A theoretical analysis of the morph frequency variation in the peppered moth over England and Wales

The migration-selection model for the spatial and temporal variation of morph frequencies of Biston betularia over England and Wales (Cook & Mani, 1980) has been extended to include effects due to non-visual selection. The parameters for non-visual selection were chosen from the recent determination by Mani (1980) and by Creed et al. (1980). The morph frequencies over England and Wales were obtained through computer simulation and the results were compared with data along the Manchester-Yorkshire, Central Wales-East Anglia and South Wales-London transects. Best fits to the data were obtained by using the non-visual selective values of Mani for carbonaria and modified values of Creed et al. for insularia. It is concluded that the observed polymorphism could be well explained through a balance of migration and visual and non-visual selections.     

A bird’s eye view of the peppered moth

Industrial melanism in Biston betularia is one of the best known examples of the role of natural selection in evolution and has received considerable scrutiny for many years. The rise in frequency of the dark form of the moth (carbonaria) and a decrease in the pale form (typica) was the result of differential predation by birds, the melanic form being more cyptic than typica in industrial areas where the tree bark was darkened by air pollution. One important aspect of early work evaluating the relative crypsis of the forms of B. betularia on tree trunks with different lichen flora was the reliance on human observers. Humans, however, do not have the same visual capabilities as birds. Birds have well‐developed ultraviolet (UV) vision, an important component of their colour processing system that affects many aspects of behaviour, including prey detection. We examined the UV characteristics of the two forms of B. betularia and a number of foliose and crustose lichens. In human visible light the speckled form typica appeared cyptic when seen against a background of foliose lichen, whereas the dark form carbonaria was conspicuous. Under UV light the situation was reversed. The foliose lichens absorbed UV and appeared dark as did carbonaria. Typica, however, reflected UV and was conspicuous. Against crustose lichens, typica was less visible than carbonaria in both visible and UV light. These findings are considered in relation to the distribution and recolonization of trees by lichens and the resting behaviour of B. betularia.     

Viola rupestris: molecular analyses to elucidate postglacial migration in Western Europe

Aim Our aim was to elucidate the postglacial migration of Viola rupestris F.W. Schmidt, with its two subspecies, rupestris and relicta Jalas, in Europe, using molecular methods. Location Norway, Sweden, Finland, England, the Netherlands, France and Switzerland. Methods Isoenzymes were analysed from 49 populations of Viola rupestris s.lat. Based on former experience of variation in the taxon, the following were included: aspartate aminotransferase, leucine/aminopeptidase, and phosphoglucomutase, yielding a total of nine interpretable and six variable (putative) loci. Results Six multilocus phenotypes were revealed (A–F). Phenotype A (only one individual) is restricted to northern Norway, B is shared by all populations in northern Norway and an isolated population on the west coast of southern Norway (Møre). Phenotype B comes close to C, co‐occurring with D in England, the latter also occurring in southern France. Phenotypes E and F occur in the rest of Europe. Main conclusions Viola rupestris ssp. relicta Jalas, previously known only from northern Scandinavia (northern unicentric), was discovered on the west coast of southern Norway (Møre), making the taxon ‘bicentric’. Populations are connected through a putative migration route from the Grands Causses in southern France (Massif Central) via England and western Norway to northern Scandinavia. The Grands Causses might represent a glacial refugium.     

Linkage map of the peppered moth,Biston betularia (Lepidoptera,Geometridae): a model of industrial melanism

We have constructed a linkage map for the peppered moth (Biston betularia), the classical ecological genetics model of industrial melanism, aimed both at localizing the network of loci controlling melanism and making inferences about chromosome dynamics. The linkage map, which is based primarily on amplified fragment length polymorphisms (AFLPs) and genes, consists of 31 linkage groups (LGs; consistent with the karyotype). Comparison with the evolutionarily distant Bombyx mori suggests that the gene content of chromosomes is highly conserved. Gene order is conserved on the autosomes, but noticeably less so on the Z chromosome, as confirmed by physical mapping using bacterial artificial chromosome fluorescence in situ hybridization (BAC-FISH). Synteny mapping identified three pairs of B. betularia LGs (11/29, 23/30 and 24/31) as being orthologous to three B. mori chromosomes (11, 23 and 24, respectively). A similar finding in an outgroup moth (Plutella xylostella) indicates that the B. mori karyotype (n=28) is a phylogenetically derived state resulting from three chromosome fusions. As with other Lepidoptera, the B. betularia W chromosome consists largely of repetitive sequence, but exceptionally we found a W homolog of a Z-linked gene (laminin A), possibly resulting from ectopic recombination between the sex chromosomes. The B. betularia linkage map, featuring the network of known melanization genes, serves as a resource for melanism research in Lepidoptera. Moreover, its close resemblance to the ancestral lepidopteran karyotype (n=31) makes it a useful reference point for reconstructing chromosome dynamic events and ancestral genome architectures. Our study highlights the unusual evolutionary stability of lepidopteran autosomes; in contrast, higher rates of intrachromosomal rearrangements support a special role of the Z chromosome in adaptive evolution and speciation.     

Allozymic Differentiation Among Geographically Distant Populations of Patella vulgata (Mollusca, Patellogastropoda)

Patella vulgata is a boreal cold temperate species and is the dominant limpet in northern Europe. Few works have focussed on the population genetics of this species. Therefore, the aim of this work was to assess the degree of genetic and morphological differentiation of P. vulgata on a macroscale by using 20 allozyme loci and 6 morphological variables. Samples were taken from the following locations: Dingle Peninsula (Southwest Ireland), Port Erin (Southwest Isle of Man), St. Bees Head (north Cumbria, England), St. Agnes Head (north Cornwall, England), Cellar Beach (south Devon, England), Whitley Bay (north Newcastle-Upon-Tyne, England), Sines (Portugal), and Pointe de Chanchardon, La Rochelle (Bay of Biscay, France). Morphological variables were analysed by the multivariate Canonical discriminant analysis. Genetic variation was assessed by diversity measures such as polymorphism and heterozygosity; genetic subdivision of P. vulgata population was determined by the estimator θ of F _ST, and the genetic similarity between populations was measured by Nei’s genetic identity. No significant morphological differentiation was observed among samples. Moderate genetic population subdivision was observed (θ = 0.137±0.074) despite great geographic distances. The minimum genetic identity observed was between Ireland and France (I = 0.942) and maximum was observed between Portugal and north-east England (I=0.998). Two main groups were shown by UPGMA cluster analysis (I = 0.965). One formed by Irish, Manx, north Cumbria, and curiously, south Devon samples, while the second includes Portuguese, French, north-Newcastle-upon-thyne, and north Cornwall samples. No association (g = 0.956; p>0.050) was found between pair-wise genetic divergence and geographic distance separating subpopulations, mainly due to an unexpected pattern of genetic heterogeneity found in Southwest England.     

Variations in Black Grouse Tetrao tetrix winter survival in a year with prolonged snow cover

Capsule Low Black Grouse survival rate in northern England during a severe winter with prolonged snow was attributed to limited availability and proximity of woodland.

Aims To compare the impact of the severe winter weather on Black Grouse in an open treeless landscape in northern England with more wooded landscapes in Scotland.

Methods We assessed the impact of severe winter conditions in relation to previous milder winters at a sample of leks counted annually in northern England and the Scottish Highlands between 2001 and 2012. In addition, following the severe winter in 2009/10 Black Grouse were surveyed at all leks in 2010 where they were surveyed the previous year. We assessed woodland habitat extent within 1 km of leks and related this to indices of Black Grouse over-winter survival.

Results Black Grouse in northern England were severely affected by the severe winter of 2009/10, declining by 38% in comparison to a 23% increase in the Scottish Highlands. Leks in northern England were at higher altitudes and had less tree cover than those in Scotland.

Conclusion The lack of woodland cover around leks in northern England implies that availability of above snow forage, shelter and cover from predators was too low for Black Grouse to survive during prolonged deep snow. Measures are required to provide emergency food sources in periods of prolonged snow, such as the provision of supplementary food at leks, the cultivation of seed-rich arable crops, and the establishment of pockets of woodland.     

North and South: Exploring isotopic analysis of bone carbonates and collagen to understand post-medieval diets in London and northern England

Objectives

We evaluate the potential of paired isotopic analysis of bone carbonate and collagen to examine the diet of post-medieval human and animal populations from England (17th–19th c.), including, for the first time, manufacturing towns in northern England. The potential for identifying C₄ crop consumption is explored alongside regional and local patterning in diet by sex and socioeconomic status.

Materials and Methods

Humans (n = 216) and animals (n = 168) were analyzed from sites in London and northern England for both carbon and nitrogen isotopes of bone collagen (𝛿¹³C_coll, 𝛿¹⁵N_coll). Isotopic analysis of bone carbonates (𝛿¹³C_carb, 𝛿¹⁸O_carb) was carried out on all humans and 27 animals, using Fourier transform infrared spectroscopy–attenuated total reflectance to assess diagenesis.

Results

Variations in diet were observed between and within different populations by geographical location and socioeconomic status. Three pigs and one cow consumed C₄ resources, indicating the availability of C₄-fed animal protein. Londoners consumed more animal and marine protein and C₄ resources. Middle- and upper-class populations from both London and northern populations also had greater access to these foods compared to those of lower status in the same regions.

Discussion

This substantial multi-isotope dataset deriving from bone carbonate and collagen combined from diverse post-medieval urban communities enabled, for the first time, the biomolecular identification of the dynamics of C₄ consumption (cane sugar/maize) in England, providing insight into the dynamics of food globalization during this period. We also add substantially to the animal dataset for post-medieval England, providing further insight into animal management during a key moment of agricultural change.

     

New and resurgent insect pests on low trellis hops

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No Evidence for Resistance to Fenbendazole in Trichostrongylus tenuis,a Nematode Parasite of the Red Grouse

ABSTRACT The parasitic nematode Trichostrongylus tenuis has a detrimental effect on red grouse (Lagopus lagopus scoticus) at the individual and population levels. Treatment using grit coated with the anthelmintic fenbendazole hydrochloride reduces parasite infection and increases grouse density. However, a frequent and low dose of anthelmintic increases selection pressure for parasite resistance, a serious practical and economic problem. We used an egg hatch assay to test resistance of T. tenuis from 12 moors in northern England, which differed in grit treatment intensity. The anthelmintic concentration that prevented 50% and 95% of T. tenuis eggs from hatching (ED₅₀ and ED₉₅, respectively) did not differ among moors and were not related to treatment. We suggest annual monitoring and responsible anthelmintic use to prevent resistance so that medicated grit continues to enhance red grouse management.     

Projecting the distribution of forests in New England in response to climate change

Aim To project the distribution of three major forest types in the northeastern USA in response to expected climate change. Location The New England region of the United States. Methods We modelled the potential distribution of boreal conifer, northern deciduous hardwood and mixed oak–hickory forests using the process‐based BIOME4 vegetation model parameterized for regional forests under historic and projected future climate conditions. Projections of future climate were derived from three general circulation models forced by three global warming scenarios that span the range of likely anthropogenic greenhouse gas emissions. Results Annual temperature in New England is projected to increase by 2.2–3.3 °C by 2041–70 and by 3.0–5.2 °C by 2071–99 with corresponding increases in precipitation of 4.7–9.5% and 6.4–11.4%, respectively. We project that regional warming will result in the loss of 71–100% of boreal conifer forest in New England by the late 21st century. The range of mixed oak–hickory forests will shift northward by 1.0–2.1 latitudinal degrees (c. 100–200 km) and will increase in area by 149–431% by the end of the 21st century. Northern deciduous hardwoods are expected to decrease in area by 26% and move upslope by 76 m on average. The upslope movement of the northern deciduous hardwoods and the increase in oak–hickory forests coincide with an approximate 556 m upslope retreat of the boreal conifer forest by 2071–99. In our simulations, rising atmospheric CO₂ concentrations reduce the losses of boreal conifer forest in New England from expected losses based on climatic change alone. Main conclusion Projected climate warming in the 21st century is likely to cause the extensive loss of boreal conifer forests, reduce the extent of northern hardwood deciduous forests, and result in large increases of mixed oak–hickory forest in New England.     

GENETIC AND MORPHOMETRIC DIVERGENCE IN ANCESTRAL EUROPEAN AND DESCENDENT NEW ZEALAND POPULATIONS OF CHAFFINCHES (FRINGILLA COELEBS)

Descendent populations of chaffinches (Fringilla coelebs) introduced to New Zealand about 120 years ago were compared with “ancestral” populations in northern Europe and with those in a broader region of Europe (including Iberia) using protein electrophoresis at 42 loci and 12 skeletal measurements. The New Zealand populations exhibit very small scale differentiation in genetics (F_st = 0.040) and morphometrics, and the haphazard pattern of among-population variation does not align with environmental variation nor is it predicted by the geographic proximity of populations. Thus random drift is implicated in the differentiation among the descendent populations. The New Zealand chaffinches have diverged only slightly in morphometrics from an extant population in southern England, and constant heritability rate tests suggest that random drift alone could account for this small shift. In sharp contrast, the European populations are subdivided genetically (F_st = 0.222) and morphometrically, and this subdivision coincides with the Pyrenees mountains between Iberia and northern Europe which act as a barrier to gene flow between these regions. Iberian populations have smaller skulls and longer wings on average than northern European populations and are characterized by high frequencies of alternative common alleles at Ada and Np. Within both the Iberian and northern European regions, however, populations are effectively panmictic in protein-encoding genes, indicating that homogenizing gene flow is apparently extensive enough to prevent among-population differentiation in allozymes by drift. Variation in body size as represented by PC I is related to environmental productivity across Europe, unlike in New Zealand. These observations jointly suggest that longer term adaptive differentiation via selection for optimal body size has evolved in Europe. Because multilocus evolution is expected to proceed slowly in populations subject to the opposing forces of selection and homogenizing gene flow, I argue that local adaptation within “ancestral” populations in northern Europe may still be evolving.     

Population‐level manipulations of field vole densities induce subsequent changes in plant quality but no impacts on vole demography

Grazing‐induced changes in plant quality have been suggested to drive the negative delayed density dependence exhibited by many herbivore species, but little field evidence exists to support this hypothesis. We tested a key premise of the hypothesis that reciprocal feedback between vole grazing pressure and the induction of anti‐herbivore silicon defenses in grasses drives observed population cycles in a large‐scale field experiment in northern England. We repeatedly reduced population densities of field voles (Microtus agrestis) on replicated 1‐ha grassland plots at Kielder Forest, northern England, over a period of 1 year. Subsequently, we tested for the impact of past density on vole life history traits in spring, and whether these effects were driven by induced silicon defenses in the voles’ major over‐winter food, the grass Deschampsia caespitosa. After several months of density manipulation, leaf silicon concentrations diverged and averaged 22% lower on sites where vole density had been reduced, but this difference did not persist beyond the period of the density manipulations. There were no significant effects of our density manipulations on vole body mass, spring population growth rate, or mean date for the onset of spring reproduction the following year. These findings show that grazing by field voles does induce increased silicon defenses in grasses at a landscape scale. However, at the vole densities encountered, levels of plant damage appear to be below those needed to induce changes in silicon levels large and persistent enough to affect vole performance, confirming the threshold effects we have previously observed in laboratory‐based studies. Our findings do not support the plant quality hypothesis for observed vole population cycles in northern England, at least over the range of vole densities that now prevail here.     

Evolution of flight morphology in a butterfly that has recently expanded its geographic range

Individuals colonizing unoccupied habitats typically possess characters associated with increased dispersal and, in insects, colonization success has been related to flight morphology. The speckled wood butterfly, Pararge aegeria, has undergone recent major expansions in its distribution: in the north of its range, P. aegeria has colonized many areas in north and east England, and in the south, it was first recorded on Madeira in 1976. We examined morphological traits associated with flight and reproduction in the northern subspecies tircis, and in the southern subspecies aegeria, from sites colonized about 20 years ago in northern England and on Madeira, respectively. Investment in flight was measured as relative wing area and thorax mass, and investment in reproduction as relative abdomen mass. All measurements were from individuals reared in a common environment and there were significant family effects in most of the variables measured. Compared with individuals from sites continuously occupied in recent history, colonizing individuals were larger (adult live mass). In the subspecies tircis, colonizing individuals also had relatively larger thoraxes and lower wing aspect ratios indicating that evolutionary changes in flight morphology may be related to colonization. However, sex by site interactions in analyses of thorax mass and abdomen mass suggest different selection pressures on flight morphology between the sexes in relation to colonization. Overall, the subspecies aegeria was smaller (adult live mass) and had a relatively larger thorax and wings, and smaller abdomen than subspecies tircis. Evolutionary changes in flight morphology and dispersal rate may be important determinants of range expansion, and may affect responses to future climate change. Received: 1 March 1999 / Accepted: 30 June 1999     

Trends in abundance of rocky intertidal seaweeds and filter feeders across gradients of elevation,wave exposure,and ice scour in eastern Canada

On NW Atlantic rocky shores, the main basal organisms in intertidal communities are seaweeds (Ascophyllum nodosum, Fucus spp. and Chondrus crispus) and filter feeders (barnacles, Semibalanus balanoides, and mussels, Mytilus spp.). Their ecology has been extensively studied in New England (United States), but knowledge gaps exist for northern shores, which are subjected to stronger environmental stress. Therefore, we studied the above organisms on Canadian shores. We quantified the summer abundance of these seaweeds and filter feeders across full vertical (intertidal elevation) and horizontal (wave exposure and winter ice scour) environmental gradients on the Gulf of St. Lawrence and open Atlantic coasts of Nova Scotia. At the regional scale along the open Atlantic coast, seaweeds showed similar abundances in Nova Scotia than values reported for New England. However, both filter feeders were considerably less abundant in Nova Scotia. At the local scale in Nova Scotia, intense winter ice scour (which only occurs on the Gulf of St. Lawrence coast) was associated with a very low abundance of all species except barnacles. Spatial trends in Nova Scotia were similar to patterns known for certain species elsewhere, such as A. nodosum being almost restricted to sheltered habitats, regardless of elevation, and C. crispus being almost restricted to low elevations, regardless of exposure. Other trends were, however, characteristic of Nova Scotia, such as C. crispus being frequent at low elevations in exposed habitats, unlike in New England, where mussels often predominate there because of competitive advantages. In Nova Scotia, mussels were always restricted to cracks and crevices, unlike in New England, where they form extensive intertidal beds on exposed shores. The direct effects of increased environmental stress and indirect effects through altered interspecific interactions might explain the regional differences in local species distribution, which will require experimental validation. Handling editor: K. Martens     

Insect melanism: the molecules matter

Insect melanism, especially in the peppered moth Biston betularia, has long been a textbook case of evolution in action. Hypotheses of the role of natural selection in maintaining melanic polymorphisms have implicated a wide range of explanations in various species, but to understand fully the ecology of melanism, we need to understand its molecular and developmental genetic basis. Because developmental genes often affect more than one trait, identifying the genes responsible for melanism is crucial for a thorough understanding of the fitnesses and selective responses of melanic alleles in nature. Molecular genetic information is also vital for elucidating the evolutionary history and possible mechanistic diversity of melanism among species. Recent studies of the developmental genetics of melanin pigmentation in Drosophila, and of the genetics of pigmentation differences among other insect species, have provided valuable insights into the underpinnings of this important source of polymorphism throughout the Insecta.     

Using stable isotopes to link breeding population trends to winter ecology in Willow Warblers,Phylloscopus trochilus

Capsule Stable isotope composition of feathers grown in Africa indicates that Willow Warblers, Phylloscopus trochilus, breeding in areas of Britain experiencing different population trends may vary in their distribution, timing of moult and/or use of prey resources during the non-breeding season.

Aims To compare stable isotope ratios of feathers of Willow Warblers breeding in parts of Britain with differing rates of population change.

Methods Feathers were collected from three regions with differing recent population trends; northern Scotland (population increased), western Wales (declined slightly) and eastern England (declined rapidly). Collection occurred at ringing sites run by British Trust for Ornithology (BTO) volunteers as part of the Constant Effort Sites (CES) scheme between April and July in 2008 and 2009.

Results Willow Warblers from northern Scotland had significantly higher δ ¹⁵N and δ ¹³C values than those from eastern England, although the regional mean values did not differ greatly. There was no significant difference in δ ¹⁵N and δ ¹³C values between the sexes, but δ ¹⁵N values were significantly greater in individuals caught later in the breeding season.

Conclusions During the premigratory moult period in Africa, Willow Warblers from different parts of Britain may vary in their distribution, timing of moult and/or use of prey resources.     

Distribution and ecology of small cow-wheat (Melampyrum sylvaticum L.; Scrophulariaceae) in the British Isles

Summary

Melampyrum sylvaticum L. is an annual, hemiparasitic herb which mainly grows in light woodland and on upland ledges in northern Britain and Ireland. Its distribution is poorly known due to confusion with the variable M. pratense. A review of its ecology and distribution has been carried out and it has now been recorded from 83 10-km squares in the British Isles, including three in England and two in Wales; an updated distribution map is given. It is probably declining slowly in the lowlands at the margins of its range due to afforestation, development of dense shade, over-grazing and intensive agriculture.     

A low-density single nucleotide polymorphism panel for brown trout (Salmo trutta L.) suitable for exploring genetic diversity at a range of spatial scales

The rivers of southern England and northern France which drain into the English Channel contain several genetically unique groups of trout (Salmo trutta L.) that have suffered dramatic declines in numbers over the past 40 years. Knowledge of levels and patterns of genetic diversity is essential for effective management of these vulnerable populations. Using restriction site-associated DNA sequencing (RADseq) data, we describe the development and characterisation of a panel of 95 single nucleotide polymorphism (SNP) loci for trout from this region and investigate their applicability and variability in both target (i.e., southern English) and non-target trout populations from northern Britain and Ireland. In addition, we present three case studies which demonstrate the utility and resolution of these genetic markers at three levels of spatial separation:(a) between closely related populations in nearby rivers, (b) within a catchment and (c) when determining parentage and familial relationships between fish sampled from a single site, using both empirical and simulated data. The SNP loci will be useful for population genetic and assignment studies on brown trout within the UK and beyond.     

Influence of genotype and geography on shell shape and morphometric trait variation among North Atlantic blue mussel (Mytilus spp.) populations

The influence of geography and genotype on shell shape (outline) and trait (morphometric) variation among North Atlantic blue mussels and their hybrids has been examined. Shape differences among reference taxa (Mytilus trossulus, Mytilus edulis and Mytilus galloprovincialis) were consistent with an association between taxon‐specific genes and shape genes. Newfoundland M. edulis × M. trossulus populations and northern Quebec M. trossulus populations exhibited an uncoupling of taxon‐specific genes from shape genes, whereas Nova Scotia M. trossulus populations and SW England M. edulis × M. galloprovincialis populations exhibited an association between taxon‐specific genes and shape genes. We found no evidence of a geographic effect (NE versus NW Atlantic) for shape variation, indicating that the genotype effect is stronger than any geographic effect at macrogeographic scales. Pronounced differences were observed in trait variability consistent with an association between taxon‐specific genes and trait genes in European populations, and trait divergence of New York M. edulis from all European mussels. Trait variability in mussels from Newfoundland, Nova Scotia and northern Quebec indicated an uncoupling of taxon genes from trait genes, whereas trait variability in SW England M. edulis × M. galloprovincialis populations was consistent with background genotype, indicating a strong association between taxon genes and trait genes. A pronounced macrogeographic split (NE versus NW Atlantic) regardless of taxonomy was observed, indicating that geography exerts a greater influence than genotype on trait variation at the macrogeographic scale. This is consistent with pronounced within‐taxon genetic divergence, indicative of different selection regimes or more likely of different evolutionary histories of mussels on either side of the North Atlantic. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 96 , 875–897.     

Delayed density‐dependent onset of spring reproduction in a fluctuating population of field voles

Delayed density‐dependent demographic processes are thought to be the basis for multi‐annual cyclic fluctuations in small rodent populations, but evidence for delayed density dependence of a particular demographic trait is rare. Here, using capture–recapture data from 22 sites collected over nine years, we demonstrate a strong effect of population density with a one‐year lag on the timing of the onset of spring reproduction in a cyclically fluctuating population of field voles Microtus agrestis in northern England. The mean date for the onset of spring reproduction was delayed by about 24 days for every additional 100 voles ha^?1 in the previous spring. This delayed density dependence is sufficient to generate the type of cyclic population dynamics described in the study system.