Disentangling the drivers of urban bird diversity in the non-breeding season: A general synthesis.

Current knowledge about the impacts of urbanisation on bird assemblages is based on evidence from studies partly or wholly undertaken in the breeding season. In comparison, the non-breeding season remains little studied, despite the fact that winter conditions at higher latitudes are changing more rapidly than other seasons. During the non-breeding season, cities may attract or retain bird species because they offer milder conditions or better feeding opportunities than surrounding habitats. However, the range of climatic, ecological and anthropogenic mechanisms shaping different facets of urban bird diversity in the non-breeding season are poorly understood. We explored these mechanisms using structural equation modelling to assess how urbanisation affects the taxonomic, phylogenetic and functional diversity of avian assemblages sampled worldwide in the non-breeding season. We found that minimum temperature, elevation, urban area and city age played a critical role in determining taxonomic diversity while a range of factors-including productivity, precipitation, elevation, distance to coasts and rivers, socio-economic (as a proxy of human facilitation) and road density-each contributed to patterns of phylogenetic and functional diversity. The structure and function of urban bird assemblages appear to be predominantly shaped by temperature, productivity and city age, with effects of these factors differing across seasons. Our results underline the importance of considering multiple hypotheses, including seasonal effects, when evaluating the impacts of urbanisation on biodiversity.

Dense city centers support less evolutionary unique bird communities than sparser urban areas.

Urbanization alters avian communities, generally lowering the number of species and contemporaneously increasing their functional relatedness, leading to biotic homogenization. Urbanization can also negatively affect the phylogenetic diversity of species assemblages, potentially decreasing their evolutionary distinctiveness. We compare species assemblages in a gradient of building density in seventeen European cities to test whether the evolutionary distinctiveness of communities is shaped by the degree of urbanization. We found a significant decline in the evolutionary uniqueness of avian communities in highly dense urban areas, compared to low and medium-dense areas. Overall, communities from dense city centers supported one million years of evolutionary history less than communities from low-dense urban areas. Such evolutionary homogenization was due to a filtering process of the most evolutionarily unique birds. Metrics related to evolutionary uniqueness have to play a role when assessing the effects of urbanization and can be used to identify local conservation priorities.

Spatial Overlap and Habitat Selection of Corvid Species in European Cities.

Understanding habitat and spatial overlap in sympatric species of urban areas would aid in predicting species and community modifications in response to global change. Habitat overlap has been widely investigated for specialist species but neglected for generalists living in urban settings. Many corvid species are generalists and are adapted to urban areas. This work aimed to determine the urban habitat requirements and spatial overlap of five corvid species in sixteen European cities during the breeding season. All five studied corvid species had high overlap in their habitat selection while still having particular tendencies. We found three species, the Carrion/Hooded Crow, Rook, and Eurasian Magpie, selected open habitats. The Western Jackdaw avoided areas with bare soil cover, and the Eurasian Jay chose more forested areas. The species with similar habitat selection also had congruent spatial distributions. Our results indicate that although the corvids had some tendencies regarding habitat selection, as generalists, they still tolerated a wide range of urban habitats, which resulted in high overlap in their habitat niches and spatial distributions.

Effects of light and noise pollution on avian communities of European cities are correlated with the species' diet.

Urbanization affects avian community composition in European cities, increasing biotic homogenization. Anthropic pollution (such as light at night and noise) is among the most important drivers shaping bird use in urban areas, where bird species are mainly attracted by urban greenery. In this study, we collected data on 127 breeding bird species at 1349 point counts distributed along a gradient of urbanization in fourteen different European cities. The main aim was to explore the effects of anthropic pollution and city characteristics, on shaping the avian communities, regarding species' diet composition. The green cover of urban areas increased the number of insectivorous and omnivorous bird species, while slightly decreasing the overall diet heterogeneity of the avian communities. The green heterogeneity-a measure of evenness considering the relative coverage of grass, shrubs and trees-was positively correlated with the richness of granivorous, insectivorous, and omnivorous species, increasing the level of diet heterogeneity in the assemblages. Additionally, the effects of light pollution on avian communities were associated with the species' diet. Overall, light pollution negatively affected insectivorous and omnivorous bird species while not affecting granivorous species. The noise pollution, in contrast, was not significantly associated with changes in species assemblages. Our results offer some tips to urban planners, managers, and ecologists, in the challenge of producing more eco-friendly cities for the future.

EVI and NDVI as proxies for multifaceted avian diversity in urban areas.

Most ecological studies use remote sensing to analyze broad-scale biodiversity patterns, focusing mainly on taxonomic diversity in natural landscapes. One of the most important effects of high levels of urbanization is species loss (i.e., biotic homogenization). Therefore, cost-effective and more efficient methods to monitor biological communities' distribution are essential. This study explores whether the Enhanced Vegetation Index (EVI) and the Normalized Difference Vegetation Index (NDVI) can predict multifaceted avian diversity, urban tolerance, and specialization in urban landscapes. We sampled bird communities among 15 European cities and extracted Landsat 30-meter resolution EVI and NDVI values of the pixels within a 50-m buffer of bird sample points using Google Earth Engine (32-day Landsat 8 Collection Tier 1). Mixed models were used to find the best associations of EVI and NDVI, predicting multiple avian diversity facets: Taxonomic diversity, functional diversity, phylogenetic diversity, specialization levels, and urban tolerance. A total of 113 bird species across 15 cities from 10 different European countries were detected. EVI mean was the best predictor for foraging substrate specialization. NDVI mean was the best predictor for most avian diversity facets: taxonomic diversity, functional richness and evenness, phylogenetic diversity, phylogenetic species variability, community evolutionary distinctiveness, urban tolerance, diet foraging behavior, and habitat richness specialists. Finally, EVI and NDVI standard deviation were not the best predictors for any avian diversity facets studied. Our findings expand previous knowledge about EVI and NDVI as surrogates of avian diversity at a continental scale. Considering the European Commission's proposal for a Nature Restoration Law calling for expanding green urban space areas by 2050, we propose NDVI as a proxy of multiple facets of avian diversity to efficiently monitor bird community responses to land use changes in the cities.

Long-Term Winter Population Trends of Corvids in Relation to Urbanization and Climate at Northern Latitudes.

Corvids (crows, magpies, jays) live in a close association with humans, and therefore knowledge about their population status and changes will be an essential part of monitoring the quality of urban environments. Wintering bird populations can track habitat and climate changes more rapidly than breeding populations. We conducted a long-term (1991-2020) winter census of corvid species in 31 human settlements along a 920 km latitudinal gradient in Finland. We observed a total of five corvid species: the Eurasian Magpie (occurring in 114 surveys out of 122; total abundance 990 ind.), the Hooded Crow (in 96 surveys; 666 ind.), the Eurasian Jackdaw (in 51 surveys; 808 ind.), the Eurasian Jay (in 5 surveys; 6 ind.) and the Rook (in 1 survey; 1 ind.). Only the numbers of the Eurasian Jackdaw differed between the study winters, being greater at the end of the study period (2019/2020) than during the earlier winters (1991/1992 and 1999/2000). The average growth rate (λ) of the Eurasian Jackdaw increased during the study period, whereas no changes were observed in the cases of the Hooded Crow or the Eurasian Magpie. The growth rate of the Eurasian Jackdaw was greater than that observed in the Finnish bird-monitoring work, probably because our data came only from the core area of each human settlement. Even though the number of buildings and their cover increased in the study plots, and the winter temperature differed between winters, the average growth rate (λ) of corvid species did not significantly correlate with these variables. These results suggest that urban settlements are stable wintering environments for the generalist corvids. The between-species interactions were all positive, but non-significant. Despite the total number of winter-feeding sites being greater during the winter of 1991/1992 than during the winter of 2019/2020, the changes in the numbers of feeding stations did not correlate with the growth rates of any corvid species. We assume that the Eurasian Jackdaw has benefitted from the decreased persecution, and probably also from large-scale climate warming that our study design was unable to take in to account. Our results indicated that wintering corvid populations succeed well in the human settlements in Finland. We recommend conducting long-term corvid research, also during breeding season, to understand more detailed causes of the population changes of corvids along an urban gradient. Without year-round long-term monitoring data, the conservation and management recommendations related to the corvid species in urban habitats may be misleading.

Using functional traits and phylogeny to understand local extinction risk in dragonflies and damselflies (Odonata).

Understanding the risk of local extinction of a species is vital in conservation biology, especially now when anthropogenic disturbances and global warming are severely changing natural habitats. Local extinction risk depends on species traits, such as its geographical range size, fresh body mass, dispersal ability, length of flying period, life history variation, and how specialized it is regarding its breeding habitat. We used a phylogenetic approach because closely related species are not independent observations in the statistical tests. Our field data contained the local extinction risk of 31 odonate (dragonflies and damselflies) species from Central Finland. Species relatedness (i.e., phylogenetic signal) did not affect local extinction risk, length of flying period, nor the geographical range size of a species. However, we found that closely related species were similar in hind wing length, length of larval period, and habitat of larvae. Both phylogenetically corrected (PGLS) and uncorrected (GLM) analysis indicated that the geographical range size of species was negatively related to local extinction risk. Contrary to expectations, habitat specialist species did not have higher local extinction rates than habitat generalist species nor was it affected by the relatedness of species. As predicted, species' long larval period increased, and long wings decreased the local extinction risk when evolutionary relatedness was controlled. Our results suggest that a relatively narrow geographical range size is an accurate estimate for a local extinction risk of an odonate species, but the species with long life history and large habitat niche width of adults increased local extinction risk. Because the results were so similar between PGLS and GLM methods, it seems that using a phylogenetic approach does not improve predicting local extinctions.

Effects of urbanization on taxonomic, functional and phylogenetic avian diversity in Europe.

Europe is an urbanized continent characterized by a long history of human-wildlife interactions. This study aimed to assess the effects of specific elements of urbanization and urban pollution on complementary avian diversity metrics, to provide new insights on the conservation of urban birds. Our study recorded 133 bird species at 1624 point counts uniformly distributed in seventeen different European cities. Our results thus covered a large spatial scale, confirming both effects of geographical and local attributes of the cities on avian diversity. However, we found contrasting effects for the different diversity components analyzed. Overall, taxonomic diversity (bird species richness), phylogenetic diversity and relatedness were significantly and negatively associated with latitude, while functional dispersion of communities showed no association whatsoever. At the local level (within the city), we found that urban greenery (grass, bush, and trees) is positively correlated with the number of breeding bird species, while the building cover showed a detrimental effect. Functional dispersion was the less affected diversity metric, while grass and trees and water (rivers or urban streams) positively affected the phylogenetic diversity of avian communities. Finally, the phylogenetic relatedness of species increased with all the main indicators of urbanization (building surface, floors, pedestrian's density and level of light pollution) and was only mitigated by the presence of bushes. We argue that maintaining adequate levels of avian diversity within the urban settlements can help to increase the potential resilience of urban ecosystems exposed to the stress provoked by rapid and continuous changes. We listed some characteristics of the cities providing positive and negative effects on each facet of urban avian diversity.

Spatial and temporal changes in occupancy frequency distribution patterns of freshwater macrophytes in Finland.

A useful method for characterizing biological numerous assemblages at regional scales is the species occupancy frequency distribution (SOFD). An SOFD shows the number or proportion of study sites each species occurred. Species that occur at only a few sites are termed satellite species, while species that occur at many sites are termed core species. This study is the first to document and assess SOFD patterns in aquatic macrophytes. It characterizes SOFD patterns of freshwater macrophyte assemblages in Finland at two spatial and two temporal scales. For this, I analyzed three published datasets on freshwater macrophyte distributions: two from studies conducted at a local scale and the third from large national surveys. One local study and the national study also included data on temporal variation in species occupancy frequencies. In the national study, the number of core and satellite species varied slightly between the older and the newer survey, respectively. Among the 113 waterbodies surveyed as part of the national study, the SOFD followed a unimodal satellite pattern. However, for the older dataset (from the 1930s), a bimodal symmetric pattern also fit the SOFD data well. At the local scale, I observed geographical variation in SOFD patterns. The dataset from southern Finland followed a unimodal satellite SOFD pattern; data from central Finland instead displayed a bimodal symmetric SOFD pattern, although they also fit equally well with a bimodal truncated pattern. Moreover, temporal patterns in central Finland seemed to demonstrate a shift from a bimodal symmetric to a bimodal asymmetric SOFD probably. Geographical variation in the SOFD pattern may be due to variation in the regional species pool. The temporal changes in SOFD pattern may be due to lake eutrophication and anthropogenic disturbance around waterbodies, which may increase number of macrophyte species.

The impact of tree crops and temperature on the timing of frugivorous bird migration.

Migration has evolved to tackle temporal changes in availability of resources. Climate change has been shown to affect the migration dates of species, which raises the question of whether the variation in the timing of migration is climate or resource dependent? The relative importance of temperature and availability of food as drivers of migration behaviour during both spring and autumn seasons has been poorly studied. Here, we investigated these patterns in frugivorous and granivorous birds (hereafter frugivorous) that are assumed to postpone their autumn migration when there is plenty of food available, which may also advance upcoming spring migration. On the other hand, especially spring migration dates have been negatively connected with increasing temperatures. We tested whether the autumn and spring migration dates of eleven common frugivorous birds depended on the crop size of trees or ambient temperatures using 29 years of data in Finland. The increased crop sizes of trees delayed autumn migration dates; whereas, autumn temperature did not show a significant connection. We also observed a temporal trend towards later departure. Increasing temperature and crop sizes advanced spring arrival dates. Our results support the hypothesis that the timing of autumn migration in the frugivorous birds depends on the availability of food and is weakly connected with the variation in temperature. Importantly, crop size can have carry-over effects and affect the timing of spring arrival possibly because birds have overwintered closer to the breeding grounds after an abundant crop year.

Odonate species occupancy frequency distribution and abundance-occupancy relationship patterns in temporal and permanent water bodies in a subtropical area.

This paper investigates species richness and species occupancy frequency distributions (SOFD) as well as patterns of abundance-occupancy relationship (SAOR) in Odonata (dragonflies and damselflies) in a subtropical area. A total of 82 species and 1983 individuals were noted from 73 permanent and temporal water bodies (lakes and ponds) in the Pampa biome in southern Brazil. Odonate species occupancy ranged from 1 to 54. There were few widely distributed generalist species and several specialist species with a restricted distribution. About 70% of the species occurred in <10% of the water bodies, yielding a surprisingly high number of rare species, often making up the majority of the communities. No difference in species richness was found between temporal and permanent water bodies. Both temporal and permanent water bodies had odonate assemblages that fitted best with the unimodal satellite SOFD pattern. It seems that unimodal satellite SOFD pattern frequently occurred in the aquatic habitats. The SAOR pattern was positive and did not differ between permanent and temporal water bodies. Our results are consistent with a niche-based model rather than a metapopulation dynamic model.

Biodiversity within the city: Effects of land sharing and land sparing urban development on avian diversity.

Urbanization, one of the most extreme human-induced environmental changes, is negatively affecting biodiversity worldwide, strongly suggesting that we should reconcile urban development with conservation. Urbanization can follow two extreme types of development within a continuum: land sharing (buildings mixed with dispersed green space) or land sparing (buildings interspersed with green patches that concentrate biodiversity-supporting vegetation). Recent local-scale studies indicate that biodiversity is typically favored by land sparing. We investigated which of these two types of urbanization is associated with a higher taxonomic (i.e. species richness), functional, and phylogenetic diversity of birds. To do so, we collected information on breeding and wintering bird assemblages in 45 land-sharing and 45 land-sparing areas in nine European cities, which provide the first attempt to explore this question using a large geographical scale and temporal replication. We found that land-sharing urban areas were significantly associated with a higher taxonomic and functional diversity of birds during winter, but not during the breeding season (with only a marginally significant effect for functional diversity). We found no association between the type of urban development and phylogenetic diversity. Our findings indicate that not all components of avian diversity are similarly affected by these two means of urban planning and highlight the importance of integrating the temporal perspective into this kind of studies. Our results also offer useful information to the current debate about the trade-off between biodiversity conservation and human well-being in the context of land sharing and sparing urban practices. In addition, we found that certain small-scale urban landscape characteristics (i.e. few impervious surfaces, high water or tree cover) and human practices (i.e. bird feeders or plants with berries) can help maintaining more diverse urban bird assemblages. We provide specific suggestions for both policymakers and citizens that hopefully will help to create more biodiversity-friendly cities in the future.

Land-sharing vs. land-sparing urban development modulate predator-prey interactions in Europe.

Urban areas are expanding globally as a consequence of human population increases, with overall negative effects on biodiversity. To prevent the further loss of biodiversity, it is urgent to understand the mechanisms behind this loss to develop evidence-based sustainable solutions to preserve biodiversity in urban landscapes. The two extreme urban development types along a continuum, land-sparing (large, continuous green areas and high-density housing) and land-sharing (small, fragmented green areas and low-density housing) have been the recent focus of debates regarding the pattern of urban development. However, in this context, there is no information on the mechanisms behind the observed biodiversity changes. One of the main mechanisms proposed to explain urban biodiversity loss is the alteration of predator-prey interactions. Using ground-nesting birds as a model system and data from nine European cities, we experimentally tested the effects of these two extreme urban development types on artificial ground nest survival and whether nest survival correlates with the local abundance of ground-nesting birds and their nest predators. Nest survival (n = 554) was lower in land-sharing than in land-sparing urban areas. Nest survival decreased with increasing numbers of local predators (cats and corvids) and with nest visibility. Correspondingly, relative abundance of ground-nesting birds was greater in land-sparing than in land-sharing urban areas, though overall bird species richness was unaffected by the pattern of urban development. We provide the first evidence that predator-prey interactions differ between the two extreme urban development types. Changing interactions may explain the higher proportion of ground-nesting birds in land-sparing areas, and suggest a limitation of the land-sharing model. Nest predator control and the provision of more green-covered urban habitats may also improve conservation of sensitive birds in cities. Our findings provide information on how to further expand our cities without severe loss of urban-sensitive species and give support for land-sparing over land-sharing urban development.

Body mass and territorial defence strategy affect the territory size of odonate species.

The territory is a distinct mating place that a male defends against intruding conspecific males. The size of a territory varies between species and most of the variation between species has been found to scale allometrically with body mass. The variation that could not be explained by body mass has been explained with several variables such as habitat productivity, trophic level, locomotion strategy and thermoregulation. All previous interspecific comparative studies have been done on vertebrate species such as birds, mammals, reptiles and fishes, meaning that studies using invertebrate species are missing. Here, we studied the relationship of a species's territory size with its fresh body mass (FBM) in addition to other ecologically relevant traits using 86 damselfly and dragonfly (Odonata) species. We found that territory size is strongly affected by species FBM, following an allometric relationship similar to vertebrates. We also found that the territory size of a species was affected by its territorial defence strategy, constantly flying species having larger territories than species that mostly perch. Breeding habitat or the presence of sexual characters did not affect territory sizes, but lotic species and species without wing spots had steeper allometric slopes. It seems that an increase in a species's body mass increases its territory size and may force the species to shift its territory defence strategy from a percher to a flier.

Brood parasitism in eusocial insects (Hymenoptera): role of host geographical range size and phylogeny.

Interspecific brood parasitism is common in many animal systems. Brood parasites enter the nests of other species and divert host resources for producing their own offspring, which can lead to strong antagonistic parasite-host coevolution. Here, we look at commonalities among social insect species that are victims of brood parasites, and use phylogenetic data and information on geographical range size to predict which species are most probably to fall victims to brood parasites in the future. In our analyses, we focus on three eusocial hymenopteran groups and their brood parasites: (i) bumblebees, (ii) Myrmica ants, and (iii) vespine and polistine wasps. In these groups, some, but not all, species are parasitized by obligate workerless inquilines that only produce reproductive-caste descendants. We find phylogenetic signals for geographical range size and the presence of parasites in bumblebees, but not in ants and wasps. Phylogenetic logistic regressions indicate that the probability of being attacked by one or more brood parasite species increases with the size of the geographical range in bumblebees, but the effect is statistically only marginally significant in ants. However, non-phylogenetic logistic regressions suggest that bumblebee species with the largest geographical range sizes may have a lower likelihood of harbouring social parasites than do hosts with medium-sized ranges. Our results provide new insights into the ecology and evolution of host-social parasite systems, and indicate that host phylogeny and geographical range size can be used to predict threats posed by social parasites, as well to design efficient conservation measures for both hosts and their parasites. This article is part of the theme issue 'The coevolutionary biology of brood parasitism: from mechanism to pattern'.

Phylogeny affects host's weight, immune response and parasitism in damselflies and dragonflies.

Host-parasite interactions are an intriguing part of ecology, and understanding how hosts are able to withstand parasitic attacks, e.g. by allocating resources to immune defence, is important. Damselflies and dragonflies show a variety of parasitism patterns, but large-scale comparative immune defence studies are rare, and it is difficult to say what the interplay is between their immune defence and parasitism. The aim of this study was to find whether there are differences in immune response between different damselfly and dragonfly species and whether these could explain their levels of gregarine and water mite parasitism. Using an artificial pathogen, a piece of nylon filament, we measured the encapsulation response of 22 different damselfly and dragonfly species and found that (i) there are significant encapsulation differences between species, (ii) body mass has a strong association with encapsulation and parasite prevalences, (iii) body mass shows a strong phylogenetic signal, whereas encapsulation response and gregarine and water mite prevalences show weak signals, and (iv) associations between the traits are affected by phylogeny. We do not know what the relationship is between these four traits, but it seems clear that phylogeny plays a role in determining parasitism levels of damselflies and dragonflies.

Risk of local extinction of Odonata freshwater habitat generalists and specialists.

Understanding the risk of a local extinction in a single population relative to the habitat requirements of a species is important in both theoretical and applied ecology. Local extinction risk depends on several factors, such as habitat requirements, range size of species, and habitat quality. We studied the local extinctions among 31 dragonfly and damselfly species from 1930 to 1975 and from 1995 to 2003 in Central Finland. We tested whether habitat specialists had a higher local extinction rate than generalist species. Approximately 30% of the local dragonfly and damselfly populations were extirpated during the 2 study periods. The size of the geographical range of the species was negatively related to extinction rate of the local populations. In contrast to our prediction, the specialist species had lower local extinction rates than the generalist species, probably because generalist species occurred in both low- and high-quality habitat. Our results are consistent with source-sink theory.

Parasite burden and the insect immune response: interpopulation comparison.

The immune response affects host's survival and reproductive success. Insurmountable immune function has not evolved because it is costly and there is a trade-off between other life-history traits. In previous studies several factors such as diet and temperature have been proposed to cause interpopulation differences in immune response. Moreover, the insect immune system may be functionally more protective upon secondary exposure, thus infection history may associate with the immune response. Here we measured how geographical location and parasite burden is related to variation in immune response between populations. We included 13 populations of the Northern Damselfly Coenagrion hastulatum (Odonata: Coenagrionidae) in Finland over a latitudinal range of 880 km to this study. We found that water mites associated strongly with the immune response at interpopulation level: the more the mites, the higher the immune response. Also, in an alternative model based on AIC, latitude and individual size associated with the immune response. In turn, endoparasitic gregarines did not affect the immune response. To conclude, a positive interpopulation association between the immune response and the rate of water mite infection may indicate (i) local adaptation to chronic parasite stress, (ii) effective 'induced' immune response against parasites, or (iii) a combined effect of both of these.

Wing shape and its influence on the outcome of territorial contests in the damselfly Calopteryx virgo.

Male mating success is often determined by territory ownership and traits associated with successful territory defense. Empirical studies have shown that the territory owner wins the majority of fights with challenging males. Several physical and physiological traits have been found to correlate with resource holding potential. In addition, in aerial insects, wing design may also have a strong influence on resource holding potential, since it determines efficiency and precision during flight. However, this possibility has not yet been thoroughly evaluated using the modern technique of geometric morphometrics to analyze shape. Therefore, this study examined whether wing shape affects the outcome of male-male contests in the territorial damselfly, Calopteryx virgo (L.) (Odonata: Calopterygidae). Wing shape and also traditional flight-related morphological measures were compared between 27 pairs of winners and losers from experimental territorial contests. Contrary to expectations, there were no differences between winners and losers in all studied wing traits (shape, length, width, total surface, aspect ratio, and wing loading). However, highly significant differences in wing shape and size were detected between the fore- and hindwing. It is currently not known how these differences relate to flight performance, since previous biomechanical studies in damselflies assumed fore- and hindwings to have an identical planform.

Predation selects for increased immune function in male damselflies, Calopteryx splendens.

Predation selects for numerous traits in many animal species, with sick or parasitized prey often being at high risk. When challenged by parasites and pathogens, prey with poor immune functions are thus likely to be at a selective disadvantage. We tested the hypothesis that predation by birds selects for increased immune function in a wild population of male damselflies Calopteryx splendens, while controlling for a trait known to be under selection by bird predation, dark wing-spots. We found that selection on both immune function and wing-spot size was significantly positive, and that selection on either trait was independent of selection on the other. We found no evidence of nonlinear quadratic or correlational selection. In contrast to previous studies, we found no phenotypic correlation between immune function and wing-spot size. There was also no difference in immune response between territorial and non-territorial males. Our study suggests that predation may be an important agent of selection on the immune systems of prey, and because the selection we detected was directional, has the potential to cause phenotypic change in populations.