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Background Climate change due to anthropogenic global warming is the most important factor that will affect future range distribution of species and will shape future biogeographic patterns. While much effort has been expended in understanding how climate change will affect rare and declining species we have less of an understanding of the likely consequences for some abundant species. The Common Grackle (Quiscalus quiscula; Linnaeus 1758), though declining in portions of its range, is a widespread blackbird (Icteridae) species in North America east of the Rocky Mountains. This study examined how climate change might affect the future range distribution of Common Grackles. Methods We used the R package Wallace and six general climate models (ACCESS1-0, BCC-CSM1-1, CESM1-CAM5-1-FV2, CNRM-CM5, MIROC-ESM, and MPI-ESM-LR) available for the future (2070) to identify climatically suitable areas, with an ecological niche modelling approach that includes the use of environmental conditions. Results Future projections suggested a significant expansion from the current range into northern parts of North America and Alaska, even under more optimistic climate change scenarios. Additionally, there is evidence of possible future colonization of islands in the Caribbean as well as coastal regions in eastern Central America. The most important bioclimatic variables for model predictions were Annual Mean Temperature, Temperature Seasonality, Mean Temperature of Wettest Quarter and Annual Precipitation. Conclusions The results suggest that the Common Grackle could continue to expand its range in North America over the next 50 years. This research is important in helping us understand how climate change will affect future range patterns of widespread, common bird species.

Urbanization can change interactions in insect communities, and the few studies of tritrophic interactions in urban settings focus on interactions between plants, herbivorous insects and their mutualists and natural enemies. Plant pathogen infection is also widespread and common, and infection may also alter such interactions, but we have no understanding of whether the ecological consequences of pathogen infection vary with urbanization. Using replicated aphid colonies on experimental plants, we investigated how infection by the plant pathogen Botrytis cinerea influences interactions between plants, aphids and the aphid natural enemies and ant mutualists in highly urbanized, suburban and rural study sites. Aphid and natural enemy abundance were highest in the suburban site, while mutualist ants were most abundant in the urban site, reversing the usual positive density-dependent relationship between natural enemies and aphids. The effect of pathogen infection varied with trait and site, mediated by natural enemy preference for hosts or prey on uninfected plants. The effect of infection on aphid abundance was only seen in the suburban site, where natural enemies were most abundant on uninfected plants and aphid numbers were greatest on infected plants. In the urban site, there was no effect of infection, while in the rural site, aphid numbers were lower on infected plants. Uninfected plants were smaller than infected plants and differed between locations. This study suggests that the effects of urbanization on ecological interactions may become more complex and difficult to predict as we study ecological assemblages and communities at greater levels of structural complexity.

Indirect effects are ubiquitous in nature, and have received much attention in terrestrial plant&ndash, insect herbivore&ndash, enemy systems. In such tritrophic systems, changes in plant quality can have consequential effects on the behavior and abundance of insect predators and parasitoids. Plant quality as perceived by insect herbivores may vary for a range of reasons, including because of infection by plant pathogens. However, plant diseases vary in their origin (viral, bacterial or fungal) and as a result may have differing effects on plant physiology. To investigate if the main groups of plant pathogens differ in their indirect effects on higher trophic levels, we performed a meta-analysis using 216 measured responses from 29 primary studies. There was no overall effect of plant pathogens on natural enemy traits as differences between pathogen types masked their effects. Infection by fungal plant pathogens showed indirect negative effects on the performance and preference of natural enemies via both chewing and piercing-sucking insect herbivore feeding guilds. Infection by bacterial plant pathogens had a positive effect on the natural enemies (parasitoids) of chewing herbivores. Infection by viral plant pathogens showed no clear effect, although parasitoid preference may be positively affected by their presence. It is important to note that given the limited volume of studies to date on such systems, this work should be considered exploratory. Plant pathogens are very common in nature, and tritrophic systems provide an elegant means to examine the consequences of indirect interactions in ecology. We suggest that further studies examining how plant pathogens affect higher trophic levels would be of considerable value.

Simple Summary The European mole is a burrowing mammal which is widely distributed across Britain and much of continental Europe. Its presence is readily confirmed by the presence of molehills, which contain the spoil heaps left behind as the mole digs its underground tunnels. Despite being easy to record, there are very few studies of moles in an urban environment. We asked how area of greenspace (largely parks, recreation areas, nature reserves and playing fields), distance to the nearest patch, human disturbance, how long the green patch had been isolated for, and degree of urban construction around the patch affected mole presence and abundance. We found that patch size affected mole presence, with a minimum greenspace of approximately 10 ha required. Where moles were found, larger patches had more signs of moles and surprisingly, mole abundance was also associated with the degree of urban construction around the greenspace. This result shows how urban planning can affect the presence of unusual species, such as the European mole. Abstract The European mole Talpa europaea is common across much of Britain. It has a unique fossorial lifestyle, and evidence of its presence is readily identified through the presence of characteristic molehills. Although molehills are often a common sight in urban greenspaces, moles are remarkably understudied, with very few studies to date exploring the urban ecology of moles. Here, we investigate if factors such as greenspace (largely urban parks and playing fields) area, intensity of management, distance to nearest patch, amount of time the patch had been isolated from other green patches, and the amount of urbanization (constructed surfaces) surrounding the patch, influence the distribution and abundance of urban moles. Mole signs (hills and surface runs) were counted in all discrete urban greenspaces (excluding domestic gardens and one private golf course) within an 89.5 km2 area in the UK town of Reading. We found that 17 out of 59 surveyed sites contained moles, with their presence being recorded in greenspaces with a minimum patch area of approximately 0.1 km2 (10 ha). Where present, the abundance of mole territories in the greenspaces was associated with both the area of greenspace and degree of urbanization within 150 m of the patch boundary. While the former was not surprising, the latter outcome may be a consequence of sites with an increased risk of flooding being home to fewer moles, and the surrounding area is also less likely to be built upon. This case study highlights how choices made in designing urban green infrastructure will determine which species survive in urban areas long into the future.

The modification of habitats in urban areas is thought to alter patterns of species interactions, by filtering specialist species and those at higher trophic levels. However, empirical studies addressing these hypotheses remain limited in scope and number. This work investigates (1) how main urban land uses affect predator-prey and mutualistic interactions, and (2) how specialist and generalist predators respond to size and availability of urban green spaces. In a large town in the UK, experimental colonies of ant-attended Black bean aphid Aphis fabae and non-ant-attended Pea aphid Acyrthosiphon pisum were monitored over two years. Ants were more frequently found in highly urbanised sites; however mutualistic ants were also more often encountered when the habitat was more plant diverse. Aphids were not affected by urban land uses, but A. fabae numbers were positively related to the presence of mutualists, and so indirectly affected by urbanisation. Predators were the only group negatively affected by increased urbanisation, and specialist species were positively related to increased proportion of urban green areas within the habitats. While this work supports the hypothesis that specialist predators are negatively affected by urbanisation, we also show that a fundamental ecological interaction, mutualism, is affected by urbanisation.

Providing food for wild birds is perhaps the most widespread intentional interaction between people and wildlife. In the UK, almost half of households feed wild birds, often as peanuts and seed supplied in hanging feeders. Such food is also taken by the introduced, invasive Grey Squirrel Sciurus carolinensis. Little is known of how Grey Squirrels utilise this resource and how they affect feeder use by wild birds. To assess this we recorded the numbers and time spent by animals visiting experimental feeding stations in suburban gardens, and also asked if exclusionary guards (to prevent Grey Squirrel access), food type (peanut, mixed seed), habitat and weather conditions influenced visits. Using automated cameras, we recorded 24,825 bird and 8577 Grey Squirrel visits. On average >44% of the time feeders were utilised, they were being visited by Grey Squirrels. Grey Squirrel presence prevented birds from feeding at the same time (>99.99%). Feeders where Grey Squirrels were dominant were less likely to be visited by birds, even in their absence. Guards reduced Grey Squirrel use to a minimum on seed feeders, and by approximately half on peanut feeders. Squirrels, food type, guard status, habitat and rainfall all influenced bird activity and timing of feeder visits. Our work suggests that Grey Squirrels reduce the availability of supplementary food to wild birds, while gaining large volumes of food resources with corresponding benefits. Given the ubiquity of supplementary feeding, it is likely that this is an important resource for urban Grey Squirrels; feeder guards mitigate this effect.

Urban gardens can harbour a high diversity of insects, which are critically important components of urban ecosystems. In this work, we investigate the richness and diversity of a major taxon of economic and ecological importance, the aphids (Homoptera: Aphididae), and their main insect predators, the hoverflies (Diptera: Syrphidae) and ladybirds (Coleoptera: Coccinellidae), in urban gardens. We examined how variation in environmental factors associated with urbanisation (garden host plant abundance, garden plant richness, garden size, proportion of impervious surfaces in the surrounding area) directly and indirectly (via prey and predator abundance) influence the local diversity and abundance of aphids, ladybirds, and hoverflies. Sixty-seven domestic gardens located in southern England were surveyed during the peak period of aphid abundance, and the numbers and identity of aphids and their predators were recorded. We observed 45 aphid species (179917 individuals in total), 15 hoverfly species (494 individuals) and 8 ladybird species (173 individuals). We found that aphid species richness and abundance were positively associated with utilised host plant abundance and garden plant species richness. Hoverfly abundance was positively correlated with garden plant richness. The abundance of ladybirds was positively correlated with aphid abundance and garden plant species richness, and negatively associated to the proportion of impervious surfaces in the surrounding environment. The difference in responses between the two major taxa of aphid predators may reflect differences in their behaviour and natural history. Our results indicate that overall increases in urban land cover are not favourable for ladybirds as a group, and that fine scale habitat variables that are determined by garden owners have the potential to greatly affect the general diversity of aphids and their main predators.

Urbanization can alter the composition of arthropod communities. However, little is known about how urbanization affects ecological interactions. Using experimental colonies of the black bean aphid Aphis fabae Scopoli reared on Vicia faba L, we asked if patterns of predator-prey, host-parasitoid and ant-aphid mutualisms varied along an urbanization gradient across a large town in southern England. We recorded the presence of naturally occurring predators, parasitoid wasps and mutualistic ants together with aphid abundance. We examined how biotic (green areas and plant richness) and abiotic features (impervious surfaces and distance to town center) affected (1) aphid colony size, (2) the likelihood of finding predators, mutualistic ants and aphid mummies (indicating the presence of parasitoids), and (3) how the interplay among these factors affected patterns of parasitoid attack, predator abundance, mutualistic interactions and aphid abundance. The best model to predict aphid abundance was the number of mutualistic ants attending the colonies. Aphid predators responded negatively to both the proportion of impervious surfaces and to the number of mutualistic ants farming the colonies, and positively to aphid population size, whereas parasitized aphids were found in colonies with higher numbers of aphids and ants. The number of mutualistic ants attending was positively associated with aphid colony size and negatively with the number of aphid predators. Our findings suggest that for insect-natural enemy interactions, urbanization may affect some groups, while not influencing others, and that local effects (mutualists, host plant presence) will also be key determinants of how urban ecological communities are formed.

Nests are a critically important factor in determining the breeding success of many species of birds. Nevertheless, we have surprisingly little understanding of how the local environment helps determine the materials used in nest construction, how this differs among related species using similar nest sites, or if materials used directly or indirectly influence the numbers of offspring successfully reared. We also have little understanding of any potential links between nest construction and the assemblage of invertebrates which inhabit nests, in particular, ectoparasites. We addressed these questions by monitoring the success rates of Blue Tits Cyanistes caeruleus and Great Tits Parus major, using nest boxes in rural, urban greenspace and urban garden settings. We collected used nests, identified the arthropods present, and measured the proportions of highly processed anthropogenic materials used in nest construction. Twenty-five percent of Great Tit nest materials were of anthropogenic origin and this was consistent across habitats, while Blue Tits used little (1–2%) anthropogenic material except in gardens (~16%), suggesting that Great Tits preferentially sought out these materials. In fledged nests, an increasing use of anthropogenic material was associated with a lower general arthropod diversity and ectoparasite predator abundance (Blue Tits only), but higher levels of Siphonaptera (fleas). Higher arthropod diversity was associated with lower flea numbers, suggesting that increased diversity played a role in limiting flea numbers. No direct link was found between breeding success and either anthropogenic material usage or arthropod diversity and abundance. However, breeding success declined with increasing urbanisation in both species and increased with nest weight in Blue Tits. The interplay between urbanisation and bird ecology is complex; our work shows that subtle anthropogenic influences may have indirect and unexpected consequences for urban birds.

In countries such as the UK, USA and Australia, approximately half of households provide supplementary food for wild birds, making this the public’s most common form of active engagement with nature. Year round supplementary feeding is currently encouraged by major conservation charities in the UK as it is thought to be of benefit to bird conservation. However, little is understood of how the provision of supplementary food affects the behaviour and ecology of target and non-target species. Given the scale of supplementary feeding, any negative effects may have important implications for conservation. Potential nest predators are abundant in urban areas and some species frequently visit supplementary feeding stations. We asked if providing supplementary food affected the likelihood of nest predation in the vicinity of the feeder, by acting as a point attractant for potential nest predators. We provided feeding stations (empty, peanut feeder, peanut feeder with guard to exclude potential nest predators) in an area of suburban parkland in the UK and monitored the predation rate experienced by eggs placed in artificial nests located at distances which replicate the size of typical suburban gardens. Nest predators (Magpies Pica pica, Grey Squirrels Sciurus carolinensis) were frequent visitors to filled feeders, and predation caused by Magpies, European Jays Garrulus glandarius and Grey Squirrels was significantly higher when nests were adjacent to filled feeders. The presence of a feeder guard did not significantly reduce nest predation. As supplementary feeding is becoming increasingly common during the breeding season in suburban habitats, we suggest that providing point attractants to nest predators at this time may have previously unconsidered consequences for the breeding success of urban birds.

Climate variability is one of the most important forces affecting the distributional range dynamics of species and consequentially plays a significant role in shaping biogeographic patterns. This study aims to infer the role of climate in the recent evolutionary history of the Common Grackle Quiscalus quiscula. Studies of other migratory North American birds have shown that their populations were isolated in two or three refugia in southern North America during the Last Glacial Maximum (LGM). In contrast, preliminary genetic work suggest that Common Grackles may have occupied a single refugium during that time. They subsequently became widespread and northern populations evolved highly migratory behaviour. We used an ecological niche modelling approach that involved the use of three general climate models for the past (the LGM, approximately 22000 years before present) and for present environmental conditions to identify climatically stable areas. Extrapolations to the past showed contraction to a large continuous refugium located in the southern part of North America, and projection to the present showed expansion that covers much of eastern and middle North America. The most important bioclimatic variable for model predictions was annual mean temperature, which explained 74% of the variation in the model. Results suggest that the Common Grackle has expanded its distributional range by more than 300% after the LGM.

Myrmecophyte plants house ants within domatia in exchange for protection against herbivores. Ant-myrmecophyte mutualisms exhibit two general patterns due to competition between ants for plant occupancy: (i) domatia nest sites are a limiting resource and (ii) each individual plant hosts one ant species at a time. However, individual camelthorn trees (Vachellia erioloba) typically host two to four ant species simultaneously, often coexisting in adjacent domatia on the same branch. Such fine-grain spatial coexistence brings into question the conventional wisdom on ant-myrmecophyte mutualisms. Camelthorn ants appear not to be nest-site limited, despite low abundance of suitable domatia, and have random distributions of nest sites within and across trees. These patterns suggest a lack of competition between ants for domatia and contrast strongly with other ant-myrmecophyte systems. Comparison of this unusual case with others suggests that spatial scale is crucial to coexistence or competitive exclusion involving multiple ant species. Furthermore, coexistence may be facilitated when co-occurring ant species diverge strongly on at least one niche axis. Our conclusions provide recommendations for future ant-myrmecophyte research, particularly in utilizing multispecies systems to further our understanding of mutualism biology.

Feeding wild bird is popular in domestic gardens across the world, with around half of households in the UK, North America and Australia doing so. Nevertheless, there is surprisingly little empirical information on many aspects of the activity. We sought to characterise garden bird feeding in a large UK urban area in two ways. First, we conduct- ed face-to-face questionnaires with a representative cross-section of residents. Just over half fed birds, the majority doing so year-round and at least weekly. Second, a 2-year, longitudinal study recorded all foodstuffs put out by house- holds on every provisioning occasion. In this way, we obtained the first year-round quantitative records of the amounts and types of wild bird food provided in individual gardens. A median of 127 g, equivalent to 628 kcal, was given daily per household (typically consisting of several food types). We estimated the daily cost of this provisioning level to be UK£0.35 per household based on the relative proportions of each food type. Provisioning level was not significantly influenced by weather or season. Comparisons between the data sets revealed significantly less frequent feeding amongst the feeders in the longitudinal study (assumed to be 'keen' feeders owing to their participation in this long- term study and numbers of food types provided) than the face-to-face questionnaire respondents, suggesting that ques- tionnaires relying upon participants' estimates rather than records of provisioning may overestimate actual provision- ing frequency. Assuming 100% uptake, the median provisioning level equates to sufficient supplementary resources across the UK to fully support 196 million individuals of a hypothetical average garden-feeding bird species (based on 10 common UK garden-feeding birds' energy requirements). This compares with an estimated total of 71 million breed- ing individuals of these 10 species in the UK (non-breeding numbers unknown). Taking the lowest provisioning level recorded (101 kcal/day) as a conservative measure, 31 million of these average individuals could theoretically be sup- ported.

Grass lawns are a ubiquitous feature of urban green-space throughout much of the temperate world. Species poor and intensively managed, lawns are ecologically impoverished, however environmentally aware lawn owners are reluctant to implement alternatives due to aesthetic concerns. Developing an alternative lawn format which is both biodiversity friendly and aesthetically pleasing is an imperative for urban greening. We suggest that such an alternative can be provided by replacing the grass lawn by a forb-based mix. To advance this, we tested the floral performance of three groups of clonal perennial forbs (native, non-native and mixed), each maintained using standard lawn management mowing regimes. Our findings show that both the frequency of mowing and the height at which mowing is applied influence floral performance and lawn aesthetics. Species origin was found to influence floral productivity, floral visibility and floral variety within grass-free lawns, with native species providing the greatest floral performance. The behaviour and management of grass lawns was not found to be a suitable analogue for the management of grass-free lawns and grass-free lawns are sufficiently different from grass lawns to require an entirely original management approach. We suggest that the grass-free lawn can provide an aesthetically and environmentally relevant replacement for the ubiquitous and ecologically poor grass lawn.

Insect diversity may be declining even more rapidly than in plants and vertebrates, particularly in areas where indigenous habitats are replaced by an anthropogenic one. The most common component of anthropogenic greenspace is the ornamental lawn. Intensively managed and offering limited habitat opportunities for both plants and insects, lawns are biodiversity poor and ecologically insensitive. An alternative lawn format that positively influences biodiversity and reduces management requirements would be a useful tool in eco-friendly urban greenspace management. In investigating the potential for a forb-only alternative to the grass lawn we sampled both trial grassfree lawn formats and turf lawns to identify any influence that lawn composition and grass-free lawn specific mowing regimes might have on the abundance and diversity of insect families. In addition to the mowing regimes, both the composition and origin of lawn flora were found to significantly influence insect abundance and diversity and these factors rarely interacted. Native-only and mixed origin grass-free lawns hosted greater numbers of adult insects than found in turf and an equivalent diversity of insect families, however the mowing regime applied was distinct from traditional turf lawn management by being substantially less intensive and our results suggest that there is the potential for even greater abundance and diversity via the grass-free format that may offer additional resources to insectivorous garden species such as birds. When the composition of grass-free lawns included native forbs the diversity of insect families was found be sufficiently different from turf lawns to form distinct assemblages and in so doing contribute to beta diversity within urban greenspace. In sum, grass-free lawns may be a useful and aesthetically appropriate tool for adding value to the generally biodiversity poor urban lawnscape.

The grass-free lawn is a novel development in modern ornamental horticulture where the traditional monoculture of grass is replaced by a variety of mowing-tolerant clonal forbs. It brings floral aesthetics and a diverse species approach to the use of lawn space. How the number of constituent forb species affects the aesthetic and structural performance of grass-free lawns was investigated using grass-free lawns composed of four, six and twelve British native clonal perennial forb species. Lawn productivity was seen to increase with increasing species number, but the relationship was not linear. Plant cover was dynamic in all lawn types, varied between years and was not representative of individual species’ floral performance. The behaviour of component species common to all lawns suggested that lawns with 12 species show greater structural stability than lawns with lower species number. Visual performance in lawns with the greatest species number was lower than in lawns with fewer species, with increasing variety in floral size and individual species floral productivity, leading to a trade-off between diversity and floral performance. Individual species were seen to have different aesthetic functions in grass-free lawns by providing flowers, ground coverage or both.

In Britain, managed grass lawns provide the most traditional and widespread of garden and landscape practices in use today. Grass lawns are coming under increasing challenge as they tend to support a low level of biodiversity and can require substantial additional inputs to maintain. Here we apply a novel approach to the traditional monocultural lawnscape by replacing grasses entirely with clonal perennial forbs. We monitored changes in plant coverage and species composition over a two year period and here we report the results of a study comparing plant origin (native, non-native and mixed) and mowing regime. This allows us to assess the viability of this construct as an alternative to traditional grass lawns. Grass-free lawns provided a similar level of plant cover to grass lawns. Both the mowing regime and the combination of species used affected this outcome, with native plant species seen to have the highest survival rates, and mowing at 4 cm to produce the greatest amount of ground coverage and plant species diversity within grass-free lawns. Grass-free lawns required over 50 percent less mowing than a traditionally managed grass lawn. Observations suggest that plant forms that exhibited: (a) a relatively fast growth rate, (b) a relatively large individual leaf area, and (c) an average leaf height substantially above the cut to be applied, were unsuitable for use in grass-free lawns. With an equivalent level of ground coverage to grass lawns, increased plant diversity and a reduced need for mowing, the grass-free lawn can be seen as a species diverse, lower input and potentially highly ornamental alternative to the traditional lawn format.

Nest site selection in arboreal, domatia-dwelling ants, particularly those coexisting on a single host plant, is little understood. To examine this phenomenon we studied the African savannah tree Vachellia erioloba, which hosts ants in swollen-thorn domatia. We found four ant species from different genera (Cataulacus intrudens, Tapinoma subtile, Tetraponera ambigua and an unidentified Crematogaster species). In contrast to other African ant plants, many V. erioloba trees (41 % in our survey) were simultaneously co-occupied by more than one ant species. Our study provides quantitative field data describing: (1) aspects of tree and domatia morphology relevant to supporting a community of mutualist ants, (2) how ant species occupancy varies with domatia morphology and (3) how ant colony size varies with domatia size and species. We found that Crematogaster sp. occupy the largest thorns, followed by C. intrudens, with T. subtile in the smallest thorns. Thorn age, as well as nest entrance hole size correlated closely with ant species occupant. These differing occupancy patterns may help to explain the unusual coexistence of three ant species on individual myrmecophytic trees. In all three common ant species, colony size, as measured by total number of ants, increased with domatia size. Additionally, domatia volume and species identity interact to predict ant numbers, suggesting differing responses between species to increased availability of nesting space. The proportion of total ants in nests that were immatures varied with thorn volume and species, highlighting the importance of domatia morphology in influencing colony structure.

In much of the English-speaking world the lawn is the most common of all garden features. For arguably a millennium it has played a significant role in the landscape and during that period it has been inextricably linked with grasses. Nevertheless other plant species have accompanied the grasses and also been used in creating lawns. From medieval wildflowers to Victorian weeds, the plants that challenge the formal concept of the perfect lawn have journeyed with it but have until recently remained only small players within the dominion of grass. By the beginning of the 21st century, with a new environmental ethos permeating the garden, the long journey of the grassy lawn and its plant companions has led to the grass monoculture being heretically rethought: by removing both the monoculture and the grass.

Domestic cats (Felis catus) are the most abundant predator in many urban ecosystems, and their ranging behaviour will help determine predation rates. To investigate how degree of urbanisation affects cat ranging behaviour, we used GPS trackers to follow 38 cats in three (urban, suburban and peri-urban) residential areas in the large town of Reading, UK. Median home range (95% KE) was 1.28 ha, but varied from 0.9 ha in the urban habitat, to 1.56 ha in the suburban habitat and 1.60 ha in the peri-urban region, with a maximum range size of 6.61 ha. The median maximum distance reached from home was 99 m, and again varied with level of urbanisation (urban: 79 m; suburban: 141 m; peri-urban: 148 m; maximum 278 m). For home and core (50% KE) ranges there were no significant differences with respect to study areas, cat sexes, cats living in the same household, or day/night range. A decreased proportion of constructed surfaces (a proxy for urbanisation) was associated with an increase in cat range size. As urban areas grow, many areas containing species of conservation importance are encroached upon by residential zones on urban fringes. To protect these species we suggest that boundary habitats should be managed to reduce rates of cat access to these areas, or that buffer zones of 300-400m should be formed between housing and areas containing vulnerable species. These management options may help mitigate the ecological consequences of cat predation.

Ants are widely employed by plants as an antiherbivore defence. A single host plant can associate with multiple, symbiotic ant species, although usually only a single ant species at a time. Different plant-ant species may vary in the degree to which they defend their host plant. In Kenya, ant–acacia interactions are well studied, but less is known about systems elsewhere in Africa. A southern African species, Vachellia erioloba, is occupied by thorn-dwelling ants from three different genera. Unusually, multiple colonies of all these ants simultaneously and stably inhabit trees. We investigated if the ants on V. erioloba (i) deter insect herbivores; (ii) differ in their effectiveness depending on the identity of the herbivore; and (iii) protect the tree against an important herbivore, the larvae of the lepidopteran Gonometa postica. We show that experimental exclusion of ants leads to greater levels of herbivory on trees. The ants inhabiting V. erioloba are an effective deterrent against hemipteran and coleopteran, but not lepidopteran herbivores. Defensive services do not vary among ant species, but only Crematogaster ants exhibit aggression towards G. postica. This highlights the potential of the V. erioloba–ant mutualism for studying ant–plant interactions that involve multiple, simultaneously resident thorn-dwelling ant species.

Parasitoids are the most important natural enemies of many insect species. Larvae of many Drosophila species can defend themselves against attack by parasitoids through a cellular immune response called encapsulation. The paper reviews recent studies of the evolutionary biology and ecological genetics of resistance in Drosophila, concentrating on D. melanogaster. The physiological basis of encapsulation, and the genes known to interfere with resistance are briefly summarized. Evidence for within- and between-population genetic variation in resistance from isofemale line, artificial selection and classical genetic studies are reviewed. There is now firm evidence that resistance is costly to Drosophila, and the nature of this cost is discussed, and the possibility that it may involve a reduction in metabolic rate considered. Comparative data on encapsulation and metabolic rates across seven Drosophila species provides support for this hypothesis. Finally, the possible population and community ecological consequences of evolution in the levels of host resistance are examined.

Herbivore dynamics and community structure are influenced both by plant quality and the actions of natural enemies. A factorial experiment manipulating both higher and lower trophic levels was designed to explore the determinants of colony growth of the aphid Aphis jacobaeae, a specialist herbivore on ragwort Senecio jacobaea. Potential plant quality was manipulated by regular addition of NPK-fertiliser and predator pressure was reduced by interception traps; the experiment was carried out at two sites. The size and persistence of aphid colonies were measured. Fertiliser addition affected plant growth in only one site, but never had a measurable effect on aphid colony growth. In both habitats the action of insect predators dominated, imposing strong and negative effects on aphid colony performance. Ants were left unmanipulated in both sites and their performance on the aphid colonies did not significantly differ between sites or between treatments. Our results suggest that, at least for aphid herbivores on S. jacobaea, the action of generalist insect predators appears to be the dominant factor affecting colony performance and can under certain conditions even improve plant productivity.

Providing supplementary food for wild birds is a globally popular pastime; almost half of the households in many developed countries participate and billions of US dollars are spent annually. Although the direct influence of this additional resource on bird survivorship and fecundity has been studied, there is little understanding of the wider ecological consequences of this massive perturbation to (what are usually) urban ecosystems. We investigated the possible effects of wild bird feeding on the size and survivorship of colonies of a widespread arthropod prey species of many small passerine birds, the pea aphid [Acyrthosiphon pisum (Harris); Hemiptera: Aphididae], in suburban gardens in a large town in southern England. We found significantly fewer aphids and shorter colony survival times in colonies exposed to avian predation compared to protected controls in gardens with a bird feeder but no such differences between exposed and protected colonies in gardens that did not feed birds. Our work therefore suggests that supplementary feeding of wild birds in gardens may indirectly influence population sizes and survivorship of their arthropod prey and highlights the need for further research into the potential effects on other species.

1. Intra-specific variation in plant defence traits has been shown to profoundly affect herbivore community structure. Here we describe two experiments designed to test whether similar effects occur at higher trophic levels, by studying pea aphid–natural enemy interactions in a disused pasture in southern England. 2. In the first experiment, the numbers and identity of natural enemies attacking different monoclonal pea aphid colonies were recorded in a series of assays throughout the period of pea aphid activity. 3. In the summer assay, there was a significant effect of clone on the numbers of aphidophagous hoverfly larvae and the total number of non-hoverfly natural enemies recruited. Clone also appeared to influence the attack rate suffered by the primary predator in the system, the hoverfly Episyrphus balteatus, by Diplazon laetatorius, an ichneumonid parasitoid. Colonies were generally driven to extinction by hoverfly attack, resulting in the recording of low numbers of parasitoids and entomopathogens, suggesting intense intra-guild predation. 4. To further examine the influence of clonal variation on the recruitment of natural enemies, a second experiment was performed to monitor the temporal dynamics of community development. Colonies were destructively sampled every other day and the numbers of natural enemies attacking aphid colonies were recorded. These data demonstrated that clonal variation influenced the timing, abundance, and identity of natural enemies attacking aphid colonies. 5. Taken together, these data suggest that clonal variation may have a significant influence on the patterns of interactions between aphids and their natural enemies, and that such effects are likely to affect our understanding of the ecology and biological control of these insect herbivores.

Traditionally, spoor (tracks, pug marks) have been used as a cost effective tool to assess the presence of larger mammals. Automated camera traps are now increasingly utilized to monitor wildlife, primarily as the cost has greatly declined and statistical approaches to data analysis have improved. While camera traps have become ubiquitous, we have little understanding of their effectiveness when compared to traditional approaches using spoor in the field. Here, we a) test the success of camera traps in recording a range of carnivore species against spoor; b) ask if simple measures of spoor size taken by amateur volunteers is likely to allow individual identification of leopards and c) for a trained tracker, ask if this approach may allow individual leopards to be followed with confidence in savannah habitat. We found that camera traps significantly under-recorded mammalian top and meso-carnivores, with camera traps more likely under-record the presence of smaller carnivores (civet 64%; genet 46%, Meller’s mongoose 45%) than larger (jackal sp. 30%, brown hyena 22%), while leopard was more likely to be recorded by camera trap (all recorded by camera trap only). We found that amateur trackers could be beneficial in regards to collecting presence data; however the large variance in measurements of spoor taken in the field by volunteers suggests that this approach is unlikely to add further data. Nevertheless, the use of simple spoor measurements in the field by a trained field researcher increases their ability to reliably follow a leopard trail in difficult terrain. This allows researchers to glean further data on leopard behaviour and habitat utilisation without the need for complex analysis.

The accidental introduction of the spiralling whitefly, Aleurodicus dispersus Russell (Homoptera: Aleyrodidae) to Seychelles in late 2003 is exploited during early 2005 to study interactions between A. dispersus, native and exotic host plants and their associated arthropod fauna. The numbers of A. dispersus egg spirals and pupae, predator and herbivore taxa were recorded for eight related native/exotic pairs of host plants found on Mahe, the largest island in Seychelles. Our data revealed no significant difference in herbivore density (excluding A. dispersus) between related native and exotic plants, which suggests that the exotic plants do not benefit from ‘enemy release’. There were also no differences in predator density, or combined species richness between native and exotic plants. Together these data suggest that ‘biotic resistance’ to invasion is also unlikely. Despite the apparent lack of differences in community structure significantly fewer A. dispersus egg spirals and pupae were found on the native plants than on the exotic plants. Additional data on A. dispersus density were collected on Cousin Island, a managed nature reserve in which exotic plants are carefully controlled. Significantly higher densities of A. dispersus were observed on Mahe, where exotic plants are abundant, than on Cousin. These data suggest that the rapid invasion of Seychelles by A. dispersus may largely be due to the high proportion of plant species that are both exotic and hosts of A. dispersus; no support was found for either the ‘enemy release’ or the ‘biotic resistance’ hypotheses.

Competition is one of the most important biotic factors determining the structure of ecological communities. In this study, we show that there is variation in competitive ability between two clones of the pea aphid, Acyrthosiphon pisum, both of which out-compete a clone of the vetch aphid, Megoura viciae, in the laboratory. We tested whether this variation in competitive ability would alter the outcome of interspecific competition in the field. White one pea aphid clone followed the pattern set in the laboratory, out-competing the Megoura viciae clone, another showed the reverse effect with Megoura viciae dominating. These differences appear to be the result of variation in early population growth rate between the pea aphid clones, rather than predation, although predation did lead to the eventual extinction of colonies. We also questioned whether intra- and interspecific differences in predator escape behaviour could affect the outcome of competition in the field. All three clones responded similarly to the presence of foraging hoverfly larvae (Episyrphus balteatus), but the Megoura viciae clone dropped from the plant significantly less often in response to the presence of a foraging two-spot ladybird (Adalia bipunctata). This work provides evidence that intraspecific variation in competitive ability can alter the outcome of interspecific competitive interactions in nature and suggests that species-specific behavioural. traits may have the potential to modify the outcome of these interactions. (c) 2005 Gesellschaft fur Okologie. Published by Elsevier GmbH. All rights reserved.

1. The presence of an across-species trade-off between dispersal ability and competitive ability has been proposed as a mechanism that facilitates coexistence. It is not clear if a similar trade-off exists within species. Such a trade-off would constrain the evolution of either trait and, given appropriate selection pressures, promote local adaptation in these traits. 2. This study found substantial levels of heritable variation in competitive ability of the pea aphid, Acyrthosiphon pisum Harris (Homoptera: Aphididae), measured in terms of relative survival when reared with a single clone of the vetch aphid, Megoura viciae Buckton (Homoptera: Aphididae). 3. Pea aphids can move to new patches by either flying (longer distance dispersal) or walking (local dispersal) from plant to plant. There was considerable clonal variation in dispersal ability, measured in terms of the proportion of winged offspring produced, and ability to survive away from their host plant. 4. Winged individuals showed longer off-plant survival times than wingless forms of the same pea aphid clone. 5. There was no evidence of a relationship between clonal competitive ability and either measure of dispersal ability, although the power of the test is limited by the number of pea aphid clones used in the trial. 6. However, there was a positive correlation between clonal fecundity and the proportion of winged offspring produced. Although speculative, it is suggested that clones that are more likely to either overwhelm their host plant or attract higher numbers of natural enemies as a result of having higher fecundity are more likely to produce winged morphs.

Typically, the relationship between insect development and temperature is described by two characteristics: the minimum temperature needed for development to occur (T-min) and the number of day degrees required (DDR) for the completion of development. We investigated these characteristics in three English populations of Thrips major and T tabaci [Cawood, Yorkshire (N53degrees49', W1degrees7'); Boxworth, Cambridgeshire (N52degrees15', W0degrees1'); Silwood Park, Berkshire (N51degrees24', W0degrees38')], and two populations of Frankliniella occidentalis (Cawood; Silwood Park). While there were no significant differences among populations in either T-min (mean for T major = 7.0degreesC; T tabaci = 5.9degreesC; F. occidentalis = 6.7degreesC) or DDR (mean for T major = 229.9; T tabaci = 260.8; F occidentalis = 233.4), there were significant differences in the relationship between temperature and body size, suggesting the presence of geographic variation in this trait. Using published data, in addition to those newly collected, we found a negative relationship between T-min. and DDR for F occidentalis and T tabaci, supporting the hypothesis that a trade-off between T-min and DDR may constrain adaptation to local climatic conditions.

We report the results of a study investigating the influence of elevated CO2 on species interactions across three trophic levels: a plant (Brassica oleracea), two aphid herbivores (the generalist Myzus persicae and the specialist Brevicoryne brassicae), and two natural enemies (the coccinellid Hippodamia convergens (ladybird) and the parasitoid wasp Diaeretiella rapae). Brassica oleracea plants reared under elevated CO2 conditions (650 ppmv vs. 350 ppmv) were larger and had decreased water and nitrogen content. Brevicoryne brassicae reared on plants grown in elevated CO2 were larger and accumulated more fat, while there was no change in M. persicae traits. Fecundity of individual aphids appeared to be increased when reared on plants grown in elevated CO2. However, these differences were generally lost when aphids were reared in colonies, suggesting that such changes in plant quality will have subtile effects on aphid intraspecific interactions. Nevertheless, CO2 treatment did influence aphid distribution on plants, with significantly fewer M. persicae found on the shoots, and B. brassicae was only found on senescing leaves, when colonies were reared on plants grown in elevated CO2. We reared B. brassicae and M. persicae in competition on plants grown at both the CO2 concentration treatments. We found a significantly lower ratio of M. persicae: B. brassicae on plants grown under elevated CO2 conditions, strongly suggesting that increasing CO2 concentrations can alter the outcome of competition among insect herbivores. This was also reflected in the distribution of the aphids on the plants. While the CO2 treatment did not influence where B. brassicae were found, fewer M. persicae were present on senescing leaves under elevated CO2 conditions. Changes in plant quality resulting from the CO2 treatments did not appear to alter aphid quality as prey species, as the number consumed by the ladybird H. convergens, and the number parasitised by the parasitoid wasp D. rapae, did not change. To our knowledge, this study provides the first empirical evidence that changes in host plant quality mediated by increasing levels of CO2 can alter the outcome of interspecific competition among insect herbivores.

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, naive female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness.

The interplay between coevolutionary and population or community dynamics is currently the focus of much empirical and theoretical consideration. Here, we develop a simulation model to study the coevolutionary and population dynamics of a hypothetical host–parasitoid interaction. In the model, host resistance and parasitoid virulence are allowed to coevolve. We investigate how trade-offs associated with these traits modify the system's coevolutionary and population dynamics. The most important influence on these dynamics comes from the incorporation of density-dependent costs of resistance ability. We find three main outcomes. First, if the costs of resistance are high, then one or both of the players go extinct. Second, when the costs of resistance are intermediate to low, cycling population and coevolutionary dynamics are found, with slower evolutionary changes observed when the costs of virulence are also low. Third, when the costs associated with resistance and virulence are both high, the hosts trade-off resistance against fecundity and invest little in resistance. However, the parasitoids continue to invest in virulence, leading to stable host and parasitoid population sizes. These results support the hypothesis that costs associated with resistance and virulence will maintain the heritable variation in these traits found in natural populations and that the nature of these trade-offs will greatly influence the population dynamics of the interacting species.

The populations of many species are structured such that mating is not random and occurs between members of local patches. When patches are founded by a single female and all matings occur between siblings, brothers may compete with each other for matings with their sisters. This local mate competition (LMC) selects for a female-biased sex ratio, especially in species where females have control over offspring sex, as in the parasitic Hymenoptera. Two factors are predicted to decrease the degree of female bias: (1) an increase in the number of foundress females in the patch and (2) an increase in the fraction of individuals mating after dispersal from the natal patch. Pollinating fig wasps are well known as classic examples of species where all matings occur in the local patch. We studied non-pollinating fig wasps, which are more diverse than the pollinating fig wasps and also provide natural experimental groups of species with different male morphologies that are linked to different mating structures. In this group of wasps, species with wingless males mate in the local patch (i.e. the fig fruit) while winged male species mate after dispersal. Species with both kinds of male have a mixture of local and non-local mating. Data from 44 species show that sex ratios (defined as the proportion of males) are in accordance with theoretical predictions: wingless male species

1. The evolution of host resistance to parasitoid attack will be constrained by two factors: the costs of the ability to defend against attack, and the costs of surviving actual attack. These factors have been investigated using Drosophila melanogaster and its parasitoids as a model system. The costs of defensive ability are expressed as a trade-off with larval competitive ability, whereas the costs of actual defence are exhibited in terms of reduced adult fecundity and size. 2. The costs of actual defence may be ameliorated by the host-choice decisions made by Pachycrepoideus vindemiae, a pupal parasitoid. If larvae that have successfully encapsulated a parasitoid develop into poorer quality hosts, then these may be rejected by ovipositing pupal parasitoids. 3. Pupae developing from larvae that have encapsulated the parasitoid Asobara tabida are smaller and have relatively thinner puparia. Thinner puparia are likely to be associated with a reduction in mechanical strength and possibly with a decrease in desiccation tolerance. 4. Pachycrepoideus vindemiae that develop in capsule-bearing pupae are smaller than those that emerge from previously unattacked hosts. This supports the prediction that ovipositing female P. vindemiae should avoid attacking capsule-bearing hosts. However, in choice experiments with 1-day-old pupae, P. vindemiae females oviposited preferentially in hosts containing a capsule, whereas there was no preference found with 4-day-old hosts. This appears to be a maladaptive host choice decision, as the female pupal parasitoids are preferentially attacking hosts that will result in a reduction of their own fitness. 5. The increased likelihood of attack by a pupal parasitoid is another cost of actual defence against larval parasitoid attack.

Larvae of the pine beauty moth Panolis flammae (Denis & Schiffermüller) were reared in sleeve cages on five different seed origins (provenances) of pole stage Pinus contorta in the field in each of four years from 1985 to 1988. Survival varied significantly between the years. In those years when survival was high, Significant differences between tree provenance were not found. However, between provenance significant differences were found in larval weight and stage of development. In the years when survival was low, the results seen in good years were reversed. significant differences attributable to provenance were found but these were not reflected in significant differences between larval weight or development. In addition, there was a significant correlation between the proportion surviving and larval weight, which was not the case in those years where larval survival was high. The results are discussed in light of the pest status of P. flammea in Britain and in view of current silvicultural policies. The use of trees resistant to insect attack as part of an integrated pest management programme is highlighted and the need to coordinate laboratory and field studies so as to control for environmental variation discussed.

Recent work has shown that the evolution of Drosophila melanogaster resistance to attack by the parasitoid Asobara tabida is constrained by a trade-off with larval competitive ability. However, there are two very important questions that need to be answered. First, is this a general cost, or is it parasitoid specific? Second, does a selected increase in immune response against one parasitoid species result in a correlated change in resistance to other parasitoid species? The answers to both questions will influence the coevolutionary dynamics of these species, and also may have a previously unconsidered, yet important, influence on community structure.

Parasitoids are among the most important natural enemies of insects in many environments.Acyrthosiphon pisum, the pea aphid, is a common pest of the leguminous crops in temperate regions. Pea aphids are frequently attacked by a range of endoparasitic wasps, including the common aphidiine,Aphidius ervi. Immunity to parasitoid attack is thought to involve secondary symbiotic bacteria, the presence of which is associated with the death of the parasitoid egg. It has been suggested that there is a fecundity cost of resistance, as individuals carrying the secondary symbionts associated with parasitoid resistance have fewer offspring. Supporting this hypothesis, we find a positive relationship between fecundity and susceptibility to parasitoid attack. There is also a negative relationship between fecundity and off-plant survival time (which positively correlates with resistance to parasitoid attack). Taken together, these results suggest that the aphids can either invest in defence (parasitoid resistance, increased off-plant survival time) or reproduction, and speculate that this may be mediated by changes in the aphids' endosymbiont fauna. Furthermore, there is a positive relationship between aphid size and resistance, suggesting that successful resistance to parasitoid attack may involve physical, as well as physiological, defences.

Costs of resistance are widely assumed to be important in the evolution of parasite and pathogen defence in animals, but they have been demonstrated experimentally on very few occasions. Endoparasitoids are insects whose larvae develop inside the bodies of other insects where they defend themselves from attack by their hosts' immune systems (especially cellular encapsulation). Working with Drosophila melanogaster and its endoparasitoid Leptopilina boulardi, we selected for increased resistance in four replicate populations of flies. The percentage of flies surviving attack increased from about 0.5% to between 40% and 50% in five generations, revealing substantial additive genetic variation in resistance in the field population from which our culture was established. In comparison with four control lines, flies from selected lines suffered from lower larval survival under conditions of moderate to severe intraspecific competition.

Urban domestic cat (Felis catus) populations can attain exceedingly high densities and are not limited by natural prey availability. This has generated concerns that they may negatively affect prey populations, leading to calls for management. We enlisted cat-owners to record prey returned home to estimate patterns of predation by free-roaming pets in different localities within the town of Reading, UK and questionnaire surveys were used to quantify attitudes to different possible management strategies. Prey return rates were highly variable: only 20% of cats returned ≥4 dead prey annually. Consequently, approximately 65% of owners received no prey in a given season, but this declined to 22% after eight seasons. The estimated mean predation rate was 18.3 prey cat⁻¹ year⁻¹ but this varied markedly both spatially and temporally: per capita predation rates declined with increasing cat density. Comparisons with estimates of the density of six common bird prey species indicated that cats killed numbers equivalent to adult density on c. 39% of occasions. Population modeling studies suggest that such predation rates could significantly reduce the size of local bird populations for common urban species. Conversely, most urban residents did not consider cat predation to be a significant problem. Collar-mounted anti-predation devices were the only management action acceptable to the majority of urban residents (65%), but were less acceptable to cat-owners because of perceived risks to their pets; only 24% of cats were fitted with such devices. Overall, cat predation did appear to be of sufficient magnitude to affect some prey populations, although further investigation of some key aspects of cat predation is warranted. Management of the predation behavior of urban cat populations in the UK is likely to be challenging and achieving this would require considerable engagement with cat owners.