The evolution of novel host use is unlikely to be constrained by trade‐offs or a lack of genetic variation

The genetic and ecological factors that shape the evolution of animal diets remain poorly understood. For herbivorous insects, the expectation has been that trade‐offs exist, such that adaptation to one host plant reduces performance on other potential hosts. We investigated the genetic architecture of alternative host use by rearing individual Lycaeides melissa butterflies from two wild populations in a crossed design on two hosts (one native and one introduced) and analysing the genetic basis of differences in performance using genomic approaches. Survival during the experiment was highest when butterfly larvae were reared on their natal host plant, consistent with local adaptation. However, cross‐host correlations in performance among families (within populations) were not different from zero. We found that L. melissa populations possess genetic variation for larval performance and variation in performance had a polygenic basis. We documented very few genetic variants with trade‐offs that would inherently constrain diet breadth by preventing the optimization of performance across hosts. Instead, most genetic variants that affected performance on one host had little to no effect on the other host. In total, these results suggest that genetic trade‐offs are not the primary cause of dietary specialization in L. melissa butterflies.     

Molecular and morphological divergence in the butterfly genus Lycaeides (Lepidoptera: Lycaenidae) in North America: evidence of recent speciation

Male genital morphology, allozyme allele frequencies and mtDNA sequence variation were surveyed in the butterfly species Lycaeides idas and L. melissa from across much of their range in North America. Despite clear differences in male genital morphology, wing colour patterns and habitat characteristics, genetic variation was not taxonomically or geographically structured and the species were not identifiable by either genetic data set. Genetic distances (Nei's D=0.002–0.078, calculated from allozyme data) between all populations of both species were within the range commonly observed for conspecific populations of other butterflies. The most frequent mtDNA haplotype was present in individuals of both species in populations from southern California to Wisconsin. We conclude that speciation has probably happened recently and the lack of genetic differentiation between the species is the product of either (1) recent or ongoing gene flow at neutral loci, and/or (2) an insufficiency of time for lineage sorting. The evolution of male genital morphology, wing colour patterns and ecological characteristics has proceeded more rapidly than allozyme or mtDNA evolution.     

A Molecular and Fitness Evaluation of Commercially Available versus Locally Collected Blue Lupine Lupinus perennis L. Seeds for Use in Ecosystem Restoration Efforts

Dependence on wild seed sources is often impractical for large‐scale habitat restoration programs. Reliance on commercial seed supplies of unknown provenance and fitness is thereby warranted. Little consideration has been given, however, to how the large volumes of seed required should be sourced. We evaluated commercial and locally collected seed sources for potential use in a New York State‐based, landscape‐scale program for restoring blue lupine Lupinus perennis. Through analysis of microsatellite markers we determined that “native” lupine designations by some commercial suppliers were in fact interspecific hybrids and therefore unreliable; at least two commercial sources, however, were genetically as close to native New York populations as native New York populations were to one other. Common garden experiments revealed that seed source influenced first‐year overwintering survival and subsequent height growth of surviving plants; seed sources more closely related genetically to native New York populations survived better and produced more stems per individual in the field in the area targeted for restoration. We conclude that (1) commercial suppliers often but not always offer reliably characterized seed sources of sufficient genetic similarity to native populations to warrant their use in restoration projects and (2) genetic affinity of potential seed stock to native populations is positively related to its fitness in the environment targeted for restoration.     

The co-occurrence of an introduced biological control agent (Coleoptera: <Emphasis Type="Italic">Coccinella septempunctata</Emphasis>) and an endangered butterfly (Lepidoptera: <Emphasis Type="Italic">Lycaeides melissa samuelis</Emphasis>)

Whether a biological control agent presents a non-target risk to a native species depends if they co-occur spatially and temporally, and if the agent will harm the native species. We sampled two study sites during 1993 in Minnesota and Wisconsin to survey predators and parasitoids of the extant populations of the United States federally endangered Karner blue butterfly, Lycaeides melissa samuelis. We found the introduced coccinellid Coccinella septempuntata co-occurring spatially and temporally with eggs, larvae and adults of L. m. samuelis. The two species were also observed together on the latters sole host plant, Lupinus perennis, and in Wisconsin, an adult C. septempunctata was observed consuming second instar larvae of L. m. samuelis. Using a simple model to hypothesize the risk that C. septempunctata presents to L. m. samuelis, we showed that increases in predator density could greatly increase mortality to L. m. samuelis. At these sites, C. septempunctata were reproducing and had access to summer aphids and suitable overwintering habitat. Nearby agricultural crops could provide spring aphids for oogenesis, and assist with C. septempunctata population build-up. Maintaining a minimum isolation distance between agricultural crops known to harbor aphids and extant L. m. samuelis populations may need to be considered as part of the butterfly management program.     

Lack of evidence for reproductive isolation among ecologically specialised lycaenid butterflies

Abstract 1. The evolution of reproductive isolation between recently diverged or incipient species is a critical component of speciation and a major focus of speciation models. In phytophagous insects, host plant fidelity (the habit of mating and ovipositing on a single host species) can contribute to assortative mating and reproductive isolation between populations adapting to alternative hosts. The potential role of host plant fidelity in the evolution of reproductive isolation was examined in a pair of North American blue butterfly species, Lycaeides idas and L. melissa .
2. These species are morphologically distinct and populations of each species utilise different host plants; however they share 410 bp haplotypes of the mitochondrial cytochrome oxidase subunit I (COI) gene, indicating recent divergence.
3. Some populations using native hosts exhibited strong fidelity for their natal host plant over the hosts used by nearby populations. Because these butterflies mate on or near the host plant, the development of strong host fidelity may create reproductive isolation among populations on different hosts and restrict gene flow.
4. Tests of population differentiation using allozyme allele frequency data did not provide convincing evidence of restricted gene flow among populations. Based on morphological differences, observed ecological specialisation, and the sharing of genetic markers, these butterflies appear to be undergoing adaptive radiation driven at least partially by host shifts. Neutral genetic markers may fail to detect the effects of very recent host shifts in these populations due to gene flow and/or the recency of divergence and shared ancestral polymorphism.     

Caterpillars on a phytochemical landscape: The case of alfalfa and the Melissa blue butterfly

Modern metabolomic approaches that generate more comprehensive phytochemical profiles than were previously available are providing new opportunities for understanding plant‐animal interactions. Specifically, we can characterize the phytochemical landscape by asking how a larger number of individual compounds affect herbivores and how compounds covary among plants. Here we use the recent colonization of alfalfa (Medicago sativa) by the Melissa blue butterfly (Lycaeides melissa) to investigate the effects of indivdiual compounds and suites of covarying phytochemicals on caterpillar performance. We find that survival, development time, and adult weight are all associated with variation in nutrition and toxicity, including biomolecules associated with plant cell function as well as putative anti‐herbivore action. The plant‐insect interface is complex, with clusters of covarying compounds in many cases encompassing divergent effects on different aspects of caterpillar performance. Individual compounds with the strongest associations are largely specialized metabolites, including alkaloids, phenolic glycosides, and saponins. The saponins are represented in our data by more than 25 individual compounds with beneficial and detrimental effects on L. melissa caterpillars, which highlights the value of metabolomic data as opposed to approaches that rely on total concentrations within broad defensive classes.     

After 60 years,an answer to the question: what is the Karner blue butterfly?

The Karner blue butterfly (KBB), Lycaeides melissa samuelis, is a federally protected taxon whose relationship to the Melissa blue, Lycaeides melissa, has been a point of contention during the 66 years since the KBB was first described. Using a large population-genomic dataset and a model of population divergence with migration, we investigated the relationship between the KBB and L. melissa, as well as the relationship between L. melissa and a third taxon, Lycaeides idas. We report that gene flow between the KBB and L. melissa is low, and comparable to gene flow between L. melissa and L. idas. Considering this population-genetic evidence, we conclude that the KBB is a unique evolutionary lineage that should be recognized as Lycaeides samuelis.     

Assessment of Potential Karner Blue Butterfly (Lycaeides melissa samuelis) (Family: Lycanidae) Reintroduction Sites in Ontario, Canada

Although species reintroduction is useful as both an integral part and a performance indicator of habitat restoration, it is not a risk‐free process. Evaluation of potential reintroduction sites is often crucially important in reducing chances of failure. Despite its importance, there is no standard way to do so. This study applied a systematic scheme to evaluate five potential reintroduction sites for the endangered Karner Blue butterfly (Lycaeides melissa samuelis Nabokov) in Ontario, Canada, by looking at both biotic and abiotic aspects. Field data were collected in 2003 from these sites, and three potential founder butterfly sites in the United States. We used data collected from the U.S. sites to determine the minimum standards for ecological requirements of the Karner Blue. Data from the Ontario sites were then compared against these standards. The results show that all five potential reintroduction sites are of lower quality, at least in certain aspects, compared with the three potential founder butterfly sites. This implies that success is not guaranteed if the Karner Blue is reintroduced into these sites under current habitat conditions. Further site restoration is required and should focus on the shortcomings identified for individual sites in this study. The lessons from this study are useful for potential reintroduction site assessment in other restoration projects because they reveal whether the sites are ready for species reintroduction and, if not, how they need to be improved.     

Variation in butterfly egg adhesion: adaptation to local host plant senescence characteristics?

Most butterflies that overwinter as diapausing eggs have evolved oviposition behaviours where egg placement is near or on perennating portions of their host plant. We describe an alternative strategy used by alpine populations of a Lycaenid butterfly species complex where eggs are not attached strongly to the leaf substrate and subsequently fall off the plant. Other populations occurring at lower elevations attach their eggs strongly to the plant. In the laboratory, we measured the amount of force required to detach eggs from the leaf substrate. Eggs of the non‐alpine populations required more than five times the force required to detach eggs of the alpine population. Field surveys of the host plant characteristics used by various populations suggest that easy detachment may be adaptive for alpine populations because, unlike the host plants used by other populations, nearly all of the preceding year's vegetative growth is blown away by strong winds during the winter months.     

The history and geography of diversification within the butterfly genus Lycaeides in North America

The Lycaeides butterfly species complex in North America consists of two nominal, morphologically defined species. These butterflies are ecologically diverse and appear to be distributed as a geographically complex mosaic of locally differentiated populations that may be undergoing adaptive radiation. We asked whether patterns of molecular genetic variation within the species complex are congruent with currently recognized morphological species and whether the distribution of molecular variation is consistent with the hypothesis that Pleistocene climate changes contributed to the process of differentiation within the genus. Variation in the form of the genitalia from 726 males from 59 populations clearly distinguishes both species with only six populations containing morphologically intermediate or ambiguous individuals. However, partitioning of molecular variance in a 236 bp section of the mitochondrial AT-rich region from 628 individuals (57 populations) surveyed using single strand conformation polymorphism analysis (SSCP) indicates that only 26% of the total genetic variation is distributed along nominal species boundaries as defined by morphology. Instead, three phylogeographical groups were detected, represented by three major haplotype clades, which account for 90% of the total genetic variance. Pleistocene glaciations appear to have fostered divergence during glacial maxima, while post-glacial range expansions created opportunities for gene exchange and reticulation along suture zones between geographical groups. Data presented here allow us to make inferences about the history of the species complex. However, evidence of ancestral polymorphism and reticulation limit our ability to define species boundaries based on mitochondrial DNA sequence variation.