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 regionÒs 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, 2008aÖc) 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 ZealandÒs 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.