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Morphological and Genetic Divergence in Three Populations of Anthocoris Antevolens (Hemiptera: Heteroptera: Anthocoridae) Author(S): David R

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Morphological and Genetic Divergence in Three Populations of Anthocoris Antevolens (Hemiptera: Heteroptera: Anthocoridae) Author(S): David R Morphological and Genetic Divergence in Three Populations of Anthocoris antevolens (Hemiptera: Heteroptera: Anthocoridae) Author(s): David R. Horton, Thomas R. Unruh, Tamera M. Lewis, and Kelly Thomsen-Archer Source: Annals of the Entomological Society of America, 100(3):403-412. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1603/0013-8746(2007)100[403:MAGDIT]2.0.CO;2 URL: http://www.bioone.org/doi/ full/10.1603/0013-8746%282007%29100%5B403%3AMAGDIT%5D2.0.CO %3B2 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/ terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. ARTHROPOD BIOLOGY Morphological and Genetic Divergence in Three Populations of Anthocoris antevolens (Hemiptera: Heteroptera: Anthocoridae) 1 DAVID R. HORTON, THOMAS R. UNRUH, TAMERA M. LEWIS, AND KELLY THOMSEN-ARCHER USDAÐARS, 5230 Konnowac Pass Road, Wapato, WA 98951 Ann. Entomol. Soc. Am. 100(3): 403Ð412 (2007) ABSTRACT Anthocoris antevolens White (Hemiptera: Heteroptera: Anthocoridae) is a widespread predatory bug in North America that exhibits substantial geographic variation in coloration, body measurements, size and shape of the male genitalia, pubescence, and sexual behavior. Earlier be- havioral studies with three populations (including two populations that are sympatric in central Washington) showed that there was limited or no successful mating between insects from nonlike populations, despite vigorous mating attempts by males. The current study shows that males from those three populations diverge also in size and shape of claspers, length of the phallus, body measurements, and pubescence. Divergence extends to the two sympatric populations. Analysis of mitochondrial DNA shows that phenotypic divergence is associated with genetic divergence. Results reported here, in combination with the earlier published mating trials, support statements made elsewhere by us that A. antevolens is actually a complex of an unknown number of externally similar species. KEY WORDS morphometrics, genitalia, cryptic species, geographic variation, mtDNA The predatory bug Anthocoris antevolens White oak, Quercus garrayana Douglas (Fagaceae), was re- (Hemiptera: Heteroptera: Anthocoridae) is a wide- productively isolated from A. antevolens collected spread species found throughout Canada and Alaska, from neighboring stands of willow, Salix spp. (Sali- south into the New England states and northern plains caceae). Males from the two populations differed in states east of the Rockies, and south into Arizona and shape of the clasper and length of the phallus. Mor- southern California west of the Rockies (Kelton 1978, phological differences and reproductive incompati- Henry 1988). Of the 12 species of Anthocoris listed for bility were maintained in the laboratory in consecu- North America (Henry 1988), A. antevolens is geo- tive generations reared on a common diet (Horton graphically the most widespread. The bug associates and Lewis 2005). Further examination of bugs from often with deciduous trees and shrubs, especially in the Salix source suggested that this population may the Salicaceae and Rosaceae (Horton et al. 2004), and itself be composed of two distinct types, based upon it can be an important source of biological control in a limited examination of male genitalia and pubes- orchard systems of the PaciÞc northwest (McMullen cence on the hemelytra (unpublished data). Behav- and Jong 1967, Horton and Lewis 2000). ioral studies were then done that conÞrmed that two A. antevolens shows substantial geographic variation reproductively incompatible populations do indeed in size, coloration, pubescence, sexual behavior, size occupy Salix (and other host plants such as pear, Pyrus or shape of the maleÕs clasper, and length of the maleÕs communis L. [Rosaceae]) throughout the Yakima Val- phallus (Horton et al. 2005, Horton and Lewis 2005, ley (Horton et al. 2005). unpublished data). The variation is now known to Here, we explore whether the two Yakima popula- encompass even sympatric populations, and popula- tions co-occurring on both Salix and Pyrus spp., and tion divergence has been shown to be associated with shown earlier to be reproductively incompatible reproductive incompatibility between those sympat- (Horton et al. 2005), also demonstrate quantiÞable ric populations (Horton et al. 2005, Horton and Lewis differences in body measurements, size and shape of 2005). Thus, it has become clear that A. antevolens is the male genitalia, pubescence, and molecular genetic actually a complex of an unknown number of repro- composition. As in the Horton et al. (2005) study, we ductively isolated species whose external character- have included for comparison a third population col- istics nonetheless key them to A. antevolens in avail- lected from Salix Ϸ120 km west of the two Yakima able keys (Hill 1957, Kelton 1978). populations. Females from that third population in Horton and Lewis (2005) showed that A. antevolens laboratory trials were shown to be successfully insem- collected in western Yakima, WA, from Oregon white inated by males from one of the two Yakima types, albeit at low rates, but they were found to be com- 1 Corresponding author, e-mail: [email protected]. pletely incompatible with males from the second 404 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA Vol. 100, no. 3 Yakima population (Horton et al. 2005). SpeciÞc ob- jectives are to determine whether the reproductive incompatibilities noted in Horton et al. (2005) are reßected in genetic and morphological traits within the three populations. Materials and Methods Source of Insects and Rearing. Insects used in the current study were male siblings of those used in the mating studies described previously (Horton et al. 2005), with some additional material collected in a subsequent year (see below). In all cases, bugs were the Þrst-generation male offspring of Þeld-collected females. Insects from three populations (described in Horton et al. 2005) were examined. The Alder Lake (AL) population was collected in June 2002 from Salix growing along a 20-km stretch of Highway 7 between Mineral Lake and Alder Lake, WA (46Њ 71Ј N, 122Њ 22Ј W). The site is west of the Cascade Crest. A second population (GC) was collected in June 2002 from Ϸ20 pear trees growing at the Apple Tree Golf Course in western Yakima, WA (46Њ 57Ј N, 120Њ 61Ј W). The site is Ϸ120 km east of the AL site and occurs east of the Cascade Crest. The third population (UG) was col- lected June 2002 from a 100-m stretch of Salix growing along the Yakima River just southeast of Union Gap, WA (46Њ 52Ј N, 120Њ 47Ј W). The UG and GC sites are separated by 12 km. However, we occasionally Þnd UG and GC type bugs co-occurring throughout the Yakima Valley on Salix spp. and Pyrus communis (un- Fig. 1. (A) Fully inßated phallus. (B) Stored phallus published data). We have examined enough insects within the genital capsule, with arrows showing points of from both the UG and GC populations that we can dissection. (C) Phallus pulled from genital capsule for mea- now differentiate between the two phenotypes based suring (also see Horton and Lewis 2005). eb, ejaculatory upon external characteristics (the GC bugs are gen- bulb; gc, genital capsule. erally darker and have less noticeable pubescence); we carefully examined our Þeld-collected material before beginning the studies described below, to en- summarized below, producing sample sizes of 15 males sure that parental bugs were not a mix of the two per population. phenotypes within either the GC or UG collections. Male Genitalia. We estimated length of the phallus For each population, 10 mated females were placed and described size and shape of the left clasper for individually in small cages (135-ml ventilated vials) each specimen (Horton and Lewis 2005). The phallus containing prey [nymphs and eggs of pear psylla, Ca- in A. antevolens is a thin, membranous two-walled copsylla pyricola (Fo¨rster) (Homoptera: Psyllidae)], organ (Fig. 1) that must be fully inßated within the and small pear seedlings. The bugs readily oviposited femaleÕs copulatory tube to allow insemination (Hor- into the pear seedlings. Offspring were reared at 22Ð ton and Lewis 2005). Phallus length was estimated by 24ЊC and a photoperiod of 16:8 (L:D) h. Large nymphs dissecting the uninßated organ from the genital cap- were removed from the cages and placed individually sule and measuring length of the dissected organ be- in petri dishes (9 cm in diameter) lined with Þlter neath a dissecting microscope. Measurement has high paper and containing psylla-infested pear leaves. As repeatability (Horton and Lewis 2005). A single indi- the nymphs eclosed to the adult stage, date of matu- vidual did
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