Diversity and Taxonomic Review Of

Diversity and taxonomic review of Leptusa Kraatz (Coleoptera: Staphylinidae: Aleocharinae) from Great Smoky Mountains National Park, U.S.A., with descriptions of four new species

JONG-SEOK PARK1,4, CHRISTOPHER E. CARLTON2 & MICHAEL L. FERRO3 Louisiana State Arthropod Museum, Department of Entomology, LSB 400, Louisiana State University Agricultural Center, Baton Rouge, LA 70803, U.S.A. E-mail: [email protected], [email protected], [email protected] 4Corresponding author

Abstract

A taxonomic review of the genus Leptusa Kraatz from Great Smoky Mountains National Park is presented. Ten species are recognized, and four new species are described and illustrated: L. (Adoxopisalia) pseudosmokyiensis Park & Carlton, sp. nov., L. (A.) stocksi Park & Carlton, sp. nov., L. (Dysleptusa) gimmeli Park & Carlton, sp. nov., and L. (Eucryptusa) ferroi Park & Carlton, sp. nov. The previously undescribed female of L. (D.) pusio (Casey) is also reported for the first time. Descriptions, habitus illustrations and line drawings of diagnostic characters of all species occurring in Great Smoky Mountains National Park are provided. A key is provided that will allow discrimination of 16 of the 18 species of Leptusa known to occur in the eastern United States. “Overlooked Syndrome”, the phenomena where undescribed species persist in a well known fauna, is defined and discussed.

Key words: taxonomy, Homalotini, Great Smoky Mountains National Park, Overlooked Syndrome

Introduction

The genus Leptusa Kraatz, 1856 includes over 400 species and is found in all geographical regions except Australia, Madagascar, and the Pacific islands (Pace 1989). After Pace (1989) published his monograph treating the world fauna of the genus, Assing described numerous species from the Palearctic fauna (Assing 2001, 2002, 2003, 2004, 2007, 2009a, 2009b, 2009c). Some North American species were treated by Gusarov (2003), and Klimaszewski et al. (2004) reviewed the Canadian species. The Coleoptera Taxonomic Working Group of the All Taxa Biodiversity Inventory (ATBI) in Great Smoky Mountains National Park (GSMNP) has been active since June 2001 (Carlton & Bayless 2007). To date 2518 beetle species have been recorded from the park including more than 40 species new to science. The ATBI and related research resulted in the collection of 194 specimens of Leptusa from GSMNP. The majority of these specimens were collected as the result of two studies by M. L. Ferro and M. L. Gimmel (unpublished). During April 2006, 90 emergence chambers were filled with hardwood coarse woody debris (CWD) of various sizes and decay classes from six locations in GSMNP. Beetles emerging from the wood were collected monthly over a period of two years. During fall, 2006 and spring, 2007 a comparison of leaf litter and coarse woody debris decay class V (see Pyle & Brown 1999 for definitions of course woody debris decay classes) habitats was conducted at the same sites. Specimens were extracted from sifted leaf litter and woody debris samples using Berlese funnels. In total ten species were identified, four of which were undescribed. We also discovered the female of L. pusio (Casey), which has not been described previously. Here we describe or provide updated redescriptions of all species of Leptusa found in the GSMNP including four new species. Redescriptions are provided to better compare and contrast newly described species with existing fauna and to facilitate recognition of additional undescribed species that might be discovered in future. Habitus illustrations and line drawings of diagnostic characters are provided for these

Accepted by V. Gusarov: 4 Oct. 2010; published: 29 Oct. 2010 1 species. A key is provided that will allow discrimination of 16 species occurring in eastern United States out of the total 18 species that are known from the region. Terminology for chaetotaxy and microstructures follows Sawada (1972) and Ashe (1984), and the terminology for abdominal segments follows Thayer (2005). Specimens collected during this study are deposited in the Louisiana State Arthropod Museum (LSAM), Baton Rouge, LA and the Great Smoky Mountains National Park Collection (GSMNP), Gatlinburg, TN, U.S.A. Depositories are indicated parenthetically. Holotypes of species described herein are deposited in the Field Museum of Natural History (FMNH), Chicago.

Genus Leptusa Kraatz, 1856

Leptusa Kraatz, 1856: 60; Fenyes, 1920: 120; Bernhauer & Scheerpeltz, 1926: 553; Smetana, 1973: 3; Lohse, 1974: 42; Moore & Legner, 1975: 483; Seevers, 1978: 164; Pace, 1989: 25; Ashe, 2001: 366; Assing, 2002: 971; Gusarov, 2003: 113; Gusarov & Herman, 2003a: 115, 2003b: 191; Klimaszewski et al., 2004: 3; Gouix & Klimaszewski, 2007: 61.

Type species. Bolitochara pulchella Mannerheim, 1830; designated by Gusarov & Herman (2003a). Diagnosis. The genus Leptusa can be recognized by the following combination of characters: body more or less convex, parallel-sided, pubescent; microsculpture present; hind wings often absent; eye size highly variable but usually small; antennae with 11 antennomeres, thickened toward apex; mandible short, right mandible with internal tooth; distal lobe of galea membranous and pubescent; maxillary palpomere 1 smallest, 2 moderately long, 3 largest, 4 subuliform; ligula narrow and long, not bifid, rounded apically; labial palpi with two apparent articles due to fusion of articles 1–2; pronotum usually widest at apical third or anterior half; mesocoxae narrowly separated; mesoventral process longer than metaventral process (approximately 2/3 length of mesocoxae); legs usually short, tarsal formula 4-4-5, first metatarsomere as long as following two articles or slightly longer. Abdominal tergites III–VI with setigerous punctures (Pace 1989, Klimaszewski et al. 2004). Remarks. All species of Leptusa collected in GSMNP have the following combination of characters: gular sutures widely separated, approximately 1/5 width of head; each side of ventral surface of head with carinae (fading before attaining gular suture or attaining gular suture); labrum bearing 3 pairs of short setae along anterior and lateral margin (Figs. 2b–11b); median area of labium distinctly narrow or lateral areas meeting at middle, and a pair of distal setae present in a longitudinal row; mesoventrite carinate or not (if carinate, then not exceeding 1/2 length of mesoventrite). Abdominal sternites VII of males possess many pores occupying anywhere from anterior one-tenth to anterior half.

Key to the Leptusa species from the eastern United States

Note. Leptusa (E.) nanula (Casey, 1893) and L. obscura Blatchley, 1910 are not included in the key. See discussion.

1. Head and pronotum with coarse setigerous punctures, distance between punctures less than their diameters (Figs. 1g, 1j) ...... 2 - Head and pronotum with moderate sized setigerous punctures, distance between punctures more than their diameters (Figs. 1b–1f, 1h, 1i) ...... 3 2. Antennomere 4 subquadrate (Fig. 8a); elytra unicolored (Fig. 1g); abdominal tergites VII–VIII with single median elongate tubercle (Figs. 1g, 8f) ...... L. (Dysleptusa) pusio (Casey) - Antennomere 4 longer than wide (Fig. 11a); posterior part of elytra paler (Fig. 1j); abdominal tergites VII–VIII without single median elongate tubercle (Figs. 1j, 11f) ...... L. (Ulitusa) cribratula (Casey) 3. Elytra not more than 1.3 times longer than pronotum (Figs. 1b, 1c, 1f) ...... 4 - Elytra at least 1.5 times longer than pronotum (Figs. 1a, 1d, 1e, 1g–1j)...... 8 4. Body length less than 2.5 mm; antennomere 4 subquadrate (Fig. 7a); hind wings absent ...... 5 - Body length more than 2.5 mm; antennomere 4 longer than wide (Figs. 3a, 4a); hind wings present ...... 6

2 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. 5. Body light reddish brown (Klimaszewski et al. 2004: Fig. 9); posterior margin of male abdominal tergite VIII without tubercles (Klimaszewski et al. 2004: Fig. 49); flagellum approximately 2 times longer than length of median lobe (Klimaszewski et al. 2004: Fig. 48)...... L. (Adoxopisalia) elegans Blatchley - Body brown (Fig. 1g); posterior margin of male abdominal tergite VIII with tubercle (Fig. 7f); flagellum as long as 1/2 length of median lobe (Fig. 7k) ...... L. (Dysleptusa) gimmeli n. sp. 6. Antennomere 4 transverse; hind wings absent ...... L. (Dysleptusa) smetanaiella Pace - Antennomere 4 longer than wide; hind wings present ...... 7 7. Antennomere 5 longer than wide (Fig. 3a); carina on each side of surface of head incomplete (fading before attaining gular suture); mesoventrite with short carina...... L. (Adoxopisalia) pseudosmokyiensis n. sp. - Antennomere 5 subquadrate (Fig. 4a); carina on each side of ventral surface of head complete (attaining gular suture); mesoventrite without carina...... L. (Adoxopisalia) smokyiensis Pace 8. Head, pronotum and elytra opaque and velutinous...... 9 - Head, pronotum and elytra somewhat glossy ...... 11 9. Antennomere 5 subquadrate (Fig. 5a); male abdominal tergites VII–VIII without single median elongate tubercle (Fig. 5f) ...... L. (Adoxopisalia) stocksi n. sp. - Antennomere 5 longer than wide (Fig. 2a); male abdominal tergites VII–VIII with single median elongate tubercle (Fig. 2f) ...... 10 10. Posterior margin of male abdominal tergite VIII without tubercle (Fig. 2f); flagellum approximately twice length of median lobe (Fig. 2j)...... L. (Adoxopisalia) opaca Casey - Posterior margin of male abdominal tergite VIII with tubercles (Klimaszewski et al. 2004: Fig. 45); flagellum at least 3 times longer than length of median lobe (Klimaszewski et al. 2004: Fig. 44)...... L. (Adoxopisalia) pseudopaca Klimaszewski & Majka 11. Abdominal tergite VI transverse (at least 1.7 times wider than long), posterior margin of abdominal tergite VIII emarginate (Figs. 6f, 6h), male abdominal tergite VIII with single median elongate tubercle (Fig. 6f) ...... 12 - Abdominal tergite VI subquadrate (less than 1.5 times wider than long), abdominal tergite VIII with rounded posterior margin (Figs. 9f, 9h, 10f, 10h), male abdominal tergite VIII without median tubercle (Figs. 9f, 10f )...... 13 12. Antennomere 4 subquadrate, 5 transverse (Fig. 6a); carina on each side of ventral surface of head complete (attaining gular suture); posterior margin of male abdominal tergite VIII without tubercle (Fig. 6f) ...... L. (Dysleptusa) carolinensis Pace - Antennomere 4 longer than wide, 5 subquadrate (Klimaszewski et al. 2004: Fig. 12); carina on each side of ventral surface of head incomplete (fading before attaining gular suture); posterior margin of male abdominal tergite VIII with several tubercles (Klimaszewski et al. 2004: Fig. 22)...... L. (Boreoleptusa) canonica Casey 13. Mesoventrite not carinate; median lobe in dorsal view somewhat thick (Pace 1989: Fig. 639), flagellum longer than 1/2 length of median lobe (Fig. 9k) ...... L. (Eucryptusa) brevicollis Casey - Mesoventrite with short carina, reaching approximately 1/3 length of mesoventrite; median lobe in dorsal view slender (Fig. 10l), flagellum shorter than 1/2 length of median lobe (Fig. 10k) ...... L. (Eucryptusa) ferroi n. sp.

Subgenus Adoxopisalia Pace, 1989

Leptusa (Adoxopisalia Pace, 1989: 130); Klimaszewski et al., 2004: 23; Gouix & Klimaszewski, 2007: 61.

Type species. Leptusa opaca Casey, 1893; designated by Pace (1989). Diagnosis. See Pace (1989) and Klimaszewski et al. (2004).

Leptusa (Adoxopisalia) opaca Casey, 1893 (Figs. 1a, 2a–k, Map 1)

Leptusa opaca Casey, 1893: 364; Fenyes, 1920: 123; Bernhauer & Scheerpeltz, 1926: 555; Pace, 1989: 130; Gusarov, 2003: 118; Klimaszewski et al., 2005: 46; Klimaszewski et al., 2007: 815, 826; Gouix & Klimaszewski, 2007: 61. Leptusa seminitens Casey, 1893: 364; Bernhauer & Scheerpeltz, 1926: 555 (as synonym of L. opaca).

Material examined (n=2). U.S.A: Tennessee: Sevier Co.: %, GSMNP, Twin Creeks, FIT #1, 26 June – 1 July 2001, C. Carlton, V. Moseley, A. Tishechkin (LSAM); North Carolina: Haywood Co.: & (slide mounted), GSMNP, AHSLC @ Purchase Knob, FIT, 17S 312021 3940123, 17 August 2005, T. Grannan (GSMNP).

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 3 FIGURE 1. Habitus of Leptusa. a: Leptusa (Adoxopisalia) opaca Casey; b: L. (A.) pseudosmokyiensis sp. nov.; c: L. (A.) smokyiensis Pace; d: L. (A.) stocksi sp. nov.; e: L. (Dysleptusa) carolinensis Pace; f: L. (D.) gimmeli sp. nov.; g: L. (D.) pusio (Casey); h: L. (Eucryptusa) brevicollis Casey; i: L. (E.) ferroi sp. nov.; j: L. (Ulitusa) cribratula (Casey). Scale bars = 1 mm.

Redescription. Length 3.1–3.2 mm. Head, pronotum, and elytra blackish brown; antennomeres 1–2, 11, mouthparts, tibia, tarsus, and posterior part of abdominal tergites II–V yellowish brown; femora brown (Fig. 1a). Head opaque, pubescent, with microsculpture. Antennomeres 1–3 elongate, 4–5 longer than wide, 6–7 subquadrate, and 8–10 transverse (Fig. 2a). Carina on each side of ventral surface of head complete, attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 2b). Labium with two distal setae in a longitudinal row, and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 2c). Labial palpi bearing 12 setae (a–h, α–δ), γ-, δ-seta at least three times longer than α-, β-seta (Fig. 2d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), and 3 pairs of additional setae; many pores present in middle area (Fig. 2e). Thorax. Pronotum approximately 1.61 times wider than long, pubescent, opaque, with microsculpture. Mesoventrite with distinct reticulate microsculpture. Metaventrite with setigerous punctures. Elytra wider than pronotum and abdomen, approximately 1.62 times longer than wide, approximately 1.95 times longer than pronotum, latero-posterior margin emarginate, opaque, and pubescent. Hind wings present. Abdomen. Abdomen glossy, as wide as elytra. Abdominal tergite VI approximately 1.79 times wider than long; tergites VII–VIII with single median elongate tubercle (Fig. 2f); tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 2f). Male abdominal sternite VIII with posterior margin produced and approximately bearing 15–16 main setae, and

4 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 2. Details of Leptusa (Adoxopisalia) opaca Casey. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: paramere; j: median lobe. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 5 many additional short setae and pores (Fig. 2g). Female abdominal tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 2h). Genitalia. Paramere as in Fig. 2i. Median lobe as in Fig. 2j. Spermatheca as in Fig. 2k. Type locality. Leptusa opaca: Pennsylvania. Distribution. CANADA: New Brunswick, Nova Scotia, Ontario, Prince Edward Island, Quebec; USA: Arkansas, Georgia, New York, North Carolina, Pennsylvania, Rhode Island, Tennessee, Wisconsin. Locations where L. opaca has been collected in GSMNP as in Map 1. Habitat. Both specimens were collected from secondary forests during mid to late summer using flight intercept traps.

Leptusa (Adoxopisalia) pseudosmokyiensis Park & Carlton, new species (Figs. 1b, 3a–i, Map 1)

Type material. Holotype: U.S.A.: Tennessee: %, “USA: TN: Sevier Co. GSMNP Laurel Falls N35°40.808’ 2 April 2007 W83°36.067’ SP1 Litter 2 of 3 -M Gimmel”, “HOLOTYPE, Leptusa pseudosmokyiensis Park and Carlton, des. 2010” (FMNH). Paratypes (n=2): U.S.A.: Tennessee: %, “USA: TN: Sevier Co. GSMNP Porters Creek N35°40.790’ 5 April 2007 W83°23.855’ SP2 Litter 3 of 3 -M Gimmel” (GSMNP); % (slide mounted), “USA: TN: Sevier Co. GSMNP Laurel Falls N35°40.808’ 2 April 2007 W83°36.067’ SP1 Litter 3 of 3 -M Gimmel” (LSAM). Diagnosis. Leptusa (Adoxopisalia) pseudosmokyiensis may be distinguished from L. (A.) smokyiensis by external morphology. We did not compare genitalia because of the absence of available specimens of L. (A.) smokyiensis. The new species differs from L. (A.) smokyiensis in having distinctively longer antennomeres 4– 5 (Fig. 3a), and a short mesoventral carina. Additionally, the mouthparts of L. (A.) pseudosmokyiensis differ from L. (A.) smokyiensis in possessing short (as long as α-, β-, δ-seta) γ-seta on the labial palpi (Fig. 3d), labrum with more setae (Fig. 3b), and an incomplete carina on each side of midline of ventral surface of head, fading before attaining the gular suture. Description of male. Length 2.8–2.9 mm. Body blackish brown; antennomeres 1–3, 11, mouthparts, legs, and tergites II–V brown (Fig. 1b). Head somewhat glossy, pubescent, with microsculpture. Antennomeres 1–3 elongate, 4–5 longer than wide, 6–7 subquadrate, and 8–10 transverse (Fig. 3a). Carina on each side of ventral surface of head incomplete, fading before attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 8 pairs of long setae, a-seta, b-seta, and many pores (Fig. 3b). Labium with two distal setae in a longitudinal row, and two pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 3c). Labial palpi bearing 12 seta (a–h, α–δ) (Fig. 3d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), and many pores (Fig. 3e). Thorax. Pronotum wider than head, and approximately 1.25 times wider than long; somewhat glossy, pubescent, with microsculpture. Mesoventrite with distinct reticulate microsculpture, and incompletely carinate, reaching 1/3 length of mesoventrite. Metaventrite with setigerous punctures. Elytra approximately 1.49 times longer than wide, and approximately 1.25 times longer than pronotum; latero-posterior margin emarginate, somewhat glossy, and pubescent; setigerous punctures present. Hind wings present. Abdomen. Abdomen glossy, wider than elytra, widest at abdominal tergites VI–VII. Abdominal tergites VI approximately 1.83 times wider than long; tergites VII with single median elongate tubercle; tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 3f). Male abdominal sternite VII with many pores in anterior one-tenth; sternite VIII with posterior margin produced, bearing 7 pairs of main setae, and many additional short setae and pores (Fig. 3g). Genitalia. Paramere as in Fig. 3h. Median lobe as in Fig. 3i. Female. Unknown. Distribution. USA: Tennessee. Locations where L. (A.) pseudosmokyiensis has been collected in GSMNP as in Map 1.

6 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 3. Details of Leptusa (Adoxopisalia) pseudosmokyiensis sp. nov. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: paramere; i: median lobe. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 7 Etymology. The specific epithet refers to the superficial similarity of this species to L. (A.) smokyiensis. Habitat. All three specimens of this species were collected by sifting leaf litter in old growth forests. No specimens were collected by sifting and emergence trapping from CWD (decay classes I–V) at the same times and locations.

Leptusa (Adoxopisalia) smokyiensis Pace, 1989 (Figs. 1c, 4a–h, Map 1)

Leptusa (Adoxopisalia) smokyiensis Pace, 1989: 252; Klimaszewski et al., 2004: 9.

Type material examined. Holotype. U.S.A.: North Carolina: &, glued on rectangular card, spermatheca dissected and mounted in balsam on a clear plastic card, “NC Gr. Sm. Mts. N. P. Clingmans Dome 1950–2020 m 2.VI.86 A. Smetana” [white label, printed]; “HOLOTYPUS Leptusa smokyiensis. Det. R. PACE 1982” [red label, handwritten]; “Leptusa smokyiensis sp. n. det. R. Pace 1987” [white label, handwritten]; “HOLOTYPE CNC No. 20977” [red label, handwritten]. Deposited in Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa. Additional material examined (n=2). U.S.A.: Tennessee: Blount Co.: &, GSMNP, middle Prong Tr at 3944389 259358, 2200`, Berlese, 17 June 2006, C. Ware, C. Shapico, J. Hudson (LSAM); Sevier Co.: & (slide mounted), GSMNP, Porters Creek N35°40.790’ W83°23.855’, SP2 Litter 3 of 3, 5 October 2006, M Gimmel (GSMNP). Redescription of female. Length 2.6–2.8 mm. Head and pronotum blackish brown; elytra and abdomen brown, abdominal tergites III–V somewhat brighter; mouthparts, legs, and antennomeres 1–3, 11 yellowish brown (Fig. 1c). Head somewhat glossy, pubescent, with microsculpture. Antennomeres 1–3 elongate, 4 longer than wide, 4 subquadrate, 6–7 subquadrate, and 8–10 transverse (Fig. 4a). Carina on each side of ventral surface of head complete, attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 4b). Labium with two distal setae in a longitudinal row, and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 4c). Labial palpi bearing 12 setae (a–h, α–δ), γ-seta close to f-seta (Fig. 4d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), a pair of additional setae, and many pores (Fig. 4e). Thorax. Pronotum wider than head, and approximately 1.23 times wider than long; somewhat glossy, with microsculpture. Mesoventrite with distinct reticulate microsculpture. Metaventrite with setigerous punctures. Elytra as wide as pronotum, approximately 1.55 times longer than wide, and approximately 1.2 times longer than pronotum; latero-posterior margin emarginate, somewhat glossy, and pubescent; setigerous punctures present. Hind wings present. Abdomen. Abdomen glossy, wider than elytra, widest at abdominal tergites V–VI. Female abdominal tergites VI approximately 1.62 times wider than long; tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 4f). Female abdominal sternite VIII bearing 5 pairs of main setae, and many additional short setae and pores (Fig. 4g). Genitalia. Spermatheca as in Fig. 4h. Male. Unknown. Type locality. North Carolina (GSMNP). Distribution. USA: North Carolina, Tennessee. Locations where L. (A.) smokyiensis has been collected in GSMNP as in Map 1. Habitat. Both specimens examined were collected from forest habitat by sifting leaf litter in mid to late summer. Comments. Pace (1989) mentioned that this species does not have hind wings, but our specimens have fully developed hind wings. We presume that the hind wings of the holotype may be damaged because wingless Leptusa species usually have short and compact elytra, such as L. (D.) gimmeli, described below.

8 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 4. Details of Leptusa (Adoxopisalia) smokyiensis Pace. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: female abdominal tergite VIII; g: female abdominal sternite VIII; h: spermatheca. Scale bars = 0.1 mm.

Holotype. U.S.A.: Tennessee: Sevier co.: %, “TENNESSEE: Sevier Co., GSMNP, Clingmans Dome ATBI Plot. Malaise trap MT 16200110816. 31 Jul–16 Aug 2001. I.C.Stocks”, “HOLOTYPE, Leptusa stocksi Park and Carlton, des. 2010”. Deposited in the Field Museum, Chicago. Paratypes (n=2). U.S.A., Tennessee: 2%% with same data as holotype GSMNP and LSAM (slide mounted specimen missing genitalia). Diagnosis. Leptusa (Adoxopisalia) stocksi has similar appearance with L. (A.) opaca, but it can be distinguished by shorter antennomeres 4–7 (Fig. 5a), shape of median lobe (Fig. 5i), male abdominal tergites VII–VIII without single median elongate tubercle (Fig. 5f), and short δ-seta (as long as α-, β-seta) of labial palpi (Fig. 5d). Description of male. Length 3.0–3.3 mm. Body brown; antennomeres 1–3, 11, mouthparts, legs, and abdominal tergites II–V yellowish brown (Fig. 1c). Head opaque, pubescent, with microsculpture. Antennomeres 1–3 elongate, 4 longer than wide, 5–6 subquadrate, 7–10 transverse (Fig. 5a). Carina on each side of ventral surface of head complete, attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 5b). Labium with two distal setae in a longitudinal row, and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 5c). Labial palpi bearing 12 setae (a–h, α–δ) (Fig. 5d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), a pair of additional setae, and many pores (Fig. 5e). Thorax. Pronotum wider than head, and approximately 1.44 times wider than long; pubescent, opaque, with microsculpture. Mesoventrite with distinct reticulate microsculpture. Metaventrite with setigerous punctures. Elytra opaque, wider than pronotum and abdomen, approximately 1.58 times longer than wide, approximately 1.56 times longer than pronotum, and latero-posterior margin emarginate. Hind wings present. Abdomen. Abdomen somewhat glossy. Abdominal tergite VI approximately 1.99 wider than long. Male abdominal tergite VIII with posterior margin emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 5f). Male abdominal sternite VIII with posterior margin produced, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 5g). Genitalia. Paramere as in Fig. 5h. Median lobe as in Fig. 5i. Female. Unknown. Distribution. USA: Tennessee. Locations where L. (A.) stocksi has been collected in GSMNP as in Map 1. Etymology. This species is named in honor of Ian Stocks (Clemson University, Clemson, SC), collector of the type series. Habitat. This species is only known from Clingmans Dome, the highest point in GSMNP. All specimens were collected using a malaise trap during August.

Subgenus Dysleptusa Pace, 1982

Leptusa (Dysleptusa Pace, 1982: 580); 1989: 106; Klimaszewski et al., 2004: 30; Gouix & Klimaszewski, 2007: 62.

Type species. Oxypoda fuliginosa Aubé, 1850; designated by Pace (1989). Diagnosis. See Pace (1989) and Klimaszewski et al. (2004).

Leptusa (Dysleptusa) carolinensis Pace, 1989 (Figs. 1e, 6a–l, Map 1)

Leptusa (Dysleptusa) carolinensis Pace, 1989: 252; Klimaszewski et al., 2004: 32; Gouix & Klimaszewski, 2007: 62; Webster et at., 2009: 207.

10 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 5. Details of Leptusa (Adoxopisalia) stocksi sp. nov. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: paramere; i: median lobe. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 11 FIGURE 6. Details of Leptusa (Dysleptusa) carolinensis Pace. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe; l: spermatheca. Scale bars = 0.1 mm.

Type material examined. Holotype. U.S.A.: North Carolina: %, glued on rectangular card, both antennae broken (left antennomeres 9–11 missing and right antennomeres 8–11 missing), aedeagus dissected and

12 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. mounted in balsam on a clear plastic card, “NC Haywood Co. Blue Rdg. Pkw. Woodfin Cascade 1400 m 28.V.1986 A. Smetana” [white label, printed], “HOLOTYPUS Leptusa carolinensis det. R. PACE 1987” [red label, handwritten], “Leptusa carolinensis sp. n. det. R. Pace 1987” [white label, handwritten], “HOLOTYPE CNC No. 20978” [red label, handwritten]. Deposited in Canadian National Collection of Insects, Arachnids and Nematodes (CNC), Ottawa. Additional material examined (n=20). U.S.A.: Tennessee: Sevier co.: 2%%, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 4 October 2006–1 April 2007, SN34C -CWD Rear 1, M. Ferro (GSMNP); Blount Co.: %, &, GSMNP, Tremont, N35°37.308’ W83°40.447’, 4 October 2006–1 April 2007, SN23B -CWD Rear 1, M. Ferro (GSMNP); 2&&, GSMNP, Tremont, N35°37.308’ W83°40.447’, 4 October 2006–1 April 2007, SN24B -CWD Rear 1, M. Ferro (GSMNP); Cocke Co.: % (slide mounted), GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 4 October 2006–1 April 2007, SP31C -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 4 October 2006–1 April 2007, SP31B - CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 1 April–28 June 2007, SP34B -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 13 April–18 May 2006, SP31C -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 13 April–18 May 2006, SP33B -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 18 May–24 June 2006, SP34C -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 18 May–24 June 2006, SP34A -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 5 October 2006, SP3 -CWD5 2 of 3, M. Ferro (LSAM); %, GSMNP, ATBI Plot: Albright Grove Pitfall, 87 83 16 50 35 43 60, 19–30 January 2001, Parker, Stocks, Peterson (LSAM); Sevier Co.: %, GSMNP, Laurel Falls, N35°40.808’ W83°36.067’, 10 April–18 May 2006, SP12C -CWD Rear 1, M. Ferro (LSAM); Cocke Co.: &, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 13 April–18 May 2006, SP34A -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Albright Grove, N35°44.173’ W83°16.647’, 13 April–18 May 2006, SP35A -CWD Rear 1, M. Ferro (LSAM); Blount Co.: &, GSMNP, Tremont, N35°37.308’ W83°40.447’, 4 October 2006–1 April 2007, SN34A -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Tremont, N35°37.308’ W83°40.447’, 4 October 2006–1 April 2007, SN35A - CWD Rear 1, M. Ferro (LSAM). Redescription. Length 2.5–2.8 mm. Body blackish brown, and somewhat glossy; posterior area of elytra and abdominal tergites II–IV somewhat brighter; antennomeres 1–3, mouthparts, legs, and abdominal tergite VIII yellowish brown (Fig. 1e). Head pubescent, with microsculpture. Antennomeres 1–3 elongate, 4 subquadrate, 5–10 transverse (Fig. 6a). Carina on each side of ventral surface of head complete, attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 6b). Labium bearing two distal setae in a longitudinal row in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 6c). Labial palpi bearing 12 setae (a–h, α–δ), γ-seta at least three times longer than α-, β-, δ-seta (Fig. 6d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), 3 pairs of additional setae, and many pores (Fig. 6e). Thorax. Pronotum approximately 1.61 times wider than long, pubescent, microsculpture present. Mesoventrite with distinct reticulate microsculpture; incompletely carinate, carina extending approximately 1/ 3 length of mesoventrite. Metaventrite with setigerous punctures. Elytra wider than pronotum, approximately 1.66 times longer than wide, approximately 1.85 times longer than pronotum, latero-posterior margin emarginate, setigerous punctures present. Hind wings present. Abdomen. Abdomen wider than elytra, widest at abdominal tergite VI. Abdominal tergites VI approximately 1.82 times wider than long. Male abdominal tergite VII–VIII with single median elongate tubercle (Fig. 6f); tergite VIII with posterior margin emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 6f). Male abdominal sternite VII with many pores in anterior one-fourth; sternite VIII with posterior margin produced, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 6g). Female abdominal tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 6h). Female abdominal sternite VIII with posterior margin rounded, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 6i).

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 13 Genitalia. Paramere as in Fig. 6j. Median lobe as in Fig. 6k. Spermatheca as in Fig. 6l. Type locality. North Carolina (Haywood County). Distribution. CANADA: New Brunswick, Nova Scotia, Quebec; USA: North Carolina, Tennessee. Locations where L. carolinensis has been collected in GSMNP as in Map 1. Habitat. This species appears to be associated with CWD. One specimen was collected using a pitfall trap and one specimen was sifted from CWD decay class V. All other specimens emerged from CWD collected in both primary and secondary forests: six from decay class I, ten from decay class II, and two from decay class III–IV. No specimens were collected by sifting leaf litter at the same times and locations.

Leptusa (Dysleptusa) gimmeli Park & Carlton, new species (Figs. 1g, 7a–l, Map 1)

Holotype. U.S.A.: Tennessee: %, “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 5 October 2006 W83°16.647’ SP3 Litter 3 of 3 –M Gimmel”, “HOLOTYPE, Leptusa gimmeli Park and Carlton, des. 2010”. Paratypes (n=107). U.S.A.: Tennessee: 13%% (2%% slide mounted), 15&& with same data as holotype (LSAM); 4%%, 10&&, “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 5 October 2006 W83°16.647’ SP3 Litter 1 of 3 –M Gimmel” (LSAM); 11%%, 15&& (3&& slide mounted), “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 5 October 2006 W83°16.647’ SP3 Litter 2 of 3 –M Gimmel” (GSMNP); 6%%, 7&&, “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 31 March 2007 W83°16.647’ SP3 Litter 2 of 3 –M Gimmel”; 8%%, 17&& (& slide mounted), “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 31 March 2007 W83°16.647’ SP3 Litter 3 of 3 –M Gimmel” (LSAM); &, “USA: TN: Cocke Co. GSMNP Albright Grove N 35°44.173’ 31 12 IV – 18 V W83°16.647’ SP34A –CWD Rear 1 –M Ferro” (LSAM). Diagnosis. Leptusa (Dysleptusa) gimmeli differs from L. (D.) carolinensis by its smaller body length, shape of the spermatheca (Fig. 7l), short elytra (apparently as long as wide, Fig. 1f), and absence of hind wings. This species differs from L. (D.) smetanaiella by its shorter body length, color, subquadrate antennomere 4 (Fig. 7a), and short elytra (apparently as long as wide, Fig. 1f). Leptusa (D.) gimmeli has a median lobe shaped similar to that of L. (D.) carolinensis and L. (D.) smetanaiella, but it can be distinguished from those species by the shape of the bulbus: the anterior parts of the median lobe (Fig. 7k). Description. Length 1.9–2.3 mm. Body brown, glossy; head and abdominal tergites V–VI blackish brown; antennomeres 1–3, 11, mouthparts, legs, and abdominal tergite VIII yellowish brown (Fig. 1f). Head pubescent and with microsculpture. Antennomeres 1–3 elongate, 4 subquadrate, 5–10 transverse (Fig. 7a). Carina on each side of ventral surface of head incomplete, fading before attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and pores (Fig. 7b). Labium bearing two distal setae in a longitudinal row in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 7c). Labial palpi bearing 12 setae (a–h, α–δ) (Fig. 7d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w) and many pores (Fig. 7e). Thorax. Pronotum wider than head, approximately 1.44 times wider than long, pubescent and with microsculpture. Mesoventrite with distinct reticulate microsculpture. Elytra as wide as pronotum, approximately 1.2 times longer than wide, approximately 1.06 times longer than pronotum, latero-posterior margin emarginate, and setigerous punctures present. Hind wings absent. Abdomen. Abdomen wider than elytra, widest at abdominal tergites V–VI. Abdominal tergite VI approximately 1.88 times wider than long. Male abdominal tergites VII–VIII with single median elongate tubercle (Fig. 7f); tergite VIII bearing 4 pairs of main setae, and many additional short setae and pores; posterior margin with several tubercles, and shallowly emarginate (Fig. 7f). Male abdominal sternite VII with many pores in anterior one-third; sternite VIII with posterior margin weakly produced, bearing 7 pairs of main setae, and many additional short setae and pores (Fig. 7g). Female abdominal tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 7h). Female abdominal sternite VIII with posterior margin rounded, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 7i).

14 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 7. Details of Leptusa (Dysleptusa) gimmeli sp. nov. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe; l: spermatheca. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 15 Genitalia. Paramere as in Fig. 7j. Median lobe as in Fig. 7k. Spermatheca as in Fig. 7l. Distribution. USA: Tennessee. Locations where L. (D.) gimmeli has been collected in GSMNP as in Map 1. Etymology. This species is named in honor of Matthew L. Gimmel (Louisiana State University, Baton Rouge), collector of all but one specimen in the type series. Habitat. This species is known only from Albright Grove, an area of old growth forest in GSMNP and is represented by more specimens than any other species in the park. Over 100 specimens were collected from sifted leaf litter from both the spring and fall. Despite an extensive survey only a single specimen was collected from CWD in Albright Grove (emergent from decay class II).

Leptusa (Dysleptusa) pusio (Casey, 1906) (Figs. 1g, 8a–l, Map 1)

Ulitusa pusio Casey, 1906: 348. Leptusa pusio: Fenyes, 1920: 123; Bernhauer & Scheerpeltz, 1926: 556; Pace, 1989: 107; Gusarov, 2003: 115; Klimaszewski et al., 2004: 9.

Material examined (n=27). U.S.A.: Tennessee: Sevier Co.: 5%%, 7&& (& slide mounted), GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 5 April 2007, SP2 Litter 2 of 3, M. Ferro (GSMNP); %, 2&& (% slide mounted), GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 12 April–18 May 2006, SP25C -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 5 October 2006, SP2 Litter 1 of 3, M. Gimmel (LSAM); %, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 12 April–18 May 2007, SP23B -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP21B -CWD Rear 1, M. Ferro (LSAM); %, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 16 April–18 May 2006, SN32B -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 5 October 2006, SP2 Litter 2 of 3, M. Gimmel (LSAM); &, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 18 May–24 June 2006, SP24A -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP24A -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 18 May–24 June 2006, SP24C -CWD Rear 1, M. Ferro (LSAM); &, GSMNP, Greenbrier, N35°43.147’ W83°23.349’, 4 October 2006–1 April 2007, SP13A -CWD Rear 1, M. Ferro (LSAM). Redescription. Length 1.9–2.2 mm. Body brown, glossy; head and abdominal tergite VI blackish brown; antennomeres 1–3, mouthparts and legs somewhat brighter (Fig. 1g). Head as wide as pronotum, with large setigerous punctures (distance between punctures more than their diameter) present. Antennomeres 1–3 elongate, 4 subquadrate, 5–10 transverse (Fig. 8a). Carina on each side of ventral surface of head complete, attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and pores (Fig. 8b). Labium bearing two distal setae in a longitudinal row in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 8c). Labial palpi bearing 12 setae (a–h, α–δ), γ-seta at least three times longer than α-, β-, δ-seta (Fig. 8d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), a pair of additional setae, and many pores (Fig. 8e). Thorax. Pronotum approximately 1.32 times wider than long, with large setigerous punctures (distance between punctures more than their diameter) present. Mesoventrite with distinct reticulate microsculpture; incompletely carinate, carina extending approximately 1/3 length of mesoventrite. Metaventrite with large setigerous punctures present. Elytra wider than pronotum, approximately 1.46 times longer than wide, approximately 1.35 times longer than pronotum, latero-posterior margin emarginate, with large setigerous punctures (distance between punctures more than their diameter) present. Hind wings present. Abdomen. Abdomen wider than elytra, widest at abdominal tergites VI–VII. Abdominal tergite VI approximately 2.1 times wider than long. Male abdominal tergites VII–VIII with single median elongate tubercle (Fig. 8f); tergite VIII bearing 4 pairs of main setae, many additional short setae and pores; posterior margin with several tubercles, and emarginate (Fig. 8f). Male abdominal sternite VII with many pores in

16 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. anterior one-fourth; sternite VIII with posterior margin produced, bearing 9 pairs of main setae, and many additional short setae and pores (Fig. 8g). Female abdominal tergite VIII with posterior margin shallowly emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 8h). Female abdominal sternite VIII with posterior margin rounded, bearing 5 pairs of main setae, and many additional short setae and pores (Fig. 8i).

FIGURE 8. Details of Leptusa (Dysleptusa) pusio(Casey). a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe; l: spermatheca. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 17 Genitalia. Paramere as in Fig. 8j. Median lobe as in Fig. 8k. Spermatheca as in Fig. 8l. Type locality. Ohio (Cincinnati). Distribution. USA: Ohio, Tennessee. Locations where L. (D.) pusio has been collected in GSMNP as in Map 1. Habitat. Specimens examined were collected from primary and secondary forest in GSMNP. Specimens were collected by sifting leaf litter and emergent from CWD decay classes I through III–IV. Comments. Our conclusion that these specimens are conspecific with L. pusio is based on comparison with the genitalia, as illustrated by Gusarov (2003: figs 157-160). He designated the lectotype of this species, and the illustrations were drawn using the paralectotype.

Subgenus Eucryptusa Casey, 1906

Eucryptusa Casey, 1906: 345. Leptusa (Eucryptusa): Fenyes, 1920: 123; Pace, 1989: 116; Klimaszewski et al., 2004: 18; Gouix & Klimaszewski, 2007: 63.

Type species. Silusa nanula Casey, 1893; fixed by monotypy. Diagnosis. See Pace (1989) and Klimaszewski et al. (2004).

Leptusa (Eucryptusa) brevicollis Casey, 1893 (Figs. 1h, 9a–l, Map 1)

Leptusa brevicollis Casey, 1893: 363; Fenyes, 1920: 123; Bernhauer & Scheerpeltz, 1926: 554; Moore & Legner, 1975: 483; Pace, 1989: 116; Gusarov, 2003: 116; Klimaszewski et al., 2004: 18; Klimaszewski et al., 2005: 46; Klimaszewski et al., 2007: 814, 826; Gouix & Klimaszewski, 2007: 63. Leptusa laticollis Notman, 1921: 153; Bernhauer & Scheerpeltz, 1926: 554; Moore & Legner, 1975: 483; Pace, 1989: 11; Gusarov, 2003: 116 (as a synonym of L. brevicollis).

Material examined (n=2). U.S.A.: North Carolina: Haywood Co.: %, & (slide mounted), GSMNP, nr. Purchase Gap house, 35°35.3’N 83°4.5’W, flight intercept, 23–27 June 2007, LSAM Team (LSAM). Redescription. Length 2.1–2.9 mm. Body brown, glossy, head somewhat darker; abdominal tergite VII blackish brown; antenna, mouthparts, and legs yellowish brown (Fig. 1h). Head pubescent, with microsculpture. Antennomeres 1–3 elongate, 4 subquadrate, 5–10 transverse (Fig. 9a). Carina on each side of ventral surface of head incomplete, fading before attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 9b). Labium with two distal setae in a longitudinal row, and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 9c). Labial palpi bearing 12 setae (a–h, α–δ), γ-seta close to f-seta (Fig. 9d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), 5 additional setae, and many pores (Fig. 9e). Thorax. Pronotum approximately 1.35 times wider than long; pubescent, with microsculpture. Mesoventrite with distinct reticulate microsculpture. Metaventrite with setigerous punctures. Elytra wider than pronotum and abdomen, approximately 1.72 times longer than wide, approximately 1.63 times longer than pronotum, latero-posterior margin emarginate, pubescent, setigerous punctures present. Hind wings present. Abdomen. Abdominal tergites VI approximately 1.39 times wider than long. Male abdominal tergite VII with single median elongate tubercle; tergite VIII with posterior margin rounded, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 9f). Male abdominal sternite VII with many pores in anterior one-half; sternite VIII with posterior margin rounded, bearing 7 pairs of main setae, and many additional short setae and pores (Fig. 9g). Female abdominal tergite VIII bearing 4 pairs of main setae, and

18 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. many additional short setae and pores (Fig. 9h). Female abdominal sternite VIII with posterior margin rounded, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 9i). Genitalia. Paramere as in Fig. 9j. Median lobe as in Fig. 9k. Spermatheca as in Fig. 9l.

FIGURE 9. Details of Leptusa (Eucryptusa) brevicollis Casey. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe; l: spermatheca. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 19 Type locality. Pennsylvania. Distribution. CANADA: New Brunswick, Nova Scotia, Ontario, Prince Edward Island, Quebec; USA: Massachusetts, New Hampshire, New York, North Carolina, Pennsylvania, Vermont, Virginia. Locations where L. (E.) brevicollis has been collected in GSMNP as in Map 1. Habitat. Specimens examined were collected with a flight intercept trap from secondary forest.

Leptusa (Eucryptusa) ferroi Park & Carlton, new species (Figs. 1i, 10a–m, Map 1)

Type material. Holotype. U.S.A.: Tennessee: %, “USA: TN: Blount Co. GSMNP Tremont N35°37.308’ 15 IV – 18 V 2006 W83°40.447’ SN22B -CWD Rear 1 -M Ferro”, “HOLOTYPE, Leptusa ferroi Park and Carlton, des. 2010”. Paratypes (n=1). U.S.A.: Tennessee: &, “USA: TN: Blount Co. GSMNP middle Prorg Tr at 3944389 259358, 2200’ Berlese 17 June 2006, C. Ware, C. Shapico, J. Hudson” (LSAM). Diagnosis. Leptusa (Eucryptusa) ferroi is similar to L. (E.) brevicollis, but it differs by having a shorter tubus of the median lobe, a distinctively more slender median lobe in dorsal view (Fig. 10l), and short mesoventral carina. Description. Length 2.8–2.9 mm. Body brown, glossy; antennomeres 1–3, legs, and abdominal tergites II–IV somewhat brighter; antennomere 11, mouthparts yellowish brown (Fig. 1i). Head pubescent, with setigerous punctures, and with microsculpture. Antennomeres 1–3 elongate, 4 longer than wide, 5 subquadrate, 6–10 transverse (Fig. 10a). Carina on each side of ventral surface of head incomplete, fading before attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, 7 pairs of long setae, a-seta, b-seta, and many pores (Fig. 10b). Labium with two distal setae in a longitudinal row and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 10c). Labial palpi bearing 12 setae (a–h, α–δ), (Fig. 10d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), 2 pairs of additional setae, and many pores (Fig. 10e). Thorax. Pronotum wider than head, approximately 1.39 times wider than long, pubescent, with setigerous punctures, and with microsculpture. Mesoventrite with distinct reticulate microsculpture. Mesoventrite incompletely carinate, carina extending approximately 1/3 length of mesoventrite. Metaventrite with setigerous punctures. Elytra wider than pronotum and abdomen, approximately 1.65 times longer than wide, approximately 1.52 times longer than pronotum, latero-posterior margin emarginate, pubescent, with setigerous punctures. Hind wings present. Abdomen. Abdominal tergite VI approximately 1.31 times wider than long. Male abdominal tergite VII with single median elongate tubercle; tergite VIII with posterior margin rounded, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 10f), sternite VIII with posterior margin rounded, bearing 7 pairs of main setae, and many additional short setae, and pores (Fig. 10g). Female abdominal tergite VIII bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 10h), sternite VIII with posterior margin rounded, bearing at least 4 pairs of main setae, and many additional short setae and pores (Fig. 10i). Genitalia. Paramere as in Fig. 10j. Median lobe as in Figs. 10k–l. Spermatheca as in Fig. 10m. Distribution. USA: Tennessee. Locations where L. (E.) ferroi has been collected in GSMNP as in Map 1. Etymology. This species is named in honor of Michael L. Ferro (Louisiana State University, Baton Rouge), collector of the holotype. Habitat. The holotype was emergent from CWD decay class II, and the female paratype was collected from sifted leaf litter. Both were from secondary forest in GSMNP.

20 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 10. Details of Leptusa (Eucryptusa) ferroi sp. nov. a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe, lateral aspect; l: median lobe, dorsal aspect; m: spermatheca. Scale bars = 0.1 mm.

Ulitusa Casey, 1906: 347. Leptusa (Ulitusa): Pace, 1989: 116; Klimaszewski et al., 2004: 21; Gouix & Klimaszewski, 2007: 63.

Type species. Ulitusa cribratula Casey, 1906; designated by Casey (1911). Diagnosis. See Pace (1989) and Klimaszewski et al. (2004).

Leptusa (Ulitusa) cribratula (Casey, 1906) (Figs. 1j, 11a–l, Map 1)

Ulitusa cribratula Casey, 1906: 347. Leptusa cribratula: Fenyes, 1920: 123; Bernhauer & Scheerpeltz, 1926: 556; Seevers, 1978: 284; Moore & Legner, 1975: 483; Pace, 1989:116; Klimaszewski et al., 2004: 22; Gouix & Klimaszewski, 2007: 63.

Material examined (n=25) (most not sexed). U.S.A: Tennessee: Sevier Co.: 4, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 2 April 2007, SN3 CWD5 1 of 3, M. Ferro (GSMNP); 1, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 2 April 2007, SN3 CWD5 3 of 3, M. Ferro (LSAM); 2 (& slide mounted), GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 6 October 2006, SN3 CWD5 3 of 3, M. Ferro (LSAM); 1, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 28 June–10 October 2007, SN31C -CWD Rear 1, M. Ferro (LSAM); 1, GSMNP, Sugarlands Quiet Walkway, N35°39.826’ W83°31.509’, 4 October 2006–1 April 2007, SN34C -CWD Rear 1, M. Ferro (LSAM); 2, GSMNP, Greenbrier, N35°43.147’ W83°23.349’, 18 May–24 June 2006, SP15B -CWD Rear 1, M. Ferro (LSAM); 1, GSMNP, Greenbrier, N35°43.147’ W83°23.349’, 4 October 2006–1 April 2007, SP13A -CWD Rear 1, M. Ferro (LSAM); & (slide mounted), GSMNP, Greenbrier, N35°43.147’ W83°23.349’, 4 October 2006–1 April 2007, SP14B -CWD Rear 1, M. Ferro (LSAM); 3, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP21C -CWD Rear 1, M. Ferro (GSMNP); 1, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP25B -CWD Rear 1, M. Ferro (LSAM); 1, GSMNP Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP24C -CWD Rear 1, M. Ferro (LSAM); 1, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP24A -CWD Rear 1, M. Ferro (LSAM); 2, GSMNP, Porters Creek, N35°40.790’ W83°23.855’, 4 October 2006–1 April 2007, SP23B -CWD Rear 1, M. Ferro (LSAM); Blount Co.: 1, GSMNP, Tremont, N35°37.308’ W83°40.447’, 4 October 2006, SN2 CWD5 2 of 3, M. Ferro (LSAM); 1, GSMNP, Tremont, N35°37.308’ W83°40.447’, 24 June–14 July 2006, SN25C -CWD Rear 1, M. Ferro (LSAM); 1, GSMNP, Goldmine trail, 35°38.454’N 83°54.484’W, 18 March 2008, under pine bark, M. Gimmel (LSAM); 1, GSMNP, Gregory’s Cave trail, 35°36.560’N 83°48.407’W, 17 March 2008, under pine bark, M. Gimmel (LSAM). Redescription. Length 2.1–3.0 mm. Body brown, glossy; abdominal tergites V–VI blackish brown; antennomeres 1–3, 11, mouthparts, legs, and posterior part of elytra yellowish brown (Fig. 1j). Head pubescent, with microsculpture and large punctures (distance between punctures more than their diameter). Antennomeres 1–3 elongate, 4 longer than wide, 5–10 transverse (Fig. 11a). Carina on each side of ventral surface of head incomplete, fading before attaining gular suture. Mouthparts. Labrum transverse, bearing 3 pairs of short setae, and 8 pairs of long setae, a-seta, b-seta, and many pores (Fig. 11b). Labium with two distal setae in a longitudinal row, and several pores in median area; a pair of setal pores, 2 pairs of real pores and several pseudopores present in lateral area (Fig. 11c). Labial palpi bearing 12 setae (a–h, α–δ), γ-seta close to f-seta (Fig. 11d). Mentum trapezoidal, bearing 4 pairs of main setae (b, u, v, w), a pair of additional setae, and many pores (Fig. 11e). Thorax. Pronotum approximately 1.25 times wider than long; pubescent, with microsculpture and large punctures (distance between punctures more than their diameter). Mesoventrite with distinct reticulate microsculpture. Mesoventrite incompletely carinate, carina extending approximately 1/3 to 1/2 length of mesoventrite. Metaventrite with setigerous punctures. Elytra wider than pronotum and abdomen;

22 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. FIGURE 11. Details of Leptusa (Ulitusa) cribratula (Casey). a: antenna; b: labrum; c: labium; d: labial palpus; e: mentum; f: male abdominal tergite VIII; g: male abdominal sternite VIII; h: female abdominal tergite VIII; i: female abdominal sternite VIII; j: paramere; k: median lobe; l: spermatheca. Scale bars = 0.1 mm.

LEPTUSA OF GREAT SMOKY MOUNTAINS NATIONAL PARK Zootaxa 2662 © 2010 Magnolia Press · 23 approximately 1.54 times longer than wide; approximately 1.46 times longer than pronotum, latero-posterior margin emarginate, pubescent, with large setigerous punctures (distance between punctures more than their diameter). Hind wings present. Abdomen. Abdominal tergite VI approximately 1.87 times wider than long. Male abdominal tergite VIII with posterior margin emarginate, bearing 4 pairs of main setae, and many additional short setae and pores (Fig. 11f). Male abdominal sternite VII with many pores in anterior one-fourth; sternite VIII with posterior margin rounded, bearing 6 pairs of main setae, and many additional short setae and pores (Fig. 11g). Female abdominal tergite VIII with posterior margin emarginate, bearing 4 pairs of main setae, many additional short setae and pores (Fig. 11h), sternite VIII with posterior margin rounded, bearing 5 pairs of main setae, and many additional short setae and pores (Fig. 11i). Genitalia. Paramere as in Fig. 11j. Median lobe as in Fig. 11k. Spermatheca as in Fig. 11l. Type locality. Ohio (Cincinnati). Distribution. CANADA: Ontario, Quebec; USA: Alabama, Florida, Ohio, Pennsylvania, Tennessee, Virginia. Locations where L. (U.) cribratula has been collected in GSMNP as in Map 1. Habitat. This species appears to be associated with CWD. Two specimens were collected from under pine bark. All other specimens were emergent or sifted from CWD collected in both primary and secondary forests: seven from decay class I, four from decay class II, four from decay class III–IV, and eight from decay class V. No specimens were collected by sifting leaf litter at the same times and locations.

MAP 1. Locations where Leptusa specimens have been collected in GSMNP. Leptusa opaca Casey: 8, 12; L. pseudosmokyiensis: 5, 9; L. smokyiensis: 4, 6, 9; L. stocksi: 6; L. carolinensis: 3, 5, 7, 11; L. gimmeli: 11; L. pusio: 7, 9, 10; L. brevicollis: 12; L. ferroi: 3, 4; L. cribratula: 1, 2, 3, 7, 9, 10.

Discussion

The four species described herein bring the total number of species of Leptusa known from eastern United States to 18. Of these, we were able to include 16 in the key based on examination of types, authoritatively identified specimens, or diagnostic characters provided in Klimaszewski et al. (2004). We have not seen specimens of L. (E.) nanula (Casey, 1893), known from Rhode Island and Iowa or L. obscura Blatchley, 1910, known from Indiana. Despite an enormous amount of collecting effort within GSMNP since 2001 (Carlton & Bayless 2007), six of the ten Leptusa species were represented by less than five specimens and eight of the ten species were collected from three or fewer locations. Local abundance and habitat specificity of some species (i.e. L.

24 · Zootaxa 2662 © 2010 Magnolia Press PARK ET AL. gimmeli, L. carolinensis) may account for their true or perceived rarity. Very few specimens were collected with passive collection techniques (pitfalls and flight intercept traps). The majority of specimens were collected from leaf litter using Berlese funnels or from CWD using emergence chambers. Species’ actual ranges will be difficult to resolve without active collecting throughout the park. At least L. gimmeli appears to be highly restricted in distribution to a small area of old growth cove forest called Albright Grove in the northeastern area of GSMNP. Habitat preferences can be proposed for a few species that are represented by sufficient sample sizes. Leptusa carolinensis and L. cribratula appear to be strongly associated with CWD habitats (45 specimens from CWD; 3 specimens from other habitats). Leptusa gimmeli appears strongly associated with non-CWD forest litter (106 specimens from sifted litter; one from other habitats). The discovery of undescribed beetle species in eastern North America, where the fauna is presumably well known, illustrates a larger problem in taxonomy and systematics that we have coined “Overlooked Syndrome” (OS) (Ferro & Carlton 2010). This syndrome presents when undescribed species across an otherwise familiar landscape persist because researchers are ignorant of their existence or are otherwise impotent to rectify the issue. This situation is not likely to come as a surprise to specialists of poorly studied insect taxa. But, it is not fully appreciated by many ecologists and conservation biologists and, thus, has not been adequately dealt with in studies of community ecology that include arthropod diversity components. Taxa with the following characteristics may be more likely to suffer from OS: 1) many species within one genus, 2) physically small, 3) slight morphological differences or diagnostic morphological differences internal, 4) obscure habits, 5) of little or no economic value, and 6) not popular among amateur collectors. Leptusa is a poster child of OS and may serve as a guide for specialists in search of easy prey for marketing species names and other forms of taxonomic exploitation. Overlooked Syndrome is easily overcome where taxonomic specialists are available, i.e. C. P. Alexander and Tipulidae (Byers 1982). However, because of the characteristics listed above, researchers are often reluctant or unable to work on the group, due in part to a lack of monetary resources and/or non-existence of a satisfactory reward system. The results are artificially anemic estimates of total diversity in the region and lack of credibility of ecological research involving OS taxa at any functional or analytical levels.

Acknowledgements

We thank Aleš Smetana and Anthony E. Davies (Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa) for the loan of type specimens. We also thank Jan Klimaszewski (Canadian Forest Service, Québec), Volker Assing (Hannover), and Matthew Gimmel (Louisiana State University, Baton Rouge) for helpful comments. This publication was approved by the Director, Louisiana Agricultural Experiment Station as manuscript number 2010-234-4168. This project is funded in part by National Science Foundation NSF DEB-0516311 to Christopher Carlton and Victoria Bayless.

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