Gloger's Rule Research Articles

Predicting biotic responses to future climate warming with classic ecogeographic rules

Models for future environmental change all involve global warming, whether slow or fast. Predicting how plants and animals will respond to such warming can be aided by using ecogeographic biological 'rules', some long-established, that make predictions based on observations in nature, as well as plausible physiological and ecological expectations. Bergmann's rule is well known, namely that warm-blooded animals are generally smaller in warm climates, but six further temperature-related rules - Allen's rule, Gloger's rule, Hesse's rule, Jordan's rule, Rapoport's rule and Thorson's rule - are also worth considering as predictive tools. These rules have been discussed in the recent ecological and physiological literature, and in some cases meta-analytical studies of multiple studies show how they are applicable across taxa and in particular physical environmental situations.

The dark side of coloration: Ecogeographical evidence supports Gloger's rule in American marsupials.

Geographical distribution of color phenotypes and associations with ecological predictors remains poorly understood. An important geographic pattern concerning this topic is Gloger's rule, which predicts the increase of pigmentation in endothermic animals from cold and dry to warm and wet environments. Didelphid marsupials exhibit a variety of color patterns, ranging from light and dark uniform to more complex colorations. However, surprisingly little is known about the adaptive significance of dark coloration in this singular group of mammals. Using a phylogenetic comparative approach, we investigated whether coloration in different body regions of didelphids (i.e., dorsum and face) is associated with variables representing heat and humidity of the environment, as predicted by Gloger's rule. We demonstrated that Gloger's rule explains the interspecific color variation in American marsupials, especially when considering the facial region. Thus, dark coloration was more frequent among didelphid species occupying warm and wet environments than cold and dry environments. We also discuss the selective forces that can potentially explain coat color variation in didelphid marsupials, including camouflage, pathogen resistance, and pleiotropy hypotheses.

A review of Gloger's rule, an ecogeographical rule of colour: definitions, interpretations and evidence

Gloger's rule is an ecogeographical rule that links animal colouration with climatic variation. This rule is named after C.W.L. Gloger who was one of the first to summarise the associations between climatic variation and animal colouration, noting in particular that birds and mammals seemed more pigmented in tropical regions. The term 'Gloger's rule' was coined by B. Rensch in 1929 and included different patterns of variation from those described by Gloger. Rensch defined the rule in two ways: a simple version stating that endothermic animals are predicted to be darker in warmer and humid areas due to the increased deposition of melanin pigments; and a complex version that includes the differential effects of humidity and temperature on both main types of melanin pigments-eu- and phaeo-melanin. The blackish eu-melanins are predicted to increase with humidity, and decrease only at extreme low temperatures, while the brown-yellowish phaeomelanins prevail in dry and warm regions and decrease rapidly with lower temperatures. A survey of the literature indicates that there is considerable variation/confusion in the way Gloger's rule is understood (based on 271 studies that define the rule). Whereas the complex version is hardly mentioned, only a quarter of the definitions are consistent with the simple version of Gloger's rule (darker where warm and wet), and most definitions mention only the effects of humidity (darker where wet). A smaller subset of studies define the rule based on other correlated climatic and environmental variables such as vegetation, latitude, altitude, solar radiation, etc., and a few even contradict the original definition (darker where cold). Based on the literature survey, I synthesised the qualitative (N=124 studies) and quantitative (meta-analytically, N=38 studies, 241 effects) evidence testing the simple version of Gloger's rule (I found no tests of the complex version). Both lines of evidence supported the predicted effects of humidity (and closely linked variables) on colour variation, but not the effects of temperature. Moreover, humidity effects are not restricted to birds and mammals, as the data indicate that these effects also apply to insects. This suggests that the simple version of Gloger's rule as originally defined may not be valid, and possibly that the rule should be re-formulated in terms of humidity effects only. I suggest, however, that more data are needed before such a reformulation, due to potential publication biases. In conclusion, I recommend that authors cite Rensch when referring to Gloger's rule and that they make clear which version they are referring to. Future research should concentrate on rigorously testing the validity and generality of both versions of Gloger's rule and establishing the mechanism(s) responsible for the patterns it describes. Since humidity seems to be the core climatic variable behind Gloger's rule, I suggest that the two most plausible mechanisms are camouflage and protection against parasites/pathogens, the latter possibly through pleiotropic effects on the immune system. Understanding the processes that lead to climatic effects on animal colouration may provide insights into past and future patterns of adaptation to climatic change.

Plumage coloration follows Gloger's rule in a ring species

AbstractAimAnimal coloration is expected to differ between populations in different habitats according to Gloger's rule, with darker animals found in more humid, vegetated or warmer environments. Yet despite considerable support across the globe, the mechanisms behind this biogeographical rule are currently still unclear. Exploiting a ring species with plumage coloration from crimson to pale yellow, we test Gloger's rule and the mechanisms behind phenotypic divergence.LocationSouth‐eastern Australia.Major taxa studiedCrimson rosellas (Platycercus elegans).MethodsWe combined three modelling approaches (spatial regression; random decision forest species distribution and conditional inference tree) to test the association between 10 environmental variables (based on long‐term climate data and remotely sensed reflectance of the land) and plumage coloration across the distribution of P. elegans. We also took in‐situ measurements of background coloration of dominant vegetation to examine the relationship between (a) background coloration measured locally and remotely, (b) P. elegans coloration and (c) known differences in visual sensitivity of the subspecies using species‐specific visual models.ResultsOn both a continental and a local scale, the distribution of yellow‐red plumage coloration was strongly predicted by average rainfall, summer temperature and the Earth's reflectance between 620–670 nm. Remotely sensed radiance measures correlated strongly and positively with reflectance of the leaves of the dominant tree species at sites across the P. elegans distribution. Visual modelling indicated that differences in background colour could affect signalling efficacy in dim‐light conditions.Main conclusionsOur study shows that the highly variable plumage coloration conforms to Gloger's rule, and indicates that background coloration of the vegetation and thermoregulation are likely to be important mechanisms. Our results also show that Gloger's rule can explain variation in pigmentary systems other than melanin, and highlight that selection from environmental variation could be an important force behind the geographic diversity found in ring species.

Do ecogeographical rules explain morphological variation in a diverse, Holarctic genus of small mammals?

AbstractAimWe use ecogeographical rules to understand the relationship between biogeography and morphological evolution in Sorex (Linnaeus, 1758) shrews. Specifically, we test for climatic correlations in body size (Bergmann's rule; larger species in colder habitats) and pelage brightness (Gloger's rule; darker species in more humid/warmer habitats).LocationNorth America and Eurasia (Holarctic).TaxonSorex spp. (Mammalia: Soricomorpha: Soricidae).MethodsWe compiled body size data for 51 species of Sorex and measured pelage brightness from museum specimens for 43 species. We sourced bioclimatic data across the geographical range of each species as well as specific to the museum specimen localities. For comparative purposes, we also estimated a Sorex phylogeny using existing sequence data. To test Bergmann's and Gloger's rules, we constructed phylogenetic least squares models considering latitude, altitude, and bioclimatic variables as predictors of interspecific variation in body size and pelage brightness across Sorex. We then performed these tests separately for the Palaearctic and Nearctic lineages.ResultsSorex exhibits wide variation in both body size and pelage brightness. In models of the entire genus and of the Nearctic clade, neither trait is significantly predicted by the variables tested. A decrease in pelage brightness in the Palaearctic clade is significantly predicted by increasing mean annual temperature and annual precipitation, but only when considering climatic data specific to the measured specimen localities.Main conclusionsSorex does not conform to Bergmann's rule. This result aligns with intraspecific studies suggesting Bergmann's rule is less evident in smaller taxa, but it counters others that found support for the rule in North American and European assemblages. Different patterns in pelage evolution across the Nearctic and Palaearctic clades may result from different historical climatic pressures; however, the significance of the relationship between climate and pelage evolution is dependent on the specificity of the climatic data. Sorex may be a useful focal taxon for exploring the relationship between intra‐ and interspecific ecogeographical patterns as well as the relative roles of morphological, behavioural, physiological, and life history characteristics in explaining the ability to persist in climatically challenging environments.

Transcriptomic analysis of skin pigmentation variation in the Virginia opossum (Didelphis virginiana).

Skin pigmentation and coat pigmentation are two of the best-studied examples of traits under natural selection given their quantifiable fitness interactions with the environment (e.g., camouflage) and signalling with other organisms (e.g., warning coloration). Previous morphological studies have found that skin pigmentation variation in the Virginia opossum (Didelphis virginiana) is associated with variation in precipitation and temperatures across its distribution range following Gloger's rule (lighter pigmentation in temperate environments). To investigate the molecular mechanism associated with skin pigmentation variation, we used RNA-Seq and quantified gene expression of wild opossums from tropical and temperate populations. Using differential expression analysis and a co-expression network approach, we found that expression variation in genes with melanocytic and immune functions is significantly associated with the degree of skin pigmentation variation and may be underlying this phenotypic difference. We also found evidence suggesting that the Wnt/β-catenin signalling pathway might be regulating the depigmentation observed in temperate populations. Based on our study results, we present several alternative hypotheses that may explain Gloger's rule pattern of skin pigmentation variation in opossum, including changes in pathogen diversity supporting a pathogen-resistant hypothesis, thermal stress associated with temperate environments, and pleiotropic and epistatic interactions between melanocytic and immune genes.

Darker where cold and wet: Australian birds follow their own version of Gloger's rule

Gloger's rule is usually interpreted as predicting darker coloured animals in warmer and more humid/vegetated regions. The relative importance of temperature and rainfall or vegetation is however unclear, and often only one variable is tested at a time, mainly through proxies. Here, I assess the predictions of Gloger's rule for interspecific achromatic plumage variation (dark to light variation) for an entire avifauna (551 species of Australian landbirds). I tested the effects of climatic variables (temperature and rainfall) and vegetation structure on plumage reflectance at species and assemblage level (100 × 100 km cells), controlling for phylogenetic relatedness and spatial autocorrelation. To assess the robustness of these results I compared observed results with those of a null distribution of effects obtained from repeatedly simulating random plumage reflectance evolution on the phylogeny. At both the species and assemblage level, darker coloured birds were found in wetter and colder regions and in more densely vegetated habitats. Simulations confirm results at the species level and the effect of temperature at the assemblage level, but rainfall and vegetation effects at the assemblage level fall within the distribution of simulated effects and should be interpreted with care. Interspecific colour variation in Australian birds supports Gloger's rule for rainfall/vegetation, but shows the opposite pattern for temperature. Darker colours in wet and vegetated environments are consistent with the role of melanin pigmentation in preventing feather degradation by bacteria, but also with background‐matching for camouflage. Darker plumage might be beneficial in colder regions or detrimental in warmer regions if it affects thermoregulation, a selective force often only assumed to be of importance for ectotherms. The data highlight the need to test the generality of biogeographic rules across levels and at broad scale. Experimental work is needed to confirm the mechanisms linking plumage achromatic variation to climate.

Ecogeographical gradients in plumage coloration among Australasian songbird clades

AbstractAimAnimal coloration often shows high degrees of evolutionary lability, producing variation among species that is easily apparent. This variation may produce consistent geographical patterns as species converge on adaptive phenotypes in similar environments. Some such geographical patterns in colour variation have been recently predicted as a response to the light environment in different habitats. Others like Gloger's rule – the negative relationship between brightness and humidity – have long been observed but still demand explanation. Finally, the conventional wisdom that tropical birds are more colourful remains largely untested.LocationAustralia, New Guinea.MethodsHere, we compared plumage coloration across two families of Australian birds (Meliphagidae, n = 97 species; Acanthizidae, n = 40 species) in a combined spatial and phylogenetic framework. We assessed the extent to which environmental variables extracted from species ranges explain variation in colour traits, while correcting for the autocorrelation inherent in spatially structured data using extensive simulations.ResultsWe found several strong effects of environment on plumage coloration. Inland species with ranges marked by high aridity and temperature seasonality showed greater colour span among acanthizids, and greater saturation among meliphagids. Gloger's rule was supported in both clades, but more strongly for dorsal plumage. The most consistent correlate in this relationship was vegetation: birds in regions with more vegetation had markedly darker plumage. Ornament hue showed no significant associations with vegetation or climate.Main conclusionsBirds living close to the equator were not more colourful, but species inhabiting arid regions were. Species may respond to the shorter and less predictable breeding seasons of arid environments by evolving increased ornamentation. The consistent relationship we observed between vegetation and dorsal brightness supports a primary role for countershading and background matching in Gloger's rule.

Gloger's rule in North American Barn Owls

ABSTRACT Studying the geographic variation of phenotypic traits can provide key information about the potential adaptive function of alternative phenotypes. Gloger's rule posits that animals should be dark- vs. light-colored in warm and humid vs. cold and dry habitats, respectively. The rule is based on the assumption that melanin pigments and/or dark coloration confer selective advantages in warm and humid regions. This rule may not apply, however, if genes for color are acting on other traits conferring fitness benefits in specific climes. Covariation between coloration and climate will therefore depend on the relative importance of coloration or melanin pigments and the genetically correlated physiological and behavioral processes that enable an animal to deal with climatic factors. The Barn Owl (Tyto alba) displays three melanin-based plumage traits, and we tested whether geographic variation in these traits at the scale of the North American continent supported Gloger's rule. An analysis of variation...

Floral pigmentation patterns provide an example of Gloger's rule in plants

Ecogeographic rules explain spatial trends in biodiversity, species interactions and phenotypes(1). Gloger's rule and its corollaries state that pigmentation of endothermic animals will increase from more polar to equatorial regions due to changing selective pressures including heat, humidity, predation and UV irradiance(2-4). In plants, floral pigmentation varies within and among taxa, yet causes of wide-scale geographic variation are lacking. We show that Gloger's rule explains patterns of variation in UV-absorbing floral pigmentation in a widespread plant, Argentina anserina (Rosaceae). Specifically, the floral pigmentation pattern unique to the UV spectrum (UV 'bullseye') increases with proximity to the Equator in both hemispheres, and larger bullseyes are associated with higher UVB incidence. Experiments confirm UV as an agent of selection and bullseye size as a target. Results extend the generality of an ecogeographic rule-formulated for animals-to plants, implicating UV as a selective agent on a floral trait generally assumed to enhance plant-pollinator interactions. Global change is expected to alter UV irradiance in terrestrial systems(5), potentially intensifying the importance of UV-mediated selection to floral evolution. Because floral UV reflectance and pattern enhance pollinator attraction(6,7), altered selective regimes could disrupt coevolved plant-pollinator interactions, weakening an important ecosystem service(8).

Flower colour: Gloger's rule isn't just for the birds.

A 180-year-old ‘law’ in zoology has found its best support so far in a study of floral colour, which not only documents darker plants growing closer to the equator, but also supports the idea that the colour stems from ultraviolet protection.

Pigmentation in Drosophila melanogaster reaches its maximum in Ethiopia and correlates most strongly with ultra-violet radiation in sub-Saharan Africa.

BackgroundPigmentation has a long history of investigation in evolutionary biology. In Drosophila melanogaster, latitudinal and altitudinal clines have been found but their underlying causes remain unclear. Moreover, most studies were conducted on cosmopolitan populations which have a relatively low level of genetic structure and diversity compared to sub-Saharan African populations. We investigated: 1) the correlation between pigmentation traits within and between the thorax and the fourth abdominal segment, and 2) their associations with different geographical and ecological variables, using 710 lines belonging to 30 sub-Saharan and cosmopolitan populations.ResultsPigmentation clines substantially differed between sub-Saharan and cosmopolitan populations. While positive correlations with latitude have previously been described in Europe, India and Australia, in agreement with Bogert's rule or the thermal melanism hypothesis, we found a significant negative correlation in Africa. This correlation persisted even after correction for altitude, which in its turn showed a positive correlation with pigmentation independently from latitude. More importantly, we found that thoracic pigmentation reaches its maximal values in this species in high-altitude populations of Ethiopia (1,600-3,100 m). Ethiopian flies have a diffuse wide thoracic trident making the mesonotum and the head almost black, a phenotype that is absent from all other sub-Saharan or cosmopolitan populations including high-altitude flies from Peru (~3,400 m). Ecological analyses indicated that the variable most predictive of pigmentation in Africa, especially for the thorax, was ultra-violet (UV) intensity, consistent with the so-called Gloger's rule invoking a role of melanin in UV protection.ConclusionOur data suggest that different environmental factors may shape clinal variation in tropical and temperate regions, and may lead to the evolution of different degrees of melanism in different high altitude populations in the tropics.

Latitudinal variation of morphological characteristics in the swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae)

AbstractPatterns of latitudinal variation in the phenotype or genotype of an organism may provide evidence for natural selection. In this study, we investigated seven populations of swallowtail Sericinus montelus Gray, 1798 (Lepidoptera: Papilionidae), a non‐migratory species, to explore the latitudinal variation of morphological characteristics in adults. The results showed that body size and the development of dark pigmentation on wings in this species responded strongly to latitude. The body size of both male and female adult of S. montelus was negatively correlated with latitude. These findings provided solid evidence to support the converse Bergmann's rule. We considered that the observed variation in morphological characteristics was most likely mediated by the seasonal length and thermoperiod to adapt to different latitudinal environment (e.g. shortened developmental time of immature stages for smaller body size at higher latitude). Moreover, the tendency towards progressively darker colour patterns (only in adult males) at increasingly low latitudes was consistent with Gloger's rule. We suggested that the observed colour variation was most likely associated with thermoregulation. Slight variation in the morphology of the W‐shaped stripe on the forewing of adult females was also found, and we presumed that the functions of sexual preferences, mimicry and thermoregulation might be involved.

Adaptation of Pelage Color and Pigment Variations in Israeli Subterranean Blind Mole Rats, Spalax Ehrenbergi

BackgroundConcealing coloration in rodents is well established. However, only a few studies examined how soil color, pelage color, hair-melanin content, and genetics (i.e., the causal chain) synergize to configure it. This study investigates the causal chain of dorsal coloration in Israeli subterranean blind mole rats, Spalax ehrenbergi.MethodsWe examined pelage coloration of 128 adult animals from 11 populations belonging to four species of Spalax ehrenbergi superspecies (Spalax galili, Spalax golani, Spalax carmeli, and Spalax judaei) and the corresponding coloration of soil samples from the collection sites using a digital colorimeter. Additionally, we quantified hair-melanin contents of 67 animals using HPLC and sequenced the MC1R gene in 68 individuals from all four mole rat species.ResultsDue to high variability of soil colors, the correlation between soil and pelage color coordinates was weak and significant only between soil hue and pelage lightness. Multiple stepwise forward regression revealed that soil lightness was significantly associated with all pelage color variables. Pelage color lightness among the four species increased with the higher southward aridity in accordance to Gloger's rule (darker in humid habitats and lighter in arid habitats). Darker and lighter pelage colors are associated with darker basalt and terra rossa, and lighter rendzina soils, respectively. Despite soil lightness varying significantly, pelage lightness and eumelanin converged among populations living in similar soil types. Partial sequencing of the MC1R gene identified three allelic variants, two of which were predominant in northern species (S. galili and S. golani), and the third was exclusive to southern species (S. carmeli and S. judaei), which might have caused the differences found in pheomelanin/eumelanin ratio.Conclusion/SignificanceDarker dorsal pelage in darker basalt and terra rossa soils in the north and lighter pelage in rendzina and loess soils in the south reflect the combined results of crypsis and thermoregulatory function following Gloger's rule.

Ecogeographic Patterns of Morphological Variation in Elepaios (Chasiempisspp.): Bergmann's, Allen's, and Gloger's Rules in a Microcosm

Abstract Animals often exhibit predictable geographic variation in morphology, and such ecogeographic patterns reflect local adaptation to varying environmental conditions. The most common of these patterns are termed Bergmann's, Allen's, and Gloger's rules. I studied morphological variation in the Hawaii Elepaio (Chasiempis sandwichensis) and the Oahu Elepaio (C. ibidis), forest birds endemic to the Hawaiian Islands. I measured body size and plumage color of 223 live elepaios captured at 36 sites on Hawaii and 132 live elepaios captured at 23 sites on Oahu, and I examined 132 museum specimens from an additional 22 locations on Hawaii. I used multiple regressions to examine relationships of elepaio body size and plumage color to elevation and annual rainfall on each island. Size of Hawaii Elepaios varied among sites and was related to elevation and rainfall. Wing chord, tail length, and body mass had positive relationships with elevation, as predicted by Bergmann's rule. Proportional bill length and propo...

Classifying nature: Constantin W. L. Gloger's (1803-1863) tapestry of a “Natural System of the Animal Kingdom ”

There has been no shortage of attempts to develop classification systems in order to capture the diversity of living forms in nature. One unpublished proposal (thus, largely unknown alternative method) was devised by Constantin Wilhelm Lambert Gloger (1803–1863), an ornithologist and zoologist from Upper Silesia who worked in Berlin for many years. Gloger described important phenomena of the geographical colour variation of birds and mammals (Gloger's Rule, 1833) and he combined as “climatic varieties” of polytypic species many representatives that had been described previously as independent species. During the early 1830s, Gloger started several large ornithological book projects which he abruptly discontinued when, in January 1834, an “enlightening inspiration” deviated his scientific endeavours towards work on a natural system of animals based on natural philosophical principles. He failed entirely in this attempt but continued work on it until the end of his life as a poor and lonely private scholar in Berlin. Gloger planned to publish this zoological classification system, which resembles a modern dichotomous identification key, under the title “Gloger's Natural System of the Animal Kingdom ”. However, his systematic tables, albeit printed and distributed “as manuscripts” by the author, were never published in coherent fashion. Five of these tables from 1834–1836, comprising his system of mammalian genera as interconnected folios of a 2.4 meter long tapestry, have recently been re-discovered in the Berlin Natural History Museum. They are here described in detail and illustrated for the first time, together with another folio depicting his system of mammalian families and one folio of Gloger's universal table of the world system. It remains unclear, though, how exactly Gloger intended to apply his “numbering system according to natural philosophy”, as he called it. Nevertheless, Gloger's approach illustrates alternative classification schemes representing the systematists' continuous endeavour to bring order into the seemingly chaotic nature, while highlighting the danger of fatal error and complete failure. (© 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim)

Gloger’s ecogeographical rule and colour variation among Willow Tits Parus montanus

The Willow Tit Parus montanus is distributed across the Palaearctic and into the Oriental zoogeographical region with 15 subspecies. P. m. borealis is the only subspecies in Fennoscandia. Based on 86 skinned museum specimens from Norway (south of 64 oN), we found relatively large variations in the colours of the mantle, the underpart and the cheeks. Birds along the coast are significantly darker and have less white on their cheeks than inland birds. Furthermore, the birds become paler on their mantle, underparts and cheeks towards north and east, and in particular the annual precipitation explains a significant part of the colour variation. This is in accordance with Gloger's climatic rule that points out that feathers tend to be darkly coloured in habitats where relative humidity is high and pale where it is low.

Variation of coat color in house mice throughout Asia

AbstractCoat color variation due to melanin pigment synthesis in house mice Mus musculus in Asia is described and found to be consistent with Gloger's rule, which states that individuals of endothermic animals are darker in humid habitats than those in drier habitats. Three properties of coat color (hue, value and chroma) were measured, and a lightness variable was derived from a principal components analysis using 428 skin specimens representing three subspecies from 85 localities. Dorsal coat color ranged from yellow through brown to black, whereas ventral coat color ranged from white to black. Dorsal coat color varied less than the ventral color. In our samples, the variation in coat color in natural populations was far less than that observed in the laboratory. We found a significant correlation between the lightness variable of dorsal coat color and precipitation. Dark coat color was observed in more humid and closed habitats (darker background color), and pale coat color in drier, more open habitats (lighter background color). This result might imply the role of concealment as a selective force affecting dorsal coat color that was observed in house mice. We also discussed other selective forces that could affect the coat color variation in house mice, such as resistance to bacterial degradation and thermoregulation. In addition, the color spectra of the dorsal pelage among the three subspecies were different, the major distinction being the environmental background color of the habitats in which they are distributed.

Gloger's Rule, Feather-Degrading Bacteria, and Color Variation Among Song Sparrows

AbstractFeathers tend to be darkly colored in habitats where relative humidity is high and pale where it is low. We suggest that this correlation, known as Gloger's rule, results, in part, from selection for dark feathers that are more resistant than light feathers to bacterial degradation, which is a severe problem in humid habitats where bacteria thrive and a lesser problem in arid habitats. In May and June 2000–2002 we sampled feather-degrading bacteria (Bacillus licheniformis) from the plumage of Song Sparrows (Melospiza melodia) in southeastern Arizona and northwestern Washington. Under standardized laboratory conditions, feather-degrading bacteria from the plumage of sparrows in the humid Northwest degraded feathers more rapidly and more completely than feather-degrading bacteria from sparrows of the arid Southwest. The differences in feather-degrading bacteria and in relative humidity produce a difference in the intensity of selection, which in turn produces the difference in color described in Gloger's rule.La Regla de Gloger, Bacterias Degradantes de Plumas y Variación de Color en Melospiza melodiaResumen. Las plumas tienden a ser de tonos obscuros en hábitats donde la humedad relativa es alta y más pálidas en hábitats donde la humedad relativa es baja. Esta correlación, conocida como la regla de Gloger, se aplica a muchas especies de aves a través del mundo. Sugerimos que la regla de Gloger es, en parte, un producto evolutivo de la selección por plumas obscuras, que son más resistentes a la degradación bacteriana que las plumas claras. La degradación bacteriana es un problema severo en hábitats húmedos donde prosperan las bacterias y un problema menor en hábitats áridos. En mayo y junio de 2000 a 2002 tomamos muestras de bacterias degradantes de plumas (Bacillus licheniformis) del Melospiza melodia fallax, que tiene plumaje pálido y reside en la parte sureste del estado de Arizona, y comparamos la incidencia y actividad de estas bacterias con las de aquellas encontradas en el plumaje obscuro de M. m. morphna, que reside de los bosques húmedos del noroeste del estado de Washington. Sin embargo, bajo condiciones estandarizadas de laboratorio, las bacterias obtenidas de M. m. morphna, degradaron las plumas más rápida y completamente que las bacterias de M. m. fallax. Las diferencias sugieren que las plumas obscuras de M. m. morphna del noroeste húmedo están sujetas a selección más intensa para resistir la degradación bacteriana que las plumas claras del gorrión del suroeste árido. La diferencia en humedad relativa produce una diferencia en la intensidad de selección, que a su vez produce la diferencia en color descrita en la regla de Gloger.

GLOGER'S RULE, FEATHER-DEGRADING BACTERIA, AND COLOR VARIATION AMONG SONG SPARROWS

Abstract Feathers tend to be darkly colored in habitats where relative humidity is high and pale where it is low. We suggest that this correlation, known as Gloger's rule, results, in part, from selection for dark feathers that are more resistant than light feathers to bacterial degradation, which is a severe problem in humid habitats where bacteria thrive and a lesser problem in arid habitats. In May and June 2000–2002 we sampled feather-degrading bacteria (Bacillus licheniformis) from the plumage of Song Sparrows (Melospiza melodia) in southeastern Arizona and northwestern Washington. Under standardized laboratory conditions, feather-degrading bacteria from the plumage of sparrows in the humid Northwest degraded feathers more rapidly and more completely than feather-degrading bacteria from sparrows of the arid Southwest. The differences in feather-degrading bacteria and in relative humidity produce a difference in the intensity of selection, which in turn produces the difference in color described in Gl...