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Determining Parasitoid Species Composition in a Host Population: a Molecular Approach Author(S): Kelley J

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Determining Parasitoid Species Composition in a Host Population: a Molecular Approach Author(S): Kelley J Determining Parasitoid Species Composition in a Host Population: A Molecular Approach Author(s): Kelley J. Tilmon, Bryan N. Danforth, William H. Day, and Michael P. Hoffmann Source: Annals of the Entomological Society of America, 93(3):640-647. 2000. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1603/0013-8746(2000)093[0640:DPSCIA]2.0.CO;2 URL: http://www.bioone.org/doi/full/10.1603/0013-8746%282000%29093%5B0640%3ADPSCIA %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. GENETICS Determining Parasitoid Species Composition in a Host Population: A Molecular Approach 1 KELLEY J. TILMON, BRYAN N. DANFORTH, WILLIAM H. DAY, AND MICHAEL P. HOFFMANN Department of Entomology, Cornell University, Ithaca, NY 14853 Ann. Entomol. Soc. Am. 93(3): 640Ð647 (2000) ABSTRACT Larvae of closely related parasitoid taxa often lack morphological differences that can be used for species level identiÞcation. Determining the parasitoid species present in a host population may require rearing, often a time-consuming process. To monitor Þeld parasitism rates by several species of Peristenus wasps (Hymenoptera: Braconidae) that are natural enemies of Lygus (Heteroptera: Miridae), we have developed a two-step molecular approach. Polymerase chain reaction (PCR) of the COI gene with wasp-targeting primers is performed on DNA extracted from a Lygus nymph and the parasitoid larva (if any) therein. A positive reaction indicates parasitoid presence. A restriction digest of the PCR product then indicates which parasitoid species is present among known alternatives, and a diagnosis is achieved in days rather than weeks or months. KEY WORDS Lygus lineolaris, Peristenus, parasitoid, DNA identiÞcation technique, PCR-RFLP DETERMINING THE IDENTITY of parasitoids attacking a clude species and even generic determination (Loan host species in different habitats and locations is rel- and Shaw 1987). A careful study of larval head sclerites evant to understanding both ecological and evolution- by Carignan et al. (1995) failed to reveal characters ary relationships between hosts and parasitoids, and to that separate three geographically overlapping assessing biological control of pestiferous hosts. The nymphal parasitoids of L. lineolaris (Palisot), P. digo- host range of a parasitoid species is similarly impor- neutis Loan, P. pallipes (Curtis), and P. pseudopallipes tant. Predictions about the host range and impact of an Loan. Also, new Peristenus species such as P. conradi introduced parasitoid in a new region, with new po- Marsh (Day et al. 1992) and P. stygicus Loan continue tential hosts, are difÞcult to make with accuracy. to be introduced from abroad. Larval morphology also Postintroduction data for already-established intro- fails to distinguish Peristenus from Leiophron Nees, duced parasitoids could prove valuable for developing another euphorine genus parasitic on Lygus and other criteria for release programs, and shed light on natural mirids (Loan and Shaw 1987, Day and Saunders 1990), processes of geographic dispersion and host range further complicating the study of this system. expansion. Studies of parasitoid populations in Lygus have re- The parasitoidÐhost associations between Lygus lied on a combination of dissection and rearing (Day Hahn (Heteroptera: Miridae: Mirini) and Peristenus 1994). The more easily identiÞed wasp adults are Foerster (Hymenoptera: Braconidae: Euphorinae) ephemeral and difÞcult to sample. Dissection of a are good models for studying the pattern and impact Lygus subsample gives the best estimate of percent of parasitoid introduction events. In North America, parasitism, and rearing of a paired subsample gives the there are both native and introduced Peristenus spe- identity of the parasitoid species present (Day 1994). cies with overlapping geographic and host ranges. This However, a postemergence diapause of up to 11 mo, system can provide important data on the abundance depending on the species, precedes adult emergence. and impact of a new parasitoid on target and nontarget With species or generations that have long diapause, hosts, and on the interaction between native and in- the wasp cocoons must be held in summer and then troduced natural enemies of a common host (Kuhl- winter conditions for several months. With parasitoids mann et al. 1998). of Lygus, it is not uncommon to wait 8 mo or more for Because the period of interaction between the adult information on the parasitoid species composition in parasitoid and its individual target is quite brief, Þeld a host population. studies must often focus on already-parasitized hosts Studies on the ecological interaction of Lygus para- and the immature parasitoids within them. However, sitoids, as well as programs to monitor the spread of a frequent obstacle to obtaining parasitoidÐhost data introduced Peristenus species, could beneÞt from a is the issue of identity. The reduced morphology of simple, rapid, and reliable laboratory assay to detect parasitic larvae such as Peristenus immatures can pre- and distinguish among known parasitoid species. Though related species may be morphologically in- 1 BeneÞcial Insects Research Laboratory, USDA-ARS, 501 S. distinct as immatures, their molecular differences can Chapel Street, Newark, DE 19713. provide useful characters for classiÞcation. Molecular 0013-8746/00/0640Ð0647$02.00/0 ᭧ 2000 Entomological Society of America May 2000 TILMON ET AL.: PARASITOID IDENTIFICATION TECHNIQUE 641 variation at the protein and DNA levels has been used third Peristenus species included in this initial study is for insect species diagnostics (Black et al. 1992; Sper- P. conradi. P. conradi prefers the introduced Adelpho- ling et al. 1995; Antolin et al. 1996; Taylor et al. 1996, coris lineolatis (Goeze), but parasitizes L. lineolaris at 1997). Furthermore, in parasitological studies, the mo- a low frequency (Day et al. 1992). A fourth native lecular genetic differences between parasite and host species parasitic on L. lineolaris in the northeast is may be used for parasite detection. Serological and Leiophron uniformis (Gahan); our initial efforts have electrophoretic techniques have proven useful for de- focused on the genus Peristenus and L. uniformis was tecting pathogens in insect vectors (Higgins and Azad not included in this study. 1995) as well as parasitism in herbivorous pests (Ho¨ller and Braune 1988, Stuart and Greenstone Materials and Methods 1997). The development of the PCR technique has further simpliÞed the detection of insect endobionts Characterization of Genetic Variability. The basis (Higgins and Azad 1995, Zhu and Greenstone 1999). of this laboratory technique to detect and distinguish The purpose of this study was to identify some of the parasitoid species is that portions of the parasitoid genetic variation between Lygus and Peristenus, and DNA will differ from the host as well as from each genetic variation among select species of Peristenus, other. We extracted DNA from adult specimens of L. and use this information to develop a diagnostic tool lineolaris, P. digoneutis, P. pallipes, and P. conradi. for the detection and identiÞcation of parasitism in L. Wasps were collected from several locations in the lineolaris. northeastern United States and authoritatively iden- Host and Parasitoid Species. The host species in this tiÞed by W.H.D., and preserved at Ϫ80ЊC. L. lineolaris study, Lygus lineolaris, is native to and occurs through- specimens were Þeld collected from alfalfa in Tomp- out North America (Kelton 1975). One of the broadest kins County, NY, and preserved in 95% ethanol at herbivore generalists, it has been documented on 328 Ϫ20ЊC. We removed the abdomen from adult L. line- plant species of 55 plant families in 30 orders (Young olaris specimens before DNA extraction to ensure 1986). In addition to its food plants, Lygus has been against parasitoid contamination (though parasitoid documented to feed opportunistically on numerous larvae are seldom found in adult Lygus). We followed live and dead insects (Wheeler 1976), and on the standard protocols for DNA extraction (Doyle and blood of faculty (Myers 1929) and graduate students Doyle 1987, 1990) as outlined below. Specimens were (K.J.T., unpublished data). ground in individual 1.5-ml Eppendorf tubes in the Peristenus digoneutis, a European species parasitic presence of 2ϫ CTAB extraction buffer and 100 ␮gof on L. rugulipennis Poppius, was successfully intro- proteinase K. Tubes were incubated for2hat55ЊC, duced in northern New Jersey by the USDA in the late homogenates were extracted with chloroform- 1970s and early 1980s for biological control
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