Rapid diversification of sexual signals in Hawaiian Nesosydne planthoppers (Hemiptera: Delphacidae): the relative role of neutral and selective forces.

Changes in sexual signals have the potential to promote rapid divergence and reproductive isolation among populations of animals. Thus, identifying processes contributing to variation in signals is key to understanding the drivers of speciation. However, it is difficult to identify the processes initiating changes in signals in empirical systems because (1) the demographic history of populations under study is usually unclear, and (2) there is no unified hypothesis-testing framework for evaluating the simultaneous contribution of multiple processes. A unique system for study in the Hawaiian Islands, the planthopper species Nesosydne chambersi, offers a clear demographic context to disentangle these factors. By measuring variation in male vibratory sexual signals across different genetic populations on the island of Hawaii, we found that that multiple signal traits varied significantly between populations. We developed a mixed modelling framework to simultaneously test competing hypotheses about which processes contribute to changes in signal traits: genetic drift, sensory drive or reproductive character displacement. Our findings suggest that signal divergence proceeds along different axes for different signal traits under the influence of both neutral and selective processes. They are the first, to our knowledge, to document the relative importance of multiple processes on divergence in sexual signals.

Isolation and characterization of nine microsatellite loci from Bemisia tabaci (Hemiptera: Aleyrodidae) biotype B.

Nine microsatellites were isolated from Bemisia tabaci (Gennadius) biotype B and screened across 60 individuals from two populations (biotype B) to examine polymorphism. Two to 12 alleles were observed per locus. Observed and expected heterozygosities ranged from 0.033 to 0.967 and 0.033 to 0.854, respectively. There was no significant deviation from Hardy-Weinberg equilibrium and no significant linkage disequilibrium between loci. One locus showed evidence for null alleles. These loci will be useful in future studies of the genetic structure of worldwide biotypes and gene flow analyses between and within biotypes of B. tabaci.

Domestication of olive fly through a multi-regional host shift to cultivated olives: comparative dating using complete mitochondrial genomes.

The evolutionary history of the olive fly, Bactrocera oleae, was reconstructed in a phylogenetic and coalescent framework using full mitochondrial genome data from 21 individuals covering the entire worldwide distribution of the species. Special attention was given to reconstructing the timing of the processes under study. The early subdivision of the olive fly reflects the Quaternary differentiation between Olea europea subsp. europea in the Mediterranean area and the two lineages of Olea europea subsp. cuspidata in Africa and Asia, pointing to an early and close association between the olive fly and its host. The geographic structure and timing of olive fly differentiation in the Mediterranean indicates a clear connection with the post-glacial recolonization of wild olives in the area, and is irreconcilable with the early historical process of domestication and spread of the cultivated olive from its Levantine origin. Therefore, we suggest an early co-history of the olive fly with its wild host during the Quaternary and post-glacial periods and a multi-regional shift of olive flies to cultivated olives as these cultivars gradually replaced wild olives in historical times.

Molecular markers reveal strong geographic, but not host associated, genetic differentiation in Aphidius transcaspicus, a parasitoid of the aphid genus Hyalopterus.

Host plant associated genetic differentiation is a common phenomenon in phytophagous insects, but the degree to which such associations sequentially drive diversification at higher trophic levels is not as well understood. A recent study examining neutral molecular markers in Hyalopterus aphids revealed that genetic structure in this genus is strongly determined by primary host plant use (Prunus spp.). In this paper, we take a similar approach to determine whether this host plant specificity has affected genetic structure in the parasitoid Aphidius transcaspicus, an important natural enemy of Hyalopterus spp. in the Mediterranean. Mitochondrial DNA (428 bp) and seven microsatellite loci were examined in parasitoids collected from aphid populations on almond, apricot, peach and plum trees from Spain and Greece. In contrast to the previous findings for Hyalopterus from the same regions, here we find no evidence for host associated diversification in A. transcaspicus at the species level or below, though geographic structure between regional populations is exceptionally high. These findings have several implications for our understanding of the ecology and evolution of A. transcaspicus as well as for its use as a biological control agent for Hyalopterus, suggesting that a consideration of host plant specificity may be less critical than factors such as climatic suitability or geographic origins of invasive populations.

Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago.

The extant endemic katydids (Orthoptera: Tettigoniidae) of the Hawaiian Archipelago include one to three species per high island and a single species on Nihoa, all currently placed in the genus Banza. These acoustic insects provide an excellent opportunity for investigating the evolution of reproductive isolation and speciation, but such studies require an understanding of phylogenetic relationships within the group. We use maximum parsimony, likelihood-based Bayesian inference, and maximum likelihood to infer phylogenetic relationships among these taxa, based on approximately 2kb of mitochondrial cytochrome oxidase I and cytochrome b. Our results strongly support two distinct high island clades: one clade ("Clade I") composed of species from Kauai, Oahu, Molokai, and Lanai and another clade ("Clade II") composed of species from Maui and Hawaii (Banza unica, from Oahu, may be basal to both these clades, but its placement is not well resolved). Within these clades, some inferred relationships are strongly supported, such as the sister status of B. kauaiensis (Kauai) and B. parvula (Oahu) within Clade I, but other relationships remain more ambiguous, such as the relative position of B. brunnea (Maui) within Clade II. Although a detailed reconstruction of the historical biogeography of the Hawaiian katydids is difficult, we use our genetic data combined with the known geological history of the Hawaiian Islands to set limits on plausible historical scenarios for diversification of this group. Beyond these historical biogeographic inferences, our results indicate possible cryptic speciation on both Oahu and Hawaii, as well as what may be unusually high average rates of nucleotide substitution. The present work sets the stage for future genetic and experimental investigations of this group.

Bioinvasions of the medfly Ceratitis capitata: source estimation using DNA sequences at multiple intron loci.

The Mediterranean fruit fly, Ceratitis capitata, is a devastating agricultural pest that threatens to become established in vulnerable areas such as California and Florida. Considerable controversy surrounds the status of Californian medfly infestations: Do they represent repeated introductions or the persistence of a resident population? Attempts to resolve this question using traditional population genetic markers and statistical methods are problematic because the most likely source populations in Latin America were themselves only recently colonized and are genetically very similar. Here, significant population structure among several New World medfly populations is demonstrated through the analysis of DNA sequence variation at four intron loci. Surprisingly, in these newly founded populations, estimates of population structure increase when measures of subdivision take into account the relatedness of alleles as well as their frequency. A nonequilibrium, likelihood-based statistical test that utilizes multilocus genotypes suggests that the sole medfly captured in California during 1996 was introduced from Latin America and was less likely to be a remnant of an ancestral Californian population. Many bioinvasions are hierarchical in nature, consisting of several sequential or overlapping invasion events, the totality of which can be termed a metainvasion. Phylogenetic data from multilocus DNA sequences will be vital to understanding the evolutionary and ecological processes that underlie metainvasions and to resolving their constituent levels.

Invasion genetics of the Mediterranean fruit fly: variation in multiple nuclear introns.

Biological invasions generally start from low initial population sizes, leading to reduced genetic variation in nuclear and especially mitochondrial DNA. Consequently, genetic approaches for the study of invasion history and population structure are difficult. An extreme example is the Mediterranean fruit fly, Ceratitis capitata (Medfly), for which successive invasions during this century have resulted in a loss of 60% of ancestral genetic variation in isozymes and 75% of variation in mitochondrial DNA. Using Medflies as an example, we present a new approach to invasion genetics that measures DNA sequence variation within introns from multiple nuclear loci. These loci are so variable that even relatively recently founded Medfly populations within California and Hawaii retain ample genetic diversity. Invading populations have only lost 35% of the ancestral genetic variation. Intron variation will allow high-resolution genetic characterization of invading populations in both natural and managed systems, although non-equilibrium methods of analysis may be necessary if the genetic diversity represents sorting ancestral polymorphism.

Speciation and phylogeography of Hawaiian terrestrial arthropods.

The Hawaiian archipelago is arguably the world's finest natural laboratory for the study of evolution and patterns of speciation. Arthropods comprise over 75% of the endemic biota of the Hawaiian Islands and a large proportion belongs to species radiations. We classify patterns of speciation within Hawaiian arthropod lineages into three categories: (i) single representatives of a lineage throughout the islands; (ii) species radiations with either (a) single endemic species on different volcanoes or islands, or (b) multiple species on each volcano or island; and (iii) single widespread species within a radiation of species that exhibits local endemism. A common pattern of phylogeography is that of repeated colonization of new island groups, such that lineages progress down the island chain, with the most ancestral groups (populations or species) on the oldest islands. While great dispersal ability and its subsequent loss are features of many of these taxa, there are a number of mechanisms that underlie diversification. These mechanisms may be genetic, including repeated founder events, hybridization, and sexual selection, or ecological, including shifts in habitat and/or host affiliation. The majority of studies reviewed suggest that natural selection is a primary force of change during the initial diversification of taxa.

Intraspecific mitochondrial DNA variation in the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae).

Restriction endonuclease analyses of mitochondrial DNA (mtDNA) were used to examine genetic variability and population structure in Leptinotarsa decemlineata (Say). A group of three enzymes, EcoRI, HpaI, and PstI, was used to reveal polymorphism both within and among some of the 10 populations tested, yielding 16 haplotypes in combination. The frequencies of these 16 haplotypes differed significantly across geographic regions, indicating some partitioning of mtDNA haplotypes. Estimates of mtDNA sequence divergence (delta) between haplotypes ranged from 0.016 to 0.135%, suggesting local differentiation of mtDNA in some populations. Analysis of these data suggests that Texas was colonized by more than one mtDNA lineage, most likely originating in Mexico. We hypothesize that a larger founder size for the initial introductions or high levels of variability in the parent population at the edge of the CPB expanding range led to the initial partitioning of haplotypes observed in samples from Texas.

Geographic structure of insect populations: gene flow, phylogeography, and their uses.

Geographic structure of populations is a fundamental component of ecology and evolution that combines both demographic and genetic processes, such as gene flow and migration, genetic drift, selection, and population extinction. Recent advances in both molecular biology and theory have revolutionized the field and have not only expanded the availability of data but also facilitated the accessibility and interpretation of current data. These new techniques allow analysis of genetic similarity among populations to be coupled with phylogeography and the distribution of genotypes within and among populations relative to the history of those genotypes. The numerous case studies described herein illustrate the growing impact of geographic structure on insect science, as well as the importance of insect model systems for understanding general concepts in ecology and evolution.

Population structure of a predatory beetle: the importance of gene flow for intertrophic level interactions.

Migration and gene flow of natural enemies play an important role in the stability of predator-prey interactions and community organization in both natural and managed systems. Yet, relative to that of their herbivorous insect prey, the genetic structure of natural enemy populations has been little studied. We present evidence that populations of the predatory coccinellid beetle Coleomegilla maculata (Coleoptera: Coccinellidae), are not genetically subdivided and that levels of gene flow among these populations are extremely high. Furthermore, in the same geographical area, gene flow of C. maculata was significantly (one order of magnitude) greater than that of an abundant prey species, the Colorado potato beetle Leptinotarsa decemlineata (Coleoptera: Chrysomelidae). The high mobility of this natural enemy relative to the insect herbivores on which it feeds may contribute to its effectiveness as a biological control agent in agricultural systems.