The Relationship between Diet Breadth and Geographic Range Size in the Butterfly Subfamily Nymphalinae – A Study of Global Scale

The “oscillation hypothesis” has been proposed as a general explanation for the exceptional diversification of herbivorous insect species. The hypothesis states that speciation rates are elevated through repeated correlated changes – oscillations – in degree of host plant specificity and geographic range. The aim of this study is to test one of the predictions from the oscillation hypothesis: a positive correlation between diet breadth (number of host plants used) and geographic range size, using the globally distributed butterfly subfamily Nymphalinae. Data on diet breadth and global geographic range were collected for 182 Nymphalinae butterflies species and the size of the geographic range was measured using a GIS. We tested both diet breadth and geographic range size for phylogenetic signal to see if species are independent of each other with respect to these characters. As this test gave inconclusive results, data was analysed both using cross-species comparisons and taking phylogeny into account using generalised estimating equations as applied in the APE package in R. Irrespective of which method was used, we found a significant positive correlation between diet breadth and geographic range size. These results are consistent for two different measures of diet breadth and removal of outliers. We conclude that the global range sizes of Nymphalinae butterflies are correlated to diet breadth. That is, butterflies that feed on a large number of host plants tend to have larger geographic ranges than do butterflies that feed on fewer plants. These results lend support for an important step in the oscillation hypothesis of plant-driven diversification, in that it can provide the necessary fuel for future population fragmentation and speciation

Why stay in a bad relationship? The effect of local host phenology on a generalist butterfly feeding on a low-ranked host

Metabolic profiling by GC-MS and LC-MS. (PDF 93 kb

The Benefit of Additional Oviposition Targets for a Polyphagous Butterfly

While the reasons for the prevalence of specialists over generalists among herbivorous insects have been at the focus of much interest, less effort has been put into understanding the polyphagous exceptions. Recent studies have suggested that these exceptions may be important for insect diversification, which calls for a better understanding of the potential factors that can lead to an increased host plant repertoire. Females of the Nymphalid butterfly, Polygonia c-album, were used to test if egg output and/or likelihood of finding a host increased with the addition of a secondary host. There was no effect of prior eggs on the host for willingness to oviposit on a plant. The main experiments were conducted both in small laboratory cages and in large outdoor experimental arenas. No positive effect was found when another oviposition target was added in small cages in the laboratory. On the other hand, in the outdoor arenas the females more often found a host to oviposit on and had a higher egg output when they had access to an additional host, even though the second host was lower in their preference hierarchy. The difference between these experiments was attributed to searching for acceptable host plants within a patch, a factor that was included in the large cages but not in the small. When host availability is limited, adding oviposition targets can potentially act to counterbalance specialization and thus favor the evolution of generalization

Phylogenetic relatedness and host plant growth form influence gene expression of the polyphagous comma butterfly (Polygonia c-album)

<p>Abstract</p> <p>Background</p> <p>The mechanisms that shape the host plant range of herbivorous insect are to date not well understood but knowledge of these mechanisms and the selective forces that influence them can expand our understanding of the larger ecological interaction. Nevertheless, it is well established that chemical defenses of plants influence the host range of herbivorous insects. While host plant chemistry is influenced by phylogeny, also the growth forms of plants appear to influence the plant defense strategies as first postulated by Feeny (the "plant apparency" hypothesis). In the present study we aim to investigate the molecular basis of the diverse host plant range of the comma butterfly (<it>Polygonia c-album</it>) by testing differential gene expression in the caterpillars on three host plants that are either closely related or share the same growth form.</p> <p>Results</p> <p>In total 120 genes were identified to be differentially expressed in <it>P. c-album </it>after feeding on different host plants, 55 of them in the midgut and 65 in the restbody of the caterpillars. Expression patterns could be confirmed with an independent method for 14 of 27 tested genes. Pairwise similarities in upregulation in the midgut of the caterpillars were higher between plants that shared either growth form or were phylogenetically related. No known detoxifying enzymes were found to be differently regulated in the midgut after feeding on different host plants.</p> <p>Conclusion</p> <p>Our data suggest a complex picture of gene expression in response to host plant feeding. While each plant requires a unique gene regulation in the caterpillar, both phylogenetic relatedness and host plant growth form appear to influence the expression profile of the polyphagous comma butterfly, in agreement with phylogenetic studies of host plant utilization in butterflies.</p

Polyphagy and diversification in tussock moths: Support for the oscillation hypothesis from extreme generalists

Theory on plasticity driving speciation, as applied to insect–plant interactions (the oscillation hypothesis), predicts more species in clades with higher diversity of host use, all else being equal. Previous support comes mainly from specialized herbivores such as butterflies, and plasticity theory suggests that there may be an upper host range limit where host diversity no longer promotes diversification. The tussock moths (Erebidae: Lymantriinae) are known for extreme levels of polyphagy. We demonstrate that this system is also very different from butterflies in terms of phylogenetic signal for polyphagy and for use of specific host orders. Yet we found support for the generality of the oscillation hypothesis, in that clades with higher diversity of host use were found to contain more species. These clades also consistently contained the most polyphagous single species. Comparing host use in Lymantriinae with related taxa shows that the taxon indeed stands out in terms of the frequency of polyphagous species. Comparative evidence suggests that this is most probably due to its nonfeeding adults, with polyphagy being part of a resulting life history syndrome. Our results indicate that even high levels of plasticity can drive diversification, at least when the levels oscillate over time.</p

Rewiring of interactions in a changing environment: nettle‐feeding butterflies and their parasitoids

Climate and land use change can alter the incidence and strength of biotic interactions, with important effects on the distribution, abundance and function of species. To assess the importance of these effects and their dynamics, studies quantifying how biotic interactions change in space and time are needed. We studied interactions between nettle‐feeding butterflies and their shared natural enemies (parasitoids) locally and across 500 km latitudinal gradient in Sweden. We also examined the potential impact of the range‐expansion of the butterfly Araschnia levana on resident butterflies via shared parasitoids, by studying how parasitism in resident butterflies covaries with the presence or absence of the newly‐established species. We collected 6777 larvae of four nettle‐feeding butterfly species (Aglais urticae, Aglais io, Ar. levana and Vanessa atalanta), over two years, at 19 sites distributed along the gradient. We documented the parasitoid complex for each butterfly species and measured their overlap, and analysed how parasitism rates were affected by butterfly species assemblage, variations in abundance, time, and the arrival of Ar. levana. Parasitoids caused high mortality, with substantial overlap in the complex of parasitoids associated with the four host butterflies. Levels of parasitism differed significantly among butterflies and were influenced by the local butterfly species assemblage. Our results also suggest that parasitism in resident butterflies is elevated at sites where Ar. levana has been established for a longer period. In our study system, variations in butterfly species assemblages were associated in a predictable way with substantial variations in rates of parasitism. This relationship is likely to affect the dynamics of the butterfly host species, and potentially cascade to the larger number of species with which they interact. These results highlight the importance of indirect interactions and their potential to reorganise ecological communities, especially in the context of shifts in species distributions in a warmer world

Odour Maps in the Brain of Butterflies with Divergent Host-Plant Preferences

Butterflies are believed to use mainly visual cues when searching for food and oviposition sites despite that their olfactory system is morphologically similar to their nocturnal relatives, the moths. The olfactory ability in butterflies has, however, not been thoroughly investigated. Therefore, we performed the first study of odour representation in the primary olfactory centre, the antennal lobes, of butterflies. Host plant range is highly variable within the butterfly family Nymphalidae, with extreme specialists and wide generalists found even among closely related species. Here we measured odour evoked Ca2+ activity in the antennal lobes of two nymphalid species with diverging host plant preferences, the specialist Aglais urticae and the generalist Polygonia c-album. The butterflies responded with stimulus-specific combinations of activated glomeruli to single plant-related compounds and to extracts of host and non-host plants. In general, responses were similar between the species. However, the specialist A. urticae responded more specifically to its preferred host plant, stinging nettle, than P. c-album. In addition, we found a species-specific difference both in correlation between responses to two common green leaf volatiles and the sensitivity to these compounds. Our results indicate that these butterflies have the ability to detect and to discriminate between different plant-related odorants

Diversity begets diversity: host expansions and the diversification of plant-feeding insects

<p>Abstract</p> <p>Background</p> <p>Plant-feeding insects make up a large part of earth's total biodiversity. While it has been shown that herbivory has repeatedly led to increased diversification rates in insects, there has been no compelling explanation for how plant-feeding has promoted speciation rates. There is a growing awareness that ecological factors can lead to rapid diversification and, as one of the most prominent features of most insect-plant interactions, specialization onto a diverse resource has often been assumed to be the main process behind this diversification. However, specialization is mainly a pruning process, and is not able to actually generate diversity by itself. Here we investigate the role of host colonizations in generating insect diversity, by testing if insect speciation rate is correlated with resource diversity.</p> <p>Results</p> <p>By applying a variant of independent contrast analysis, specially tailored for use on questions of species richness (MacroCAIC), we show that species richness is strongly correlated with diversity of host use in the butterfly family Nymphalidae. Furthermore, by comparing the results from reciprocal sister group selection, where sister groups were selected either on the basis of diversity of host use or species richness, we find that it is likely that diversity of host use is driving species richness, rather than vice versa.</p> <p>Conclusion</p> <p>We conclude that resource diversity is correlated with species richness in the Nymphalidae and suggest a scenario based on recurring oscillations between host expansions – the incorporation of new plants into the repertoire – and specialization, as an important driving force behind the diversification of plant-feeding insects.</p