The Coleopterists Bulletin, 68(3): 441–599. 2014. THE GROUND BEETLE (COLEOPTERA:CARABIDAE)FAUNA OF MAINE, USA RICHARD G. DEARBORN Forest Insect & Disease Laboratory, Maine Dept. of Agriculture, Conservation and Forestry 168 State House Station, Augusta, ME 04333-0168, U.S.A. ROBERT E. NELSON Department of Geology, Colby College 5804 Mayflower Hill, Waterville, ME 04901-8858, U.S.A. [email protected] CHARLENE DONAHUE Forest Insect & Disease Laboratory, Maine Dept. of Agriculture, Conservation and Forestry 168 State House Station, Augusta, ME 04333-0168, U.S.A. ROSS T. BELL Department of Biology, Marsh Life Science Building, University of Vermont Burlington, VT 05405-0086, U.S.A. AND REGINALD P. W EBSTER 24 Mill Stream Drive, Charters Settlement, NB E3C 1X1, CANADA ABSTRACT A survey of the modern carabid fauna of Maine has shown that the fauna consists of 425 documented species, 14 more than previously documented for the Maine fauna in the latest catalog for the family in North America or in the most recent checklist on the state beetle fauna. New state records are Agonum cupreum Dejean, Amara anthobia Villa and Villa, Anisodactylus laetus Dejean, Bembidion intermedium Kirby, Bembidion sejunctum sejunctum Casey, Brachinus vulcanoides Erwin, Diplocheila impressicollis Dejean, Elaphropus dolosus LeConte, Lebia grandis Hentz, Myas coracinus Say, Olisthopus micans LeConte, Panagaeus fasciatus Say, Pentagonica picticornis Bates, and Tachyta parvicornis Notman. The record for B. sejunctum sejunctum is the first for the species in the eastern United States. A recent record for Omophron labiatum (F.) in the state could not be substantiated by any specimen. Notes on biology are presented for species for which that knowledge exists. Distributions are presented for all taxa based on standard biophysical regions for the state and the knowledge of those distributions; distribution maps are presented for all species for which township records are known and for which we have specimen records in our database. Work on better defining the current distributional limits is ongoing. Several adventive European taxa have already exhibited very rapid dispersal across the state. Key Words: biodiversity, Nearctic, distribution, biology, maps Knowledge of the distribution of organisms on European honeybees in California (Core on Earth is intrinsically important to under- et al. 2012), for example, are helping to focus standing the total biosphere, its functions, and attention on the roles of native pollinators – its potential interactions with the climatic and long ignored by most, even in the entomo- human environment. Knowledge of the insect logical communities. fauna in particular is important, as insects and The family Carabidae among the beetles is birds are the two major groups of organisms in one of the largest in the world, with approxi- which most species are capable of flight, and mately 33,905 described species thus far, including thus sufficiently mobile to be able to respond Cicindelini (Bousquet 2012), with 2,635 species- rapidly to environmental change. The ecological group taxa reported from North America north habits of many insects are also not at all under- of Mexico by Bousquet and Larochelle (1993). stood, yet they play pivotal roles in ecosys- Bousquet (2012), however, estimated that the tem nutrient cycling or other critical functions. true number of North American carabid species- Recent episodes of parasitism by native flies group taxa would eventually number about 441 442 THE COLEOPTERISTS BULLETIN 68(3), 2014 2,560 (2,676 including all Geadephaga), given 2008) and butterflies in Massachusetts (Breed assumptions about the potential synonymization et al. 2013). Exploration of previously ignored of numerous Casey species with future revisions or insufficiently sampled microhabitats can of various groups. also yield new species to the list, as was found Lindroth (1961– 1969b) provided the first with the discovery of the previously unknown major review of North American Carabidae species Platynus indecentis by Liebherr and across all species, though his studies addressed Will (1996). only the Canadian-Alaskan fauna and excluded Major recent works (Larochelle and Larivière the Cicindelini. Nonetheless, most of the species 1990; Bousquet 2010, 2012; Majka et al. 2011) in the Maine fauna were included in this valu- have documented well the overall carabid fauna able major work. Although they clearly stated of the state, though detailed distributions within that it was not to be considered a state list, the the state have not been attempted previously; for annotated catalogue of the Maine Forest Service many species, the numbers of specimens known (MFS) collection by Dearborn and Donahue are too few to allow definitive statements to be (1993) was apparently the first systematic list made of where a species is or is not to be found. of the carabid fauna of Maine and included some It should be noted that Bousquet (2010, 2012) 145 species that could be found in the state col- listed a number of species for the state for which lection (including Cicindelidae, which they listed we have been unable to locate specimens, but which as a separate family). Bousquet and Larochelle presumably are present in more recent additions (1993) listed 343 species in the state fauna in their to the Canadian National Collection in Ottawa catalogue of the family for all of North America, than were there when we conducted our survey of which Bousquet (2010) subsequently expanded to their holdings. Majka et al. (2011) accepted all 393 species and then (Bousquet 2012) to 411 species of Bousquet’s (2010) records and added 36 new by accepting the additions contributed by Majka records to the known state fauna. et al. (2011) in their valuable synthesis of the This project was undertaken in 1995 as a entire Maine beetle fauna. companion study to a survey of the Vermont This rich diversity is made possible by habitat and New Hampshire carabid faunas undertaken and/or food preference specializations among by Ross and Joyce Bell, similar to the way in many taxa, allowing closely related species to which Chris Majka and colleagues (Majka 2005; occupy numerous discrete microhabitats within Majka et al. 2006, 2007, 2008; Majka and Bousquet a small area. It is this habitat specialization that 2008; Majka and Gilhen 2008; Majka and Vickery makes their potential as environmental indicators 2008; Webster and Bousquet 2008; Webster so significant. and DeMerchant 2012) Majka et al. 2006, 2007, Geographically, Maine is also particularly well 2008 have been very efficiently and effectively situated as a potential biothermometer of the clarifying our knowledge of the carabid fauna of impacts of regional environmental change. Situ- the adjacent Maritime Provinces of Canada. Our ated in the northeastern corner of the USA, the long-term goals are to document as thoroughly as state borders only one other US state (New possible what is known of the modern carabid Hampshire, to the west) but two Canadian prov- beetle fauna of the state of Maine, based on actual inces to the north (Québec and New Brunswick). specimens (or recent highly reliable literature Since this location also lies in the ecotonal sources) and to be able to delimit the range of each boundary between the boreal conifer forest and of these species, in a manner comparable to that the eastern hardwood forest of North America, achieved by Larochelle (1975) for the carabid many eastern North American species reach their fauna of Québec and Labrador. Thus, our study northern or southern distributional limits within will complement that of Krinsky and Oliver (2001) the state. for Connecticut and that of Sikes (2004) for Knowledge of current distributions of native Rhode Island; when the Vermont-New Hampshire species is also critical for understanding future fauna is published, only Massachusetts will remain changes in their distributions due to climate change. without its own statewide carabid survey among This is also particularly important with regards the New England states. to documenting rates of spread for adventive taxa This paper represents a preliminary report on that have readily become established in the state that study, based on a total of over 5,400 indi- (e.g., Harpalus rufipes (DeGeer) and Harpalus vidual species records (= species × townships rubripes (Duftschmid)). of record) for the state as a whole. Work is con- A warming climate may also result in north- tinuing and the long-term plan is to produce a ward spread of taxa previously restricted to more more comprehensive report when we believe southern latitudes, as has already been docu- the overall fauna is much better understood. This mented for deer ticks in Maine (Lubelczyk current work is intended as a status report on the THE COLEOPTERISTS BULLETIN 68(3), 2014 443 results of the first decade of study undertaken. documentation, including Bousquet (2012) and Specimen data are complete as of the year 2000, Majka et al. (2011). Locality records in recent and thus must be seen as a “snapshot”; of what taxonomic revisions of Harpalus Latreille (Noonan was known of the fauna at that time, although 1991), Anisodactylus Dejean (Noonan 1996), and recent literature records of major significance Pseudoperyphus Hatch (Maddison, 2008) were (Maddison 2008; Ward and Mays 2010) have accepted, as were records from the recent syn- also been included. Although work is continuing thesis on Cicindelinae (Leonard and Bell 1999) by several workers on Maine specimens, revisiting and the intensive early faunal surveys of Mount all collections to include newer records, however, Desert Island (Procter 1946), with the taxonomy was not feasible. Undoubtedly, much remains to of this last source brought into conformity with be found and documented. modern usage. The Procter Collection is currently Some recent taxonomic changes have, for prac- housed at Acadia National Park in Bar Harbor, tical reasons, not been incorporated in our paper. Maine, with annual additions to the fauna being TheseincludethereturnofAcalathus advena made through Entomological BioBlitzes usually (LeConte) to the genus Calathus Stephens, the focused on individual orders (Chandler et al.