects approaching from the air, or possibly mammalian predators during the hunt, particularly when two or more zebras are observed moving together as a herd.     

Fear of humans as apex predators has landscape‐scale impacts from mountain lions to mice

Apex predators such as large carnivores can have cascading, landscape‐scale impacts across wildlife communities, which could result largely from the fear they inspire, although this has yet to be experimentally demonstrated. Humans have supplanted large carnivores as apex predators in many systems, and similarly pervasive impacts may now result from fear of the human ‘super predator’. We conducted a landscape‐scale playback experiment demonstrating that the sound of humans speaking generates a landscape of fear with pervasive effects across wildlife communities. Large carnivores avoided human voices and moved more cautiously when hearing humans, while medium‐sized carnivores became more elusive and reduced foraging. Small mammals evidently benefited, increasing habitat use and foraging. Thus, just the sound of a predator can have landscape‐scale effects at multiple trophic levels. Our results indicate that many of the globally observed impacts on wildlife attributed to anthropogenic activity may be explained by fear of humans.     

Inferring Predator Behavior from Attack Rates on Prey-Replicas That Differ in Conspicuousness

Behavioral ecologists and evolutionary biologists have long studied how predators respond to prey items novel in color and pattern. Because a predatory response is influenced by both the predator’s ability to detect the prey and a post-detection behavioral response, variation among prey types in conspicuousness may confound inference about post-prey-detection predator behavior. That is, a relatively high attack rate on a given prey type may result primarily from enhanced conspicuousness and not predators’ direct preference for that prey. Few studies, however, account for such variation in conspicuousness. In a field experiment, we measured predation rates on clay replicas of two aposematic forms of the poison dart frog Dendrobates pumilio, one novel and one familiar, and two cryptic controls. To ask whether predators prefer or avoid a novel aposematic prey form independently of conspicuousness differences among replicas, we first modeled the visual system of a typical avian predator. Then, we used this model to estimate replica contrast against a leaf litter background to test whether variation in contrast alone could explain variation in predator attack rate. We found that absolute predation rates did not differ among color forms. Predation rates relative to conspicuousness did, however, deviate significantly from expectation, suggesting that predators do make post-detection decisions to avoid or attack a given prey type. The direction of this deviation from expectation, though, depended on assumptions we made about how avian predators discriminate objects from the visual background. Our results show that it is important to account for prey conspicuousness when investigating predator behavior and also that existing models of predator visual systems need to be refined.     

Bottom-up and top-down processes in African ungulate communities: resources and predation acting on the relative abundance of zebra and grazing bovids

African ungulate populations appear to be limited principally by their food resources. Within ungulate communities, plains zebras coexist with grazing bovids of similar body size, but rarely are the dominant species. Given the highly effective nutritional strategy of the equids and the resistance of zebras to drought, this is unexpected and suggests that zebra populations may commonly be limited by other mechanisms. Long-term research in the Serengeti ecosystem and in the Kruger National Park suggests that zebra could be less sensitive to food shortage, and more sensitive to predation, than grazing bovids: if this is a general principle, then, at a larger scale, resource availability should have a weaker effect on the abundance of zebra than on grazing ruminants of similar body size (wildebeest and buffalo), and zebras should be relatively more abundant in ecosystems where predators are rare or absent. We test these expectations using data on 23 near-natural ecosystems in east and southern Africa. The abundance of wildebeest is more closely related to resources than is that of zebra; buffalo are intermediate. We show that hyena densities are closely correlated with those of lions, and use the abundance of lions as an index of predation by large predators. The numerical response of lions to increases in the abundance of their prey was linear for mesoherbivores, and apparently so for the three species alone. Finally, the abundance of zebra relative to grazing bovids is lower in ecosystems with high biomasses of lions. These results indicate that zebras may commonly be more sensitive to top-down processes than grazing bovids: the mechanism(s) have not been demonstrated, but predation could play a role. If it is true, then when numbers of the large mammalian predators decline, zebra populations should increase faster than buffalo and wildebeest.     

Landscapes of Coexistence for terrestrial carnivores: the ecological consequences of being downgraded from ultimate to penultimate predator by humans

Fear of predation can have major impacts on the behaviour of prey species. Recently the concept of the ecology of fear has been defined and formalised; yet there has been relatively little focus on how these ideas apply to large carnivore species which, although not prey sensu stricto, also experience fear as a result of threats from humans. Large carnivores are likely also subject to a Landscape of Fear similar to that described for prey species. We argue that although fear is generic, ‘human‐caused mortality’ represents a distinct and very important cause of fear for large carnivores, particularly terrestrial large carnivores as their activities overlap with those of humans to a greater degree. We introduce the idea of a ‘Landscape of Coexistence’ for large carnivores to denote a subset of the Landscape of Fear where sufficient areas of low human‐caused mortality risk are present in the landscape for long term coexistence of large carnivores and humans. We then explore aspects of terrestrial large carnivore behavioural ecology that may be best explained by risk of human‐caused mortality, and how the nature of a Landscape of Coexistence for these large carnivores is likely to be shaped by specific factors such as habitat structure, wild and domestic prey base, and human distribution and behaviour. The human characteristics of this Landscape of Coexistence may be as important in determining large carnivore distribution and behavioural ecology as the distribution of resources. Understanding the Landscape of Coexistence for terrestrial large carnivores is therefore important for their biology and conservation throughout large parts of their remaining ranges. Synthesis The Landscape of Fear concept describing the relationship between predator and prey also applies to the relationship between humans and top carnivores. We synthesise current research to introduce the Landscape of Coexistence concept, arguing that top predators respond to the risks of human‐caused mortality through spatiotemporal partitioning of activities to reduce contact with people. The character of the Landscape of Coexistence may be more important than the distribution of resources in determining large carnivore distribution and behavioural ecology in human dominated landscapes. Understanding their behavioural responses to human threats is crucial to successful conservation of large carnivores.     

Fear, human shields and the redistribution of prey and predators in protected areas

Protected areas form crucial baselines to judge ecological change, yet areas of Africa, Asia and North America that retain large carnivores are under intense economic and political pressures to accommodate massive human visitation and attendant infrastructure. An unintended consequence is the strong modulation of the three-way interaction involving people, predators and prey, a dynamic that questions the extent to which animal distributions and interactions are independent of subtle human influences. Here, I capitalize on the remarkable 9-day synchronicity in which 90% of moose neonates in the Yellowstone Ecosystem are born, to demonstrate a substantive change in how prey avoid predators; birth sites shift away from traffic-averse brown bears and towards paved roads. The decade-long modification was associated with carnivore recolonization, but neither mothers in bear-free areas nor non-parous females altered patterns of landscape use. These findings offer rigorous support that mammals use humans to shield against carnivores and raise the possibility that redistribution has occurred in other mammalian taxa due to human presence in ways we have yet to anticipate. To interpret ecologically functioning systems within parks, we must now also account for indirect anthropogenic effects on species distributions and behaviour.     

The dominantly expressed class II molecule from a resistant MHC haplotype presents only a few Marek’s disease virus peptides by using an unprecedented binding motif

Viral diseases pose major threats to humans and other animals, including the billions of chickens that are an important food source as well as a public health concern due to zoonotic pathogens. Unlike humans and other typical mammals, the major histocompatibility complex (MHC) of chickens can confer decisive resistance or susceptibility to many viral diseases. An iconic example is Marek’s disease, caused by an oncogenic herpesvirus with over 100 genes. Classical MHC class I and class II molecules present antigenic peptides to T lymphocytes, and it has been hard to understand how such MHC molecules could be involved in susceptibility to Marek’s disease, given the potential number of peptides from over 100 genes. We used a new in vitro infection system and immunopeptidomics to determine peptide motifs for the 2 class II molecules expressed by the MHC haplotype B2, which is known to confer resistance to Marek’s disease. Surprisingly, we found that the vast majority of viral peptide epitopes presented by chicken class II molecules arise from only 4 viral genes, nearly all having the peptide motif for BL2*02, the dominantly expressed class II molecule in chickens. We expressed BL2*02 linked to several Marek’s disease virus (MDV) peptides and determined one X-ray crystal structure, showing how a single small amino acid in the binding site causes a crinkle in the peptide, leading to a core binding peptide of 10 amino acids, compared to the 9 amino acids in all other reported class II molecules. The limited number of potential T cell epitopes from such a complex virus can explain the differential MHC-determined resistance to MDV, but raises questions of mechanism and opportunities for vaccine targets in this important food species, as well as providing a basis for understanding class II molecules in other species including humans.

This study shows that chicken MHC class II molecules present peptides from only a handful of the more than 100 genes of the oncogenic herpesvirus Marek’s disease virus, explaining the strong genetic association of chicken MHC with resistance and susceptibility to this and other economically-important pathogens.     

What aspects of autism predispose to talent?

In this paper, we explore the question, why are striking special skills so much more common in autism spectrum conditions (ASC) than in other groups? Current cognitive accounts of ASC are briefly reviewed in relation to special skills. Difficulties in ‘theory of mind’ may contribute to originality in ASC, since individuals who do not automatically ‘read other minds’ may be better able to think outside prevailing fashions and popular theories. However, originality alone does not confer talent. Executive dysfunction has been suggested as the ‘releasing’ mechanism for special skills in ASC, but other groups with executive difficulties do not show raised incidence of talents. Detail-focused processing bias (‘weak coherence’, ‘enhanced perceptual functioning’) appears to be the most promising predisposing characteristic, or ‘starting engine’, for talent development. In support of this notion, we summarize data from a population-based twin study in which parents reported on their 8-year-olds'' talents and their ASC-like traits. Across the whole sample, ASC-like traits, and specifically ‘restricted and repetitive behaviours and interests’ related to detail focus, were more pronounced in children reported to have talents outstripping older children. We suggest that detail-focused cognitive style predisposes to talent in savant domains in, and beyond, autism spectrum disorders.     

Cry-wolf signals emerging from coevolutionary feedbacks in a tritrophic system

For a communication system to be stable, senders should convey honest information. Providing dishonest information, however, can be advantageous to senders, which imposes a constraint on the evolution of communication systems. Beyond single populations and bitrophic systems, one may ask whether stable communication systems can evolve in multitrophic systems. Consider cross-species signalling where herbivore-induced plant volatiles (HIPVs) attract predators to reduce the damage from arthropod herbivores. Such plant signals may be honest and help predators to identify profitable prey/plant types via HIPV composition and to assess prey density via the amount of HIPVs. There could be selection for dishonest signals that attract predators for protection from possible future herbivory. Recently, we described a case in which plants release a fixed, high amount of HIPVs independent of herbivore load, adopting what we labelled a ‘cry-wolf’ strategy. To understand when such signals evolve, we model coevolutionary interactions between plants, herbivores and predators, and show that both ‘honest’ and ‘cry-wolf’ types can emerge, depending on the assumed plant–herbivore encounter rates and herbivore population density. It is suggested that the ‘cry-wolf’ strategy may have evolved to reduce the risk of heavy damage in the future. Our model suggests that eco-evolutionary feedback loops involving a third species may have important consequences for the stability of this outcome.     

Pigeons (Columba livia) as Trainable Observers of Pathology and Radiology Breast Cancer Images

Pathologists and radiologists spend years acquiring and refining their medically essential visual skills, so it is of considerable interest to understand how this process actually unfolds and what image features and properties are critical for accurate diagnostic performance. Key insights into human behavioral tasks can often be obtained by using appropriate animal models. We report here that pigeons (Columba livia)—which share many visual system properties with humans—can serve as promising surrogate observers of medical images, a capability not previously documented. The birds proved to have a remarkable ability to distinguish benign from malignant human breast histopathology after training with differential food reinforcement; even more importantly, the pigeons were able to generalize what they had learned when confronted with novel image sets. The birds’ histological accuracy, like that of humans, was modestly affected by the presence or absence of color as well as by degrees of image compression, but these impacts could be ameliorated with further training. Turning to radiology, the birds proved to be similarly capable of detecting cancer-relevant microcalcifications on mammogram images. However, when given a different (and for humans quite difficult) task—namely, classification of suspicious mammographic densities (masses)—the pigeons proved to be capable only of image memorization and were unable to successfully generalize when shown novel examples. The birds’ successes and difficulties suggest that pigeons are well-suited to help us better understand human medical image perception, and may also prove useful in performance assessment and development of medical imaging hardware, image processing, and image analysis tools.     

A model for generating aspects of zebra and other mammalian coat patterns

A model is put forward which is capable of generating chemical maps whose concentration contours are similar to the patterns seen on the flanks of zebras, cats and other mammals. The model derives from the reaction-diffusion kinetics invented by Turing (1952) and it is assumed that the necessary molecular apparatus is present in each cell of a two-dimensional array and that the cells are in diffusion contact. The model was expressed in differential equation form and solved digitally under a range of different initial, boundary and other conditions. The main forms of pattern that the model generated were spots of variable complexity, rings, and both vertical and horizontal stripes. If morphogen concentration levels are assumed to act as melanin-production switches, then a common basic mechanism is capable of generating a variety of skin patterns. Simple spots such as those found in the fallow deer or the serval, F. serval, are generated if the kinetics are initiated simultaneously in each cell and interpretation depends only on the presence or absence of morphogen, which is assumed for the deer to be an activator and for the cat a suppressor of pigment formation. The reticulated pattern of the giraffe is generated if there is a single high-value threshold. Complex spots typical of the leopards can be produced if there are different concentration thresholds for different colours. Rings of pattern typical of those found on cat tails are generated if the cellular array is a very narrow cylinder. Horizontal stripes are generated if the kinetics in each cell are initiated by a diffusion gradient whose source is the dorsal line of cells and these stripes may break up into spots to give a pattern very similar to that of, for example, the fishing cat, F. viverina. The vertical stripes of the caffre cat, F. caffra, or the zebras are formed if the kinetics are initiated by a vertically-moving constant-velocity wave which also allows morphogen diffusion between previously uncoupled cells. Thus far, the mechanism has generated neither the triradii that are commonly found on forelimbs nor the rings often observed on mammalian limbs. It does however incorporate the randomness that characterizes skin pattern, its operation is of the scale required in embryogenesis, it can be made stable to growth and it can explain certain degenerate patterns. Analysis of a spotted zebra in the light of the model provides evidence that zebra stripes arise from the inhibition rather than the stimulation of melanin; their pattern is thus of white stripes on a black background.     

A novel camera trapping method for individually identifying pumas by facial features

Camera traps (CTs), used in conjunction with capture–mark–recapture analyses (CMR; photo‐CMR), are a valuable tool for estimating abundances of rare and elusive wildlife. However, a critical requirement of photo‐CMR is that individuals are identifiable in CT images (photo‐ID). Thus, photo‐CMR is generally limited to species with conspicuous pelage patterns (e.g., stripes or spots) using lateral‐view images from CTs stationed along travel paths. Pumas (Puma concolor) are an elusive species for which CTs are highly effective at collecting image data, but their suitability to photo‐ID is controversial due to their lack of pelage markings. For a wide range of taxa, facial features are useful for photo‐ID, but this method has generally been limited to images collected with traditional handheld cameras. Here, we evaluate the feasibility of using puma facial features for photo‐ID in a CT framework. We consider two issues: (1) the ability to capture puma facial images using CTs, and (2) whether facial images improve human ability to photo‐ID pumas. We tested a novel CT accessory that used light and sound to attract the attention of pumas, thereby collecting face images for use in photo‐ID. Face captures rates increased at CTs that included the accessory (n = 208, χ ² = 43.23, p ≤ .001). To evaluate if puma faces improve photo‐ID, we measured the inter‐rater agreement of 5 independent assessments of photo‐ID for 16 of our puma face capture events. Agreement was moderate to good (Fleiss’ kappa = 0.54, 95% CI = 0.48–0.60), and was 92.90% greater than a previously published kappa using conventional CT methods. This study is the first time that such a technique has been used for photo‐ID, and we believe a promising demonstration of how photo‐ID may be feasible for an elusive but unmarked species.     

Colors of primate pelage and skin: objective assessment of conspicuousness

We present a quantitative means of assessing the conspicuousness of animal coats or other objects in terms of the color vision of each possible observer. We measured reflectance spectra from the fur and skin of many primate species in order to provide an objective survey of the possibilities of pelage coloration found in extant primates. We show that the orange coloration displayed by many platyrrhine and some strepsirhine primates, while being conspicuous to humans, would be cryptic amongst foliage to all males and many females of their own species. In relation to this finding, we briefly review what is known of the color vision of birds that prey on primates, and assess how conspicuous the orange pelage would be to these predators.