Review of the rove beetle species of the subtribe Gyrophaenina Kraatz (Coleoptera, Staphylinidae) from New Brunswick, Canada: new species, provincial records and bionomic information

A comprehensive species review of the genera Gyrophaena Mannerheim and Eumicrota Casey is presented for New Brunswick, Canada. Twenty-four species of Gyrophaena are reported from New Brunswick including two new species, and two species of Eumicrota. Nineteen previously described species are newly recorded in New Brunswick, eight of which are newly reported in Canada. New species are: Gyrophaena meduxnekeagensis Klimaszewski & Webster, sp. n., and G. pseudocriddlei Klimaszewski & Webster, sp. n. New Canadian records are: G. chippewa Seevers, G. corruscula Erichson, G. dybasi Seevers, G. fuscicollis Casey, G. illiana Seevers, G. involuta Casey, G. laetula Casey, and G. lobata Casey. New records for New Brunswick: G. fuscicollis Casey, G. caseyi Seevers, G. chippewa Seevers, E. corruscula Erichson, G. criddlei Casey, G. dybasi Seevers, G. fl avicornis Melsheimer, G. gaudens Casey, G. gilvicollis Casey, G. gracilis Seevers, G. illiana Seevers, G. involuta Casey, G. laetula Casey, G. lobata Casey, G. sculptipennis Casey, E. socia (Erichson), G. subnitens Casey, G. uteana Casey, and G. vitrina Casey. All species are documented by colour habitus images, black/white images of genital structures, and distributional maps. All female genital structures are presented and illustrated here for the fi rst time. A key to the New Brunswick species is provided. ZooKeys 22: 81–170 (2009) doi: 10.3897/zookeys.22.219 www.pensoftonline.net/zookeys Copyright Her Majesty the Queen in Right of Canada. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Launched to accelerate biodiversity research A peer-reviewed open-access journal


Introduction
recorded 62 valid species of Gyrophaena from America north of Mexico. Th e genus is very poorly known in Canada and the only key to identifi cation of species was provided by Seevers (1951) in his North American revision published over half a century ago. Th is outstanding contribution requires updating, particularly at the specifi c level, for the Canadian fauna. Moore and Legner (1975) recorded 13 species from Canada and none from New Brunswick. Campbell and Davies (1991) listed 20 species of Gyrophaena and one of Eumicrota for Canada, and 5 species of the former genus for the province of New Brunswick. Ashe (1984) published a thorough generic revision of the subtribe Gyrophaenina with a review of described subgenera but it lacked species level treatment. Here we present the fi rst comprehensive species review of the genera Gyrophaena and Eumicrota from New Brunswick, Canada. We report 26 species including two new species, 8 new national and 19 new provincial records. Th ere are now 29 species of Gyrophaena and two species of Eumicrota known to occur in Canada, considering new data provided in this contribution, and records provided by Campbell and Davies (1991) and Dollin et al. (2008). Intensive collecting of aleocharine rove beetles in New Brunswick since 2004 by the second author (RPW) has yielded many new species, and new provincial and national records. Th ese records will be published in a series of papers. Th e goal of the present contribution is to describe two new species, publish new collection and bionomic data on Gyrophaena, and to provide a revised key for the identifi cation of species.

Method and conventions
Collection method. Various kinds of mushrooms and forest litter were sifted using the method described by Smetana (1971) and other methods described below. Specimens of Gyrophaena are easily collected from various kinds of mushrooms by breaking the mushrooms into pieces in a plastic box and then collecting the beetles as they move. Th is was the primary method employed for collecting most of the specimens reported in this study.
Specimen preparation. More than 300 adult specimens of Gyrophaena were examined and most specimens were dissected. Th e genital structures were dehydrated in absolute alcohol and mounted in Canada balsam on celluloid microslides and pinned with the specimens from which they originated. Th e photographs were taken using an image processing system (Nikon SMZ 1500 stereoscopic microscope; Nikon Digital Camera DXM 1200F; and Adobe Photoshop software).
Terminology mainly follows that used by Seevers (1951) and Ashe (1984). Th e ventral part of the median lobe of the aedeagus is considered to be the part of the bul- in time and space, and extremely heterogeneous in physical and chemical characteristics (Ashe 1984). Adults of gyrophaenes have developed adaptations to fi nd and live on desirable mushrooms. Th e principal structural adaptations of the adults are modi-1 2 3 1mm fi cations of their mouthparts. Th e beetles feed by "grazing" maturing spores, basidia, cystidea, and hyphae from the hymenium layer of fresh mushrooms and their maxillae are highly modifi ed for this purpose (Ashe 1984). Th e galeal setae form a cap over the apex of the lacinial spore brush for effi cient collecting of material removed from the hymenium. Other adaptations of the gyrophaenes to their ephemeral and unpredictable habitat and food source are a short life cycle and effi ciency in locating and colonizing their mushroom hosts (for details, see Ashe 1984). Adults may also be found in moist forest litter and under logs, which may be an adaptation for survival when few suitable mushroom habitats are available (Ashe 1984). Some adults were collected by one of us (RPW) quite early in the season, which would suggest that some species probably overwinter in litter as adults. Geographic distribution. Sixty-two valid species of Gyrophaena and seven species of Eumicrota are known from America north of Mexico (Ashe 2001); 31 species, including present records, are known to occur in Canada.

Checklist of Gyrophaena Mannerheim species occurring in Canada with the United States records
Conventions. Junior synonyms are indented. Th e United States records, particularly from the states bordering Canada, are also included. Countries and provinces in bold font represent new records. Species follow the taxonomic order established by Seevers (1951). * Holarctic species; † adventive species introduced into North America.
Description. Body length 1.6-2.0 mm, approximately broadly subparallel; head rufous, rufo-piceous to (rarely) black; pronotum rufo-fl avate, sometimes darker basally; elytra fl avate or fl avo-testaceous, mottled with darker brown posterocentral section of disc; abdomen fl avo-testaceous with slightly darker apex. Punctation: vertex of head with at least six umbilicate punctures on each side; pronotum moderately densely punctured; elytra fi nely, irregularly punctate. Microsculpture: fi nely meshed and strong on head and elytra, obsolete on pronotum. Antennae as illustrated ( Bionomics. Macrohabitat: mixed forest, mixed forest with hemlock, hardwood forest, yellow birch (Betula alleghaniensis Britt.) and spruce forest, red spruce forest, mature red maple and red spruce forest. Microhabitat: on gilled mushrooms, on small gilled mushrooms on side of rotten logs, on Pleurotus sp. growing on side of logs. Collecting period: August and September. Collecting method: sifting mushrooms, aspirating, and hand collecting specimens.

III. Gyrophaena (Gyrophaena) laetula species group
Bionomics. Macrohabitat: Mixed forests, mature mixed forests, regenerating mixed forest, red spruce and yellow birch forest, mature red spruce and red maple forest, forested black spruce (Picea mariana (Mill.) BSP) bog with red maple. Microhabitat: on/in gilled mushrooms, on Pleurotus sp. on dead standing Populus tremuloides Michx. Collecting period: June, July, August, and September. Collecting method: sifting mushrooms and aspirating specimens.
Distribution ( Description. Body length 1.7-1.8 mm, stout and compact; head piceous; pronotum rufo-piceous to piceous; elytra uniformly brown or light brown with piceous posterior angles; abdomen rufo-testaceous to dark brown. Punctation: vertex of head with some eight small punctures on each side, pronotum with one conspicuous puncture in median row on each side and with a few scattered punctures elsewhere, elytra fi nely and sparsely punctate. Microsculpture: fi nely meshed throughout. Antennae as illustrated (Fig. 7). Pronotum 1.6 times as wide as long. MALE: tergite 8 with two large and rounded teeth and some small median tuberosities (Fig. 55); sternite 8 pointed apically (Fig. 56). Median lobe of aedeagus with narrowly elongate and tapering tubus and ventrally projecting apex, apical projection small and narrow, directed anteriad in lateral view (Fig. 53), dorsal projection of internal sac narrow and sinuate (Fig. 53). Paramere as illustrated (Fig. 54). FEMALE. Unknown.
Bionomics. Macrohabitat: red maple and red oak (Quercus rubra L.) forest. Microhabitat: on a polypore fungus growing on a log. Collecting period: August. Collecting method: sifting mushrooms and aspirating specimens.
Distribution (Map 5). CANADA: New Brunswick and Nova Scotia, Quebec; UNITED STATES: New Hampshire, New York, and Wisconsin.
Comments. Our specimens agree in all respects with the description and illustrations of this species by Seevers (1951), but tergite 8 of males in our specimens has only 2-4 median small teeth while in Seever's illustration there are 5 small median teeth. We attribute this diff erence to intra-specifi c variation. Description. Body length 1.8-2.0 mm, subparallel; head brown to piceous; pronotum fl avate; elytra brown to dark brown; abdomen rufo-fl avate, apical part of abdomen some-times darker. Punctation: vertex of head with at least seven small umbilicate punctures on each side, pronotum with two weakly defi ned median rows of punctures, elytra fi nely and sparsely punctate. Microsculpture: reticulate throughout, often weakly so on elytra. Antennae as illustrated (Fig. 9). Pronotum 1.3 times as wide as long. MALE: tergite 8 with two narrowly elongate and rounded lateral teeth and two slightly smaller rounded median teeth (Fig. 66); sternite 8 broadly rounded apically (Fig. 67). Median lobe of aedeagus with broad and ventrally angular tubus, apex narrow and acute (Fig. 64), dorsal projection of internal sac short and irregular in shape (Fig. 64). Paramere as illustrated (Fig. 65). FEMALE. Tergite 8 truncate posteriorly (Fig. 69); sternite 8 broadly rounded and slightly emarginate medially (Fig. 70); spermatheca as illustrated (Fig. 68). Bionomics. Macrohabitat: hardwood forest, hardwood forest with hemlock, mature red spruce and red maple forest, and mature mixed forest. Microhabitat: in gilled mushrooms on forest fl oor, small gilled mushrooms on log, on Pleurotus sp. on log. Collecting period: August. Collecting method: sifting mushrooms and aspirating adults.
Bionomics. Macrohabitat: mixed forest, 8.5-year-old regenerating mixed forest, eastern white cedar swamp, mature red spruce, and red maple forest. Microhabitat: on/ in gilled mushrooms on forest fl oor, on log, and on a stump, on small gilled mushrooms on side of decayed log. Th is species has also been found in rotting mushrooms, on orange bracket (polypore) fungus, on bracket fungus on white birch, and on Pleurotus sp. on dead standing Populus tremuloides. Collecting period: June, July, August, and September. Collecting method: sifting mushrooms, aspirating, and hand collecting specimens.
Bionomics. Macrohabitat: mixed forest, white and red spruce forest, eastern white cedar swamps, yellow birch and spruce forest, on ridge with oak in silver maple forest, and red oak and red maple forest. Microhabitat: on/in gilled mushrooms on forest fl oor, on Russula virescens. Collecting period: August and September. Collecting method: sifting mushrooms, aspirating, and hand collecting specimens.
Distribution (Map 12). CANADA: British Columbia, New Brunswick, Ontario; UNITED STATES: Michigan. punctures on each side, pronotum with two irregular median rows of large punctures and scattered punctures elsewhere, elytra with fi ne, sparse punctures. Microsculpture: reticulate throughout except for pronotum. Antennae as illustrated (Fig. 16). Pronotum 1.4 times as wide as long. MALE: tergite 8 with two lobe-shaped lateral teeth and usually two (but occasionally one or three) small median teeth, apical margin of disc emarginate (Fig. 109); sternite 8 broadly rounded apically with minute apical emargination (Fig. 110). Median lobe of aedeagus with elongate and slightly arcuate tubus bearing two characteristic (long and short) ventral projections (Fig. 107), fl agellum sinuate and projecting externally (Fig. 107). Paramere as illustrated (Fig. 101)   Etymology. Named by adding prefi x "pseudo" to the specifi c name "criddlei". Gyrophaena criddlei Casey is externally similar to this new species.

Gyrophaena (Gyrophaena) criddlei Casey
Description. Th is species is externally and genitally similar to G. criddlei but may be easily distinguished by its smaller body size, short elytra, and the characteristic shape of male tergite 8 and median lobe of aedeagus with diff erently shaped projections on the tubus.
Bionomics. Macrohabitat: red oak and red maple forest, mature red spruce and red maple forest. Microhabitat: on stalked polypore mushroom on forest fl oor, one specimen in gilled mushroom on forest fl oor. Collecting period: September. Collecting method: sifting mushrooms and aspirating specimens.
Bionomics. Macrohabitat: Mixed forest, hardwood forest, red spruce forest, red spruce and yellow birch forest, mature red spruce and red maple forest, mature red spruce and eastern white cedar forest, oak and red maple forest. Th is species has been collected in association with grasses. Microhabitat: on fresh gilled mushrooms on forest fl oor, in decaying gilled mushroom (one occurrence), and on Pleurotus sp. on log. Collecting period: August, September, and October. Collecting method: sifting mushrooms, aspirating, and hand collecting specimens.
Bionomics. Macrohabitat: mixed forest and hardwood forest with hemlock. Th is species has also been found in spruce/pine/fi r forests (80-120 years old), old deciduous forest, hemlock forest (120+ years old), and in a meadow. Microhabitat: in/on gilled mushrooms and small gilled mushrooms on log. Th is species has also been reported from a rotting mushroom, on Heterobasidium annosum, on white fungus on a white birch, on red maple log and in a decayed log. Collecting period: July to September. Collecting method: sifting mushrooms, aspirating, and hand collecting.
Bionomics. Macrohabitat: mixed forest, hardwood forest, rich Appalachian hardwood forest, fl ood plain forest, and eastern white cedar swamps, and mature red spruce and red maple forest. Microhabitat: on gilled mushrooms on forest fl oor, on small gilled mushrooms on side of log, on bracket fungi, and on Trametes hirsuta on a poplar log. Collecting period: June, July, August, and September. Collecting method: sifting mushrooms and aspirating specimens.
Body length 1.4-1.7 mm, narrowly oval; head piceous; pronotum brown to piceous; elytra brown or reddish-brown with piceous sutural region and posterior angles; abdomen reddish-brown, apical part often darker and piceous. Punctation: vertex of head with at least six moderately-sized umbilicate punctures on each side, pronotum with six punctures in two confused median rows and additional punctures elsewhere, elytra with fine, sparse punctures. Microsculpture: reticulate throughout except for pronotum, most evident on head. Antennae as illustrated (Fig. 22). Pronotum 1.5 times as wide as long. MALE: tergite 8 with two long and narrow lateral teeth and two very small median teeth close to each other (Fig. 147); sternite 8 rounded apically (Fig. 148). Median lobe of aedeagus with long and narrowly produced tubus bearing large angular swelling in the middle of ventral margin, apex with subapical minute projection (Fig. 145), apical projection of internal sac elongate and irregular in shape (Fig. 145). Paramere as illustrated (Fig. 146). FEMALE. Tergite 8 truncate apically (Fig. 150); sternite 8 rounded apically and slightly pointed (Fig. 151); spermatheca as illustrated (Fig. 149). Bionomics. Macrohabitat: mixed forest, on ridge with red oaks surrounded by a silver maple forest, mature red spruce and red maple forest. Microhabitat: in/on gilled mushrooms and on bracket fungi. Collecting period: June, August and September. Collecting method: sifting mushrooms and aspirating specimens.
Distribution (Map 19). CANADA: New Brunswick, Quebec, and British Columbia; UNITED STATES: California, Colorado, and Utah.
Body length 1.4-1.7 mm, narrowly oval; head piceous; pronotum dark rufo-testaceous to piceous; elytra reddish-brown with piceous sutural region and posterior angles; abdomen yellowish-or reddish-brown, apical portion often darker and piceous. Punctation: vertex of head with at least six moderately-sized umbilicate punctures on each side, pronotum with six punctures in two confused median rows and additional punctures elsewhere, elytra with fi ne and sparse punctures. Microsculpture: reticulate throughout, weaker on pronotum, most evident on head. Antennae as illustrated (Fig. 23). Pronotum 1.5 times as wide as long. MALE: tergite 8 with two long and narrow lateral teeth and with two (rarely three) minute median teeth [smaller and more sharply pointed than those of G. uteana] (Fig. 154); sternite 8 rounded apically (Fig. 155). Median lobe of aedeagus with long and narrowly produced tubus which is lacking angular swelling in the middle of ventral margin, apex with subapical minute projection (Fig. 152), apical projection of internal sac elongate and irregular in shape (Fig. 152). Paramere as illustrated (Fig. 153). FEMALE. Tergite 8 truncate apically (Fig. 157); sternite 8 rounded apically and slightly pointed (Fig. 158); spermatheca as illustrated (Fig. 156).
Bionomics. Macrohabitat: mixed forest and conifer forest. Microhabitat: on/in gilled mushrooms, on small gilled mushroom on log, on Pleurotus sp. on dead standing Populus tremuloides, on polypore fungi, and on bracket fungi. Th is species has also been found on Cantharellus deliciosus. Collecting period: June, July, and August. Collecting method: sifting mushrooms, aspirating, and hand collecting.
Distribution (Map 20). CANADA: New Brunswick, Ontario, and Quebec; UNITED STATES: Illinois, Indiana, Massachusetts, Michigan, Pennsylvania, and Wisconsin. Description. Body length 2.0-2.7 mm, broadly subparallel; head piceous; pronotum light to dark reddish-brown; elytra fl avo-testaceous or reddish-brown with some small darker spots laterally and in scutellar region; abdomen reddish-brown, apical portion often darker. Punctation: vertex of head with at least 10 moderately-sized coarse punctures on each side, pronotum with two median rows of coarse punctures and additional punctures scattered elsewhere, elytra with fi ne, sparse punctures. Microsculpture: reticulate throughout but weaker on elytra. Antennae as illustrated (Fig. 24). Pronotum 1.4 times as wide as long. MALE: tergite 8 with two small lateral teeth and usually with two minute median teeth (occasionally only one tooth present or the teeth reduced to small tuberosities), apical margin slightly emarginate medially (Fig. 161); sternite 8 rounded apically (Fig. 162). Median lobe of aedeagus with complex tubus, its apex with subapical hook-like projection (Fig. 159), apical projection of internal sac elongate and truncate apically (Fig. 159). Paramere as illustrated (Fig. 160). FEMALE. Tergite 8 truncate apically (Fig. 164); sternite 8 slightly produced apically (Fig. 165); spermatheca as illustrated (Fig. 163). Bionomics. Macrohabitat: Mixed forest, hardwood forest, rich Appalachian hardwood forest, red oak and red maple forest, 8.5-year-old regenerating mixed forest, mature red spruce and red maple forest. Microhabitat: on/in gilled mushrooms on forest fl oor, in gilled mushroom on stump, on Pleurotis species growing on a log, and in Porodaedalea piceina (Peck) Niemalä on dead standing beech tree. Th is species has also been found in decaying mushrooms, on white fungus on white birch. Collecting period: June, July, August, and September. Collecting method: sifting mushrooms, aspirating, and hand collecting.

XII. Gyrophaena (Phaenogyra) strictula
Distribution (Map 22). CANADA: New Brunswick, Ontario, and Manitoba; UNITED STATES: Illinois, Kansas, Maine, Michigan, Minnesota, Missouri, New York, and Wisconsin. Comments. Gyrophaena subnitens is similar externally to G. meduxnekeagensis but may be distinguished from the latter species by paler, light-yellow antennae, short elytra (Fig. 25), and the shape of tubus of median lobe of aedeagus without basal swelling (Fig. 166). From Palaearctic G. polita (Gravenhorst) (Figs 27,(180)(181)(182)(183), it diff ers in having a lighter body coloration, shorter and more rapidly converging postocular temples of head, and broader pronotum (Fig. 25). Th e genital structures are generally similar in these species. Etymology. In reference to Meduxnekeag in the Meduxnekeag Valley Nature Preserve where the type and most of the paratypes were collected. This private protected area, adjacent to the Meduxnekeag River and established by the Meduxnekeag River Valley Association, is home to an unsually high number of rare plant and Coleoptera species. The word "Meduxnekeag" is a Maliseet Indian one meaning "rough or rocky at its mouth" presumably applied to the mouth of the river itself.
Bionomics. Macrohabitat: hardwood forest, mature mixed forest, mixed forest, wet alder (Alnus sp.) swamp. Microhabitat: on small white gilled mushroom on side of decaying log, and on Pleurotus sp. growing on dead standing Populus tremuloides, partially dried polypore fungus at base of dead standing Populus tremuloides. Collecting period: May and July. Collecting method: sifting mushrooms and aspirating specimens.
Bionomics. Macrohabitat: Silver maple swamp, fl ood plain forest, red oak forest. Microhabitat: on Trametes hirsuta (Wolfen) Pilat growing on Populus tremuloides log (two sites), partially dried Pleurotus sp. on dead standing trembling aspen (one specimen) and one specimen from nest contents of barred owl (Strix varia Barton) nest box with small chicks. Collecting period: May, June, and August. Collecting method: sifting mushrooms and aspirating specimens.
Bionomics. Macrohabitat: hardwood forest, mixed forest, silver maple forest, fl ood plain forest with butternut trees, mature red spruce and red maple forest, and oak forest. Th is species has also been found in a conifer forest, red spruce forest (80-120 years old), hemlock forest (120+ years old), hemlock/balsam fi r/black spruce forest, and stream margin forest. Microhabitat: on/in Pleurotus sp. on logs and dead standing trembling aspens, in partially dried Pleurotus sp. on dead standing sugar maple, in slightly decayed polypore fungi on log, in decaying polypore fungi on dead standing spruce, in bracket fungi, in/on gilled mushrooms on logs and forest fl oor, on Climatodon septentrionale growing on dead standing sugar maple. Also found on Heterobasidium annosum on Picea rubens. Collecting period: June, July, August, and September. Collecting method: sifting mushrooms, aspirating and hand collecting specimens.