Vertical stratification of ithomiine butterfly (Nymphalidae: Ithomiinae) mimicry complexes: the relationship between adult flight height and larval host-plant height

Data are presented which confirm previous findings that sympatric mimicry complexes dominated by unpalatable Neotropical ithomiine butterflies (Nymphalidae: Ithomiinae) are vertically stratified by height of flight. Flight height of ithomiine species is positively correlated with the height of their larval host-plants. Thus members of a mimicry complex utilize host-plants of similar heights. Non-mimetic British woodland butterflies also show a positive relationship between flight height and host-plant height, which suggests that the relationship is independent of mimicry. I propose that female butterflies fly at heights which maximize the probability of encountering their larval host-plants, and that males fly at similar heights to females in order to maximize the probability of encountering potential mates. Female butterflies probably encounter plants of similar heights to their larval hosts more frequently than they encounter plants of other heights. I suggest that butterfly species may therefore be more likely to make host shifts to plant species of a similar height to their current host-plants. Finally, I discuss how the relationship between flight height and height of larval host-plants, coupled with microhabitat-dependent selection on colour pattern, could lead to the evolution in sympatry of vertically stratified mimicry complexes.     

The number of butterfly species in woodlands

Summary The number of butterfly species present in 22 woods in eastern England was recorded during fortnightly visits between May to October 1978. A total of 26 species was observed. The number of species present was significantly correlated with the area of woodland but not with the area of glades and rides, which provided both larval food plants and adult foraging sites. The number of vagile species, but not the number of poorly vagile species, present was correlated with the area of woodland within 1 km of the study wood, this being a measure of the sources of potential colonizers. It was predicted that a woodland area of about 460 ha would be required to support all the species recorded in the study.     

Evolutionary responses by butterflies to patchy spatial distributions of resources in tropical environments

The greatest diversity of butterflies and their host plants occurs in tropical regions. Some groups of butterflies in the tropics exhibit monophagous feeding in the larval stage, exploiting only one family of plants; others are polyphagous, feeding on plants in two or more distinct families. The two major types of tropical habitats for butterflies, namely primary and secondary forests, offer very different evolutionary opportunities for the exploitation of plants as larval food. Butterflies are faced with the major logistical problem, as are many other herbivorous insects, of depositing eggs on the correct plant for successful larval feeding. This paper, using the concepts of phenotype set and spatial patchiness of resources, attemps to make some predictions as to the optimal phenotypic systems for monophagous and polyphagous feeding in tropical butterflies, as related to the spatial patchiness of larval host plants in primary and secondary forests. In addition to the secondary compound chemistry of larval host plants as playing a role in the evolution of monophagy and polyphagy, the assumption is made that the spatial patchiness of host plants within and among different families also acts as a major factor in determining optimal ranges of phenotypes for different patterns of larval feeding. Owing to the high spatial patchiness of primary forest species of canopy trees and vines, it is predicted that butterflies exploiting these will be mostly polyphagous, whereas secondary forests having stable formations of fewer plant species and larger patches of these plants, will have mostly monophagous species. Forest understories may have both monophagous and polyphagous species, depending upon the layer of forest and the general type of understory (i.e. palmaceous or dicotyledonous). Field data on some groups of butterflies from tropical America support these predictions. Polyphagous butterflies are predicted to possess a genetic system of mixed morphs with a population being polymorphic as a whole; monophagous butterflies are predicted to have individuals all more or less similar genetically, and with a high amount of genic variation within individuals. Other forms of monophagy may evolve in species that are essentially monomorphic but with various mechanisms (physiological, developmental, behavioral) of phenotypic flexibility at the individual level. Although the environment is essentially coarse-grained for larvae since most are sedentary and polymorphism is an optimal adaptive strategy, the oviposition strategy of the adult must also be considered and some situations (i.e. forest canopy) have resources (host plants) distributed in a fine-grained fashion. Other forms of limited polyphagy may result from monomorphic genetic systems in which there is considerable phenotypic flexibility.     

Habitat complementarity and butterfly traits are essential considerations when mitigating the effects of exotic plantation forestry

Habitat loss threatens insect diversity globally. However, complementary vegetation types in remaining habitat increases opportunities for species survival. We assess the extent to which indigenous forest patches moderate the impact of exotic commercial afforestation on grassland butterflies. Butterflies were sampled in grassland along uncorrelated gradients of landscape-scale indigenous forest and plantation cover, while controlling for variation in local vegetation composition. We separately assessed responses by butterfly groups differing in habitat preference, larval diet, and mobility. There was no effect of landscape- or local-scale variables on species richness, but there was a strong interactive effect of forest and plantation cover on butterfly assemblage structure. The effect varied according to species traits. When forest cover was high, assemblages did not differ at different levels of plantation cover. However, plantation cover significantly influenced assemblage structure when forest cover was low. Grassland with limited forest cover in the protected area supported unique assemblages with high frequency of less mobile, specialized species with herbaceous larval host plants, whereas grassland with low forest cover near plantations had a prevalence of mobile, generalist species. A positive association between forest cover and butterflies with woody larval host plants suggests that indigenous forest patches improved the suitability of fragmented grassland for a subset of butterflies, emphasising the value of natural heterogeneity in transformed areas. However, certain butterfly traits associated with large, open grassland were under-represented in grassland between plantations, underscoring the importance of open areas in the broader landscape to conserve the full diversity of species.

     

Spillover of Insects from Rain Forest into Adjacent Oil Palm Plantations

Conversion of natural forest to oil palm plantations is a major threat to biodiversity in Southeast Asia. The retention of natural forest habitats within plantations has been proposed as a method to reduce biodiversity losses in agricultural areas, and we examined whether forest areas resulted in spillover of species into adjacent oil palm plantations. We sampled ants and butterflies along two 2‐km transects across an ecotone from plantation into adjacent forest in Sabah, Malaysian Borneo. Species richness of both taxa was reduced in plantations, but to a greater extent in butterflies (54% reduction) than in ants (25% reduction). Butterfly diversity increased in plantations with increasing proximity to forest primarily due to spillover of ‘vagrant’ forest species (whose larval host plants do not occur in plantations), although richness of species that could potentially breed in plantations also increased near to forest. By contrast, ants showed no spillover effects and were less sensitive to land‐use changes, with much higher levels of similarity in species assemblages across habitats than for butterflies. Our results for butterflies suggest that despite the negative impacts of plantations on diversity, proximity to forest could improve diversity in adjacent plantations for some taxa. Spillover of forest species implies that retaining forest areas within plantations may be important for facilitating dispersal of some species through the landscape.     

The adaptiveness of searching and host selection behaviour in Pieris rapae (L.)

This study evaluates the adaptive significance of host preferences and searching behaviour in Vancouver and Canberra populations of the cabbage butterfly Pieris rapae (L.). As a result of a complex of responses to plant age, the butterflies concentrate their eggs on middle-aged plants. Young larvae develop faster and survive better on young plants than old ones, but larvae on smaller plants are more susceptible to crowding effects. Thus a preference for plants which are well-grown but not too old is selectively advantageous. By contrast, the butterflies’ host species preferences appear non-adaptive, and are unrelated to the quality of the host as larval food. Vancouver butterflies change their flight direction often and are very responsive to hosts, thereby generating a very aggregated distribution at a low cost in flight time. Canberra butterflies tend to fly in straight lines and are less responsive to hosts; their egg distribution is consequently more nearly random, but they fly further for each egg they lay. The relative costs of aggregation and increased flight time differ between the populations in a manner consistent with the observed behavioural differences.     

Oviposition preference and larval performance of North American monarch butterflies on four Asclepias species

Monarch butterflies, Danaus plexippus L. (Lepidoptera: Nymphalidae), occur world‐wide and are specialist herbivores of plants in the milkweed family (Asclepiadaceae). In North America, two monarch populations breed east and west of the continental divide in areas populated by different host plant species. To examine the population variation in monarch responses to different Asclepias species, we measured oviposition preference and larval performance among captive progeny reared from adult butterflies collected in eastern and western North America. Host plant use was evaluated using two milkweed species widely distributed in eastern North America (A. incarnata and A. syriaca), and two species common to western North America (A. fascicularis and A. speciosa). We predicted that exposure to different host plant species in their respective breeding ranges could select for divergent host use traits, so that monarchs should preferentially lay more eggs on, and larvae should perform better on, milkweed species common to their native habitats. Results showed that across all adult female butterflies, oviposition preferences were highest for A. incarnata and lowest for A. fascicularis, but mean preferences did not differ significantly between eastern and western monarch populations. Larvae from both populations experienced the highest survival and growth rates on A. incarnata and A. fascicularis, and we again found no significant interactions between monarch source population and milkweed species. Moreover, the average rank order of larval performance did not correspond directly to mean female oviposition preferences, suggesting that additional factors beyond larval performance influence monarch oviposition behavior. Finally, significant family level variation was observed for both preference and performance responses within populations, suggesting an underlying genetic variation or maternal effects governing these traits.     

Relationship of butterfly diversity with nectar plant species richness in and around the Aokigahara primary woodland of Mount Fuji,central Japan

We examined the relationships between the diversities of vegetation, adult nectar plants, and butterflies in and around the Aokigahara primary woodland on the northwestern footslopes of Mount Fuji, central Japan. The results showed that the nectar resource utilization by adult butterflies was significantly biased to herbaceous plants, especially to perennials, compared to woody species, although most of the study area was in and near a primary woodland. There were greater nectar plant species in sites with greater plant species richness. Among the butterfly community indices analyzed, the strongest correlation was detected between butterfly species richness and nectar plant species richness at each site. Another close correlation was detected between the species richness of nectar plants and herbaceous plants at each site. These results suggest that herbaceous plant species richness in a habitat plays a central role in its nectar plant species richness, and the nectar plant richness is a highly important factor supporting its adult butterfly species richness. Consequently, we propose that the maintenance and management of herbaceous plant species richness in a butterfly habitat, which lead to those of its nectar plant species richness, are very important for conservation of butterfly diversity even in and around woodland landscapes of temperate regions.     

Progressive bias in species status is symptomatic of fine-grained mapping units subject to repeated sampling

Atlas maps of butterflies invariably fail to distinguish the status of records, that is, whether they are observations of breeding populations or vagrant individuals. Yet, for conservation purposes, it is clearly important to know whether records relate to breeding populations in suitable habitats or not. The high mobility of butterfly adults carries two expectations. First, vagrants will frequently be recorded within grid squares, particularly when mapping is fine grained. Second, the frequency of vagrants is likely to be higher in mapping units comprising biotopes with few butterfly habitats, than in those rich in butterfly habitats, if only because there is a greater regional pool of potential vagrants for biotopes depleted in butterfly habitats. It follows that with repeated sampling of squares or biotopes differences in species breeding status will be cumulative between those rich and poor in butterfly habitats. In this paper, these predictions have been tested using data from 30, 1 ha grid squares on Alderley Edge Site of Special Scientific Interest (SSSI) (UK), half of which are in woodland (few butterfly habitats) and half with open habitats (non-woodland; rich in butterfly habitats). Butterflies have been recorded during 38 visits over 4 years. Within squares, vagrants have been distinguished from breeding species on the basis of the presence of larval hostplants and other suitable resources for seasonal maintenance. Owing to the occurrence of vagrants, number of breeding species was found to be significantly fewer than total species in mapping units. Vagrants were found to be significantly more abundant in woodland squares, which contain few species larval hostplants, than in squares dominated by open biotopes with many species larval hostplants. Repeated sampling resulted in a cumulative increase in the frequency of vagrant species. Furthermore, the relative frequency of vagrants increased at a greater rate in woodland squares compared to non-woodland squares. Suggestions are made as to how recording for atlases may be improved.     

Resource-based habitat definition, niche overlap and conservation of two sympatric glacial relict butterflies

The precise knowledge of ecological resources and conditions required by species threatened by rapidly changing environmental conditions is of prime importance for conservation biology. Transferability of this knowledge between species with similar ecological requirements is often assumed, but rarely tested. This is especially the case for glacial relict populations confined to climate‐habitat traps from where they cannot move to rejoin areas with suitable environmental conditions. Using two glacial relict butterflies as model organisms, we first quantitatively define larval and adult resource‐based habitat use of each species. Secondly, we test the transferability of ecological profiles (both habitat and ecological niche) between these two species that share both the same biotope and the same host plant. Our results show that both species have markedly different ecological requirements relating to differences in life history and behavioural traits (i.e. egg‐laying strategies and mate‐locating behaviour). Although the two species share many ecological features, they use different functional habitats within our study site. The high degree of interspecific niche overlap should indicate a high interspecific competition. However, we argue that their co‐existence can be explained by the non‐limiting abundance of some resources (e.g. host plants), by the partial separation in time of adult flight periods and by the territorial behaviour of one of the species. We discuss the following general messages: (1) functional habitat of a (threatened) species should be defined in a spatial context corresponding to individual station keeping, and (2) quick diagnosis based on similar ecological requirements may be misleading for the design of reliable conservation and restoration strategies. Detailed mechanistic and quantitative ecological understanding of resource‐use and environmental tolerances across an organism's life cycle is essential for effective conservation in changing environments, like for glacial relict species.     

A study of fungus flies (Dipteral Mycetophilidae) in beech woodland

Abstract. 1. A study was made of larval and adult Mycetophilidae in coppiced beech woodland in southern England. Adult mycetophilids were trapped throughout 1968 using sticky traps at ground level. Larvae inhabiting terrestrial fungi were collected from all sporophores from a defined area of the woodland floor between August and October 1968.
2. Larval mycetophilids were extracted from only 12.6% of all sporophores collected and from fifteen of thirty-eight species of fungi present. Some species of fungi were never inhabited by larval mycetophilids while others appeared to be highly attractive to them. Since it was not possible to identify the larvae beyond the level of tribe it was impossible to ascertain host preferences for different species. Peak numbers of larvae occurred at the beginning of September and the beginning of October.
3. Adult mycetophilids had two peaks of activity, the first in March and April when the dominant species were Boletina gripha and Phronia basalis and the second in autumn when species of the genus Mycetophila were dominant. Phronia basalis is a species whose larvae inhabit dead wood and the abundance of this species (40% of all adults) probably reflects the amount of rotting wood in this type of habitat. The autumn peak of adult activity was due to species known to inhabit agaric sporophores and came about 1 month after the peak of larval numbers in fungi.
4. The adult fauna was rich with 107 species representing about a quarter of the known British fauna. However, only twelve species were trapped in sufficient numbers to allow deductions concerning seasonal activity.
5. The results were discussed with respect to the problems of assessing populations of insects inhabiting fungal sporophores and to the previous work on this neglected group.     

N equals two (times five). Exploring the effects of horse rewilding on five congeneric adult butterflies

Rewilding incomplete ecosystems by using ungulate megaherbivores represents a significant potential for sustainable management of habitats of declining species. Two xeric grasslands patches in the Podyjí National Park, Czech Republic, were rewilded by a feral horse breed, the Exmoor pony, in 2018. Before this in 2017, demography, mobility, and adult habitat use of five congeneric Melitaea butterflies co-occurring at the grasslands were investigated (Vodickova et al., J. Nature Conserv. 2019). In 2021, four seasons after the rewilding, we replicated the survey to assess the effects of the horse on the butterflies. Here, we compare the results of the two surveys and investigate changes in spatial patterns of adult distribution using Ripley’s K-functions.Total numbers of captures, and estimated population sizes, were consistently lower in 2021, with the largest drop for spring-flying M. cinxia. We cannot discern whether this was due to the cold 2021 spring, or due to reduction of grasses by the horse, possibly contributing to desiccation of M. cinxia host plants. Demographic parameters such as residency/longevity and capture probability changed only little. Mobility ranking among species remained identical, but within species, some mobility characteristics changed among years. Among early summer species, M. britomartis, second most abundant in 2017, switched to the first position with M. aurelia, and these two species displayed the most notable shift in adult habitat use between the two seasons. Short thorny shrubs avoided by the horse protect M. britomartis host plants; this threatened butterfly thus did not suffer from horse presence. M. athalia, a species of woodland edges, profited from decay of conifers caused by a series of dry years; and M. didyma, forming multiple generations, from increase of its host plant. Contrary to expectations, spatial distribution of most butterflies became more aggregated within rewilded pastures, probably due to regularities in home ranges use by the horses, which restructured the vegetation in a zonal, rather than patchy, way. A considerably larger areas should be rewilded by the herbivores to fully achieve the desired beneficial effects.     

Butterflies as an indicator group of riparian ecosystem assessment

Riparian ecosystems play an important role in modulating a range of ecosystem processes that affect aquatic and terrestrial organisms. Butterflies are a major herbivore in terrestrial ecosystems and are also common in riparian ecosystems. Since butterflies use plants for larval food and adult nectar sources in riparian ecosystems, butterfly diversity can be utilized to evaluate riparian ecosystems. We compiled butterfly data from 33 sites in three riparian ecosystem types across the country and compared butterfly diversity in terms of number of species and quality index in relation to riparian environmental variables. Number of butterfly and plant species was not different among three riparian habitat types. Additionally, there was no significant ecological variable to distinguish the butterfly communities on three riparian habitats. Non-metric multi-dimensional scaling ordination showed that butterfly communities in three riparian ecosystem types differed from each other, and butterfly riparian quality index was the main variable for butterfly assemblages. Five indicator species for moor and another five species for riverine riparian ecosystems were identified. Three and one indicator species for moor and riparian ecosystems, respectively, were plant specialists, while 44 butterflies were general feeders, feeding on a wide range of hostplants in several habitats. These results suggest that butterfly species use actively riparian habitats for nectar and larval food, and the butterfly riparian quality index can be employed to track faunal change in riparian habitats, which are frequently threatened by disturbances such as water level and climate changes, and invasive species.     

Butterfly species and traits associated with selectively logged forest in Borneo

Logging can significantly change the structure of rainforest communities. To better understand how logging drives this change, butterflies and environmental variables were assessed within both unlogged and logged forest in Indonesian Borneo. In the whole dataset, we found local environmental variables and geographic distance combined captured 53.1% of the variation in butterfly community composition; 29.6% was associated with measured local environmental variables, 13.6% with geographic distance between sites, and 9.9% with covariation between geographic distance and environmental variables. The primary axis of variation in butterfly community composition represented a disturbance gradient from unlogged to logged forest. Subsequent axes represented gradients influenced by variables such as canopy cover and total tree density. There were significant associations between environmental variables and geographic range and larval host plant use of species. Specifically, butterflies using trees as larval host plants and those with distributions limited to Borneo were more likely to be present in unlogged forest. By contrast, species that tended to be more abundant in logged forest were those with widespread distributions and those using lianas and grasses as larval host plants. The results of this study highlight the importance of environmental variables and disturbance, e.g., selective logging, in structuring rainforest community diversity. Moreover, they confirm how species traits, such as larval food use and geographic distributions can determine patterns of species abundance following environmental change.     

The effect of coppice management on moth assemblages in an English woodland

Coppice woodlands in Britain may become the target of increased management due to the rise in demand for woodfuel. The biodiversity value of sweet chestnut (Castanea sativa) coppice and the effect of coppice management upon this has received limited study. Moths were the focus of this study in an actively coppiced sweet chestnut woodland in southern England. Coupes, with between one and 20 years of coppice regrowth, were systematically sampled for night flying micro- and macro-moths and coppice structure and ground vegetation were described. Using differences in moth assemblage, three stages of coppice development were distinguished: with one-four, five-nine and more than 10 years of coppice regrowth. Differences in moth assemblage related to habitat conditions within each coppice stage. The young coppice stage moth assemblage was characterised by species typically associated with open habitats; moths of the middle coppice stage assemblage fed on trees and were species typically associated with open woodland and scrub habitats; moths of the mature coppice stage assemblage were species typically associated with closed canopy woodland and contained specialist species whose larval food consists of material such as lichen and decaying leaves. All three coppice stages supported species of listed conservation status; the mature coppice stage contained a distinctive range of scarce and threatened species. The study showed that active coppicing promotes a change in moth assemblage but consequently will temporarily eliminate many species of mature stage coppice. Management which provides a range of coppice age classes within a woodland, appears key in promoting moth diversity.     

Metamorphosis of a Butterfly-Associated Bacterial Community

Butterflies are charismatic insects that have long been a focus of biological research. They are also habitats for microorganisms, yet these microbial symbionts are little-studied, despite their likely importance to butterfly ecology and evolution. In particular, the diversity and composition of the microbial communities inhabiting adult butterflies remain uncharacterized, and it is unknown how the larval (caterpillar) and adult microbiota compare. To address these knowledge gaps, we used Illumina sequencing of 16S rRNA genes from internal bacterial communities associated with multiple life stages of the neotropical butterfly Heliconius erato. We found that the leaf-chewing larvae and nectar- and pollen-feeding adults of H. erato contain markedly distinct bacterial communities, a pattern presumably rooted in their distinct diets. Larvae and adult butterflies host relatively small and similar numbers of bacterial phylotypes, but few are common to both stages. The larval microbiota clearly simplifies and reorganizes during metamorphosis; thus, structural changes in a butterfly''s bacterial community parallel those in its own morphology. We furthermore identify specific bacterial taxa that may mediate larval and adult feeding biology in Heliconius and other butterflies. Although male and female Heliconius adults differ in reproductive physiology and degree of pollen feeding, bacterial communities associated with H. erato are not sexually dimorphic. Lastly, we show that captive and wild individuals host different microbiota, a finding that may have important implications for the relevance of experimental studies using captive butterflies.     

Role of larval host plants in the climate-driven range expansion of the butterfly Polygonia c-album

1. Some species have expanded their ranges during recent climate warming and the availability of breeding habitat and species' dispersal ability are two important factors determining expansions. The exploitation of a wide range of larval host plants should increase an herbivorous insect species' ability to track climate by increasing habitat availability. Therefore we investigated whether the performance of a species on different host plants changed towards its range boundary, and under warmer temperatures. 2. We studied the polyphagous butterfly Polygonia c-album, which is currently expanding its range in Britain and apparently has altered its host plant preference from Humulus lupulus to include other hosts (particularly Ulmus glabra and Urtica dioica). We investigated insect performance (development time, larval growth rate, adult size, survival) and adult flight morphology on these host plants under four rearing temperatures (18-28.5 degrees C) in populations from core and range margin sites. 3. In general, differences between core and margin populations were small compared with effects of rearing temperature and host plant. In terms of insect performance, host plants were generally ranked U. glabra > or = U. dioica > H. lupulus at all temperatures. Adult P. c-album can either enter diapause or develop directly and higher temperatures resulted in more directly developing adults, but lower survival rates (particularly on the original host H. lupulus) and smaller adult size. 4. Adult flight morphology of wild-caught individuals from range margin populations appeared to be related to increased dispersal potential relative to core populations. However, there was no difference in laboratory reared individuals, and conflicting results were obtained for different measures of flight morphology in relation to larval host plant and temperature effects, making conclusions about dispersal potential difficult. 5. Current range expansion of P. c-album is associated with the exploitation of more widespread host plants on which performance is improved. This study demonstrates how polyphagy may enhance the ability of species to track climate change. Our findings suggest that observed differences in climate-driven range shifts of generalist vs. specialist species may increase in the future and are likely to lead to greatly altered community composition.     

Life history of the Glanville fritillary butterfly in fragmented versus continuous landscapes

Habitat loss and fragmentation threaten the long‐term viability of innumerable species of plants and animals. At the same time, habitat fragmentation may impose strong natural selection and lead to evolution of life histories with possible consequences for demographic dynamics. The Baltic populations of the Glanville fritillary butterfly (Melitaea cinxia) inhabit regions with highly fragmented habitat (networks of small dry meadows) as well as regions with extensive continuous habitat (calcareous alvar grasslands). Here, we report the results of common garden studies on butterflies originating from two highly fragmented landscapes (FL) in Finland and Sweden and from two continuous landscapes (CL) in Sweden and Estonia, conducted in a large outdoor cage (32 by 26 m) and in the laboratory. We investigated a comprehensive set of 51 life‐history traits, including measures of larval growth and development, flight performance, and adult reproductive behavior. Seventeen of the 51 traits showed a significant difference between fragmented versus CL. Most notably, the growth rate of postdiapause larvae and several measures of flight capacity, including flight metabolic rate, were higher in butterflies from fragmented than CL. Females from CL had shorter intervals between consecutive egg clutches and somewhat higher life‐time egg production, but shorter longevity, than females from FL. These results are likely to reflect the constant opportunities for oviposition in females living in continuous habitats, while the more dispersive females from FL allocate more resources to dispersal capacity at the cost of egg maturation rate. This study supports theoretical predictions about small population sizes and high rate of population turnover in fragmented habitats selecting for increased rate of dispersal, but the results also indicate that many other life‐history traits apart from dispersal are affected by the degree of habitat fragmentation.     

Ecology of tropical butterflies in rainforest gaps

Tropical forest gaps are ephemeral and patchily distributed within forest areas and have very different light environments compared with closed-canopy forest. We used fruit-baited traps to investigate if gaps are exploited by more opportunistic butterfly species compared with closed-canopy forest. Gaps supported a higher diversity of butterflies in terms of species evenness but closed-canopy sites contained species with more restricted geographical distributions. There was little similarity between the assemblages of butterflies trapped in the canopy and those in either gap or closed-canopy sites, but the greater similarity was with gaps, and increased diversity in gaps was partly due to canopy species turning up in gaps. Dispersal rates (as measured by recapture rates) were higher in gaps and there was evidence that butterflies in gaps had relatively larger and broader thoraxes, indicating a flight morphology adapted for faster flight. These results support the notion of a distinctive gap fauna comprising more widespread, mobile species. Habitat modification that opens up the canopy is likely to result in an increase in these widespread species and a decline in understorey species with restricted distributions.     

The impact of forest spread on a marginal population of a protected peony (<Emphasis Type="Italic">Paeonia officinalis</Emphasis> L.): the importance of conserving the habitat mosaic

In the Mediterranean region of Europe, land-use changes have allowed for rapid colonisation of open habitats by woody species. As a result, it is critical to gather information on how protected species in open habitats respond to forest spread in such areas. Our objective is to quantify whether spatial heterogeneity of the vegetation associated with recent forest closure influences demographic structure and maternal fertility in a population of the protected Paeonia officinalis L. In closed woodland, adult plants of P. officinalis are almost exclusively vegetative, in open habitats seedlings are rare and on the woodland edge there is a relative over-representation of flowering plants and seedlings. Forest closure dramatically reduces flowering frequency, but has no significant effect on maternal fertility of flowering plants. The spatial aggregation of seedlings close to the maternal plants suggests that dispersal is spatially restricted. Together, these results suggest that the viability of the population requires a transitional habitat between open garrigues or grassland with spaced trees and woodland. A management programme incorporating tree and shrub thinning and cutting of parcels in rotation to maximise the length of the forest edge could maintain a habitat mosaic that favours the persistence of this species in the study site.