Large-scale Geographic Variation Research Articles

Modelling the spatial risk of malaria through probability distribution of Anopheles maculipennis s.l. and imported cases

ABSTRACT Malaria remains one of the most important infectious diseases globally due to its high incidence and mortality rates. The influx of infected cases from endemic to non-endemic malaria regions like Europe has resulted in a public health concern over sporadic local outbreaks. This is facilitated by the continued presence of competent Anopheles vectors in non-endemic countries. We modelled the potential distribution of the main malaria vector across Spain using the ensemble of eight modelling techniques based on environmental parameters and the Anopheles maculipennis s.l. presence/absence data collected from 2000 to 2020. We then combined this map with the number of imported malaria cases in each municipality to detect the geographic hot spots with a higher risk of local malaria transmission. The malaria vector occurred preferentially in irrigated lands characterized by warm climate conditions and moderate annual precipitation. Some areas surrounding irrigated lands in northern Spain (e.g. Zaragoza, Logroño), mainland areas (e.g. Madrid, Toledo) and in the South (e.g. Huelva), presented a significant likelihood of A. maculipennis s.l. occurrence, with a large overlap with the presence of imported cases of malaria. While the risk of malaria re-emergence in Spain is low, it is not evenly distributed throughout the country. The four recorded local cases of mosquito-borne transmission occurred in areas with a high overlap of imported cases and mosquito presence. Integrating mosquito distribution with human incidence cases provides an effective tool for the quantification of large-scale geographic variation in transmission risk and pinpointing priority areas for targeted surveillance and prevention.

Extensive regional variation in the phenology of insects and their response to temperature across North America.

Climate change models often assume similar responses to temperatures across the range of a species, but local adaptation or phenotypic plasticity can lead plants and animals to respond differently to temperature in different parts of their range. To date, there have been few tests of this assumption at the scale of continents, so it is unclear if this is a large-scale problem. Here, we examined the assumption that insect taxa show similar responses to temperature at 96 sites in grassy habitats across North America. We sampled insects with Malaise traps during 2019-2021 (N = 1041 samples) and examined the biomass of insects in relation to temperature and time of season. Our samples mostly contained Diptera (33%), Lepidoptera (19%), Hymenoptera (18%), and Coleoptera (10%). We found strong regional differences in the phenology of insects and their response to temperature, even within the same taxonomic group, habitat type, and time of season. For example, the biomass of nematoceran flies increased across the season in the central part of the continent, but it only showed a small increase in the Northeast and a seasonal decline in the Southeast and West. At a smaller scale, insect biomass at different traps operating on the same days was correlated up to ~75 km apart. Large-scale geographic and phenological variation in insect biomass and abundance has not been studied well, and it is a major source of controversy in previous analyses of insect declines that have aggregated studies from different locations and time periods. Our study illustrates that large-scale predictions about changes in insect populations, and their causes, will need to incorporate regional and taxonomic differences in the response to temperature.

Geographical variation in nest defense among cinereous tit populations in China.

Behavioral divergence among populations is common across taxonomic groups, still we know very little about anti-predator behaviors. Animal exposure to predation risk is variable in different ecological contexts. In addition, reproduction value of animals in different geographical regions usually varies. In this study, we tested whether cinereous tits Parus cinereus in different populations exhibited nest defense behaviors similar to those of nest or adult predators and whether their nest defense behaviors showed geographical variation. By using field dummy experiments, we observed tits' nest defense behavior in nest predator common chipmunk Tamias sibiricus and red squirrel Sciurus vulgaris, adult predator Eurasian sparrowhawk Accipiter nisus and nonthreatening species Oriental turtle dove Streptopelia orientalis in the ZJ (44° N), DZ (31° N), and DLS (18° N) populations, respectively. The response scores varied significantly across the 4 dummies in ZJ-tits and DLS-tits but did not in DZ-tits. When facing the chipmunk, ZJ-tits showed the highest response score and DZ-tits showed the lowest response score. When facing the squirrel, ZJ-tits showed a higher response score than tits in the other 2 populations. However, tits among the 3 populations responded similarly to a sparrowhawk or dove with slight response behaviors. In addition, response scores to nest predators were positively correlated with brood size across the 3 populations, but no trend was found for sparrowhawks or doves. Our results indicated that the nest defense behaviors of cinereous tits have evolved in response to large-scale geographical variation in ecological contexts and reproduction value.

Transgenerational Genetic Effects Help Explain Latitudinal Variation in Seed Mass and Germination Timing in Plantago lanceolata.

We know little about the underlying genetic control of phenotypic patterns of seed traits across large-scale geographic and environmental gradients. Such knowledge is important for understanding the evolution of populations within species and for improving species conservation. Therefore, to test for genetic variation in Plantago lanceolata, we made reciprocal crosses between northern and southern genotypes that span the species’ range in Europe. The results provide evidence of transgenerational genetic effects on seed mass and germination timing. Northern mothers produced larger seeds with delayed germination, in contrast to southern mothers, which produced smaller seeds with accelerated germination. A maternal latitude affected both the seed coat, solely maternal tissue, and embryo/endosperm tissues. Thus, latitudinal variation in seed size and germination timing can be explained, in part, by the direct influence of maternal genotype, independent of zygotic genes that parents pass directly to the embryo and endosperm. Data suggest that researchers exploring the existence and evolution of large-scale geographic variation within species test for transgenerational genetic effects. In addition, data suggest that transgenerational control of seed traits should be considered when developing procedures designed to facilitate species conservation and restoration.

Geographic Variation in Floral Color and Reflectance Correlates With Temperature and Colonization History.

Petal color variation within species is common and may be molded by abiotic or biotic selection pressures, or neutral population structure. For example, darker flowers may be favored in cooler environments because they absorb more solar radiation, elevating the temperature of reproductive structures. Additionally, flower color may evolve to attract the dominant or most efficient pollinator type in a given population. Here, we evaluate geographic variation in petal coloration across the range of Campanula americana in Eastern North America and test whether color covaries with abiotic factors, the pollination community, and genetic structure established through post-glacial expansion. Consistent with other studies, flowers from cooler, higher latitude populations were less reflective across the UV-NIR spectrum than those from warmer populations. Local temperature explained variation in petal reflectance better than the pollinator community or colonization history. Petal color perceived by trichromatic bee pollinators displayed a strong longitudinal pattern but was unassociated with climatic factors and the pollinator community. Instead, pollinator-perceived color was tightly correlated with the geographic distance from C. americana's glacial refugium. In total, abiotic conditions appear to shape large-scale geographic variation in the intensity of petal reflectance while genetic structure is the strongest driver of pollinator-perceived petal coloration. This study highlights the importance of abiotic factors and historical processes associated with range expansion as major evolutionary forces shaping diversity of flower coloration on large geographic scales.

Competition influences tree growth, but not mortality, across environmental gradients in Amazonia and tropical Africa.

Competition among trees is an important driver of community structure and dynamics in tropical forests. Neighboring trees may impact an individual tree’s growth rate and probability of mortality, but large‐scale geographic and environmental variation in these competitive effects has yet to be evaluated across the tropical forest biome. We quantified effects of competition on tree‐level basal area growth and mortality for trees ≥10‐cm diameter across 151 ~1‐ha plots in mature tropical forests in Amazonia and tropical Africa by developing nonlinear models that accounted for wood density, tree size, and neighborhood crowding. Using these models, we assessed how water availability (i.e., climatic water deficit) and soil fertility influenced the predicted plot‐level strength of competition (i.e., the extent to which growth is reduced, or mortality is increased, by competition across all individual trees). On both continents, tree basal area growth decreased with wood density and increased with tree size. Growth decreased with neighborhood crowding, which suggests that competition is important. Tree mortality decreased with wood density and generally increased with tree size, but was apparently unaffected by neighborhood crowding. Across plots, variation in the plot‐level strength of competition was most strongly related to plot basal area (i.e., the sum of the basal area of all trees in a plot), with greater reductions in growth occurring in forests with high basal area, but in Amazonia, the strength of competition also varied with plot‐level wood density. In Amazonia, the strength of competition increased with water availability because of the greater basal area of wetter forests, but was only weakly related to soil fertility. In Africa, competition was weakly related to soil fertility and invariant across the shorter water availability gradient. Overall, our results suggest that competition influences the structure and dynamics of tropical forests primarily through effects on individual tree growth rather than mortality and that the strength of competition largely depends on environment‐mediated variation in basal area.

A century later: Adaptive plasticity and rapid evolution contribute to geographic variation in invasive mosquitofish

One century after their introduction to Europe, eastern mosquitofish (Gambusia holbrooki) represent a natural experiment to determine the relative contributions of adaptive plasticity and rapid evolutionary change in creating large-scale geographic variation in phenotypes. We evaluated the population-genetic structure and invasion history based on allele length polymorphisms of 15 nuclear microsatellites, which we quantified for N = 660 individuals from 23 populations sampled in 2013 across the invasive range of G. holbrooki in Europe. We analysed body-shape and life-history variation in N = 1331 individuals from 36 populations, sampled in 2013 and 2017, and tested heritability of phenotypic differences in a subset of four populations using a common-garden experiment. The genetic structure of wild-caught individuals suggested a single introduction for all European mosquitofish, which were genetically impoverished compared to their native counterparts. We found some convergent patterns of phenotypic divergence across native and invasive climatic gradients (e.g., increased body size in colder/more northern populations); however, several phenotypic responses were not consistent between sampling years, pointing towards plastic phenotypes. Our analysis of common-garden reared individuals uncovered moderate heritability estimates only for two measures of male body size (intraclass correlation coefficient, ICC = 0.628 and 0.556) and offspring fat content (ICC = 0.734), while suggesting high levels of plasticity in most other phenotypic traits (ICC ≤ 0.407). Our results highlight the importance of phenotypic plasticity in invasive species during range expansions and demonstrate that strong selective pressures—in this case towards increased body size in colder environments—simultaneously promote rapid evolutionary divergence.

Discrete or indiscrete? Redefining the colour polymorphism of the land snail Cepaea nemoralis

Biologists have long tried to describe and name the different phenotypes that make up the shell polymorphism of the land snail Cepaea nemoralis. Traditionally, the view is that the ground colour of the shell is one of a few major colour classes, either yellow, pink or brown, but in practise it is frequently difficult to distinguish the colours, and define different shades of the same colour. To understand whether colour variation is in reality continuous, and to investigate how the variation may be perceived by an avian predator, we applied psychophysical models of colour vision to shell reflectance measures. We found that both achromatic and chromatic variation are indiscrete in Cepaea nemoralis, being continuously distributed over many perceptual units. Nonetheless, clustering analysis based on the density of the distribution did reveal three groups, roughly corresponding to human-perceived yellow, pink and brown shells. We also found large-scale geographic variation in the frequency of these groups across Europe, and some covariance between shell colour and banding patterns. Although further studies are necessary, the observation of continuous variation in colour is intriguing because the traditional theory is that the underlying supergene that determines colour has evolved to prevent phenotypes from “dissolving” into continuous trait distributions. The findings thus have significance for understanding the Cepaea polymorphism, and the nature of the selection that acts upon it, as well as more generally highlighting the need to measure colour objectively in other systems.

Variation in ectomycorrhizal fungal communities associated with Silver linden (Tilia tomentosa) within and across urban areas.

Trees in urban areas face harsh environmental conditions. Ectomycorrhizal fungi (EcM) form a symbiosis with many tree species and provide a range of benefits to their host through their extraradical hyphal network. Although our understanding of the environmental drivers and large scale geographical variation of EcM communities in natural ecosystems is growing, our knowledge of EcM communities within and across urban areas is still limited. Here, we characterized EcM communities using Illumina miseq sequencing on 175 root samples of the urban tree Tilia tomentosa from three European cities, namely Leuven (Belgium), Strasbourg (France) and Porto (Portugal). We found strong differences in EcM richness and community composition between cities. Soil acidity, organic matter and moisture content were significantly associated with EcM community composition. In agreement, the explained variability in EcM communities was mostly attributed to general soil characteristics, whereas very little variation was explained by city and heavy metal pollution. Overall, our results suggest that EcM communities in urban areas are significantly associated with soil characteristics, while heavy metal pollution and biogeography had little or no impact. These findings deliver new insights into EcM distribution patterns in urban areas and contribute to specific inoculation strategies to improve urban tree vitality.

The ecological success of a social parasite increases with manipulation of collective host behaviour.

Many parasites alter the behaviour of their host to their own advantage, yet hosts often vary in their susceptibility to manipulation. The ecological and evolutionary implications of such variation can be profound, as resistant host populations may suffer lower parasite pressures than those susceptible to manipulation. To test this prediction, we assessed parasite-induced aggressive behaviours across 16 populations of two Temnothorax ant species, many of which harbour the slavemaker ant Protomognathus americanus. This social parasite uses its Dufour's gland secretions to manipulate its hosts into attacking nestmates, which may deter defenders away from itself during invasion. We indeed find that colonies that were manipulated into attacking their Dufour-treated nestmates were less aggressive towards the slavemaker than those that did not show slavemaker-induced nestmate attack. Slavemakers benefited from altering their hosts' aggression, as both the likelihood that slavemakers survived host encounters and slavemaker prevalence in ant communities increased with slavemaker-induced nestmate attack. Finally, we show that Temnothorax longispinosus colonies were more susceptible to manipulation than Temnothorax curvispinosus colonies. This explains why T.curvispinosus colonies responded with more aggression towards invading slavemakers, why they were less likely to let slavemakers escape and why they were less frequently parasitized by the slavemaker than T.longispinosus. Our findings highlight that large-scale geographic variation in resistance to manipulation can have important implications for the prevalence and host preference of parasites.

Latitudinal variation in behaviors linked to risk tolerance is driven by nest-site competition and spatial distribution in the ant Temnothorax rugatulus

Geographic range has long been noted to be associated with many organismic and ecological traits such as body size and species richness. However, much less is known about whether and how ecological variation across latitudinal gradients reflects behavioral variation. Ant colonies may also show behavioral variation, and Temnothorax rugatulus show a colony-level behavioral syndrome that seems to reflect risk tolerance across their North American range. While it is presumed that this pattern is the result of adaptation to local environmental conditions, which ecological factors are driving this variation are unknown. Here, we test if colony risk tolerance is affected by competition, predation, resource availability, or environmental stress at each site. Our results show that increased competition, specifically for nest sites, as well as increased spatial clustering of colonies predicts higher risk tolerance. Additionally, the spatial clustering of colonies influences the structure of the risk-taking syndrome, i.e., which colony-level behaviors are correlated and how strongly. This emphasizes the need for understanding large-scale geographic variation in behavior, as it may explain how ecological factors drive the evolution and maintenance of intraspecific behavioral variation across populations.

Large-scale geographic variation in distribution and abundance of Australian deep-water kelp forests.

Despite the significance of marine habitat-forming organisms, little is known about their large-scale distribution and abundance in deeper waters, where they are difficult to access. Such information is necessary to develop sound conservation and management strategies. Kelps are main habitat-formers in temperate reefs worldwide; however, these habitats are highly sensitive to environmental change. The kelp Ecklonia radiate is the major habitat-forming organism on subtidal reefs in temperate Australia. Here, we provide large-scale ecological data encompassing the latitudinal distribution along the continent of these kelp forests, which is a necessary first step towards quantitative inferences about the effects of climatic change and other stressors on these valuable habitats. We used the Autonomous Underwater Vehicle (AUV) facility of Australia’s Integrated Marine Observing System (IMOS) to survey 157,000 m2 of seabed, of which ca 13,000 m2 were used to quantify kelp covers at multiple spatial scales (10–100 m to 100–1,000 km) and depths (15–60 m) across several regions ca 2–6° latitude apart along the East and West coast of Australia. We investigated the large-scale geographic variation in distribution and abundance of deep-water kelp (>15 m depth) and their relationships with physical variables. Kelp cover generally increased with latitude despite great variability at smaller spatial scales. Maximum depth of kelp occurrence was 40–50 m. Kelp latitudinal distribution along the continent was most strongly related to water temperature and substratum availability. This extensive survey data, coupled with ongoing AUV missions, will allow for the detection of long-term shifts in the distribution and abundance of habitat-forming kelp and the organisms they support on a continental scale, and provide information necessary for successful implementation and management of conservation reserves.

Incorporating site and year-specific deuterium ratios (δ2 H) from precipitation into geographic assignments of a migratory bird

The study of migratory connectivity is rapidly growing in ornithology, as is the technology used to measure it. While use of extrinsic markers, such as archival tags, is becoming more prevalent, for many small species the best tool available for tracking birds remains intrinsic markers, such as stable-hydrogen isotope ratios (H-2). Many researchers have raised concerns that spatial and temporal environmental variation introduces a large amount of error into isotope-based assignments, limiting their utility. Here, using feathers, we sought to address these issues in developing H-2 base maps for assigning pied flycatchers Ficedula hypoleuca of known origin to 15 sites across the breeding range (approx. 4 020 800 km(2)). We evaluated the effects of including random site variation and year-specific precipitation H-2 (H-2(P)) maps on assignments, compared to using mean annual growing season H-2(p) and no site effects. We found a positive correlation between feather H-2 (H-2(F)) and mean annual H-2(P,) resulting from large scale geographic variation. Repeatability of feather H-2 for individuals sampled in multiple years was strong overall, but variable among populations. Annual variation in H-2(P) explained 21% of within individual variation in H-2(F). Neither year- nor site-specific methods improved assignment precision or accuracy. All three methods assigned flycatchers of unknown origin captured at an African overwintering site to similar breeding areas. However, methods using long-term means of H-2(p) assigned birds more precisely than year-specific methods. Our results suggest that annual variation in this system is primarily a result of food web or individual level processes and that random site effects are not strong enough to drastically impact accuracy. We conclude that improvements in isotope based geographic assignments will rely on the addition of prior information, such as relative abundance in a Bayesian framework, or additional intrinsic markers.

Large-scale geographical variation in eggshell metal and calcium content in a passerine bird (Ficedula hypoleuca)

Birds have been used as bioindicators of pollution, such as toxic metals. Levels of pollutants in eggs are especially interesting, as developing birds are more sensitive to detrimental effects of pollutants than adults. Only very few studies have monitored intraspecific, large-scale variation in metal pollution across a species' breeding range. We studied large-scale geographic variation in metal levels in the eggs of a small passerine, the pied flycatcher (Ficedula hypoleuca), sampled from 15 populations across Europe. We measured 10 eggshell elements (As, Cd, Cr, Cu, Ni, Pb, Zn, Se, Sr, and Ca) and several shell characteristics (mass, thickness, porosity, and color). We found significant variation among populations in eggshell metal levels for all metals except copper. Eggshell lead, zinc, and chromium levels decreased from central Europe to the north, in line with the gradient in pollution levels over Europe, thus suggesting that eggshell can be used as an indicator of pollution levels. Eggshell lead levels were also correlated with soil lead levels and pH. Most of the metals were not correlated with eggshell characteristics, with the exception of shell mass, or with breeding success, which may suggest that birds can cope well with the current background exposure levels across Europe.

Moose body mass variation revisited: disentangling effects of environmental conditions and genetics

Large-scale geographical variation in phenotypic traits within species is often correlated to local environmental conditions and population density. Such phenotypic variation has recently been shown to also be influenced by genetic structuring of populations. In ungulates, large-scale geographical variation in phenotypic traits, such as body mass, has been related to environmental conditions and population density, but little is known about the genetic influences. Research on the genetic structure of moose suggests two distinct genetic lineages in Norway, structured along a north-south gradient. This corresponds with many environmental gradients, thus genetic structuring provides an additional factor affecting geographical phenotypic variation in Norwegian moose. We investigated if genetic structure explained geographical variation in body mass in Norwegian moose while accounting for environmental conditions, age and sex, and if it captured some of the variance in body mass that previously was attributed to environmental factors. Genetic structuring of moose was the most important variable in explaining the geographic variation in body mass within age and sex classes. Several environmental variables also had strong explanatory power, related to habitat diversity, environmental seasonality and winter harshness. The results suggest that environmental conditions, landscape characteristics, and genetic structure should be evaluated together when explaining large-scale patterns in phenotypic characters or life history traits. However, to better understand the role of genetic and environmental effects on phenotypic traits in moose, an extended individual-based study of variation in fitness-related characters is needed, preferably in an area of convergence between different genetic lineages.

Morph-specific and sex-specific temperature effects on morphology in the colour polymorphic damselfly Ischnura elegans

Colour polymorphic species with extensive ranges often exhibit large-scale geographic patterns of morph frequency variation. Because colour polymorphism is associated with correlated differences in multiple traits, such as thermal performance, a likely proximate explanation for such patterns is morph-specific responses to temperature variation. The colour polymorphic Blue-tailed damselfly Ischnura elegans exhibits large-scale geographic variation in morph frequencies, but the possibility that temperature is a proximate explanation for the latitudinal cline in morph frequencies has only ever been tested within a single developmental stage (egg survival and hatching time), where no difference between the morphs was found. I therefore carried out a temperature manipulation on larvae of I. elegans which I raised to maturity in the laboratory. I found that individuals exhibited incomplete compensatory growth after being exposed to cold temperatures, and that individuals which did not emerge successfully and those that experienced cold temperatures had more juvenile morphology in the last instar. In addition, there were sex-specific and morph-specific effects of temperature on adult morphology, such that sexual size dimorphism was increased when individuals experienced warm temperatures throughout the larval stage, and that cold temperatures tended to result in larger size of androchromes and their offspring compared to the other morphs. These results are generally consistent with the large-scale geographic variation in morph frequencies found in this species.

Behaviour and physiology shape the growth accelerations associated with predation risk, high temperatures and southern latitudes in Ischnura damselfly larvae

1. To better predict effects of climate change and predation risk on prey animals and ecosystems, we need studies documenting not only latitudinal patterns in growth rate but also growth plasticity to temperature and predation risk and the underlying proximate mechanisms: behaviour (food intake) and digestive physiology (growth efficiency). The mechanistic underpinnings of predator-induced growth increases remain especially poorly understood. 2. We reared larvae from replicated northern and southern populations of the damselfly Ischnura elegans in a common garden experiment manipulating temperature and predation risk and quantified growth rate, food intake and growth efficiency. 3. The predator-induced and temperature-induced growth accelerations were the same at both latitudes, despite considerably faster growth rates in the southern populations. While the higher growth rates in the southern populations and the high rearing temperature were driven by both an increased food intake and a higher growth efficiency, the higher growth rates under predation risk were completely driven by a higher growth efficiency, despite a lowered food intake. 4. The emerging pattern that higher growth rates associated with latitude, temperature and predation risk were all (partly or completely) mediated by a higher growth efficiency has two major implications. First, it indicates that energy allocation trade-offs and the associated physiological costs play a major role both in shaping large-scale geographic variation in growth rates and in shaping the extent and direction of growth rate plasticity. Secondly, it suggests that the efficiency of energy transfer in aquatic food chains, where damselfly larvae are important intermediate predators, will be higher in southern populations, at higher temperatures and under predation risk. This may eventually contribute to the lengthening of food chains under these conditions and highlights that the prey identity may determine the influence of predation risk on food chain length.

Biogeographical Survey Identifies Consistent Alternative Physiological Optima and a Minor Role for Environmental Drivers in Maintaining a Polymorphism

The contribution of adaptive mechanisms in maintaining genetic polymorphisms is still debated in many systems. To understand the contribution of selective factors in maintaining polymorphism, we investigated large-scale (>1000 km) geographic variation in morph frequencies and fitness-related physiological traits in the damselfly Nehalennia irene. As fitness-related physiological traits, we investigated investment in immune function (phenoloxidase activity), energy storage and fecundity (abdomen protein and lipid content), and flight muscles (thorax protein content). In the first part of the study, our aim was to identify selective agents maintaining the large-scale spatial variation in morph frequencies. Morph frequencies varied considerably among populations, but, in contrast to expectation, in a geographically unstructured way. Furthermore, frequencies co-varied only weakly with the numerous investigated ecological parameters. This suggests that spatial frequency patterns are driven by stochastic processes, or alternatively, are consequence of highly variable and currently unidentified ecological conditions. In line with this, the investigated ecological parameters did not affect the fitness-related physiological traits differently in both morphs. In the second part of the study, we aimed at identifying trade-offs between fitness-related physiological traits that may contribute to the local maintenance of both colour morphs by defining alternative phenotypic optima, and test the spatial consistency of such trade-off patterns. The female morph with higher levels of phenoloxidase activity had a lower thorax protein content, and vice versa, suggesting a trade-off between investments in immune function and in flight muscles. This physiological trade-off was consistent across the geographical scale studied and supports widespread correlational selection, possibly driven by male harassment, favouring alternative trait combinations in both female morphs.

Combination of local selection pressures drives diversity in aposematic signals

The diversity of aposematic signals is one of the most difficult phenomena for understanding the evolution of such signals because aposematic animals are most effectively protected when they are common. Theoretical and experimental studies predict that a combination of local selection pressures could maintain variation in aposematic signals. However, the application of this hypothesis to large-scale geographic variation in aposematic signals, other than mimicry systems, is yet to be tested empirically. I investigated geographic variation in morphological and behavioural aposematic signals of the newts, Cynops pyrrhogaster, and in predation pressures on them in populations ranging over 800 km of latitude. Field experiments demonstrated that local differences in predation pressures explain well the island-mainland variation in the aposematic colouration and behaviour of newts. Furthermore, I found a latitudinal gradient in aposematic colouration but not in behaviour, independent of predation pressures. The results suggested that island-mainland variation in aposematic signals resulting from local differences in predation pressures might also be shaped by several factors, such as temperature, body size variation, and genetic differences, and such factors might act on each aposematic trait differently.

An index of unhealthy lifestyle is associated with coronary heart disease mortality rates for small areas in England after adjustment for deprivation

Indices of socio-economic deprivation are often used as a proxy for differences in the health behaviours of populations within small areas, but these indices are a measure of the economic environment rather than the health environment. Sets of synthetic estimates of the ward-level prevalence of low fruit and vegetable consumption, obesity, raised blood pressure, raised cholesterol and smoking were combined to develop an index of unhealthy lifestyle. Multi-level regression models showed that this index described about 50% of the large-scale geographic variation in CHD mortality rates in England, and substantially adds to the ability of an index of deprivation to explain geographic variations in CHD mortality rates.