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Characterization of Two Non-Native Invasive Bark Beetles, Scolytus Schevyrewi and Scolytus Multistriatus (Coleoptera: Curculionidae: Scolytinae) Patricia L

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Characterization of Two Non-Native Invasive Bark Beetles, Scolytus Schevyrewi and Scolytus Multistriatus (Coleoptera: Curculionidae: Scolytinae) Patricia L 527 Characterization of two non-native invasive bark beetles, Scolytus schevyrewi and Scolytus multistriatus (Coleoptera: Curculionidae: Scolytinae) Patricia L. Johnson 1 United States Department of Agriculture, Forest Service, Wallowa Valley Office, Box A, 88401 Highway 82, Enterprise, Oregon 97828, United States of America Jane L. Hayes United States Department of Agriculture, Forest Service, Pacific Northwest Forestry and Range Sciences Laboratory, 1401 Gekeler Lane, La Grande, Oregon 97850, United States of America John Rinehart Eastern Oregon University, One University Boulevard, La Grande, Oregon 97850, United States of America Walter S. Sheppard Department of Entomology, FSHN Building 166, Washington State University, Pullman, Washington 99164-6382, United States of America Steven E.· Smith School of Natural Resources, University of Arizona, 325 Biological Sciences East, 1311 East 4th Street, Tucson, Arizona 85721, United States of America Abstract-Scolytus schevyrewi Semenov, the banded elm bark beetle, and S. multistrlatus Marsham, the smaller European elm bark beetle, are morphologically similar. Reliance on adult ex- ternal morphological characters for identification can be problematic because of wide within- species variability and the need for good-quality specimens. The inability to identify developmental stages can also hamper early-detection programs. Using two character identification systems, geni- talic (aedeagus) morphology, and DNA markers (random amplified polymorphic DNA polymerase chain reaction (RAPD-PCR)) to distinguish S. schevyrewi from S. multistriatus, we examined spec- imens from geographically distinct populations of both species collected from infested host trees or semiochemical-baited funnel traps. We found that aedeagus morphology can be used to identify the two species, The use of two oligonucleotide primers in the RAPD-PCR analysis yielded distinct DNA banding patterns for the two species. Species identification using RAPD-PCR analysis was validated by a blind test and used to make species identitications of larval specimens. These tools improve the ability to differentiate between S. schevyrewi and S. multistriatus at immature and adult stages, and could be developed and used for other scolytines as well. Resume-Scolytus schevyrewi Semenov, le scolyte asiatique de l'orme, et S. multistriatus Mars- ham, Ie petit scolyte europeen de l'orme, se ressemblent morphologiquement. L'utilisation des seuls caracteres externes des adultes pourI'Identification pose un probleme, car il y a un fort de- gre de variabilite au sein de chacune des especes et il est necessaire d' obtenir des specimens de bonne qualite. L'impossibilite d'identifier les stades immatures peut aussi nuire aux programmes de detection hative. En utilisant deux systemes de caracteres diagnostiques, la morphologie geni- tale (edeage) et des marqueurs ADN (ADN polymorphe amplifie aleatoirement en chaine par po- lymerase (RAPD-PCR)), pour distinguer S. schevyrewi et S. multistriatus, nous avons examine des specimens proven ant de populations geographiquemenr distinctes des deux especes et recol- tes sur des arbres hotes infestes ou dans des pieges appates de produits semiochimiques. Nous Received 13 May 2007. Accepted 6 May 2008. Corresponding author (e-mail: [email protected]). Can. Entomol. 140: 527-538 (2008) © 2008 Entomological Society of Canada 528 Can. Entomol. Vol. 140, 2008 trouvons que la morphologie de I'edcage pennet d'identifier les deux especes. L'utilisation de deux sondes doligonucleotides dans I'analyse RAPD-PCR produit des patrons de bandes ADN distincts chez les deux especes. Nous avons valide I'identification par l'analyse RAPD-PCR dans un test aveugle; nous I'avons aussi appliquee a I'identification specifique de larves. Ces outils permettent de mieux differencier S. schevyrewi de S. multistriatus aux stades immatures et adulte; il serait possible d'en mettre au point et d'en utiliser de semblables pour d'autres scolyti- nes. [Traduit par la Redaction] Introduction in North America in 2003 even though it had been collected from Denver, Colorado in 1994 Scolytus schevyrewi Semenov (Coleoptera: (Negron et al. 2005) and again from New Mex- Curculionidae: Scolytinae), the banded elm ico in 1997 (LaBonte et al. 2003). By 2004, es- bark beetle, recently introduced into the United tablished populations of S. schevyrewi were States of America, is morphologically similar to found in 21 states across the United States of another non-native invasive species that is well America (Negron et al. 2005). Harris (2004) established in North America, Scolytus found that up to 96% of S.· schevyrewi collected multistriatus (Marsham), the smaller European from American elm bolts were infected with elm bark beetle (Fig. 1). The two species utilize Dutch elm disease. Scolytus schevyrewi is a po- the same preferred hosts (species of Ulmus L. tential vector of Dutch elm disease (Seybold and (Ulmaceae)) and have been found in the same Lee 2004). The similar morphology and preferred host tree (Negron et al. 2005; P.L. Johnson, un- hosts of S. multistriatus and S. schevyrewi are the published data). Scolytus schevyrewi has been most likely reasons why S. schevyrewi was not known to attack other hosts such as species of identified until several years after it became es- Salix L. and Populus L. (Salicaceae), Prunus L. tablished in the United States of America (Liu (Rosaceae), Elaeagnus L. (Elaeagnaceae), Alnus and Haack 2003). Mill. (Betulaceae), and Quercus L. (Fagaceae) The negative impacts of non-native invasive (Wang 1992; Allen and Humble 2002; North scolytines such as S. multistriatus and American Plant Protection Organization (July S. scolytus F. (Weber 1990; Haack and Cavey 2003)). Scolytus multistriatus has attacked or- 2000; Allen and Humble 2002) and the estab- namental cypresses (Cupressus L. (Cupres- lishment of S. schevyrewi years before its detec- saceae)) in Australia (Neuman 1987). The tion underscore the need to develop additional known morphological differences between tools to distinguish between these species in. S. schevyrewi and S. multistriatus include the particular and other morphologically similar general size, shape, and color of the adults, and non-native scolytines at all life stages (Haack the size, shape, and location of the apical spine 2006) in general. The purpose of this study was on the underside of the abdomen (LaBonte et al. to examine alternative techniques and develop 2003). tools to characterize S. schevyrewi that could be Native to western Europe, the Middle East, used to distinguish it more easily from and northern Africa (Bellows et al. 1998), S. multistriatus and other scolytines, particu- S. multistriatus was first recorded in North larly at early life stages and in different condi- America in Boston, Massachusetts, in 1909 tions. The study included morphometric (Chapman 1910) and subsequently spread west quantification of the male aedeagus (external across the United States of America and into genitalia) and examination of diagnostic molec- southern Canada (Baker 1972). Scolytus ular markers using random amplified polymor- multistriatus is a primary vector of the Dutch phic DNA polymerase chain reaction (RAPD- elm disease pathogen Ophiostoma novo-ulmi PCR). Male genitalic morphology has been Brasier in the United States of America and used qualitatively to identify bark beetle species Canada (Baker 1972; Furniss and Carolin (e.g., Sharp and Muir 1912; Cerezke 1964; Vite 1977). The native range of S. schevyrewi in- et al. 1974). Molecular genetic markers identi- cludes Asian Russia, Mongolia, Turkmenistan, fied through RAPD-PCR have also been used to Uzbekistan, Tajikistan, Kazakhstan, southern differentiate variation within and between spe- Kyrgyzstan, Korea, and most of China (Wang cies, including non-native, invasive insects. For 1992). Scolytus schevyrewi was first identified example, the genetic relatedness of populations © 2008 Entomological Society of Canada Johnson et al. 529 and Utah, and S. multistriatus from Kansas, Maryland, and Oregon. Specimens of each spe- cies were reared from American elm (Ulmus americana (L.) (Ulmaceae)) limbs or collected from semiochemical-baited Lindgren funnel traps. Identification of all specimens was con- firmed by experts using current external mor- phological criteria (La Bonte et al. 2003). Samples used for morphometries were from all five states, with five populations (n = 10 per population) for each species. Samples of. S. schevyrewi were reared from elm limbs ob- tained from Lakewood (2004) and Golden (2005), Colorado (CO1 (39°42'N, 105°06'W) and C02 (39°78'N, 105°22'W)), and La Grande (2005), Oregon (OR3 (45°20'N, 118°06'W)), and trap-collected from Salt Lake City (2005) and Tremonton (2006), Utah (UT1 (40o45'N, 111o52'W) and UT2 (41°73'N, 112° 19'W)). Samples of S. multistriatus populations were obtained from trap collections in 2005 and 2006 from Crawford County, Kansas (KSI and KS2 (37°30'N, 94°51'W)); Prince George County (2006), Maryland (MD (38°84'N, 76°83'W)); and La Grande, Oregon (OR1), plus a popula- of the non-native bark beetle Tomicus piniperda tion reared from elm limbs from La Grande (L.) (Curculionidae: Scolytinae) was traced us- (2005), Oregon (OR3). Scolytus schevyrewi ing RAPD-PCR by Carter et al. (1996), who used for analysis of molecular genetics were determined that United States populations re- collected from Lakewood, Colorado (COl), La sulted from multiple introductions. Similarly, Grande, Oregon (OR3), and Salt Lake
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