Introduced Leaf Beetles of the Maritime Provinces, 3: the Viburnum Leaf Beetle, Pyrrhalta Viburni (Paykull) (Coleoptera: Chrysomelidae)

Home , Galerucinae, Longitarsus luridus, Neogalerucella, Sphaeroderma testaceum, Viburnum acerifolium, Viburnum leaf beetle

PROC. ENTOMOL. SOC. WASH. 109(2), 2007, pp. 454–462

INTRODUCED LEAF BEETLES OF THE MARITIME PROVINCES, 3: THE VIBURNUM LEAF BEETLE, PYRRHALTA VIBURNI (PAYKULL) (COLEOPTERA: CHRYSOMELIDAE)

CHRISTOPHER G. MAJKA AND LAURENT LESAGE

(CGM) Nova Scotia Museum of Natural History, 1747 Summer Street, Halifax, Nova Scotia, Canada B3H 3A6 (e-mail: [email protected]); (LL) Agriculture and Agri-Food Canada, ECORC, K.W. Neatby Building, 960 Carling Avenue, Ottawa, Ontario, Canada K1A 0C6

Abstract.—Pyrrhalta viburni (Paykull), the viburnum leaf beetle, is an introduced Palearctic leaf beetle established in northeastern and northwestern North America. It was first reported in Ontario in 1947 but did not create problems until 1978 when destruction of European varieties of viburnums was observed. It has subsequently been found in a number of Canadian provinces and northeastern United States. It is herein newly reported from Prince Edward Island and Newfoundland. In Nova Scotia, several localities are reported for the first time, while specimens collected in Annapolis Royal, in 1924, establish a new timeline for its occurrence on the continent. Two possible scenarios with respect to the origin and dispersal of P. viburni in the Maritimes are proposed and discussed. The systematic history of the genus is briefly recounted and the diagnostics of the genus and species are given. The occurrence of P. viburni on native and exotic species in the plant genus Viburnum is examined. and concerns are expressed with respect to its impact on native and cultivated varieties. Key Words: Coleoptera, Chrysomelidae, Galerucinae, Pyrrhalta viburni, viburnum leaf beetle, Canada, Maritime Provinces, introduced species

Pyrrhalta viburni (Paykull) is an acci- Pyrrhalta viburni was first reported on dentally introduced Palearctic leaf beetle, the North American continent from which is now established in northeastern specimens collected in Fonthill in the and northwestern North America. In the Niagara Peninsula of Ontario in 1947 Old World, it is found throughout much (Sheppard 1955). In 1978, Becker (1979) of Europe (including Ireland and Great found the species established on the Britain) north to Fennoscandia and east southern side of the Ottawa River in across Russian Karelia to the Urals, Ottawa (Ontario), and on the northern Orenburg, and Kazan. The southern side in Hull (Que´bec). Old scars on twigs limit of its distribution ranges from the of infested bushes indicated that the Pyrenees east through Liguria and the beetle had been present for several years southern Carpathians to the Black Sea, before the infestations were noticed. In Crimea, Georgia, and Dagestan in the 1993, it was reported in Halifax and Caucasus (Warchalowski 1994, Borowiec Dartmouth, Nova Scotia (Wheeler and 2005) Hoebeke 1994), and in 1994 it was found VOLUME 109, NUMBER 2 455 in Maine (Weston and Hoebeke 2003). It Hoebeke 2003, Weston and Diaz 2005). was recorded in New York (1996), Small and Catling (2005: 50) wrote that, Vermont (2000), Pennsylvania (2000), ‘‘Biologists have speculated that certain and Ohio (2002) by Weston and Hoe- species of Viburnum may become extir- beke (2003). In 2001, it was recorded in pated over parts of their range. The British Columbia (Gillespie 2001). More situation has been described as verging recently it has been reported in Connecti- on ecological disaster.’’ cut (2004), Massachusetts (2004), New In this context, we examined the Hampshire (2005) and Vermont (2004) published records of this species from (Anonymous 2005). Weston and Diaz the Maritime Provinces of Canada. We (2005) predicted that ‘‘given its broad also studied recent and historical speci- distribution in Europe, it seems likely mens housed in various collections (see that this pest will become widely distrib- below). Our survey reveals that P. viburni uted in the United States, given the is more broadly distributed in this region abundance of suitable native host plants than hitherto known, and it has been such as Viburnum dentatum and V. present for a significantly longer period opulus variety americanum throughout than previously reported. the eastern and northern halves, respec- tively, of the United States.’’ SYSTEMATICS Pyrrhalta viburni has been recorded from a large variety of hosts whose Pyrrhalta has a complicated taxonom- susceptibility to the beetle varies consid- ic history. De Joannis (1866) originally erably. Based on field observations, created it as a subgenus of Galeruca Weston and Desurmont (2002) reported Geoffroy, 1792. In the first edition of the that V. trilobum Marshall, V. opulus L., Coleopterorum Catalogus, Weise (1924) V. sargentii Koehne, V. rafinesquianum considered Pyrrhalta as monotypic, in- Schultes, V. recognitum Fernald, and V. cluding only P. viburni. Wilcox (1965) dentatum L. were the most susceptible enlarged the genus considerably by in- hosts. In laboratory trials, they found V. cluding the subgenera Galerucella trilobum, V. sargentii, and V. prunifolium Crotch, Neogalerucella Chuˆjoˆ, Xantho- L. susceptible in varying degrees to larval galeruca Laboissie`rre, and Tricholoch- feeding, while V. wrightii Miq., V. maea Laboissie`re. He maintained rough- plicatum (Thunb.), V. rhytidophylloides ly the same conception in the second J. Sur., V. carlesii Hemsl., and V. edition of the Coleopterorum Catalogus sieboldii Miq were resistant. Adults (Wilcox 1971). Ten years later, however, additionally fed to a substantial degree he included in Pyrrhalta only the Pale- on V. wrightii, V. rhytidophylloides, and arctic species and elevated to generic V. carlesii. Clark et al. (2004) addition- level the subgenera of his first contribu- ally record P. viburni from V. acerifolium tion (Seeno and Wilcox 1982). LeSage L., V. dilatatum Thnb., V. lantana L., V. (1991) adopted these changes but lentago L., V. x pragense Decker, and V. Downie and Arnett (1996) continued to tinus L., although they did not specify use the broad conception of Pyrrhalta. susceptibility nor larval versus adult Recently Riley et al. (2002) largely consumption for these species. Concern followed the generic arrangement of has been expressed by a number of Seeno and Wilcox (1982), with only authors about the potential impact of slight modifications regarding Galeru- P. viburni on populations of Viburnum in cella Crotch and Neogalerucella Chuˆjoˆ. North America (Weston and Desurmont This new classification which was 2002, Weston et al. 2002, Weston and adopted for the Catalog of Leaf Beetles 456 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON of America North of Mexico (Riley et al. DHWC David H. Webster Collection, 2003) reaches a broad consensus among Kentville, Nova Scotia those interested in leaf beetles in North GSC Gary Selig Collection, Bridge- America. water, Nova Scotia NSAC Nova Scotia Agricultural Col- IDENTIFICATION lege, Bible Hill, Nova Scotia The genus Pyrrhalta can be identified NSMC Nova Scotia Museum Collec- using a combination of external char- tion, Halifax, Nova Scotia acters: antennomere 4 longer than 2, the NSNR Nova Scotia Department of front coxae closed and not separated by Natural Resources, Shubena- the prosternum, tibial spurs present on cadie, Nova Scotia hind tibiae in both sexes, the inner margin of epipleuron is distinct to apex, elytra without distinct vittae, and elytral LOCALITY RECORDS punctation fine. Pyrrhalta viburni has now been found At a specific level, P. viburni is in the four Maritime Provinces: New distinguished by a golden pubescence Brunswick, Nova Scotia, Prince Edward on the head, pronotum, and elytra. Island, and Newfoundland (Fig. 2). In Species in the genus Tricholochmaea total, 76 specimens were examined. Un- Laboissie`re are also pubescent, but the less otherwise indicated, specimens are hairs are never golden. On the pronotum adults. In addition to the locality records of P. viburni, the pair of dark markings obtained from the specimens examined reach the lateral margins, but not in by the authors, the list below includes Tricholochmaea. In addition, the prono- localities reported in the literature. tum is twice as wide as long in Pyrrhalta, NEW BRUNSWICK: Charlotte Co.: whereas it is only 1.5 as wide as long in St. Stephen, 1995–2001, R.E. Hoebeke Tricholochmaea. The elytral punctures and A.G. Wheeler, (Hoebeke and Wheel- are coarse in Tricholochmaea but fine in er 2003); Saint John Co.: Saint John, Pyrrhalta. There are no vittae on the 1995–2001, R.E. Hoebeke and A.G. elytra as in most specimens of Ophraella Wheeler, (Hoebeke and Wheeler 2003); Wilcox, although the shoulders are often York Co.: Fredericton, 1995–2001, R.E. darkened (Fig. 1). Hoebeke and A.G. Wheeler, (Hoebeke and Wheeler 2003). NEWFOUND- CONVENTIONS LAND: St. John’s, Long Pond, IX– Abbreviations of collections referred X.2001, D.J. Larson (4, CNC); St. to in the text are: John’s, 2002, D.J. Larson (2, CNC). NOVA SCOTIA: Annapolis Co.: Anna- ACNS Agriculture and Agri-Food polis Royal, 19.IX.1924, J.P. Spitall (1, Canada, Kentville, Nova Sco- ACNS); 1.X.1924, J.P. Spitall (4, ACNS); tia Halifax Co.: Halifax, 26.VI.1993, R.E. ACPE Agriculture and Agri-Food Hoebeke & A.G. Wheeler, (on) Vibur- Canada, Charlottetown, num opulus (1, CUIC); Dartmouth, Prince Edward Island 26.VI.1993, R.E. Hoebeke & A.G. Wheel- CGMC Christopher G. Majka Collec- er, (on) Viburnum opulus, (1, CUIC); tion, Halifax, Nova Scotia Point Pleasant Park: 9.IX.2001, C.G. CNC Canadian National Collection, Majka, swamp, low vegetation (1, Ottawa, Ontario CNC); ibid., 23.IX.1993, Barry Wright, CUIC Cornell University Insect Col- (on) Viburnum nudum (9, NSMC); ibid., lection, Ithaca, New York 25.IX.1993, Barry Wright, (on) Viburnum VOLUME 109, NUMBER 2 457

Fig. 1. Habitus photograph of Pyrrhalta viburni. 458 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Distribution of Pyrrhalta viburni in the Maritime Provinces. Closed circles indicate collection localities. Inset map: collection locality in Newfoundland. nudum (3, NSMC); ibid., 1.IX.2000, C.G. (2, ACPE); Charlottetown, 1995–2001, Majka, C.G. Majka, (on) Viburnum R.E. Hoebeke and A.G. Wheeler, (Hoe- nudum (1, CGMC); ibid., 10.VIII.2001, beke and Wheeler 2003); Cornwall, C.G. Majka, (on) Viburnum nudum (2, 28.VI.2005, defoliating snowballs, M.E. CGMC); ibid., 9.IX.2001, C.G. Majka, Smith (36 larvae, CNC). boggy area, deciduous vegetation (3, CGMC); ibid., 21.IX.2001, C.G. Majka, DISCUSSION deciduous forest (3, CGMC); Herring An examination of Fig. 2 reveals that Cove: 29.IX.2002, C.G. Majka, coastal P. viburni has now been recorded from barrens (2, CGMC); Hants Co.: Windsor, a number of widely distributed localities 28.VII.1999, J. Ogden (2, NSNR); Pictou in the southern portions of the Maritime Co.: New Glasgow, 22.VIII.2002, S. Provinces and in Newfoundland. On the Wallis (8, NSNR); Lyons Brook, mainland, it has not been recorded in 30.VIII.2002, E. Georgeson (1, NSNR); northern New Brunswick (although col- Salt Springs, 16.VIII.2006, C.G. Majka, lecting effort in this area has been slight) defoliating Viburnum opulus (20, CGMC); nor on Cape Breton Island. Of particular Lunenburg Co.: Bridgewater, 23.IX.2002, note and interest are specimens collected G.D. Selig, defoliating Viburnum opulus by J.P. Spitall (a researcher with the (2, GSC). PRINCE EDWARD IS- Dominion Entomological Station [1911– LAND: Queens Co.: Charlottetown, 52] in Annapolis Royal) in Annapolis 6.VIII.1997, M.E.M. Smith, on Viburnum Royal, NS in 1924. These specimens VOLUME 109, NUMBER 2 459 extend by 23 years the known timeline of Table 1. Early records (date first recorded) of this species in North America. The adventive Coleoptera from Annapolis Royal, NS. question thus arises as to the possible origin of this population. In this context Anobiidae two possibilities present themselves. Lasioderma serricorne Fabricius 1929 1) Annapolis Royal is the site of one Ptinus fur Linne´ 1912 of the earliest European settlements in Bostrichidae North America, established by French Lyctus linearis (Goeze) 1932 settlers in 1605. It was the military and Cantharidae administrative center of Acadia during Cantharis rufa Linne´ 1930 parts of 17th century, and between 1710 Carabidae Agonum muelleri Herbst 1909 and 1749 (until the establishment of Amara familiaris Duftschmid 1909 Halifax) was the capital of Nova Scotia Pterostichus melanrius (Illiger) 1909 (which then included much of the Clivinia fossor (Linne´) 1935 Maritime Provinces). During this peri- Cerambycidae od, there was an extensive trans-Atlan- Callidium violaceum Linne´ 1929 tic commerce between Annapolis Royal Chrysomelidae and (particularly) French seaports (Cal- Longitarsus luridus (Scopoli) 1924 nek 1897). Indirect evidence indicates Pyrrhalta viburni (Paykull) 1924 that this commerce had an effect on the Cryptophagidae fauna of the area. Of 161 species of Cryptophagus subfumatus Kraatz 1941 beetles which have been collected in the Curculionidae Annapolis Royal area (ACNS, CNC, Sitophilus granarius (Linne´) 1910 CGMC, DHWC, NSAC, NSMC), 47 Otiorhynchus ovatus (Linne´) 1932 species (29.2%) are introduced, Pale- Otiorhynchus sulcatus (Fabricius) 1928 arctic species, double the 14.8% overall Sitona flavenscens Gyllenhal 1928 ratio of introduced species in Nova Sitona hispidulus (Fabricius) 1928 Scotia (C. Majka, unpublished data). Sciaphilus asperatus (Bonsdorff 1910 Strophosoma melanogrammum (Forster) 1924 Many of these are synanthropic spe- Trachyphloeus asperatus (Beck) 1924 cies or those associated with agricul- Hypera zoilus (Scopoli) 1924 ture and/or horticulture. In many in- Dermestidae stances these specimens represent some Attagenus pellio Linne´ 1930 of the earliest known records of these Dermestes lardarius Linne´ 1909 species in the province (Table 1), in- Latridiidae dicating an early and extensive history Corticaria saginata Mannerheim 1924 of introductions to the area. Thus, it is Cortinicara gibbosa (Herbst) 1928 possible that P. viburni may have been Melyridae imported to the Annapolis Royal area Malachius aeneus Linne´ 1928 sometime during this early history of Oedemeridae settlement. Nacerdes melanura (Linne´) 1935 2) Annapolis Royal was also the site Scarabaeidae (from 1885 to , 1945) of the Annapolis Aphodius prodromus (Brahm) 1915 Royal Nurseries, the largest and most comprehensive horticultural nursery in Staphylinidae eastern Canada (Clarke 1927). This Tasgius ater (Gravenhorst) 1938 nursery imported horticultural stock Tenebrionidae from throughout the world, propagated Tenebrio molitor Linne´ 1913 it, and sold plants throughout the Maritime Provinces (A. Wilson, pers. 460 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON comm.). In the nursey’s catalogue as naturalized shrubs, and other species (Clarke 1927) three exotic Palearctic of Viburnum are grown in horticultural species of Viburnum were offered for sale contexts (Roland 1998, Hinds 1986). In (V. opulus L., V. tomentosum Shasta, and the Maritime Provinces P. viburni has V. plicatum Thunb.), suggesting the been recorded on V. opulus and V. possibility that P. viburni could have nudum, and V. acerifolium, V. dentatum, been imported in association with nurs- V. lanata, V. lentago, and V. recognitum ery stock of one of these species. are recorded hosts (Clark et al. 2004), Whether this early introduction to indicating that a substantial proportion Annapolis Royal was responsible for of species in this region could be affected the subsequent occurrence of P. viburni by this beetle. at other locations in the Maritime Thus far, the impact of P. viburni in Provinces is uncertain. this region has not been noted to be Pyrrhalta viburni is one of a substantial significant, although occasional signifi- number of introduced leaf beetles known cant defoliation of shrubs has been in the Maritime Provinces. In Nova reported (G. Selig pers. comm.). Weston Scotia, 25 of the 178 species of Chry- and Desurmont (2002) wrote that, ‘‘Vi- somelidae (14%) recorded in the prov- burnum leaf beetle has the potential to ince are introduced species, similar to the become a major landscape pest because 14.8% of the overall beetle fauna which of its ability to kill susceptible viburnums is introduced (C. Majka, unpublished if allowed to defoliate shrubs for several data). Chrysolina staphylea L. (Evans years in succession.’’ Young (2004) wrote 1899), Cassida rubiginosa Mu¨ller (Brown that, ‘‘Heavy infestations … could de- 1940), Longitarsus luridus (Goeze) foliate shrubs, cause dieback, and even- (Brown 1967, LeSage 1988), L. ferrugi- tually kill plants. Costs to homeowners, neus (Foudras) (LeSage 1988), L. praten- parks, arboretums, municipalities, and sis (Panzer) (LeSage 1988), Sphaero- nurseries to manage heavy infestations derma testaceum (Fabricius) (Hoebeke … and to replace killed plants could be and Wheeler 2003, Majka and LeSage high. Quarantines imposed … would 2006) and Oulema melanopus (Linne´) represent an economic burden to the (LeSage et al. 2007) are all introduced many growers who export nursery stock Palearctic chrysomelids whose presence ….’’ Given such concerns, ongoing in the region has been discussed in monitoring of populations of P. viburni publications. The impact of this sub- in the region would not be imprudent. stantial number of introduced species on the native fauna and the environment in ACKNOWLEDGMENTS general has been rather little investigat- ed. Species such as Lilioceris lilii (Sco- Thanks are extended to Christine poli), Crioceris duodecimpunctata (L.), C. Noronha and Mary Smith (Agriculture asparagi (L.), and Oulema melanopus are and Agri-Food Canada, Charlottetown, all potentially serious pests. PEI), Susan Westby and Cory Sheffield In the Maritime Provinces, the native (Agriculture and Agri-Food Canada, Viburnum alnifolium Marsh., V. edule Kentville, NS), Gary Selig, Jean-Pierre (Michx.) Raf., and V. nudum L. are LeBlanc (Nova Scotia Agricultural Col- common, generally distributed species. lege), and Jeffrey Ogden (Nova Scotia Viburnum acerifolium, V. lentago, and V. Department of Natural Resources) for recognitum have been found in New making specimens and records available, Brunswick. Exotic species such as V. to Klaus Bolte for executing the habitus dentatum, V. lantana. and V. opulus grow photograph, and to Alex Wilson (Nova VOLUME 109, NUMBER 2 461

Scotia Museum) for his assistance. Evans, J. D. 1899. List of Coleoptera from Halifax. Shawn Clark (Brigham Young Universi- The Canadian Entomologist 31: 32–321. Geoffroy, E. L. 1792. Histoire abre´ge´e des insectes ty) read an earlier draft of this manu- qui se trouvent aux environs de Paris dans script and made many constructive laquelle les animaux sont range´s suivant un ordre suggestions. C. G. Majka thanks Calum me´thodique. Durand, Paris, Vol. 1, 523 pp. Ewing and Andrew Hebda (Nova Scotia Gillespie, D. R. 2001. Arthropod introductions Museum) for continuing support and into British Columbia – the past 50 years. encouragement. This work has been Journal of the Entomological Society of British Columbia 98: 91–97. assisted by a research grant from the Hinds, H. R. 1986. Flora of New Brunswick. Nova Scotia Museum of Natural Primrose Press, Fredericton, N.B. 460 pp. History. Hoebeke, E. R. and A. G. Wheeler, Jr. 2003. Sphaeroderma testaceum (F.) (Coleoptera: Chrysomelidae), a Palearctic flea beetle new to LITERATURE CITED North America. Proceedings of the Entomo- Anonymous. 2005. Pest Tracker Invasive Insect: logical Society of Washington 105: 990–994. Viburnum Leaf Beetle, Pyrrhalta viburni. Kerzhner, I. M. 1991. Histoire abre´ge´e des insectes Available from http://ceris.purdue.edu/napis/ qui se trouvent aux environs de Paris (Geof- pests/vlb/ (accessed 12 May 2006). froy, 1792): proposed conservation of some Becker, E. C. 1979. Pyrrhalta viburni (Coleoptera: generic names (Crustacea and Insecta). Bulle- Chrysomelidae), a Eurasian pest of Viburnum tin of Zoological Nomenclature 48: 107–134. recently established in Canada. The Canadian LeSage, L. 1988. Notes on European Longitarsus Entomologist 111: 417–419. species introduced in North America (Co- Borowiec, L. 2005. Chrysomelidae: The Leaf leoptera: Chrysomelidae: Alticinae). The Ca- Beetles of Europe and the Mediterranean nadian Entomologist 120: 1133–1145. Subregion (Checklist and Iconography). Uni- ———. 1991. Family Chrysomelidae leaf beetles. versity of Wroclaw, Wroclaw, Poland, Avail- pp. 301–322. In Y. Bousquet ed. Checklist of able from http://culex.biol.uni.wroc.pl/cassidae/ beetles of Canada and Alaska. Research Branch, European%20Chrysomelidae/index.htm (ac- Agriculture Canada, Publication 1861/E, 430 pp. cessed 12 May 2006). LeSage, L., E. J. Dobesberger, and C. G. Majka. Brown, W. J. 1940. Notes on the American 2007. Introduced leaf beetles of the Maritime distribution of some species of Coleoptera Provinces, 2: The cereal leaf beetle Oulema common to the European and North American melanopus (Linnaeus) (Coleoptera: Chrysome- continents. The Canadian Entomologist 72: lidae). Proceedings of the Entomological Soci- 65–78. ety of Washington 109: 286–294. ———. 1967. Notes on the extralimital distribution Majka, C. G. and L. LeSage. 2006. Introduced leaf of some species of Coleoptera. The Canadian beetles of the Maritime Provinces, 1: Sphaer- Entomologist 99: 85–93. oderma testaceum (F.) (Coleoptera: Chrysome- Calnek, W. A. 1897. History of the County of lidae). Proceedings of the Entomological Soci- Annapolis. Mika Studio, Bellville, Ontario, ety of Washington 108: 243–247. (reprint). 660 pp. Riley, E. G., S. M. Clark, R. W. Flowers, and A. J. Clarke, E. R. 1927. Catalogue, spring 1927: Gilbert. 2002. Chrysomelidae Latreille 1802. Annapolis Royal Nurseries. Annapolis, Nova pp. 617–691. In R. H. Arnett, Jr., M. C. Scotia. 51 pp. Thomas, P. E. Skelley, and J. H. Frank, (eds). Clark, S. M., D. G. LeDoux, T. N. Seeno, E. G. American Beetles, Volume 2: Polyphaga: Scar- Riley, A. J. Gilbert, and J. M. Sullivan. 2004. abaeoidea through Curculionoidea. CRC Host plants of leaf beetle species occurring in Press, Boca Raton, USA. the United States and Canada (Coleoptera: Riley, E. G., S. M. Clark, and T. N. Seeno. 2003. Megalopodidae, Orsodacnidae and Chrysome- Catalog of the leaf beetles of America north of lidae, excluding Bruchinae). Coleopterists So- Mexico. Coleopterists Society Special Publica- ciety Special Publication 2. 476 pp. tion No. 1. 290 pp. de Joannis, M. L. 1866. Monographie des Gale´r- Roland, A. E. 1998. Roland’s Flora of Nova ucides d’Europe, du Nord de l’Afrique et de Scotia. Revised by M. Zinck. Nova Scotia l’Asie. L’Abeille 3: 1–168. Museum, Halifax, Nova Scotia. 1297 pp. Downie, N. M. and R. H. Arnett, Jr. 1996. The Seeno, T. N. and J. A. Wilcox. 1982. Leaf beetles Beetles of Northeastern North America. Sand- genera (Coleoptera: Chrysomelidae). Entomo- hill Crane Press, Gainesville, Florida. 1721 pp. graphy 1: 1–221. 462 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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