Observations on the taxonomy and natural history of Ophraella Wilcox (Coleoptera: Chrysomelidae), with a description of a new species.

-Morphological evidence indicates that the North American galerucine genus Ophraella is a strictly monophyletic taxon, closely related to Monoxia LeConte and Erynephala Blake. Host associations for all species, including several hitherto lacking host records, are listed; all hosts are in the Asteraceae. Distinguishing morphological features of adults are described, and descriptions of immature stages of most of the species are provided. 0. ma crovittata LeSage is placed in synonymy with 0. sexvittata (LeConte), and the possibility of conspecificity of 0. pilosa LeSage and 0. americana (Fabricius) is raised. A new species, Ophraella artemisiae, is described from Texas. Several features of the life history are described. Phenological data indicate that northern populations and species are univoltine, whereas more southern populations are generally mul tivoltine. Cocoon formation differs from previous reports in that the cocoon material issues from the rectum. Several natural enemies were observed, including, in 0. communa LeSage, a eulophid genus not previously recorded in North America. Geographically isolated populations of 0. communa display partial sexual isolation, as do closely related sympatric species (0. communa and 0. notulata [Fabricius]) which occupy different host plants. A difference in host affiliation is therefore not the only barrier to gene exchange. Species of Ophraella Wilcox, a North American genus of leaf beetles (Chryso melidae: Galerucinae, tribe Galerucini), are locally abundant throughout much of their range. Nevertheless, little is known of the biology of the genus, perhaps because the host plants have little economic importance. A few comments on host associations and other aspects of the biology of some species are provided by Blatchley (1910), Woods (1924), Wilcox (1965), Messina and Root (1980), and LeSage (1986); the life history of only one species has been described in any detail (Welch, 1978; Goeden and Ricker, 1985). The genus was revised by LeSage in 1986. In the course of an ongoing analysis of the phylogeny and of the evolution and genetics of host association in the genus, I have made a variety of observations that increase the taxonomic and biological knowledge of Ophraella. Although some of these observations are not quantitative, investigators embarking on the study of little known taxa will find qualitative observations more helpful than none at all. In subsequent publications I shall treat the morphology of Ophraella and related genera, and the phylogeny and evolution of host associations within the genus, based on morphological and electrophoretic analyses. In another publication (Futuyma, in press), I have described feeding responses of many of the species to their congeners" host plants. This content downloaded from 157.55. 39.162 on Thu, 11 Aug 2016 05:34:19 UTC All use subject to http://about.jstor. org/terms 164 JOURNAL OF THE NEW YORK ENTOMOLOGICAL SOCIETY Vol. 98(2) MONOPHYLY OF OPHRAELLA Wilcox (1965) segregated Ophraella from Galerucella, and considered it most closely related to Erynephala Blake, Monoxia LeConte, Ophraea Jacoby, and several Neo tropical genera. White (1979) referred the Neotropical Trirhabda dilatipennis Jacoby to Ophraella, but LeSage (1986) later placed this species in Neolochmaea Laboissiere. I have examined the morphology of this species, as well as species of Erynephala, Monoxia, and the holarctic taxon Pyrrhalta (subgenus Tricholochmaea) Laboissiere. With reference to Neolochmaea and Pyrrhalta, the close relationship of Ophraella to Monoxia and Erynephala is supported by several synapomorphies not hitherto noted, including (1) spermathecal duct inserted subbasally (rather than extending rectilinearly from base); (2) sternum VIII of the female lacking the long anterior rod like apodeme present in the other genera; (3) styli of the female (referred to as "accessory glands" by LeSage (1986)) reduced to flat setiferous discs (rather than elongate); (4) internal sac of male lacking ornamentation; (5) internal scuto-scutellar sclerites of larva each bearing laterally what appears to be a setaless socket (absent in the other genera). Brivio (1977) has noted that the above differences in female genitalia between Ophraella and Pyrrhalta also distinguish Ophraella from Galeru cella (s.s.) and, for the most part, from Xanthogaleruca. Ophraella shares with Mo noxia, its apparent sister group, the following synapomorphies: (1) pupation within a loosely woven cocoon rather than free; (2) sternum VIII of female with lateral extensions of the base (absent in the other genera); (3) apex of the male sternum deeply, conically invaginated (vs. shallowly or not at all); (4) aedeagus (median lobe) basally with a dorsal ring (vs. absent), this laterally confluent with the basal spurs. In addition to the features noted by LeSage (1986), the species of Ophraella share the following synapomorphic characters with respect to the condition in Erynephala and Monoxia: (1) mandible of adult with three rather than five teeth; (2) teeth IV and V of larval mandible only shallowly (rather than fully) separated, hence appearing fused basally; (3) larval cuticle lacking discrete aggregations of pigment ("melano phores"); (4) setae of dorsal sclerites of larva almost uniform in length (strongly variable in the other genera). Electrophoresis of 18 enzyme loci also supports the monophyly of Ophraella relative to Monoxia and Erynephala (Futuyma and Mc Cafferty, in prep.). TAXONOMY AND HOST AFFILIATION LeSage (1986) recognized 13 species of Ophraella, of which he described five as new, renaming two others. He listed host records for 11 species. My records confirm LeSage's (1986) and Wilcox's (1965) supposition that all the hosts of Ophraella are Asteraceae (Compositae) (Table 1). Below I present an account of my present un derstanding of the taxonomic status and host affiliation of each of the species. (1) 0. americana (Fabricius) and 0. pilosa LeSage. LeSage reported host records for 0. americana as "Solidago spp. ," and distinguished 0. pilosa from 0. americana on the basis of slight differences in genitalia, markings, average body size, the density of erect setae on the elytra, and association with Aster. Collections from Aster species (especially A. urophyllus) in central New York, Missouri (Dade Co.), Montana (Lake Co.), and Ontario (Leeds Co.) all conform morphologically to the description of 0. pilosa. I have collected these forms on Solidago only in Ithaca, New York, where This content downloaded from 157.55. 39.162 on Thu, 11 Aug 2016 05:34:19 UTC All use subject to http://about.jstor. org/terms 1990 TAXONOMY AND BIOLOGY OF OPHRAELLA 165 Table 1. Known host plants of Ophraella species. Notations following host records: L, literature records (cf. LeSage, 1986; Goeden and Ricker, 1985); P, personal record; *, host not previously reported. Localities for host records are indicated in some instances. Attributions to other collectors are based on reliable personal communications.