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Coconut crabs Birgus latro have recently been reclassified from Data Deficient to Vulnerable on the IUCN Red List. This is a somewhat unusual case of temporal and spatial information being used to change the IUCN status of an arthropod and it draws attention to the paucity of biological data on most invertebrate species. To be listed, two or more scientific criteria need to be documented but such data are unavailable for many invertebrates. This raises the question as to whether certain invertebrates receive more scientific attention and are hence more likely to be listed if, like the coconut crab, they are large, slow-reproducing or a dual-biome species (characteristics which make them inherently vulnerable) and whether being an indicator or a flagship species is important.

Stripes deter horseflies (tabanids) from landing on zebras and, while several mechanisms have been proposed, these hypotheses have yet to be tested satisfactorily. Here, we investigated three possible visual mechanisms that could impede successful tabanid landings (aliasing, contrast and polarization) but additionally explored pattern element size employing video footage of horseflies around differently patterned coats placed on domestic horses. We found that horseflies are averse to landing on highly but not on lightly contrasting stripes printed on horse coats. We could find no evidence for horseflies being attracted to coats that better reflected polarized light. Horseflies were somewhat less attracted to regular than to irregular check patterns, but this effect was not large enough to support the hypothesis of disrupting optic flow through aliasing. More likely it is due to attraction towards larger dark patches present in the irregular check patterns, an idea bolstered by comparing landings to the size of dark patterns present on the different coats. Our working hypothesis for the principal anti-parasite features of zebra pelage are that their stripes are sharply outlined and thin because these features specifically eliminate the occurrence of large monochrome dark patches that are highly attractive to horseflies at close distances.

Comparative studies indicate that several mammalian clades obey Gloger’s rule in that they exhibit darker coloration in humid warm climates, although the mechanisms responsible for this association still are poorly understood. We surveyed external appearances of a single species, the feral pig (Sus scrofa), shot at 48 hunting lodges across North America and matched these to potential abiotic drivers, namely: relative humidity, temperature, precipitation, and ultraviolet (UV) radiation, and to biotic factors of habitat shade and predation pressure. We found that darker animals occupy locations of greater precipitation and warmer temperatures, as expected from Gloger’s rule. The recent range expansion of S. scrofa implies selection for pelage coloration has occurred very rapidly. Separating pelage coloration into eumelanin- and phaeomelanin-based pigmentation, we found more pronounced eumelanin-based pelage in areas of higher rainfall and temperatures and UV radiation, whereas pelage phaeomelanin is related to cool dry climates with lower UV radiation. This implies that humidity or UV protection but not crypsis are the mechanisms underlying Gloger’s rule in this species and the factors driving eumelanin and phaeomelanin expression in mammalian pelage are different, reinforcing new interpretations of this venerable rule.

Primates are noted for their varied and complex pelage and bare skin coloration but the significance of this diverse coloration remains opaque. Using new updated information, novel scoring of coat and skin coloration, and controlling for shared ancestry, we reexamined and extended findings from previous studies across the whole order and the five major clades within it. Across primates, we found (i) direct and indirect evidence for pelage coloration being driven by protective coloration strategies including background matching, countershading, disruptive coloration, and aposematism, (ii) diurnal primates being more colorful, and (iii) the possibility that pelage color diversity is negatively associated with female trichromatic vision; while (iv) reaffirming avoidance of hybridization driving head coloration in males, (v) darker species living in warm, humid conditions (Gloger’s rule), and (vi) advertising to multiple mating partners favoring red genitalia in females. Nonetheless, the importance of these drivers varies greatly across clades. In strepsirrhines and cercopithecoids, countershading is important; greater color diversity may be important for conspecific signaling in more diurnal and social strepsirrhines; lack of female color vision may be associated with colorful strepsirrhines and platyrrhines; whereas cercopithecoids obey Gloger’s rule. Haplorrhines show background matching, aposematism, character displacement, and red female genitalia where several mating partners are available. Our findings emphasize several evolutionary drivers of coloration in this extraordinarily colorful order. Throughout, we used coarse but rigorous measures of coloration, and our ability to replicate findings from earlier studies opens up opportunities for classifying coloration of large numbers of species at a macroevolutionary scale.

Natural habitats are increasingly affected by anthropogenically driven environmental changes resulting from habitat destruction, chemical and light pollution, and climate change. Organisms inhabiting such habitats are faced with novel disturbances that can alter their modes of signaling. Coloration is one such sensory modality whose production, perception and function is being affected by human-induced disturbances. Animals that acquire pigment derivatives through diet are adversely impacted by the introduction of chemical pollutants into their environments as well as by general loss of natural habitat due to urbanization or logging leading to declines in pigment sources. Those species that do manage to produce color-based signals and displays may face disruptions to their signaling medium in the form of light pollution and turbidity. Furthermore, forest fragmentation and the resulting breaks in canopy cover can expose animals to predation due to the influx of light into previously dark environments. Global climate warming has been decreasing snow cover in arctic regions, causing birds and mammals that undergo seasonal molts to appear conspicuous against a snowless background. Ectotherms that rely on color for thermoregulation are under pressure to change their appearances. Rapid changes in habitat type through severe fire events or coral bleaching also challenge animals to match their backgrounds. Through this review, we aim to describe the wide-ranging impacts of anthropogenic environmental changes on visual ecology and suggest directions for the use of coloration both as an indicator of ecological change and as a tool for conservation.

The strategies underlying different forms of protective coloration are well understood but little attention has been paid to the ecological, life-history and behavioural circumstances under which they evolve. While some comparative studies have investigated the ecological correlates of aposematism, and background matching, the latter particularly in mammals, few have examined the ecological correlates of other types of protective coloration. Here, we first outline which types of defensive coloration strategies may be exhibited by the same individual; concluding that many protective coloration mechanisms can be employed simultaneously, particularly in conjunction with background matching. Second, we review the ecological predictions that have been made for each sort of protective coloration mechanism before systematically surveying phylogenetically controlled comparative studies linking ecological and social variables to antipredator defences that involve coloration. We find that some a priori predictions based on small-scale empirical studies and logical arguments are indeed supported by comparative data, especially in relation to how illumination affects both background matching and self-shadow concealment through countershading; how body size is associated with countershading, motion dazzle, flash coloration and aposematism, although only in selected taxa; how immobility may promote background matching in ambush predators; and how mobility may facilitate motion dazzle. Examination of nearly 120 comparative tests reveals that many focus on ecological variables that have little to do with predictions derived from antipredator defence theory, and that broad-scale ecological studies of defence strategies that incorporate phylogenetics are still very much in their infancy. We close by making recommendations for future evolutionary ecological research.

Biologists usually pursue the adaptationist paradigm in trying to explain the functional significance of animal coloration. Here I collate instances in which coloration may be a poor match in the context of background matching, Batesian mimicry, aposematism, and colour polymorphisms. This can occur because of trade-offs with other functions, relaxed selection from predation, or colour trait neutrality. Also, mechanistic, pleiotropic and chance genetic effects can all result in a poor match to the background environment or to signaling efficiently. While biologists implicitly recognise these constraints placed on adaptive coloration, they rarely explicitly acknowledge the heterodox notion that coloration might be under weak selection or no selection at all. Unfortunately, it is difficult to show this definitively, as illustrated in an investigation into the function of colour polymorphisms in coconut crabs.

Objectives Neonates of several primate species are born with very different pelage coloration from their parents but then assume adult coat color within weeks or months of birth. We set out to test the three conventional functional hypotheses for this distinctive transient natal coat coloration in primates (infanticide, allomothering, and confusion of paternity) as well as solicitation of maternal care, and across all neonate primates, background matching, and Gloger's rule (dark coats being associated with humid environments). Materials and Methods To examine the functional significance of neonate pelage coloration, we employed five different measures of distinctive coat coloration, new updated information on primate life histories, and a complete primate phylogeny of 286 species to control for shared ancestry. Results Across primates, we found a strong association between distinctive natal coats and the prevalence of infanticide but no support for allomothering or confusing paternity. We also found that species in which young have distinct natal coats have relatively shorter interbirth intervals, as well as a tendency to be weaned earlier which is unrelated to allomothering. Darker primate natal coats (whether distinctive or not) are not found in shadier, warmer, or wetter habitats. Discussion While distinct natal coats in primates are generally found in infanticidal species, we argue on logical grounds that this is not an advertisement to incoming males but instead neonate coloration solicits greater maternal care. This enables infants to pass through this infanticide-induced mortality-prone infant phase more rapidly. Conspicuous neonate coats in comparison to adults appear to act as a super-normal stimulus for mothers to hasten post-natal development of their offspring.

Cheetahs are thought to exhibit low genetic variability and, as a consequence, may suffer high juvenile mortality, impaired reproduction, and susceptibility to disease (O'Brien et al. 1983, 1985, 1986, 1987; Wildt et al. 1983; Marker & O'Brien 1989; Menotti-Raymond & O'Brien 1993). The species is now regularly quoted as an example of how lack of genetic diversity affects the persistence of species. Recently, however, independent criticisms have been leveled at this body of work questioning both the analysis and the standards of evidence for lack of genetic diversity (Caughley 1994; Merola 1994) as well as the evidence that cheetahs are suffering as a consequence (Laurenson et al. 1992; Caro 1994; Caro & Laurenson 1994; Caughley 1994; Laurenson 1994; Merola 1994). It is becoming clear that a loosely strung chain of poorly established findings will no longer convince a skeptical conservation community of the importance of genetics in the fate of this species. O'Brien (1994a) combines a number of points in his first rejoinder to these criticisms, including novel interpretations of ecological data that we obtained and concern about their collection and analysis. Here, we first show that this interpretation is seriously flawed and that our conclusion that predation is the key source of mortality in the wild is valid. Then, extending previous criticisms and continuing to focus on this one consequence of lack of genetic variability, we discuss O'Brien et al.'s (1985) interspecific comparison of juvenile mortality in captivity by highlighting analytical problems and presenting new data on captivity. In the Serengeti National Park, Tanzania, cheetah cubs suffered 95% mortality before reaching independence, with predation by lions and spotted hyenas accounting for the majority (73.2%) of cub deaths whose cause of death was definitely or probably known ( Laurenson 1994, Caro & Laurenson 1994). O'Brien (1994a) sug

The coconut crab Birgus latro, the largest terrestrial decapod, is under threat in most parts of its geographical range. Its life cycle involves two biomes (restricted terrestrial habitats near the coast, and salt water currents of the tropical Indian and Pacific Oceans). Its dependence on coastal habitat means it is highly vulnerable to the habitat destruction that typically accompanies human population expansion along coastlines. Additionally, it has a slow reproductive rate and can reach large adult body sizes that, together with its slow movement when on land, make it highly susceptible to overharvesting. We studied the distribution and population changes of coconut crabs at 15 island sites in coastal Tanzania on the western edge of the species' geographical range. Our aim was to provide the data required for reassessment of the extinction risk status of this species, which, despite indications of sharp declines in many places, is currently categorized on the IUCN Red List as Data Deficient. Pemba Island, Zanzibar, in Tanzania, is an important refuge for B. latro but subpopulations are fragmented and exploited by children and fishers. We discovered that larger subpopulations are found in the presence of crops and farther away from people, whereas the largest adult coconut crabs are found on more remote island reserves and where crabs are not exploited. Remoteness and protection still offer hope for this species but there are also opportunities for protection through local communities capitalizing on tourist revenue, a conservation solution that could be applied more generally across the species' range.

Aposematic coloration is traditionally considered to signal unpalatability or toxicity. In mammals, most research has focused on just one form of defense, namely, noxious anal secretions, and its black-and-white advertisement as exemplified by skunks. The original formulation of aposematism, however, encompassed a broader range of morphological, physiological, and behavioral defenses, and there are many mammal species with black-and-white contrasting patterns that do not have noxious adaptations. Here, using Bayesian phylogenetic models and data from 1726 terrestrial nonvolant mammals we find that two aspects of conspicuous coloration, black-and-white coloration patterns on the head and body, advertise defenses that are morphological (spines, large body size), behavioral (pugnacity), and physiological (anal secretions), as well as being involved with sexual signaling and environmental factors linked to crypsis. Within Carnivora, defensive anal secretions are associated with complex black-and-white head patterns and longitudinal black-and-white body striping; in primates, larger bodied species exhibit irregular patches of black-and-white pelage; and in rodents, pugnacity is linked to sharp countershading and irregular blocks of white and black pelage. We show that black-and-white coloration in mammals is multifunctional, that it serves to warn predators of several defenses other than noxious anal secretions, and that aposematism in mammals is not restricted to carnivores.

Aposematic coloration is commonly considered to signal unpalatability, yet animals advertise malodour, spines, and weaponry as well as toxins, some of which can be seen at a distance whereas others are hidden from predators. Separating defences into overt and covert categories in this way and whether they act before, during contact, or following ingestion generates new insights into the evolution of aposematism. Signals drawing attention to overt defences are difficult to fake whereas signals advertising covert defences can deceive would-be predators, and those acting later in the predatory sequence are more likely to be dishonest. These two orthogonal defence categorizations help to frame where dishonest signalling occurs in nature, set limits on deception by dishonest Batesian mimics, and prompt new questions.

Phenotypic colour polymorphisms are common in nature but their adaptive significance is often difficult to interpret. Terrestrial coconut crabs Birgus latro have cohabiting distinct red and blue co...

What happens when conservation interventions ostensibly fail? We outline a REDD+ intervention on Zanzibar, Tanzania which is adapting to a failure to implement carbon compensation payments and to the increased global price of cloves. Using a mix of quantitative and qualitative methods we provide preliminary evidence that well-managed shehia (wards) with registered Community Forestry Management Agreements (CoFMAs) are slowing their rates of deforestation. We also find an increase in the number of shehia with CoFMAs despite the absence of carbon payments. Using this island-wide case study we make inferences about the mechanisms whereby institutional expansion has occurred in ways consistent with cultural evolutionary dynamics of institutional change. We draw lessons for planning and practice that may help with the design of future conservation interventions and with bolstering the morale and effectiveness of disappointed partners.

Tim Caro introduces why zebras are striped.

Of all hypotheses advanced for why zebras have stripes, avoidance ofbiting fly attack receives by far the most support, yet the mechanisms bywhich stripes thwart landings are not yet understood. A logical and popularhypothesis is that stripes interfere with optic flow patterns needed by flyinginsects to execute controlled landings. This could occur through disruptingthe radial symmetry of optic flow via the aperture effect (i.e. generation offalse motion cues by straight edges), or through spatio-temporal aliasing(i.e. misregistration of repeated features) of evenly spaced stripes. By recordingand reconstructing tabanid fly behaviour around horses wearingdifferently patterned rugs, we could tease out these hypotheses using realistictarget stimuli. We found that flies avoided landing on, flew faster near,and did not approach as close to striped and checked rugs compared togrey. Our observations that flies avoided checked patterns in a similarway to stripes refutes the hypothesis that stripes disrupt optic flow via theaperture effect, which critically demands parallel striped patterns. Ourdata narrow the menu of fly-equid visual interactions that form the basisfor the extraordinary coloration of zebras.

Conspicuous coloration in animals has many possible functions including signaling to conspecifics, or predator deterrence through confusion, intimidation, and duping; the last includes flash behavior where predators are deceived into looking for conspicuous cues exhibited in flight but that are hidden when the animal comes to rest. In an effort to see if flash behavior occurs in mammals, we made predictions about situations where conspicuous coloration (as based on human assessment) might occur in artiodactyls and lagomorphs, and other predictions as to where such coloration might be found under an intraspecific signaling hypothesis. Using phylogenetically controlled analyses, we found that across species of artiodactyls, conspicuous rumps are more likely to have evolved in larger-sized group-living species supporting an intraspecific signaling function; this was not replicated in lagomorphs. Examining those artiodactyls that can facultatively expose color patches (putative flash behavior), we discovered that this trait occurred in artiodactyls that are solitary or living in very small groups irrespective of their body size. It is therefore possible that species such as white- and black-tailed deer, which display white rumps and tails during pursuit but hide them when stationary, are using flash behavior to confuse the predator into looking for the wrong object and thereby avoid detection and suggests that this form of antipredator defense in mammals needs greater attention. We found no effects of group size or body mass on conspicuous tail or ear markings in these taxa. Many mammals have conspicuous markings on their appendages and hindquarters, the function of which is mostly unknown. We matched these markings in rabbits, hares, and pikas and in bovids and cervids to both body size and group size across species. We found that conspicuous rumps are found in group living ungulates but when we separated these into conspicuous hindquarters always on display or that could be hidden, we found that hidden markings were principally found in species living alone or in very small groups irrespective of their body size. These species may expose conspicuous patches during flight but hide them at rest fooling the predator into searching for the wrong object, a relatively newly researched defense mechanism called flash behavior.

We scored pelage coloration of pigs and peccaries and matched them to socioecological variables to assess the relative strengths of protective coloration and signaling in driving coat coloration. Using phylogenetically controlled analyses, we found that faces, bellies, and perhaps flanks are lighter in larger species; ear coloration is lighter in less-shady environments and in more-social species; white facial spots are associated with less shade and with nighttime activity; and white body spots are associated with greater sociality. There is a marginal association between striped natal coats, a classically cryptic pelage, and litter size. These findings indicate that the body coloration of Suiformes has been selected to match overall background lighting conditions in both adults and neonates, but that specific areas and color patches on the body are associated with signaling. Our study suggests that small areas of contrasting pelage coloration are superimposed on classically cryptic body plans to allow both protective coloration and signaling to operate simultaneously.

Color polymorphisms are widespread in nature and can be maintained by several evolutionary processes. We used the coconut crab (Birgus latro) red/blue color polymorphism as a test case to explore the functional significance of intraspecific variation in crab coloration. Across our study sites on Pemba and Chumbe Islands, Tanzania, and Christmas Island, Australian Territory, red:blue morph ratios were 76.5%, 66.7% and 72.0% respectively, or approximately 3:1 in each case. To investigate whether coloration is sex or size dependent, signals strength or behavior, or is a physiological pleiotropic effect, we measured body weights, linear dimensions, pinch-force, behavioral dispositions and took crude environmental variables in the field. Except for a slight tendency for red crabs to be found in more open situations, we found no differences between color morphs for any of these variables. Other factors associated with color polymorphisms in other species, including frequency-dependent predation seems unlikely in such a large species with few natural predators, while niche separation in space seems improbable as morphs are typically found together. Assortative mating is plausible but mating is rapid, suggesting little choice is involved. Our findings suggest that neither morphological nor behavioral factors can explain color polymorphism in this species and that explanations must be found elsewhere.

Habitat fragmentation can play a major role in the reduction of genetic diversity among wildlife populations. The Ruaha-Rungwa and Katavi-Rukwa ecosystems in south-western Tanzania comprise one of the world’s largest remaining African savannah elephant metapopulation but are increasingly threatened by loss of connectivity and poaching for ivory. To investigate the genetic structure of populations, we compared the genotypes for nine microsatellite loci in the western, central and eastern populations. We found evidence of genetic differentiation among the three populations, but the levels were low and mostly concerned the younger cohort, suggesting recent divergence probably resulting from habitat loss between the two ecosystems. We identified weak isolation by distance, suggesting higher gene flow among individuals located less than 50km apart. In a long-lived species with overlapping generations, it takes a long time to develop genetic substructure even when there are substantial obstacles to migration. Thus, in these recently fragmented populations, inbreeding (and the loss of heterozygosity) may be less of an immediate concern than the loss of adults due to illegal hunting.

Flash behavior, in which otherwise cryptic prey exhibit conspicuous coloration or noise when fleeing from potential predators, has been postulated to hinder location of prey once they become stationary. Here, using artificial computer-generated prey and humans as visual predators, we show that human subjects are more likely to abandon their search for prey that flash, compared to continuously cryptic fleeing controls. Survivorship of flashing prey was an additional 20% higher than the survivorship of continuously cryptic prey, depending on the background against which it was depicted. This survivorship advantage was consistent regardless of whether prey showed flash colors continuously or intermittently during flight. The advantage over continuously cryptic prey was highest when the flashing prey was presented first. Likewise, the more search areas containing no prey that the volunteers had initially viewed, the more likely they were to give up when there was a cryptic prey present. Collectively, these 3 findings indicate that volunteers inferred the flashing prey was absent from the search area when they failed to see a prey in the same form as they saw it move. Our results demonstrate first proof of concept: flash behavior, widely seen in taxa from insects to mammals, is an effective antipredator escape mechanism.