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Gene Flow in the Exotic Colonizing Ladybeetleharmonia Axyridisin North America E

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Gene Flow in the Exotic Colonizing Ladybeetleharmonia Axyridisin North America E View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Digital Repository @ Iowa State University Entomology Publications Entomology 3-1997 Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America E. S. Krafsur Iowa State University, [email protected] T. J. Kring University of Arkansas, Fayetteville J. C. Miller Oregon State University P. Nariboli Iowa State University J. J. Obrycki Iowa State University SeFoe nelloxtw pa thige fors aaddndition addal aitutionhorsal works at: http://lib.dr.iastate.edu/ent_pubs Part of the Entomology Commons, Evolution Commons, Other Ecology and Evolutionary Biology Commons, and the Population Biology Commons The ompc lete bibliographic information for this item can be found at http://lib.dr.iastate.edu/ ent_pubs/407. For information on how to cite this item, please visit http://lib.dr.iastate.edu/ howtocite.html. This Article is brought to you for free and open access by the Entomology at Iowa State University Digital Repository. It has been accepted for inclusion in Entomology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Gene Flow in the Exotic Colonizing LadybeetleHarmonia axyridisin North America Abstract Gene flow was studied inHarmonia axyridis(Pallas) (Coleoptera: Coccinellidae), an exotic, arboreal ladybeetle predator that recently spread rapidly throughout North America. A survey of isozyme polymorphisms showed 30 of 52 resolved putative loci were polymorphic (58%), and the mean heterozygosity was 16.75 ± 2.98% among all loci and 26.31 ± 4.37% at only the polymorphic loci. The mean number of alleles at the 52 loci was 2.01 ± 1.97. Gene frequencies were estimated in populations from Georgia, Virginia, Delaware, Rhode Island, Arkansas, Illinois, Iowa, and Oregon and differed significantly (P < 0.02) at 11 of 16 loci. Three of 16 loci (Fbp, Est-1, Tre) were not in Hardy–Weinberg equilibrium in all populations and are not included in theFstatistics. Random mating was indicated within populations (FIS = −0.005 ± 0.014) but not among populations (FST = 0.025 ± 0.005). According to Wright's island model,FSTestimates the average number of reproducing immigrants to be equivalent to ca. 10 beetles per population per generation. Thus there was a measure of genetic differentiation caused by drift, but this differentiation was small with respect to the large geographical distances among the sampled populations. The genetic data suggest that the founding North American population(s) was substanti Keywords Harmonia axyridis, isozyme variation, breeding structure, gene flow, colonizing species Disciplines Entomology | Evolution | Other Ecology and Evolutionary Biology | Population Biology Comments This article is from Biological Control 8 (1997); 207, doi: 10.1006/bcon.1997.0508. Rights Works produced by employees of the U.S. Government as part of their official duties are not copyrighted within the U.S. The onc tent of this document is not copyrighted. Authors E. S. Krafsur, T. J. Kring, J. C. Miller, P. Nariboli, J. J. Obrycki, J. R. Ruberson, and P. W. Schaefer This article is available at Iowa State University Digital Repository: http://lib.dr.iastate.edu/ent_pubs/407 BIOLOGICAL CONTROL 8, 207–214 (1997) ARTICLE NO. BC970508 Gene Flow in the Exotic Colonizing Ladybeetle Harmonia axyridis in North America E. S. Krafsur,* T. J. Kring,† J. C. Miller,‡ P. Nariboli,* J. J. Obrycki,* J. R. Ruberson,§ and P. W. Schaefer\ *Department of Entomology, Iowa State University, Ames, Iowa 50011-3222; †Department of Entomology, University of Arkansas, Fayettville, Arkansas 72701; ‡Department of Entomology, Oregon State University, Corvallis, Oregon 97331; §Department of Entomology, University of Georgia, Tifton, Georgia 31793; and \ USDA-ARS, Beneficial Insect Introduction Research Laboratory, Newark, Delaware 19713 E-mail: [email protected] Received November 4, 1996; accepted January 21, 1997 herbaceous habitats in the United States (Tedders and Gene flow was studied in Harmonia axyridis (Pallas) Schaefer, 1994; T. Kring, unpublished data). (Coleoptera: Coccinellidae), an exotic, arboreal lady- Numerous attempts have been made to add H. beetle predator that recently spread rapidly through- axyridis to the North American fauna, the first of which out North America. A survey of isozyme polymor- was in 1916 in California (Gordon, 1985). From 1978 to phisms showed 30 of 52 resolved putative loci were 1986, 11 H. axyridis collections, ranging from 35 to 438 polymorphic (58%), and the mean heterozygosity was adults (mean 167), were made in Japan, Korea, and the 16.75 6 2.98% among all loci and 26.31 6 4.37% at only the polymorphic loci. The mean number of alleles at former USSR for propagation and release in the United the 52 loci was 2.01 6 1.97. Gene frequencies were States. Fifty to 75 adults were used to begin cultures in estimated in populations from Georgia, Virginia, Dela- USDA quarantine. In many cases, eggs from the found- ware, Rhode Island, Arkansas, Illinois, Iowa, and Or- ing beetles were transferred from quarantine and egon and differed significantly (P F 0.02) at 11 of 16 shipped to the cooperating agencies for propagation. loci. Three of 16 loci (Fbp, Est-1, Tre) were not in The number and life stages of H. axyridis released have Hardy–Weinberg equilibrium in all populations and varied. In 1981, releases were made in Maryland, are not included in the F statistics. Random mating Maine, and Washington of eggs, larvae, or adults from was indicated within populations (FIS 520.005 6 0.014) cultures established from Japanese and Russian beetles. but not among populations (FST 5 0.025 6 0.005). Ac- More releases were made in Maryland and Washington cording to Wright’s island model, FST estimates the in 1982. Coulson (1992), Coulson et al. (1988), and average number of reproducing immigrants to be Tedders and Schaefer (1994) should be consulted for a equivalent to ca. 10 beetles per population per genera- more comprehensive list of releases. No H. axyridis tion. Thus there was a measure of genetic differentia- were detected, despite repeated sampling, after the tion caused by drift, but this differentiation was small with respect to the large geographical distances among foregoing releases were made. In 1979, 32 cultured H. the sampled populations. The genetic data suggest axyridis were released in Louisiana, and in 1980, there that the founding North American population(s) was were eight releases totaling 3781 specimens in Missis- substantial. r 1997 Academic Press sippi; these releases were descendants of beetles col- KEY WORDS: Harmonia axyridis; isozyme variation; lected in Japan (Chapin and Brou, 1991). About 1000 breeding structure; gene flow; colonizing species. cultured H. axyridis were released in Connecticut pine forests in 1985 and 1986 (McClure, 1987), and nearly 87,000 larvae were release in Byron, Georgia, pecan INTRODUCTION orchards from 1978 to 1981 (Tedders and Schaefer, 1994). Intensive sampling from 1981 to 1991 failed to Harmonia axyridis (Pallas) is native to eastern Asia show any postrelease recoveries of H. axyridis in Geor- (Chapin and Brou, 1991) and has attracted attention gia (Tedders and Schaefer, 1994). from geneticists because its color and spotting patterns The first detections of H. axyridis in North America are among the most variable of the Coccinellidae were made in 1988 in Louisiana, 1990 in Mississippi, (Dobzhansky, 1933; Ayala, 1978). H. axyridis is a preda- and 1991 in Georgia (Tedders and Schaefer, 1994), tor of aphids, but it also feeds on scales and coleopteran hundreds of kilometers from the sites where they had and lepidopteran eggs (Hodek, 1973); this ladybeetle is been released 8 to 10 years earlier. These reports typically arboreal, but it has been found often in suggest a sudden appearance and explosive increase in 207 1049-9644/97 $25.00 Copyright r 1997 by Academic Press All rights of reproduction in any form reserved. 208 KRAFSUR ET AL. population density together with a rapid spread of the Three buffer systems were used: Tris–borate EDTA, ladybeetle. Tedders and Schaefer (1994) suggested that pH 9 (TBE), NAM-citrate, 6.5 (Clayton and Tretiak, two independent populations of H. axyridis were estab- 1972), and, for cationic proteins, the Mops system lished, one in Louisiana and the other in Georgia. In (Thomas and Hodes, 1981). The TBE electrode and gel contrast, Day et al. (1994) argued that H. axyridis buffer consisted of 81 mM Tris, 20 mM borate, and 1.5 establishment in the United States likely resulted from mM disodium EDTA. The NAM-citrate gel buffer was accidental introductions occurring at seaports. We re- 2.7 mM citrate, and the electrode buffer was 5.35 mM cord here the abrupt appearance of H. axyridis in citrate, both adjusted to pH 6.5 with N-(3-aminopropyl)- Arkansas (1992), Iowa (1994), Illinois (1994), and Or- morpholine. The Mops system included 3-(N-morpho- egon (1993). This is the second recent instance of a lino)propanesulfonic acid (Mops) titrated with KOH to surprising, explosive colonization of North America by pH 6.8. ladybeetles. The first such was the exotic Coccinella Staining recipes were as prescribed by Murphy et al. septempunctata L., which spread rapidly over the United (1990), and agar overlays were used for all stains States after first being detected in 1973, despite exten- requiring coupling enzymes. sive earlier releases of cultured beetles (Schaefer et al., 1987). Genetic Statistics The rapid spread of H. axyridis raises interesting and important questions about the roles of selection and Gene diversity (the expected frequency of heterozy- gotes under Hardy–Weinberg assumptions) at a locus genetic drift among populations. What effect has the 2 expansion in geographic range had on genetic differen- was estimated as he 5 1 2 op i , where p is the frequency tiation? If the distribution of H.
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