J.Klimaszewski et al. 2010. Review of adventive species of Coleoptera... recorded from eastern Canada

Jan Klimaszewski, David Langor, Christopher G. Majka, Patrice Bouchard, Yves Bousquet, Laurent LeSage, Ales Smetana, Patricia Sylvestre, Georges Pelletier, Anthony Davies, Pierre DesRochers, Henri Goulet, Reginald Webster & Jon Sweeney. 2010.
Review of adventive species of Coleoptera (Insecta) recorded from eastern Canada.
Pensoft Publishers. Sofia-Moscow.
First published 2010 ISBN 978-954-642-552-2 (hardback)
Pensoft Series Faunistica, No 94. 272 pp.
Printed in Bulgaria, April 2010.

Introduction

For as long as entomologists have turned their attention to Coleoptera in eastern Canada, the subject of adventive species has been a topic of considerable interest. In the first paper about Coleoptera from eastern Canada, Kirby (1837) listed eleven adventive species including Pterostichus melanarius (Illiger), Philonthus politus (Linnaeus), Philonthus ventralis (Gravenhorst), Attagenus pellio Linnaeus, Dermestes lardarius Linnaeus, Gastrophysa polygoni (Linnaeus), Omosita discoidea (Fabricius), Nitidula rufi pes (Linnaeus), Microbregma emarginatum (Duftschmid), Tenebrio molitor (Linnaeus), and Necrobia violacea (Linnaeus). This was an indication that a significant number of adventive species had already established themselves in the region. Early reports of adventive beetles in Nova Scotia include nine Palaearctic species (Jones 1870), four weevil species (Harrington 1891), and eight additional species (Evans 1899).

Recent archaeo-entomological work now makes it clear that a substantial number of adventive species were established on the continent much earlier that we had previously known. In Newfoundland and Labrador, Prevost and Bain (2007) found remains of adventive species such as Quedius mesomelinus (Marsham), Cercyon analis (Paykull), Ptinus fur (Linnaeus), Tipnus unicolor (Piller and Mitterpacher), Oryzaephilus surinamensis (Linnaeus), Mycetaea subterranea (Fabricius), Bruchus pisorum (Linnaeus), Latridius minutus (Linnaeus), and Sitophilus granarius (Linnaeus) in archeological excavations of latrines that dated from ca. 1620.

In Quebec, Bain (1999) found 61 species of adventive beetles in excavations of latrines dating from ca. 1850, a strong indication of the scale of the synanthropic beetle fauna that had already established itself in this region over 150 years ago. In similar excavations in Boston, Massachusetts dating from the 17th century, Bain (1998) found 25 adventive species of Coleoptera, all of which are also established in eastern Canada.

Although insect collections from the 17th century that could document importations are lacking, Majka and LeSage (2007) pointed out that sites such as Annapolis Royal in Nova Scotia have been more or less continuously inhabited by European settlers since 1605, and were the hub of a considerable trans-Atlantic commercial trade. They further pointed out that in contemporary collections of Coleoptera from Annapolis Royal, the proportion of adventive species collected there is exactly double that of the provincial average, an indirect indication that the area may have had a long history of importation and establishment of adventive species.

Many writers have contributed to our understanding of adventive species in North America; however, two deserve particular mention. In a series of papers, the Canadian entomologist, William J. Brown (1940a, 1950, 1967), documented the presence, distribution, and early timelines of over 110 species of Coleoptera in Canada, most of them from eastern provinces. This substantial undertaking helped to delineate the composition and the scale of this adventive fauna. Brown (1940a, 1950) also helped to develop the theory that the transport of dry ballast (bulky rock, sand, and soil) in trans-Atlantic shipping was responsible for many such introductions. Brown (1950) noted that large quantities of dry ballast were unloaded at ports in the Maritime Provinces by British vessels in search of lumber, commencing with the Napoleonic Wars (1799-1815). Such practices persisted in varying degrees until the late 19th and early 20th centuries.

The great Swedish zoogeographer and coleopterist, Carl Lindroth (1954, 1955, 1957, 1963), continued to develop these ideas, in part as a result of the extensive fi eldwork that he conducted in Atlantic Canada on the regionТs carabid fauna. Lindroth (1957) conducted vegetational and faunal surveys at sites in Great Britain known to have been sources of dry ballast in the trans-Atlantic shipping trade, and found many species that had been introduced into North America. The ideas developed by these two pioneering coleopterists, with their strong interests in trans-Atlantic zoogeography, remain a pillar on which subsequent research has been based.

In Atlantic Canada, investigators such as Hoebeke and Wheeler (1996a, 1996b, 2000, 2003), Wheeler and Hoebeke (1994), Johnson (1990b), Bousquet (1992), many papers by C.G. Majka and J. Klimaszewski and their colleagues cited in the references, and a considerable number of other studies, have reported many additional adventive species in many families of Coleoptera. Recent fi eldwork and examination of hitherto unidentified specimens in collections have focused attention on several unreported adventive species in the region and have suggested that many species have a wider distribution than previously documented.

In the research that has been conducted to date, it is clear that there are a number of mechanisms that are responsible for the introduction of adventive species:

1. Dry ballast. As documented by the studies of Lindroth and Brown, dry ballast consisted of rocks, soil, rubble, and associated material from a number of quarries in southwestern England, which was placed in the holds of ships to give them stability as they crossed the Atlantic from east to west in pursuit of the lumber and fish trade. Th e quantities of ballast were so sizeable that British naval authorities forbade the captains of vessels from simply dumping this material overboard in harbours, and so it was unloaded on land. Since no quarantine measures existed in those times, all associated plants and animals living in this material were simultaneously unloaded. It is clear that this mechanism was a pathway for the introduction of many adventive species of fl ora and fauna, including substantial numbers of Coleoptera, especially ground and rove beetles.

2. Agricultural and horticultural importations. There is a long history of importation of agricultural and horticultural plants into North America. Majka and Klimaszewski (2004) suggested that species such as Chrysolina staphylaea (Linnaeus) and Meligethes viridescens (Fabricius), associated with various imported plant species, may have been introduced into North America via this mechanism. LeSage et al. (2007, 2008), LeSage and Majka (2009), and Majka and LeSage (2006, 2007, 2008aЦc) have documented a variety of leaf beetles associated with agricultural and horticultural plants introduced into Atlantic Canada.

Majka and LeSage (2007) also noted that Annapolis Royal, Nova Scotia, was the site of the largest plant nursery in eastern Canada from 1885-1945. This town is also the site of the earliest known records of the adventive Palaearctic leaf beetle, Pyrrhalta viburni (Paykull), in North America (first detected in 1924). During this period Annapolis Nurseries were importing at least three exotic European species of Viburnum, the host plant of P. viburni, so it is certainly possible that the leaf beetle was introduced in association with these nursery plants.

3. Importations of silvicultural products. There is a long history of silvicultural importations of seedlings of a large variety of trees and shrubs into eastern Canada, both for commercial as well as horticultural purposes. Maj ka and Klimaszewski (2004) noted a variety of beetle species such as Phloeocharis subtilissima Mannerheim, Cephennium gallicum Ganglbauer, Dromius fenestratus (Fabricius), and Scymnus suturalis Th unberg that are specifically associated with coniferous trees and pointed out the long history (commencing in 1878) of importation of tens of thousands of forest seedlings of a variety of species of trees from various points of origin in Europe. Recently the biodiversity of adventive insects and mites on woody plants in Canada was reviewed by Langor et al. (2008).

4. Importations associated with livestock. There is similarly a long history of importation of livestock into eastern Canada, a trade that offered the possibility of direct and indirect importations of a variety of beetles associated with such stock, or with their manure. Majka and Klimaszewski (2004) noted that species of scarab beetles in the Aphodiini such as Aphodius fi metarius (Linnaeus) and Teuchestes fossor (Linnaeus), which are commonly associated with cattle manure, and have been documented, in Halifax since at least 1869, could have been imported in association with livestock shipments. Horses were imported to Sable Island commencing in 1783 and it is probable that the adventive scarab beetles Onthophagus nuchicornis (Linnaeus), Eupleurus subterraneus (Linnaeus), and Aphodius fi metarius, associated with horse dung and abundant throughout the island, came to the island in association with such livestock shipments (Wright 1989).

Documenting a very different livestock associated importation, Majka et al. (2007h) reported the importation and establishment of the Palaearctic ptinid, Ptinus sexpunctatus Panzer, in association with the importation of blue orchard mason bees, Osmia lignaria Say (Hymenoptera: Megachilidae), from colonies in Utah in the United States. They also noted the importation of the meloid, Tricrania stansburyi Haldeman, to Nova Scotia via the same mechanism, although this latter species has apparently not become established in the province.

5. Importations associated with wood products. Phloeophagous and xylophagous species in the Ptinidae, Bostrichidae, Buprestidae, Cerambycidae, and Curculionidae have been intercepted numerous times in wood "dunnage" used to pack shipping containers arriving at North American ports from overseas (Haack 2006). Th is pathway may be responsible for the introduction of Anoplophora glabripennis (Haack et al. 2010) and Tetropium fuscum (Smith and Hurley 2000) into North America. Softwood "dunnage" used to pack heavy goods in ships and shipping containers has long been recognized as a significant pathway for movement of bark- and wood-boring beetles (Ridley et al. 2000). More than 100 species of bark and ambrosia beetles (Curculionidae: Scolytinae) have been intercepted at New ZealandТs borders, mainly in international shipments associated with wood dunnage, crating or sawn timber (Brockerhoff et al. 2006). In a similar study, Haack (2006) found interception records of 147 different species of bark- and wood-feeding Buprestidae, Cerambycidae, and Scolytinae at United States ports between 1985 and 2005. Both studies found that there was a relationship between interception frequency and probability of establishment of bark beetles (Brockerhoff et al. 2006; Haack 2006).

6. Importations associated with stored products such as grain, cereal and other organic materials. Bousquet (1990a) found that nearly two-thirds of the 120 species associated with stored products in Canada are adventive species, a clear illustration of the scale of this adventive fauna and of its economic importance. Majka (2007a) found that 50% (37 of 74 species) of Bostrichiformia fauna (species in the Derodontidae, Dermestidae, Bostrichidae, and Ptinidae, many of which are stored product pests) of the Maritime Provinces were adventive species.

7. Intentional introductions (e.g., for biological control of pest plants and animals). Sampson and Ingraham (1989) and Sampson and MacSween (1993) surveyed the extensive recent programs of intentional introductions of insects for biocontrol purposes in the Maritime Provinces of Canada, primarily of beetles in the Chrysomelidae, Brentidae, and Curculionidae. Gordon (1985) provided documentation and a survey of the 144 species of exotic Coccinellidae that have, at various times, been intentionally introduced for biocontrol purposes in North America. Of these, 11 species have been intentionally introduced in eastern Canada.

The ecological impacts of adventive insect species are not well understood; there have been a number of studies on the effects of invasive insects on native biodiversity but very few on their effects on ecosystem processes (Kenis et al. 2009). Cormier et al. (2000) found that 95% of the coccinellids captured in fields on Cape Breton Island were introduced species ( Hippodamia variegata (Goeze), Coccinella septempunctata Linnaeus, and Propylaea quatuordecimpunctata (Linnaeus), whereas only 5% of the individuals belonged to six native species. In Nova Scotia, there are records of the native Hippodamia parenthesis (Say) from 1927 to 1961, but virtually no recent specimens have been collected. It is possible that this species has declined in response to competition with introduced species (Majka and McCorquodale 2006).

In Manitoba, populations of the native Hippodamia tredecimpunctata tibialis (Say), H. convergens Guerin-Meneville, H. parenthesis, Coccinella transversoguttata richardsoni Brown, and Coccinella trifasciata perplexa Mulsant have all declined after the establishment of the adventive C. septempunctata (Turnock et al. 2003). Wheeler and Hoebeke (1995b) suggested that C. septempunctata may adversely aff ect Coccinella novemnotata Herbst. In the orchards of the Niagara Peninsula of Ontario, Stethorus p. punctum (Le-Conte) was displaced by the adventive S. punctillum Weise (Putman 1955). Maerz et al. (2005) found that the introduced weevil Barypeithes pellucidus (Boheman) has had a signifi cant eff ect on the diet of red-backed salamanders, Plethedon cinereus (Green), in New York and Pennsylvania where beetles, particularly B. pellucidus, comprised the largest proportion of food items for salamanders in upland forests and the second largest proportion (after earthworms) in lowland forests. The authors conclude that "the seasonally hyper-abundant B. pellucidus had a strong effect on seasonal fluctuations in P. cinereus diet" and further hypothesized that the "influence of introduced prey on temporal and geographic food resources contributes to temporal and geographic demographic and phenotypic variation among P. cinereus populations."

In a study of the fauna of owl nests in mainland Nova Scotia and Cape Breton Island, Majka et al. (2006b) found that 50% of the 14 Coleoptera found in such habitats were adventive species. They further remarked on the degree to which adventive species of Coleoptera appear to have penetrated indigenous habitats, even in remote areas where these owl nests were found, far from any human habitation, and even in habitats (owl nests in undisturbed coniferous forests) that one might expect would be relatively free of anthropogenic influences. Similar investigations and documentation of the impacts of adventive species clearly need to be undertaken in relation to other groups of beetles.

The emerald ash borer, Agrilus planipennis Fairmaire, is an adventive species from Asia that has killed tens of millions of ash trees (Fraxinus spp.), and spread to thirteen states and two provinces since its discovery in the Detroit-Windsor area in 2002 (Haack et al. 2002; http://www.emeraldashborer.info/map.cfm). It likely arrived in North America via either infested wood packing material or live ash nursery stock, and unlike its native congeners (e.g., Agrilus anxius and A. bileneatus), A. planipennis is not acting as a secondary pest, but is attacking and killing healthy trees, and is considered a serious threat to ash trees in North America (Poland and McCullough 2006). Th e easternmost location in Canada where the emerald ash borer is known to be established is near Montreal, Quebec, where it was discovered in June 2008 (http://www.inspection. gc.ca/english/plaveg/pestrava/surv/sit2008e.shtml#agrpla ).

Improved monitoring programs are needed to detect further importations of adventive species. This requires the ongoing support and development of taxonomic resources (materials, reference collections, training, personnel) to be able to recognize such species. Museum research in historical collections of Coleoptera is important for establishing early timelines of the detection of adventive species. Without such information it would not be possible to trace the dispersal patterns of adventive species within the continent such as the detailed work on adventive Amara species in North America undertaken by Hieke (1990). In this volume we have compiled the available information on early dates of detection of adventive species with the objective of motivating further research on this topic. It is also increasingly apparent that archaeo-entomological research such as that being conducted by Allison Bain and her colleagues at Universite Laval (e.g., Bain 1998, 1999; Bain and Prevost 2010) off ers important opportunities for extending these timelines back hundreds of years before voucher specimens were preserved in museum collections.

This volume reviews our current understanding of adventive Coleoptera in eastern Canada and resulted from the collective efforts of entomologists from the Canadian Forest Service, the Nova Scotia Museum, and the Canadian National Collection of Insects, Agriculture and Agri-Food, Canada. Its focus is on the changing natural world around us, mainly due to human commercial activities. It brings together all available knowledge on adventive species of Coleoptera recorded from eastern Canada. Th e adventive fauna is increasingly penetrating native habitats and mixing with native species. It is clear that many species have become established in the region in the past 400 years since the time European settlement of the continent began. Adventive species now constitute a large component of our fauna. This is the first comprehensive contribution providing baseline data on adventive species of the region. It is our hope that it will enhance an understanding of these adventive species and promote further research on their role in our ecosystems, their negative and positive impacts on our economy, and on the native fauna.