Photographic key to the Pseudoscorpions of Canada and the...

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Canadian Journal of Arthropod Identification No.12 (January 2011) BRUNKE ET AL. doi: 10.3752/cjai.2011.12 1 Staphylinidae of Eastern Canada and Adjacent United States. Key to Subfamilies; Staphylininae: Tribes and Subtribes, and Species of Staphylinina Adam Brunke*, Alfred Newton**, Jan Klimaszewski***, Christopher Majka**** and Stephen Marshall* * University of Guelph, 50 Stone Road East, School of Environmental Sciences, 1216/17 Bovey Building, Guelph, ON, N1G 2W1. [email protected], [email protected]. ** Field Museum of Natural History, Zoology Department/Insect Division, 1400 South Lake Shore Drive, Chicago IL, 60605. [email protected]. *** Laurentian Forestry Centre, 1055, rue du P.E.P.S., Stn. Sainte-Foy Québec, QC, G1V 4C7. [email protected] **** Nova Scotia Museum, 1747 Summer St., Halifax, NS, B3H 3A6. [email protected]. Abstract. Rove beetles (Staphylinidae) are diverse and dominant in many of North America’s ecosystems but, despite this and even though some subfamilies are nearly completely revised, most species remain difficult for non-specialists to identify. The relatively recent recognition that staphylinid assemblages in North America can provide useful indicators of natural and human impact on biodiversity has highlighted the need for accessible and effective identification tools for this large family. In the first of what we hope to be a series of publications on the staphylinid fauna of eastern Canada and the adjacent United States (ECAS), we here provide a key to the twenty-two subfamilies known from the region, a tribe/subtribe level key for the subfamily Staphylininae, and a species key to the twenty-five species of the subtribe Staphylinina. Within the Staphylinina, the Platydracus cinnamopterus species complex is defined to include P. praetermissus Newton spec. nov., P. cinnamopterus (Gravenhorst) and P. zonatus (Gravenhorst). Lectotypes are designated for Staphylinus cinnamopterus Gravenhorst, Staphylinus zonatus Gravenhorst and Staphylinus badius Mannerheim. One new Canadian record, one new record from eastern Canada, and thirty-nine new provincial or state records are presented. Introduction Rove beetles (Coleoptera: Staphylinidae) constitute the largest family of insects worldwide, with more than 55,440 described species (Grebennikov and Newton 2009) found in a great diversity of terrestrial and periaquatic habitats. In Canada, more than 1400 species are known and some large subfamilies (Staphylininae, Tachyporinae) have been nearly completely revised. An excellent synthesis of the staphylinid literature is given by Thayer (2005); however, our understanding of staphylinid ecology and habitat requirements is still very limited. Recent work has revealed that staphylinids are dominant organisms in Canadian forest ecosystems (Paquin and Duperre 2001) and because many species require continuous, mature or old growth stands, the composition of their species assemblages effectively communicates the degree of natural or human impact upon these systems (Pohl et al. 2008). In a recent review of the use of beetles in conservation, New (2010) highlighted the critical importance of species-level identifications in surveys aiming to document changes to ecosystems via human development or climate change. The continued ability of insect surveys to address important ecological and conservation-themed questions depends primarily on the correct identification of specimens, which in turn depends on the availability of effective keys. Although several recent works such as Klimaszewski (2000) and Newton et al. (2000) include keys to the genus or subfamily level, accompanied by line drawings of important characters, keys to most staphylinid groups are poorly illustrated and cover broad geographic areas (e.g. North America north of Mexico).

Transcript of Photographic key to the Pseudoscorpions of Canada and the...

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Canadian Journal of Arthropod Identification No.12 (January 2011) BRUNKE ET AL.

doi: 10.3752/cjai.2011.12 1

Staphylinidae of Eastern Canada and Adjacent UnitedStates. Key to Subfamilies; Staphylininae: Tribes andSubtribes, and Species of StaphylininaAdam Brunke*, Alfred Newton**, Jan Klimaszewski***, ChristopherMajka**** and Stephen Marshall**University of Guelph, 50 Stone Road East, School of Environmental Sciences, 1216/17 BoveyBuilding, Guelph, ON, N1G 2W1. [email protected], [email protected]. **FieldMuseum of Natural History, Zoology Department/Insect Division, 1400 South Lake Shore Drive,Chicago IL, 60605. [email protected]. *** Laurentian Forestry Centre, 1055, rue duP.E.P.S., Stn. Sainte-Foy Québec, QC, G1V 4C7. [email protected] **** NovaScotia Museum, 1747 Summer St., Halifax, NS, B3H 3A6. [email protected].

Abstract. Rove beetles (Staphylinidae) are diverse and dominant in many of NorthAmerica’s ecosystems but, despite this and even though some subfamilies arenearly completely revised, most species remain difficult for non-specialists toidentify. The relatively recent recognition that staphylinid assemblages in NorthAmerica can provide useful indicators of natural and human impact on biodiversityhas highlighted the need for accessible and effective identification tools for thislarge family. In the first of what we hope to be a series of publications on thestaphylinid fauna of eastern Canada and the adjacent United States (ECAS), wehere provide a key to the twenty-two subfamilies known from the region, atribe/subtribe level key for the subfamily Staphylininae, and a species key to thetwenty-five species of the subtribe Staphylinina. Within the Staphylinina, thePlatydracus cinnamopterus species complex is defined to include P. praetermissusNewton spec. nov., P. cinnamopterus (Gravenhorst) and P. zonatus (Gravenhorst).Lectotypes are designated for Staphylinus cinnamopterus Gravenhorst, Staphylinuszonatus Gravenhorst and Staphylinus badius Mannerheim. One new Canadianrecord, one new record from eastern Canada, and thirty-nine new provincial orstate records are presented.

IntroductionRove beetles (Coleoptera:

Staphylinidae) constitute the largest family ofinsects worldwide, with more than 55,440described species (Grebennikov and Newton2009) found in a great diversity of terrestrial andperiaquatic habitats. In Canada, more than 1400species are known and some large subfamilies(Staphylininae, Tachyporinae) have been nearlycompletely revised. An excellent synthesis ofthe staphylinid literature is given by Thayer(2005); however, our understanding ofstaphylinid ecology and habitat requirements isstill very limited. Recent work has revealed thatstaphylinids are dominant organisms in Canadianforest ecosystems (Paquin and Duperre 2001)and because many species require continuous,mature or old growth stands, the composition oftheir species assemblages effectively

communicates the degree of natural or humanimpact upon these systems (Pohl et al. 2008). Ina recent review of the use of beetles inconservation, New (2010) highlighted the criticalimportance of species-level identifications insurveys aiming to document changes toecosystems via human development or climatechange. The continued ability of insect surveysto address important ecological andconservation-themed questions dependsprimarily on the correct identification ofspecimens, which in turn depends on theavailability of effective keys.

Although several recent works such asKlimaszewski (2000) and Newton et al. (2000)include keys to the genus or subfamily level,accompanied by line drawings of importantcharacters, keys to most staphylinid groups arepoorly illustrated and cover broad geographicareas (e.g. North America north of Mexico).

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Furthermore, many important staphylinid keysare scattered in the primary literature and can betime-consuming or difficult to locate. Theseobstacles can now be addressed using online,open-access publication to provide richlyillustrated, digital keys to the species ofStaphylinidae known to inhabit eastern Canadaand the adjacent United States (ECAS). ECASis defined here as Ontario eastward and includesthe following states that are adjacent to easternCanada: Michigan, Indiana, Ohio, Pennsylvania,New York, Vermont, New Hampshire andMaine. Although broader regions have beenused in previous works (i.e., Northeastern NorthAmerica of Downie and Arnett (1996)), ECAScorresponds well to a naturally delimitedstaphylinid fauna and has allowed thesimplification of the keys through exclusion ofsouthern or western taxa.

Despite the largest and some of themost conspicuous species being in theStaphylinina (Staphylininae: Staphylinini), thetaxonomy of this subtribe is currently far fromsettled. The largest genus, Platydracus C. G.Thomson, remains under revision by the secondauthor and thus no working key exists for thefauna of ECAS. To remedy this, we provide aregionalized Platydracus key in advance of theupcoming revision and describe a new speciesthat occurs in ECAS. Multiple accidentalintroductions (Newton 1987; Majka andKlimaszewski 2008), a history of incorrectnames (Newton 1987), inadequate speciesdescriptions, colour variation, and the presenceof several rare or typically ‘southern’ species hasfurther complicated the recognition ofStaphylinina in this region. Despite this, weshow here that most Staphylinina can be easilyidentified to species using high-resolutionphotographs of readily visible characters.

In this first publication we provide anoverview of, and a key to, the twenty-twostaphylinid subfamilies occurring in ECAS, andthen review the tribes and subtribes ofStaphylininae and the twenty-five species ofStaphylinina occurring in ECAS. Keyspresented here will also work for the fauna ofConnecticut, Illinois, Massachusetts, Minnesota,Rhode Island and Wisconsin. We envision thisfirst key as a nexus for future keys to link to,thus creating an integrated guide to theStaphylinidae of ECAS. Future publications areplanned, with the most immediate ones covering

the remaining groups of Staphylininae.

MethodsSpecimen photographs were taken with a

Visionary Digital imaging system and CombineZor Helicon Focus software was used to combinemultiple photographs into high depth-of-fieldimages. Most photographs of living specimenswere taken with digital SLR cameras and a105mm macro lens, often with the addition of a2X teleconverter or extension rings, but livephotographs were also contributed by a numberof collaborators using a variety of equipment(see acknowledgments). Online keys werecreated with Adobe Creative Suite 3 software,including Photoshop, Illustrator, Dreamweaverand Fireworks. Maps of species distributionswere prepared using ARC MAP, with recordsprevious to 1970 mapped separately from thoseoccurring on or after this year in order tohighlight possible distribution changes over time(this was not done for common species). Mostspecies in potential decline reviewed hereinshowed a noticeable change beginning around1970, with a few species ‘declining’ in recordsafter 1980. Thus, 1970 was chosen as a standarddivision point with special cases discussed underthose species. Measurements were taken with aneyepiece micrometer using various dissectingmicroscopes. A list of institutions from whichmaterial was examined and a list of contributingphotographers are given in theacknowledgements. Throughout, ‘abdominalsegments 1-6’ corresponds to the segmentsnumbered as visible and therefore representanatomical segments 2-8. Unless otherwisestated, we follow the higher classification ofNewton et al. (2000).

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Identification Keys

1. Key to the subfamilies in ECAS

1 Body with one of the distinctive shapes and all corresponding important features:

Oxyporinae (Fig. 1.1.1-1.1.2): Mandibles about as long as head, protruding forward andcrossing; last segment of labial palpi crescent shaped (examine ventral surface of head)

Scaphidiinae (Fig. 1.2.1-1.2.3): Head partly concealed from above; less than half the abdomenextending past the elytra; antennae originate between the eyes

Micropeplinae (Fig.1.3.1): Antennae with 9 antennomeres, the last enlarged to form a club;entire dorsal surface with sculpturing of ridges and pits; body elongate oval in shape

Pselaphinae (Fig. 1.4.1): Characteristic body shape: elytra and abdomen appear as an ‘apicalsection’, always distinctly wider than head and pronotum; elytra far from reaching the apex ofthe abdomen; antennae often clubbed; often with deep pits on the dorsal surface

Scydmaeninae (Fig. 1.5.1): Body characteristically ant-like; posterior of head without ocelli,elytra reaching or nearly reaching apex of abdomen; always smaller than 2.5mm

- Body unlike the shapes shown above ......................................................................................................2

2 Dorsal surface of elytra with ridges (Fig. 1.6.1, 1.7.1); pronotum approximately equal in width at baseand apex ..................................................................................................................................................3

- If elytra with ridges (rarely), then pronotum distinctly narrower at apex OR at base .............................4

3 With ridges on both pronotum and elytra (Fig. 1.6.1); two known species in ECAS, identical inappearance....................................................................................................... Pseudopsinae: Pseudopsis

- With ridges on elytra only; two species, both red and black (Fig. 1.7.1) ........ Olisthaerinae: Olisthaerus

4 With a unique habitus as in Fig. 1.8.1; antennae with a two-segmented club and inserted in front of theeyes (Fig. 1.8.1); eyes protruding from lateral head outline and highly convex (Fig. 1.8.1); two speciesin ECAS, differing mostly in color...........................................................Megalopsidiinae: Megalopinus

- Unlike Megalopinus ................................................................................................................................5

5 With a distinctive habitus (Fig. 1.9.1), rarely with eyes slightly less prominent (Dianous) (Fig. 1.9.2);antennae originate between the eyes (Fig. 1.9.3); second last segment of maxillary palpus swollen,wider apically than preceding segment ....................................................................................... Steninae

- Unlike Fig. 1.9.1 or Dianous; if antennae originate between eyes, then eyes not distinctly protrudingfrom lateral head outline..........................................................................................................................6

6 Habitus similar to Fig. 1.10.1; eyes located at the hind margin of the head, globular; second lastsegment of maxillary palpus swollen, wider apically than preceding segment (Fig. 1.10.2); mandibleslong and thin, without teeth (Fig. 1.10.2); antennae with a two-segmented club (Fig. 1.10.2); alwayssmaller than 2.5 mm ............................................................................................................Euaesthetinae

- Unlike Fig. 1.10.1; if similar to 1.10.1 and smaller than 2.5mm, then antennae without a two-segmented club ........................................................................................................................................7

7 Abdomen with 7 visible abdominal sternites (Fig. 1.11.1) OR with a unique habitus as in Fig. 1.11.2 orFig. 1.11.3 .................................................................................................................... Oxytelinae (most)

- Abdomen with 6 visible abdominal sternites (Fig. 1.12.1)......................................................................8

8 Antenna inserted behind the anterior margin of the eye (Fig. 1.13.1) ...................... Aleocharinae (most)- Antenna inserted in front of the anterior margin of the eye (Fig. 1.12.2)................................................9

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9 Metasternum with small, rounded lobes near the apex (Fig. 1.13.2); appearance distinct (Fig. 1.13.3);found with termites of the genus Reticulotermes (Fig. 1.13.4)...............................................................................................................................................................Aleocharinae:Trichopseniini: Trichopsenius

- Metasternum without lobes (Fig. 1.14.1); not exactly like Trichopsenius ............................................10

10 Ocelli present behind compound eyes (Fig. 1.15.1), often obscured in Eusphalerum and Pycnoglypta;see Fig. 1.15.2-3 to check if your specimen belongs to these two genera ................................ Omaliinae

- Ocelli absent (Fig. 1.12.3); not similar to Eusphalerum or Pycnoglypta ..............................................11

11 Body oval with elytra ≥ 2 times as long as the pronotum and covering one or more tergites (Fig.1.16.1); pronotum strongly transverse, about 2 times as wide as long or wider (Fig. 1.16.1);antennomeres I and II expanded (Fig. 1.16.1); small beetles, 1-3mm in length.............. ........Proteininae

- Not with the above combination of characters ......................................................................................12

12 Antennae with 10 antennomeres (Fig. 1.13.5); size always < 2mm .............Aleocharinae: Hypocyphtini- Antennae with 11 antennomeres (Fig. 1.12.4); always >2mm ..............................................................13

13 Abdominal tergites unmargined (dorsally, without a longitudinal suture at each side) (Fig. 1.17.1);abdomen always parallel-sided..............................................................................................................14

- Abdominal tergites margined: most segments with a longitudinal suture at each side (Fig. 1.12.5); ifunmargined then abdomen tapered posteriorly (Fig. 1.14.2); abdomen parallel sided or not ...............15

14 Abdominal surface with fish scale-like sculpturing (Fig. 1.18.1); last maxillary palpomere much largerthan the preceding and slipper-shaped (Fig. 1.18.2)............................................. Paederinae: Palaminus

- Abdominal surface unlike fish scales but often with other textures (Fig. 1.17.2); last maxillarypalpomere smaller than the preceding and not slipper-shaped (Fig. 1.17.3) ............................. Osoriinae

15 Procoxa much smaller and differently shaped than femur (Fig. 1.19.1)................................................16- Procoxa large and elongate, similar to femur (Fig. 1.12.6) ...................................................................19

16 Elytra with at least one distinct row of fused punctures (Fig. 1.20.1); protibia with thick spines alongoutside edge (Fig. 1.20.2) ........................................................................................ Piestinae: Siagonium

- Punctures of elytra not arranged in distinct rows (Fig. 1.19.2); protibia smooth, with only setae onoutside edge (Fig. 1.19.3) ......................................................................................................................17

17 Pronotum, measured at midline, about as long as elytra or longer (Fig. 1.11.4); elytra convex.............................................................................................................................................. Oxytelinae: Euphaniini

- Pronotum, measured at midline, about half as long as elytra (Fig. 1.19.4); elytra flattened .................18

18 Sides of pronotum serrate; pronotum evenly covered with setae-bearing punctures (Fig. 1.19.5)................................................................................................................Phloeocharinae: Charhyphus picipennis

- Sides of pronotum smooth; pronotum with only a few setae-bearing punctures (Fig. 1.21.1) ............................................................................................................................. Habrocerinae: Habrocerus magnus

19 Antennomeres 3-11 extremely elongate, with prominent long black setae much longer than the widthof their corresponding segment (Fig. 1.21.2); body distinctly widest at elytra; antennomeres 1 and 2much wider than 3-11............................................................................................................................20

- Antennomeres 5-11 with antennal setae almost always shorter than the width of their correspondingsegment (Fig. 1.14.3); if with antennae similar to Fig. 1.21.2 (one genus), then body linear in form,with all body sections subequal in width...............................................................................................21

20 Dorsal surface smooth, with only a few large bristles; without a ‘neck’ (Fig. 1.21.3) ...................................................................................................................................................... Habrocerinae: Habrocerus

- Dorsal surface covered in punctures with hairs; with a constriction behind the eyes to form a ‘neck’(Fig. 1.22.1)................................................................................... Trichophyinae: Trichophya pilicornis

21 Head constricted behind eyes to form a 'neck' clearly visible from above (Fig. 1.12.7) or sometimesforming a broad, weakly defined neck (Fig. 1.12.8) .............................................................................22

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- Without a 'neck': head either straight behind the eyes or gradually meeting pronotum such that an extrasection is not present (Fig. 1.11.5) ........................................................................................................23

22 Pronotum without a postcoxal process (Fig. 1.12.9) or with a small, translucent one (Fig. 1.12.10)..........................................................................................................................................Staphylininae (most)

- Pronotum with a sclerotized, opaque postcoxal process (Fig. 1.18.3).......................... Paederinae (most)23 Elytral epipleuron with a carina (Fig. 1.14.4)...................................................................... Tachyporinae- Elytral epipleuron without a carina (Fig. 1.12.11).................................................................................2424 Habitus as in Fig. 1.12.12; pronotum mostly glabrous....................Staphylininae: Atanygnathus bicolor- Habitus as in Fig. 1.19.6; pronotum heavily pubescent......... Phloeocharinae: Phloeocharis subtilissima

2. Key to the tribes and subtribes of Staphylininae in ECAS

1 Elytra overlapping at midline; inside margin of elytra arcuate (Fig. 2.1.1)...........................Xantholinini- Elytra meeting evenly at midline; inside margin of elytra straight (Fig. 2.2.1) .......................................2

2 Neck narrow: about 1/4 the width of the head (Fig. 2.3.1)............................ Diochini: Diochus schaumi

- Neck, if present, broader: at least one half the width of the head (Fig. 2.2.2) .........................................3

3 Antennal base separated by a distance less than that between the eye and antennal base (Fig. 2.4.1);mandibles (when closed) directed forward and blade-like; with a pair of plates anterior to theprosternum (Fig. 2.1.2).................................................................................................... Othiini: Atrecus

- Antennal base separated by a distance greater than that between eye and antennal base (Fig. 2.2.3);mandibles (when closed) not directed forward; without a pair of plates anterior to the prosternum(Fig.2.2.4)........................................................................................................................ (Staphylinini) 4

4 With a unique appearance (Fig. 1.12.12); without a visible neck; maxillary and labial palpi extremelyelongate (Fig. 2.2.5); tarsal formula 5-4-4....................................Tanygnathinina: Atanygnathus bicolor

- Unlike A. bicolor; always with at least a weakly formed neck; maxillary and labial palpi not extremelyelongate (Fig. 1.12.8); tarsal formula 5-5-5.............................................................................................5

5 Dorsal surface of neck completely without setose punctures, often with non-setose micropunctures(Fig. 1.12.8).............................................................................................................................................6

- Dorsal surface of neck with setose punctures on at least lateral portions (Fig. 1.12.7) ...........................8

6 Pronotum with dorsal rows of ≥4 punctures (Fig 2.2.6) OR with many punctures not arranged in pairs;without empodial setae between tarsal claws (Fig. 2.2.8) .......................................................Philonthina

- Pronotum with dorsal rows of 1-3 punctures (Fig. 2.2.7) OR impunctate; with empodial setae betweentarsal claws (Fig. 2.2.9), sometimes very long (Fig. 2.2.10) ...................................................................7

7 Last segment of maxillary palpus reduced, much narrower than the previous segment (Fig. 2.2.11)...................................................................................................................... Amblyopinina: Heterothops

- Last segment of maxillary palpus normally developed, not markedly narrower than previous segment(Fig. 1.12.8)................................................................................................................................ Quediina

8 Head narrow, about half the width of the pronotum (Fig. 2.2.12); last segment of labial palpuscharacteristically hoof-shaped, with a broad apical face (Fig. 2.2.14) .......................................................................................................................................................Anisolinina: Tympanophorus puncticollis

- Head much wider than half the width of the pronotum (Fig. 2.2.13); last segment of labial palpus notexactly like that of T. puncticollis, apical face narrow (Fig. 2.2.15) .....................................Staphylinina

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3. Key to the genera of Staphylinina in ECAS

1 Pronotum smooth and glossy, except for margins (Fig. 3.1.1) ....................... Creophilus (C. maxillosus)- Most of pronotum punctate, often with an impunctate midline (Fig. 2.2.13) ..........................................2

2 Anterior angles of pronotum sharp, approximately 90 degrees or less (Fig. 3.2.1) ................ Ontholestes

- Anterior angles of pronotum rounded, greater than 90 degrees (Fig. 3.3.1)............................................3

3 Large lateral setae of head not restricted to the anterior half of the temple (Fig. 3.3.2) .........Platydracus- Large lateral setae of head restricted to the anterior half of the temple (Fig. 3.4.1) ................................4

4 Body and femora completely black AND with elytra (measured at middle) longer than pronotum (Fig.3.5.1); last segment of labial palpus stout with its apical margin at a distinct angle (Fig. 2.2.15) ...................................................................................................................................................................Tasgius

- Not completely black OR with elytra, at middle, distinctly shorter than pronotum (Fig. 3.6.1); lastsegment of labial palpus more elongate with its apical margin either truncate or pointed (Fig. 3.6.2) ...5

5 Elytra red; head with eyebrow-like patches of golden setae (Fig. 3.7.1)..... Staphylinus (S. ornaticauda)

- Elytra not red; head without eyebrow-like patches of golden setae (Fig. 3.4.2) .....................................6

6 Base of some abdominal segments with gold or silver setae (best seen ventrally) (Fig. 3.4.3); lastmaxillary palpomere spindle-shaped (Fig. 3.4.4) ................................................................ Dinothenarus

- Abdominal segments never with silver or gold setae (Fig. 3.6.3); last maxillary palpomere elongate andrectangular (Fig. 3.6.2) ..................................................................................................................Ocypus

4. Key to the Dinothenarus species in ECAS:

1 Head yellowish-red (Fig. 3.4.2), colour sometimes heavily muted in discoloured specimens (Fig.4.1.1); pronotum grey and orange; abdomen with gold and silver setae ............. Dinothenarus capitatus

- Head and pronotum black (Fig. 4.2.1); abdominal segments with gold setae only ..................D. badipes

5. Key to the Ocypus species in ECAS

1 Legs black, concolourous with rest of body (Fig. 3.6.1) .................................................... Ocypus nitens

- Legs brownish red, contrasting with the rest of body (Fig. 5.1.1) ........................................ O. brunnipes

6. Key to the Ontholestes species in ECAS

1 Legs bicoloured, black and yellow (Fig. 6.1.1); apical half of antennae black; base of fifth abdominaltergite with golden setae........................................................................................Ontholestes cingulatus

- Legs completely dark; antennae entirely orange; base of fifth abdominal tergite with silver setae (Fig.6.2.1) ....................................................................................................................................... O. murinus

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7. Key to the Platydracus species in ECAS

1 Eye about twice as long as the temple (Fig. 7.1.1) ................................................Platydracus femoratus

- Eye smaller, no more than marginally longer than the temple (Fig. 2.2.13) ...........................................2

2 Punctures of pronotum spaced unevenly, creating smooth, glossy areas in addition to a medianimpunctate line (Fig. 7.2.1) .....................................................................................................................3

- Punctures of pronotum spaced evenly, with the exception of a median impunctate line in some species(Fig.7.3.1)................................................................................................................................................4

3 Pronotum about as long as wide; at middle, impunctate line of pronotum obsolete (Fig. 7.2.2)................................................................................................................................................................. P. exulans

- Pronotum longer than wide; impunctate line of pronotum at least as wide as 5 punctures for most of itslength (Fig. 7.4.1); found along or near ocean coastline .....................................................P. praelongus

4 Elytra reddish, with distinct, elongate patches of dark setae (Fig. 7.5.1); scutellum with a median goldline bordered by a pair of dark patches (Fig. 7.5.1).................................................................................5

- Elytra at most, with broad, faint darker areas (Fig. 7.6.1); scutellum without a median gold line (Fig.7.6.1) .......................................................................................................................................................6

5 Large species (22-35mm); impunctate area of pronotum interrupted over much of its length (Fig. 7.7.1)..............................................................................................................................................P. maculosus

- Smaller species (13-19mm); impunctate area complete along the entire length of pronotum (Fig. 7.5.2).....................................................................................................................................................P. comes

6 Each elytron with an orange or gold spot (Fig. 7.8.1) ................................. maculate form of P. fossator

- Elytra without such a spot (Fig. 7.9.1) ....................................................................................................7

7 Abdominal tergites with areas of dense yellow or blonde setae (Fig. 7.10.1); impunctate median line ofpronotum always absent or interrupted as in Fig. 7.7.1 ...........................................................................8

- Abdominal tergites with either sparse areas of yellow setae or none at all (Fig. 7.9.2); impunctatemedian line of pronotum variable..........................................................................................................10

8 Body surface dull and without color (i.e., no part metallic) (Fig. 7.11.1); elytra red ....... P. immaculatus

- At least pronotum metallic greenish-bronze to purple (Fig. 7.10.2); elytra not red ................................9

9 Abdominal tergites 1-5 each with distinct tripartite yellow pubescence (Fig. 7.10.2); pronotum metallicbronze-green; ventral margin of elytral epipleuron pale (Fig. 7.10.3)................................... P. viridanus

- At least one of abdominal tergites 1-5 with a different pattern of yellow setae (Fig. 7.12.1); pronotumwith a purple reflection (bronze in discoloured specimens); ventral margin of elytral epipleuron black(Fig. 7.12.2)........................................................................................................................... P. violaceus

10 Pronotum without a complete smooth median line (Fig. 7.6.2); if smooth area present, then interruptedalong its length ......................................................................................................................................11

- Pronotum with a complete impunctate median line, its narrowest point at least one puncture wide (Fig.7.9.3) .....................................................................................................(P. cinnamopterus Complex) 13

11 Body dark brown, with reddish-brown elytra and paler abdominal apex (Fig. 7.6.3); antennomere Idark dorsally and pale ventrally (Fig. 7.6.4); eyes small, distinctly smaller than the temple (Fig. 7.6.3) ..................................................................................................................................................P. mysticus

- Body entirely black (Fig. 7.13.1), rarely with sparse yellow setae at the base of the abdominalsegments; antennomere I completely black (Fig. 7.13.2); eyes larger, subequal to distinctly longer thanthe temple (Fig. 7.13.2) .........................................................................................................................12

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12 At least basal abdominal tergites with a pair of dark velvet spots (Fig. 7.13.1); eye subequal in lengthto temple..............................................................................................................................P. tomentosus

- Abdominal tergites without dark velvet spots (Fig. 7.8.2); eye distinctly longer than temple ..................................................................................................................................... melanistic form of P. fossator

13 Head, pronotum and elytra completely bright red (Fig. 7.9.4) ..............rufous form of P. cinnamopterus

- Some of these areas darker (Fig. 7.9.1) .................................................................................................14

14 Impunctate median line of pronotum wide: 3 or more punctures could fit across its narrowest point(Fig. 7.14.1); antennomere 9 subquadrate: its apical width nearly equal or equal to its length (Note:rotate antennae so that their segments are widest in profile) (Fig. 7.14.2); apex of median lobe ofaedeagus widely truncate in parameral view (Fig. 7.14.3) .............. P. praetermissus Newton spec. nov.

- Impunctate median line of pronotum narrow: 2 or fewer punctures could fit across its narrowest point(Fig. 7.9.5); antennomere 9 at least slightly transverse: its apical width greater than its length (Note:rotate antennae so that their segments are widest in profile) (Fig. 7.3.2); apex of median lobe ofaedeagus obtusely to acutely projected in parameral view (Fig. 7.9.6) .................................................15

15 Antennomere 9 distinctly transverse (Note: rotate antennae so that their segments are widest in profile)(Fig. 7.9.7); pronotum usually evenly coloured, light to dark red (Fig. 7.9.8); apex of median lobe ofaedeagus acutely projected in parameral view (Fig. 7.9.6) ...........................................P. cinnamopterus

- Antennomere 9 slightly transverse (Note: rotate antennae so that their segments are widest in profile)(Fig. 7.3.2); pronotum light to dark red with dark brown to black anterior angles, almost never evenlycoloured (Fig. 7.3.3); apex of median lobe of aedeagus obtusely projected in parameral view (Fig.7.3.4) ........................................................................................................................................ P. zonatus

8. Key to the Tasgius species in ECAS

1 Pronotum with punctures of two different sizes, most punctures well-separated (Fig. 8.1.1).................................................................................................................................................................. Tasgius ater

- Pronotum with punctures subequal in size, most punctures almost touching (Fig. 8.2.1) .......................2

2 Pronotum with sides widest just before anterior angle, sides strongly converging (Fig. 8.3.1); basal halfof antennomere 2 bicoloured (Fig. 8.3.2) ............................................................................T. melanarius

- Pronotum widest at the anterior one third, sides weakly converging (Fig. 8.2.2); basal half ofantennomere 2 solid red (Fig. 8.2.3).........................................................................................T. winkleri

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9. The subfamilies of Staphylinidae in ECAS

9.1 Aleocharinae Fleming 1821

Most aleocharines can be recognized byantennae that are inserted posterior to a linedrawn along the anterior margins of the eyes(Fig. 1.13.1), non-bulging eyes (unlike Steninae),male genitalia with complex parameres and theiroverall appearance which is unlike that of theScaphidiinae (Fig. 1.1.1). The following twoaleocharine groups of ECAS are exceptions,because their antennae are inserted eitheranterior to the level of the anterior eye margin orambiguously close to it:

Trichopsenius Horn, with characteristicmetasternal plates (Fig. 1.13.2) and adistinct habitus (Fig. 1.13.3)

The tribe Hypocyphtini, minute beetles(<2mm) with only 10 antennomeres(Fig. 1.13.5)

Aleocharinae is the most diverse subfamilyof Staphylinidae in ECAS and also the leastknown. This subfamily is widely distributed inNorth America and occurs in almost allterrestrial habitats, but most species are forestdwelling. There they occur in leaf litter, underbark, in fungi, in moss and within the nests ofants, termites, mammals and birds. Elsewhere inECAS they inhabit seashores, edges of waterbodies, wetlands and prairies. Drusillacanaliculata Fabricius is a common species oftenfound in disturbed areas under leaves and stones,usually near ants but often near water(Fig.9.1.1). Species of Gymnusa Gravenhorst areuncommonly collected aleocharines that areassociated with Sphagnum in wetlands and alongstreams. In undisturbed fens and bogs, lightlypressing on water-logged vegetation willsometimes reveal specimens of this genus (Fig.9.1.2). Tachyusa Erichson are characteristicallyshaped beetles found along the margins of lakes,ponds, wetlands, and streams; they aresometimes collected from beaver lodges (Fig.9.1.3). Xenodusa Wasmann species live withcolonies of Formica ants during the warmermonths and overwinter with colonies of the antgenus Camponotus (Wheeler 1911) (Fig. 9.1.4).In ECAS, Aleocharinae are primarily predaceouson a wide variety of invertebrates, however somegroups are strictly mycophagous (Ashe 1981) orparasitoids of cyclorraphous Diptera(Klimaszewski 1984).

9.2 Euaesthetinae Thomson 1859Euaesthetines can be recognized by the

globular eyes that are located at the hind marginof the head (Fig.1.10.1), antennae that end in atwo-segmented club, thin toothless mandibles,and a swollen second last maxillary palpomere(Fig. 1.10.2). Euaesthetines are also distinctivefor their relatively small size: the largest inECAS are 2.3mm long (Fig. 9.2.1).

This subfamily is widespread in NorthAmerica, with two western endemic genera;three genera are known from ECAS. The mostdiverse is Euaesthetus Gravenhorst (Fig. 9.2.2),which can be found in leaf litter, moist fields,and wet meadows dominated by sedges, wherethey often co-exist with the related Steninae.Stictocranius puncticeps LeConte is the onlyeastern species of the genus and can be found inleaf litter. Several species of EdaphusMotschulsky are known from the northeasternUS but are not yet recorded from Canadaalthough they likely occur in the Carolinianforests of southern Ontario and Appalachianforests of New Brunswick and Nova Scotia.Edaphus occur in leaf and wood debris,treeholes, and stumps in moist forests (Puthz1974). All members of this subfamily are bestcollected by sifting litter and then processing itin a Berlese funnel, but can also be captured bypan and pitfall trapping. Euaesthetines arepredators of small invertebrates but their exactdiet is unknown.

9.3 Habrocerinae Mulsant and Rey 1877:Habrocerus Erichson 1839

Species of Habrocerus can berecognized by antennomeres 3-11 that areextremely slender, with long black macrosetae(Fig. 1.21.2), and by their dorsal pronotum withfew setae (Fig. 1.21.3). Trichophya Mannerheim(Trichophyinae) has similar antennae; howeverits dorsal surface is covered with many setae(Fig. 1.22.1). Habrocerus magnus LeConte hasbroader antennomeres and lacks other importantmorphological features of the genus (Fig. 9.3.1)(Assing and Wunderle 1995), and probablybelongs in a lineage of Tachyporinae (Assingand Wunderle 1995) or Olisthaerinae (Newton etal. 2000). We treat this species here asHabrocerus pending its formal transferelsewhere.

In North America, this subfamily occursin the northeast and southwest. There arecurrently three species found in ECAS: therelatively common, introduced Habrocerus

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capillaricornis Gravenhorst, the native H.schwarzi Horn, and the native but unplaced‘Habrocerus’ magnus (see above). Virtuallynothing is known about the rarely collected H.magnus, although it shares features with knownsubcortical staphylinids (relatively flattenedbody, small globular coxae). H. capillaricornis ismost often collected from forested areas in litter(Fig. 9.3.2), under bark, in fungi (Assing andWunderle 1995) and in compost. Habrocerusschwarzi is also collected from forested areas,most often from decaying fungi in the laterstages of decomposition but also from mooseand grouse dung and leaf litter (Assing andWunderle 1995). It has also been captured ingreat numbers in pitfall traps placed in parts ofthe ‘southern’ boreal forest of Québecundergoing the deciduous stage of succession(Paquin and Duperre 2001). Most records of H.schwarzi are from Canada and this species wasfound to be dependent on old growth forests(Spence et al. 1997). The diet of habrocerines isunknown.

9.4 Megalopsidiinae Leng 1920:Megalopinus Eichelbaum 1915

Megalopinus is immediatelyrecognizable by its distinct habitus, bulging eyesoccupying most of the lateral head margin,orange elytral markings, and clubbed antennae(Fig.1.8.1). They are somewhat similar toSteninae but with antennae inserted in front of aline drawn at the anterior margin of the eyes(Fig.1.8.1).

These beetles appear to be microhabitatspecialists, living under dead logs with fungalgrowth (Leschen and Newton 2003). As theyfeign death and drop to the ground afterdisturbance (Leschen and Newton 2003), theyare probably best collected by sifting fungusydebris under logs. Specimens have also beencollected from pan traps near forest and hand-collected from fungus. Two species of thismostly tropical genus occur in the southeasternand south-central United States and reach theirknown northern limit in Indiana. However, theyare very rarely encountered and they mayeventually be found elsewhere in ECAS.

Individuals of Megalopinus caelatusGravenhorst have been observed feeding onlarval Diptera in the laboratory, slowly locatingand capturing their prey during the day (Leschenand Newton 2003). They likely huntinvertebrates associated with fungus growing onthe undersides of logs.

9.5 Micropeplinae Leach 1815:Micropeplus Latreille 1809

Micropeplus is easily recognized due toits nine segmented antennae with an enlargedterminal segment, forming a club (Fig.1.3.1). The body is elongate oval in shape andthe entire dorsal surface is covered with a varietyof ridges (Fig. 1.3.1). The genus is widelydistributed in North America.

In ECAS, species of Micropeplus canbe sifted from leaf litter on the forest floor, inboggy areas (Campbell 1968), or along streams(Campbell 1973). They sometimes occur oncarrion and in beaver lodges (Campbell, 1968).Sweeping field vegetation in the evening oftenyields specimens (Campbell 1968) that areprobably dispersing. The species of Micropeplusare mycophagous, feeding on the spores andconidia of moulds and other fungi (Hinton andStephens 1942; Thayer 2005).

9.6 Olisthaerinae C.G. Thomson 1858:Olisthaerus Dejean 1833

The genus Olisthaerus can berecognized by an elongate and parallel body, adark head, red pronotum and abdomen, aparallel-sided pronotum, and the longitudinalstriae of the elytron (Fig. 1.7.1). The genusSiagonium Kirby and Spence is loosely similarin appearance but is easily separated by the rowsof coarse punctures on the elytra and thepresence of cephalic horns in males.

Two rare Holarctic species occur inECAS: O. megacephalus (Zetterstedt) andO. substriatus (Gyllenhal). Olisthaerusmegacephalus differs from O. substriatus by thewider head and smaller, flat eyes (Fig. 9.6.1).They are found under the bark of dead conifers(Newton et al. 2000) within the boreal forestecozone (both species) or in relict boreal forestsfound on mountains in New York and NewHampshire (O. substriatus only) (Fig. 9.6.2). Inboreal forest surveys, O. substriatus (Paquin andDuperre 2001) and O. megacephalus (Hammondet al. 2004) were captured exclusively within‘old’ (>100 years old) or old growth forests. Thefeeding habits of Olisthaerus are unknown.Olisthaerus megacephalus and the subfamilyOlisthaerinae are newly recorded for theprovince of Newfoundland and Labrador.

CANADA: NL: Labrador, Nain, 56.50 -61.75,2-VII-1983, R. Sexton, 1 (DEBU).

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9.7 Omaliinae MacLeay 1825

Omaliinae can be recognized easily bythe presence of a pair of ocelli behind thecompound eyes (Fig. 1.15.1). However,individuals of Pycnoglypta Thomson andEusphalerum Kraatz sometimes lackocelli. Eusphalerum species in ECAS are alwaysyellowish-brown with a habitus as in Fig.1.15.2. The two Pycnoglypta species in ECASare similar in habitus to Fig. 1.15.3.

Omaliines are widely distributed inNorth America and can be collected year-roundalthough the greatest diversity occurs in fall toearly spring. They can be collected by siftinglitter in forests and along wet areas (Fig. 9.7.1),in fungi, on flowers, in sweeps and pan trapsplaced in grassy areas, under stones near water(Fig. 9.7.2), and on the snow surface in latewinter (Fig. 9.7.3). Some species aggregate onsurfaces in fall and can be relatively conspicuous(Fig. 9.7.4). Many Omaliinae are predators (Fig.9.7.5) or omnivores, but others seem to bestrictly saprophagous or pollen-feeding.

9.8 Osoriinae Erichson 1839

Osoriines are characterized by aparallel-sided abdomen that lacks sutures on thedorsal surface (Fig. 1.17.1). The abdomen caneither be flattened and with a single lateral suture(Fig. 1.17.1), or cylindrical and formed of fusedtergites and sternites (Fig. 1.17.2). The onlystaphylinids with a similar abdomen are speciesof Palaminus Erichson (Paederinae), which donot have the last maxillary palpomere smallerthan the previous one (as in Fig. 1.17.3) andsome Stenus Latreille (Steninae), which havelarge convex eyes that occupy most of the lateralhead outline.

The Osoriinae is a widespreadsubfamily in North America but it remainstaxonomically and biologically poorly known.The species inhabiting ECAS are generallyfound under bark, in leaf litter and in ant nests indecaying wood. Osoriinae are very poorlyrepresented in collections of ECAS insects,possibly because they are really rare, or possiblybecause they are merely secretive; adults of someRenardia Motschulsky are winter-active underbark. The most common species in ECAS,Thoracophorus costalis Erichson (Fig. 9.8.1), isoften found under bark (especially of large beechlogs), but sometimes occurs in leaf litter and onconk fungi. Motter (1898) found a large number

(up to 160 individuals) of Eleusis pallidaLeConte adults and larvae in human graves insandy soil, although this species too is normallyfound under bark. This strange report mayindicate subterranean larval and/or pupal stages.Osoriines are saprophagous or mycophagous asfar as known.

9.9 Oxyporinae Fleming 1821:Oxyporus Fabricius 1775

Species of Oxyporus are easilyrecognized by their characteristic labial palpi(Fig. 1.1.2) and prognathous mandibles, whichnormally cross (Fig. 1.1.1). Their unique habitusmakes them difficult to confuse with any othersubfamily.

Widely distributed in North America,species of Oxyporus are associated with fleshyfungi and are almost exclusively collected fromthem. Both adults and larvae tear pieces of thestem and cap with their mandibles, digesting thefungus pre-orally (Hanley and Goodrich 1994).Females construct egg chambers within the hostfungus and remain there after oviposition(Hanley and Goodrich 1995). Both females andmales have been observed to guard the eggchamber (Hanley and Goodrich 1995). Topupate, mature larvae drop off their host andbury themselves in the soil below (Hanley andGoodrich 1994). The common O. vittatusGravenhorst (Fig. 9.9.1) utilizes a broad range offungal host genera while O. quinquemaculatusLeConte (Fig. 9.9.2) exhibits a strong preferencefor fungi in the genus Pluteus Fries (Hanley andGoodrich 1995). Other species, such as O. majorGravenhorst (Fig. 9.9.3), are intermediate in thebreadth of their host range (Hanley and Goodrich1995). However, the larvae of all species seem tobe found only from one or two host fungi(Hanley and Goodrich 1995). The males of someOxyporus are dimorphic for mandible size, withthe difference more pronounced in species withnarrower host preferences (Hanley 2001).Hanley (2001) hypothesized that the two maletypes corresponded to differences in mateacquisition strategies.

9.10 Oxytelinae Fleming 1821

Most oxytelines can be recognized bytheir abdomens, which have seven fullydeveloped sternites rather than the usual six (Fig.1.11.1). The small tribes Euphaniini andCoprophilini do not have seven sternites

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but, fortunately, the genera of these two tribesare distinctive in habitus (Figs 1.11.2, 1.11.3,9.10.1). Mitosynum vockerothi Campbell is notillustrated in this paper but has only beencollected once at the margin of a sphagnum bogin New Brunswick (Campbell 1982) and will keyout with Euphaniini: Syntomium Curtis.

The Oxytelinae is widely distributed inNorth America. Species of Bledius Leach, thelargest genus of Oxytelinae and the onlymembers of tribe Blediini in ECAS (Makranczy2006) burrow in sun-exposed, non-vegetatedfreshwater and marine shorelines where theygraze upon algae scraped from particles ofsubstrate (Herman 1986) (Fig. 9.10.3). Femalesof some Bledius species provision for their larvaein these burrows (Wyatt 1986). Bledius areeasily found by locating their small burrows andscooping the surrounding substrate into a pail ofwater; the beetle will then float to the surface.

The tribe Oxytelini contains mostECAS oxytelines (tribe Thinobiini is treated hereas part of Oxytelini sensu Makranczy 2006)which are generally associated with decayingorganic matter, leaf litter and moss. The genusThinodromus Kraatz can be sifted from the litteraround streams, rivers, and ponds (Fig. 9.10.2).Anotylus Thomson is a diverse genus in ECAS,with at least five species introduced from Europeand Asia (Fig. 9.10.4). Apocellus Erichson has adistinctive habitus and can be found in opengrassy areas and along streams near moss (Fig.9.10.5). Oxytelinae, in general, are best collectedby bait trapping with dung, carrion, and compost,UV lighting, and sifting wet litter along riparianareas. This subfamily contains species ofpredators, algivores, and coprophages; some areprobably saprophages or highly opportunisticomnivores.

9.11 Paederinae Fleming 1821

The species of Paederinae in ECAS canbe recognized by the following combination ofcharacters: a well-sclerotized postcoxal process(Fig. 1.18.3), an apical maxillary palpomere thatis 1/3 the length of the previous segment orshorter (Fig. 1.18.4), or distinctively flattenedand broadened (Fig. 1.18.2), a well-defined neck(Fig. 2.2.2), and the pronotal base distant fromthe humeral angles of the elytra (Fig. 1.18.5).One paederine genus, Palaminus, does not keyout with the rest because it lacks sutures on theabdominal tergites. However, thecharacteristically shaped apical maxillary

palpomere (Fig. 1.18.2) and the fish scale-likesculpturing of the abdomen (Fig. 1.18.1) willdistinguish it from all others.

Paederinae are widely distributed inNorth America in a variety of habitats, with alarge number of species living at the margins ofor near water. Two of the most conspicuousriparian genera are Homaeotarsus Hotchhuth(Fig. 9.11.1) and Paederus Fabricius (Fig.9.11.2). Endosymbionts in the haemolymph ofthe latter genus produce the toxin pederin(Kellner 2001), with females of some speciesnormally possessing up to ten times that of males(Kellner and Dettner 1995). Pederin can causesevere dermatitis if the beetle is crushed againstthe skin (Gelmetti and Grimalt 1993). AstenusDejean is a distinctive genus normally found inwetlands and along shoreline habitat althoughsome species prefer drier, open areas (Fig.9.11.3). Occurring in a variety of litter anddecaying organic matter, Achenomorphuscorticinus Gravenhorst is a common species insouthern Canada and eastern United States (Fig.9.11.4); it is also attracted to lights. One of themost common staphylinids under bark in ECASis Sunius confluentus (Say), a species sometimesbrought indoors with recently cut firewood (Fig.9.11.5). As far as known, all paederines arepredaceous.

9.12 Phloeocharinae Erichson 1839

The subfamily Phloeocharinae is poorlydefined and is therefore best recognized by acombination of characters shared by many othergroups. The two species found in ECAS key outseparately but each can be recognized by habitusalone (Fig. 1.19.6, 9.12.1). Additionally,Charhyphus picipennis LeConte is distinctive forits small, globular procoxae (Fig. 1.19.1),extremely flattened body and serrate lateralpronotal margins (Fig. 1.19.5). Phloeocharissubtilissima Mannerheim can be distinguishedfrom similar groups by its loosely clubbedantennae lacking long black macrosetae (aspresent in Trichophyia), lack of ocelli, andelytral epipleuron without a carina (Fig. 1.12.11).

Phloeocharines occur in western andeastern North America, but do not occur in mostof the central United States. For most of ECAS,Phloeocharinae is represented only by C.picipennis, a species frequently found under thebark of hardwood trees (oak, beech, and others),especially in the earlier stages of decay when thesap is fermenting. Recently, the widespread

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Palearctic species P. subtilissima has beenreported as established in a small park in Halifax,Nova Scotia (Majka and Klimaszewski 2004). InNova Scotia it was collected from under and onthe bark of dead trees and sometimes within theburrows of scolytine beetles (Majka andKlimaszewski 2004). Future collecting isnecessary to confirm its long-term establishmentin Nova Scotia and its possible establishmentelsewhere. The diet of C. picipennis is unknownbut P. subtilissima has been reported (in itsnative range) to feed on scolytine beetles withintheir galleries (Mazur 1995).

9.13 Piestinae Erichson 1839:Siagonium Kirby and Spence 1815

Siagonium can be recognized by thesmall, globular procoxae (Fig. 1.19.1), protibiawith thick spines (Fig. 1.20.2) and the hornlikeprojections in front of the eyes in males (Fig.9.13.1, major male; 9.13.2, minor male). Thegenus Olisthaerus DeJean is loosely similar inappearance but lacks the rows of coarsepunctures on the elytra and does not possesscephalic horns in either sex.

The subfamily Piestinae is widelydistributed in North America and represented inECAS by three species of Siagonium. Very littleis known about the biology of this group. Allspecies occur under the bark of dead trees,especially conifers, and are sometimes collectedat lights. Siagonium stacesmithi Hatch appears tobe trans-boreal in distribution and one individualwas captured in a 'northern' boreal forestconsisting of mature black spruce trees (Paquinand Duperre 2001). Siagonium americanum(Melsheimer) and the more common S.punctatum (LeConte) are found in a greatervariety of forest types, with the latter speciesextending into Mexico. Although the exact dietof Siagonium has never been experimentallydetermined, its mandibles and maxillae bearstriking similarity to those of the mycophagoussilvanid Dendrophagus crenatus (Paykull) andgut contents were found to contain fungalfragments(Crowson and Ellis 1969). Themandibles of Siagonium and other Piestinae alsopossess invaginations, similar to the fungalspore-transmitting mycangia of scotyline andcucujoid beetles (Crowson and Ellis 1969).

9.14 Proteininae Erichson 1839

Proteininae can be recognized by thefollowing combination of characters (Fig.

1.16.1): size small (1-3mm long), body oval withelytra at least twice as long as the pronotum andcovering at least one tergite, pronotum stronglytransverse (at least twice as wide as long at thebase), and antennomeres 1 and 2 much widerthan 3 and 4. Proteinines somewhat resemblesome omaliines but always lack ocelli(proteinines do not resemble those fewOmaliinae that sometimes lack ocelli).

Two genera are widely distributed inNorth America: Proteinus Latreille andMegarthrus Stephens. The former has the hindangles of the pronotum rounded and in the latterthey are pointed (Fig. 9.14.1). Both genera canbe found on decaying fungi and in leaf litter;however Megarthrus can also be collected fromcarrion and dung (Cuccodoro and Löbl1996). Megarthrus has been collectedsuccessfully with window, flight intercept, andpan traps, especially when baited with fungi,carrion, or dung. Several species of Proteinuswere captured in pitfall traps in New Brunswickred spruce forests; two of these were recentlydescribed as new (P. pseudothomasiKlimaszewski and P. acadiensis Klimaszewski)(Klimaszewski et al. 2005). The proteininespecies of ECAS are probably mainlymycophagous and/or saprophagous (Thayer2005) but some species are at least partlypredaceous on nematodes (Cuccodoro 1995).

9.15 Pselaphinae Latreille 1802

Pselaphinae are distinctive for theirbody shape (9.15.1), where the elytra andabdomen appear as an ‘apical section’, which isalways distinctly wider than the head andpronotum (Fig. 1.4.1). Deep pits are usuallypresent on the pronotum and elytra; they are alsooccasionally found on the head (Fig.1.4.1). Alltarsi possess three segments. Pselaphinae looselyresemble the Scydmaeninae in body outline butthe elytra of pselaphines are shorter and do notcover most of the abdomen. Pselaphinae arewidely distributed in North America.

A great diversity of pselaphines can becollected in damp, forested microhabitats, andwetlands, especially if the debris is removed,sifted, and processed in a Berlese funnel. Theyare sometimes collected at lights. Variousfavoured microhabitats include leaf litter, flooddebris, under bark (Fig. 9.15.2), grass clumps, inrotten logs or stumps (9.15.3), treeholes, inSphagnum, and under stones (Fig. 9.15.4). Manyspecies live occasionally or exclusively in thenests of ants and termites, feeding on the host

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larvae or engaging in trophallaxis with the hostants themselves (Akre and Hill 1973). Allpselaphines are predators; mostly of smallarthropods such as springtails and mites but themyrmecophilous and termitophilous speciesderive nutrients from their host colonies (seeabove).

9. 16 Pseudopsinae Ganglbauer 1895:Pseudopsis Newman 1834

Pseudopsis Newman can be recognizedby its distinctive habitus (Fig. 9.16.2),longitudinal ridges on the pronotum and elytra(Fig. 1.6.1), and a last (sixth visible) abdominaltergite that is notched apically with an apicalcomb of setae (Fig. 9.16.1).

This genus is widely distributed inNorth America, with two northerntranscontinental species (Herman 1975). Twospecies of Pseudopsis (P. subulata Herman andP. sagitta Herman) are known from ECAS andcan only be separated by differences in malegenitalia. However, individuals south of Canadaare most likely to be P. subulata. Twoundescribed species were reported from borealforest in Québec but no diagnosis was given todistinguish them from described species (Paquinand Duperre 2001). In ECAS, Pseudopsis hasbeen collected from sifted leaf litter and fromunderneath dung. Elsewhere the genus has beenfound in mammal burrows, fungi, mouldy pinecones, and in streamside litter (Herman 1975).One species, P. sagitta, was found to bedependent on old growth forests (Spence et al.1997) and both species are collected in numbersin mature forests (A. Brunke personalobservations). The feeding habits of Pseudopsisare unknown.

9.17 Scaphidiinae Latreille 1807

The Scaphidiinae have a distinctivebody shape, oval in outline and highly convex(Fig. 1.2.1), with long elytra that conceal half ofthe abdomen or more (dead specimens oftenhave very little abdomen visible dorsally) (Fig.1.2.2). Other distinguishing features include thehead, which is partially concealed dorsally (Fig.1.2.1) and the antennae that arise between theeyes (Fig. 1.2.3).

Scaphidiines are widely distributed inNorth America and inhabit decaying wood andfungi, especially fungus-covered logs. They alsooccur generally in leaf litter, under bark, and incompost. This subfamily may be profitably

collected by gathering leaf litter and fungi besideand under old rotting logs or stumps, sifting thecoarse material, and finally processing the siftatein a Berlese funnel. Larger species can becollected easily by hand as they are slow toescape. Some species can be collected by lightlytapping on the fruiting bodies of polypore fungiand mushrooms (Fig. 9.17.1). All scaphidiinesfeed on fungi and/or slime moulds (Newton1984) (Fig. 9.17.2).

9.18 Scydmaeninae Leach 1815

Scydmaeninae are best recognized bytheir habitus (Fig. 1.5.1) and small size (2.5mmor less) (Fig. 9.18.1), but some are similar toPselaphinae and even some Omaliinae in whichthe elytra nearly cover the abdomen. However,pselaphines always have short elytra and thoseOmaliinae with elytra nearly reaching theabdominal apex are always much larger than2.5mm and possess ocelli.

Scydmaeninae are widely distributed inNorth America and can be found in leaf litter,rotting logs and stumps, in treehole debris, moss,and under stones (Fig. 9.18.2). Some species areassociated with ant colonies, but are probablypreying upon other arthropods rather than theants themselves (O’Keefe 2000). They arecommon locally (Campbell and Davies 1991)but, due to their small size, they are most oftencollected when deep pockets of damp litter ormoss are processed in a Berlese funnel. Untilrecently, the Scydmaeninae were consideredtheir own family within Staphylinoidea but arenow placed within the staphylinine group ofsubfamilies in Staphylinidae (Grebennikov andNewton 2009). Scydmaenines are predators oforbatid mites (O’Keefe 2000), and some usetheir modified mandibles like a can-opener onthe genital and anal plates of their prey (Schmid1988).

9.19 Steninae McLeay 1825

Steninae are easily recognized by theirhabitus, particularly their globular eyes andcylindrical pronotum (Fig. 1.9.1). They looselyresemble Megalopinus (Megalopsidiinae) but inSteninae the antennae insert between the eyesand are not strongly clubbed (Fig. 1.9.3).

Two genera are widespread in NorthAmerica, the very diverse Stenus Latreille andthe rarely seen Dianous Leach (two similarspecies). Dianous differ from Stenus in havingsmaller but still globular eyes (Fig. 1.9.2). Stenus

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species can be collected from a variety of moisthabitats but Dianous seems to prefer the edges ofsmaller, headwater streams. Stenus can be sweptor beaten from various periaquatic plants (Fig.9.19.1), hand-collected from the shores of lakesand ponds, sifted from debris beside streams andrivers (Fig. 9.19.2), and found under bark (Fig.9.19.3). Some species are highly generalist intheir habitat choice but others appear to prefercertain substrates or wetlands such as fens orseeps. Individuals of some species escape bypropelling themselves across the water surfaceusing secretions from their pygidial glands(Stevens and DeKimpe 1993). Other species canalso slowly swim at the surface using lateralundulations of the abdomen. Stenines arepredators of small arthropods includingspringtails and most use a protrusable, adhesivelabium to capture prey; some species simply usetheir mandibles (Betz 1998). Use of the modifiedlabium is most prevalent in species which huntfor prey in complex vegetation (Betz 1998).

9.20 Tachyporinae McLeay 1825

Tachyporines can be usually recognizedby their tapering abdomen, but identificationshould be confirmed by the followingcombination of characters: abdomen with sixsternites (Fig. 1.12.1), elytral epipleuron with acarina (Fig. 1.14.5), neck absent (Fig. 1.11.3),procoxae similar in shape to profemora (Fig.1.12.6), antennomeres not extremely elongate(i.e., unlike Habrocerus), antennae inserted infront of the eyes (Fig. 1.12.2), pronotal marginssmooth, and dorsal abdomen margined (Fig.1.12.5) (except in one genus with a stronglytapered abdomen [Fig.1.14.2]).

Tachyporines are widespread in NorthAmerica and these small to medium-sizedStaphylinidae are most often noticed as theydisperse to their preferred microhabitat.Common species that are relatively general intheir habitat requirements include Tachinuscorticinus Gravenhorst, found in leaf litter,fields, decomposing organic matter, and understones (Fig. 9.20.1), and Sepedophilus littoreus(L)., found in rotting wood, compost, flooddebris, under loose bark, and in rotting fungi(Fig. 9.20.2); both species are native toEurope. Coproporus ventriculus (Say), is a verycommon native species found under the bark ofboth deciduous and coniferous trees, especiallywhen slime moulds are present (Fig.9.20.3). Sifting or flooding patches of leaves orvery wet sphagnum moss near creeks, bogs, or

waterfall areas will sometimes yieldNitidotachinus scrutator (Gemminger &Harold), (Fig. 9.20.4). Many genera ofTachyporinae, including the common Lordithonkelleyi (Mälkin) (Fig. 9.20.5), can be collectedfrom various types of fungal fruiting bodies.Some species of Tachyporus Gravenhorst inhabitprairies, old fields, and agricultural crops wherethey can by swept from the vegetation in theevening (Fig. 9.20.6). Tachyporines can also befound in dung, in carrion, and along the marginsof lakes and ponds. They are readily sampledfrom litter and fungi using a sifter. Sometachyporines are exclusively predaceous ormycophagous but many appear to be largelyopportunistic on both arthropods and fungi.

9.21 Trichophyinae C.G Thomson 1858:Trichophya pilicornis (Gyllenhal 1810)

Trichophya pilicornis can be recognizedby a combination of the characteristic antennae(Fig. 1.21.2), the heavily setose dorsal surface,body widest at elytra and the distinct neck (Fig.1.22.1). This species resembles someTachyporinae but possesses specialized antennaeand strongly enlarged first and secondantennomeres (Fig. 1.21.2).

This species also occurs in thePalearctic region and is suspected of beingintroduced into northeastern North America viahuman activity (Ashe and Newton 1993). It wasfirst recorded in 1895 from New York and wasknown from Canada (Ontario) by 1905 (Asheand Newton 1993). Unlike most other non-nativeStaphylinidae, it is very common in the borealforest (Paquin and Duperre 2001) and apparentlyless common south of this ecozone. Trichophyapilicornis can be collected by sifting leaf litter(Fig. 9.21.1) or by pitfall trapping. Species ofTrichophya are mycophagous and facultativelypredaceous (Ashe and Newton 1993).

10. The tribes and subtribes of Staphylininaein ECAS

10.1 Diochini Casey 1906:Diochus schaumi Kraatz 1860

Diochus schaumi is distinctive amongStaphylininae for its combination of a thin neck(about ¼ the width of the posterior head) (Fig.2.3.1), broad and flattened pronotum, and elytrathat do not overlap (Fig. 10.1.1). This species isvariable in colour (Fig. 10.1.2).

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Map 1. Distribution of Diochus schaumi Kraatz in eastern Canada and adjacent United States. Recordsinclude data from Smetana (1982).

This species is widely distributed ineastern and southern North America, but is notfound west of the Rocky Mountains (Smetana1982). Its distribution is distinctly south of theboreal forest region and is most typically foundin forests with southern characteristics (Map 1).Data from Map 1 includes localities found inSmetana (1982).

Eastern Canada: ON, QCAdjacent U.S: WI, MI, IN, OH, PA, NY, NH,ME

Diochus schaumi has been collected inECAS from March to December, with most

individuals collected in April and May.However,copulation occurs in fall, as late as November(Fig. 10.1.3).

This species is frequently collected bysifting litter and other debris in wet areasincluding slough forests, swamps, marshes, andbogs. Smetana (1982) also reported it from thenests of Microtus pennsylvanicus (meadow vole)and from ‘debris at the base of grass tufts’.

10.2 Othiini C.G. Thomson:Atrecus Jacquelin du Val 1856

Atrecus is distinguished from otherStaphylininae by the following combination of

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characters: a pair of plates anterior to theprosternum (Fig. 2.1.2), thick neck (Fig. 2.4.1),antennal bases closer to each other than to thenearest eye (Fig. 2.4.1), mandibles projectingforward (Fig. 2.4.1), and elytra not overlapping(Fig. 10.2.1).

Two species occur in ECAS (A.americanus (Casey) and A. macrocephalus(Nordmann)) mainly under the bark ofconiferous trees (Fig. 10.2.2). They cansometimes be collected in debris around thebases of old trees (Smetana 1982). Atrecus istranscontinental in Canada and occurs in easternand western United States. This genus will becovered in greater detail in a future publicationon Othiini and Xantholinini.

10.3 Anisolinina Hayashi 1993:Tympanophorus puncticollis (Erichson 1840)

Tympanophorus puncticollis can bedistinguished from other Staphylininae by thecombination of a neck with coarse lateralpunctures (Fig. 2.2.12), lack of plates anterior tothe prosternum, elytra that do not overlap (Fig.10.3.1), small head (Fig 2.2.12), and hoof-likeapical labial palpomere (Fig. 2.2.14). Thedistinctive labial palpi will quickly distinguishthis species from all other Staphylininae.

This species is transcontinental inCanada in the boreal ecozone; it inhabits moresouthern forests in eastern North America,occurring west to Minnesota and as south asAlabama. Its range in ECAS is given by Map 2.Tympanophorus puncticollis is newly recordedfrom Indiana, Pennsylvania and New York:

UNITED STATES: IN: Tippcanoe Co., 40.42 -86.89, 25-VII-1971, N.M. Downie, 2 (FMNH);31-VII-1971, N.M. Downie, 1 (FMNH).PA: Allegheny Co., Pittsburgh, 40.43 -80.00,VII, 1 (CMNH). Monroe Co., Delaware WaterGap, 40.9667 -75.1167, A. T. Slosson, 1(AMNH). NY: Madison Co., Darts Corners, 42.8-75.5, VIII, K.W. Cooper, 1 (AMNH). QueensCo., Far Rockaway, 40.60 -73.75, 22-VI-1904,L.B. Woodruff, 1 (AMNH).

Eastern Canada: ON, QC, NB, NS, NLAdjacent U.S.: MI, IN, PA, ME, NY, NH

Tympanophorus puncticollis has beencollected in ECAS from May to September withan increase in abundance occurring in June-July.

This species is rarely collected and

consequently very little has been published aboutits natural history. The scant habitat data suggestthat it occurs in moist (with ferns) or boggy areaswithin forested habitat but it has also beencollected at the mouth of a fox burrow, along acobblestone shoreline of a lake, in a 'lumberyard', and in rotting sawdust in an abandonedsawmill (Fig. 10.3.2). Specimens have beencaught by pitfall, window pane, and raisedmalaise (six feet high) traps, suggesting that it ishighly mobile. One individual was extractedfrom moss on a dead balsam fir using a Berlesefunnel (Paquin and Duperre 2001) and T.puncticollis was newly reported from Maine byMajka et al. (2011) from a black spruceplantation. Paquin and Duperre (2001) suggesteda possible association with the forest canopy asseveral individuals were captured in their canopyfunnel traps.

10.4 Philonthina Kirby 1837

Members of the subtribe Philonthinacan be recognized among the Staphylininae bythe following combination of characters: elytranot overlapping, without plates anterior to theprosternum, neck present and impunctate (Fig.2.2.2), antennae closer to the nearest eye than toeach other (Fig. 2.2.3), punctures of pronotumeither not arranged in dorsal rows or these rowsconsisting of at least 4 punctures (Fig. 2.2.6), andwithout empodial setae between the tarsal claws(Fig. 2.2.8).

Philonthina is a widespread and diversesubtribe of Staphylininae in North America andcontains many abundant species that areassociated with human activity. One largespecies, Philonthus politus (L.), is often found incompost, around livestock, and on carrion (Fig.10.4.1). Philonthus caeruleipennis Mannerheimis a common species found in rotting organic(especially fungi) matter in or near forests; thisone (Fig. 10.4.2) was found in a compost heap ina wooded backyard. Bisnius blandus(Gravenhorst), a forest specialist, occurs mainlyin rotting fungi but also at carrion (Fig. 10.4.3).These and many other Philonthina can becaptured in abundance using pitfall or pan trapsbaited with dung, mushrooms, or carrion. Somespecies are associated with older trees: adults andlarvae of Hesperus apicialis (Say) can becollected from debris in old tree holes andstumps (Fig. 10.4.4), and Laetulonthus laetulus

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Map 2. Distribution of Tympanophorus puncticollis (Erichson) in eastern Canada and adjacent UnitedStates.

(Say) is attracted to large, freshly killed treeswhere it hunts invertebrates attracted to sap(Fig.10.4.5). The majority of Philonthina speciesin ECAS inhabit moist debris near or at the edgeof lakes, ponds, rivers, and streams. Sifting thisdebris often yields many Erichsonius Fauvelspecies, the largest in ECAS being E. rosellusFrank (Fig. 10.4.6). This subtribe will be coveredcompletely in a future publication.

10.5 Quediina Kraatz 1857

Members of the subtribe Quediina canbe recognized among the Staphylininae by thefollowing combination of characters: elytra notoverlapping, without plates anterior to theprosternum, neck present, impunctate andusually very broad (Fig. 1.12.8), antennae closerto the nearest eye than to each other (Fig. 2.2.3),pronotum impunctate or with dorsal rows ofthree or fewer punctures (Fig. 2.2.7), withempodial setae between the tarsal claws (Fig

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2.2.9) and last maxillary palpomere not distinctlynarrower than the preceding (Fig. 1.12.8).

The Quediina in the classic sense havebeen shown to be an artificial grouping of genera(Solodovnikov and Schomann 2009,Chatzimanolis et al. 2010) but all includedgenera occurring in ECAS, except Heterothops,belong in this subtribe sensu Chatzimanolis et al.(2010); Heterothops has been placed in thesubtribe Amblyopinina. Species of this groupinhabit a diversity of habitats similar to thosefrequented by Philonthina but most are quiteinfrequently collected relative to the latter. Onecommonly encountered species is Quediusplagiatus (Mannerheim), which occurs under thebark of conifers, beech, and larger birch trees(Fig 10.5.1). Quedius mesomelinus (Marsham) isa European species often found in compost,gardens, and rotting fungi (Fig. 10.5.2). SomeQuediina are strongly associated with the edgesof lakes, wetlands, and streams. Sifting ortreading old rotting leaves, moss, and otherdebris at these edges will often yield manyindividuals of Acylophorus Nordmann,distinctive in Staphylinini for its elbowedantennae (Fig. 10.5.3). Several species ofQuediina are obligate or frequent inhabitants ofbeaver and muskrat lodges; Hemiquedius ferox(LeConte) is commonly encountered this waybut also occurs in debris along lakeshores and inbogs (Fig. 10.5.4). The species of Quediina willbe covered in greater detail in a futurepublication.

10.6 Tanygnathinina Reitter 1909:Atanygnathus bicolor (Casey 1915)

Atanygnathus bicolor can be easilyrecognized among the Staphylininae by the lackof a visible neck, elongate maxillary palpi (Fig.2.2.5), and its unique habitus (Fig. 1.12.12). It isalso unique for its 5-4-4 tarsal formula. In overallappearance, it loosely resembles someTachyporinae but lacks an epipleural ridge on thelateral portion of the elytron (Fig. 1.12.11).

This species is distributed mainly in thesouth-central and southeastern United States,extending along the east coast to New Hampshireand westward to Michigan. It is newly recordedfrom Canada (Ontario) from a single specimencollected in Algonquin Provincial Park, and fromthe state of Michigan, both considerabledistances from the northernmost recordpreviously known in New Hampshire (Smetana1990) (Map 3).

CANADA: ON: Nipissing Distr., AlgonquinProv. Pk., Broadwing Lake, 45.5977 -78.5275,22-VIII-2008, island with sphagnum bog, pitfall,B. Wells, 1 (DEBU).UNITED STATES: MI: Berrien Co., Mud LakeBog, 28-IV-1984, sphagnum, L. E. Watrous, 10(FMNH).

Eastern Canada: ONAdjacent U.S.: NH, MI

Based on the small number ofspecimens from ECAS, this species appears tohave both a spring and late summer/fall peak inadult activity.

This species is strongly associated withvery wet moss and debris in wetlands and lakes(Smetana 1971). Specimens have been siftedfrom grass, sphagnum and litter in bogs, 'sweptfrom grass by a pond' (Smetana 1990), andcollected in a pan trap placed in a sphagnum maton a lake. The specimens from Ontario andMichigan likely represent true populations for anotherwise southeastern species, as they werecollected in the typical specialized habitat andfurther collecting may reveal that A. bicoloroccurs broadly across ECAS.

10.7 Amblyopinina Seevers 1944:Heterothops Stephens 1829

The genus Heterothops can berecognized by the combination of: elytra notoverlapping, without plates anterior to theprosternum, neck present, impunctate andusually very broad (Fig. 1.12.8), antennae closerto the nearest eye than to each other (Fig. 2.2.3),pronotum impunctate or with dorsal rows ofthree or fewer punctures (Fig. 2.2.7), withempodial setae between the tarsal claws (Fig2.2.9) and last maxillary palpomere distinctlynarrower than the preceding (Fig. 2.2.11) –compared with that of Quediina (Fig. 1.12.8).

Heterothops is the only representativein ECAS of the otherwise south-temperateAmblyopinina. This genus is broadly distributedin North America and has the highest diversity inthe West. Heterothops are generally rarelycollected and sifting moist litter is the mostproductive method to obtain specimens.Heterothops fusculus (LeConte) is an easilyrecognized, large Heterothops (Fig. 10.7.1) thatcan be found in yard compost in addition to morenatural habitats, while the other four species inECAS are darker and more obscure. One of thesespecies, Heterothops marmotae Smetana, has

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Map 3. Distribution of Atanygnathus bicolor (Casey) in eastern Canada and adjacent United States.Records include data from Smetana (1990).

only been collected in groundhog (Marmotamonax) burrows in the vicinity of Ottawa,Ontario, Canada (Smetana 1971b). It is surelymore widespread but its apparently restrictedhabits have probably prevented its collectionelsewhere. The species of Heterothops will becovered in greater detail in a future publication.

10.8 Xantholinini Erichson 1839

Xantholinini may be easily recognizedamong the Staphylininae by their slender, linearhabitus (Fig.10.8.2), thin neck (Fig.2.1.2),

overlapping elytra (Fig.2.1.1), elbowed antennae(Fig.10.8.2), and a pair of plates anterior to theprosternum (also in Othiini) (Fig.2.1.2).

This is a poorly collected group that isbroadly distributed in North America. Species ofXantholinini generally live within litter or indecaying organic matter and are therefore bestcollected by sifting different types of litter ordebris. Passive methods of collection such asWinkler extractors or Berlese funnels are evenmore effective as many species adopt a coiledposition upon disturbance and remain motionless

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for a long time (for a detailed account of thisbehaviour see Yamazaki 2007). Stictolinusflavipes (LeConte) (Fig. 10.8.1) andLithocharodes longicollis (LeConte) (Fig.10.8.2) are both commonly collected in varioustypes of litter using these passive techniques. Incontrast, species associated with livestock orcompost are more active and are easily collectedin great numbers using an aspirator or softforceps. Two such species include the introducedGyrohypnus fracticornis (O.F. Müller) (Fig.10.8.3) and Gauropterus fulgidus (Fab.) (Fig.10.8.4). Peeling the bark of conifer or beech treeswill usually yield individuals of the nativeNudobius cephalus (Say) (Fig. 10.8.5). Somespecies including Oxybleptes kiteleyi Smetanahave only be captured in small numbers by usingflight intercept or pan traps in open areas withlarge trees (Fig. 10.8.6). The biology of theseand most Xantholinini remains very poorlyknown. The species will be covered in greaterdetail in an upcoming publication.

11. The species of Staphylinina in ECAS

11.1 Creophilus maxillosus (Linnaeus 1758)

Creophilus maxillosus is easilydistinguished from other Staphylinina by itslargely impunctate pronotum (Fig. 3.1.1) andthick covering of black and yellow setae (Fig.11.1.1). It cannot be confused with any otherspecies of the subtribe.

This is a very widespread species,which occurs across North America, northernCentral America, the West Indies, the entirePalearctic region (Newton et al. 2000), Chile,Argentina (Navarrete-Heredia et al. 2002), andPeru (Asenjo and Clarke 2007). Its range inECAS is given by Map 4.

Eastern Canada: ON, QC, NB, NL, NS, PE (AllECAS Provinces)Adjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS States)

In ECAS, Creophilus maxillosus hasbeen collected from February to December, withno apparent seasonality. This species overwintersas an adult.

Adults and larvae of this species arefound primarily on carrion of all kinds and rarelyon dung, compost, or at UV lights. Larvae aretypically found on larger, more persistent

carrion. C. maxillosus inhabits grassy and openforested habitats and is often encountered on lakeand ocean shorelines (Majka et al. 2008).

Adults and larvae are predators, mainlyof maggots and adult flies, but also of otherarthropods attracted to carrion. Adults will formtheir bodies into a tight ball and roll off carrioninto leaf litter when disturbed (Fig. 11.1.2). Fordefense against other arthropods, C. maxillosushas an eversible Y-shaped gland at the apex ofthe abdomen that produces iridodiols and otherrepellant compounds (Huth and Dettner 1990).Other species of Staphylinina also have thisgland and produce similar secretions dominatedby iridodial (Huth and Dettner 1990).

Two subspecies occur in ECAS: thenative and widespread villosus Gravenhorst andthe Palearctic maxillosus L., which is knownfrom scattered records from Ontario, Québec,and Massachusetts as early as 1929 (Québec).Creophilus m. maxillosus can be most reliablydifferentiated from C. m. villosus by the darkpubescence on the hind angles of the head (Fig.11.1.3), which is pale yellow in the nativesubspecies (Fig. 11.1.4).

11.2 Dinothenarus badipes (LeConte 1863)

Dinothenarus badipes is distinguishedfrom other species of Staphylinina in ECAS byits entirely black body (Fig. 4.2.1), entirelyorange legs (Fig. 11.2.1), golden setae at thebases of each abdominal segment (Fig. 3.4.3)and a spindle-shaped apical maxillary palpomere(Fig. 3.4.4). Ocypus brunnipes Fab. is similar incolouration but lacks the golden abdominal setaeand its apical maxillary palpomere is rectangular.

This species is broadly distributed overeastern North America. Its range in ECAS isgiven in Map 5. It is newly recorded from theprovince of Prince Edward Island:

CANADA: PE: Queens Co., Harrington, barleyfield, pitfall trap, 14-VI-2004, C. Noronha, 6(ACPE); Johnstons River, rutabaga field, 14-XI-1983, M.E.M. Smith, 1 (ACPE); Kinlock, 2-VI-1962, B. Wonnacott, 1 (UPEI).

Eastern Canada: ON, QC, NB, NS, PEAdjacent U.S.: MI, IN, PA, NY, VT, NH, ME

Dinothenarus badipes has beencollected in ECAS from January to December,with a single peak in abundance in April-Junedepending on latitude.

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Map 4. Distribution of Creophilus maxillosus (L.) in eastern Canada and adjacent United States.

This is a common species that has beencollected from under logs and stones, leaf litter,moss, and loose bark in a wide variety of habitatsincluding forests, the shores of wetlands andlakes, along streams and rivers, orchards, oaksavannah, and tallgrass prairies. However, D.badipes does not occur in the boreal forest,though it may occasionally occur in transitionareas. This species is rarely found at carriontraps. Snider (1984) successfully cultured D.badipes on a diet consisting solely of millipedes,but this species is likely a generalist predator ofground-dwelling arthropods and it is also knownto consume adult click beetles (Hawkins 1936).

11.3 Dinothenarus capitatus (Bland 1864)

Dinothenarus capitatus is easilydistinguished from other Staphylinina by itsreddish-orange head and darker body (Fig.3.4.2). Specimens with heads secondarilydarkened due to chemical killing agents ordrying (Fig. 4.1.1) can be identified toDinothenarus using the characters in the key,and distinguished from D. badipes (LeConte) bythe variegated pattern of light and dark setae onthe dorsal pronotum, and the dark tibiae (Fig.3.4.2).

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Map 5. Distribution of Dinothenarus badipes (LeConte) in eastern Canada and adjacent United States.

This species is transcontinental inCanada and occurs as far south as West Virginiain the eastern United States. Its range in ECAS isgiven in Map 6.

Eastern Canada: ON, QC, NB, NL, NSAdjacent U.S.: MI, OH, PA, NY, VT, NH, ME

Dinothenarus capitatus is collected inECAS from April to November, with peaks inabundance in May and August-September.

This forest-inhabiting species has beencollected primarily on the dung of variousmammals or on carrion, but sometimes occurs ondecaying fungi. It can be sifted from forest litter

and, although it has occasionally been collectedin disturbed areas as it disperses between forestpatches, it is a strong indicator of undisturbedforest conditions (Klimaszewski et al. 2008) andmany specimen labels bear the words "mature"or "old growth forest". The current range ofDinothenarus capitatus in ECAS appears to beshrinking relative to its historical distribution,especially near developed areas.

11.4 Ocypus brunnipes (Fabricius 1781)

Ocypus brunnipes can be easilyrecognized by its all black body and dull orangelegs (Fig. 5.1.1). The only similar species is

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Map 6. Distribution of Dinothenarus capitatus (Bland) in eastern Canada and adjacent United States.

Dinothenarus badipes (LeConte), from which itcan be distinguished by the lack of golden setaeat the bases of the abdominal segments (Fig.3.6.3), and by its rectangular apical maxillarypalpomere (Fig. 3.6.2).

This species is native to Europe, Russia,the Caucasus, and Turkey (Herman 2001) andhas been accidentally introduced into NorthAmerica, where it is so far recorded only fromNew Hampshire and Massachusetts (newlyreported) (Map 7).

UNITED STATES: MA: Essex Co., Ipswitch,42.68 -70.85, 26-VII-1973, L.L. Pechuman, 1(CUIC).

Both North American collections of O.brunnipes have been in late summer.

This non-native species was firstrecorded from North America by Newton (1987),based on a single collection of one male and onefemale from New Hampshire in 1966. In 1973this species was collected again (1 male) but thistime from Massachusetts. Ocypus brunnipes maybe established in New England at very lowdensities. This species is flightless and inhabitsboth forested and open, disturbed areas in itsnative Palearctic range (Deichsel 2006).

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Map 7. Distribution of Ocypus brunnipes (Fab.) in eastern Canada and adjacent United States.

11.5 Ocypus nitens (Schrank 1781)

Ocypus nitens is unique among theStaphylinina in ECAS for its combination ofblack body, black legs, and distinctively shortelytra (distinctly shorter than the pronotum whenmeasured at the middle) (Fig. 3.6.1).

This species is native to Europe, Russia,the Caucasus, Turkey, and Iran (Herman 2001).Its range in North America is mostly limited tothe coastal northeastern United States and werecord it here for the first time from Maine andNew York. Its range in ECAS is given in Map 8.

UNITED STATES: ME: York Co., Kittery,43.0833 -70.7333, 5-IV-1989, G. Clark, 1(DENH). NY: Saratoga Co., Ballston Lake, atporch light, 21-IV-2010, K. Hillig, 1, based onphotographic record frombugguide.net/node/view/387557.

Adjacent U.S.: NY, NH, ME

This species has been collected in NorthAmerica year-round, with a large peak inabundance in April-May and a smaller peak inSeptember.

Ocypus nitens is a non-native speciesthat was first reported in North America from

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Map 8. Distribution of Ocypus nitens (Schrank) in eastern Canada and adjacent United States.

Massachusetts and southeastern New Hampshire(Newton 1987). It was first collected inMassachusetts in 1944 from sifted debris and hassince become common in forested and openhabitat in Massachusetts and New Hampshire(Fig. 11.5.1). This species is also found atcarrion-baited pitfall traps and often wandersinto dwellings. It has similar habits in its nativerange (Balog et al. 2003) and is pronouncedlysynanthropic. All examined specimens withexposed hind wings were brachypterous. Ocypusnitens has since expanded its range to Maine by1989, Rhode Island by 1995 (Sikes 1995) andNew York by 2010. This species will likely

continue to spread westwards into easternCanada in the near future.

11.6 Ontholestes cingulatus(Gravenhorst 1802)

Ontholestes cingulatus can be distinguished fromother Staphylinina by its distinctive goldenabdominal apex (Fig. 6.1.1). Older, contracted,or dirty specimens may be identified toOntholestes by the sharp anterior angles of thepronotum (Fig. 3.2.1) and distinguished from O.murinus (L.) by the bicoloured legs and antennae(Fig. 11.6.1). This species is transcontinental in

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Map 9. Distribution of Ontholestes cingulatus (Gravenhorst) in eastern Canada and adjacent United States.

Canada and occurs in the central and easternUnited States. Although this species iswidespread, it had not been officially recordedfrom Michigan, Ohio, Vermont, or PrinceEdward Island. Its range in ECAS is given byMap 9. The occurrence of Ontholestes cingulatusin these states and province is confirmed basedon numerous historical and recent collections.

CANADA: PE: Queens Co., Harrington, barleyfield, pitfall trap, 16-IX-2004, C. Noronha, 3(ACPE); St. Patricks, compost, 20-VIII-2002,C.G. Majka, 1 (CGMC); St. Patricks, old field,compost, 25-VI-2003, C.G. Majka, 1 (CGMC).

UNITED STATES: MI: 207 specimens.OH: 95 specimens. VT: Addison Co., Bristoland Middlebury townline, route 116, pitfall traps,4 to12-XI-1976, R. Davidson, 6 (CMNH).Bennington Co., Dorset, Polyporus sulfureus, 14-VII-1970, J.F. Lawrence, 3 (MCZ); Manchester,beech-maple forest, ex. ‘several clusters ofConopholis americana’, 17-VIII-1969, C.T.Parsons, 1, 12-VIII-1971, 3 (MCZ). LamoilleCo., Stowe, 1 (AMNH). Orleans Co., EastCharleston, 21-VI-1967, M.A. Deyrup, 1, oldgrass heap, 27-VI-1967, 1 (AMNH). WindsorCo., Woodstock, 20-VIII, 2 (MCZ).

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Eastern Canada: ON, QC, NB, NL, NS, PE (AllECAS provinces)Adjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Ontholestes cingulatus has beencollected in ECAS from February to September,with a peak in abundance in May-June.

This species is common and most oftencollected on dung and smaller carrion but alsoregularly occurs in decaying fungi and incompost piles. This species is a strong flier andis captured frequently in flight-intercept andmalaise traps, and infrequently at UV lights.

Males guard recently mated females asthey oviposit, chasing and 'grappling' with anymales that approach her (Alcock 1991). Matingoccurs with males on top of females and bothsexes facing the same direction (Alcock 1991);quite different from smaller staphylinids thattypically mate end-to-end. Both larvae and adultscan be reared on a diet of muscoid flies (Schmidt1999). Ontholestes cingulatus was considered amoderate indicator of undisturbed forest byKlimaszewski et al. (2008) and was observed toprefer shaded carrion by Hobischack et al.(2006); however, collection data suggest it alsooccurs in open forests and backyards.

11.7 Ontholestes murinus (Linnaeus 1758)

Among the species of Staphylinina,Ontholestes murinus can be identified to genusby its acute anterior pronotal angles (Fig. 3.2.1)and distinguished from O. cingulatus(Gravenhorst) by the dark legs, completely paleantennae and apex of the abdomen with silversetae (Fig. 6.2.1).

This species is native to Europe, Russia,Georgia, Armenia, Turkey, Iran, Kazakhstan,Kyrgyzstan, and China (Herman 2001). In NorthAmerica, it is only known from the AvalonPeninsula of Newfoundland (Map 10) where ithas been collected in May to September.

North American specimens ofOntholestes murinus were first collected in 1949in a meadow, on cow dung and in a barn(Smetana 1981). One additional specimen wascollected in 1986 and the continuedestablishment of this exotic species needsreaffirmation.

The Platydracus cinnamopterus Complex

Diagnosis:Small (12-19 mm), slender, parallel-

sided Platydracus; body and appendagesgenerally light to dark reddish brown, uniformlycolored or not on head and pronotum to alwaysvariegated on abdomen, with darker venter(except abdominal apex, more or less reddish)and black scutellum; pubescence light to dark, onabdomen forming more or less distinct, pairedblack pubescent lines near middle and diffuselighter pubescence laterally; head and pronotumwith dense umbilicate punctures except for acomplete mid-longitudinal impunctate line on thepronotum at least one puncture diameter wide;antenna pubescent from 4th antennomere;mandibles stout, apically acute, each bearingthree stout median teeth (two dorsal, one ventralon left, one dorsal and two ventral on rightmandible, in both cases with only two teethvisible in dorsal view); mentum brown,sclerotized, anterior edge emarginate; pronotumabout as wide as long; sutural length of elytraabout 0.6 times as long as pronotum; protibiausually with two stout setae in row on externaledge and one posteriorly; apex of 6th visibleabdominal sternite in male with broadly roundedemargination, in female with broadly rounded,non-emarginate apex, sexual dimorphismotherwise absent externally and limited to venterof genital segment and genitalia; aedeagus not orvery inconspicuously asymmetrical.

A species known as Staphylinus orPlatydracus cinnamopterus (Gravenhorst)meeting the above diagnosis was for nearly twocenturies considered the most common andwidespread Platydracus species in eastern NorthAmerica. In the early stages of a revision of thisgenus for the New World, it became apparent tothe second author (Newton 1973) that threespecies with quite different male genitalia werebeing confused under this name. After furtherstudy of type material of all relevant names,including several that were long consideredsynonyms of P. cinnamopterus, it wasestablished that two of the species had availablenames and the third was undescribed. The threespecies are broadly sympatric in eastern NorthAmerica and all occur within at least thesouthern part of ECAS. They are closely similarexternally and difficult to distinguish (especiallyolder or darker specimens in poor condition)without use of male genitalia (but see externalcharacters in key). In order to clarify the status ofthe available names and allow recognition of

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Map 10. Distribution of Ontholestes murinus (L.) in eastern Canada and adjacent United States.

these species, the new species (P. praetermissusNewton, spec. nov.) is formally described hereand the other two species (P. cinnamopterus andP. zonatus, both Gravenhorst 1802) areredescribed and their identities fixed by reviewof type material, including designation oflectotypes where necessary.

11.8 Platydracus cinnamopterus(Gravenhorst 1802)

Staphylinus cinnamopterus Gravenhorst, 1802:164; Platydracus cinnamopterus: Newton1973, Smetana and Davies 2000, Herman

2001, etc. (see Herman 2001: 3447 foradditional literature citations)Type material: Lectotype, male, in ZMHB,here designated by Newton, with labels:"5959" [white printed label];"cinnamopterus Gr. Am. Spt." [green,bordered, handwritten label]; and redlectotype designation label by Newton dated1978. Paralectotypes: 1 male and 2 females,in ZMHB, with no original labels, each withyellow paralectotype designation labels byNewton dated 1978.

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Map 11. Distribution of Platydracus cinnamopterus (Gravenhorst).

Other material examined: 1386 specimensthroughout the range of the species, all but a fewbeing males with examined aedeagus. Full datawill be presented in the pending revision ofPlatydracus by Newton, but all records thatcould be georeferenced are shown in Map 11.

Description:With the characters of the P.

cinnamopterus Complex (see above), plus:average pronotal length 2.53 mm (n = 10); head,pronotum and elytra usually light to mediumreddish, less often dark red, pronotum evenlycolored; antenna not reaching middle ofpronotum when extended posteriorly,antennomere 9 distinctly transverse; impunctatemedian line of pronotum at least one puncturediameter wide at narrowest point, usually abouttwo puncture diameters wide; emargination ofsixth visible male sternite in ventral viewmoderately deep, about 1/4 as deep as wide; apexof median lobe of aedeagus projected, theconverging sides of the projection in parameral

view slightly convex and forming an acute angle,the apex broadly rounded; sides of median lobein lateral view even, not toothed.

Platydracus cinnamopterus may bedistinguished from all other species of the genusin ECAS by the combination of generally reddishcolor (light to very dark) with uniformly coloredhead and pronotum (Fig. 7.9.1, 7.9.4), blackscutellum, and variegated abdomen with sparsegold setae only, and paired, patches of blackvelvet setae on the first five visible segments(Fig. 7.9.2); antennae with distinctly transverseantennomere 9 (Fig. 7.9.7); and the narrow butcomplete impunctate medial line of pronotumabout 2 puncture diameters wide at narrowest(Fig. 7.9.5). However, it is definitivelydistinguished from the other members of the P.cinnamopterus Complex, P. praetermissus andP. zonatus, by the structure of the aedeagus (Fig.7.9.6 and 11.8.2).

The species is widely distributed in theforested areas of eastern North America: fromNova Scotia, west to southern Manitoba, south to

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Map 12. Distribution of Platydracus cinnamopterus (Gravenhorst) in eastern Canada and adjacent UnitedStates.

Texas and east to the Florida panhandle (Map11). Its range in ECAS is shown in greater detailin Map 12. Note that historical records of P.cinnamopterus in the literature up to the presenttime are unreliable since they may refer to any ofthe three species of the P. cinnamopterusComplex.

Eastern Canada: ON, QC, NB, NSAdjacent U.S.: MI, IN, OH, PA, NY, NH, ME

In ECAS, P. cinnamopterus has beencollected from January through November, witha majority (55%) found in May or June.

This common species is mostfrequently found under the loose bark ofhardwood trees but is also collected in leaf litter,under stones and logs and in rotten wood. It alsofrequents decaying plant matter such as flooddebris (Fig. 11.8.1) and grass clumps. Otherspecimens have been collected on beaches of theGreat Lakes under drift, from fungi, at a UVlight, and from a beaver lodge. Unlike the othertwo species of the complex, P. cinnamopterus isoften found in disturbed areas under patio stones,in gardens and in compost. The relatively shortantennae compared to the other members of thecinnamopterus complex may be correlated with

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Map 13. Distribution of Platydracus comes (LeConte) in eastern Canada and adjacent United States.

greater maneuverability within subcorticalmicrohabitats and under objects. The presence ofa rufous morph (Fig. 7.9.4) is unique to P.cinnamopterus within the cinnamopterusComplex. The description of larvae and pupaeof this species by LeSage (1977) may becorrectly assigned to P. cinnamopterus sinceover 85% of the specimen records of the P.cinnamopterus Complex from Ontario andQuébec are P. cinnamopterus, and P.praetermissus n. sp. is not known from Canada.

11.9 Platydracus comes (LeConte 1863)

Platydracus comes may bedistinguished from other species of the genus inECAS by the combination of elongate, darkspots on the elytra (Fig. 7.5.1), the median line ofgolden setae on the scutellum (Fig. 7.5.1), andthe complete impunctate median line of thepronotum (Fig. 7.5.2). This species is easilyconfused with the similarly coloured P.maculosus (Gravenhorst) (Fig. 11.9.1) but it ismuch smaller (13-19 mm) and has a completeimpunctate median line of the pronotum.

This species is eastern in distribution,with most specimens collected in the

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southeastern United States. It reaches thenorthern limit of its distribution in ECAS (Map13), with most of the northeastern records pre-1940. Platydracus comes is newly recorded fromOhio based on these records.

UNITED STATES: OH: Athens Co., WaterlooTwp., 39.3166 -82.2166, 24-VII-1935, 1 (CNC).Hamilton Co., Cincinnati, 39.1 -84.5, 22-VII-1900, C. Drury, 2 (CNMH); 31-VII-1900, T.Drury, 1 (MCZ); 7-VIII-1900, C. Drury, 1(CNMH).

The tentative record of Platydracuscomes from the province of Québec listed inCampbell and Davies (1991) as '?' represents amisidentification of P. viridanus. Platydracuscomes probably does not occur regularly inCanada, but occasional migrant individualsmight be found in southernmost Ontario.

Adjacent U.S.: IN, OH, PA, NY

In ECAS, Platydracus comes has beencollected in July to September, with nearly allspecimens taken in July.

This species lives in pine or mixed-pineforests, usually occurring with dung and carrioncommunities and infrequently collected atcompost-baited traps or UV lights.

11.10 Platydracus exulans (Erichson 1839)

Platydracus exulans is distinguishedfrom congeners by its bronze-green metallichead and pronotum (Fig. 11.10.1), and theunevenly spaced pronotal punctures (Fig.7.2.1). The only other Platydracus with unevenpronotal punctures is P. praelongus, however, P.exulans differs by the narrower medianimpunctate area of the pronotum (Fig. 7.5.2) andby general habitus. The metallic colour gives itsome similarity to P. viridanus but the latter hasevenly spaced pronotal punctures and a differentabdominal pubescence pattern.

Platydracus exulans is an easternspecies, occurring as far north as Ontario andextending southwest to Louisiana. Its range inECAS is given by Map 14. This species is newlyrecorded from Michigan, Ohio, andPennsylvania. It was reported from Québec byDownie and Arnett (1996) without data, but wehave seen no specimens from that province andthis record is probably based on a misidentifiedspecimen.

UNITED STATES: MI: Ingham Co., EastLansing, 42.7333 -84.4833, Date Unknown,C.F.B, 1 (FMNH). OH: Hamilton Co.,Cincinnati, 39.1 -84.52, V-1934, 1 (UMMZ).Hocking Co., Ash Cave Area, 39.4003 -82.5384,1-V-1986, bark, R.L. Aalbu, 1 (FMNH).PA: Delaware Co., Manoa, 40 -75.3, IV-1946,E.J. Ford, 1 (BPBM). Westmoreland Co.,Jeannette, 40.3333 -79.6166, 14-IV-1931, 1(FMNH); 16-IV-1931, 2 (AMNH), 1 (FMNH).

Eastern Canada: ONAdjacent U.S.: IN, MI, NY, OH, PA

Platydracus exulans has been collectedfrom late March to late June, with the majority ofrecords in April and May.

Most available specimens withcollection data are from dung or dung-baitedpitfall traps in mixed oak-pine forest. However,Newton (1973) reported it from carrion,including opossum, chicken, and rabbit. Watson(2004) found this species in Louisiana oncarcasses of alligator, bear, deer, and pig duringthe winter and spring. Platydracus exulans wasfound most frequently on pig carcasses in theadvanced to dry stage of decay (Watson2004). Newton (1973) suggested that therelatively early season occurrence and scantappearance at baits may indicate that P. exulansdwells primarily within vertebrate nests or asimilar microhabitat.

This species has been collected twice inOntario, both times in the same location nearOttawa but forty-four years apart. Furthercollecting is needed to determine whether theCanadian specimens of P. exulans representsporadic migrants or part of a stable Canadianpopulation.

11.11 Platydracus femoratus (Fabricius 1801)

Platydracus femoratus is easilyrecognized by the large eyes, which are abouttwice as long as the temple (Fig. 7.1.1).Although the specimen figured in Fig. 11.11.1has its abdominal apex paler, most NorthAmerican individuals of this species have auniformly coloured abdomen.

Platydracus femoratus is distributedbroadly from Pennsylvania to Brazil, but occursonly sporadically in ECAS (Map 15); it isunlikely that it will be encountered in Canada.Individuals collected in ECAS are probablymigrants from the southern United States where

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Map 14. Distribution of Platydracus exulans (Erichson) in eastern Canada and adjacent United States.

they regularly occur. This species is newlyrecorded from Pennsylvania:

UNTED STATES: PA: Allegheny Co.,Pittsburgh, 40.43 -80.00, 15-V-1961, Legerman,1 (CMNH); Tarentum (4km N), 40.6 -79.77, 7-VII-1991, W. Zanol, 1 (CMNH). Franklin Co.,Mount Alto, 39.85 -77.55, Schaeffer, 1 (CUIC).Philadelphia Co., Chestnut Hill, 40.07 -75.2, 24-VIII, 1 (MCZ); Mount Airy, 40.07 -75.18, VII-1935, 1 (FMNH).

Adjacent U.S.: IN, OH, PA

Specimens in ECAS have been taken inMay to October with no apparent pattern inseasonality.

This species is most frequent at dungbut also often occurs on carrion. Severalspecimens have been collected in rotting fruitand at lights.

11.12 Platydracus fossator (Gravenhorst 1802)

Individuals with the typical apicalorange spot on the elytron are distinguished fromother Staphylinina (Gravenhorst) by thatcharacter alone (Fig. 7.8.1). Specimens lackingthe spot (with a completely black body) are

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Map 15. Distribution of Platydracus femoratus (Fab.) in eastern Canada and adjacent United States.

identified as Platydracus using the characters inthe key, and to fossator using the followingcombination of characters: elytra completelyblack (Fig. 7.8.2), abdomen lacking densepatches of golden or black velvet setae (Fig.7.8.2), pronotum lacking complete impunctatemedian line (Fig. 7.6.2), and eyes distinctlylonger than the temple.

This species is widespread, occurringfrom central North America east to the coast andsouth to Arizona, Texas and Florida. Specimensfrom west of the Great Plains were all collectedat high elevations. Platydracus fossator is quitecommon south of the Canadian border andoccurs regularly at some sites in southern Canada

(Map 16). This species is newly recorded fromNova Scotia, Ohio, and Pennsylvania:

CANADA: NS: Annapolis Co., Middleton, 20-VII-1928, W.J. Brown, 1 (CNC). Antigonish Co.,Brierly Brook, 20-VIII-1954, Douglas C.Ferguson, 1 (NMSC). Colchester Co.,Glenholme (Roger's Field), 45.42 -63.51, wildblueberry field, pitfall trap, 25-VIII-2009, C.Cutler, 2 (NSAC); Glenholme (Webb Field),45.41 -63.51, wild blueberry field, pitfall trap,25-VIII-2009, C. Cutler, 3 (NSAC); Kemptown,4-VIII-1999, J. Ogden, 1 (NSNR); Portapique, incow dung, 31-VII-1929, Frost, 1 (MCZ). DigbyCo., Digby, 14-XI-1960, C.J.S. Fox, 1 (ACNS);

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Map 16. Distribution of Platydracus fossator (Gravenhorst) in eastern Canada and adjacent United States.

South Range/Porter’s Lake, ‘Crawling in mowedlawn during midday’, 3-VIII-2003, G.D. Selig,(GSC); Weymouth, 14-VIII-1990, 1(MCZ).Hants Co., Monte Vista Farm, 14-VIII-1966,Kenneth A. Neil, 1 (NSMC).UNITED STATES: OH: Adams Co., 20-VIII-1967, 1 (Hamilton). Franklin Co., Columbus, 1(OSU). Stark Co., 1-VII-1937, 1 (INHS).PA: 69 specimens.

Eastern Canada: ON, QC, NB, NSAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Platydracus fossator has been collectedin ECAS from March to October, with a markedincrease in abundance during July andAugust.While this species inhabits a variety offorest types (except boreal), a large proportion ofspecimens were collected from open forests withconifers and/or oak trees, including oaksavannah. Platydracus fossator occurs at carrion,dung, and at decaying soft fungi includingPleurotus ostreatus Fries (Cline and Leschen2005), and is often encountered flying orwalking as it disperses between thesemicrohabitats (Fig. 11.12.1). Both adults andlarvae have been reared on a diet of muscoidflies (Schmidt 1999).

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Map 17. Distribution of Platydracus immaculatus (Mannerheim) in eastern Canada and adjacent UnitedStates.

11.13 Platydracus immaculatus(Mannerheim 1830)

Platydracus immaculatus is adistinctive species that can be differentiated fromother Staphylinina by a combination of a reddishhead, red elytra without distinct and elongatedark spots, abdominal tergites with dense, pairedpatches of golden setae, and a non-metallic body(Fig. 7.11.1). Species of the P. cinnamopteruscomplex are similar in colouration but lack thedistinct, paired patches of golden setae on theabdominal tergites.

This species is eastern in distribution:Ontario to North Carolina and west to Utah.Historically, it was quite commonly collected inECAS but Platydracus immaculatus is nowinfrequently collected over much of its range.Although this is only subtly apparent in Map 17,the change is quite apparent after the early1980’s.

Eastern Canada: ON, QCAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

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Map 18. Distribution of Platydracus maculosus (Gravenhorst) in eastern Canada and adjacent UnitedStates.

In ECAS, P. immaculatus has beencollected from February to November, with arelatively early peak in abundance during April-May.

Platydracus immaculatus is an openhabitat species, occurring especially in stonyareas with sandy soil. It can be collected fromunder stones, leaves, and other debris in oldfields, savannah, edges of open wetlands, and thegrassy areas along the shores of lakes and rivers.This species' range in Canada has always beenrestricted to the southern half of Ontario andQuébec; however, P. immaculatus now seems tobe rare in this area, with only 5 records from

Canada after 1980 (Fig. 11.13.1). Furthercollecting in sandy, grassy habitat with stones isnecessary to verify this species' apparent declinein Canada. Unlike many other species ofPlatydracus, P. immaculatus does not visitbaited traps and is primarily captured by liftingdebris or in pitfall traps. Platydracusimmaculatus was generally known by theyounger name P. vulpinus (Nordmann, 1837)until Smetana and Davies (2000) noted that thename immaculatus was older, and Herman(2001) adopted immaculatus as the valid name.

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11.14 Platydracus maculosus(Gravenhorst 1802)

Platydracus maculosus can be easilydistinguished from all other species ofStaphylinina in ECAS (LeConte) by its large size(22-35 mm) and the distinct, dark and elongatespots on its elytra (Fig. 11.14.1). It can also bedifferentiated from the similar P. comes by theincomplete impunctate median line of thepronotum (Fig. 7.7.1) and from P. femoratus byits smaller eyes (Fig. 11.14.1).

This species is widespread in easternNorth America, ranging south to the Floridapanhandle. Its range in ECAS is given in Map18.

Eastern Canada: ON, QCAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Platydracus maculosus has beencollected in ECAS from January to November,with a large peak in abundance in May and asecond, smaller increase in September.

Platydracus maculosus is the largestand one of the most commonly encounteredPlatydracus species in ECAS. It primarilyinhabits deciduous forests and open areas, whereit occurs on carrion (Fig. 11.14.2), on dung, inleaf litter and sometimes in rotting fungi. A fewindividuals have been collected under bark and atlights. They are often observed in flight and theirdark blue hindwings give them a wasp-likeappearance. Adults overwinter under logs androcks and can be found there in the late fall andearly spring (Voris 1939). Smetana and Davies(2000) noted that P. maculosus had an oldersynonym, P. viduatus (Fabricius, 1801), andused that as the valid name for this species, butP. maculosus was conserved as the valid nameby Opinion 2039 of the InternationalCommission on Zoological Nomenclature (ICZN2003).

11.15 Platydracus mysticus (Erichson 1840)

Platydracus mysticus can be recognizedby the combination of small eyes (much smallerthan the temple) (Fig. 7.6.3), elytra withoutelongate dark spots, pronotum without animpunctate median line (Fig. 7.6.2), abdominaltergites lacking distinct patches of golden setae,

and first antennomere distinctly bicoloured (Fig.7.6.4). The darkest individuals resemble P.tomentosus, but that species has eyes that aresubequal in length to the temple.

This species ranges over most ofnortheastern North America: southern Ontarioand Quebec, southwest to central Texas. Itsrange in ECAS is given by Map 19. Specimensreported by Klimaszewski et al. (2005) as P.mysticus were later found to be misidentified P.viridanus (A. Smetana pers. comm.). Thus P.mysticus does not occur in the MaritimeProvinces of Canada as far as known.Platydracus mysticus is newly recorded fromMichigan and Maine.

UNITED STATES: MI: 57 specimens. ME:Penobscot Co., Orono, 44.8833 -68.6667, 24-V-1948, 1 (UAIC).

Eastern Canada: ON, QCAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Platydracus mysticus has been collectedin ECAS in March to December, with arelatively early peak in abundance in April-May.

This species is typically found underrocks and logs in forested or field habitat,although the vast majority of specimens lacklabel data. It may also be found on beaches andshoreline in 'washup' and under drift. Newton(1973) suggested that Platydracus mysticus maygenerally be cursorial and surface active; the factthat most collections are made singly and manylabels state 'on ground' or 'on path' (or similar)supports this. This species is a competent flier asit has been collected by a flight intercept trap(Watrous 2008). Platydracus mysticus iscommon in collections until about 1960 and laterrecords are increasingly less numerous,suggesting that this species is in decline overmuch of its range in ECAS, especially in denselypopulated areas; focused collecting is needed toverify this. Possible factors contributing to thisapparent decline include urbanization andcompetition with introduced species of nearlyidentical adult habits (i.e. Tasgius spp.).

11.6 Platydracus praelongus(Mannerheim 1830)

Platydracus praelongus is distinctiveamong Platydracus species for its pronotum withan uneven punctation and wide (at least as wideas five punctures) impunctate line (Fig. 7.4.1);

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both the head and pronotum are dull-metallic(Fig. 11.16.1).

This species occurs along the easterncoast of the United States and Mexico, north toMaine, but it may be eventually found in theextreme southern portions of Atlantic Canada. Itsrange in ECAS is given by Map 20. It is recordedfrom the state of New York for the first timebased on study of 77 specimens collected until1975.

Adjacent U.S.: NY, NH, ME

Platydracus praelongus has beencollected in ECAS from February to December,with a large increase in abundance in May andsmaller peak in August.

Platydracus praelongus is a seashorespecialist of and is the only North AmericanPlatydracus associated with marine coastalenvironments. In ECAS it has been collected onbeaches under debris and logs (Fig. 11.16.2),swept from dune grass, and found along theedges of salt marshes under stones and in litter.

11.17 Platydracus praetermissusNewton, spec. nov.

Type material: Holotype, male, in FMNH, withlabels: "ARK: Wash. Co., 3 mi S Devils Den St.Park, 28-31.V.79 litter, S&J Peck, oak-hickory"[white printed label]; "A. Newton collection"[green printed label]; "♂ GENITALIA 51, to J.Klimaszewski for drawing, 1993" [red printedlabel]; and red handwritten holotype label by A.Newton dated 2010 [note: genital segment andaedeagus removed and stored in glycerin in amicrovial pinned with the specimen].Paratypes, 198, all males, all but a few withaedeagus examined:

UNITED STATES: AL: Dale Co., Ft. RuckerMil. Res., 31.33 -85.71, barrier pitfall trap, 4-I to16-IV-1999, R. Turnbow, 1 (Turnbow); Ft.Rucker Mil. Res., 31.33 -85.71, berlese leaflitter, 3-VII-2005, R. Turnbow, 1 (Turnbow);Lee Co., Auburn, 32.62 -85.48, sifting forestlitter, 20-III-1972, E. J. Kiteley, 1 (CNC);Auburn, 32.62 -85.48, sifting forest litter, 25-III-1972, E. J. Kiteley, 1 (CNC). AR: Arkansas Co.,White River N.W.R., 34.33 -91.12, woodlandpitfall trap, 9-VII-1969, Underwood and Brown,1 (UAAM); Ashley Co., pine forest, pitfall trap,Line 15 (A-15), 5-VIII-1974, Heiss, Howard andChenowith, 1 (UAAM); 13.5 mi S of Crossett,33.00 -91.83, pitfall trap, 74-2459 Line 13, 24-

VI-1974, Heiss, Howard and Chenowith, 1(UAAM); Bradley Co., 16-V-1946, 1 (UAAM);19-VI-1963, 1 (UAAM); 5-VI-1965, 1 (UAAM);24-VII-1965, 1 (UAAM); 30-VII-1965, 2(UAAM); Franklin Co., Shores Lake, OzarkNational Forest, 35.63 -93.95, 20-V-1983, J.M.Campbell, 1 (CNC); Grant Co., junction ofhighway 167 and 35 (2.7 mi SE on 35), 34.23 -92.37, pine, berlese, 77-06, 19-II-1977,Chenowith, 1 (UAAM); Hempstead Co., 12-III-1959, 1 (FMNH); 12-III-1959, leaf trash,GO.848, 1 (FMNH); 12-III-1959, leaf trash,GO.942, 1 (FMNH); Johnson Co., 25.3 mi W ofjunction of highways 21 and 1003, 35.68 -93.67,pitfall trap, 73-534 Line 14, 31-V-1973, R.T.Allen, 1 (UAAM); Newton Co., hardwood forest,pitfall trapline, 75-0149, 16-VII-1975, 1(UAAM); pitfall, 75-0159, 22-VII-1975, 1(UAAM); 2.7 mi W of Buffalo River at highway74, 36.00 -93.38, pitfall trap, Line 2, 12-VII-1973, D. Carlyle, 1 (UAAM); Pope Co., Popeand Van Buren Co. Lines, 12.4 mi W onhighway 16, 35.70 -92.92, pitfall trap, Line 10,21-VI-1973, 1 (UAAM); 17.3 mi W ofintersection of Pope and Van Buren Co. Lines,35.70 -92.92, pitfall trap, Line 11, 21-VII-1973,2 (UAAM); 17.3 mi W of intersection of Popeand Van Buren Co. Lines, 35.70 -92.92, pitfalltrap, 73-651 Line 11, 28-VI-1973, R.T. Allen, 1(UAAM); Pulaski Co., Little Rock (Marsh Rd.),34.72 -92.48, berlese leaf litter, 14-II-2000, B.Baldwin, 1 (Baldwin); Little Rock (Marsh Rd.),34.72 -92.48, hanging sugar traps, 16-IV-2003,Baldwin, 1 (Baldwin); Union Co., pitfall trap,75-0222 Line 3, 28-VII-1975, 1 (UAAM); 0.5 miSE of Lapile, 33.07 -92.27, pitfall trap, 74-0136Line 3, 10-VI-1974, Heiss, Howard andChenowith, 1 (UAAM); 4 mi S of Mount Holly,33.23 -92.97, pitfall trap, 74-0106 Line 15, 4-VI-1974, Heiss, Howard and Chenowith, 1(UAAM); Washington Co., leaf trash, GO.96, 5-XII-1958, 1 (UAAM); Devils Den State Park,35.78 -94.23, in crevice, 8-VI-1946, M.W.Sanderson, 1 (INHS); Farmington, 36.05 -94.25,black light trap, 73-46, 22-VI-1973, W.D. Wylie,1 (UAAM); Fayetteville, N slope of MarkhamHill, 36.07 -94.15, under boulder, 76-97, 4-XII-1976, R. Chenowith, 1 (UAAM).DC: Washington, Rock Creek Park, 38.97 -77.05, oak hillsides, gulley floor/log, 67-2a, 24-I-1967, W. Suter, 1 (FMNH). DE: Sussex Co.,Angola, 38.68 -75.18, 20-VI-1970, 1 (AMNH).FL: Alachua Co., 4 mi SW Gainesville,Hogtown Creek, 29.65 -82.38, pitfall, 27-V-1983, R.M. Reeves, 3 (DENH); Gadsden Co., 1mi E of Havana, 30.62 -84.41, pine-hardwood

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litter, 6-I-1977, C.W. O’Brien and G.B.Marshall, 2 (FMNH); Hamilton Co., 8 mi S ofJasper, Hwy 129, 30.40 -82.93, mixed hardwoodlitter, 24-III-1977, C.W. O’Brien et al., 1(FMNH); Leon Co., Tallahassee, 30.43 -84.28,mixed hardwood litter, 8-IV-1976, C.W. O’Brienand G.B. Marshall, 1 (FMNH); Tallahassee,30.43 -84.28, mixed hardwood litter, 76-115,28-VI-1976, Marshall and Justice, 1 (UAAM);Tallahassee, 30.43 -84.28, hardwood litter, 4-III-1976, G.B. Marshall, 1 (FMNH); Tallahassee,30.43 -84.28, hardwood litter, blacklight trap, 6-XII-1976, G.B. Marshall, 1 (FMNH);Tallahassee, 30.43 84.28, mixed hardwood litter,20-IV-1976, C.W. O’Brien and Justice, 1(FMNH); Tallahassee, 30.43 -84.28, mixedhardwood litter, 2-IV-1976, C.W. O’Brien andG.B. Marshall, 1 (FMNH); Tallahassee, 30.43 -84.28, hardwoods, berlesate hardwoods, 77-204,14-X-1977, L. Justice and E. Healy, 1 (UAAM);Tallahassee, 30.43 -84.28, mixed hardwoodlitter, 11-III-1977, G.B. Marshall, 1 (FMNH);Tallahassee, 30.43 -84.28, mixed hardwoodlitter, 6-X-1977, C.W. O’Brien, 1 (FMNH);Okaloosa Co., Destin, 30.40 -86.50, 13-II-1976,E.J. Kiteley, 1 (CNC); Destin, 30.40 -86.50, 10-III-1980, E.J. Kiteley, 1 (CNC); Destin, 30.40 -86.50, 23-III-1982, E.J. Kiteley, 1 (CNC);Orange Co., Winter Park, 28.60 -81.33, 4-III-1939, F.E. Lutz, 1 (AMNH); Polk Co., in steinertrap, 14-III-1961, R.E. Vild, 1 (FSCA);Suwannee Co., 15 mi SW of Live Oak, Hwy 51,pine and hardwood litter, 24-III-1977, C.W.O’Brien et al., 1 (FMNH); Taylor Co., 5 mi E ofGainesville, Hatchet Creek, 29.65 -82.25, UVlight, 27-V-1983, R.M. Reeves, 2 (DENH).GA: Fulton Co., Atlanta, 33.75 -84.38, 1-VI-1929, P.W. Fattig, 1 (INHS); Liberty Co.,Riceboro, 31.73 -81.43, 27-IV-1937, 1 (CAS).IL: Champaign Co., Mahomet, Hart MemorialWoods, 40.20 -88.40, trap 4, 4-VI-1960, 1(INHS); Mahomet, Hart Memorial Woods, 40.20-88.40, trap 13, 1-VII-1966, 1 (INHS); Mahomet,Hart Memorial Woods, 40.20 -88.40, trap 3, 7-VI-1967, R.T. Allen and J.D. Unzicker, 1(INHS); Clark Co., Rocky Branch, 39.47 -87.77,pitfall trap, 1 to 13-V-1996, M.A. Goodrich, 1(EIU); Rocky Branch, 39.47 -87.77, pitfall trap,12 to 26-VII-1998, M.A. Goodrich, 1 (EIU);Rocky Branch, 39.47 -87.77, pitfall trap, 14 to28-VI-1998, M.A. Goodrich, 3 (EIU); RockyBranch, 39.47 -87.77, pitfall trap, 24 to 31-V-1998, M.A. Goodrich, 1 (EIU); Coles Co.,Charleston vicinity, 39.50 -88.18, 6-VI-2001,N.L. Owens, 1 (EIU); Cook Co., Des Plaines,Carlé Woods, 42.03 -87.88, rotten oak log, 12-X-

1957, O. Park, 1 (FMNH); Edgar Co., 4 mi SSEof Kansas, 39.50 -87.92, ‘from part of forestNOT burned in fall 1999’, pitfall trap, 25-VIII to1-IX-2000, M.A. Goodrich, 1 (EIU); 4 mi SSEof Kansas, 39.50 -87.92, ‘from part of forestburned in 1999’, pitfall trap, 13 to 20-X-2000,M.A. Goodrich, 1 (EIU); 4 mi SSE of Kansas,39.50 -87.92, ‘from part of forest NOT burned in1999’, pitfall, 31-V to 7-VI-2000, M.A.Goodrich, 1 (EIU); Jackson Co., Carbondale,37.73 -89.22, 5-V-1959, M. Stibitz, 1 (SIUC);Carbondale, 37.73 -89.22, 28-V-1963, C.S.Adams, 1 (SIUC); Piatt Co., White Health, 40.08-88.52, ground no. 24, 1-I-1983, J.C. Dirks, 1(INHS); Pope Co., Dixon Springs Park, GhostDance Canyon Trail, 37.38 -88.67, 125m, mixedhardwood forest in ravine near stream, underrock, 1043, 22-VI-2002, A. Newton, 1 (FMNH);St. Clair Co., 4-VII-1901, G.W. Block, 1(UMRM); 5-VI-1902, G.W. Block, 1 (UMRM);Vermillion Co., Catlin, Camp Drake, 40.10 -87.75, ground, 12-VI-1940, J.E. Porter, 1(INHS). IN: Jasper Co., Jasper-Pulaski StateGame Area, 41.15 -86.93, sweeping vegetation,grassy area, 19-V-1996, R.M. Brattain, 1(Brattain); Tippecanoe Co., 17-V-1959, N.M.Downie, 1 (FMNH); Vigo Co., 8-VI-1893, W.S.Blatchley, 1 (PURC). KS: Douglas Co.,Lecompton, 39.05 -95.40, 10-X-1933, 1(FMNH); Jefferson Co., University of KansasNatural History Reserve, 39.05 -95.18, forest,litter, 6-V-1986, J. Pakaluk, 1 (KSEM);Leavenworth Co., 18-V-1956, G.W. Byers, 1(KSEM). KY: Meade Co., 0.5 mi S of Route1158 on Route 333, 7-VI-1986, R. A. Mattingly,1 (ULKY). LA: West Feliciana Co., FelicianaPreserve near Freeland, 30.78 -91.25, pitfalltraps, 17-V-1995, D. Pashley, 4 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,pitfall traps, 5-X-1995, D. Pashley, 4 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,pitfall traps, 5-IX-1995, D. Pashley, 3 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,16-V-1995, D. Pashley, 1 (LSAM); FelicianaPreserve near Freeland, 30.78 -91.25, pitfalltraps, 23-V-1995, D. Pashley, 1 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,pitfall traps, 24-V-1995, D. Pashley, 3 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,pitfall traps, 6-I-1995, D. Pashley, 1 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,pitfall traps, 5-II-1995, D. Pashley, 1 (LSAM);Feliciana Preserve near Freeland, 30.78 -91.25,6-II-1995, D. Pashley, 7 (LSAM); Kakowi 2(T35.R1W, S72), pitfall trap, 9-VIII-1995, D.Pashley, 1 (LSAM); Tunica Hills WMA1, 30.92

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-91.49, pitfall trap, 23-V-1995, D. Pashley, 1(LSAM); Tunica Hills WMA1, 30.92 -91.49,pitfall trap, 6-II-1995, D. Pashley, 3 (LSAM);Tunica Hills WMA1, 30.92 -91.49, pitfall trap,18-V-1995, D. Pashley, 6 (LSAM); Tunica HillsWMA1, 30.92 -91.49, pitfall trap, 17-V-1995, D.Pashley, 4 (LSAM); Tunica Hills WMA1, 30.92-91.49, pitfall trap, 5-XI-1995, D. Pashley, 1(LSAM); Tunica Hills WMA1, 30.92 -91.49,pitfall trap, 25-VIII-1995, D. Pashley, 1(LSAM); Tunica Hills WMA1, 30.92 -91.49,pitfall trap, 15-VII-1995, D. Pashley, 2 (LSAM);Tunica Hills WMA 2, 30.92 -91.49, 5-XI-1995,D. Pashley, 1 (LSAM). MI: Barry Co., BarryGame Area (T3N, R10Q, Sec. 26), 42.62 -85.47,barrier pitfall, 8-V-1980, 1 (DENH); BerrienCo., Lakeside, Warren Woods, 41.83 -86.63, 6-X-1973, W. Suter, 1 (FMNH); Warren WoodsState Park, 41.83 -86.63, virgin beech-mapleforest, berlese, leaf and log litter, 892, 30-V-1992, A. Newton and M. Thayer, 1 (FMNH).MO: Boone Co., Ashland, 38.77 -92.25, pitfalltraps, 16 to 26-VIII-1984, N. Stebbins, 1(CMNH); Columbia, 38.95 -92.33, 9-1999,students of general entomology, (LSAM); IronCo., Annapolis, 37.37 -90.70, forest debris, 1-IV-1956, G. Ulrich, 1 (INHS); Reynolds Co., 10 miSW of Graniteville, Johnson Shut-ins State Park,37.53 -90.85, in dead leaves, 22-IV-1956, J.Kingsolver, 1 (INHS); St. Louis Co., Ranken,38.53 -90.52, 1-V-1983, E. P. Meiners, 1 (CAS);Ranken, 38.53 -90.52, 6-VII-1941, E. P.Meiners, 1 (UMRM); St. Louis, 38.63 -90.20,29-IV-1906, G.W. Bock, 1 (UMRM); WayneCo., Sam A Baker State Park, 37.25 -90.50, 11 to12-V-1951, 1 (EIU). MS: Benton Co., Ashland,34.83 -89.18, 8-IV-1977, S.C. Elliot, 1 (UMIC);Covington Co., ground trash, 18-XII-1973, J.R.McCoy, 1 (MEM); George Co., Lucedale, 30.93-88.58, 26-V-1930, H. Deitrich, 1 (CUIC);Leakesville, 31.15 -88.58, 30-IX-1931, H.Deitrich, 1 (CUIC); Grenada Co., Grenada,33.77 -89.82, trash samples peripheral tocultivated cotton, 17-IV-1981, R. Seward, 1(MEM); Lafayette Co., 6 mi E of Oxford, 34.37 -89.42, 25-II-1977, M.T. McCraine, 1 (UMIC);Marion Co., ground trash, 30-XI-1973, J.R.McCoy, 2 (MEM); Oktibbeha Co., 5 mi W ofStarkville, Adaton, 33.48 -88.92, oak buttressbasal tree hole, 4-I-1982, W. Suter, 1 (FMNH);Perry Co., Richton, 31.35 -88.93, 23-IV-1930,H. Deitrich, 1 (CUIC); Pontotoc Co., 1 mi SE ofEcru, 34.33 -89.00, deciduous woods, pitfalltrap, 18-VI-1980, W.H.Cross, 1 (MEM); WayneCo., 2.5 mi SSE of Buckatunna, 31.5 -88.52, pittrap, 21-V-1983, P.K. Lago, 1 (UMIC).

NE: Otoe Co., Nebraska City, 29-IV-1916, R.W.Dawson, 1 (UNSM). NJ: Atlantic Co., AtlanticCity, 39.37 -74.42, 15-VI, 1 (MCZ); BurlingtonCo., Centerton, E along Rancocas Creek, 40.00 -74.87, mixed hardwood forest, dung trap(human), 2 to 16-VI-1969, A. Newton, 1(FMNH); Rancocas State Park, 40.00 -74.85,mixed hardwood forest, litter, berlese, 28-VIII-1976, A. Newton and M. Thayer, 1 (FMNH);Cape May Co., Anglesea, 39.02 -74.80, 30-V-1930, 1 (MCZ). NY: Suffolk Co., Long Island,Bellport, 40.75 -72.93, 8-VI-1913, A. Nicolay, 1(AMNH); Long Island, Sag Harbor, 41.00 -72.30, 2-V-1929, R.L., 1 (CUIC); Long Island,Wildwood State Park, 40.97 -72.80, 7-VI-1951,H. Dietrich, 1 (CUIC); Long Island, Wyandanch,40.75 -73.37, 10-X-1915, F. M. Schott, 1(AMNH). OH: Preble Co., Hueston WoodsState Park, 39.58 -82.77, litter, beech-mapleforest, 19-V-1990, P. Kovarik, 1 (FMNH). OK:Latimer Co., V-1987, K. H. Stephan, 1(Stephan); V-1995, K. H. Stephan, 1 (Stephan);S of Red Oak, 34.95 -95.08, IV-1997, K.H.Stephan, 1 (TAMU). PA: Bucks Co., NE ofJamison, Neshaminy Creek, Horseshoe Bend,40.27 -75.08, 1-IV-1955, W. Ivie, 1 (AMNH);Delaware Co., Chester, 39.85 -75.35, IV-1904, 1(CUIC); Philadelphia Co., Chestnut Hill, 40.07 -75.20, 6-VII, 1 (MCZ); Chestnut Hill, 40.07 -75.20, 24-VI, 1 (MCZ). SC: Florence Co.,Florence, 34.2 -79.77, 10-I-1938, 1 (CAS);Florence, 34.2 -79.77, 46m, 25-II-1938, 1(MCZ); Florence, 34.2 -79.77, 31m, 14-II-1938,C.W.P., 1 (MCZ); Florence, 34.2 -79.77, woodstrash, 16-V-1963, V.M. Kirk, 1 (FMNH); HorryCo., 3.5 mi NW of Longs, Nelson Property,33.97 -78.77, wet mixed conifer-hardwoodforest, ex. leaf litter, 1-4pm, 19-III-2000, A.S.Ramsdale, 1 (IRCW). TX: Angelina Co., 8 mi Wof Lufkin, 31.33 -94.83, 21 to 25-IV-1976, A.Smetana, 1 (CNC); 8 mi W of Lufkin, 31.33 -94.83, 21-IV-1976, A. Smetana, 1 (CNC); JasperCo., 3 mi N of Buna, US 96, 30.43 -93.97, 6-XII-1968, G.E. Ball, 1 (UASM); San Jacinto Co.,Big Creek Scenic Area, 30.52 -95.10, rotting log,18-II-1955, D.K. Rudd, 1 (TAMU); Trinity Co.,2.5 mi NW of Apple Springs, 31.22 -94.97, 23-IV-1976, A. Smetana, 2 (CNC); Tyler Co., 2.8mi W of Spurger, Big Thicket Nat. Pres., BeechWoods Trail, 30.70 -94.18, beech-magnoliaforest, berlese leaf litter, 8-III-1989, R.Anderson, 2 (FMNH). VA: Henrico Co., 1 mi Wof Elko, Elko Nat. Area, 37.47 -77.23, 15-VI-1990, C.A. Pague, 1 (VMNH); Mecklenburg Co.,Elm Hill S.G.M.A., DF site near Kerr Dam,36.60 -78.30, 5 to 19-VI-1991, VMNH survey, 2

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(VMNH); Prince William Co., Occoquan, 38.68-77.27, 2-V-1943, E.S. Ross, 1 (CNC); VirginiaBeach Co., Little Creek Amphibious Base, 36.92-76.17, 12-V-1989, K.A. Buhlmann, VDNHsurvey, 1 (VNMH); Virginia Beach, SeashoreState Park, 36.90 -76.02, scrub pitfall site, 21-VI-1989, K.A. Buhlmann VDNH survey, 5(VNMH).

Description: With the characters of the P.cinnamopterus Complex (see above), plus:average pronotal length 2.51 mm (n = 13); head,pronotum and elytra usually dark reddish,pronotum not bicolored; antenna reaching orexceeding middle of pronotum when extendedposteriorly, antennomere 9 subquadrate;impunctate median line of pronotum at leastthree puncture diameters wide at narrowest point,usually about four puncture diameters wide;emargination of sixth visible male sternite inventral view shallow, about 1/6 as deep as wide;apex of median lobe of aedeagus in parameralview broadly and slightly convexly truncate;sides of median lobe in lateral view distinctlytoothed near apex.

Etymology: From the Latin adjectivepraetermissus, meaning "neglected" or"overlooked", in reference to the late discoveryof this widespread species which had beengenerally misidentified as P. cinnamopterus.

Platydracus praetermissus may bedistinguished from all other species of the genusin ECAS by the combination of generally darkreddish color (Fig. 11.17.1); black scutellum;variegated abdomen with only sparse gold setaeand paired areas of black velvet setae on the firstfive visible segments (Fig. 7.9.2); antennae withsubquadrate antennomere 9 (Fig. 7.14.2); and thecomplete impunctate medial line of pronotumabout 3-4 puncture diameters wide at narrowest(Fig. 7.14.1). It is definitively distinguished fromthe other members of the P. cinnamopterusComplex, P. cinnamopterus and P. zonatus, bythe structure of the aedeagus (Fig. 7.14.3,11.17.2).

Platydracus praetermissus is widelydistributed in the forested areas of eastern UnitedStates: from Long Island, New York, west toNebraska, south to Texas and east to peninsularFlorida (Map 21). Its distribution in ECAS isshown in greater detail in Map 22. Thedistribution of this species is generally moresouthern than the other species of thecinnamopterus Complex and is not yet known to

include Canada. However, it may occur in thefew, relatively mature hardwood forest patchesleft in southern Ontario. Adjacent U.S.: MI, IN,OH, PA, NY

In ECAS, P. praetermissus specimenshave been collected from April through October,with most (10 of 15 records) found in May orJune.

The four specimens of P. praetermissusfrom ECAS with collection data were obtainedfrom forest leaf litter (2), in a barrier pitfall trap(1), and sweeping vegetation in a grassy area (1).In its entire range, this species has been collectedmost often in ground pitfall traps or in forest leaflitter, and rarely associated with decaying logs orin UV light traps at night (twice each). Unlike P.cinnamopterus, this species is not known tooccur under bark. Note that some historicalrecords of P. cinnamopterus in the literature mayrefer to this species.

11.18 Platydracus tomentosus(Gravenhorst 1802)

This species is distinguished from otherPlatydracus by the combination of a completelyblack body and setae, eyes subequal to thetemple (Fig. 7.13.2) and paired patches of blackvelvet setae on the abdominal tergites (Fig.7.13.1). The immaculate form of P. fossatorcould be confused with P. tomentosus, but theformer lacks patches of black velvet setae on theabdominal tergites and has eyes that aredistinctly longer than the temple. The darkestindividuals of P. mysticus are also similar, buthave eyes that are distinctly shorter than thetemple.

This species occurs in eastern NorthAmerica, from Ontario to Florida, and west toNebraska; it also occurs in Cuba. Its range inECAS is given by Map 23. Platydracustomentosus is newly recorded from New Yorkand Ohio.

UNITED STATES: NY: 37 specimens. OH:Logan Co., 40.39 -83.85, 8-VIII-1932, 1(UMSP). Mercer Co., Celina Spring, 40.55 -84.57, IV-1951, E. Klee, 2 (CAS).

Eastern Canada: ONAdjacent U.S.: MI, IN, OH, NY

Platydracus tomentosus has beencollected in ECAS from January to November,with a peak in abundance in May.

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Map 19. Distribution of Platydracus mysticus (Erichson) in eastern Canada and adjacent United States.

Platydracus tomentosus (Gravenhorst)primarily inhabits the edges of lakes and ponds,and slow-moving waterways, where it can befound under stones, wood, and flood debris. Itsometimes occurs in forests or in prairies butthese individuals may only be dispersing throughto the nearest water body. Schmidt (1994) foundmany adults within 3 meters of a large creek in apasture, where they were found singly or in pairsin small 'discus-shaped' excavations underobjects. This species is probably a generalistpredator of riparian arthropods, and Schmidt(1994) fed them lampyrid larvae, isopods, adultflies, and elaterid larvae collected from the samelocation. Larvae burrow readily in soil (Schmidt

1994), unlike the adults, and could be primarilyfossorial. There are very few recent records ofthis species from the northeastern portion of itsrange (Map 20) and further collecting in favoredhabitat is necessary to determine whether P.tomentosus is in decline.

11. 19 Platydracus violaceus(Gravenhorst 1802)

This species is easily recognized by thecombination of a purple-blue metallic dorsalbody and characteristic pattern of pale abdominalsetae (Fig. 7.12.1). It may be confused with

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Map 20. Distribution of Platydracus praelongus (Mannerheim) in eastern Canada and adjacent UnitedStates.

P. viridanus (Horn), which normally has abronze metallic reflection but can appear purplein specimens from traps with killing agents.Platydracus viridanus also has a more transversepronotum that is distinctly wider than the head,while in P. violaceus, the pronotum is subequalin width to the head. Platydracus violaceus alsohas a dark elytral epipleuron (Fig. 7.12.2) andcompletely dark legs, while in P. viridanus theventral margin of the epipleuron is paler than therest of the elytron and the legs are usually at leastpartially pale.

This species is distributed broadly ineastern North America. Its range in ECAS is

given by Map 24. It is newly recorded from thestate of Vermont based on a historical record; itvery likely still occurs there:

UNITED STATES: VT: Addison Co.,Salisbury, 43.9 -73.1, 5-VII-1895, 1 (MCZ)

Eastern Canada: ON, QC, NB, NS, PEIAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Platydracus violaceus has beencollected in ECAS from April to December, withan increase in abundance occurring in May.

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Map 21. Distribution of Platydracus praetermissus sp. nov.

Platydracus violaceus is a commonspecies found in mesic to swampy forests,primarily under the loose bark of dead hardwoodtrees including oaks, maples, basswood, horsechestnut, hackberry, beech, and hickory. Itoccurs less frequently under the bark of whitepine, in rotting wood, and under logs. The fewrecords in rotting fungi, on carrion, or on dungprobably do not reflect habitat preferences.Larvae were found to readily consume variousinsect larvae occurring under bark (Newton1973) and it is likely that adults have similardietary habits. Mature larvae are collected in theearly spring (Hoebeke 1978; A. Brunke personalobservations) indicating that this speciesoverwinters in this stage. Larvae form pupalcells out of frass and wood debris and aresometimes parasitized by the proctotrupidCodrus carolinensis (Ashmead) (Hoebeke 1978).Adults appear to emerge later in spring (April-May). Jennings and Tallamy (2006) found this

species in a relatively undisturbed secondaryforest but not in an isolated woodlot of similartree species composition. Similarly, Majka(2010) found this species in an old growth forestremnant in Prince Edward Island, a provincewhere it was not known previously. It washypothesized that forestry practices wereresponsible for this species’ absence in previouscollections made elsewhere in the province.Based on our collection data, this species doesnot require old growth but is usually found inforests with at least some mature trees. Smetanaand Davies (2000) noted that P. violaceus was apreoccupied name and replaced it with a youngersynonym, P. cupripennis (Melsheimer, 1844),but P. violaceus was conserved as the valid namefor this species by Opinion 2039 of theInternational Commission on ZoologicalNomenclature (ICZN 2003).

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Map 22. Distribution of Platydracus praetermissus sp. nov. in eastern Canada and adjacent United States.

11.20 Platydracus viridanus (Horn 1879)

The combination of the small eyes (no longerthan the temple), bronze metallic dorsalreflection, evenly punctate pronotum, elytrawithout spots (Fig. 7.10.2), an elytral epipleuronwith a pale ventral margin (Fig. 7.10.3), and thedistinctive pattern of abdominal setae (Fig.7.10.1) distinguish Platydracus viridanus fromall congeners. For characters differentiating thisspecies from P. exulans, see comments underthat species. Specimens of P. violaceussometimes appear bronze if discoloured bykilling agents but have a different pattern of

abdominal setae, a dark elytral epipleuron, andother differences noted under that species.

Platydracus viridanus occurs acrossnortheastern North America, from Minnesota toNova Scotia, extending south in the Appalachianmountains to northern Georgia.

UNITED STATES: OH: Franklin Co.,Columbus, 39.97 -83.00, 1940, Rings, 1(BMNH). Hocking Co., Rock House State Park,39.5 -82.62, 26-VII-1979, L.E. Watrous, 2(FMNH). Ross Co., Tar Hollow State Forest,39.3577 -82.7746, 5-VIII-1989, riparian woods,malaise trap, R.S. Miller family, 1 (MTEC).

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Map 23. Distribution of Platydracus tomentosus (Gravenhorst) in eastern Canada and adjacent UnitedStates.

PA: 90 specimens. VT: Addison Co., Salisbury,43.93 -73.1, 5-VII-1895, 3 (MCZ). BenningtonCo., 43.03 -73.1, 2 (FMNH). Essex Co.,Ferdinand, Route 105, 44.7333 -71.7667, 8-VII-1976, L.L. Pechuman, 1 (CUIC).

Eastern Canada: ON, QC, NB, NSAdjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

This species has been collected inECAS from May to September, with nearly allrecords occurring in July-August.

Platydracus viridanus is a mesic forest-dwelling species that is typically localized indistribution and occurs in decaying fungi, oncarrion, and dung. Individuals have also beencollected in decaying plant matter (with larvae),under logs, in a buzzard nest (with larvae), inforest floor pitfall traps, along a grassy rivermargin (Fig. 11.20.1), and in a dry field.Jennings and Tallamy (2006) found this speciesin a relatively undisturbed secondary forest butnot in an isolated woodlot of similar tree speciescomposition. This species was also commonlycollected in pitfall traps in an old-growth whitepine forest in Ontario.

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Map 24. Distribution of Platydracus violaceus (Gravenhorst) in eastern Canada and adjacent United States.

11.21 Platydracus zonatus (Gravenhorst 1802)

Staphylinus zonatus Gravenhorst, 1802: 162;synonym of S. cinnamopterus, Erichson1839; as valid species of Platydracus:Newton in Smetana and Davies 2000,Herman 2001, etc. (see Herman 2001:3473 for additional literature citations)Type material: Lectotype, male, inZMHB, here designated by Newton,with labels "7087" [white printed label];"America s. Topp" [green handwrittenlabel]; and red lectotype designationlabel by Newton dated 1978 [note: rightantennomeres 4+, most of right elytron

and both metathoracic legs missing,genital segment and aedeagus removedand glued on card with abdomen]. Asecond specimen, female, in ZMHBassociated with the lectotype, withgreen handwritten label "var. zonatusGr. A. spt.", in poor condition with partsof abdomen glued to card belowspecimen, was not designated aparalectotype although assumed to beconspecific.

Staphylinus badius Mannerheim, 1830: 23(attributed to Dejean, 1821: 21 [nomennudum]); synonym of S. cinnamopterus:Erichson 1839; synonym of Platydracus

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Map 25. Distribution of Platydracus viridanus (Horn) in eastern Canada and adjacent United States.

zonatus: Newton in Smetana and Davies 2000,Herman 2001, etc.Type material: Lectotype, female, inMZHF, here designated by Newton,with labels: "♀"; "Dejean."; "Amer.bor."; "Mus. Zool. Helsinki, Loan No.C11426"; and red lectotype designationlabel by Newton dated 1978 [note:associated with label "Staphylinusbadius Mannh." in Mannerheimcollection in MZHF, teste H.Silfverberg 1978].

Staphylinus quadraticeps Casey, 1924: 149[preoccupied name, not Staphylinus

quadraticeps Ménétries 1832]; synonymof S. caseyi: Scheerpeltz 1933; synonymof Platydracus zonatus: Newton inSmetana and Davies 2000, Herman2001, etc.Type material: Holotype, female, inUSNM, with labels "Fla."; "Caseybequest 1925"; "TYPE USNM 48271","quadraticeps Csy"; and red holotypelabel dated Newton 1978 [note:considered holotype because Caseyindicated only one female was seen].

Staphylinus caseyi Scheerpeltz 1933: 1392[replacement name for Staphylinusquadraticeps Casey 1924]; synonym

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Map 26. Distribution of Platydracus zonatus (Gravenhorst).

of Platydracus zonatus: Newton inSmetana and Davies 2000, Herman2001, etc.

Note on synonymy: The only located typematerial of P. badius (Mannerheim) and P.quadraticeps (Casey) consists of single females.These specimens are identified as P. zonatusbased on general agreement with the externalcharacters cited in the Platydracus species keyand the diagnostic description of P. zonatus, andbecause their pronotal punctation does not agreewith that of P. praetermissus.

Other material examined: 445 specimensthroughout the range of the species, all beingmales with an examined aedeagus. Full data willbe presented in the pending revision ofPlatydracus by Newton, but all records thatcould be georeferenced are shown in Map 26.

Description: With the characters of the P.cinnamopterus Complex (see above), plus:average pronotal length 2.31 mm (n = 9); head,pronotum and elytra usually medium to darkreddish, less often light red, pronotum more orless distinctly bicolored (darkest at anteriorangles and sides and sometimes along median,rest of disc lighter); antenna reaching middle ofpronotum when extended posteriorly,antennomere 9 slightly transverse; impunctatemedian line of pronotum at least one puncturediameter wide at narrowest point, usually abouttwo puncture diameters wide; emargination ofsixth visible male sternite in ventral viewdeep,about 1/3 as deep as wide; apex of medianlobe of aedeagus projected in parameral view,the converging sides of the projection in ventralview usually slightly concave and forming anobtuse angle, the apex narrowly rounded; sidesof median lobe in lateral view distinctly toothednear base of projection.

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Platydracus zonatus may bedistinguished from all other species of the genusin ECAS by the combination of generally reddishcolor (medium to very dark) (Fig. 11.21.1) with amore or less distinctly bicolored pronotum (Fig.7.3.3), black scutellum, and variegated abdomenwith only sparse gold setae and paired areas ofblack velvet setae on the first five visiblesegments; antennae with slightly transverseantennomere 9 (Fig. 7.3.2) ; and the narrow butcomplete impunctate medial line of pronotumabout 2 puncture diameters wide at narrowest(Fig. 7.9.5). It is definitively distinguished fromthe other members of the P. cinnamopterusComplex, P. cinnamopterus and P.praetermissus, by the structure of the aedeagus(Fig. 7.3.4, 11.21.3).

Platydracus zonatus is widelydistributed in the forested areas of eastern NorthAmerica: from southern Québec, west to Kansas,south to Texas and east to peninsular Florida.Unlike P. cinnamopterus, its northern range isapparently constrained by the distribution of theCarolinian forest and is therefore regularlyknown in Canada only from a few sites insouthern Ontario.

Eastern Canada: ON, QCAdjacent U.S.: MI, IN, OH, PA, NY, NH

In ECAS, P. zonatus specimens havebeen collected from March through November,with a majority (56%) found in May or June.

A majority of specimens of P. zonatusfrom ECAS with microhabitat data have beencollected in forests in various kinds of grounddebris including forest leaf litter (Fig. 11.21.2)and under stones and logs; one specimen wasfound in Sphagnum moss, one at blacklight atnight, several in prairie and old field areas, andseveral were collected in flight intercept traps.Unlike P. cinnamopterus, this species is notknown to occur under bark. Note that somehistorical records of P. cinnamopterus in theliterature may refer to this species. The partiallyillustrated larva of P. zonatus in Newton (1990)is based on a series of larvae collected with fourconfirmed adults of this species.

11.22 Staphylinus ornaticauda LeConte 1863

This species is unique among thespecies of Staphylinina with red elytra for itshead with patches of golden setae (Fig. 3.7.1). Itcannot be confused with any other species of thesubtribe in ECAS.

This species is spottily distributed inNorth America and records exist for Minnesota(historical), Iowa (no date), Michigan, Manitoba,Ontario, Québec and Nova Scotia. Itsdistribution in ECAS is given by (Map 28).

Eastern Canada: ON, QC, NSAdjacent U.S.: MI

Staphylinus ornaticauda has beencollected in ECAS from May to September, witha majority of the specimens being collected inJune to mid-July.

Staphylinus ornaticauda is the onlyspecies of the genus in the New World. It isrestricted to wetlands (bogs and fens) with anabundance of sphagnum moss, that occur southof the boreal forest zone; therefore its entireknown range is relatively small (Map 24) andmost of the known habitat exists in Canada.Specimens with label data indicate it can becaptured by pitfalls at the edges of bogs/fens andby sifting sphagnum moss. It was last collectedin the United States in 1922 (Michigan) and mostrecently from Canada in 1981 (Nova Scotia).Ten individuals (the most collected at one time)were collected by pitfall trapping, south of theRichmond area, Ottawa, Ontario, Canada in1978. This species is flightless and probably haslimited dispersal capabilities. Its restricteddistribution, narrow habitat requirements andapparent rarity suggest that this species warrantsfurther study as a possible species at risk ofextinction.

11.23 Tasgius ater (Gravenhorst 1802)

Tasgius ater is easily recognized amongthe species of Tasgius and other all-blackStaphylinina by the presence of smallerpunctures between the normal, larger puncturesof the head and pronotum (Fig. 8.1.1), and thewide spacing of these punctures; this produces anoverall glossy appearance (Fig. 11.23.1).

This species is native to the Palearcticregion but was first described by Gravenhorst(1802) from North America. Thus, it has been onthe continent for a very long time and has sincebecome widely distributed and established inECAS (Map 29). Tasgius ater is newly recordedfrom Michigan, Ohio, and Vermont based on thestudy of 68 specimens.

Eastern Canada: ON, QC, NB, NS, PE, NL (AllECAS provinces)

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Map 27. Distribution of Platydracus zonatus (Gravenhorst) in eastern Canada and adjacent United States.

Adjacent U.S.: MI, IN, OH, PA, NY, VT, NH,ME (All ECAS states)

Tasgius ater has been collected inECAS from January to November, with anincrease in abundance in July-August.

Tasgius ater is found under rocks andlogs in a variety of environments includingwoodlands, near water, and in open fields.Specimens have also been collected under loosebark, in rotting grass piles, and on beaches. Thisspecies often occurs in human-disturbed areas ingardens and occasionally in dwellings (Fig.11.23.2) but is less synanthropic than the otherTasgius occurring in ECAS.

11.24 Tasgius melanarius (Heer 1839)

Tasgius melanarius may bedistinguished from other Tasgius occurring inECAS by the combination of the pronotalpunctures of subequal size (Fig. 8.3.1), thepronotum widest near the apex (Fig. 8.3.1) andthe second antennomere darkened at its base(Fig. 8.3.2). The dense and subequal punctationof the pronotum causes an overall dullappearance (Fig. 3.5.1).

North American specimens of thisspecies were misidentified as T. globulifer until

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Map 28. Distribution of Staphylinus ornaticauda (LeConte) in eastern Canada and adjacent United States.

Newton (1987) clarified the matter, and reportedthat all previous North American records of T.globulifer were actually T. melanarius or T.winkleri (Bernhauer). Tasgius melanarius hasbeen collected on both east and west coasts andwas taken as early as 1935 in Québec. TheQuébec record is still the oldest known and T.melanarius has since expanded its range ineastern North America. Consequently, T.melanarius is here newly recorded from NewYork, Vermont, and Maine. Its distribution inECAS is given by Map 30.

UNITED STATES: ME: Waldo Co., Camden4mi N, 44.2833 -69.0667, 16 to 17-VII-1987,

J.K. Liebherr, 1 (CUIC). NY: Columbia Co.,Kinderhook, 42.4 -73.7, 9-VI-1978, N.M.Downie, 1 (FMNH). Essex Co., St. Armand,44.3969 -74.0594, 6-VII-1999, R. Turnbow, 1(Turnbow Coll.). Ontario Co., Geneva, 42.8667 -76.9833, 3 to 9 -VII-2002, J. Huether, 1(Brattain). Suffolk Co., Hauppauge, betw.Wheeler Rd. and N.Y.S. Office Bldg., 40.8 -73.2, 25-III-1975, under bark of dead fallen tree,F.C. Schlauch, 1 (CUIC); Long Island, CaumsettState Park, 40.9333 -73.4667, 21-VIII-1987,under board on beach, A. Newton and M.Thayer, 1 (FMNH); Long Island, East Hampton,40.9667 -72.1833, 27-VII-1965, A.J. Kistler, 1(DENH). Tompkins Co., Ithaca Vicinity, 42.4

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Map 29. Distribution of Tasgius ater (Gravenhorst) in eastern Canada and adjacent United States.

-76.5, 16-IX-1988, inlet, G.C. Eickwort, 1(CUIC). Westchester Co., Armonk, 41.1333 -73.7167, 15-VIII-1985, 2 (AMNH). VT:Addison Co., Ferrisburg, Lewis Creek, 44.22 -73.25, 10-IX-1980, J. Leonard, 1 (CUIC).Chittenden Co., Shelburne, Laplatte R. mouth,44.3833 -73.2333, 20-X-1984, S. Tschupp, 1(CUIC).

Eastern Canada: ON, QC, NB, NSAdjacent U.S.: OH, NY, VT, NH, ME

Tasgius melanarius has been collectedin ECAS from March to December, with peaksin abundance occurring in May and September.

This species is native to Europe and isfound in disturbed woodlands, coastal areas,urban areas and old fields under rocks, boards,litter, loose bark, and logs. It often occurs inyards (Fig. 11.24.1), sometimes falling intoswimming pools. In autumn this species,together with T. winkleri, is regularly observedon sidewalks and lawns, likely looking forfavorable overwintering sites.

11.25 Tasgius winkleri (Bernhauer 1906)

Tasgius winkleri is distinguished fromother Tasgius occurring in ECAS by thecombination of the pronotal punctures of

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Map 30. Distribution of Tasgius melanarius (Heer) in eastern Canada and adjacent United States.

subequal size (Fig. 8.2.2), the pronotum widestnear the middle (Fig. 8.2.2) and the secondantennomere not darkened at its base (Fig. 8.2.3).As in T. melanarius, the dense and subequalpunctation of the pronotum causes an overall dullappearance (Fig. 11.25.1).

This species was first correctlyrecognized in eastern and western NorthAmerica by Newton (1987), who indicated that ithad been collected as early as 1938 in NewYork. Prior to 1987, both this species and T.melanarius had been misidentified as T.globulifer. Additional specimens from Syosset,NY indicate that Tasgius winkleri has been inNorth America at least since 1931 and new

records from eastern Canada (Ontario),Michigan, and Pennsylvania suggest that it isexpanding its range in North America. Its rangein ECAS is given by Map 31.

CANADA: ON: 176 specimens.UNITED STATS: MI: Calhoun Co., nearMarshall, I-69 rest stop, litter, 42.3278 -84.9901,10-X-2008, A. Brunke and S.A Marshall, 1(DEBU). Gratiot Co., Sumner Twp. Sec. 24,43.3333 -84.7833, 8-IX-1977, R. D. Ward, 1(CUIC). PA: Montgomery Co., Cold Point,40.1167 -75.2667, 23-IX-1956, 1 (AMNH).

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Map 31. Distribution of Tasgius winkleri (Bernhauer) in eastern Canada and adjacent United States.

Eastern Canada: ONAdjacent U.S.: MI, PA, NY, NH

Tasgius winkleri has been collected inECAS in April to November, with an increase inabundance during September.

Tasgius winkleri inhabits openwoodlots, old fields, shorelines, wetland edges,and backyards, where it is found under rocks andin leaf litter. It has also been collected from apile of rotting grass and in a compost heap. It ispronouncedly synanthropic and generally onlyoccupies habitats that have been degraded byhuman activity.

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Acknowledgements

We thank the following photographers for the use of their work, which contributed greatly to the aestheticand scientific value of this publication:A. M. Bradford Alexandria, VA, USA. http://www.plantbonz.comC. Eisemann Pelham, MA, USA. http://www.charleyeiseman.comD.K.B. Cheung Guelph, ON, Canada. http://www.dkbdigitaldesigns.comJ. McClarin Nashua, NH. http://bugguide.net/user/view/1611T. Murray Groton, MA. http://www.pbase.com/tmurray74S. Justis Virginia Beach, VA, USA. http://mysite.verizon.net/psjustis/index.htmS. Perraut Hammond, ON, Canada. http://bugguide.net/user/view/9196______________________________________________________________________________________We are grateful to the following institutions and individuals for making specimens, including some types,available for study:

ACNS Agriculture and Agri-Food Canada, Nova Scotia, Canada (Susan Westby)ACPE Agriculture and Agri-Food Canada, Prince Edward Island, Canada (Christine Noronha)AFC Atlantic Forestry Centre, Canadian Forest Service, New Brunswick, Canada (Jonathan

Sweeny)AMNH American Museum of Natural History, New York, New York, USA (Lee H. Herman)ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA (W. Wayne Moss)ANIC Australian National Insect Collection, Canberra, Australia (John F. Lawrence, Tom Weir)Baldwin Brian Baldwin Collection, Mena, Arkansas, USA (Brian Baldwin)BMNH The Natural History Museum, London, United Kingdom (Martin Brendell, Peter

Hammond)Brattain R. Michael Brattain Collection, Lafayette, Indiana, USA (R. Michael Brattain)BU Boston University, Boston, Massachusetts, USA (James F. A. Traniello)CAS California Academy of Sciences, San Francisco, California, USA (David H. Kavanaugh)CASM Chicago Academy of Sciences, Chicago, Illinois, USA (Douglas J. Taron)CBU Cape Breton University, Nova Scotia, Canada (David. B. McCorquodale)CGMC Christopher G. Majka Collection, Nova Scotia, Canada (Christopher G. Majka)Chantal Claude Chantal Collection, Varennes, Québec, Canada (Claude Chantal)Cicero Joseph Cicero Collection, Gainesville, Florida, USA (Joseph Cicero)CMNC Canadian Museum of Nature, Québec, Canada (Robert S. Anderson, François Génier)CMNH Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA (George Wallace,

John Rawlins)CNC Canadian National Collection of Insects, Ottawa, Ontario, Canada (J. Milton Campbell,

Anthony Davies, Patrice Bouchard)CNHM Cincinnati Museum of Natural History, Cincinnati, Ohio, USA (Kevina Vulinec)CSCA California State Collection of Arthropods, Sacramento, California (Fred G. Andrews)CUIC Cornell University, Ithaca, New York, USA (L.L. Pechuman, James K. Liebherr)Curtis R. Curtis Collection, Chicago, Illinois, USA (R. Curtis)DAL Dalhousie University, Nova Scotia, Canada (Tatiana Rossolimo)DEBU University of Guelph Insect Collection, Guelph, Ontario, CanadaDEI Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (Lothar

Zerche)DENH University of New Hampshire, Durham, New Hampshire, USA (Donald S. Chandler)DEPAUW Depauw University, Greencastle, Indiana, USADHWC David H. Webster Collection, Nova Scotia, Canada (David. H. Webster)EIU Eastern Illinois University, Charleston, Illinois, USA (Michael Goodrich)

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EMEC Essig Museum of Entomology, Berkeley, California, USA (Cheryl B. Barr, John T.Doyen)

FMNH Field Museum of Natural History, Chicago, Illinois, USA (Henry Dybas, Rupert Wenzel)FSCA Florida State Collection of Arthropods, Gainesville, Florida, USA (Michael C. Thomas)GSC Gary Selig Collection, Nova Scotia, Canada (Gary Selig)Gusarov Vladimir I. Gusarov Collection, Oslo, Norway (Vladimir I. Gusarov)Hamilton Robert Hamilton Collection, Chicago, Illinois, USA (Robert Hamilton)HNHM Hungarian Natural History Museum, Budapest, Hungary (Zoltan Kaszab, Ottó Merkl)Huben Michael Huben [formerly Huybensz] Collection, Massachusetts, USA (Michael Huben)INHS Illinois Natural History Survey, Champaign, Illinios, USA (Milton W. Sanderson,

Kathryn C. McGiffen)IRCW University of Wisconsin, Madison, Wisconsin, USA (Steven Krauth)IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium (Leon Baert,

Didier Drugmand)ISMC Indiana State Museum, Indianapolis, Indiana, USA (Robert Waltz)ISUI Iowa State University, Ames, Iowa, USA (Ray Miller)Ivie Michael A. Ivie Collection, Bozeman, Montana, USA (Michael A. Ivie)JOC Jeffrey Ogden Collection, Nova Scotia, Canada (Jeffrey Ogden)JohnsonPJ Paul J. Johnson Collection, Brookings, South Dakota, USA (Paul J. Johnson)Kistner David H. Kistner Collection, Chico, California, USA (gradually to FMNH) (David H.

Kistner)KSEM Snow Entomological Museum, Lawrence, Kansas, USA (James S. Ashe, Robert G.

Brooks)KSUC Kansas State University, Manhattan, Kansas, USA (H. Derrick Blocker)LACM Natural History Museum of Los Angeles County, Los Angeles, California, USA (Charles

L. Hogue, Roy R. Snelling)LEMQ Lyman Entomological Museum, Ste-Anne-de-Bellevue, Québec, Canada (Terry A.

Wheeler)LFC Laurentian Forestry Centre, Sainte-Foy, Québec, Canada (Jan Klimaszewski)LSAM Louisiana State Arthropod Museum, Baton Rouge, Louisiana, USA (Victoria Moseley

Bayless)MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA (John F. Lawrence,

Philip D. Perkins)MEM Mississippi Entomological Museum, Mississippi State, Mississippi, USA (Richard L.

Brown, Patricia R. Miller)MHNG Museum d’Histoire Naturelle, Geneva, Switzerland (Ivan Löbl)Miliotis Paul S. Miliotis Collection, Dunstable, Massachusetts, USA (Paul S. Miliotis)MPM Milwaukee Public Museum, Milwaukee, Wisconsin, USA (Gerald R. Noonan)MSUC Michigan State University, East Lansing, Michigan, USA (Roland L. Fischer)MUNC Memorial University, St. John's, Newfoundland, CanadaMZHF Finnish Museum of Natural History, Helsinki, Finland (Hans Silfverberg)MZLU Lund University, Lund, Sweden (Roy Danielsson)MZPW Museum of the Institute of Zoology, Warsaw, Poland (S. Adam Ślipiński)MZSP Museu de Zoologia da Universidade de São Paulo, São Paulo, Brazil (Hans Reichardt)NBM New Brunswick Museum, New Brunswick, Canada (Donald McAlpine)NelsonG Gayle H. Nelson Collection, Blue Springs, Missouri, USA (Gayle H. Nelson)NMW Naturhistorisches Museum Wien, Wien, Austria (Harald Schillhammer, Heinrich

Schönmann)NSAC Nova Scotia Agricultural College, Nova Scotia, Canada (Christopher Cutler)

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NSMC Nova Scotia Museum, Nova Scotia, CanadaOSAC Oregon State Arthropod Collection, Corvallis, Oregon, USA (Michael D. Schwartz)OSEC Oklahoma State University, Stillwater, Oklahoma, USA (W. A. Drew)Patrick L. Brian Patrick Collection, Mitchell, South Dakota, USA (L. Brian Patrick)PURC Purdue University, West Lafayette, Indiana, USA (Arvin Provonsha)RGC Rebecca Gorham Collection, Nova Scotia, Canada (Rebecca Gorham)ROM Royal Ontario Museum, Toronto, Ontario, Canada (Glenn B. Wiggins)RWC Reginald Webster Collection, New Brunswick, Canada (Reginald Webster)SBMN Santa Barbara Museum of Natural History, Santa Barbara, California, USA (Scott E.

Miller)Schulke Michael Schülke Collection, Berlin, Germany (Michael Schülke)SDMC San Diego Natural History Museum, San Diego, California, USA (David Faulkner)SDSU South Dakota University, Brookings, South Dakota, USA (Edward U. Balsbaugh Jr.)Sikes Derek S. Sikes Collection, Fairbanks, Alaska, USA (Derek S. Sikes)SIUC Southern Illinois University, Carbondale, Illinois, USA (Jay McPherson)SMNS Staatliches Museum fur Naturkunde, Stuttgart, Germany (Wolfgang Schawaller)SMU Saint Mary’s University, Nova Scotia, Canada (Doug Stongman)Stephan Karl H. Stephan Collection, Red Oak, Oklahoma, USA (to TAMU) (Karl H. Stephan)Suter Walter R. Suter Collection, Kenosha, Wisconsin, USA (to FMNH) (Walter R. Suter)TAMU Texas A&M University, College Station, Texas, USA (Edward G. Riley)Thomas Paul N. Thomas Collection, Lombard, Illinois, USA (Paul N. Thomas)Turnbow Robert H. Turnbow Collection, Fort Rucker, Alabama, USA (Robert H. Turnbow)UAAM Arthropod Museum, University of Arkansas, Fayetteville, Arkansas (Robert T. Allen)UAIC University of Arizona, Tucson, Arizona, USA (Carl A. Olson)UASM University of Alberta, Edmonton, Alberta, Canada (Danny Shpeley)UBCZ Spencer Museum, University of British Columbia, Vancouver, British Columbia, Canada

(Robert A. Cannings)UCDC University of California at Davis, California, USA (Robert Schuster, Lynn Kimsey)UCR University of California at Riverside, California, USA (Saul I. Frommer, Ian Moore)UGCA University of Georgia, Athens, Georgia, USA (Warren T. Atyeo)ULKY University of Louisville, Louisville, Kentucky, USA (Charles V. Covell Jr.)UMIC University of Mississippi, University, Mississippi, USA (Paul K. Lago)UMMZ University of Michigan, Ann Arbor, Michigan, USA (Richard D. Alexander, Barry M.

O'Connor)UMRM University of Missouri, Columbia, Missouri, USA (Wilbur R. Enns)UMSP University of Minnesota, St. Paul, Minnesota, USA (Philip J. Clausen)UNSM University of Nebraska, Lincoln, Nebraska, USA (Brett C. Ratcliffe)UMNB Université de Moncton, New Brunswick, Canada (Pauline Duerr)USNM Smithsonian Institution, Washington D.C., USA (Terry L. Erwin, Gloria M. House)Valentine Barry D. Valentine Collection, Columbus, Ohio, USA (Barry D. Valentine)VMNH Virginia Museum of Natural History, Martinsville, Virginia, USA (Richard L. Hoffman)Wappes James E. Wappes Collection, Bulverde, Texas, USA (James E. Wappes)WFBM W. F. Barr Entomological Collection, University of Idaho, Moscow, Idaho, USA

(William F. Barr)ZIN Zoological Institute, Russian Academy of Sciences, St. Petersburg, Russia (Vladimir

Gusarov, Alexey Solodovnikov)ZMHB Museum fur Naturkunde der Humboldt Universitat, Berlin, Germany (Fritz Hieke,

Manfred Uhlig)ZSMC Zoologische Sammlung des Bayerischen, München, Germany (Martin Baehr)

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A. Brunke would like to thank D. K. B.Cheung (DEBU) for technical assistance andessential training used to construct the interactivekeys. A. Brunke also thanks D. K. B. Cheungand M. Jackson (DEBU) for assistance withmapping software and creation of custom maps.Thanks to J. Renkema (NSAC) and D. McAlpine(NBM) for bringing the presence of P.cinnamopterus in Nova Scotia and NewBrunswick to our attention, respectively. Wethank R. Webster (AFC) and A. Smetana (CNC)for recognizing the misidentification present inKlimaszewski et al. (2005). M. Thayer (FMNH)read an earlier version of the manuscript andprovided many helpful comments. Funding forthis project was provided in part by an NSERCPSG-M awarded to A. Brunke. A. Newtonacknowledges U.S. National Science Foundationgrant BSR-8906825 for support of databasing,dissection, and other activities in support of hisongoing revision of Platydracus, and thanks J.Klimaszewski for producing the aedeagaldrawings of the P. cinnamopterus complexspecies used here (a more complete set ofacknowledgments related to this revision willaccompany its publication). C. Majka thanks theBoard of Governors of the Nova Scotia Museumfor ongoing support.

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Stevens, C. and DeKimpe, N. 1993. Synthesis of stenusine, the spreading agent of the beetle Stenus comma.The Journal of Organic Chemistry 58(1): 132-134.

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Figures 1.1.1-1.3.1: 1.1.1 Oxyporus rufipennis LeConte, dorsal habitus and mandibles; 1.1.2 O. rufipennis,labial palpi; 1.2.1 Scaphidium quadriguttatum Say, dorsal habitus; 1.2.2 S. quadriguttatum, lateral habitus.1.2.3 S. quadriguttatum, head, showing antennal insertion; 1.3.1 Micropeplus browni Campbell, dorsalhabitus.

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Figures 1.4.1-1.8.1. 1.4.1 Rybaxis mystica (Casey), below, Adranes lecontei Brendel, above; 1.5.1Euconnus Thomson sp., dorsal habitus; 1.6.1 Pseudopsis sulcata Herman complex, dorsal habitus; 1.7.1Olisthaerus substriatus Gyllenhal, dorsal habitus; 1.8.1 Megalopinus caelatus Gravenhorst, dorsal habitus.

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Figures 1.9.1-1.11.1. 1.9.1 Stenus laccophilus Casey, dorsal habitus; 1.9.2 Dianous chalybaeus LeConte,from moss beside a headwater stream; 1.9.3 S. laccophilus, close-up of dorsal head; 1.10.1 EuaesthetusGravenhorst sp., dorsal habitus; 1.10.2 Euaesthetus sp., ventral head; 1.11.1 Bledius semiferrugeneusLeConte, ventrolateral view of abdomen.

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Figures 1.11.2-1.12.2. 1.11.2 Coprophilus striatulus Fabricius, dorsal habitus; 1.11.3 Deleaster dichrousGravenhorst, dorsal habitus; 1.11.4 Syntomium grahami Hatch, dorsal view of forebody; 1.11.5. C.striatulus, head; 1.12.1 Philonthus cognatus Stephens, ventrolateral view of abdomen; 1.12.2 P. cognatus,dorsolateral view of head.

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Figures 1.12.3-1.12.8. 1.12.3 Philonthus cognatus Stephens, dorsal head, showing lack of ocelli; 1.12.4 P.cognatus, right antenna; 1.12.5 Diochus schaumi Kraatz, dorsal abdomen with paratergites; 1.12.6Dinothenarus badipes (LeConte), anteroventral view of prolegs; 1.12.7 Platydracus viridanus (Horn),dorsal view of head showing neck; 1.12.8. Quedius peregrinus Gravenhorst, dorsal view of head showingneck.

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Figures 1.12.9-1.13.2. 1.12.9 Philonthus cognatus Stephens, lateral view of pronotum and proleg; 1.12.10Erichsonius nanus (Horn), ventrolateral view of pronotum; 1.12.11 Atanygnathus bicolor (Casey), rightelytron; 1.12.12 A. bicolor, habitus; 1.13.1 Aleochara bilineata Gyllenhal, head; 1.13.2 Trichopseniusxenoflavipes Seevers, ventral.

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Figures 1.13.3-1.14.3. 1.13.3 Trichopsenius xenoflavipes Seevers, dorsal habitus; 1.13.4 Reticulotermesvirginicus (Banks); 1.13.5 Oligota Mannerheim sp., left antenna; 1.14.1 Tachyporus maculicollis LeConte,ventral view of thorax; 1.14.2 Sepedophilus littoreus (L.), dorsal abdomen; 1.14.3 Tachinus fimbriatusGravenhorst, right antenna.

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Figures 1.14.4-1.17.1. 1.14.4 Tachinus fimbriatus Gravenhorst, lateral view of elytron; 1.15.1 Acidotasubcarinata, dorsal head with ocelli; 1.15.2 Eusphalerum pothos Mannerheim, habitus; 1.15.3 Pycnoglyptacampbelli Gusarov, habitus; 1.16.1 Proteinus Latreille sp., dorsal habitus; 1.17.1 Renardia nigrellaLeConte, dorsal abdomen.

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Figures 1.17.2-1.17.3. 1.17.2 Thoracophorus costalis Erichson, dorsal abdomen; 1.17.3 Eleusis pallidaLeConte, head and pronotum; 1.18.1 Palaminus Erichson sp., abdomen; 1.18.2 Palaminus sp., anteroventalview of forebody showing maxillary palpus; 1.18.3 Homaeotarsus bicolor (Gravenhorst), lateral view ofthorax; 1.18.4 Lobrathium grande (LeConte), dorsal head with maxillary palpus.

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Figures 1.18.5-1.19.5. 1.18.5 Lathrobium Gravenhorst sp., dorsal view of thorax; 1.19.1 Charhyphuspicipennis LeConte, anteroventral view of procoxae; 1.19.2 C. picipennis, dorsal pronotum and elytra,showing punctation; 1.19.3 C. picipennis, left proleg; 1.19.4 C. picipennis, dorsal pronotum and elytra;1.19.5 C. picipennis, dorsal pronotum with serrate margins.

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Figures 1.19.6-1.21.3. 1.19.6 Phloeocharis subtilissima Mannerheim, habitus; 1.20.1 Siagonium punctatum(LeConte), dorsal elytra showing punctures; 1.20.2, S. punctatum, left protibia; 1.21.1 Habrocerus magnusLeConte, dorsal pronotum; 1.21.2 H. capillaricornis Gravenhorst, right antenna; 1.21.3. H. capillaricornis,dorsal habitus.

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Figures 1.22.1-2.2.3. 1.22.1 Trichophya pilicornis (Gyllenhal), dorsal habitus. 2.1.1 Gyrohypnus campbelliSmetana, elytra; 2.1.2 Xantholinus linearis (Olivier), prosternum; 2.2.1 Platydracus maculosus(Gravenhorst), elytra; 2.2.2 Philonthus cognatus Stephens, head showing impunctate neck; 2.2.3 P.cognatus, head showing antennae insertions.

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Figures 2.2.4-2.2.9. 2.2.4 Philonthus cognatus Stephens, prosternum without plates; 2.2.5 Atanygnathusbicolor (Casey), forebody; 2.2.4 P. caeruleipennis Mannerheim, dorsal view of pronotum; 2.2.7 Quediuscanadensis Casey, dorsal view of pronotum; 2.2.8 P. cognatus, hind tarsus without empodial setae; 2.2.9 Q.peregrinus Gravenhorst, mesotarsus.

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Figures 2.2.10-2.2.15. 2.2.10 Anaquedius vernix (LeConte), hind tarsus; 2.2.11 Heterothops fusculus(LeConte), head with maxillary palpus; 2.2.12 Tympanophorus puncticollis (Erichson), forebody; 2.2.13Platydracus viridanus Horn, forebody; 2.2.14 T. puncticollis, head with labial palpi; 2.2.15 Tasgiusmelanarius (Heer), head with labial palpi.

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Figures 2.3.1-3.3.2. 2.3.1 Diochus schaumi (Kraatz), dorsal forbody; 2.4.1 Atrecus americanus (Casey),head; 3.1.1 Creophilus maxillosus villosus Gravenhorst, pronotum; 3.2.1 Ontholestes cingulatus(Gravenhorst), pronotum; 3.3.1 Platydracus comes (LeConte), pronotum; 3.3.2 P. cinnamopterus(Gravenhorst), dorsal view of head showing lateral setae

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Figures 3.4.1-3.6.1. 3.4.1. Dinothenarus capitatus (Bland), dorsal view of head showing lateral setae; 3.4.2D. capitatus, habitus.3.4.3 D. badipes (LeConte), ventral abdomen; 3.4.4 D. badipes, maxillary palpus;3.5.1 Tasgius melanarius (Heer), dorsal habitus; 3.6.1 Ocypus nitens (Schrank), habitus.

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Figures 3.6.2-5.1.1. 3.6.2 Ocypus nitens (Schrank), palpi; 3.6.3 O. nitens, ventral view of abdomen; 3.7.1Staphylinus ornaticauda LeConte, dorsal habitus; 4.1.1 Dinothenarus capitatus (Bland), darkenedspecimen habitus; 4.2.1 D. badipes (LeConte), habitus; 5.1.1 O. brunnipes (Fabricius), dorsal habitus.

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Figures 6.1.1-7.3.1. 6.1.1 Ontholestes cingulatus (Gravenhorst), habitus; 6.2.1 O. murinus (L.), dorsalhabitus;7.1.1 Platydracus femoratus (Fabricius), forebody; 7.2.1 P. exulans (Erichson), pronotum, showinguneven punctation; 7.2.2 P. exulans, prontum, showing impuncte line; 7.3.1 P. zonatus (Gravenhorst),lateral portion of pronotum, showing even punctation.

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Figures 7.3.2-7.5.2. 7.3.2 Platydracus zonatus (Gravenhorst), apical antennomeres; 7.3.3 P. zonatus, lateralportion of pronotum, showing coloration;7.3.4 Platydracus zonatus, aedeagus, parameral view; 7.4.1 P.praelongus (Mannerheim), pronotum. 7.5.1 P. comes (LeConte), showing scutellum and elytral spots. 7.5.2P. comes, showing complete impunctate line of pronotum.

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Figures 7.6.1-7.8.1. 7.6.1 Platydracus mysticus (Erichson), dorsal view of forebody, showing darkscutellum and spotless elytra; 7.6.2 P. mysticus, dorsal view of forebody, showing incomplete impunctateline of pronotum;7.6.3 P. mysticus, dorsal habitus; 7.6.4 P. mysticus, right antenna; 7.7.1 P. maculosus(Gravenhorst), pronotum; 7.8.1 P. fossator (Gravenhorst), dorsal habitus of maculate form.

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Figures 7.8.2-7.9.5. 7.8.2 Platydracus fossator (Gravenhorst), dorsal habitus of immaculate form; 7.9.1 P.cinnamopterus (Gravenhorst), dorsal habitus of regular colour morph;7.9.2 P. cinnamopterus, dorsalabdomen; 7.9.3 P. cinnamopterus, pronotum with impunctate median line complete; 7.9.4 P.cinnamopterus, dorsal habitus of rufous color morph; 7.9.5 P. cinnamopterus, pronotum with narrowimpunctate median line.

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Figure 7.9.6-7.10.3. 7.9.6 Platydracus cinnamopterus (Gravenhorst), aedeagus, parameral view; 7.9.7 P.cinnamopterus, apical antennomeres;7.9.8 P. cinnamopterus, pronotum, showing coloration; 7.10.1 P.viridanus (Horn), dorsal abdomen; 7.10.2 P. viridanus, dorsal habitus; 7.10.3 P. viridanus, apex of lateralelytron.

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Figures 7.11.1-7.14.1. 7.11.1 Platydracus immaculatus (Mannerheim), dorsal habitus; 7.12.1 P. violaceus(Gravenhorst), dorsal habitus;7.12.2 P. violaceus, lateral view of left elytron; 7.13.1 P. tomentosus(Gravenhorst), dorsal habitus; 7.13.2 P. tomentosus, head and right antenna; 7.14.1 P. praetermissusNewton, spec. nov., pronotum.

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Figures 7.14.2-8.2.3. 7.14.2 Platydracus praetermissus Newton spec nov., apical antennomeres; 7.14.3 P.praetermissus, aedeagus, parameral view; 8.1.1 Tasgius ater (Gravenhorst), pronotum; 8.2.1 T. winkleri(Bernhauer), pronotum showing close punctation; 8.2.2 T. winkleri (Bernhauer), pronotal shape; 8.2.3 T.winkleri, base of right antenna.

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Figures 8.3.1-9.1.4. 8.3.1 Tasgius melanarius (Heer), pronotum; 8.3.2 T. melanarius, base of right antenna;9.1.1 Drusilla canaliculata Fabricius; 9.1.2 Gymnusa atra Casey; 9.1.3 Tachyusa Erichson sp., from beaverlodge; 9.1.4 Xenodusa cava LeConte, from Formica sp. colony in dry log.

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Figures 9.2.1-9.6.2. 9.2.1 Euaesthetus Gravenhorst sp., from sifting deep piles of damp leaf litter; 9.2.2Euaesthetus sp; 9.3.1 Habrocerus magnus LeConte, dorsal habitus; 9.3.2. H. capillaricornis, from leaflitter; 9.6.1 Olisthaerus megacephalus Zetterstedt, dorsal habitus; 9.6.2 O. substriatus Gyllenhal, fromunder bark in a relict boreal forest (Jefferson's Notch, NH).

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Figures 9.6.3-9.7.4. 9.6.3 Olisthaerus substriatus Gyllenhal, from under pine bark; 9.7.1 Lesteva pallipesLeConte, found in flood debris; 9.7.2. Brathinus varicornis LeConte, from under a rock near a smallstream; 9.7.3 Boreaphilus henningianus Sahlberg; 9.7.4. Acidota subcarinata Erichson aggregating in fall.

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Figures 9.7.5-9.9.3. 9.7.5 Coryphium nigrum Campbell with springtail prey; 9.8.1 Thoracophorus costalisErichson, found under wood in leaf litter; 9.9.1 Oxyporus vittatus Gravenhorst, extremes of colourvariability from the same locality; 9.9.2 O. quinquemaculatus LeConte, from fungus in floodplain forest;9.9.3 O. major Gravenhorst, from a white gilled mushroom in a moist hardwood forest.

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Figures 9.10.1-9.10.5. 9.10.1 Syntomium grahami Hatch, dorsal habitus; 9.10.2 Thinodromus Kraatz sp.,sifted from deep pockets of litter between boulders near a river; 9.10.3 Bledius Leach sp., at porch light;9.10.4 Anotylus Thomson sp.; 9.10.5 Apocellus Erichson sp.

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Figures 9.11.1-9.11.5. 9.11.1 Homaeotarsus bicolor, from under stones in a dry streambed; 9.11.2Paederus Fabricius sp.; 9.11.3 Astenus discopunctatus (Say); 9.11.4 Achenomorphus corticinus(Gravenhorst), sifted from leaf litter; 9.11.5 Sunius confluentus (Say), from under bark.

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Figures 9.12.1-9.15.3. 9.12.1 Charhyphus picipennis LeConte; 9.13.1 Siagonium punctatum (LeConte),major male, dorsal habitus; 9.13.2 S. punctatum, minor male from under beech bark; 1.14.1 MegarthrusStephens sp., pronotum; 9.15.1 Batrisodes lineaticollis (Aube), male; 9.15.2 Tyrus semiruber Casey, male,found under pine bark; 9.15.3 Ceophyllus monilis LeConte, male, found on a freshly cut pine stump.

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Figures 9.15.4-9.18.1. 9.15.4 Ctenisodes piceus (LeConte), male, found under a rock; 9.16.1 Pseudopsissulcata Herman complex, dorsal apex of abdomen; 9.16.2 P. sulcata complex; 9.17.1 Scaphisoma rubensCasey, from fungus on a tree; 9.17.2 Baeocera Erichson sp., feeding on the fruiting body of a slime mould;9.18.1 Stenichnus Thomson sp. (~1.2mm long).

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Figures 9.18.2-9.20.3. 9.18.2 Euconnus Thomson sp., found under a stone; 9.19.1 Stenus Latreille sp.,swept from sedges in a lakeside marsh; 9.19.2 St. insperatus Puthz, from debris next to a creek; 9.19.3 St.morio Gravenhorst, from under bark of a tree overhanging water; 9.20.1 Tachinus corticinus Gravenhorst;9.20.2 Sepedophilus littoreus, from flood debris; 9.20.3 Coproporus ventriculus (Say), from under bark.

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Figures 9.20.4-10.1.2. 9.20.4 Nitidotachinus scrutator (Gemminger & Harold), from wet sphagnum in aboggy area; 9.20.5 Lordithon kelleyi (Malkin); 9.20.6 Tachyporus pulchrus Blatchley, from a tallgrassprairie; 9.21.1 Trichophya pilicornis (Gyllenhal), from sifting leaf litter; 10.1.1 Diochus schaumi Kraatz,dorsal habitus; 10.1.2 D. schaumi, showing dark abdomen.

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Figures 10.1.3-10.4.1. 10.1.3 Diochus schaumi (Kraatz), in copula; 10.2.1 Atrecus macrocephalus(Nordmann), habitus; 10.2.2 A. macrocephalus; 10.3.1 Tympanophorus puncticollis (Erichson), dorsalhabitus; 10.3.2. T. puncticollis, from rotting sawdust in an abandoned sawmill; 10.4.1 Philonthus politus(L.), from compost.

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Figures 10.4.2-10.5.1. 10.4.2 Philonthus caeruleipennis, from compost; 10.4.3 Bisnius blandus(Gravenhorst); 10.4.4 Hesperus apicialis (Say), from a beech treehole, collected with larvae; 10.4.5Laetulonthus laetulus (Say), in wood shavings from a freshly cut pine tree; 10.4.6 Erichsonius rosellusFrank, sifted from leaf litter in a marsh; 10.5.1 Quedius plagiatus Mannerheim.

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Figures 10.5.2-10.8.1. 10.5.2 Quedius mesomelinus Marsham, found in rotting mushrooms; 10.5.3Acylophorus sp., captured by treading debris from beaver dam; 10.5.4 Hemiquedius ferox LeConte, from abeaver lodge; 10.7.1 Heterothops fusculus (LeConte); 10.8.1 Stictolinus flavipes (LeConte).

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Figures 10.8.2-11.1.2. 10.8.2 Lithocharodes longicollis (LeConte), from leaf litter on sandy substrate;10.8.3 Gyrohypnus fracticornis (O.F. Müller), from yard compost; 10.8.4 Gauropterus fulgidus, fromgarden compost; 10.8.5 Nudobius cephalus (Say), from under pine bark; 10.8.6 Oxybleptes kiteleyiSmetana; 11.1.1 Creophilus maxillosus villosus Gravenhorst, habitus; 11.1.2 C. m. villosus, escapemechanism on dead raccoon.

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Figures 11.1.3-11.6.1. 11.1.3 Creophilus maxillosus maxillosus (L.), forebody with mostly dark setae;11.1.4 C. m. villosus Gravenhorst, forebody with mostly pale setae; 11.2.1 Dinothenarus badipes(LeConte), from under rocks near a stream; 11.3.1 D. capitatus (Bland), from carrion trap; 11.5.1 Ocypusnitens (Schrank); 11.6.1 Ontholestes cingulatus (Gravenhorst), from compost.

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Figures 11.8.1-11.12.1. 11.8.1 Platydracus cinnamopterus (Gravenhorst), from flood debris; 11.8.2 P.cinnamopterus, aedeagus, lateral view; 11.9.1 P. comes (LeConte), dorsal habitus; 11.10.1 P. exulans(Erichson); 11.11.1 P. femoratus (Fabricius), dorsal habitus; 11.12.1 P. fossator (Gravenhorst), swept fromtall grass along a driveway.

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Figure 11.13.1-11.17.1. 11.13.1 Platydracus immaculatus (Mannerheim), found crossing a driveway;11.14.1 P. maculosus (Gravenhorst), dorsal habitus; 11.14.2 P. maculosus, from early-stage carrion;11.16.1 P. praelongus (Mannerheim), dorsal habitus; 11.16.2 P. praelongus, from under a board alongocean shoreline; 11.17.1 P. praetermissus Newton spec. nov., dorsal habitus.

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Figures 11.17.2-11.21.3. 11.17.2 Platydracus praetermissus Newton spec nov., aedeagus, lateral view;11.19.1 P. violaceus (Gravenhorst), from under the bark of a fallen maple; 11.20.1. P. viridanus (Horn),from tall grass near a river; 11.21.1 P. zonatus (Gravenhorst), dorsal habitus; 11.21.2 P. zonatus, from leaflitter; 11.21.3 P. zonatus, aedeagus, lateral view.

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Figures 11.23.1-11.25.2. 11.23.1 Tasgius ater (Gravenhorst), dorsal habitus; 11.23.2 T. ater, from thebasement of an old building; 11.24.1 T. melanarius (Heer), from under a composter; 11.25.1 T. winkleri(Bernhauer), dorsal habitus; 11.25.2 T. winkleri, from an urban garden.