New Staphylinidae (Coleoptera) records with new collection data from New Brunswick and eastern Canada: Tachyporinae

Abstract Twenty-three species of Tachyporinae are newly recorded from New Brunswick. This brings the total number of Tachyporinae known from the province to 70. Lordithon campbelli Schülke is newly recorded for Canada and we provide the first documented records of Tachinus addendus Horn and Tachinus frigidus Erichson for New Brunswick. Collection and habitat data are presented and discussed for each species. A list of Tachyporinae species currently known from the province of New Brunswick is presented.


Introduction
Intensive collecting of rove beetles (family Staphylinidae) in New Brunswick by the first author since 2003 and records from by-catch samples obtained during a study to develop a general attractant for the detection of invasive species of Cerambycidae have yielded many new provincial and national records. These are being published in a series of papers, each focusing on one or more subfamilies. This paper treats the subfamily Tachyporinae. The Tachyporinae of Canada and North America are fairly well known taxonomically thanks to various revisions by J.M. Campbell; Tachinus (Campbell 1973(Campbell , 1975b(Campbell , 1988, Coproporus and Cilea (Campbell 1975a), Sepedophilus (Campbell 1976), Tachyporus (Campbell 1979), Carphacis (Campbell 1980), Lordithon (Campbell 1982), Mycetoporus and Ischnosoma (Campbell 1991), Nitidotachinus (Campbell 1993a), and Bryoporus and Bryophacis (Campbell 1993b).
Tachyporinae can be found in a wide variety of habitats. Tachinus species are often found in decaying organic materials such as dung, rotting mushrooms, carrion, and compost, although some species are found in leaf litter and moist debris near streams, e.g., Tachinus limbatus Melsheimer (Campbell 1973). Nitidotachinus species are usually found in leaf litter or moss near streams, often in cool, shaded sites (Campbell 1993a). Tachyporus, Mycetoporus, Bryoporus, Bryophacis, and Ischnosoma species are usually associated with various kinds of litter and moss in forests and various wetland types, depending on the species (Campbell 1979(Campbell , 1991(Campbell , 1993b. Tachyporus species are often swept from vegetation in fields and other open habitats (Campbell 1979). Larvae and adults of Lordithon are associated with mushrooms and are active predators of Diptera larvae (Campbell 1982). Some Sepedophilus species are found in rotting wood, under loose bark, and in decaying and often moldy organic materials, such as rotting leaves (Campbell 1976); others are associated with polypore fungi or fleshy fungi on trees, depending on the species (Newton et al. 2000). Some species may be mycetophagous (Newton et al. 2000). Our only Coproporus species is subcortical (Campbell 1975b). However, in general, little is known about the biology of our North American Tachyporinae.
Thirty-six species of Tachyporinae were reported from New Brunswick by Campbell and Davies (1991). Nine species were added to the faunal list in revisions by Campbell (1991Campbell ( , 1993a and from general surveys by Klimaszewski et al. (2005) and Majka and Klimaszewski (2008). Majka et al. (2011) reported Tachinus addendus Horn and Tachinus frigidus Erichson as occurring in New Brunswick but did provide any supporting references or data for the records. Here, we report an additional 23 species, bringing the total number of Tachyporinae known from New Brunswick to 70.

Methods and conventions
The following records are based on specimens collected as part of a general survey by the first author to document the Coleoptera fauna of New Brunswick and from bycatch samples obtained during a study to develop a general attractant for the detection of invasive species of Cerambycidae.

Collection methods
Various collection methods were employed to collect the Tachyporinae reported in this study. Details are outlined in Campbell (1973) and Webster et al. (2009, Appendix). See Webster et al. (2012) for details of the methods used for deployment of Lindgren 12-funnel traps and sample collection. A description of the habitat was recorded for all specimens collected during this survey. Locality and habitat data are presented exactly as on labels for each record. This information, as well as additional collecting notes, is summarized and discussed in the collection and habitat data section for each species.

Specimen preparation
Examples of males of most species were dissected to confirm their identity. The genital structures were dehydrated in absolute alcohol and mounted in Canada balsam on celluloid microslides and pinned with the specimens from which they originated. tia, Prince Edward Island) of Canada, including Lordithon campbelli Schülke, which is newly recorded for Canada. The first documented records of T. addendus and T. frigidus from New Brunswick are provided. This brings the total number of species known from New Brunswick to 70 (Table 1).

Species accounts
All species below are newly recorded for New Brunswick, Canada. Species followed by ** are newly recorded from the Maritime provinces; species followed by *** are newly recorded for Canada. The classification of the Tachyporinae follows Bouchard et al. (2011). son Falls, Bell Forest, 46.2200°N, 67.7231°W, 16.IX.2006, R. P. Webster, mature hardwood forest, on fleshy polypore fungi on beech log (8 ♂, 7 ♀, NBM, RWC); same locality and forest type, 23-28.IV.2009, 14-20.V.2009, 20-26.V.2009, 8-16. VI.2009 Campbell (1976) reported that most specimens of this species were collected from under bark. Others were found in mushrooms, Polyporus betulinus (Bull.) Fr., on dead logs, dead beech (Fagus grandifolia Ehrh.), and tree trunks and in tree holes. In New Brunswick, specimens were found on fleshy polypore fungi on a beech log, in Piptoporus betulinus (Bull.) P. Karst. (birch polypore), on Polyporus varius Fr., and in a well-rotted log. This species was commonly collected in Lindgren funnel traps in various forest types; mature hardwood forests with sugar maple (Acer saccharum Marsh.) and beech, old red oak (Quercus rubra L.) forest, old-growth hardwood forest with sugar maple and yellow birch (Betula alleghaniensis Britt.), 110-year-old red spruce (Picea rubens Sarg.) forest with red maple (Acer rubrum L.), old red pine (Pinus resinosa Ait.) forest, and an old mixed forest. Adults were collected during April, May, June, July, August, and September.
Distribution in Canada and Alaska. ON, QC, NB, NS (Campbell 1976;Bishop et al. 2009 Collection and habitat data. Campbell (1976) reported that adults were frequently collected from rotten wood, from deep layers of decaying leaves, and from bracket fungi and mushrooms. Most specimens from New Brunswick were collected from fleshy polypore fungi and bracket fungi on standing dead trees and logs. Some adults were also collected from Lindgren funnel trap samples. Two specimens were collected from Polyporus varius Fr. on a rotten log. This species was found in sugar maple and beech forests, a red spruce forest, an old-growth hemlock (Tsuga canadensis (L.)) forest, and mixed forests. Adults were collected during June, July, August, and September.
Distribution in Canada and Alaska. ON, QC, NB, NS (Campbell 1976;Bishop et al. 2009 Collection and habitat data. In the United States, this species has been collected from under bark, under a brush pile, and by sifting humus (Campbell 1976). The New Brunswick specimens were collected from the inside of well-rotted, fungus-covered logs and from polypore fungi on dead, standing Populus sp. This species was found in an old-growth eastern white cedar (Thuja occidentalis L.) swamp and in mature to old mixed forests. Adults were collected during June and August. Collection and habitat data. Campbell (1976) reported this species from mushrooms. In New Brunswick, specimens were collected from polypore fungi on logs, from under bark, and among moldy corncobs and cornhusks. This species was found in red oak and red maple forests, old silver maple (Acer saccharinum L.) forests, and near a mixed forest. Adults were collected during May, August, and September.

Distribution in Canada and
Distribution in Canada and Alaska. ON, NB. (Brunke and Marshall 2011). Campbell (1976) did not report this species from Canada. However, there are two specimens of this species in the Canadian National Collection from Turkey Point, Ontario collected in 1975 that first establish this species as a member of the Canadian fauna. Brunke and Marshall (2011) reported an additional record from Rondeau Provincial Park, Ontario. In the United States, this species occurs from New Hampshire west to Iowa and south to Florida (Campbell 1976 Collection and habitat data. This species has been collected from dung, rotting mushrooms, deciduous leaf litter, and pine duff (Campbell 1973). Most adults from New Brunswick were collected from decaying mushrooms in hardwood and mixed forests. One individual was collected from decaying sea wrack under alders (Alnus sp.) on the upper margin of a sea beach, another was found in moose dung. Adults were collected during May, June, July, August, and September.
Distribution in Canada and Alaska. MB, ON, QC, NB, NS (Campbell 1973(Campbell , 1988. Tachinus addendus was listed as occurring in New Brunswick by Majka et al. (2011)  Collection and habitat data. Little was previously known about the habitat associations of this species other than some specimens having been collected from mushrooms (Campbell 1973). The two specimens from New Brunswick were collected from gilled mushrooms near a silver maple swamp during September. Campbell (1973) commented that the late period of annual activity (September and October) was quite different from other species occurring in eastern North America.
Distribution in Canada and Alaska. ON, QC, NB (Campbell 1973 Collection and habitat data. Tachinus fimbriatus is usually collected from rotting mushrooms (Campbell 1973), as were the two specimens from New Brunswick. Adults were collected in a mature mixed forest during September.
Distribution in Canada and Alaska. ON, QC, NB, NS (Campbell 1973(Campbell , 1988 Campbell (1973Campbell ( , 1988 reported that most specimens of this northern transcontinental species were collected from under animal dung or decaying mushrooms. Adults were also collected from the mouth of animal burrows, in leaf litter and other kinds of decaying organic matter. In Alberta, T. frigidus was considered to be a mature forest (conifer-dominated) specialist (Pohl et al. (2007). The only specimen from New Brunswick was collected from a decaying fleshy mushroom during September in a mature, coastal red spruce forest.
Distribution in Canada and Alaska. AK, YT, NT, BC, AB, MB, ON, QC, NB, NS, LB (Campbell 1973 Collection and habitat data. Specimens of this species from New Brunswick were collected from a decaying fleshy polypore mushroom on a standing, dead spruce in a mature red spruce forest, from a decaying Climacodon septentrionale (Fr.) P. Karst. on a dead, standing yellow birch in a recent clearcut (boreal forest area), and from a (fresh) Climacodon septentrionale (Fr.) P. Karst. on a living sugar maple in an old-growth northern hardwood forest. Four individuals were collected in company with Lordithon niger (Gravenhorst) from a decaying fleshy polypore fungus on a standing, dead Populus sp. in a hardwood forest (sugar maple and American beech) in Saint-Raphaël (15.VII.2006), Quebec (Webster, unpublished). One specimen from Tennessee (USA) was sifted from leaf litter. Little was previously known about the habitat and biology of this rare species. Campbell (1973) suggested that this species lived in some restricted habitat, such as mammal burrows. The habitat data above suggest that this species may be associated with decaying fleshy polypore or polypore-like fungi on standing dead and living trees. Adults were collected during July and August.
Distribution in Canada and Alaska. QC, NB, NS (Campbell 1973(Campbell , 1988 Little is known about the habitat associations of this species. Two adults of this rare species were collected from deciduous leaf litter along a small stream and from alder litter on a lake margin (Campbell 1975b). Others were taken from flight intercept traps (Campbell 1988). In New Brunswick, adults were collected from under bark in a wood pile, from leaf litter near a stream, and with a net during an evening flight. Some individuals were collected in Lindgren funnel traps deployed in old red oak forest, an old-growth red pine forest, an old-growth white spruce and balsam fir forest, and an old-growth northern hardwood forest. Adults were collected during April, June, July, August, and September but most during April.
Distribution in Canada and Alaska. AB, ON, QC, NB (Campbell 1973(Campbell , 1975b(Campbell , 1988Pohl et al. 2007). Collection and habitat data. Campbell (1991) reported this species from river banks, flood debris on rivers, under logs, and decaying vegetation. Most New Brunswick specimens were found in litter in crotches of silver maples with multiple trunks in an old silver maple swamp (floodplain forest) early in April. This habitat was probably an overwintering site for this species. Many other staphylinid adults of various species were found in the debris in these tree crotches. Other individuals were sifted from leaf litter at the bases of large silver maples. Adults were collected during April and May.

Tachyporus lecontei
Distribution in Canada and Alaska. BC, AB, SK, MB, ON, QC, NB (Campbell 1991). Collection and habitat data. Campbell (1991) reported this species from a variety of habitats, including forest leaf litter, fungi on an old tree stump, among grass roots, damp moss near a pond, under a log on a lakeshore, and from Microtus pennsylvanicus (Ord) nests. In New Brunswick, specimens were found in a various microhabitats including leaf litter and moss near brooks, margins of vernal ponds and a river, leaf litter at bases of trees, and among decaying corncobs and cornhusks. This species was found in a Carex marsh, mature hardwood forests, silver maple swamps, mixed forests, and river and brook margins. Adults were collected during April, May, June, and September and were active early in the spring on bare patches around bases of trees when over 30 cm of snow was still on the ground.

Tachyporus maculicollis
Distribution in Canada and Alaska. BC, AB, SK, MB, ON, QC, NB, NS (Campbell 1979 Collection and habitat data. This rare species has been collected from the fallen nest of a squirrel and a Berlese sample of decaying moldy material from the base of a tree (Campbell 1979). The three specimens from New Brunswick were captured in Lindgren funnel traps deployed in a 110-year-old red spruce forest and an old (120-to 180-year-old) red pine forest. Adults were collected during May.
Distribution in Canada and Alaska. ON, QC, NB (Campbell 1979 Collection and habitat data. This uncommon species was reported from dead swamp grass, among leaves, from moss, and from an entrance to a Marmota burrow by Campbell (1979). In New Brunswick, T. pulchrus was found in various wetland habitats, including eastern white cedar swamps, red maple swamps, an open grassy alder swamp, a Carex marsh/fen, a Carex marsh/flowage, and a stream margin near an eastern white cedar swamp. Adults occurred in leaves and moss, grass litter and moss on hummocks, sphagnum hummocks, leaf litter at bases of trees, and in grass litter and were collected by sifting or treading. This species was collected during April, May, June, and September (most during April).
Distribution in Canada and Alaska. MB, ON, NB (Campbell 1979 Collection and habitat data. This is a hygrophilous species in both Europe and North America and is usually found in marshes and bogs in moss (especially sphagnum) and debris (Campbell 1991). In New Brunswick, this species was found in various wetland habitats, including a forested black spruce (Picea mariana (Mill.) B.S.P.) bog, an eastern white cedar swamp, sedge (Carex) fens and marshes, freshwater marshes, stream margins, the margin of a vernal pond, and in an abandoned North American beaver Castor canadensis Kuhl) lodge occupied by muskrats (Ondatra zibethicus L.).
Adults occurred in moist leaves, sphagnum, litter, and moss, in Carex hummocks, and in debris in the roof of a beaver lodge. In New Brunswick, this species was collected during March, April, May, and June.
Distribution in Canada and Alaska. ON, QC, NB (Campbell 1991). Campbell (1991) considered the distribution and habitat of this species in the Nearctic region unusual for a Holarctic species because of the specialized habitat preferences (found in sphagnum and debris in marshes and bogs) and pattern of distribution (restricted to Ottawa Valley of Ontario and Quebec). He suggested that the species may have been overlooked by most collectors and may actually have a broader distribution in North America than the records indicate. Klimaszewski et al. (2010)  Collection and habitat data. Campbell (1991) reported that this species was often found in drier habitats than other members of the Pictum group of species. Adults were reported from various kinds of forest litter in pine, hardwood, and mixed pine and hardwood forests as well as cypress forests (Campbell 1991). In New Brunswick, this species was most often found in and near wetland habitats, such as calcareous fens, Carex marshes, tamarack (Larix laricina (Du Roi) Koch) bogs, alder swamps, and red maple swamps. One adult was found near a stream in a mature hardwood forest. Adults occurred in moss and sphagnum hummocks, moss, leaf and grass litter at bases of trees, and Carex hummocks. Adults were collected during April, May, June, and September.
Distribution in Canada and Alaska. ON, NB (Brunke and Marshall 2011). In the United States, I. flavicolle occurs throughout the southeast north to New Hampshire along the eastern seaboard (Campbell 1991). Collection and habitat data. Campbell (1991) reported this species from various of wetland habitats as well as forests. Adults were taken from flood debris along rivers, margins of beaver ponds, beaver lodges, muskrat nests, mallard (Anas platyrhynchos L.) nests, moss near seepage areas, leaf litter along margins of marshes, streams, and bogs, and various kinds of grass and leaf litter from conifer and deciduous forests (Campbell 1991). Recent studies in Alberta (mid-boreal ecoregion) by Buddle et al. (2006) indicated that this species was associated with old (>70-year-old) fire-origin, mixed wood forest stands. Later, Pohl et al. (2007) reported that this species was also associated with regenerating mixed wood stands in the western foothills of Alberta. The specimens from New Brunswick were collected in moldy conifer duff and fleshy gilled mushrooms in a small balsam fir (Abies balsamea (L.) Mill.) (regenerating) stand adjacent to a hardwood forest. Campbell (1991) reported most specimens of this species were taken in June to September. The specimens from New Brunswick were collected during May and September.
Distribution in Canada and Alaska. AK, YT, NT, BC, AB, SK, MB, ON, QC, NB, NS, LB, NF (Campbell 1991 Collection and habitat data. Little is known about the habitat requirements of this rare species. Campbell (1982) reported that adults have been taken on rotting gilled mushrooms. Most of the specimens from New Brunswick were collected from the fleshy polypore fungus, Bjerkandera adusta (Willd.) P. Karsten growing on a dead, standing American beech tree and a beech log. One individual was found in a mass of Pholiota sp. mushrooms at the base of a dead, standing Populus sp. Another individual was captured in a Lindgren funnel trap. Adults were found in mature hardwood and adjacent mixed forests, and in an old-growth white spruce and balsam fir forest. This species was collected during September in New Brunswick.
Distribution in Canada and Alaska. AK, BC, AB, ON, PQ, NB, NS (Campbell 1982;Campbell and Davies 1991). This northern species has a very broad distribution from Alaska to Nova Scotia, but with large distributional gaps between known localities (Campbell 1982 Collection and habitat data. Little was previously known about the habitat associations of this rare species (Campbell 1982). In New Brunswick, adults of Lordithon quaesitor were most frequently found on Bjerkandera adusta (Willd.) P. Karsten (a fleshy polypore fungus) on the side of beech logs and standing dead beech trees in mature hardwood forests. This species was also found in Pleurotus sp., among a group of Polyporus varius Fr. on a large sugar maple log and on a dead standing American beech in an old-growth hardwood forest with sugar maple and yellow birch, in polypore fungi on sides of logs, in Pleurotus sp. on a dead standing balsam poplar (Populus balsamifera L.) tree, and in a gilled mushroom on the forest floor of an old-growth northern hardwood forest with sugar maple and yellow birch. One adult was collected in a Lindgren funnel trap. Some adults were collected in a mature red spruce forest, a mixed forest, and an alder swamp adjacent to a mixed forest. Lordithon quaesitor was sometimes found together with L. niger, L. axillaris, and L. longiceps and probably has a similar biology to those species. Adults were collected during May, June, July, August, and September.
Distribution in Canada and Alaska. ON, QC, NB, NS (Campbell 1976;Campbell and Davies 1991;Bishop et al. 2009 Collection and habitat data. Little was previously known about the habitat associations of this rare species. One specimen from Quebec was collected from large gilled mushrooms on the side of a log (Campbell 1982). Four specimens of this species were collected from Pleurotus sp. mushrooms on the side of Populus logs in a hardwood forest in Saint-Raphaël (15.VII.2006), Quebec (Webster, unpublished). Most specimens from New Brunswick were collected from Pleurotus sp. mushrooms on standing dead Populus sp. trees or on the side of logs in mature hardwood forests with sugar maple and beech. Some adults were also collected from a large orange-gilled mushroom on the side of a log. One individual was collected from inside a Laetiporus sulphureus (Fr.) Murr. (Polyporaceae). These data suggest that this species may be specialized on Pleurotus sp. and other large gilled mushrooms that grow on standing dead trees or logs. Campbell (1982) suggested that this species, like L. niger and the European L. bicolor (Gravenhorst), may be associated with old-growth hardwood forests. Adults of L. axillaris were collected during June, July, and August in New Brunswick.
Distribution in Canada and Alaska. QC, NB (Campbell 1982).  Campbell (1982) reported collecting adults from various gilled and pore mushroom species. In New Brunswick, adults were collected from gilled mushrooms on forest floor and on Pleurotus sp. on a dead, standing Populus sp. Adults were collected during June and July.
Distribution in Canada and Alaska. NB (first Canadian record). In the United States, this species (as L. angularis (Saches) in Campbell 1982) is distributed from Massachusetts to Florida, west to Missouri (Campbell 1982). This species probably occurs in intervening areas between Massachusetts and New Brunswick. Collection and habitat data. Nothing was previously known about the habitat associations of this rare species (Campbell 1982). The New Brunswick specimens were collected from Bjerkandera adusta (Willd.) P. Karsten and Porodaedalea sp. (both are fleshy polypore fungi) on a beech log, or on dead, standing beech trees in a mature to old-growth and predominantly hardwood forest. One individual was captured in a Lindgren funnel trap in a mature to old red oak forest. Several specimens of this species were found in company with T. schwarzi in a decaying fleshy polypore fungus on a standing, dead Populus sp. in a hardwood forest in Saint-Raphaël (15.VII.2006), Quebec (Webster, unpublished). Adults from New Brunswick were collected during June, August, and September. The habitat data above suggest that this species might be specialized on fleshy polypore fungi and related species that grow on dead standing trees or logs. Campbell (1982) noted that this species appeared to be becoming increasingly rare and suggested that it might be associated with old-growth hardwood forests, which are disappearing from most of eastern North America. However, more sampling should be done in forests of various ages to establish if this species is indeed an old-growth associate.
Distribution in Canada and Alaska. ON, QC, NB (Campbell 1982 Collection and habitat data. Campbell reported that most adults of this species were collected along stream and lake margins. In Alberta, this species was associated with mature forests (Pohl et al. (2007). The New Brunswick specimen was collected from moss on a hummock at the base of a tree in an old-growth eastern white cedar swamp during May.
Distribution in Canada and Alaska. AK, YT, AB, BC, SK, ON, QC, NB, LB, NF (Campbell 1991 Collection and habitat data. Adults of M. rugosus have been collected from a wide variety of moist (often deep and moldy) litter and moss, including both decidu-ous and conifer litter of various species in forested habitats, as well as lake, stream, and river margins (Campbell 1991). New Brunswick specimens were collected from similar habitats, most frequently from moss and litter near brooks in old-growth eastern white cedar swamps. Adults were also collected from drift material and oak leaf litter along a lakeshore and from sphagnum in a black spruce forest. Adults from New Brunswick were collected during May and June.
Distribution in Canada and Alaska. AK, NT, YT, BC, AB, SK, MB, ON, QC, NB, NS, LB (Campbell 1991) New Brunswick Wildlife Trust Fund for funding various insect surveys over the past 7 years, and the Meduxnekeag River Association for permission to sample beetles at the Meduxnekeag Valley Nature Preserve (which includes the Bell Forest). The New Brunswick Department of Natural Resources (Fish and Wildlife Branch) is thanked for issuing permits for sampling in the Protected Natural Areas and for logistical support. Survey work in the Jacquet River Gorge and Caledonia Gorge Protected Natural Areas was organized through the New Brunswick Museum with external funding from the New Brunswick Environmental Trust Fund, Salamander Foundation, and the New Brunswick Wildlife Trust Fund.