Geographical variation in bill size across bird species provides evidence for Allen's rule

Allen’s rule proposes that the appendages of endotherms are smaller, relative to body size, in colder climates, in order to reduce heat loss. Empirical support for Allen’s rule is mainly derived from occasional reports of geographical clines in extremity size of individual species. Interspecific evidence is restricted to two studies of leg proportions in seabirds and shorebirds. We used phylogenetic comparative analyses of 214 bird species to examine whether bird bills, significant sites of heat exchange, conform to Allen’s rule. The species comprised eight diverse taxonomic groups—toucans, African barbets, Australian parrots, estrildid finches, Canadian galliforms, penguins, gulls, and terns. Across all species, there were strongly significant relationships between bill length and both latitude and environmental temperature, with species in colder climates having significantly shorter bills. Patterns supporting Allen’s rule in relation to latitudinal or altitudinal distribution held within all groups except the finches. Evidence for a direct association with temperature was found within four groups (parrots, galliforms, penguins, and gulls). Support for Allen’s rule in leg elements was weaker, suggesting that bird bills may be more susceptible to thermoregulatory constraints generally. Our results provide the strongest comparative support yet published for Allen’s rule and demonstrate that thermoregulation has been an important factor in shaping the evolution of bird bills.

Geographical variation and population structure in the White-rumped Sandpiper Calidris fuscicollis as shown by morphology, mitochondrial DNA and carbon isotope ratios

We studied the population structure of a high arctic breeding wader bird species, the White-rumped Sandpiper Calidris fuscicollis. Breeding adults, chicks and juveniles were sampled at seven localities throughout the species' breeding range in arctic Canada in 1999. The mitochondrial control region was analysed by DNA sequencing, feathers were analysed for carbon isotope ratios (C13/C12) by isotope ratio mass spectrometry, and morphological measurements were analysed using principal component analyses, taking the effect of sex into account (identified by molecular genetic methods). In general, our results support the notion that the White-rumped Sandpiper is a monotypic species with no subspecies, and most of the morphological and genetic variation occurs within sites. Nevertheless, some differences between sites were found. Birds from the two northernmost sites (Ellesmere and Devon Islands) had relatively longer bill and wing and shorter tarsus than birds sampled further south, possibly reflecting genetic differences between populations. The carbon isotope ratios were higher at the easternmost site (Baffin Island), revealing differences in the isotope content of the food. The mtDNA sequences showed no significant differentiation between sites and no pattern of isolation-by-distance was found. Based on the mtDNA variation, the species was estimated to have a long-term effective population size of approximately 9,000 females. The species shows no clear evidence of any population expansion or decline. Our results indicate that carbon isotope ratios, and possibly also certain mtDNA haplotypes, may be useful as tools for identifying the breeding origin of White-rumped Sandpipers on migration and wintering sites.

Does the ring species concept predict vocal variation in the crimson rosella, Platycercus elegans, complex?

Vocal variation may be important in population divergence. We studied geographical variation in contact calls of parrots of the crimson rosella, Platycercus elegans, complex, which is characterized by striking geographical plumage coloration variation. This complex has long been considered a rare example of a ring species (where two divergent forms coexist in sympatry but are connected by a chain of intermediate populations forming a geographical ring). We tested whether contact call variation is consistent with the ring species hypothesis. We recorded calls throughout the ring, including several sites from the three main population groups forming the ring and interfaces between them. We analysed duration, peak frequency, fundamental frequency and frequency modulation. We found significant differences, particularly in fundamental frequency and frequency modulation, at multiple biogeographical scales ranging from local populations to subspecies level. Discriminant function analyses showed some populations could be reliably discriminated from call structure. However, our results provided little support for three key predictions of the ring species hypothesis: (1) calls of the terminal, most divergent forms were not significantly different in three of the four acoustic variables, and differences did not appear to be maintained in sympatry, (2) phenotypically/geographically intermediate populations were not characterized by intermediate calls, and (3) call variation was not concordant with geographical sequence around the ring from one terminal form to the other. Our results underscore the emerging view that the evolutionary histories and phenotypic variability of many long-held ring species may be inadequately described by the ring species hypothesis and require alternative explanations. (C) 2008 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.

Large-herbivore distribution and abundance : intra-and interspecific niche variation in the tropics

Determining the biological and environmental factors that limit the distribution and abundance of organisms is central to our understanding of the niche concept and crucial for predicting how species may respond to large-scale environmental change, such as global warming. However, detailed ecological information for the majority of species has been collected only at a local scale, and insufficient consideration has been given to geographical variation in intraspecific niche requirements. To evaluate the influence of environmental and biological factors on patterns of species distribution and abundance, we conducted a detailed, broadscale study across the tropical savannas of northern Australia on the ecology of three large, sympatric marsupial herbivores (family Macropodidae): the antilopine wallaroo (Macropus antilopinus), common wallaroo (M. robustus), and eastern grey kangaroo (M. giganteus). Using information on species abundance, climate, fire history, habitat, and resource availability, we constructed species' habitat models varying from the level of the complete distribution to smaller regional areas. Multiple factors affected macropod abundance, and the importance of these factors was dependent on the spatial scale of analyses. Fire regimes, water availability, geology, and soil type and climate were most important at the large scale, whereas aspects of habitat structure and interspecific species abundance were important at smaller scales. The distribution and abundance of eastern grey kangaroos and common wallaroos were strongly influenced by climate. Our results suggest that interspecific competition between antilopine wallaroos and eastern grey kangaroos may occur. The antilopine wallaroo and eastern grey kangaroo (grazers) preferred more nutrient-rich soils than the common wallaroo (grazer/browser), which we relate to differences in feeding modes. The abundance of antilopine wallaroos was higher on sites that were burned, whereas the abundance of common wallaroos was higher on unburned sites. Future climate change predicted for Australia has the capacity to seriously affect the abundance and conservation of macropod species in tropical savannas. The results of our models suggest that, in particular, the effects of changing climatic conditions on fire regimes, habitat structure, and water availability may lead to species declines and marked changes in macropod communities.

Copy number variation and transposable elements feature in recent, ongoing adaptation at the Cyp6g1 locus

The increased transcription of the Cyp6g1 gene of Drosophila melanogaster, and consequent resistance to insecticides such as DDT, is a widely cited example of adaptation mediated by cis-regulatory change. A fragment of an Accord transposable element inserted upstream of the Cyp6g1 gene is causally associated with resistance and has spread to high frequencies in populations around the world since the 1940s. Here we report the existence of a natural allelic series at this locus of D. melanogaster, involving copy number variation of Cyp6g1, and two additional transposable element insertions (a P and an HMS-Beagle). We provide evidence that this genetic variation underpins phenotypic variation, as the more derived the allele, the greater the level of DDT resistance. Tracking the spatial and temporal patterns of allele frequency changes indicates that the multiple steps of the allelic series are adaptive. Further, a DDT association study shows that the most resistant allele, Cyp6g1-[BP], is greatly enriched in the top 5% of the phenotypic distribution and accounts for ~16% of the underlying phenotypic variation in resistance to DDT. In contrast, copy number variation for another candidate resistance gene, Cyp12d1, is not associated with resistance. Thus the Cyp6g1 locus is a major contributor to DDT resistance in field populations, and evolution at this locus features multiple adaptive steps occurring in rapid succession.

Phenological response of sea turtles to environmental variation across a species' northern range

Variations in environmental parameters (e. g. temperature) that form part of global climate change have been associated with shifts in the timing of seasonal events for a broad range of organisms. Most studies evaluating such phenological shifts of individual taxa have focused on a limited number of locations, making it difficult to assess how such shifts vary regionally across a species range. Here, by using 1445 records of the date of first nesting for loggerhead sea turtles (Caretta caretta) at different breeding sites, on different continents and in different years across a broad latitudinal range (25-39 degrees ' N), we demonstrate that the gradient of the relationship between temperature and the date of first breeding is steeper at higher latitudes, i.e. the phenological responses to temperature appear strongest at the poleward range limit. These findings support the hypothesis that biological changes in response to climate change will be most acute at the poleward range limits and are in accordance with the predictions of MacArthur's hypothesis that poleward range limit for species range is environmentally limited. Our findings imply that the poleward populations of loggerheads are more sensitive to climate variations and thus they might display the impacts of climate change sooner and more prominently.

Selenium content of Australian foods: a review of literature values

The aim of this study was to review the published literature values for the selenium content of Australian foods. A secondary aim was to compare the results for Australian foods with food composition data from international sources to investigate the extent of geographical variation. Published food composition data sources for the selenium content in Australian foods were identified and assessed for data quality using established criteria. The selenium content is available for 148 individual food items. The highest values found are for fish (12.0–63.2 μg/100 g), meats (4.75–37.9 μg/100 g) and eggs (9.00–41.4 μg/100 g), followed by cereals (1.00–20.3 μg/100 g). Moderate levels are seen in dairy products (2.00–7.89 μg/100 g) while most fruits and vegetables have low levels (trace—3.27 μg/100 g). High selenium foods show the greatest level of geographical variation, with foods from the United States generally having higher selenium levels than Australian foods and foods from the United Kingdom and New Zealand having lower levels. This is the first attempt to review the available literature for selenium composition of Australian foods. These data serve as an interim measure for the assessment of selenium intake for use in epidemiological studies of diet–disease relationships.

Vocal mimicry in male bowerbirds : who learns from whom?

Vocal mimicry is one of the more striking aspects of avian vocalization and is widespread across songbirds. However, little is known about how mimics acquire heterospecific and environmental sounds. We investigated geographical and individual variation in the mimetic repertoires of males of a proficient mimic, the spotted bowerbird Ptilonorhynchus maculatus. Male bower owners shared more of their mimetic repertoires with neighbouring bower owners than with more distant males. However, interbower distance did not explain variation in the highly repeatable renditions given by bower owners of two commonly mimicked species. From the similarity between model and mimic vocalizations and the patterns of repertoire sharing among males, we suggest that the bowerbirds are learning their mimetic repertoire from heterospecifics and not from each other.

Disease mapping and regression with count data in the presence of overdispersion and spatial autocorrelation: a Bayesian model averaging approach

This paper applies the generalised linear model for modelling geographical variation to esophageal cancer incidence data in the Caspian region of Iran. The data have a complex and hierarchical structure that makes them suitable for hierarchical analysis using Bayesian techniques, but with care required to deal with problems arising from counts of events observed in small geographical areas when overdispersion and residual spatial autocorrelation are present. These considerations lead to nine regression models derived from using three probability distributions for count data: Poisson, generalised Poisson and negative binomial, and three different autocorrelation structures. We employ the framework of Bayesian variable selection and a Gibbs sampling based technique to identify significant cancer risk factors. The framework deals with situations where the number of possible models based on different combinations of candidate explanatory variables is large enough such that calculation of posterior probabilities for all models is difficult or infeasible. The evidence from applying the modelling methodology suggests that modelling strategies based on the use of generalised Poisson and negative binomial with spatial autocorrelation work well and provide a robust basis for inference.

Phytophthora cinnamomi in native vegetation communities of southern Victoia - morphological variation and paragyny among isolates

Morphology has often been used as an indicator of variability within species. The present study investigated morphological and physiological characteristics of isolates of Phytophthora cinnamomi collected from diseased vegetation communities at Anglesea, Victoria, and isolates collected from other regions in the State. Characteristics studied included growth rate on potato-dextrose agar (PDA), corn-meal agar and V8-juice agar at 24°C, growth rate on V8 agar at 15°C, colony morphology on PDA, sporangial and gametangial morphology, sporangial production and mating type. Phenotypic variation was demonstrated in radial growth rate, colony morphology and sporangial dimensions. Sporangial and oogonial dimensions and sporangial production were not significantly different between isolates from different geographical regions. All isolates were found to be of the A2 mating type suggesting variation was derived asexually. Paragynal associations, in an organism characteristically defined as amphigynal, were observed following crossing with A1 isolates. This is the first such study undertaken in southern Victoria. The findings highlight the importance of appropriate management of an area of such high conservation value as the Anglesea Heath to contain the current infection and to prevent introduction of new isolates into the area.

Climatic variation and breeding in the Australian magpie (Gymnorhina Tibicen) : a case study using existing data

To anticipate the effects of climate change on Australia’s avifauna, it is first necessary to understand the current effects of climate (including climate variability) on life histories, and to examine the scope and nature of existing data that may provide the necessary historical context to anticipate the effects of climate change. This study examines naturally occurring geographical gradients (altitude, latitude) and the Southern Oscillation Index (SOI) as integrated measures of climate. These are then compared with the timing and ‘amount’ of breeding recorded for the Australian Magpie (Gymnorhina tibicen) using data from Birds Australia’s Nest Record Scheme and Atlas of Australian Birds, the NSW Bird Atlassers Inc.’s NSW Bird Atlas, and the Canberra Ornitholgists Group’s Garden Bird Survey. For this common, easily identified species, these data suggest links between Australian Magpie breeding and all three environmental variables. Breeding became later as altitude increased, the proportion of breeding records increased from north to south, and years of high SOI corresponded to more (and earlier) breeding in this species. That annual climatic fluctuations have a direct, immediate and substantial effect on breeding in the Australian Magpie, particularly on the amount of breeding that occurs, implies that longer term changes in climate will have substantial impacts on populations. Results were not solely temperature-driven, which makes predicting climate change impacts difficult. For rainfall, predictions are far less precise and regional variation is higher. The results also highlight the potential and limitations of current survey techniques for documenting the impacts of climate change on birds; in particular, the Nest Record Scheme does not measure the amount of breeding that occurs, but a useful index of this can be derived from bird atlassing data

Geographic variation in allometry in the guppy (Poecilia reticulata)

Variation in static allometry, the power relationship between character size and body size among individuals at similar developmental stages, remains poorly understood. We tested whether predation or other ecological factors could affect static allometry by comparing the allometry between the caudal fin length and the body length in adult male guppies (Poecilia reticulata) among populations from different geographical areas, exposed to different predation pressures. Neither the allometric slopes nor the allometric elevations (intercept at constant slope) changed with predation pressure. However, populations from the Northern Range in Trinidad showed allometry with similar slopes but lower intercepts than populations from the Caroni and the Oropouche drainages. Because most of these populations are exposed to predation by the prawn Macrobrachium crenulatum, we speculated that the specific selection pressures exerted by this predator generated this change in relative caudal fin size, although effects of other environmental factors could not be ruled out. This study further suggests that the allometric elevation is more variable than the allometric slope.

Microsatellite variation in the yellowhammer Emberiza citrinella : population structure of a declining farmland bird

In recent years, there has been much concern in the UK about population declines of widespread species in agricultural habitats. Conservation-orientated research on declining birds has focused on vital rates of survival and productivity. However, the environmental factors which may influence movements between populations of widespread species is poorly understood. Population genetic structure is an indirect description of dispersal between groups of individuals. To attempt to develop an understanding of genetic structuring in a widespread, but declining, farmland bird, we therefore investigated the yellowhammer, Emberiza citrinella, population in England and Wales using microsatellite data. Our first aim was to investigate whether there was genetic substructuring in the population. A second aim was to investigate if there was a relationship between genetic distances and various environmental variables. Finally, we analysed the microsatellite data for evidence of loss of genetic variation due to population decline. Our data showed a slight but significant structure within the yellowhammer population. This therefore cannot be considered a panmictic population. Our example from South Cumbria implies that high-altitude barriers may have a slight influence on population structure. However, on the whole, genetic distances between sample sites were not significantly correlated with geographical distances, degrees of population connectivity, high altitudes, or differences in precipitation between sites. Finally, we detected departures from mutation-drift equilibrium (excess heterozygosity), which is indicative of a loss of genetic variation through recent decline.

Climate-related spatial and temporal variation in bill morphology over the past century in Australian parrots

Aim: Allen's rule posits that the appendages of endothermic organisms will be larger in warmer climates to allow for dumping of heat loads. Given a link between appendage size and climate, we tested the prediction that climate change has driven the evolution of larger bills in birds, resulting in measurable changes over the recent past. Location: Australia. Methods: We explored geographical and temporal variation in bill surface area of five Australian parrot species to determine whether individuals from warmer climates have larger bills, and whether there have been increases in bill surface area over time, consistent with climatic warming. Measurements were obtained from museum specimens dating from 1871 to 2008. These data were then related to geographical location, collection date and locality-specific climate data, in order to construct and compare models of spatio-temporal and climate-related variation in bill morphology. Results: There have been increases in bill surface area in mulga parrots (Psephotus varius), gang-gang cockatoos (Callocephalon fimbriatum), red-rumped parrots (Psephotus haematonotus) and male crimson rosellas (Platycercus elegans), equating to a c. 4-10% increase in bill surface area since 1871. Average maximum summer temperature in the 5 years prior to specimen collection also positively predicted bill surface area in mulga parrots, red-rumped parrots and crimson rosellas, consistent with Allen's rule. With the exception of red-rumped parrots, however, models with geographical location and year of collection were still better predictors of bill surface area than local climate at the date of collection. Main conclusions: Our analysis provides evidence that four species of parrot have exhibited adaptive change in bills over the past century potentially mitigating the thermal stress caused by climatic warming. Although consistent with the predicted effects of climate change, the temporal patterns we observe may have additional causes, however, such as changes in primary productivity, habitat or food availability.