Gloger's Rule Research Articles

Melanism in Polymorphic Terrestrial Snakes: A Meta‐Analysis and Systematic Review

ABSTRACTBackground and AimColour polymorphic populations constitute excellent model systems for testing ecological and evolutionary hypotheses, as alternate colourations are associated with differences in various biological, behavioural and life‐history traits. Melanism is the most common and most obvious form of polymorphism. In reptiles in general (and snakes in particular), thermal melanism has been a popular hypothesis used to explain the appearance of black individuals. It suggests that a trade‐off exists between superior thermoregulation and reduced protection through the loss of crypsis. Surprisingly, despite the growing body of literature available, to date, there are only two qualitative reviews on the evolutionary significance of melanism in reptiles and no quantitative synthesis. We conducted the first systematic review and meta‐analysis summarising the current knowledge on melanism in polymorphic terrestrial snakes and synthesised the evidence for an adaptive advantage of the melanistic morph.LocationNorthern Hemisphere.TaxonPolymorphic terrestrial snakes.ResultsMean prevalence of melanism is 31% with no significant differences between species. Annual precipitation was a significant moderator of melanism prevalence. Our results revealed no significant differences in odds ratios of melanism between sexes or in mean body size.Main ConclusionsEnvironmental plasticity can be considered the primary cause of melanism, possibly as a result of seasonal climatic variations, a result corroborated by the formal meta‐analysis conducted. Conclusions from the meta‐analysis are that melanism in snakes follows Gloger's rule, as is the case in birds and mammals, and as opposed to the thermal melanism hypothesis. Further, our findings do not lend support for other predictions from the thermal melanism hypothesis, such as skewed frequency of melanistics in favour of males or females, or larger body sizes in black individuals. Our results hold implications for the future diversity of animal populations, as climate change is predicted to decrease the degree of colour variation.

Spatio‐temporal shift in body size and plumage coloration is associated with the magnitude of climate change in a cosmopolitan owl

AbstractAimClimatic gradients shape geographic variation in phenotypic traits that are involved in animal thermoregulation. Therefore, increasing temperatures under global warming are expected to cause change over time in traits that show predictable spatial patterns according to environmental clines (body and appendage size, tegument coloration), as well exemplified by biogeographic rules and shown by increasing literature at a local scale. However, whether temporal shifts in phenotypic traits vary spatially according to the magnitude of the change in local climatic conditions is still unknown.LocationWorld.TaxonBarn owl species complex (Tyto alba group).MethodsBy using thousands of museum specimens collected across the globe from 1901 to 2018, we calculated within‐population change over time in wing length, bill length and melanin‐based plumage coloration and examined whether these trends differed across geographic regions, latitudes and gradients of climate changes.ResultsWe showed that populations exposed to an increasing warmer climate were subjected to a more marked decrease in body size, as gauged by wing length, and also absolute bill length that displayed a similar, although less steep, pattern of shrinking over time. In addition, phaeomelanin‐based ventral plumage colour has become paler in regions where the climate changed into warmer and drier, and darker where both temperature and precipitation increased.Main ConclusionsThese trends are generally coherent with the predictions based on Bergmann's and Gloger's rules, but not Allen's rule, and suggest that temporal shifts in body size and tegument colour depend on the magnitude of the alteration in climatic conditions, with populations living in regions where the climate has changed the most that are subjected to more marked phenotypic changes. Spatio‐temporal variation in climate has driven within‐species phenotypic clines, and it is expected to increase differences among populations according to the predicted further climate change.

The Colouration in Dicrurus caerulescens (White-bellied Drongo) Follows the Precipitation and Temperature of the Environment

The variation in climatic conditions over space and time is considered a major driving force in speciation. Gloger's rule is one such rule that broadly explains the variation in the colouration of endotherms (birds and mammals) with the effects of climatic parameters. This predicts that endothermic animals tend to have darker colouration in warm and rainy climates. Here we have tried to explain the variation in the belly white colour in Dicrurus caerulescens (White-bellied Drongo), an overall black coloured bird with a variable white belly. The dark bellied birds are considered the subspecies D. c., and the pale bellied birds are considered D. c.insularis (in dryzone of Sri Lanka). A total number of 112 individuals of adult White-bellied Drongo, including field (n=23) and museum samples (n=36) from Sri Lanka (Total 59), and museum samples from India (n=45) and Nepal (n=8) which were deposited at the National Museum of Sri Lanka, Field Museum of Natural History, USA and the Natural History Museum, UK were taken into consideration. The length of the white belly was measured using a dial calliper adopting a standardized method to minimize inconsistencies. To understand how the extent of white belly varied with respect to precipitation and temperature, we adopted generalized least squares (GLS) methods accounting for spatial autocorrelation between data points. The climatic variables were extracted from the world climatic data through QGIS software. The GLS methods were carried out using the ―nlme‖ package in the R Software for different orders of Autoregressive-moving average (ARMA) for the squared value of white belly length. To find the best fitting model, we used the Akaike information criterion (AIC) using the package ―AICcmodavg‖ in the R. From this, we found a negative significant effect of the interaction between the temperature and precipitation for the white belly length over the studied area in ARMA (1, 1) correlation (-3.72e-01±-0.022, t= -2.23, p<0.05). The white belly is smaller; hence the birds are darker where there is higher temperature and precipitation. The reverse (pale belly) is true when lower temperatures and/or humidity. Therefore, this follows the Gloger's rule of becoming darker in colouration with the high rainfall and temperature in D. caerulescens. 
 Keywords: Dicrurus caerulescens, Colouration, Gloger's rule, Temperature, Precipitation

Rodents show darker and redder coloration in warm and rainy environments

AbstractAimAnimal coloration varies in response to environmental conditions. One well‐known principle, Gloger's rule, suggests that warmer and wetter environments lead to more pigmented animals. Yet, the original formulation lacks differentiation between the two primary melanin pigments: eumelanin and pheomelanin. We examined spatial eumelanin and pheomelanin variation to unravel the impact of various ecological factors on pigment deposition, and to assess support for the complex version of Gloger's rule.LocationSouth America.Time PeriodContemporary.Major Taxa StudiedSigmodontine rodents.MethodsWe extracted pelage color data from 231 species and quantified the geographic variation in eu‐ and pheomelanin deposition. We performed linear multiple regression to investigate the influence of temperature, precipitation, predator diversity, and UVA‐B radiance in eumelanin (lightness) and pheomelanin (redness).ResultsOur findings support the original formulations of Gloger's rule. Rodents in warmer and rainier regions, which also entails greater exposure to UV radiation and a diverse range of predators, exhibit darker‐colored pelage. In addition, redder rodents prevail in warmer environments. However, contrary to the rule predictions, we observe a reversal for reddish patterns in relation to precipitation, with rainier regions showcasing more intense red rodents.Main ConclusionsOur study breaks new ground by investigating previously unexplored facets of Gloger's rule in a continental mammalian group. We discovered compelling evidence that darker and redder coloration aligns closely with temperature and rainfall gradients. Although we found support for eumelanin–pelage predictions, expectations for pheomelanin pigmentation were only partially met. Our results might suggest that selective pressures act differently on dark and reddish coloration, revealing that coloration patterns in response to climate are more intricate than previously formulated.

Predation Risk Drives Aposematic Signal Conformity.

Contrary to expectations regarding efficient predator education mediated by lack of ambiguity and enhanced prey recognition, aposematic signals often show considerable intraspecific variability. For example, some striped skunks (Mephitis mephitis) are almost entirely white, others have black-and-white stripes of equivalent thicknesses, yet others are mostly black. We tested the ecological correlates of this variation in patterning using 749 museum skins collected across North America. Skunks had longer white-black borders and more bilaterally symmetrical stripes in areas with a greater number of potential predator species, and this effect was more marked for mammalian than avian predators, the latter of which may be less deterred by noxious defenses. Skunks from locations with greater predator diversity were less variable in extent of whiteness on their dorsa and less variable in the length of their white-black borders, suggesting strong selection from predators leads to greater conformity in stripe patterns, even at the same location, but weak selection from predators leads to relaxed selection on pattern conformity. Skunks exhibited greater areas of black pelage in areas of greater humidity conforming to Gloger's rule. Our results indicate that relaxed predation pressure is key to warning signal variation in this iconic species, whereas stronger pressure leads to signal conformity and stronger signals.

Range-wide phylogenomics of the Great Horned Owl (Bubo virginianus) reveals deep north-south divergence in northern Peru.

The Great Horned Owl (Bubo virginianus) inhabits myriad habitats throughout the Americas and shows complex patterns of individual and geographic morphological variation. The owl family Strigidae is known to follow ecogeographic rules, such as Gloger's rule. Although untested at the species level, these ecogeographic rules may affect B. virginianus plumage coloration and body size. Previous studies have indicated that, despite this species' morphological variability, little genetic differentiation exists across parts of their range. This study uses reduced representation genome-wide nuclear and complete mitochondrial DNA sequence data to assess range-wide relationships among B. virginianus populations and the disputed species status of B. v. magellanicus (Magellanic or Lesser Horned Owl) of the central and southern Andes. We found shallow phylogenetic relationships generally structured latitudinally to the north of the central Andes, and a deep divergence between a southern and northern clade close to the Marañón Valley in the central Andes, a common biogeographic barrier. We identify evidence of gene flow between B. v. magellanicus and other subspecies based on mitonuclear discordance and F-branch statistics. Overall differences in morphology, plumage coloration, voice, and genomic divergence support species status for B. v. magellanicus.

Environment drives color pattern polymorphism in sand lizards beyond the Gloger's rule

AbstractSpecies color and color pattern vary geographically contributing to environmental tolerance of the species to the fluctuating climate. In a constantly changing environment, the population remains polymorphic, when individuals that are not acclimated to the current environment can survive adverse time periods. Factors influencing color morph frequencies in populations affect spatial variation through local adaptation, which is in turn linked to large‐scale environmental gradients. The influence of environmental factors has not been adequately studied in many polymorphic organisms where the influence of sexual selection on the persistence of polymorphisms is widely recognized. We hypothesized that different color morphs of sand lizard are distributed throughout the Palearctic depending on different environmental conditions. The goal of this study was to examine if the range of morph composition in a color polymorphic lizard can be explained by geographic and climatic variation in the Palearctic. We used publicly available data on sand lizard occurrence from the Global Biodiversity Information Facility and environmental variables from WorldClim and SEDAC databases. We categorized sand lizards' photos to 10 discrete color morphs based on color and color pattern variation. We predicted the color morph distribution using maximum entropy models. We found that variations in morph distributions were mostly related to temperature seasonality, precipitation, elevation, and anthropogenic activities. Our findings support the relationship between environmental conditions and color morph distribution, implying that environmental selection acts differently on color morphs, most likely in conjunction with sexual selection.

Regional variation in climate change alters the range-wide distribution of colour polymorphism in a wild bird.

According to Gloger's rule, animal colouration is expected to be darker in wetter and warmer climates. Such environmental clines are predicted to occur in colour polymorphic species and to be shaped by selection if colour morphs represent adaptations to different environments. We studied if the distribution of the colour polymorphic tawny owl (Strix aluco) morphs (a pheomelanic brown and a pale grey) across Europe follow the predictions of Gloger's rule and if there is a temporal change in the geographical patterns corresponding to regional variations in climate change. We used data on tawny owl museum skin specimen collections. First, we investigated long-term spatiotemporal variation in the probability of observing the colour morphs in different climate zones. Second, we studied if the probability of observing the colour morphs was associated with general climatic conditions. Third, we studied if weather fluctuations prior to the finding year of an owl explain colour morph in each climate zone. The brown tawny owl morph was historically more common than the grey morph in every studied climate zone. Over time, the brown morph has become rarer in the temperate and Mediterranean zone, whereas it has first become rarer but then again more common in the boreal zone. Based on general climatic conditions, winter and summer temperatures were positively and negatively associated with the proportion of brown morph, respectively. Winter precipitation was negatively associated with the proportion of brown morph. The effects of 5-year means of weather on the probability to observe a brown morph differed between climate zones, indicating region-dependent effect of climate change and weather on tawny owl colouration. To conclude, tawny owl colouration does not explicitly follow Gloger's rule, implying a time and space-dependent complex system shaped by many factors. We provide novel insights into how the geographic distribution of pheomelanin-based colour polymorphism is changing.

Ecological factors drive the divergence of morphological, colour and behavioural traits in cactus wrens (Aves, Troglodytidae).

The study of ecological mechanisms influencing organisms' phenotypic variation is a central subject of evolutionary biology. In this study, we characterized morphological, plumage colour and acoustic variation in cactus wrens Campylorhynchus brunneicapillus throughout its distribution. We assessed whether Gloger's, Allen's and Bergmann's ecogeographical rules, and the acoustic adaptation hypothesis relate to geographical trait variation. We analysed specimen coloration in belly and crown plumage, beak shape and structural song characteristics. We tested whether the subspecific classification or the peninsular/mainland groups mirrored the geographical variation in phenotypes and whether ecological factors were associated with patterns of trait variation. Our results suggest that colour, beak shape and acoustic traits varied across the range, in agreement with two lineages described by genetics. The simple versions of Gloger's and Allen's rules are related to variations in colour traits and morphology. Conversely, patterns of phenotypic variation did not support Bergmann's rule. The acoustic adaptation hypothesis supported song divergence for frequency-related traits. Phenotypic variation supports the hypothesis of two taxa: C. affinis in the Baja California peninsula and C. brunneicapillus in the mainland. The ecological factors are associated with phenotypic trait adaptations, suggesting that divergence between lineages could result from ecological divergence.

A practical guide to collections-based research on ecogeographic rules.

Ecogeographic research into how species' forms vary across space, time, and climate has taken on new urgency due to contemporary global climate change. Research using museum specimens and other records to study biological rules like Bergmann's, Allen's, and Gloger's Rules has a long history and continues to generate publications and robust scientific debates. Despite the prevalence and history of the field, however, no simple guide on how to carry out such work has ever been published. To lower the barriers of entry for new researchers, this review was created as a practical guide on how to perform ecogeographic research. The guide consolidates disparately published methodologies into a single, convenient document that reviews the history and present of the field of ecogeographic rule research, and describes how to generate appropriate hypotheses, design experiments, gather, and analyze biotic and geographic data, and interpret the results in an ecologically meaningful manner. The result is a semi-standardized guide that enables scientists at all levels from any institution to carry out an investigation from start to finish on any biological rule, taxon, and location of their choice.

Plumage Balances Camouflage and Thermoregulation in Horned Larks (Eremophila alpestris).

AbstractAnimal coloration serves many biological functions and must therefore balance potentially competing selective pressures. For example, many animals have camouflage in which coloration matches the visual background that predators scan for prey. However, different colors reflect different amounts of solar radiation and may therefore have thermoregulatory implications as well. In this study, we examined geographic variation in dorsal patterning, coloration, and solar reflectance among horned larks (Eremophila alpestris) of the western United States. We found that plumage brightness was positively associated with soil granularity, aridity, and temperature. Plumage redness-both in terms of saturation (i.e., chroma) and hue-was positively associated with soil redness and temperature, while plumage patterning was positively associated with soil granularity. Together, these plumage-environment associations support both background matching and Gloger's rule, a widespread ecogeographic pattern in animal coloration. We also constructed thermoregulatory models that estimated cooling benefits provided by solar reflectance profiles of the dorsal plumage of each specimen based on the collection site. We found increased cooling benefits in hotter, more arid environments. Finally, cooling benefits were positively associated with residual brightness, such that individuals that were brighter than expected based on environmental conditions also had higher cooling benefits, suggesting a trade-off between camouflage and thermoregulation. Together, these data suggest that natural selection has balanced camouflage and thermoregulation in horned larks, and they illustrate how multiple competing evolutionary pressures may interact to shape geographic variation in adaptive phenotypes.

Latitudinal gradients of reproductive traits in Japanese woody plants

AbstractLatitudinal gradients in the variability of plant trait expression are insufficiently studied. Here, we infer the expression and variability of major reproductive traits in Japanese angiosperm woody plants using geographical distributions of 773 woody plant species and 4682 grid cells at the 10 × 10 km2 spatial resolution. We focus on flower color, flower size, fruit size, fruit type, seed mass, and dispersal type and relate trait mean values and within cell variability to latitude, forest area and fragmentation, phylogenetic relatedness, and socioeconomic factors. To account for the correlation between variability and mean trait values, we use a recently developed model based on Taylor's power law (TPL). All studied reproductive traits exhibited strong although contrasting latitudinal gradients. Contrary to Gloger's rule, flowers tended to be darker at higher latitudes. Flower sizes increased and fruit and seed sizes decreased at higher latitudes. Color variability increased while variability in fruit size decreased toward higher latitudes. Higher human impact influenced fruit size positively and seed size negatively. We confirmed the usefulness of the TPL in the study of trait variability. We interpret the darker flower colors at higher latitudes as an adaptation to predominant bee pollination and thus as an effect of ecological filtering. Our study does not unequivocally corroborate the hypothesis that color variability within woody plant communities is governed by the intensity of ultraviolet radiation. We interpret the observed latitudinal trends toward higher variation in flower color and size in terms of increased numbers of generalist pollinators at higher latitudes.

COAT COLOR OF WESTERN LONG-EARED BATS (MYOTIS EVOTIS) LIVING IN DIFFERENT ENVIRONMENTS: A TEST OF GLOGER'S RULE

Gloger's rule is an ecogeographical pattern observed in many vertebrates whereby populations in more humid environments have darker pigmentation than populations in more arid environments. The coat color for several species of temperate bats exhibits this pattern, including the Western Long-eared Bat (Myotis evotis) in coastal versus interior populations. We tested Gloger's rule by comparing the coat of M. evotis in the mountains and prairies of Alberta, areas with similar humidity. We predicted that both sets of bats would have similar pigmentation if humidity is the main factor driving variation in coat coloration. Discriminant function analysis of color attributes readily separated M. evotis in the mountains from those in the prairies. Individuals were darker in the mountains, violating Gloger's rule. Differential selection by nocturnal predators against conspicuous coat may explain the color differences we found, and perhaps in other species of temperate bats.

Climate and body size have differential roles on melanism evolution across workers in a worldwide ant genus.

One of the main aspects associated with the diversity in animal colour is the variation in melanization levels. In ectotherms, melanism can be advantageous in aiding thermoregulation through heat absorption. Darker bodies may also serve as a shield from harmful UV-B radiation. Melanism may also confer protection against parasites and predators through improving immunity responses and camouflage in regions with high precipitation, with complex and shaded vegetations and greater diversity of pathogens and parasites. We studied melanism evolution in the globally distributed ant genus Pheidole under the pressures of temperature, UV-B radiation and precipitation, while considering the effects of body size and nest habit, traits that are commonly overlooked. More importantly, we account for worker caste polymorphism, which is marked by distinct roles and behaviours. We revealed for the first time distinct evolutionary trajectories for each worker subcaste. As expected, major workers from species inhabiting locations with lower temperatures and higherprecipitation tend to be more melanised. Curiously, we show a slight trend where minor workers of larger species also tend to have darker bodies when inhabiting regions with higher precipitation. Lastly, we did not find evidence for the effects of UV-B radiation and nest habit in the lightness variation of workers. Our paper explores the evolution of ant melanization considering a marked ant worker polymorphism and a wide range of ecological factors. We discuss our findings under the light of the Thermal Melanism Hypothesis, the Photoprotection Hypothesis and the Gloger's Rule.

Hair phenotype diversity across Indriidae lemurs

AbstractObjectivesHair (i.e., pelage/fur) is a salient feature of primate (including human) diversity and evolution—serving functions tied to thermoregulation, protection, camouflage, and signaling—but wild primate pelage evolution remains relatively understudied. Specifically, assessing multiple hypotheses across distinct phylogenetic scales is essential but is rarely conducted. We examine whole body hair color and density variation across Indriidae (Avahi, Indri, Propithecus)—a lineage that, like humans, exhibits vertical posture (i.e., their whole bodies are vertical to the sun).Materials and methodsOur analyses consider multiple phylogenetic scales (family‐level, genus‐level) and hypotheses (e.g., Gloger's rule, the body cooling hypotheses). We obtain hair color and density from museum and/or wild animals, opsin genotypes from wild animals, and climate data from WorldClim. To analyze our data, we use phylogenetic generalized linear mixed models (PGLMM) using Markov chain Monte Carlo algorithms.ResultsOur results show that across the Indriidae family, darker hair is typical in wetter regions. However, within Propithecus, dark black hair is common in colder forest regions. Results also show pelage redness increases in populations exhibiting enhanced color vision. Lastly, we find follicle density on the crown and limbs increases in dry and open environments.DiscussionThis study highlights how different selective pressures across distinct phylogenetic scales have likely acted on primate hair evolution. Specifically, our data across Propithecus may implicate thermoregulation and is the first empirical evidence of Bogert's rule in mammals. Our study also provides rare empirical evidence supporting an early hypothesis on hominin hair evolution.

On the evolution of distinctive natal coat coloration in primates

AbstractObjectivesNeonates 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 MethodsTo 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.ResultsAcross 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.DiscussionWhile 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.

Morphological variations in a widely distributed Eastern Asian passerine cannot be consistently explained by ecogeographic rules.

Ecogeographic rules that describe quantitative relationships between morphologies and climate might help us predict how morphometrics of animals was shaped by local temperature or humidity. Although the ecogeographic rules had been widely tested in animals of Europe and North America, they had not been fully validated for species in regions that are less studied. Here, we investigate the morphometric variation of a widely distributed East Asian passerine, the vinous‐throated parrotbill (Sinosuthora webbiana), to test whether its morphological variation conforms to the prediction of Bergmann's rule, Allen's rules, and Gloger's rule. We at first described the climatic niche of S. webbiana from occurrence records (n = 7838) and specimen records (n = 290). The results of analysis of covariance (ANCOVA) suggested that the plumage coloration of these parrotbills was darker in wetter/warmer environments following Gloger's rule. However, their appendage size (culmen length, beak volume, tarsi length) was larger in colder environments, the opposite of the predictions of Allen's rule. Similarly, their body size (wing length) was larger in warmer environments, the opposite of the predictions of Bergmann's rule. Such disconformity to both Bergmann's rule and Allen's rule suggests that the evolution of morphological variations is likely governed by multiple selection forces rather than dominated by thermoregulation. Our results suggest that these ecogeographic rules should be validated prior to forecasting biological responses to climate change especially for species in less‐studied regions.

Ecogeography of Plumage Pigmentation in Great Horned Owls

ABSTRACT Plumage pigmentation is fundamental to a bird's phenotype, with pigment deposition causing relative crypsis or conspicuousness, depending on the environmental context. Geographic variation in plumage melanin tends to be predictable, suggesting that aspects of climate cause local matching of plumage to environment via genetic adaptation. Ecogeographic rules describe this predictability: Gloger's rule predicts that populations in wetter and warmer environments will be more pigmented; Bogert's Rule predicts more pigmentation in cold environments. The Great Horned Owl (Bubo virginianus) exhibits extensive geographic variation in the degree of melanin-based pigmentation. We examined fine-scale spatial variation in owl plumage melanism along environmental gradients in southwestern North America. We tested whether variation is explained by either of two non-mutually exclusive hypotheses: (1) a history of allopatric divergence between subspecies or (2) in situ local adaptation consistent with ecogeographic rules. The allopatric divergence hypothesis predicts a bimodal distribution of plumage melanism, with a geographic cline across a zone of secondary contact, whereas the local adaptation hypothesis predicts that climate explains variation independently of geography. Using a colorimeter, we measured coloration in 101 museum specimens of breeding-season Great Horned Owls that had been obtained from variable environments and elevations. Specimens previously identified as separate subspecies were distinguishable by colorimetry. Plumage lightness, however, was continuously distributed, rather than bimodal. While accounting for males having reduced pigmentation relative to females, linear models revealed that lighter plumage was associated with low latitude, low elevation, high temperature, and low precipitation. These findings suggest that variation in Great Horned Owl plumage pigmentation is best understood as continuous ecogeographic variation, consistent with ecogeographic predictions, and currently maintained in situ along multiple environmental gradients that characterize the “sky island” topography of the southwestern USA.

Color lightness of velvet ants (Hymenoptera: Mutillidae) follows an environmental gradient

Color traits are highly influenced by environmental conditions along the distributional range of many species. Studies on the variation of animal coloration across different geographic gradients are, therefore, fundamental for a better understanding of the ecological and evolutionary processes that shape color variation. Here, we address whether color lightness in velvet ants (Hymenoptera: Mutillidae) responds to latitudinal gradients and bioclimatic variations, testing three ecogeographic rules: The Thermal melanism hypothesis; the Photoprotection hypothesis; and Gloger's rule. We test these hypotheses across the New World. We used photographs of 482 specimens (n=142 species) of female mutillid wasps and extracted data on color lightness (V). We analyzed whether variation in color is determined by bioclimatic factors, using Phylogenetic Generalized Least Square analysis. Our explanatory variables were temperature, ultraviolet radiation, humidity, and forest indicators. Our results were consistent with the Photoprotection hypothesis and Gloger's rule. Species with darker coloration occupied habitats with more vegetation, higher humidity, and UV-B radiation. However, our results refute one of the initial hypotheses suggesting that mutillids do not respond to the predictions of the Thermal melanism hypothesis. The results presented here provide the first evidence that abiotic components of the environment can act as ecological filters and as selective forces driving the body coloration of velvet ants. Finally, we suggest that studies using animals with melanin-based colors as a model for mimetic and aposematic coloration hypotheses consider that this coloration may also be under the influence of climatic factors and not only predators.

Rethinking Gloger's Rule: Climate, Light Environments, and Color in a Large Family of Tropical Birds (Furnariidae).

AbstractEcogeographic rules provide a framework within which to test evolutionary hypotheses of adaptation. Gloger's rule predicts that endothermic animals should have darker colors in warm/rainy climates. This rule also predicts that animals should be more rufous in warm/dry climates, the so-called complex Gloger's rule. Empirical studies frequently demonstrate that animals are darker in cool/wet climates rather than in warm/wet climates. Furthermore, sensory ecology predicts that, to enhance crypsis, animals should be darker in darker light environments. We aimed to disentangle the effects of climate and light environments on plumage color in the large Neotropical passerine family Furnariidae. We found that birds in cooler and rainier climates had darker plumage even after controlling for habitat type. Birds in darker habitats had darker plumage even after controlling for climate. The effects of temperature and precipitation interact so that the negative effect of precipitation on brightness is strongest in cool temperatures. Finally, birds tended to be more rufous in warm/dry habitats but also, surprisingly, in cool/wet locales. We suggest that Gloger's rule results from complementary selective pressures arising from myriad ecological factors, including crypsis, thermoregulation, parasite deterrence, and resistance to feather abrasion.