The Hawaiian Drosophilidae is one of the best examples of rapid speciation in nature. Nearly 1,000 species of endemic drosophilids have evolved in situ in Hawaii since a single colonist arrived over 25 million years ago. A number of mechanisms, including ecological adaptation, sexual selection, and geographic isolation, have been proposed to explain the evolution of this hyperdiverse group of species. Here, we examine the known ecological associations of 326 species of endemic Hawaiian Drosophilidae in light of the phylogenetic relationships of these species. Our analysis suggests that the long-accepted belief of strict ecological specialization in this group does not hold for all taxa. While many species have a primary host plant family, females will also oviposit on non-preferred host plant taxa. Host shifting is fairly common in some groups, especially the grimshawi and modified mouthparts species groups of Drosophila, and the Scaptomyza subgenus Elmomyza. Associations with types of substrates (bark, leaves, flowers) are more evolutionarily conserved than associations with host plant families. These data not only give us insight into the role ecology has played in the evolution of this large group, but can help in making decisions about the management of rare and endangered host plants and the insects that rely upon them for survival.
BackgroundThe past several years have seen a flurry of papers seeking to clarify the utility and limits of DNA barcoding, particularly in areas such as species discovery and paralogy due to nuclear pseudogenes. Heteroplasmy, the coexistence of multiple mitochondrial haplotypes in a single organism, has been cited as a potentially serious problem for DNA barcoding but its effect on identification accuracy has not been tested. In addition, few studies of barcoding have tested a large group of closely-related species with a well-established morphological taxonomy. In this study we examine both of these issues, by densely sampling the Hawaiian Hylaeus bee radiation.ResultsIndividuals from 21 of the 49 a priori morphologically-defined species exhibited coding sequence heteroplasmy at levels of 1-6% or more. All homoplasmic species were successfully identified by COI using standard methods of analysis, but only 71% of heteroplasmic species. The success rate in identifying heteroplasmic species was increased to 86% by treating polymorphisms as character states rather than ambiguities. Nuclear pseudogenes (numts) were also present in four species, and were distinguishable from heteroplasmic sequences by patterns of nucleotide and amino acid change.ConclusionsHeteroplasmy significantly decreased the reliability of species identification. In addition, the practical issue of dealing with large numbers of polymorphisms- and resulting increased time and labor required - makes the development of DNA barcode databases considerably more complex than has previously been suggested. The impact of heteroplasmy on the utility of DNA barcoding as a bulk specimen identification tool will depend upon its frequency across populations, which remains unknown. However, DNA barcoding is still likely to remain an important identification tool for those species that are difficult or impossible to identify through morphology, as is the case for the ecologically important solitary bee fauna.
The only bees native to the Hawaiian Islands form a single clade of 60 species in the genus Hylaeus. The group is understudied and relatively poorly known. A data set consisting of 1201 base pairs of the mitochondrial genes cytochrome oxidase I and II and tRNALeucine, and 14 morphological characters was used to construct a phylogenetic tree for 48 of the 60 known species. Genetic variation was high, including amino acid changes, and a number of species showed evidence of heteroplasmy. Tree support was low due to high levels of homoplasy. Biogeographical analysis using DIVA indicates that early radiation took place on the island of Hawaii. This places an upper age limit of only 0.4-0.7 Myr for the group, an unusually short time for such a large radiation. Moreover, it is an unusual biogeographical pattern among the Hawaiian biota.
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