Does herbivore diversity depend on plant diversity? The case of California butterflies

It is widely believed that the diversity of plants influences the diversity of animals, and this should be particularly true of herbivores. We examine this supposition at a moderate spatial extent by comparing the richness patterns of the 217 butterfly species resident in California to those of plants, including all 5,902 vascular plant species and the 552 species known to be fed on by caterpillars. We also examine the relationships between plant/butterfly richness and 20 environmental variables. We found that although plant and butterfly diversities are positively correlated, multiple regression, path models, and spatial analysis indicate that once primary productivity (estimated by a water-energy variable, actual evapotranspiration) and topographical variability are incorporated into models, neither measure of plant richness has any relationship with butterfly richness. To examine whether butterflies with the most specialized diets follow the pattern found across all butterflies, we repeated the analyses for 37 species of strict monophages and their food plants and found that plant and butterfly richness were similarly weakly associated after incorporating the environmental variables. We condude that plant diversity does not directly influence butterfly diversity but that both are probably responding to similar environmental factors.     

Butterfly diversity in Mediterranean islands and in Pentadaktylos <Emphasis Type="Italic">Pinus</Emphasis><Emphasis Type="BoldItalic"> </Emphasis><Emphasis Type="Italic">brutia</Emphasis> forests of Cyprus

We analysed the influence of contemporary geography on butterfly diversity for islands in the Mediterranean Basin. We found that island size and distance from the mainland has a significant effect on the number of species. We also used butterflies as an indicator group to identify the importance of forest habitats for biodiversity conservation in the island of Cyprus. To understand the relative importance of local vegetation characteristics of butterflies in the Pentadaktylos mountains transect counts were used to assess the abundance and butterfly diversity in two different forest types. A total of 1,602 butterflies and 23 species were recorded during this research. We observed highly significant effects of forest type on abundance and species richness of butterflies. For example, number of butterflies was significantly higher in old forest than young pine forest. Also, the abundance of endemic butterflies was highest in old forest habitats. Therefore, the survival of the majority of endemic butterflies in Cyprus may depend on conservation of old forests and their understorey plants.     

蝴蝶在生境和生物小区之间的移位伴随着对外来蜜源的利用

在发育成熟的温带环境中,植食性昆虫群落能迅速适应引入植物并将其作为幼虫的食物和花蜜.我们研究了经过森林砍伐的热带环境中蝴蝶对利用引入植物作为蜜源植物的适应快慢程度,并研究了蝴蝶-显花植物在新的生物小区中出现的范围,发现蝴蝶对引入显花植物的利用和探访多于本地植物,这与引入植物在调查地点、生物小区和植物丰度中的普遍性有关.此外,取食花蜜的蝴蝶和显花植物与正在形成的的生物小区有关,例如路边、农田、集约耕地以及花园.在这些新生物小区中,引入植物很重要,因为它们为蝴蝶提供了蜜源.     

Recovery of indigenous butterfly community following control of invasive alien plants in a tropical island’s wet forests

Invasive alien species pose one of the highest threats to biodiversity, especially in isolated oceanic islands where high rates of both endemism and extinction risk also usually prevail. Few studies have investigated the impact of invasive alien plants on butterflies in insular ecosystems, despite butterflies representing a key indicator group for terrestrial arthropod diversity. Using the Pollard Technique, we quantified butterfly species richness and abundance in eight wet lowland forest areas invaded by alien plants, principally the strawberry guava (Psidium cattleianum Sabine) on the tropical volcanic island of Mauritius, and compared the results with paired adjacent forest plots that had been weeded of alien plants between 2 and 12 years previously. Butterfly assemblages in weed-infested and weeded forests were distinctly different with higher species richness and much higher butterfly abundance in the latter. At least some of these differences seemed attributable to weed removal effects on forest structure, but understanding the precise mechanisms involved will require further study. The results suggest that alien plant invasion may have contributed to the extinction of certain endemic taxa and can increase the likelihood of butterfly species extinction by reducing population sizes through reduced habitat quality. Such a shift in a forest’s butterfly assemblage is likely to have negative effects on both their indigenous predators and the plants they pollinate. It is argued that in order to maintain butterfly and other arthropod diversity and function in these forests, alien plant control must be maintained and extended beyond the current 1% of surviving forest remnants.     

Butterfly species diversity and their floral preferences in the Rupa Wetland of Nepal

Floral attributes often influence the foraging choices of nectar‐feeding butterflies, given the close association between plants and these butterfly pollinators. The diversity of butterflies is known to a large extent in Nepal, but little information is available on the feeding habits of butterflies. This study was conducted along the periphery of Rupa Wetland from January to December 2019 to assess butterfly species diversity and to identify the factors influencing their foraging choices. In total, we recorded 1535 individuals of 138 species representing all six families. For our examination of butterfly–nectar plant interactions, we recorded a total of 298 individuals belonging to 31 species of butterfly visiting a total of 28 nectar plant species. Overall, total butterfly visitation was found to be significantly influenced by plant category (herbaceous preferred over woody), floral color (yellow white and purple preferred over pink), and corolla type (tubular preferred over nontubular). Moreover, there was a significant positive correlation between the proboscis length of butterflies and the corolla tube length of flowers. Examining each butterfly family separately revealed that, for four of the families (Lycaenidae, Nymphalidae, Papilionidae, and Pieridae), none of the tested factors (flower color, plant category, and corolla type) were shown to significantly influence butterfly abundance at flowers. However, Hesperidae abundance was found to be significantly influenced by both flower color (with more butterflies observed at yellow flowers than purple) and flower type (with more butterflies observed at tubular flowers than nontubular flowers). Our results reveal that Rupa Lake is a suitable habitat for butterflies, providing valuable floral resources. Hence, further detailed studies encompassing all seasons, a greater variety of plants, and other influential factors in different ecological regions are fundamental for creating favorable environments to sustain important butterfly pollinators and help create balanced wetland ecosystems.     

Patterns of butterfly distribution in the Andaman islands: implications for conservation

Twenty-five islands of different sizes were rapidly surveyed in the Andaman islands for patterns of butterfly distribution and abundance. The surveys were conducted in the dry seasons of 1992 in the South Andaman islands, 1994 in the North Andaman islands and on both these years on the Little Andaman Island. Different habitat types were identified on each island and butterflies were sampled by the line transect method in each habitat type. Sixty-five species of butterflies were recorded from six families. Fifty-one species were less common and contributed to 25 % of the total count. Six species were very common. The overall distribution patterns of the species were nested. This suggests that small islands share their species with the larger islands but not vice versa. Many uncommon species were found exclusively on large islands. The presence of evergreen forest on islands significantly influenced the species encountered. Small and medium sized islands with evergreen forests had significantly more species than those without evergreen forests. Loss of primary forests due to logging and encroachment will result in the loss of many butterfly species. It is recommended that the large patches of primary evergreen forests be protected on a priority basis on large islands.     

Using a phenological network to assess weather influences on first appearance of butterflies in the Netherlands

Phenological responses of butterflies to temperature have been demonstrated in several European countries by using data from standardized butterfly monitoring schemes. Recently, phenological networks have enabled volunteers to record phenological observations at project websites. In this study, the quality of the first appearance data of butterflies from the Dutch phenological network ‘De Natuurkalender’ was examined and these data were then used to analyze trends in butterfly appearance between 2001 and 2013, the effects of climatic factors on appearance of butterflies as well as the phenological interaction of one butterfly species, Anthocharis cardamines, and its two major host plants. Although phenological networks are relatively unstructured, this study shows that data from De Natuurkalender were highly comparable to more standardized data collected by the Dutch Butterfly Monitoring Scheme. No trend in first appearance of any butterfly species was found during the time period 2001–2013. The first appearance dates of most butterflies showed, however, a clear relationship with spring temperature. Higher temperatures, especially in March and April, advanced the first appearance of butterflies. Therefore, with climatic warming in the future, earlier appearance of butterflies is expected. Although climate warming is a potential threat for phenological mismatches between different trophic levels, this study shows a similar temperature response of A. cardamines and its main host plants in the Netherlands. However, as only few phenological interactions between species are examined, further research including rarer monophagous butterfly species and their host plants is needed.     

Agonistic display or courtship behavior? A review of contests over mating opportunity in butterflies

Male butterflies compete over mating opportunities. Two types of contest behavior are reported. Males of various butterfly species compete over a mating territory via aerial interactions until one of the two contestants retreats. Males of other butterfly species fly around larval food plants to find receptive females. Males of some species among the latter type can find a conspecific pupa, and they gather around it without expelling their rivals. Scramble competition over mating occurs when a female emerges from the pupa. Many studies have been performed on territorial species, and their contest resolution has often been understood from the point of view of contest models based on game theory. However, these models cannot explain why these butterflies perform contest displays despite the fact that they do not have the ability to attack their opponent. A recent study based on Lloyd Morgan’s Canon showed that territorial contests of male butterflies are better understood as erroneous courtship between sexually active males. In this paper, I review research on contests over mating opportunity in butterflies, and show that the erroneous courtship framework can explain not only territorial contests of butterflies but also why males do not determine the owner of a conspecific pupa.     

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.     

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.     

What can sown wildflower strips contribute to butterfly conservation?: an example from a Swiss lowland agricultural landscape

The objective of this study was to compare butterfly abundances and diversity between wildflower strips and extensively used meadows to identify which butterfly species can be supported by establishing wildflower strips. Butterflies were recorded along transects during one season in twenty-five sown wildflower strips and eleven extensively used meadows in a Swiss lowland agricultural landscape (600 ha). In total 1,669 butterflies of 25 species were observed (25 in the strips, 18 in meadows). This can be related to 38 species recorded in the region (lowland part of Kanton Fribourg) within the Swiss Biodiversity Monitoring Programme. In wildflower strips the number of butterflies per transect meter was significantly higher than in the meadows, but there was no significant difference in species richness. Butterfly communities, though, were quite different between the two habitat types. Habitat type, abundances of flowering plants and presence of forest within 50 m were identified as factors influencing butterfly species richness. Butterfly abundances were affected by habitat type and abundance of flowering plants. In wildflower strips, 65% of all flower visits by butterflies were observed on Origanum. It can be concluded that sown wildflower strips can support a substantial part of a regions species pool. This is mostly true for common species, but can apply to rare species when, for example, larval food plant requirements are met.     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.     

Butterfly biogeography and endemism on tropical Pacific islands

Butterfly distributions on 26 tropical Pacific archipelagos were analysed to examine the effects of geography on diversity and endemism. The total butterfly fauna for each archipelago was divided into continental (found also on continental areas), Pacific (found within more than one archipelago but not outside of the study area), and endemic species (restricted to a single archipelago). Numbers and proportions of each species were related to eight geographic variables by stepwise multiple linear regression analysis. Total area of an archipelago and distance from other land masses were important predictors of the number of species within an archipelago. Proportions of butterfly species in each category were related differently to the geographic variables, with endemism being promoted by the number of large islands within an archipelago. Relative to birds, butterflies have been less successful in colonizing remote archipelagos and have much lower levels of endemism. Even if colonization is successful, butterfly speciation may be constrained by the mechanics of coevolution with available host plants.     

Amino acids in nectar enhance butterfly fecundity: a long-awaited link

Thirty years ago, researchers discovered that flowers pollinated by butterflies are consistently rich in nectar amino acids, and more recent findings have shown that butterflies prefer nectar with high amino acid content. These observations led to speculation that amino acids in nectar enhance butterfly fitness and that butterflies have acted as agents of natural selection on nectar composition. Despite a number of experimental efforts over the years, convincing proof that nectar amino acids affect butterfly fitness has been lacking. Here, we provide the first evidence that amino acids in nectar have a positive effect on fecundity of one butterfly species, supporting the existence of a relationship between nectar preferences and fitness benefits. Map butterflies (Araschnia levana L.) raised under natural larval food conditions laid more eggs when they were fed nectar containing amino acids, whereas nectar amino acids had no effect on the number of eggs laid by butterflies raised on larval food rich in nitrogen. Uptake and utilization of nectar amino acids by map butterflies appear to be compensatory mechanisms enabling them to override impacts of poor larval food. These results provide strong support for the long-standing postulate that nectar amino acids benefit butterflies.     

Flower visitation and pollination efficiency of some North European butterflies

Summary Nectar plant utilization by butterflies was compared with the abundance of flowering plants on some SW Swedish meadows. The transportation of pollen grains by individual butterflies was analysed using a scanning electron microscope.For the majority of plant species, butteflies were of minor importance as pollinators. Some flowers with protruding sexual organs, e.g. Knautia arvensis, when heavily utilized by butterflies, might also be pollinated by these.For Dianthus deltoides and Viscaria vulgaris, butterfly pollination was found to be important.Several butterfly species, e.g. Plebicula amanda, visited legumes regularly. In spite of this, legume pollen was rarely transported by the butterflies. This suggests these butterflies act as nectar robbers among the legumes.     

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.     

Can the growing of transgenic maize threaten protected Lepidoptera in Europe?

We evaluated whether protected European butterflies can potentially be at risk if transgenic maize is extensively grown in Central Europe. We explored potential consequences of both insect resistant (IR) and herbicide resistant (HR) transgenic maize. IR maize can produce pollen that is toxic to lepidopteran larvae, and this puts butterfly species at possible risk if the presence of young larvae coincides with maize flowering, during which large quantities of maize pollen can be deposited on vegetation. By considering the timing of maize flowering in Europe and the phenology of the protected Lepidoptera species, we found that 31 species had at least one generation where 50% of the larval stage overlapped with maize flowering, and 69 species for which first instar larvae were present during maize pollen shedding. HR maize allows high concentration herbicide treatments on fields without seasonal limitation, which can drastically reduce weed densities. In cases where such weed species are host plants for protected butterflies, reduced host plant/food availability can result, causing population decreases. By using published information, we first identified the important weed species in major maize-growing European countries. Subsequently, we checked whether the host plants of protected Lepidoptera included species that are common maize weeds. We identified 140 protected species having food plants that are common weeds in one or more of the major European maize-growing countries. If HR maize is grown in Europe, there is a potential hazard that their food plants will seriously decline, causing a subsequent decline of these protected species.     

Preferences and Food Quality of Fruit‐Feeding Butterflies in Kibale Forest,Uganda1

Many butterflies in tropical forests feed on fruits that have fallen to the forest floor. This substrate differs in many ways from floral nectar, and therefore fruit‐feeding butterflies are expected to possess adaptations for efficient foraging, choice, and ingestion of their food. Differences in food quality and in spatial and temporal availability are also likely to have led to life history evolution. Here we describe the sugar and nitrogen content of fruits that butterflies feed on in a tropical forest in Uganda, and measure the attractiveness of these fruits to the local butterfly fauna together with the role that decay plays in the attraction. These data are supplemented with feeding observations at fruit falls in the forest. Our results show that (1) fruits contain significant and variable concentrations of sugar and nitrogen, and constitute a nutritious food source for butterflies in tropical forests; (2) fruit‐feeding butterflies use cues from the fruits and fermentation products to locate their food; (3) different classes of fruit‐feeding butterflies may vary in their preferences for certain fruits, and differ in their ability to find preferred food; and (4) fruit choice is not strongly correlated with attractiveness or nutrient content. The results are discussed in the light of the evolution of food searching and life history strategies.     

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.     

舟山群岛蝶类群落嵌套分布格局及其影响因素

片断化生境中群落的物种组成常呈现嵌套分布格局。2013年7-8月, 我们在浙江舟山群岛采用截线法对28个岛屿上的蝴蝶群落进行了野外调查, 探讨了岛屿物种嵌套分布格局及其影响因素。通过测量采集标本获得蝶类的生活史特征(最小需求面积、翅展和体重), 查阅文献资料获得蝶类的栖息地特征(岛屿面积、距最近大陆距离和距最近大岛距离), 分析了影响蝶类群落嵌套结构的因素。研究结果显示: (1)舟山群岛蝶类群落符合嵌套分布格局; (2)岛屿面积和物种最小需求面积对嵌套格局的形成有显著影响; (3)舟山群岛蝶类群落嵌套格局的形成支持选择性灭绝假说; (4)随机检验零模型结果显示该嵌套分布格局并非采样偏差造成的。因此, 在制定舟山群岛区域蝶类保护措施时, 应优先考虑那些分布在面积较大岛屿的和最小需求面积较大的物种。