The Distribution, Zoogeography, and Composition of Prince Edward Island Carabidae (Coleoptera)

128 The distribution, zoogeography, and composition of Prince Edward Island Carabidae (Coleoptera)

Christopher G. Majka1 Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia, Canada B3H 3A6

Yves Bousquet Central Experimental Farm, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada K1A 0C6

Christine Noronha, Mary E. Smith Agriculture and Agri-food Canada, 440 University Avenue, Charlottetown, Prince Edward Island, Canada C1A 4N6

Abstract—Fourteen species of Carabidae are added to Prince Edward Island’s (P.E.I.) faunal list, bringing the known fauna to 167 species. Bembidion nitidum (Kirby) and Bembidion obtusum Audinet-Serville are newly recorded for the Maritime Provinces. Six species are removed from P.E.I.’s faunal list. The history of collecting of Carabidae on P.E.I. is briefly re- counted. Despite differences in land area and distance from the mainland between P.E.I., Cape Breton Island, and insular Newfoundland, their carabid faunas exhibit many similarities in size and composition. The native carabid fauna of P.E.I. comprises 49% of the species in the combined Maritime Provinces fauna, perhaps reflecting an island-related diminution of species diversity. The proportion of flightless species on P.E.I. (4.9%) is less than that in the Maritime Provinces as a whole (7.1%), an apparent indication that the Northumberland Strait has been a barrier to colonization. Twenty-seven introduced species are found on P.E.I., 26 of which can be classified as synanthropic and may have originated in dry-ballast quarries in southwestern Eng- land. Although the earliest dates of detection of many introduced species on P.E.I. are substan- tially later than elsewhere in the Maritimes, this reflects the paucity of early collecting. Land-management practices on P.E.I. (large-scale and early forest clearances, intensive agriculture, and the extensive use of biocides) may have had an impact on P.E.I.’s carabid fauna.

Résumé—On dénote 14 nouvelles mentions de Carabidae (Coleoptera) pour l’Ile du Prince Édouard, ce qui porte à 167 le nombre d’espèces connues pour la province. Deux de ces espèces, Bembidion nitidum (Kirby) et Bembidion obtusum Audinet-Serville, sont aussi des premières mentions pour les Maritimes. Six espèces sont enlevées de la liste faunistique de Î.-P.-É. On dis- cute brièvement de l’historique des récoltes de Carabidae sur l’île. Malgré des différences notoi- res de superficie et d’éloignement avec le continent, la taille et les caractéristiques de la faune carabidologique de l’Ile du Prince Édouard, du Cap-Breton et de l’île de Terre-Neuve montrent des similitudes évidentes. La faune carabidologique indigène de l’Ile du Prince Édouard repré- sente 49 % de la faune des Maritimes dans son ensemble. La proportion d’espèces incapables de vol sur l’Ile du Prince Édouard (4.9 %) est plus faible que celle des Maritimes (7.1 %) ce qui suggère que le détroit de Northumberland a pu être un obstacle à la colonisation de l’île. On dé- note 27 espèces introduites sur l’île et 26 sont synanthropes et ont pu atteindre l’île en prove- nance du sud-ouest de l’Angleterre via le lest de navires. Le fait que plusieurs espèces introduites aient été détectées sur l’île plus tard qu’ailleurs dans les Maritimes pourrait être le reflet du nombre peu élevé de récoltes faites au 19ième siècle et au début du 20ième siècle. Les pratiques de gestion des terres (déforestation hâtive et à grande échelle, exploitation agricole intensive, et uti- lisation intense d’insecticides) ont pu avoir un impact sur la faune des Carabidae de l’île.

Majka141 et al.

Received 13 April 2007. Accepted 26 August 2007.

1Corresponding author (e-mail: [email protected]).

Introduction (1908) (of Amara impuncticollis (Say)), and Morrison (1941) (of H. rufipes), that included The Carabidae (Coleoptera, ground beetles) P.E.I. records. are the third most species-rich beetle family on In the years following Lindroth’s work, a the North American continent (Marske and Ivie number of collections were made. R. Wenn at 2003). Bousquet and Larochelle (1993) listed the University of Prince Edward Island collected 2623 species in North America north of Mex- 16 species of carabids in 1970 as part of ico, 930 of which occur in Canada. Of these, biodiversity studies on P.E.I. During the 1970s they recorded 154 species as occurring on and 1980s, researchers such as L.S. Thompson Prince Edward Island (P.E.I.), the smallest and F.M. Cannon with Agriculture and province in Canada and one of the least investi- Agri-Food Canada in Charlottetown and students gated areas in the country in terms of the beetle at the University of Prince Edward Island made fauna. A long history of forestry and agriculture additional collections. During the 1980s, re- has greatly affected the environment of the searchers with Agriculture and Agri-Food Can- province. Seventy percent of P.E.I.’s forests ada in Ottawa visited P.E.I. on a periodic basis were cleared during the 20th century (Loo and and collected additional material. Y. Bousquet Ives 2003) and there has been extensive use of (1987) then added two species (Dyschirius insecticides and other biocides in connection sellatus (LeConte) and Oxypselaphus pusillus with land-management practices. (LeConte)) to the P.E.I. fauna. A major contribu- Attention in many quarters has focused on tion to the knowledge of P.E.I. Carabidae was ground beetles as bioindicators of environmen- made by A. Larochelle and M.-C. Larivière, who tal change. Frequently, numbers of large, poorly collected on P.E.I. in 1987. A. Larochelle and Y. dispersing species decrease with increasing dis- Bousquet then returned in 1988. One hundred turbance, whereas those of small generalist spe- and eight species of carabids were newly re- cies with good dispersal ability increase (Rainio corded from P.E.I. during these field trips and Niemelä 2003). In a survey of a large num- (Larochelle and Larivière 1990). Consequently, ber of studies, Rainio and Niemelä (2003) con- when Bousquet’s (1991) checklist of Carabidae cluded that carabids are useful bioindicators. was published, it included records of 137 species Consequently, in terms of building a fuller un- from P.E.I. Subsequently, Bousquet and derstanding of the biodiversity of P.E.I., as well Larochelle’s (1993) catalogue added 17 species, as for environmental monitoring purposes, it is increasing the known total fauna to 154 species; useful to review the past knowledge of this the most recent checklist (Goulet and Bousquet group on P.E.I. and add information on the 2004) recorded 153 species from P.E.I. composition of the province’s carabid fauna. In recent years C.G. Majka has collected The benchmark of research on Canadian Carabidae as part of biodiversity studies on Carabidae is the work of Lindroth (1961, P.E.I., and C. Noronha and M.E. Smith have ex- 1963b, 1966, 1968, 1969a, 1969b), who drew amined the fauna of agricultural fields. Majka on his own extensive fieldwork and research as (2005) added Amara communis (Panzer) to well as compiling what was then known about P.E.I.’s faunal list and Majka et al. (2006) Canadian Carabidae from other sources. Al- added Notiophilus biguttatus (Fabricius), though he collected extensively in both insular Carabus granulatus hibernicus Lindroth, and Newfoundland and Labrador (Lindroth 1955, Ophonus puncticeps (Stephens). The present 1963a) and Nova Scotia (Lindroth 1954), he account adds records of a further 14 species to did not do so on P.E.I. Consequently, only 18 the Carabidae of P.E.I. and removes records of species of carabids are recorded as occurring on 5 species as erroneous, as well as considering P.E.I. by Lindroth (1961, 1963b, 1966, 1968, the overall composition of P.E.I.’s fauna. 1969a, 1969b). These records were based on small numbers of specimens in the Canadian Methods and conventions National Collection, the Harvard University Museum of Comparative Zoology, and the Cor- Specimens of Carabidae originating on P.E.I. nell University insect collection. Lindroth also were examined in a number of regional collec- integrated information from published reports, tions. These included both historical materials such as Benshoter and Cook (1956) (of as well as specimens collected recently as part Omophron americanum Dejean), Brown (1950) of contemporary studies by C.G. Majka, C. (of Harpalus rufipes (DeGeer)), Hayward Noronha, and M.E. Smith. A total of 1136

© 2008 Entomological Society of Canada 130 Can. Entomol. Vol. 140, 2008 specimens of 142 species were examined. This Results information was integrated with data from pre- viously published studies (specified above) and Fourteen species of Carabidae are added to the historical collections in the Canadian National faunal list of P.E.I. Two of these, Bembidion Collection of Insects, Arachnids, and Nema- nitidum (Kirby) and Bembidion obtusum todes and the Museum of Comparative Zool- Audinet-Serville, are newly recorded in the Mari- ogy. The methodology has varied from time Provinces as a whole. Two species, collection to collection, and study to study, but B. obtusum and Harpalus rubripes (Duft- has relied largely on pitfall-trapping and schmid), are introduced Palearctic species, the hand-collecting. The following codens of col- others are native species. The status of six spe- lections (following Evenhuis 2007) were con- cies, Miscodera arctica (Paykull), Amara sulted as part of the study: impuncticollis (Say), Amara torrida (Panzer), Harpalus solitaris Dejean, Calathus gregarius ACNS Agriculture and Agri-Food Canada, (Say), and Cymindis cribricollis Dejean, is re- Kentville, Nova Scotia, Canada viewed and they are removed from the faunal list of P.E.I. Localities mentioned in the text are ACPE Agriculture and Agri-Food Canada, indicated in Figure 1. Details of species newly Charlottetown, Prince Edward Island, recorded follow. Canada CGMC Christopher G. Majka collection, Hali- fax, Nova Scotia, Canada Miscodera arctica (Paykull, 1798) CNC Canadian National Collection of In- Although this species was recorded as occur- sects, Arachnids, and Nematodes, Ot- ring on P.E.I. by Bousquet (1991), it was not tawa, Ontario, Canada listed from P.E.I. in Bousquet and Larochelle JOC Jeffrey Ogden Collection, Truro, Nova (1993). We have not been able to find any evi- Scotia, Canada dence that the species has ever been found on MCZ Museum of Comparative Zoology, P.E.I. and consequently remove it from the fau- Harvard University, Cambridge, Mas- nal list of P.E.I. Carabidae. It was apparently sachusetts, United States of America listed in Bousquet (1991) by mistake. NBM New Brunswick Museum, Saint John, New Brunswick, Canada Bembidion nitidum (Kirby, 1837) NSMC Nova Scotia Museum Collection, Hali- fax, Nova Scotia, Canada Queens Co.: Charlottetown, 1958, D.C. UMNB Université de Moncton, Moncton, New Read, potato field, pitfall trap, (13, ACPE). Brunswick, Canada In the northeast recorded from Massachu- UPEI University of Prince Edward Island, setts, Maine, New York, Ontario, Quebec, and Charlottetown, Prince Edward Island, Vermont (Bousquet and Larochelle 1993). Canada Newly recorded on P.E.I. and in the Maritime Provinces as a whole. Found in a great variety The number of specimens is indicated in pa- of upland and lowland habitats including culti- rentheses together with the collection coden. vated fields (corn, oats, flax, wheat, barley, cab- The distribution of newly recorded species in bage); mostly nocturnal; synanthropic; feeds on northeastern North America is given. For the Diptera larvae; macropterous, a frequent flier purposes of this treatment, northeastern North and strong burrower (Larochelle and Larivière America is taken to consist of the following ju- 2003). risdictions: Connecticut, Labrador, Massachu- setts, Maine, New Brunswick, Newfoundland, Bembidion obtusum New Hampshire, Nova Scotia, New York, On- Audinet-Serville, 1821 tario, P.E.I., Quebec, Rhode Island, Saint-Pierre et Miquelon, and Vermont. The systematics em- Queens Co.: Harrington, 14.vi.2004, C. ployed follow that of Ball and Bousquet (2001). Noronha, barley field, pitfall trap, (5, ACPE). The two components of the province of An introduced Palearctic species recorded in Newfoundland and Labrador are treated here as North America from New York, Ohio, Ontario, separate geographical entities: insular New- Quebec, and Vermont (Bousquet and Larochelle foundland and mainland Labrador. 1993). Newly recorded on P.E.I. and in the

Fig. 1. Map of Prince Edward Island showing localities mentioned in the text.

Maritime Provinces as a whole. Found in a Goulet and Bousquet (2004). However, it is not great variety of upland and lowland habitats recorded by Larochelle and Larivière (1990) including cultivated fields (onion, alfalfa, cab- from P.E.I. Subsequent investigation has re- bage); mostly nocturnal; synanthropic; an vealed no voucher specimens, nor are there pub- effective colonist; North American populations lished records for the province. Consequently, are brachypterous and incapable of flight we regard past reports of this species from (Larochelle and Larivière 2003). Prince Edward Island as having been unsubstan- tiated and therefore have removed it from the faunal list of P.E.I. Carabidae. Pterostichus tristis (Dejean, 1828) Queens Co.: St. Patricks, 19.vii.2001, C.G. Majka, old field, under rock, (1, CGMC). Amara latior (Kirby, 1837) In the northeast recorded from Connecticut, Massachusetts, Maine, New Brunswick, New Prince Co.: Tryon, 2.viii.2001, C. Noronha, Hampshire, Nova Scotia, New York, Ontario, potato field, pitfall trap, (3, ACPE); Tryon, Quebec, and Vermont (Bousquet and Larochelle 15.viii.2001, C. Noronha, potato field, pitfall 1993). Found in lowlands and uplands, in de- trap, (3, ACPE); Queens Co.: Cornwall, ciduous, coniferous, and mixed forests, on 8.vii.1982 and 11.vii.1982, C. Wenn, (4, UPEI); shaded ground and moist soil; nocturnal; feeds Hampshire, 14.vii.1982, Vicki B., (1, UPEI). on Lepidoptera larvae; brachypterous and inca- In northeastern North America recorded from pable of flight (Larochelle and Larivière 2003). Connecticut, Labrador, Massachusetts, Maine, New Brunswick, Newfoundland, New Hamp- shire, Nova Scotia, New York, Ontario, Quebec, Amara torrida (Panzer, 1797) and Vermont (Bousquet and Larochelle 1993). Found in a great variety of upland and lowland Majka et al. (2007a) included this species in habitats including cultivated fields (corn, oats, their checklist of the P.E.I. Carabidae on the alfalfa, cabbage, flax, barley, wheat); nocturnal; strength of its inclusion for P.E.I. in Bousquet feeds on grasshopper eggs; macropterous, and a (1991), Bousquet and Larochelle (1993) and frequent flier (Larochelle and Larivière 2003).

Amara otiosa Casey, 1918 In northeastern North America recorded from Massachusetts, Maine, New Brunswick, Amara impuncticollis (Say, 1823) was reported Newfoundland, New Hampshire, New York, from P.E.I. by Hayward (1908); however, this was Nova Scotia, Ontario, Quebec, and Vermont prior to the revalidation of Amara otiosa by Hieke (Bousquet and Larochelle 1993). Found in low- (2000), formerly considered a synonym of A. im- lands and uplands, primarily on open, dry, and puncticollis. The P.E.I. records would certainly be often sandy ground; mostly nocturnal; attributable to A. otiosa since A. impuncticollis macropterous, an occasional flier and strong (sensu Hieke 2000) does not occur in the region. burrower (Larochelle and Larivière 2003). However, we have not been able to locate these specimens to re-examine them. Harpalus rubripes (Duftschmid, 1812) Amara lunicollis Schiøde, 1837 Queens Co.: Harrington, 26.vii.2004, C. Queens Co.: Harrington, 19.v.2006, C. Noronha, barley field, pitfall trap, (1, ACPE). Noronha, potato field near hedgerow, pitfall An introduced Palearctic species recorded trap, (1, ACPE). from Connecticut, New Hampshire, Nova Sco- In northeastern North America this Holarctic tia, and Rhode Island (Bousquet and Larochelle species is recorded from Labrador, Massachu- 1993; Majka et al. 2006). Found in lowland ar- setts, Maine, New Brunswick, Newfoundland, eas in cultivated fields (hay), vacant lots, and New Hampshire, Nova Scotia, Ontario, Quebec, roadsides on open, dry, sandy ground; mostly and Vermont (Bousquet and Larochelle 1993). nocturnal; synanthropic; an effective colonist; Found in grasslands, meadows, cultivated macropterous, an occasional flier and strong fields, vacant lots, moraines, gravel pits, or- burrower (Larochelle and Larivière 2003). chards, forest clearings, thickets, and open forests; diurnal; feeds on earwigs; macropterous, Harpalus solitaris Dejean, 1829 an occasional flier (Larochelle and Larivière 2003). Majka et al. (2007a) included this species in their checklist of the P.E.I. Carabidae on the strength of its inclusion for P.E.I. in Bousquet Xestonotus lugubris (Dejean, and Larochelle (1993) and Goulet and Bousquet 1829) (2004). However, subsequent investigation has revealed no voucher specimens, nor are there Queens Co.: St. Patricks, 19.vii.2001, C.G. published records for the province. Conse- Majka, old field, under rock, (1, CGMC). quently, we regard past reports of this species In northeastern North America recorded from from Prince Edward Island as having been un- Connecticut, Massachusetts, Maine, New substantiated and therefore remove it from the Hampshire, New York, Nova Scotia, Ontario, faunal list of P.E.I. Carabidae. Quebec, Rhode Island, and Vermont (Bousquet and Larochelle 1993). Found in lowlands and uplands, primarily in deciduous forests and ad- Chlaenius niger Randall, 1838 jacent habitats, on shaded ground; nocturnal; Prince Co.: North Enmore, 10.vii.1988, A. gregarious; predaceous; macropterous, an occa- Larochelle, (1, NSMC); Summerside, 9.vii.1988, sional flier and strong burrower (Larochelle and A. Larochelle, (1, NSMC). Larivière 2003). In the northeast recorded from Connecticut, Massachusetts, Maine, New Brunswick, New- Harpalus plenalis Casey, 1914 foundland, New Hampshire, New York, Nova Scotia, Ontario, Quebec, Rhode Island, and Prince Co.: Tryon, 2.viii.2001, C. Noronha, Vermont (Bousquet and Larochelle 1993). potato field, pitfall trap, (2, ACPE); Tryon, Found in lowland and upland areas, in swamps, 15.viii.2001, C. Noronha, potato field, pitfall on the borders of eutrophic marshes, ponds, irri- trap, (1, ACPE); Queens Co.: Harrington, gation channels, roadside ditches, saline marshes 14.vi.2004, C. Noronha, barley field, pitfall and meadows; nocturnal; macropterous, a fre- trap, (1, ACPE); Harrington, 9.viii.2004, C. quent flier, moderate runner and excellent swim- Noronha, barley field, pitfall trap, (2, ACPE). mer and diver; synanthropic (Larochelle and

Larivière 2003). This species was newly re- strength of its inclusion for P.E.I. in Bousquet corded from P.E.I. by Larochelle and Larivière and Larochelle (1993) and Goulet and Bousquet (1990) and listed from there in Bousquet (2004). However, subsequent investigation has (1991), but was not subsequently included from revealed no voucher specimens, nor are there the province in either Bousquet and Larochelle published records for the province. Conse- (1993) or Goulet and Bousquet (2004). The quently, we regard past reports of this species present records establish its presence in the from Prince Edward Island as having been un- province. substantiated and therefore remove it from the faunal list of P.E.I. Carabidae. Calathus gregarius (Say, 1823) Dromius piceus Dejean, 1831 Majka et al. (2007a) included this species in their checklist of the P.E.I. Carabidae on the Queens Co.: Appin Road, 6.vii.1982, C. Wenn, strength of its inclusion for P.E.I. in Bousquet in grass, (1, UPEI); Argyle Shore, 27.vii.1991, and Larochelle (1993) and Goulet and Bousquet D.F. McAlpine, (2, NBM). (2004). However, subsequent investigation has In the northeast recorded from Connecticut, revealed no voucher specimens, nor are there Massachusetts, Maine, New Brunswick, New published records for the province. Conse- Hampshire, New York, Nova Scotia, Ontario, quently, we regard past reports of this species Quebec, Rhode Island, and Vermont (Bousquet from Prince Edward Island as having been un- and Larochelle 1993). Found in upland and substantiated and therefore remove it from the lowland areas in deciduous, coniferous, and faunal list of P.E.I. Carabidae. mixed forests on shaded ground and under loose bark and bark scales; nocturnal; a fre- Agonum metallescens (LeConte, quent climber on trees, where eggs are laid; 1854) macropterous, a frequent flier (Larochelle and Larivière 2003). PRINCE EDWARD ISLAND: no locality indicated, 1974–83, (1, UPEI). Lebia moesta LeConte, 1850 In the northeast recorded from Massachusetts, Maine, New Brunswick, Newfoundland, New Kings Co.: Woodville Mills, 30.vi.2003, Hampshire, New York, Nova Scotia, Ontario, C.G. Majka, shore of pond, (1, CGMC); Prince Quebec, and Vermont (Bousquet and Larochelle Co.: St. Felix, 2.vi.1996, M.E.M. Smith, blue- 1993). Found in lowland and upland areas, on the berry field, (1, ACPE); Tignish, 28.vi.1995, borders of marshes, ponds, swamps, bogs, rivers, M.E.M. Smith, blueberry field, (1, ACPE); and brooks; mostly nocturnal; macropterous, a Queens Co.: Belle River, 17.vi.1993, M.E.M. frequent flier (Larochelle and Larivière 2003). Smith, blueberry field, (1, ACPE). In the northeast recorded from Labrador, Mas- Platynus tenuicollis (LeConte, sachusetts, Maine, New Brunswick, Newfound- 1848) land, New Hampshire, New York, Nova Scotia, Ontario, Quebec, Rhode Island, and Vermont Kings Co.: Woodville Mills, 6.ix.2001, C.G. (Bousquet and Larochelle 1993). Found in up- Majka, mixed forest: along stream, (1, CGMC). land and lowland areas in meadows, abandoned In the northeast recorded from Connecticut, fields, roadsides, clearings, and open forests; di- Massachusetts, Maine, New Brunswick, New urnal, active on flowering plants and shrubs; Hampshire, New York, Nova Scotia, Ontario, macropterous, an occasional flier and frequent Quebec, and Vermont (Bousquet and Larochelle climber (Larochelle and Larivière 2003). 1993). In deciduous and mixed forests, along turbulent brooks and rivers; on shaded ground Lebia pumila Dejean, 1831 and wet soil; macropterous, a frequent flier (Larochelle and Larivière 2003). Prince Co.: Summerside, 4.ix.2001, C.G. Majka, wet meadow, (1, CGMC); Queens Co.: Cymindis cribricollis Dejean, 1831 Belle River, 17.vi.1993, M.E.M. Smith, blueberry field, (1, ACPE). Majka et al.(2007a) included this species in In the northeast recorded from Connecticut, their checklist of the P.E.I. Carabidae on the Massachusetts, Maine, New Brunswick, New

Hampshire, New York, Nova Scotia, Ontario, Table 1. Checklist of Prince Edward Island Carabidae. Quebec, Rhode Island, and Vermont (Bousquet Omophroninae and Larochelle 1993). Found in upland and Omophronini lowland areas in meadows, cultivated fields, va- Omophron americanum Dejean cant lots, gravel pits, roadsides, and forest clear- Omophron tesselatum Say ings; diurnal, active on flowering plants; Nebriinae gregarious; macropterous, an occasional flier Nebriini and frequent climber (Larochelle and Larivière Nebria pallipes Say 2003). Notiophilini Notiophilus aeneus (Herbst) Notiophilus biguttatus (Fabricius)† Discussion Notiophilus palustris (Duftschmid)† Carabinae Fourteen carabids are newly recorded on Carabini P.E.I., bringing the species of Carabidae now Calosoma frigidum Kirby known to occur there to 167 (Table 1). Table 2 Carabus granulatus hibernicus Lindroth† illustrates the zoogeographic composition of Carabus nemoralis Müller† P.E.I.’s carabid fauna in comparison with that of Carabus serratus Say the carabid faunas of Cape Breton Island, insu- Carabus maeander Fischer von Waldheim* lar Newfoundland (including Saint-Pierre et Cychrini Miquelon), Nova Scotia as a whole, the Nova Sphaeroderus stenostomus lecontei Dejean Cicindelinae Scotia mainland (i.e., excluding Cape Breton), Cicindelini New Brunswick, the Maritime Provinces, and Cicindela duodecimguttata Dejean Atlantic Canada. There is considerable compo- Cicindela hirticollis rhodensis Calder sitional similarity between the island faunas of Cicindela longilabris longilabris Say 2 P.E.I. (which has a land area of 5660 km and Cicindela repanda repanda Dejean lies 13 km from the mainland), Cape Breton Is- Cicindela repanda novascotiae Vaurie land (which has a land area of 10 311 km2 and Cicindela tranquebarica tranquebarica Herbst lies 1.5 km from the mainland), and insular Loricerinae Newfoundland (which has a land area of Loricerini 111 390 km2 and lies 18 km from Labrador and Loricera pilicornis pilicornis (Fabricius)* 110 km from Cape Breton Island), despite their Elaphrinae significant differences in size and distance to Elaphrini the neighbouring continental mainland. The to- Blethisa multipunctata aurata Fisher von Waldheim* tal numbers of species are similar, as are the Blethisa quadricollis Haldeman number of introduced Palearctic species. Insular Elaphrus clairvillei Kirby Newfoundland differs in having double the pro- Elaphrus olivaceus LeConte portion of Holarctic species, reflecting its Elaphrus americanus americanus Dejean greater proximity to circumboreal environments Elaphrus californicus Mannerheim and land masses such as Labrador, Greenland, Scaritinae and Iceland. It is of interest to note that three of Clivinini the four carabids found in Greenland (Böcher Clivina fossor (Linnaeus)† 1988) are also present in Atlantic Canada, and one Dyschirius sphaericollis (Say) of these, Dicheirotrichus cognatus (Gyllenhal), oc- Dyschirius sellatus (LeConte) curs on P.E.I. P.E.I. does have the highest pro- Dyschirius setosus (LeConte) Dyschirius globulosus (Say) portion (15.8%) of introduced carabids of any Dyschirius integer (LeConte) area in the region. Trechinae P.E.I.’s carabid fauna is diminished in com- Broscini parison with that on the neighbouring mainland. Broscus cephalotes cephalotes (Linnaeus)† The native fauna comprises only 47% of the Trechini species found in the combined Maritime Prov- Trechus rubens (Fabricius)† inces. This may represent an island-associated Blemus discus (Fabricius)† diminution, an area effect, a paucity of collect- Bembidiini ing, or a combination of these factors. Of the Bembidion inaequale inaequale Say Bembidion nitidum (Kirby) Maritime Provinces beetle fauna, 39% of † Coccinellidae (Majka and McCorquodale Bembidion properans (Stephens)

Table 1 (continued). Table 1 (continued). Bembidion obtusum Audinet-Serville† Amara angustata (Say) Bembidion nigrum Say Pseudamara arenaria (LeConte) Bembidion stephensii Crotch† Harpalini Bembidion bruxellense Wesmael† Notiobia terminata (Say) Bembidion obscurellum (Motschulsky)* Xestonotus lugubris (Dejean) Bembidion petrosum petrosum Gebler* Anisodactylus kirbyi Lindroth Bembidion sejunctum sejunctum Casey Anisodactylus nigrita Dejean Bembidion tetracolum Say† Anisodactylus rusticus (Say) Bembidion scopulinum (Kirby)* Anisodactylus sanctaecrucis (Fabricius) Bembidion incrematum LeConte Stenolophus fuliginosus Dejean Bembidion nigripes (Kirby)* Stenolophus comma (Fabricius) Bembidion patruele Dejean Stenolophus lineola (Fabricius) Bembidion constrictum LeConte Stenolophus conjunctus (Say) Bembidion contractum Say Bradycellus lecontei Csiki Bembidion mimus Hayward Bradycellus neglectus (LeConte) Bembidion versicolor (LeConte) Bradycellus nigrinus (Dejean) Bembidion quadrimaculatum oppositum Say Bradycellus lugubris (LeConte) Bembidion musicola Hayward Dicheirotrichus cognatus (Gyllenhal)* Bembidion transparens (Gebler)* Acupalpus canadensis Casey Bembidion concretum Casey Acupalpus carus (LeConte) Bembidion fortestriatum (Motschulsky) Acupalpus pumilius Lindroth Bembidion frontale (LeConte) Acupalpus pauperculus Dejean Elaphropus incurvus (Say) Ophonus puncticeps Stephens† Patrobini Harpalus pensylvanicus (DeGeer) Patrobus longicornis (Say) Harpalus rufipes (DeGeer)† Harpalinae Harpalus affinis (Schrank)† Pterostichini Harpalus rubripes (Duftschmid)† Poecilus lucublandus lucublandus (Say) Harpalus fulvilabris Mannerheim Stomis pumicatus (Panzer)† Harpalus laevipes Zetterstedt* Pterostichus commutabilis (Motshulsky) Harpalus plenalis Casey Pterostichus patruelis (Dejean) Harpalus herbivagus Say Pterostichus adstrictus Eschscholtz* Harpalus somnulentus Dejean Pterostichus mutus (Say) Licinini Pterostichus pensylvanicus LeConte Diplocheila obtusa (LeConte) Pterostichus corvinus (Dejean) Badister neopulchellus Lindroth Pterostichus luctuosus (Dejean) Badister grandiceps Casey Pterostichus tenuis (Casey) Badister ocularis Casey Pterostichus coracinus (Newman) Chlaeniini †Pterostichus melanarius (Illiger)† Chlaenius sericeus sericeus (Forster) Pterostichus tristis (Dejean) Chlaenius pensylvanicus pensylvanicus Say Zabrini Chlaenius lithophilius lithophilus Say Amara aulica (Panzer)† Chlaenius niger Randall Amara apricaria (Paykull)† Platnyini Amara avida (Say) Calathus ingratus Dejean Amara fulva (Müller) Laemostenus terricola (Herbst)† Amara latior (Kirby) Synuchus impunctatus (Say) Amara obesa (Say) Oxypselaphus pusillus (LeConte) Amara aenea (DeGeer)† Agonum anchomenoides Randall Amara bifrons (Gyllenhal)† Agonum canadense Goulet Amara communis (Panzer)† Agonum gratiosum (Mannerheim) Amara cupreolata Putzeys Agonum lutulentum (LeConte) Amara familiaris (Duftschmid)† Agonum picicornoides Lindroth Amara laevipennis Kirby Agonum retractum LeConte Amara littoralis Mannerheim Agonum sordens Kirby Amara lunicollis Schiøde* Agonum superioris Lindroth Amara otiosa Casey Agonum thoreyi Dejean* Amara patruelis Dejean* Agonum nigriceps LeConte* Amara quenseli (Schönherr)* Agonum affine Kirby Amara sinuosa (Casey) Agonum cupripenne Say

Table 1 (concluded). Agonum fidele Casey Agonum harrisii LeConte Agonum melanarium Dejean (% of total)

Agonum metallescens (LeConte) Atlantic fauna Agonum muelleri (Herbst)† Agonum octopunctatum (Fabricius) Agonum placidum (Say) Agonum propinquum (Gemminger and Harold) no.

Agonum tenue (LeConte) Total Agonum trigeminum Lindroth Agonum mutatum (Gemminger and Harold) Agonum crenistriatum (LeConte) Platynus decentis (Say) Platynus tenuicollis (LeConte) Platynus mannerheimii (Dejean)* Lebiini Syntomus americanus (Dejean) Dromius piceus Dejean Lebia fuscata Dejean Lebia moesta LeConte (2006), and C.G. Majka (unpublished data). Numbers of spe- Lebia pumila Dejean Lebia viridis Say et al. *Holarctic species.

† ) and Larson and Langor (1982).

Introduced Palearctic species. b

2006), 40% of Mordellidae (Majka and Jackman 2006), and 32% of Cerambycidae (Majka et al. 2007b) occur on P.E.I. Further- more, of the families of saproxylic beetles found in the Maritime Provinces, 27% of Tetratomidae, Melandryidae, Synchroidae, and Scraptiidae (Majka and Pollock 2006) and 33% of Mycteridae, Boridae, Pythidae, Pyrochro- idae, and Salpingidae (Majka 2006) are recorded on P.E.I. Thus, the Carabidae appears proportionately well represented, perhaps indicating a relatively more complete sampling ef- fort for this group. There are six species of P.E.I. carabids that have not been recorded in Nova Scotia: B. nitidum, B. obtusum, Amara angustata (Say), Stenolophus lineola (Fabricius), Agonum superioris Lindroth, and Agonum nigriceps LeConte. All except B. obtusum are native spe- Nearctic species Holarctic species Native species Palearctic species cies, perhaps indicating that they have not No. % No. % No. % No. % crossed the Isthmus of Chignecto to colonize Nova Scotia. There are also six P.E.I. carabids that have not been recorded in New Brunswick: Notiophilus palustris (Duftschmid), Broscus cephalotes cephalotes (Linnaeus), B. obtusum, Stomis pumicatus (Panzer), S. lineola, Acupalpus Comparison of the zoogeographic composition of the Carabidae of Atlantic Canada.

pumilius Lindroth, and O. puncticeps. All except Numbers of species in the Maritime Provinces are derived from Bousquet and Larochelle (1993), Majka S. lineola and A. pumilius are introduced species, Table 2. reflecting differences in the introduction histo- Note: Prince Edward IslandCape Breton IslandInsular NewfoundlandNova Scotia mainland 123Nova ScotiaNew 112 Brunswick 146Maritime Provinces 214 73.7Atlantic Canada 65.9 75.6 80.5 17 268 228 227 36 24 280 20 81.0 10.2 83.5 79.1 21.2 12.4 78.7 140 29 7.5 21 26 148 170 38 83.8 234 8.8 7.7 9.1 87.1 88.1 10.7 88.0 27 297 249 253 22 23 318 32 16.2 89.7 91.2 88.2 12.9 11.9 89.3 167 12.0 34 24 34 170 193 38 266 10.3 11.8 8.8 47 10.7 48 54 331 287 273 75 356 93 81 77 100 ries of P.E.I. and New Brunswick. Acupalpus cies in insular Newfoundland (including Saint-Pierre et Miquelon) are derived from Lindroth (1955, 1963

Table 3. Numbers and percentages of native Carabidae of Atlantic Canada according to wing state and frequency of flight. Prince Edward Maritime Insular Island Nova Scotia Cape Breton Provinces Newfoundland No. % No. % No. % No. % No. % Wing state* Macropterous 108 77.1 197 77.9 123 72.4 229 77.1 102 68.9 Wing-dimorphic 9 6.4 14 5.5 11 6.5 18 6.1 12 8.1 Wing-dimorphic (M) 1 0.7 4 1.6 3 1.8 5 1.7 3 2.0 Wing-dimorphic (B) 15 10.7 19 7.5 18 10.6 22 7.4 19 12.8 Submacropterous 3 2.1 3 1.2 2 1.2 4 1.3 2 1.4 Brachypterous 4 2.9 16 6.3 13 7.6 19 6.4 10 6.8 Total 140 253 170 297 148 Frequency of flight Frequent 77 55.0 138 54.5 92 54.1 157 52.9 70 47.3 Occasional 50 35.7 78 30.8 54 31.8 92 30.8 49 33.1 Incapable of flight 7 5.0 17 6.7 15 8.8 21 7.1 12 8.1 Unknown 6 4.3 20 7.9 9 5.3 27 9.1 17 11.5 Total 140 253 170 297 148 *M, primarily macropterous; B, primarily brachypterous. pumilius, which is present in Nova Scotia, may a barrier to the colonization of P.E.I. by flight- have thus far been overlooked in New Bruns- less carabids. wick. Stenolophus lineola has not otherwise These flightless species include Sphaero- been recorded in the Maritime Provinces; hence, derus stenostomus lecontei Dejean, Pterostichus its presence on P.E.I. (Lindroth 1968) is particu- coracinus (Newman), Pterostichus tristis, larly noteworthy. The introduced B. obtusum and O. pusillus, A. superioris, Platynus decentis the native B. nitidum are the only other Maritime (Say), and Platynus mannerheimii (Dejean). It carabids recorded solely from P.E.I. is possible that these species colonized P.E.I. by P.E.I. and Cape Breton Island also show sub- crossing emergent land in the Gulf of St. Law- stantial commonalities in the composition of rence between 10 000 and 8000 years BP. At their faunas. There are 131 species that have that time, deglaciation, changing sea levels, and been found in both areas, 111 native and 20 in- isostatic rebound contributed to major portions troduced species. Thus, 79% of native species of the Gulf being above sea level for a period of found on P.E.I. are also present on Cape Breton circa 2000 years (Shaw and Gareau 2002). Island. Cape Breton Island’s fauna does have a This extensive area of emergent land con- distinctive component: 18 species (17 native nected P.E.I. and Cape Breton Island with the and 1 introduced) have not been found on the present-day eastern shore of New Brunswick Nova Scotia mainland, and 2 of these, and the northern shore of Nova Scotia. All Bembidion fortestriatum (Motschulsky) and seven of these species are found in New Bruns- Harpalus laevipes Zetterstedt, are found on wick and six (except for A. superioris) occur in P.E.I. Nova Scotia. An interesting feature of the P.E.I. fauna is Twenty-seven carabids found on P.E.I. are in- the proportion of native flightless (brachypter- troduced Palearctic species. Table 4 indicates ous and some submacropterous) species (Ta- the year the species were first recorded on ble 3). Although the overall proportion of such P.E.I., the year and province they were first re- species in the Maritime Provinces is 7.1%, on corded in the Maritime Provinces, and the year P.E.I. there are only seven native flightless spe- they were first recorded in North America. Both cies, 5.0% of the fauna. On Cape Breton Island B. c. cephalotes, and H. rufipes were first de- the proportion is 8.8%, whereas on insular tected in North America on P.E.I. Bembidion Newfoundland it is 8.1%. It would appear that obtusum was first reported in the Maritime the 13 km wide Northumberland Strait has been Provinces from P.E.I. Although the mean year

© 2008 Entomological Society of Canada 138 Can. Entomol. Vol. 140, 2008 ‡ Bionomics † 20062006 4 Synanthropic Synanthropic 2006 3 Synanthropic et al. et al. et al. Maritime Provinces* P.E.I. Source of P.E.I. records No. of quarries Year of first detection re (2003). P North America Lindroth 1890 1890; N.B. 1987 Majka (Illiger) 1926 1926; N.S. 1981 ACPE Synanthropic Wesmael 1907 1910; N.S. 1987 Larochelle and Larivière 1990 Synanthropic (Herbst) 1894 <1894; N.S. ? CNC Synanthropic (Fabricius) 1923 1976; N.S. 2002 Majka Say 1823 1902; N.B. <1991 Bousquet 1991 1 Synanthropic (Linnaeus) 1987 1987; N.S. and P.E.I. 1987 Bousquet 1992 1 Coastal (Stephens) 1942 1942; N.S. 1970 UPEI 3 Synanthropic Crotch 1891 1980; N.S. 1988 CNC Synanthropic (Duftschmid) 1967 1967; N.S. 1987 NSMC 1 Synanthropic? Stephens 1954 1986; N.S. 2002 Majka Audinet-Serville 1956 2004; P.E.I. 2004 ACPE; this paper Synanthropic Müller 1870 1870: N.B. 1970 UPEI Synanthropic (Duftschmid) 1981 1997; N.S. 2004 ACPE; this paper 4 Synanthropic (Panzer) 1984 1984; N.S. <1993 ACPE 1 Synanthropic (Panzer) 1988 1988; N.B. 2001 Majka 2005 1 Synanthropic? (Herbst) 1840 1890; N.S. 1920 MCZ; Lindroth 1955 2 Synanthropic (Duftschmid) 1915 1945; N.S. 1957 ACPE 3 Synanthropic (DeGeer) 1937 1937; P.E.I. 1937 MCZ; Morrison 1941 4 Synanthropic (Paykull) 1875 1899; N.S. 1970 UPEI Synanthropic (Schrank) 1798 1899; N.S. 1970 UPEI 5 Synanthropic (Fabricius) 1875 <1875; N.S. 1988 CNC Synanthropic (Gyllenhal) 1929 1929; N.S. 1987 Larochelle and Larivière 1990 Synanthropic (Fabricius) 1933 1970; N.S. 1988 CNC Synanthropic (Linnaeus) 1915 1926; N.B. 1953 ACPE Synanthropic (Panzer) 1929 1929; N.S. 1987 Larochelle and Larivière 1990 1 Synanthropic (DeGeer) 1828 1947; N.S. 1983 ACPE 6 Synanthropic Year of first detection of introduced species of P.E.I. Carabidae. Number of ballast quarriesInformation in derived southwestern from England Larochelle in and which Larivi the species was found by Lindroth (1957). *N.B., New† Brunswick; N.S., Nova Scotia;‡ P.E.I., Prince Edward Island. Table 4. Notiophilus biguttatus Notiophilus palustris Carabus granulatus hibernicus Carabus nemoralis Clivina fossor Broscus c. cephalotes Trechus rubens Blemus discus Bembidion properans Bembidion obtusum Bembidion stephensii Bembidion bruxellense Bembidion tetracolum Stomis pumicatus Pterostichus melanarius Amara aulica Amara apricaria Amara aenea Amara bifrons Amara communis Amara familiaris Ophonus puncticeps Harpalus rufipes Harpalus affinis Harpalus rubripes Laemostenus terricola Agonum muelleri Mean year of first detection 1913 1939 1981

© 2008 Entomological Society of Canada Majka et al. 139 of first detection of introduced species on P.E.I. proportionately larger numbers of introduced is 1981 (compared with 1913 in North America species found there. and 1939 in the Maritime Provinces), because In view of recent concerns with respect to the of the paucity of early collecting on P.E.I. one impact of anthropogenic activities on the envi- cannot conclude that introductions on P.E.I. ronment (climate change, presence of intro- necessarily occurred later than in other areas. duced and invasive species, effects of biocides), Of these 27 species, all except B. c. cephalotes monitoring ground beetle populations on P.E.I. (a characteristically coastal species) can be cat- would be worthwhile. Remaining stands of egorized as synanthropic. precolonization forest types should be investi- Brown (1950) and Lindroth (1957) developed gated to ascertain whether relict species of the theory that ships’ dry ballast was a probable Carabidae are present in these sites. Such inves- source of entry of many introduced ground- tigations might shed light on early forest condi- dwelling Coleoptera. Of the 27 introduced spe- tions on P.E.I., and consequently the degree to cies found on P.E.I., 15 were found by Lindroth which the natural environment has changed un- (1957) in quarries in southwestern England der human stewardship. where dry ballast destined for Atlantic Canada originated, a suggestive indication that many Acknowledgments may have been introduced via this mechanism. Thanks are extended to Christopher Buddle Indeed, except for C. g. hibernicus Lindroth and (McGill University), Pauline Duerr (Université Blemus discus (Fabricius), all the other intro- de Moncton), Donna Giberson (University of duced P.E.I. carabids are found in England. Prince Edward Island), Donald F. McAlpine Because of the lack of early collecting, there (New Brunswick Museum), Jeff Ogden (Nova may be no ready way to determine how the his- Scotia Department of Natural Resources), and tory of land use on P.E.I. may have affected na- Susan Westby (Atlantic Food and Horticulture tive carabids. Although both the boreal and Research Centre) for making specimens, records, northern deciduous elements of the Acadian and information available. Particular thanks go Forest on P.E.I. are well represented (Erskine to André Larochelle and Marie-Claude Larivière 1960), by the early 18th century significant cut- (Landcare Research, Auckland, New Zealand) ting of the forests had commenced and large ar- for their important contributions to the under- eas of P.E.I. had been burnt. Seventy percent of standing of the province’s carabid fauna. Robb P.E.I.’s forests were cleared in the 20th century Bennett, Christopher Buddle, and two anony- (Loo and Ives 2003). In 1960, 60% of the land mous reviewers read earlier versions of the on P.E.I. was devoted to agriculture and a fur- manuscript and made many constructive sugges- ther 8% was otherwise open (unimproved tions. C.G. Majka thanks David Christianson, wasteland, marsh, barren land, etc.) leaving Calum Ewing, and Andrew Hebda (Nova Scotia only 32% as forest (Erskine 1960). Since then Museum) for continuing support and encourage- farm abandonment has led to some regrowth so ment. This work has been assisted by the Board that by 1992, 57% of the land was forested of Governors of the Nova Scotia Museum. (Anonymous 1992). Where land has not been completely cleared the best quality trees have References been removed for generations, leaving a forest Anonymous. 1992. 1990/1992 Prince Edward Island that is impoverished in terms of genetic quality forest inventory: summary. Forest Branch, Prince and species diversity (Loo and Ives 2003). Only Edward Island Department of Energy and For- relicts of the original vegetation of P.E.I. still estry, Charlottetown, Prince Edward Island. exist. A projection by Sobey and Glen (2004) Ball, G.E., and Bousquet, Y. 2001. Carabidae Latreille, of the pre-European settlement forest of P.E.I. 1810. In American beetles. 1. Archeostemata, indicates that the province was mostly covered Myxophaga, Adephaga, Polyphaga: Staphyliniformia. by upland hardwood forest, a stand type that Edited by R.H. Arnett, Jr. and M.C. Thomas. CRC composes only 21.7% of its present forested Press, Boca Raton, Florida. pp. 32–132. area. It is possible that this long history of Benshoter, C.A., and Cook, E.F. 1956. A revision of the genus Omophron (Carabidae: Coleoptera) of anthropogenic activities (which in the past North America north of Mexico. Annals of the 60 years have included the extensive use of var- Entomological Society of America, 49: 411–429. ious biocides) may have affected P.E.I.’s ground Böcher, J. 1988. The Coleoptera of Greenland. beetle fauna, a fact that may be reflected by the Meddelelser om Grønland Bioscience, 26: 1–100.

Bousquet, Y. 1987. The carabid fauna of Canada and Lindroth, C.H. 1957. The faunal connections be- Alaska: range extensions, additions, and descrip- tween Europe and North America. Almqvist & tions of two new species of Dyschirus (Coleo- Wiksell, Stockholm, Sweden. ptera: Carabidae). The Coleopterists Bulletin, 41: Lindroth, C.H. 1961. The ground-beetles (Carabidae, 111–135. excl. Cicindelinae) of Canada and Alaska. Part 2. Bousquet, Y. 1991. Family Carabidae: ground bee- Opuscula Entomologica Supplementum, 20: 1–200. tles, In Checklist of beetles of Canada and Alaska. Lindroth, C.H. 1963a. The fauna history of New- Edited by Y. Bousquet. Publication 1861/E, Agri- foundland illustrated by carabid beetles. Opuscula culture Canada, Ottawa, Ontario. pp. 8–60. Entomologica Supplementum, 23: 1–112. Bousquet, Y. 1992. Bembidion femoratum Sturm and Lindroth, C.H. 1963b. The ground-beetles (Carabidae, Amara communis (Panzer) (Coleoptera: Carabi- excl. Cicindelinae) of Canada and Alaska. Part 3. dae) new to North America. Journal of the New Opuscula Entomologica Supplementum, 24: 201–408. York Entomological Society, 100: 503–509. Lindroth, C.H. 1966. The ground-beetles (Carabidae, Bousquet, Y., and Larochelle, A. 1993. Catalogue of excl. Cicindelinae) of Canada and Alaska. Part 4. the Geadephaga (Coleoptera: Trachypachidae, Opuscula Entomologica Supplementum, 29: 409–648. Rhysodidae, Carabidae including Cicindelini) of Lindroth, C.H. 1968. The ground-beetles (Carabidae, America north of Mexico. Memoirs of the Ento- excl. Cicindelinae) of Canada and Alaska. Part 5. mological Society of Canada, 167: 1–397. Opuscula Entomologica Supplementum, 33: 649–944. Brown, W.J. 1950. The extralimital distribution of Lindroth, C.H. 1969a. The ground-beetles (Carabidae, some species of Coleoptera. The Canadian Ento- excl. Cicindelinae) of Canada and Alaska. Part 6. mologist, 82: 197–205. Opuscula Entomologica Supplementum, 34: 945–1192. Erskine, D.S. 1960. Plants of Prince Edward Island. Lindroth, C.H. 1969b. The ground-beetles (Carabidae, Publication 1088, Research Branch, Canada De- excl. Cicindelinae) of Canada and Alaska. Part 1. partment of Agriculture, Ottawa, Ontario. Opuscula Entomologica Supplementum, 35: I–XLVII. Evenhuis, N.L. 2007. Abbreviations for insect and Loo, J., and Ives, N. 2003. The Acadian forest: his- spider collections of the world [online]. Available torical condition and human impacts. The Forestry from: http://hbs.bishopmuseum.org/codens/codens- Chronicle, 79: 462–472. inst.html [accessed 21 June 2007]. Majka, C.G. 2005. The Palearctic species Bembidion Goulet, H., and Bousquet, Y. 2004. The ground bee- femoratum and Amara communis (Coleoptera: tles of Canada [online]. Available from Carabidae): new records and notes on modes of http://digir.agr.gc.ca/spp_pages/carabids/phps/ introduction to North America. The Canadian En- index_e.php [accessed 21 June 2007]. tomologist, 137: 532–538. Hayward, R. 1908. Studies in Amara. Transactions Majka, C.G. 2006. The Mycteridae, Boridae, Pythidae, Pyrochroidae, and Salpingidae (Coleoptera: of the American Entomological Society, 34: Tenebrionoidea) of the Maritime Provinces of Can- 13–65. ada. Zootaxa, 1250: 37–51. Hieke, F. 2000. Revision einiger Gruppen und neue Majka, C.G., and Jackman, J.A. 2006. The Arten der Gattung Amara Bonelli, 1810 (Coleop- Mordellidae (Coleoptera) of the Maritime Prov- tera: Carabidae). Annales Historico-Naturales inces of Canada. The Canadian Entomologist, 138: Musei Nationalis Hungarici, 92: 41–143. 292–304. Larochelle, A. and Larivière, M.-C. 1990. Premiéres Majka, C.G., and McCorquodale, D.B. 2006. The mentions de Carabidae (Coleoptera) pour le Maine, Coccinellidae (Coleoptera) of the Maritime Prov- le Nouveau-Brunswick, la Nouvelle-Écosse et inces of Canada: new records, biogeographic L’Île-du-Prince-Édouard. Fabreries, 15(2): 25–37. notes, and conservation concerns. Zootaxa, 1154: Larochelle, A., and Larivière, M.-C. 2003. A natural 49–68. history of the ground-beetles (Coleoptera: Majka, C.G., and Pollock, D.A. 2006. Understanding Carabidae) of America north of Mexico. Pensoft, saproxylic beetles: new records of Tetratomidae, Sofia, Bulgaria. Melandryidae, Synchroidae, and Scraptiidae from Larson, D.J., and Langor, D.W. 1982. The carabid the Maritime Provinces of Canada (Coleoptera: beetles of insular Newfoundland (Coleoptera: Tenebrionoidea). Zootaxa, 1248: 45–68. Carabidae: Cicindellidae) — 30 years after Majka, C.G., Cook, J., and Westby, S. 2006. Intro- Lindroth. The Canadian Entomologist, 114: duced Carabidae (Coleoptera) from Nova Scotia 591–597. and Prince Edward Island: new records and eco- Lindroth, C.H. 1954. Carabid beetles from Nova logical perspectives. The Canadian Entomologist, Scotia. The Canadian Entomologist, 86: 299–310. 138: 602–609. Lindroth, C.H. 1955. The carabid beetles of Newfound- Majka, C.G., Bousquet, Y., and Westby, S. 2007a. land including the French islands St. Pierre and The ground beetles (Coleoptera: Carabidae) of the Miquelon. Opuscula Entomologica Supplementum, Maritime Provinces of Canada: review of collect- 11: 1–160. ing, new records, and observations on composition,

zoogeography, and historical origins. Zootaxa, Rainio, J., and Niemelä, J. 2003. Ground beetles 1590: 1–36. (Coleoptera: Carabidae) as bioindicators. Biodi- Majka, C.G., McCorquodale, D.B., and Smith, M.E. versity and Conservation, 12: 487–506. 2007b. The Cerambycidae (Coleoptera) of Prince Shaw, J., and Gareau, P. 2002 Changing sea levels in Edward Island: new records and further lessons in Atlantic Canada. CoastWeb: Geological Survey of biodiversity. The Canadian Entomologist, 139: Canada [online]. Available from http://gsc.nrcan. 258–268. gc.ca/coast/sealevel/index_e.php [accessed 21 June 2007]. Marske, K.A., and Ivie, M.A. 2003. Beetle fauna of the United States and Canada. The Coleopterists Sobey, D.G., and Glen, W.M. 2004. A mapping of the Bulletin, 57: 495–503. present and past forest-types of Prince Edward Is- Morrison, F.O. 1941. Imported carabid beetle may land. The Canadian Field-Naturalist, 118: 504–520. be a potential pest. The Canadian Entomologist, 73: 217–218.