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Phoretic Mites and Nematode Associates of Scolytus Multistriatus and Scolytus Pygmaeus (Coleoptera: Scolytidae) in Austria

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Phoretic Mites and Nematode Associates of Scolytus Multistriatus and Scolytus Pygmaeus (Coleoptera: Scolytidae) in Austria Agricultural and Forest Entomology (2005) 7, 169–177 Phoretic mites and nematode associates of Scolytus multistriatus and Scolytus pygmaeus (Coleoptera: Scolytidae) in Austria John C. Moser, Heino Konrad*, Thomas Kirisits* and Lynn K. Carta† USDA Forest Service, Southern Research Station, 2500 Shreveport Highway, Pineville, LA 71360, U.S.A., *Institute of Forest Entomology, Forest Pathology, and Forest Protection (IFFF), Department of Forest and Soil Sciences, BOKU – University of Natural Resources and Applied Life Sciences, Vienna, Hasenauerstrasse 38, A-1190 Vienna, Austria and yUSDA-ARS, Nematology Laboratory, Beltsville, MD 20705, U.S.A. Abstract 1 The species assemblages and abundance of phoretic mites and nematodes associated with the elm bark beetles, Scolytus multistriatus and Scolytus pygmaeus, were studied in Austria. 2 A total of 3922 individual mites were recorded from 144 adults of S. multi- striatus and 178 adults of S. pygmaeus. The species spectrum was identical and the relative abundance of mites was very similar for both species of scolytids. Nine mite species, Pyemotes scolyti, Pseudotarsonemoides eccoptogasteri, Trichouropoda bipilis, Tarsonemus crassus, Proctolaelaps eccoptogasteris, Proctolaelaps scolyti, Chelacheles michalskii,nr.Eueremaeussp. and Elattoma sp. were detected. Two of the nine species, nr. Eueremaeus sp. and Elattoma sp., are documented here as new associates of Scolytus spp. 3 Pyemotes scolyti was the most frequent mite species, and Ps. eccoptogasteri and T. bipilis were relatively common, whereas the other mites occurred occasion- ally or were rare. 4 The trophic roles of most of the mites associated with S. multistriatus and S. pygmaeus are poorly known, but they may include fungivores, parasitoids of bark beetle broods, predators of bark beetle broods and/or mites and/or nematodes. 5 Besides phoretic mites, two nematode associates were seen on the investigated insects. A species of Cryptaphelenchus occurred under the elytra of both scolytid species, whereas the adults of a Neoparasitylenchus sp. were present inside abdomens of S. multistriatus, but absent from S. pygmaeus. Keywords Dutch elm disease, elm, nematode, Ophiostoma novo-ulmi, phoresy, Scolytidae, mite, Scolytus multistriatus, Scolytus pygmaeus, Ulmus minor. Introduction and ssp. novo-ulmi Brasier) of Ophiostoma novo-ulmi Brasier and hybrids between them (Brasier, 1991, 2000; Subsequent to the appearance of Dutch elm disease (DED) Brasier & Kirk, 2001; Konrad et al., 2002). Ophiostoma at the beginning of the 20th century, the elms (Ulmus spp.) ulmi and O. novo-ulmi are amongst the best known exam- in Europe, North America and parts of Asia have been ples of tree pathogens that are vectored by insects, in this seriously affected by this destructive vascular wilt disease particular case by elm bark beetles in the genus Scolytus that is caused by two closely related species of Ophiostoma Geoffroy and in North America also by Hylurgopinus (Ascomycota). Although the first wave of DED was caused rufipes (Eichhoff) (Lanier & Peacock, 1981; Webber & by Ophiostoma ulmi (Buism.) Nannf., the second and still Brasier, 1984; Webber & Gibbs, 1989; Webber, 1990, 2000). ongoing pandemic from the 1940s onwards has been trig- Scolytus multistriatus (Marsham) and Scolytus pygmaeus gered by the two subspecies (ssp. americana Brasier & Kirk (Fabricius) are among the guild of eight Scolytus species infesting elm trees in Europe (Pfeffer, 1995). In the 20th century, S. multistriatus has also been introduced and Correspondence: John C. Moser. Tel.: þ1 318 4737258; fax: established in North America (Lanier & Peacock, 1981), þ1 318 4737222; e-mail: [email protected] Australia (Lanier & Peacock, 1981) and New Zealand # 2005 The Royal Entomological Society 170 J. C. Moser et al. (Gadgil et al., 2000). Both Scolytus species are regarded as during their saprotrophic phase in and around bark beetle secondary bark beetles that breed in stressed elm trees, elm galleries in the bark of elm trees, thereby promoting sexual logs and branches (Postner, 1974). The most important reproduction and recombination (Brasier, 1978). impact of S. multistriatus and S. pygmaeus is related to As an initial step in the study of the trophic relationships their role as vectors of the DED pathogens from diseased between Scolytus spp., phoretic mites, O. novo-ulmi and to healthy trees during maturation feeding on the bark of their elm hosts, we present the first comprehensive list of twigs and in twig crotches in the crown of healthy elm trees mites phoretic on S. multistriatus and S. pygmaeus attack- (Lanier & Peacock, 1981; Webber & Brasier, 1984; Webber ing Ulmus spp. and provide data on their relative phoretic & Gibbs, 1989; Webber, 1990, 2000; Basset et al., 1992; abundance on S. multistriatus and S. pygmaeus. In add- Faccoli & Battisti, 1997). Scolytus species do not possess a ition, we report on two nematode associates of the two mycangium and, instead, transmit ascospores and conidia Scolytus species found on elm in Austria. of Ophiostoma novo-ulmi on their body surface (Lanier & Peacock, 1981; Webber & Brasier, 1984; Webber & Gibbs, 1989). Considering the importance of Scolytus spp. in the Materials and methods transmission of the DED pathogens, control measures On 17 May 2002, five stem sections, each approximately against the insects form an essential part of integrated man- 120 cm in length and 20 cm in diameter, were cut from a agement strategies against Dutch elm disease (Campana & European field elm, Ulmus minor Mill., near Gu¨ssing, Stipes, 1981; Gadgil et al., 2000). Burgenland, Austria, at 300 m a.s.l. This tree had been Eight species of phoretic mites, Chelacheles michalskii declining due to DED, and was subsequently attacked by Samsinak, Histiostoma ulmi Scheucher, Pseudotarsonemoides S. multistriatus and S. pygmaeus. Originally, we also eccoptogasteri Vitzthum, Proctolaelaps eccoptogasteris intended to include Scolytus scolytus Olivier for investiga- (Vitzthum), Pr. scolyti Evans, Pyemotes scolyti (Oudemans), tion of phoretic mites, but this scolytid had not infested the Tarsonemus crassus (Schaarschmidt), and Trichouropoda elm tree from which samples were collected. The stem sec- bipilis (Vitzthum) have been recorded in Europe phoretic on tions were placed in a laboratory rearing cage at S. multistriatus (Marsham) and/or S. pygmaeus (Fabricius), 20 Æ 1 C. Beetles emerged from 28 May to 30 June or from Scolytus galleries on Ulmus spp. (Vitzthum, 1920, 2002, and flew to the cage screens where they were collected 1921, 1923; Oudemans, 1936; Scheucher, 1957; Evans, 1958; on 10 dates, and placed in vials containing 70% alcohol. Schaarschmidt, 1959; Samsinak, 1962). Except for Pr. scolyti The two beetle species were separated and each placed in and Py. scolyti, a somewhat different guild of mite species separate vials, totalling 10 vials for each scolytid. The may be associated with S. multistriatus in North America. beetles were then transferred to lactophenol for clearing. Smiley & Moser (1974) described the following new Mites still attached to the beetles were tallied separately taxa: Heterotarsonemus milleri Smiley & Moser, Pseudo- from those that fell off the beetles into the lactophenol, tarsonemoides scolyti Smiley & Moser, Tarsonemus kennedyi and those that separated from the beetles as they were Smiley & Moser and Ununguitarsonemus peacocki Smiley & placed in alcohol after being picked off the cage screen Moser, all of which were either phoretic on the insects or (Tables 1 and 2). Voucher specimens (slides) of all mite found in galleries of S. multistriatus on Ulmus americana L. species detected in this study are maintained in the collec- To date, none of these four mites has been recorded in tions of the authors, and in other collections of mite tax- Europe where the insect is native. onomists mentioned in the acknowledgements. Although a great deal of literature has been devoted to Nematodes were fixed within the insects in 70% alcohol, the bark beetles attacking Ulmus and the pathogenic fungi transferred to lactophenol for clearing, mounted in Berlese that these beetles transmit, the literature is scant and frag- fluid (Humason, 1972) or glycerin, and measured with an mented concerning the mites known to be carried by these ocular micrometer. Specimens were viewed with differential scolytids. Except for Py. scolyti, little is known concerning interference optics, images were captured with Image- the biologies of the above mentioned mites, except for brief Pro PlusTM, version 4.1 (Silver Spring, Maryland), and accounts of their phoretic habits and/or gallery associa- images stitched together in Adobe Photo Shop, version tions. Studies on the phoretic mites of conifer bark beetles 7.0 (Adobe Systems Inc., San Jose, California). Nematode have revealed that these arthropods have a number of slides and beetles with aphelenchid nematodes in alcohol trophic roles in the insect galleries, including beetle para- were deposited in the USDA Nematode Collection, sitoids, insect and nematode predators, fungivores, and Nematology Laboratory, Beltsville, Maryland. omnivores (Kinn, 1983; Klepzig et al., 2001a, 2001b). Perhaps the most intriguing role that mites play on conifers is related to the transmission of tree pathogens, mycangial Results and discussion symbionts and fungal antagonists of bark beetles (Bridges & Moser, 1983; Moser et al., 1989b, 1995, 1997; Klepzig Phoretic mites and nematodes from Scolytus multi- et al., 2001a, 2001b). Although the mite
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