Arthur M. Shapiro

Homoploid Hybrid Speciation in an Extreme Habitat

According to theory, homoploid hybrid speciation, which is hybrid speciation without a change in chromosome number, is facilitated by adaptation to a novel or extreme habitat. Using molecular and ecological data, we found that the alpine-adapted butterflies in the genus Lycaeides are the product of hybrid speciation. The alpine populations possess a mosaic genome derived from both L. melissa and L. idas and are differentiated from and younger than their putative parental species. As predicted, adaptive traits may allow for persistence in the environmentally extreme alpine habitat and reproductively isolate these populations from their parental species.

Compounded effects of climate change and habitat alteration shift patterns of butterfly diversity.

Climate change and habitat destruction have been linked to global declines in vertebrate biodiversity, including mammals, amphibians, birds, and fishes. However, invertebrates make up the vast majority of global species richness, and the combined effects of climate change and land use on invertebrates remain poorly understood. Here we present 35 years of data on 159 species of butterflies from 10 sites along an elevational gradient spanning 0–2,775 m in a biodiversity hotspot, the Sierra Nevada Mountains of Northern California. Species richness has declined at half of the sites, with the most severe reductions at the lowest elevations, where habitat destruction is greatest. At higher elevations, we observed clear upward shifts in the elevational ranges of species, consistent with the influence of global warming. Taken together, these long-term data reveal the interacting negative effects of human-induced changes on both the climate and habitat available to butterfly species in California. Furthermore, the decline of ruderal, disturbance-associated species indicates that the traditional focus of conservation efforts on more specialized and less dispersive species should be broadened to include entire faunas when estimating and predicting the effects of pervasive stressors.

Exotics as host plants of the California butterfly fauna

Abstract Introduced species may impact native species and communities in many ways. One which has received relatively little attention is by serving as resources for natives, thereby altering their ecology. We address such impacts on the California butterfly fauna as currently understood. Eighty-two of Californias approximately 236 butterfly species (34%) are reported as ovipositing or feeding on introduced plant taxa. Many more utilize introduced plants as nectar sources. Interactions with introduced plant taxa are not distributed evenly among butterfly species. Alpine and desert butterflies interact with relatively few introduced plants because few exotic plant species have reached and successfully colonized these habitats. Other California butterfly species are specialists on particular plant families or genera with no exotic representatives in California and have thus far failed to recognize any introduced plants as potential foodplants. Some California butterflies have expanded their geographic ranges and/or extended their flight seasons by feeding on exotic plants. However, negative impacts of exotic plant species can also occur. At least three of the states butterfly species currently lay eggs on introduced taxa that are toxic to larvae. Impacts of introduced plant taxa on Californias butterflies are expected to increase as both habitat conversion and alien introductions accelerate.

The significance of wing pattern diversity in the Lycaenidae: mate discrimination by two recently diverged species

Closely related species of lycaenid butterflies are determinable, in part, by subtle differences in wing pattern. We found that female wing patterns can act as an effective mate‐recognition signal in some populations of two recently diverged species. In field experiments, we observed that males from a Lycaeides idas population and an alpine population of L. melissa preferentially initiate courtship with conspecific females. A morphometric study indicated that at least two wing pattern elements were important for distinguishing the two species: hindwing spots and orange crescent‐shaped pattern elements called aurorae. We deceived male L. idas into initiating courtship with computer generated paper models of heterospecific females when these pattern elements were manipulated, indicating that the wing pattern elements that define the diversity of this group can be effective mate recognition signals.

Identifying units for conservation using molecular systematics: the cautionary tale of the Karner blue butterfly

The federally endangered North American Karner blue butterfly (Lycaeides melissa samuelis) and the closely related Melissa blue butterfly (L. m. melissa) can be distinguished based on life history and morphology. Western populations of L. m. samuelis share mitochondrial haplotypes with L. m. melissa populations, while eastern populations of L. m. samuelis have divergent haplotypes. Here we test two hypotheses concerning the presence of L. m. melissa mitochondrial haplotypes in western L. m. samuelis populations: (i) mitochondrial introgression has occurred from L. m. melissa populations into western L. m. samuelis populations, or (ii) western populations of the nominal L. m. samuelis are more closely related to L. m. melissa than to eastern L. m. samuelis populations, yet are phenotypically similar to the latter. A Bayesian algorithm was used to cluster 190 L. melissa individuals based on 143 informative amplified fragment length polymorphism (AFLP) loci. This method clearly differentiated L. m. samuelis and L. m. melissa. Thus, genomic divergence was greater between western L. m. samuelis populations and L. m. melissa populations than it was between western and eastern populations of L. m. samuelis. This supports the hypothesis that the presence of L. m. melissa mitochondrial haplotypes in western L. m. samuelis populations is the result of mitochondrial introgression. These data provide valuable information for conservation and management plans for the endangered L. m. samuelis, and illustrate the risks of using data from a single locus for diagnosing significant units of biodiversity for conservation.

The Role of Sexual Behavior in Density-Related Dispersal of Pierid Butterflies

Sexual interactions in dense populations of a butterfly, Pieris protodice, lead to emigration of gravid females which colonize new habitats. Some other species of Pierids respond to high population density and skewed sex ratios early in each generation by aggregation of males outside the breeding area.

THE PONTIA DAPLIDICE-EDUSA HYBRID ZONE IN NORTHWESTERN ITALY

The pierid butterflies Pontia daplidice and P. edusa, parapatrically distributed in southern Europe, have very similar morphologies and life histories, but show fixed differences at four allozyme markers. We sampled these allozymes in a 28‐population transect north of Genoa in Italy, through the hybrid zone where these taxa meet. We used the numerical techniques developed for hybrid zone analysis to study the patterns of genetic differentiation and their underlying evolutionary causes. The hybrid zone is characterized by a very short and steep central region, flanked by broad tails of introgression extended up to 100 km in either direction. From mean two‐locus disequilibium of D = 0.148 (maximum‐likelihood two‐unit support limits 0.139‐0.153), and after accounting for minor differences in the center locations of the single‐locus clines, which act to bias the dispersal estimate, we estimated a dispersal rate of σ = 4.4 (3.7‐5.5) km/gen1/2. The effective selection needed to maintain the steep central portion is strong, 0.47 < s∗ < 0.64, when combined over potential intrinsic (genetic background) and extrinsic (ecological) sources of selection. The clines in allozyme loci showed variation that was significantly different between the most divergent shapes, and the differences are attributable to different degrees of introgression on the edusa side of the zone. The average selection acting on individual allozyme loci was high at s∗∗∗e  1.5%, but because of the narrowness of the central region of the cline, we suspect that this estimate is somewhat biased by selection on loci closely linked to the allozyme markers. A common question for taxa that show fixed allozyme differences in parapatry is whether or not they are genetically isolated. A fairly general measure of genetic isolation across hybrid zones is the time, T, that it takes a neutral allele to cross the hybrid zone and recombine into the opposite genetic background, given by T = (β/σ)2, where β is the barrier strength of the hybrid zone. Genetic isolation in the Pontia zone is weak, with T  25 generations for most allozyme markers. By this measure, populations of daplidice and edusa on opposite sides of the hybrid zone share more identical‐by‐descent alleles than do populations of phenotypically pure daplidice in, say, France and Morocco. Accordingly, we think it best for systematists to consider edusa as a well‐marked subspecies of P. daplidice.

Hypersensitivity reaction of Brassica nigra L. (Cruciferae) kills eggs of Pieris butterflies (Lepidoptera: Pieridae)

SummarySome individual plants of the mustard Brassica nigra in lowland California kill eggs of the Crucifer-specialist herbivores Pieris rapae and P. napi by producing a necrotic zone at the base of the egg, thereby apparently desicating it. This is a typical hypersensitivity reaction, but to an atypical stimulus. The eggs can be “rescued” by maintaining them in a saturated atmosphere. Attempts to demonstrate a bacterial or fungal agent associated with the reaction were unsuccessful.

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 (Neis 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.

ANOTHER PERSPECTIVE ON THE SLOW-GROWTH/HIGH-MORTALITY HYPOTHESIS: CHILLING EFFECTS ON SWALLOWTAIL LARVAE

The slower-growth/higher-mortality hypothesis proposes that reduced herbivore growth rates benefit plants because slower growing herbivores remain vulnerable to predator attack for an extended time period, resulting in a lower reduction in plant fitness attributed to herbivory. We propose cooling events as an alternative mechanism leading to higher mortality for slower growing larvae. We observed that, in the absence of predators, slower growing pipevine swallowtail (Battus philenor) larvae had higher mortality compared to faster growing larvae during the unseasonably cool spring temperatures of 1998. However, this was not true for the warmer spring of 1999. Laboratory experiments showed that the probability of surviving a chill coma increased with larval mass. We propose that smaller larvae are susceptible to chilling events because they have less energy reserved for metabolism during a chill coma, and that “warm” chill events near the activity threshold may be more lethal than “cold” chill events. Weath...