Host associations of <Emphasis Type="Italic">Coenonympha hero</Emphasis> (Lepidoptera: Nymphalidae) in northern Europe: microhabitat rather than plant species

Understanding ecological requirements of endangered species is a primary precondition of successful conservation practice. Regrettably, we know surprisingly little about the life history of numerous threatened insects, and about their use of larval host plants in particular. The brown butterflies (Nymphalidae: Satyrinae) have traditionally been considered polyphagous on grasses and indiscriminatory in their oviposition behavior. However, detailed studies on several species have revealed local specialization in host plant use as well as the decisive role of microlimatic conditions as determinants of habitat quality. The present study addresses host plant relationships in the endangered brown butterfly Coenonympha hero (L.) at the northern limit of its European distribution. We combine laboratory-based host preference and performance tests with an analysis of microhabitat use by adult butterflies in the field. Both lines of evidence suggest that C. hero is polyphagous enough not to be associated with one particular host species. Oviposition choices of C. hero are not driven by host plant species but rather by structural characteristics of the substrate. The preferred rigid needle-like structures may serve as cues of ‘transparent’ vegetation which allows the larvae to benefit from sunlight reaching the lower strata of the tuft. Our results suggest that conservation efforts should prioritize microclimatic parameters, rather than the presence of any particular host plant species, as decisive determinants of habitat quality in C. hero.     

Phenological matching rather than genetic variation in host preference underlies geographical variation in host plants used by orange tip butterflies

An insect species that shows variation in host species association across its geographical range may do so either because of local adaptation in host plant preference of the insect or through environmentally or genetically induced differences in the plants, causing variation in host plant suitability between regions. In the present study, we experimentally investigate the host plant preference of Anthocharis cardamines (orange tip butterfly) in two populations from the UK and two from Sweden. Previous reports indicate that A. cardamines larvae are found on different host plant species in different regions of the UK, and some variation has been reported in Sweden. Host plant choice trials showed that females prefer to oviposit on plants in an earlier phenological stage, as well as on larger plants. When controlling for plant phenological stage and size, the host species had no statistically significant effect on the choice of the females. Moreover, there were no differences in host plant species preference among the four butterfly populations. Based on our experiment, the oviposition choice by A. cardamines mainly depends on the phenological stage and the size of the host plant. This finding supports the idea that the geographical patterns of host–plant association of A. cardamines in the UK and Sweden are consequences of the phenology and availability of the local hosts, rather than regional genetic differences in the host species preference of the butterfly.     

Restoration of an endangered plant,Hygrophila pogonocalyx,leads to an adaptive host shift of the chocolate pansy (Junonia iphita iphita)

Anthropogenic introduction of a plant species may cause novel encounters between the plant and local herbivores, and initiate evolutionary changes in host plant usage by herbivores. Until recently the endemic aquatic plant Hygrophila pogonocalyx was endangered and had a restricted distribution in Taiwan. Massive restoration efforts since 1997 have led to an expansion of the plant's distribution and a novel encounter between it and an Asian butterfly, the chocolate pansy, Junonia iphita (Nymphalidae). This butterfly appears to have colonized H. pogonocalyx, switching from its original host, Strobilanthes penstemonoides var. formosana. In the present study, we aimed to investigate whether the utilization of H. pogonocalyx as a host plant has initiated a differentiation between butterflies using the novel and the original hosts. To this purpose we collected butterflies from patches of the two host plants which grow sympatrically. We tested oviposition preference for the two hosts and larval performance on them. Female adults exhibited distinct oviposition preference toward the host plant their mothers preferred. Offspring showed greater survivorship and pupal weight when fed on the host plant their mothers preferred. Male adults displayed territorial behaviors on the host plant that their mothers had preferred. Finally, the survival rate of offspring produced from cross-mating between individuals with different host plant preference was lower than that of non-hybrids. Taken together, we suggest that genetic differentiation has occurred between individuals preferring H. pogonocalyx versus S. penstemonoides as host plants via host shifting. This process was likely induced by the mass restoration of the formerly rare and endangered plant species.     

Innate preference hierarchies coupled with adult experience,rather than larval imprinting or transgenerational acclimation,determine host plant use in Pieris rapae

The evolution of host range drives diversification in phytophagous insects, and understanding the female oviposition choices is pivotal for understanding host specialization. One controversial mechanism for female host choice is Hopkins’ host selection principle, where females are predicted to increase their preference for the host species they were feeding upon as larvae. A recent hypothesis posits that such larval imprinting is especially adaptive in combination with anticipatory transgenerational acclimation, so that females both allocate and adapt their offspring to their future host. We study the butterfly Pieris rapae, for which previous evidence suggests that females prefer to oviposit on host individuals of similar nitrogen content as the plant they were feeding upon as larvae, and where the offspring show higher performance on the mother's host type. We test the hypothesis that larval experience and anticipatory transgenerational effects influence female host plant acceptance (no‐choice) and preference (choice) of two host plant species (Barbarea vulgaris and Berteroa incana) of varying nitrogen content. We then test the offspring performance on these hosts. We found no evidence of larval imprinting affecting female decision‐making during oviposition, but that an adult female experience of egg laying in no‐choice trials on the less‐preferred host Be. incana slightly increased the P. rapae propensity to oviposit on Be. incana in subsequent choice trials. We found no transgenerational effects on female host acceptance or preference, but negative transgenerational effects on larval performance, because the offspring of P. rapae females that had developed on Be. incana as larvae grew slower on both hosts, and especially on Be. incana. Our results suggest that among host species, preferences are guided by hard‐wired preference hierarchies linked to species‐specific host traits and less affected by larval experience or transgenerational effects, which may be more important for females evaluating different host individuals of the same species.     

Extinction-colonization dynamics and host-plant choice in butterfly metapopulations

Species living in highly fragmented landscapes often occur as metapopulations with frequent population turnover. Turnover rate is known to depend on ecological factors, such as population size and connectivity, but it may also be influenced by the phenotypic and genotypic composition of populations. The Glanville fritillary butterfly (Melitaea cinxia) in Finland uses two host-plant species that vary in their relative abundances among distinct habitat patches (dry meadows) in a large network of approximately 1,700 patches. We found no effect of host species use on local extinction. In contrast, population establishment was strongly influenced by the match between the host species composition of an empty habitat patch and the relative host use by larvae in previous years in the habitat patches that were well connected to the target patch. This "colonization effect" could be due to spatially variable plant acceptability or resistance or to spatially variable insect oviposition preference or larval performance. We show that spatial variation in adult oviposition preference occurs at the relevant spatial scale and that the other possible causes of the colonization effect can be discounted. We conclude that the colonization effect is generated by host preference influencing the movement patterns of ovipositing females. Migrant females with dissimilar host preferences have different perceptions of relative patch quality, which influences their likelihood of colonizing patches with particular host composition.     

Host plant use by the Heath fritillary butterfly, Melitaea athalia: plant habitat, species and chemistry

We present a study of habitat use, oviposition plant choice, and food plant suitability for the checkerspot butterfly Melitaea athalia Rottemburg (Lepidoptera: Nymphalidae) in Åland, Finland. We found that in Åland, unlike in the mainland of Finland and many parts of its range, M. athalia flies mainly in open meadows. When offered an array of plants in a large (32 × 26 m) field cage, they predominately oviposited upon Veronica chamaedrys L., V. spicata L. and Plantago lanceolata L. (Plantaginaceae), which grow in open meadows. The relative abundance of the butterfly in Åland, and its habitat and host plant use there, may reflect local adaptation to land use practices and geology that maintain clusters of small open meadows with little successional change. At the scale of a plant patch, preferred species were used as frequently in mixed species patches as in mono-specific patches, and more oviposition occurred in open than in grassy patches. All of the host plants used by M. athalia are defended by iridoid glycosides (IGs). However, oviposition choice among species and among individual plants within species was largely independent of IG concentration. This contrast with the more discerning congener, M. cinxia, supports the idea that host discrimination decreases with increasing host range. Finally, although the adult butterflies chose specific plant species for oviposition, as larvae they performed well on twelve out of thirteen species of plants, including both known hosts and related novel plants that occur in Åland, indicating a much wider range of larval food plant species than adult oviposition species.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Specific network and phylosymbiosis pattern in endophyte community of coastal halophytes

Halophytes colonize stressed environments of high salinity. Endophytic symbionts improve growth performance and thus adaptability of host plants in stressed environments. Nevertheless, studies focused on the distribution and assembly patterns of fungal endophyte assemblages of halophytes in high salinity environments are limited. We selected 27 common non-mangrove halophytes across 4 geographic sites along the eastern coastline of China for our investigation on endophyte assemblages. We found a significant amount of basidiomycetous fungi in the endophyte assemblage. The endophyte community compositions were significantly affected by plant species and geographic locations. Some halophyte hosts showed significant preference for endophytic fungi in host preference analysis. The network structures of host-microbe bipartite graphs at each site had low connection and were highly nested, modular and specialized. Current results also indicated significant correlation and congruency between host phylogeny and compositions of endophyte local communities and meta-community. Our research showed that host is the dominant factor shaping the fungal endophytic communities in aerial tissues of halophytes in high salinity environments, and the host influence on endophytic community relates to plant phylogeny.     

Spatial and temporal patterns of caterpillar performance and the suitability of two host plant species

Abstract. 1. The butterfly Melitaea cinxia uses two host plant species in the Åland Islands of south-west Finland. Survey data show that host plant use is spatially variable and that the two species are not used (fed on by M . cinxia larvae) in proportion to their abundances. The pattern of host plant use by M . cinxia has been attributed in part to plant distribution and spatial variation of butterfly oviposition preference.
2. The additional roles that may be played by spatial variation in host plant quality and larval physiology (performance ability) were investigated. Seven years of field survey data and a series of laboratory experiments were used to demonstrate that neither of these variables contributed measurably to the observed pattern of host plant use.
3. Specifically, while there was great variation among individuals in both the performance ability of caterpillars and host plant suitability, the two plant species appeared equally suitable, there was no within-species geographic variation in plant suitability, and there was no evidence for adaptation of caterpillars to the locally used host plants.     

Patch dynamics ofGlaucopsyche lygdamus (Lycaenidae): correlations between butterfly density and host species diversity

Glaucopsyche lygdamus egg densities were surveyed over a 2000-m section of Gold Creek and at 30 different isolated patches in the Gold Basin drainage in Colorado. Host plant numbers and diversity were quantified, as well as other variables potentially influencing butterfly population size, such as patch size and isolation. Egg densities correlated significantly only with measures of host species diversity. Patches consisting of a single host species, no matter how large, did not support high butterfly densities, but patches of multiple, equitably distributed host species did. The most likely explantation, in light of oviposition preference and larval performance data accumulated for this butterfly species, is that host species diversity is necessary for the persistence ofG. lygdamus populations, because alternative host species buffer population losses during poor or unusual years. The dependence of both ovipositing butterflies and developing larvae on the ephemeral, young, host plant flowers make the butterfly especially vulnerable to year-to-year variation in host plant availability and quality.     

PREFERENCE HIERARCHIES AND THE ORIGIN OF GEOGRAPHIC SPECIALIZATION IN HOST USE IN SWALLOWTAIL BUTTERFLIES

Four allopatric populations of the widely distributed western anise swallowtail butterfly, Papilio zelicaon, use different plant genera as hosts, but simultaneous choice experiments showed that these populations have diverged only slightly in oviposition preference. Of the four populations—two from southeastern Washington State, one from coastal southwestern Washington State, and one from central California—three use hosts that are not available to any of the others. Although variation for the degree of preference for particular plant species occurred within and among populations, all four populations ranked hosts in the same overall order. Monophagy on a local, low-ranking host outside the range of high-ranking hosts did not necessarily lead to the loss of preference for those high-ranking hosts, thereby indicating that the high-ranking hosts would still be accepted, and in some cases even preferred, if a population encountered them again. Hence, the overall preference hierarchy among P. zelicaon populations appears to be evolutionarily conservative. Analyses of differences among families within the California population indicated that increased preference for some hosts is inversely correlated, whereas preference for other hosts may be uncorrelated. Positive correlations may also occur but were not observed among the plant species tested. Overall, the results indicate local monophagy on different plant species in P. zelicaon has not involved major reorganizations in the preference hierarchy of ovipositing females, even in populations that may have fed on a low-ranking host for many generations. Instead, small increases in preference for local hosts have occurred within an evolutionarily conservative preference hierarchy.     

Hidden diversity and evolutionary trends in malacosporean parasites (Cnidaria: Myxozoa) identified using molecular phylogenetics

Malacosporeans represent a small fraction of myxozoan biodiversity with only two genera and three species described. They cycle between bryozoans and freshwater fish. In this study, we (i) microscopically examine and screen different freshwater/marine fish species from various geographic locations and habitats for the presence of malacosporeans using PCR; (ii) study the morphology, prevalence, host species/habitat preference and distribution of malacosporeans; (iii) perform small subunit/large subunit rDNA and Elongation factor 2 based phylogenetic analyses of newly gathered data, together with all available malacosporean data in GenBank; and (iv) investigate the evolutionary trends of malacosporeans by mapping the morphology of bryozoan-related stages, host species, habitat and geographic data on the small subunit rDNA-based phylogenetic tree. We reveal a high prevalence and diversity of malacosporeans in several fish hosts in European freshwater habitats by adding five new species of Buddenbrockia and Tetracapsuloides from cyprinid and perciform fishes. Comprehensive phylogenetic analyses revealed that, apart from Buddenbrockia and Tetracapsuloides clades, a novel malacosporean lineage (likely a new genus) exists. The fish host species spectrum was extended for Buddenbrockia plumatellae and Buddenbrockia sp. 2. Co-infections of up to three malacosporean species were found in individual fish. The significant increase in malacosporean species richness revealed in the present study points to a hidden biodiversity in this parasite group. This is most probably due to the cryptic nature of malacosporean sporogonic and presporogonic stages and mostly asymptomatic infections in the fish hosts. The potential existence of malacosporean life cycles in the marine environment as well as the evolution of worm- and sac-like morphology is discussed. This study improves the understanding of the biodiversity, prevalence, distribution, habitat and host preference of malacosporeans and unveils their evolutionary trends.     

Does prescribed burning mean a threat to the rare satyrine butterfly Hipparchia fagi? Larval-habitat preferences give the answer

The ecological effects of fire management, especially regarding arthropods are poorly investigated. Burning in winter was assumed to pose a threat to butterfly species hibernating as larvae. To assess the impact of prescribed burning on population viability, we analysed larval-habitat preferences of the highly endangered, xero-thermophilous butterfly Hipparchia fagi in vineyards of the Kaiserstuhl region (southern Germany). Microhabitat preference analyses for mature larvae and egg-laying females revealed a preference of H. fagi for Bromus erectus-dominated communities with sparse vegetation coverage and a distinct tuft growth of the host plant B. erectus on microclimatically benefited slopes. We explain the preference of B. erectus by a preference of vegetation structure. The grass tufts offer a suitable climatically buffered living space for larvae. Egg deposition took place on dry substrate at positions of high solar radiation, thus adapted to hot and dry microclimate. As the larval habitat was sparsely vegetated as well as generally legally protected, fire management was not applicable and therefore not affecting the populations. We think it is conceivable that H. fagi, occurring here at its northern range limit, might expand its larval habitat into denser, combustible B. erectus stands in the course of global warming. A change in habitat preferences would necessitate a re-evaluation of management options.     

Coincidence in the distributions of butterflies and their foodplants

The relationship between the geographic distribution of consumers and of their hosts (foodplants) is examined using the resident butterfly fauna of Britain. On average, butterfly species that feed on more widely distributed hosts are themselves more widely distributed. However, the relationship is approximately triangular and the upper constraint imposed by the range sizes of hosts is not closely followed; some species have much more restricted ranges than their hosts have. There is no relationship between the proportion of the range of the foodplant that is occupied and the size of the range of the foodplant. Monopbagous butterfly species have smaller range sizes than polyphagous species, probably as a consequence of the greater potential range sizes of the latter. Those plant species that are used as hosts by butterflies have larger range sizes than expected by chance, and individual polyphagous butterfly species tend disproportionately to be found in areas containing larger numbers of their host plant species. In sum, this study reveals a complex relationship between the distribution of butterflies and that of their resources (foodplants).     

Host plant choice in the comma butterfly-larval choosiness may ameliorate effects of indiscriminate oviposition

In most phytophagous insects, the larval diet strongly affects future fitness and in species that do not feed on plant parts as adults, larval diet is the main source of nitrogen. In many of these insect-host plant systems, the immature larvae are considered to be fully dependent on the choice of the mothers, who, in turn, possess a highly developed host recognition system. This circumstance allows for a potential mother-offspring conflict, resulting in the female maximizing her fecundity at the expense of larval performance on suboptimal hosts. In two experiments, we aimed to investigate this relationship in the polyphagous comma butterfly, Polygonia c-album, by comparing the relative acceptance of low- and medium-ranked hosts between females and neonate larvae both within individuals between life stages, and between mothers and their offspring. The study shows a variation between females in oviposition acceptance of low-ranked hosts, and that the degree of acceptance in the mothers correlates with the probability of acceptance of the same host in the larvae. We also found a negative relationship between stages within individuals as there was a higher acceptance of lower ranked hosts in females who had abandoned said host as a larva. Notably, however, neonate larvae of the comma butterfly did not unconditionally accept to feed from the least favorable host species even when it was the only food source. Our results suggest the possibility that the disadvantages associated with a generalist oviposition strategy can be decreased by larval participation in host plant choice.     

Fine-Grained Distribution of a Non-Native Resource Can Alter the Population Dynamics of a Native Consumer

New interactions with non-native species can alter selection pressures on native species. Here, we examined the effect of the spatial distribution of a non-native species, a factor that determines ecological and evolutionary outcomes but that is poorly understood, particularly on a fine scale. Specifically, we explored a native butterfly population and a non-native plant on which the butterfly oviposits despite the plant’s toxicity to larvae. We developed an individual-based model to describe movement and oviposition behaviors of each butterfly, which were determined by plant distribution and the butterfly''s host preference genotype. We estimated the parameter values of the model from rich field data. We simulated various patterns of plant distributions and compared the rates of butterfly population growth and changes in the allele frequency of oviposition preference. Neither the number nor mean area of patches of non-native species affected the butterfly population, whereas plant abundance, patch shape, and distance to the nearest native and non-native patches altered both the population dynamics and genetics. Furthermore, we found a dramatic decrease in population growth rates when we reduced the distance to the nearest native patch from 147 m to 136 m. Thus changes in the non-native resource distribution that are critical to the fate of the native herbivore could only be detected at a fine-grained scale that matched the scale of a female butterfly’s movement. In addition, we found that the native butterfly population was unlikely to be rescued by the exclusion of the allele for acceptance of the non-native plant as a host. This study thus highlights the importance of including both ecological and evolutionary dynamics in analyses of the outcome of species interactions and provides insights into habitat management for non-native species.     

Oviposition,larval preference,and larval performance in two polyphagous species: does the larva know best?

It is expected that females preferentially oviposit on plant hosts that allow for optimal larval performance. However, this expectation contradicts empirical evidence where adults do not always choose the best host for their descendants. Recent evidence suggests that females’ host selection depends on the number of potential hosts. Females from oligophagous species seem to be able to choose an appropriate host in terms of larval performance, whereas in polyphagous species, adult oviposition preference is not related with larval performance. This suggests that larvae in polyphagous species could be taking a more active role in host selection than their mothers. Here, we evaluated the oviposition preference and the larval preference and performance of two polyphagous species of economic importance, Copitarsia decolora (Guenée) (Lepidoptera: Noctuidae: Cuculliinae) and Peridroma saucia (Hübner) (Lepidoptera: Noctuidae: Noctuinae), on eight species of cultivated plants. In laboratory and greenhouse choice assays, we tested adult preference for oviposition and larval preference at 1 and 24 h. Larval performance was measured in terms of survival to adulthood, length of larval period, and pupal weight. We found that both adult females and larvae actively choose their hosts and that the larval preference toward the hosts is related to the females’ preference in both herbivore species. However, the females and larvae did not preferentially select the host with the best larval performance, indicating that larval performance is not related to female or larval preference and that other selective pressures are influencing the choice of the host plant in these two species.     

Feeding preferences and their modification in early and late instar larvae of the bagworm,Thyridopteryx ephemeraeformis (Lepidoptera: Psychidae)

Feeding preferences in naive first instar and penultimate-final instar bagworm (Thyridopteryx ephemeraeformis)larvae were investigated using two choice tests. Naive first instar larvae showed a general (though not exclusive) preference for black willow (Salix nigra)over other hosts tested, regardless of the host of origin. Penultimate-final instar larvae showed strong preferences for the previous rearing host, suggesting induction of host preference; there was no evidence of host origin effects. The ability to discriminate among hosts is important if it is actually used to make host choices based on host quality. In bagworms, where host choices (if any) must be made by larvae in habitats where several hosts are often present, host discriminatory ability is likely important.     

Oviposition preference and larval performance within a diverging lineage of lycaenid butterflies

Abstract. 1. The butterfly genus Mitoura in Northern California includes three nominal species associated with four host plants having parapatric or interdigitated ranges. Genetic analyses have shown the taxa to be very closely related, and adults from all host backgrounds will mate and produce viable offspring in the laboratory. Oviposition preference and larval performance were investigated with the aim of testing the hypothesis that variation in these traits can exist in a system in which non‐ecological barriers to gene flow (i.e. geographic barriers and genetic incompatibilities) appear to be minimal. 2. Females were sampled from 12 locations throughout Northern California, including sympatric and parapatric populations associated with the four different host‐plant species. Oviposition preference was assayed by confining wild‐caught females with branches of all four host species and counting the number of eggs laid on each. Offspring were reared on the same host species and two measures of larval success were taken: per cent survival and pupal weight. 3. For populations associated with one of the hosts, incense cedar, the preference–performance relationship is simple: the host that females chose is the plant which results in the highest pupal weights for offspring. The preference–performance relationship for populations associated with the other hosts is more complex and may reflect different levels of local adaptation. The variation in preference and performance reported here suggests that these traits can evolve when non‐ecological barriers to gene flow are low, and that differences in these traits may be important for the evolution of reproductive isolation within Mitoura.     

Relative importance of quantity, quality and isolation of patches for butterfly diversity in fragmented urban forests

The majority of forests in urban areas are small and isolated. Improving habitat quality of small forests instead of increasing habitat size and connectivity could be an effective means of conserving the biodiversity of such highly fragmented landscapes. In this study, we investigated the relative importance of habitat quantity, quality and isolation on butterfly assemblages in urban fragmented forests in Tokyo, Japan. We used four habitat geographic parameters: (1) fragment size, (2) shape index, (3) isolation (distance to the mainland), and (4) connectivity; and three habitat quality parameters: (1) herbaceous nectar plant abundance, (2) herbaceous nectar plant diversity, and (3) larval host plant diversity. We surveyed butterfly assemblages along transects in 20 forest fragments that ranged in size from 1 to 122 ha. We used generalized linear models to relate the number of species in a fragment to four habitat geographic parameters and three habitat quality parameters. The averaged models based on AIC_c showed that fragment size had a strong positive effect on butterfly species richness. There was also a positive effect of herbaceous nectar plant abundance on species diversity. These findings suggest that improving the habitat quality of small and isolated forests in highly fragmented landscapes may be capable of maintaining levels of butterfly diversity comparable to those of large fragments.