PROC. ENTOMOL. SOC. WASH. 110(1), 2008, pp. 79–86

INTRODUCED LEAF OF THE MARITIME PROVINCES, 4: STAPHYLAEA (LINNAEUS) (COLEOPTERA: CHRYSOMELIDAE)

CHRISTOPHER G. MAJKA AND LAURENT LESAGE

(CGM) Nova Scotia Museum, 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.—The origins and status of Chrysolina staphylaea (Linnaeus), first recorded in North America in Halifax, Nova Scotia, Canada in 1897, is reviewed. Additional records from Nova Scotia and Newfoundland are reported which establish that the species is more widely distributed than has hitherto been known. The status of the species on the continent is discussed, as are possible modes of introduction, dispersal, and potential host plants. Key Words: Chrysomelidae, Chrysolina staphylaea, Atlantic Canada, adventive species, introduced Coleoptera

The flightless Palearctic chrysomelid Knab (1911b: 184) wrote that at Chrysolina staphylaea (Linnaeus 1758) a meeting of the Entomological Society was first recorded in North America by of Washington, ‘‘Mr. Schwarz said that Evans (1899) from a specimen he col- this occurrence (i.e., of C. staphylaea) lected in June 1897 in Halifax, Nova was known by the early Massachusetts Scotia, Canada while attending the entomologists, but has been lost sight annual meeting of the Royal Society of of.’’ He was referring to Eugene A. Canada. Knab (1911a, b) subsequently Schwarz (1844–1928), a German ento- turned his attention to the species since mologist who came to the Museum of Evans (1899) remarked on differences Comparative Zoology in 1872, subse- between the specimen he collected and quently became one of the founders of those from Europe, and consequently the Cambridge Entomological Club, and there was some question as to its was then with U.S. Department of identity. Knab (1911a) examined three Agriculture in Washington (Matthews specimens collected by Joseph Perrin on 1974). There is, unfortunately, no further 18 June 1910 and 5 August 1910 on information provided, nor are there McNabs Island (located in Halifax Har- published records or specimens from bour) and confirmed the determination. Massachusetts so this account cannot Brown (1950) reported further specimens be further validated. from across the harbour in Dartmouth, Chrysolina staphylaea was subsequent- Nova Scotia, restricted to an area of ly reported from St John’s, Newfound- meadows that he estimated to be land by Brown (1962) (apparently from 25 square miles. specimens collected by Lindroth in 1949) 80 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON as well as Lunenburg and Liverpool in was eaten and used for oviposition. The Nova Scotia. Brown (1962) stated that larvae described by Marshall (1979) were the species is restricted to very small reared on Achillea sp. and Ranunculus areas in the vicinity of these seaports. repens L. The plants reported by Read Chantal (1972) then reported it in Le´vis, (1984) can be considered true hosts since near Que´bec City from specimens col- both adults and larvae were found on lected in 1965. Since the species is flight- Plantago lanceolata L., P. maritima L., less, Brown (1962) and Chantal (1972) and Aster tripolium L. Very little is concluded that the distribution indicated known with respect to the biology of several independent introductions. Chrysolina staphylaea in North America In the Old World, C. staphylaea has except that it prefers moist open habitats a boreo-alpine distribution which ranges and is primarily nocturnal. from Iceland and the Faeroes, east to In the following account, we present Ireland, Great Britain, and Fennoscan- additional records of C. staphylaea that dia, south to France, northern Spain, indicate that its distribution is greater and Italy, thence eastward across Siberia than previously supposed, in addition to south to central Turkey, the Caucasus, further considering its possible origins on Kazakhstan, and northwestern Mongo- the continent. lia. In the Far East it is found in Kamchatka, and the Kurile Islands IDENTIFICATION (Jolivet 1990), with an isolated popula- Chrysolina staphylaea is one of 17 tion on the island of Cheju-Do in Korea species in the genus Chrysolina (Jolivet 1975). Motschulsky, 1860 that have been re- Clark et al. (2004) listed a large corded in North America (Riley et al. number of plants on which Chrysolina 2003). However, only four are known to staphylaea had been recorded including occur in Eastern Canada, three of these species in the Asteraceae, Betulaceae, being introduced Palearctic species (Le- Clusiaceae, Lamiaceae, Lythraceae, Sage 1991). It is the only North Amer- Plantaginaceae, Ranunculaceae, and ican representative in the nominate sub- Scrophulariaceae. According to Jolivet genus, Chrysolina Motschulsky which (1990) several of these records are was characterized in detail by Bechyne´ probably in error, the true hosts being (1950). restricted to Lamiaceae, Plantaginaceae Chrysolina staphylaea is a large (6.2– and Ranunculaceae, with preferences for 8.1 mm long) and distinctive leaf Plantago spp. and Ranunculus spp. Later, that is readily identified by its entirely Jolivet (1991) reinterpreted these data reddish-brown body with paler appen- and stated that the Ranunculaceae are dages (Fig. 1). In the Maritime Pro- probably the first choice of C. staphylea, vinces, the native C. hudsonica Brown is others representing a secondary adapta- dark brown, whereas the introduced St. tion. John’s wort leaf beetles, C. hyperici Direct observation of the feeding of (Forster) and C. quadrigemina (Suffrian) the adults is difficult because they feed at are coppery, metallic green, or metallic dusk or at night (Jolivet 1990) or hide at blue. The pronotal sulci are strongly the base of plants (Read 1984). Drum- impressed in C. staphylaea and the elytra mond (1952) collected C. staphylaea, on are not striate, but some elytral punctu- Centaurea nigra L., Hypericum quadran- res are often arranged in more or less gulum L., Plantago lanceolata L., and regular series (Fig. 1). In the male, the Ranunculus repens L. in Askham Bog. In apex of the aedeagus is broadly rounded, a subsequent experiment only R. repens the angle of each being obtuse, and the VOLUME 110, NUMBER 1 81

Fig. 1. Habitus photograph of Chrysolina staphylaea. 82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

MUNC Memorial University of New- foundland collection, Edmon- ton, Alberta, Canada NSAC Nova Scotia Agricultural Col- lege, Bible Hill, Nova Scotia, Canada NSMC Nova Scotia Museum Collec- tion, Halifax, Nova Scotia, Canada USNM National Museum of Natural History, Smithsonian Institu- tion, Washington, DC, USA.

The number of specimens is indicated in parentheses together with the collec- tion coden.

RESULTS

Fig. 2. Aedeagus of Chrysolina staphylaea. Records of this species in Atlantic Canada are compiled below. The present distribution is based on records of 116 flagellum is slightly exposed (Fig. 2). The specimens. The following list also in- adults can be identified by employing the cludes records from the literature. keys in Brown (1962), Wilcox (1972), NEWFOUNDLAND: Burin Peninsu- Downie and Arnett (1996), and Riley et la, Burin, 12.IV.1996, (1, MUNC); Har- al. (2002). bour Grace, 11.VII.2004, E.R. Hoebeke Although Chrysolina staphylaea was and A.G. Wheeler, Chrysanthemum flow- described by Linnaeus in 1758, the egg ers, (1, CUC); Logy Bay, 11.VI.1984, D. and the pupa are still unknown and only Larson, (1, MUNC); Northern Bay, IX– the egg bursters of the first larval instar X.2000, (2, MUNC); Portugal Cove, (Cox 1988), and the claw of the mature 24.VI.1975, G.D. Fenwick, grassy hill- larva (Marshall 1979) have been illus- side, (1, MUNC); St. John’s, 4.VI.1949 trated. and 30.VIII.1949, C.W. Lindroth, (3, CNC); St. John’s, Brown (1962: 71); St. METHODS AND CONVENTIONS John’s, 27.VI.1965, W.J. Brown, (25, Codens (following Evenhuis, 2007) of CNC); St. John’s, 11.VII.2004, E.R. collections referred to in this study are: Hoebeke and A.G. Wheeler, (1, CUC); St. John’s, 1978–2001, (22, MUNC); Seal CBU Cape Breton University, Syd- Cove, 14.XI.1980, J. Sinois, (1, MUNC); ney, Nova Scotia, Canada Topsail, 2.VII.1965, W.J. Brown, (3, CNC Canadian National Collection CNC). NOVA SCOTIA: Cape Breton of , Arachnids, and Co.: Christmas Island, 10.V.1978, B. Nematodes, Ottawa, Ontario, Wright, (1, NSMC); Georges River, Canada 25.IV.1995, D.B. McCorquodale, (1, CUIC Cornell University Col- CBU); Georges River, 14.IV.1995, D.B. lection, Ithaca, New York, McCorquodale, (1, CBU); Fortress of USA Louisbourg, 16.VI.1999, Schrage and JOC Jeffrey Ogden Collection, Roach, woodpile, (1, CBU); Colchester Truro, Nova Scotia, Canada Co.; Shubenacadie, 21.IX.2000. J. Og- VOLUME 110, NUMBER 1 83 den, (1, JOC); Halifax Co.: Armdale, Cove, St. John’s, Topsail, Louisbourg, 26.IV.1943, D.C. Ferguson, (1, NSMC); Halifax, Dartmouth, Eastern Passage, Armdale, 8.V.1945, D.C. Ferguson, (1, Chester, Lunenburg, Mahone Bay, Li- NSMC); Cow Bay, 7.VI.1947 and verpool, and Lydgate) (Fig. 3). Many of 12.VI.1947, W.J. Brown, (8, CNC); these are seaports with a considerable Cow Bay, 18.VI.1961, C.J.S. Fox, (1, maritime history which could have pro- ACNS); Cow Bay, 18.VI.1947, W.J. vided introduction opportunities. Brown, (2, USNM); Dartmouth, Brown In Nova Scotia, there are collections (1962: 71); Dartmouth, 3.VI.1947, from Christmas Island (in the Bras d’Or 5.VI.1947, 9.VI.1947, W.J. Brown, (8, Lakes), Shubenacadie, and Caledonia CNC); Eastern Passage, 2.VI.1947 and that are a considerable distance from 22.VI.1947, W.J. Brown, (5, CNC); the seacoast and may indicate secondary, Halifax, June 1897, J.D. Evans, (Evans human-assisted transport. How many 1899: 321); Halifax, 4.VI.1915 and independent introductions the current 19.VI.1915, J. Perrin (2, CNC); Halifax, distribution represents is subject to 26.VIII.1945, D.C. Ferguson, (1, question. The species is now clustered NSMC); Halifax, 18.X.1985, C. Cooke, into four areas: a) along the south shore (1, NSMC); Halifax, no date indicated, of Nova Scotia; b) in the southern (1, NSMC); McNab’s Island, 18.VI.1910 portion of Cape Breton Island; c) on and 5.VIII.1910, J. Perrin, Knab (1911a: the Burin peninsula; and d) on the north 306); McNab’s Island, 21.VI.2001, J. portion of the Avalon peninsula (Fig. 1). Ogden, (1, JOC); McNab’s Island, Although being flightless would doubt- 18.VI.1911, 24.VI.1911, and 5.VIII.1911, less provide limitations to the dispersion no collector indicated, (5, USNM); of C. staphylaea, this in itself is not Lunenburg Co.: Chester, 16.VI.1966, B. a determinate of dispersal ability. For Wright, (1, NSMC); Lunenburg, Brown instance, in the Maritime Provinces the (1962: 71); Mahone Bay, 13.VI.1950, D. flightless, brachypterous carabids, Car- Eidt, (1, NSAC); Queens Co.: Caledonia, abus granulatus hibernicus Lindroth and 1.VI.1953, D.C. Ferguson, (1, NSMC); C. nemoralis Mu¨ller, have become very Liverpool, Brown (1962: 71); no locality widely distributed throughout the region given, 9.VIII.1906, W.P. Henderson (2, (Majka et al. 2006 and unpublished USNM); Shelburne Co.: Lydgate, data). 2.VI.1968 and 30.VI.1968, P. Doleman, Chrysolina staphylaea is a hydrophi- (2, NSMC). lous species of open habitats. Drum- The North American distribution of mond (1952) collected many specimens the species is given in Fig. 3. in Askham Bog. Brown (1962) stated that, near Dartmouth (Nova Scotia) and DISCUSSION southeastwards, it was restricted to In Nova Scotia and Newfoundland C. meadows. Chantal (1972) found many staphylaea is clearly well established and specimens under pieces of wood lying the above records indicate that it is more over moist soil. European observations widely distributed than has hitherto been would indicate that there is likely only known. Since it is flightless, Brown one generation per year in the Maritime (1962) and Chantal (1972) both conclud- Provinces. ed that the Canadian distribution repre- Within the families identified by Joli- sented several independent introduc- vet (1990) as principal host plants, there tions. Most collection sites in both are many representatives in Nova Scotia provinces are at coastal sites (i.e., Burin, including Ranunculus (ten species, six Harbour Grace, Northern Bay, Portugal adventive including R. cymbalaria Pursh, 84 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Distribution of Chrysolina staphylaea in Atlantic Canada.

R. flammula L., R. gmelini DC, R. Lindroth (1957) collected Achillea scleratus L., R. acris L., and R. bulbosus millefolium, Artemesia vulgaris L., Plan- L.) (Ranunculaceae), Mentha arvensis L., tago lanceolata L., P. major L., Ranun- M. aquatica L., M. spicata L., and M. culus repens L., and Ballota nigra L., – all lonigolia L., Lamium purpurem L., L. associated with C. staphylaea – from all amplexicaule L., Galeopsis tetrahit L., seven principal quarries in southern Prunella vulgaris L., (Lamiaceae), and England where dry-ballast destined for Plantago (six species, five adventive or Atlantic Canada originated, a possible Holarctic including P. maritima L., P. indication that C. staphylaea could have laneolata L., P. major L., and P. rugelli been introduced to North America via Decne, and P. psyllium L.) (Roland this mechanism. Brown (1940, 1950) and 1998). There are also 18 native species Lindroth (1957) both proposed that the of Aster (as per Read 1984) and Achillea transport of dry ballast (bulky rock, ptarmica L. and A. millefolium L. (as per sand, and soil) in trans-Atlantic shipping Marshall 1979). All this indicates that was responsible for the introduction of there are large number of possible hosts many species of adventive Coleoptera. for C. staphylaea in the region. Brown (1950) noted that large quantities VOLUME 110, NUMBER 1 85 were unloaded at ports in Atlantic ment. This work has been assisted by the Canada during the Napoleonic Wars Board of Governors of the Nova Scotia (1799–1815) by British vessels in search Museum. of lumber. Alternatively, plants such as Arteme- LITERATURE CITED sia, Inula, and Veronica are grown as Be´chyne´, J. 1950. 7e contribution a` la connaissance bedding plants, while Achillea and du genre Chrysolina Motsch. (Col. Chrysomelidae. Entomologischen Arbeiten aus Mentha are cultivated as garden herbs; dem Museum G. Frey 1: 47–185. so, it is possible that C. staphylaea could Brown, W. J. 1940. Notes on the American have been accidentally introduced in distribution of some species of Coleoptera association with horticultural practices common to the European and North American and the importation of bedding stock. continents. The Canadian Entomologist 72: 65–78. Chrysolina staphylaea is most fre- ———. 1950. The extralimital distribution of some quently found in moist, coastal meadows species of Coleoptera. The Canadian Ento- similar to those reported for it in Iceland mologist 72: 197–205. by Larsson and Giga (1959). Although ———. 1962. The American species of Chrysolina established in Atlantic Canada it remains Mots., (Coleoptera: Chrysomelidae). The Ca- nadian Entomologist 94: 1–16. an uncommon species. Jolivet (1990) Chantal, C. 1972. Additions a` la faune Cole´opte´r- regarded its distribution in Eurasia as ique du Que´bec. Le Naturaliste canadien 99: boreo-alpine and it remains to be de- 243–244. termined how suitable the habitats and Clark, S. M., D. G. LeDoux, T. N. Seeno, E. G. environmental regimen of Atlantic Ca- Riley, A. J. Gilbert, and J. M. Sullivan. 2004. Host plants of leaf beetle species occurring in nada are for it. Continuing to monitor the United States and Canada (Coleoptera: the species in North America would be Megalopodidae, Orsodacnidae and Chrysome- desirable to ascertain how this popula- lidae, excluding Bruchinae). Coleopterists So- tion will fare. ciety Special Publication 2. 476 pp. Cox, M. L. 1988. Egg bursters in the Chrysome- lidae, with a review of their occurrence in the ACKNOWLEDGMENTS Chrysomeloidea and Curculionoidea (Coleop- Thanks to Claude Chantal (Associa- tera). Systematic Entomology 13: 393–432. tion des entomologistes amateurs du Downie, N. M. and R. H. Arnett. 1996. The Beetles Que´bec), David Langor (Canadian For- of Northeastern North America. The Sandhill Crane Press, Gainesville, Florida, Volume II, est Service, Northern Forestry Research pp. 891–1721. Centre), David B. McCorquodale (Cape Drummond, D. C. 1952. Records of some Chry- Breton University), E. Richard Hoebeke somelidae (Col.) and their food plants. En- (Cornell University), Jean-Pierre Le tomologist’s Monthly Magazine 88: 19. Blanc (Nova Scotia Agricultural Col- Evans, J. D. 1899. List of Coleoptera from Halifax. The Canadian Entomologist 31: 32–321. lege), Jeffrey Ogden (Nova Scotia De- Evenhuis, N. L. 2007. Abbreviations for insect and partment of Natural Resources), and spider collections of the world. Available from: Alexander Konstantinov (Systematic En- http://hbs.bishopmuseum.org/codens/codens-inst. tomology Laboratory, USDA [USNM]) html (accessed 13 June 2007). for making specimens and records avail- Jolivet, P. 1975. Une excursion entomologique a` l’ile de Quelpart (Cheju-Do) et de´couverte able, and to Klaus Bolte for executing d’une espe´ce nouvelle pour la Core´e de the habitus photograph. Shawn M. Chrysolina Motschulsky (Col. Chrysomelidae) Clarke read an earlier draft of the paper re´flexions sur la plantophagie. Bulletin de la and made many constructive sugges- Socie´te´ Linne´enne de Lyon 44: 57–64 and 72– tions. C. G. Majka thanks his colleagues, 81. ———. 1990. Distribution et plantes-hoˆtes de Calum Ewing, David Christianson, and Chrysolina staphylaea (Linne´, 1758). Bulletin Andrew Hebda (Nova Scotia Museum) et Annales de la Socie´te´ royale belge d’En- for continuing support and encourage- tomologie 126: 123–130. 86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON

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