Lasioglossum (Hemihalictus) inconditum (Cockerell 1916)

Gibbs, Jason, Packer, Laurence, Dumesh, Sheila & Danforth, Bryan N., 2013, Revision and reclassification of <i> Lasioglossum </ i> (<i> Evylaeus </ i>), <i> L. </ i> (<i> Hemihalictus </ i>) and <i> L. </ i> (<i> Sphecodogastra </ i>) in eastern North America (Hymenoptera: Apoidea: Halictidae), Zootaxa 3672 (1), pp. 1-116 : 31-35

Lasioglossum (Hemihalictus) inconditum (Cockerell 1916)

( Figs. 4D , 5D , 6D , 7D , 8D , 21 , 22 , 76A , 82C , 84C )

Halictus inconditus Cockerell 1916b, p. 101 .

Holotype. ♀ USA, Colorado, Tolland , vii.1915 on Frasera (L. A. Kenoyer) [ NMNH: 27994]. Examined by JG 2012.

(Labels read “Tolland, Colo. [illegible, 10?] July ’15 on Frasera L. A. Kenoyer [handwritten on folded label]/ Type No. 27794 U.S. N.M. [numbers handwritten on red label]/ Halictus inconditus Ckll. TYPE. [handwritten]).

Halictus tracyi Cockerell 1936, p. 161 . [new synonymy]

Holotype. ♀ USA, California, Humboldt Co., Eureka 17.iv.1935 (T. D. A. Cockerell) [ CAS: 04277]. Examined by JG 2012. (Labels read “ Eureka Cal Humboldt Co. IV-17-35/ Pres by TDA Cockerell / Halictus tracyi Ckll TYPE. [handwritten]/ California Academy of Sciences Type No. 4277 [number handwritten]”).

Evylaeus inconditus (in Hurd 1979: catalogue; Moure & Hurd 1987: catalogue)

Evylaeus rufitarsis (in Mitchell 1960: redescription, key ♂; Hurd 1979: catalogue; Moure & Hurd 1987: catalogue)

Evylaeus tracyi (in Hurd 1979: catalogue; Moure & Hurd 1987: catalogue)

Lasioglossum (Evylaeus) inconditum (in Michener 1951: catalogue)

Lasioglossum (Evylaeus) rufitarse (in Michener 1951: catalogue; Krombein 1967: catalogue)

Lasioglossum (Evylaeus) tracyi (in Michener 1951: catalogue)

Diagnosis. Both sexes of L. inconditum can be recognised by the combination of head long (L/W ratio = 1.03– 1.11) ( Figs. 4D , 6D ); mesepisternum finely rugulose ( Fig. 76A ); metapostnotal rugae reaching the sharply edged posterior margin of dorsal surface ( Fig. 82C ); and lateral propodeal carina not extending to dorsal surface. Lasioglossum inconditum can be easily distinguished from most species, except L. foxii . Both sexes of L. foxii have metapostnotal rugae not reaching the smoothly rounded posterior margin ( Fig. 82B ).

Lasioglossum in the fulvicorne-fratellum species-group ( Ebmer 1995, 2002; Pesenko 2007a) are superficially similar to L. inconditum but can usually be distinguished by possession of complete lateral propodeal carinae ( Figs. 79 D-79F), except some L. boreal e ( Fig. 79C ). Males of the fulvicorne-fratellum species-group can be distinguished by yellow metabasitarsi ( Figs. 54 , 57 , 66 , and 69), shorter gonostyli, and distinctively shaped retrorse lobes ( Fig. 87 ; see diagnoses below for L. boreale , L. comagenense , L. quebecense , and L. seillean ).

Redescription. FEMALE. Length 5.9–6.7 mm. Head length 1.69–1.80 mm. Head width 1.60–1.70 mm. Wing length 4.6–5.1 mm. (n=5)

Colour. Head and mesosoma black. Antenna black, except ventral surface of flagellum reddish brown. Tegula dark reddish brown. Legs dark brown, except medio- and distitarsi reddish brown. Wing membrane hyaline, faintly dusky. Pterostigma amber. Metasomal terga black-brown, apical margins pale brown.

Structure. Head long (L/W ratio = 1.03–1.06). Clypeus ¾ below suborbital line. Eyes convergent below (UOD:LOD = 1.14–1.22). Gena narrower than eye. Ocelli normal. Pronotum smoothly rounded. Protibial spur with serrations as long as width of malus. Inner metatibial spur pectinate, teeth 4–8, basal teeth subequal to width of rachis. Propodeal lateral carina not reaching dorsal margin, oblique carina absent.

Surface sculpture. Supraclypeal area polished medially, punctures sparse (i=1–2d). Gena lineolate, postgena polished-weakly imbricate. Mesoscutum imbricate, polished posteromedially; punctures dense laterally (i≤d), separated between parapsidal lines (i=1–1.5d). Mesepisternum finely rugulose, imbricate ventrally. Metapostnotum anastomosingly rugose throughout, posterior margin sharply angled. Propodeum lateral surface weakly ruguloseimbricate, posterior surface tessellate. Metasomal terga polished-weakly coriarious; punctures fine, dense basally, obscure apically.

Pubescence. Head and mesosoma with sparse plumose hairs. Metafemoral scopa with dense plumose hairs. Propodeum with sparse plumose hairs. Metasomal terga with apical fimbriae very sparse. T1 with sparse, erect plumose hairs. Metasomal sterna with plumose scopa.

MALE. Length 5.8–6.4 mm. Head length 1.72–1.74 mm. Head width 1.55–1.58 mm. Wing length 4.2–4.3 mm. (n=3)

Colour. Head and mesosoma black. Clypeus yellow distally. Mandible brown. Labrum brown. Antenna black, except ventral surface of flagellum orange. Legs dark brown, except tarsi reddish brown. Wing membrane hyaline. Pterostigma pale brown. Metasomal terga dark brown, apical margin reddish brown.

Structure. Head long (L/W ratio = 1.10–1.11). Mandible short, reaching opposing clypeal angle. Flagellomeres, except F1 elongate, F2 longer than F1 and pedicel combined, subequal to scape. Eyes strongly convergent below (UOD:LOD = 1.43–1.50). Gena narrower than eye. Pronotum smoothly rounded. Propodeal lateral carina reaching less than half distance to dorsolateral slope.

Surface sculpture. Supraclypeal area polished medially, punctures relatively dense (i≤d). Gena lineolate, postgena polished distally. Mesoscutum weakly imbricate, polished posteromedially; punctures dense laterally (i≤d), distinctly separated between parapsidal lines (i=1–2d). Mesepisternum weakly rugulose, imbricate ventrally. Metapostnotum anastomosingly rugose, posterior margin sharply angled. Propodeum weakly rugulose laterally, posterior surface rugulose. Metasomal terga polished; punctures deep, close basally. T1–T6 apical impressed areas impunctate.

Pubescence. Head and mesosoma with sparse plumose hairs. Paraocular area below eye emargination with dense tomentum. Propodeum largely bare, with scattered plumose hairs. Metasomal terga nearly bare, apical fimbriae absent. Metasomal sterna with sparse, plumose hairs (1.5–2.5 OD).

Terminalia . As shown in Figs. 7D , 8D . Gonostylus long and slender, with short setae. Retrorse lobe long, narrow, attenuated apically.

Taxonomic notes. Lasioglossum rufitarse ( Zetterstedt 1838) is widely considered to be a Holarctic species ( Michener 1951; Mitchell 1960; Ebmer & Sakagami 1985; Moure & Hurd 1987). Previous study of Nearctic and Palaearctic populations suggested there were no morphological differences between them ( Ebmer & Sakagami 1985). We examined a number of L. rufitarse females from Europe and found the medial punctation of T1 and T2 to be sparser than in Nearctic specimens, but this character is extremely subtle and needs to be tested with additional specimens. DNA barcodes from Palaearctic and Nearctic individuals are too divergent for them to likely be conspecific (6.7 % genetic distance) and are consistent with an absence of gene flow between Nearctic and Palaearctic populations. We therefore consider the Nearctic representatives of L. “ rufitarse ” to be a species distinct from the Palaearctic populations.

The oldest available name for North American L. “ rufitarse ” is L. inconditum . DNA barcodes of L. inconditum throughout its range, including localities relatively close to the type localities of Halictus inconditus and H. tracyi , do not show variation consistent with multiple species. Morphological examination revealed that L. inconditum is conspecific with Halictus tracyi . In Cockerell’s (1936) original description of Halictus tracyi he separated it from L. inconditum based on the inner metatibial spur, claiming the latter taxon has fewer teeth. Examination of the types revealed this to be true. However, tooth number on the spur seems to vary continuously so we consider this difference to be insufficient for recognising both species.

Biology. Lasioglossum inconditum is a polylectic ( Moure & Hurd 1987), ground-nesting species ( Cockerell 1936). No published information is available for the social level of L. inconditum , but one of us (LP) excavated a nest of L. inconditum in Newfoundland; the nest was shallow, with the burrow being almost parallel to the soil surface, and contained a single female. Its close relative in Europe, L. rufitarse , is solitary ( Knerer 1968). The range of L. inconditum is mostly limited to alpine habitats or northern regions with short flight seasons that may preclude the formation of annual eusocial colonies.

DNA barcodes. Twenty-nine individuals were sequenced (maximum intraspecific p-distance: 1.7%). Comparison of DNA barcode consensus sequences from L. rufitarse (n=3, specimens from Germany and Russia) and L. inconditum (n=29) revealed 35 fixed nucleotide substitutions (i. e., 6.7% genetic distance). Such a high level of genetic divergence strongly suggests a lack of gene flow between populations. In comparison, there are 17 variable sites within the North American population, which includes specimens collected from geographically distant sites, such as Arizona and Newfoundland. Additional study, particularly of eastern Palaearctic populations would be beneficial. It is conceivable that L. rufitarse and L. inconditum could both occur in parts of Alaska and Siberia so study in both these regions would be of interest. Lasioglossum inconditum differs from other North American nitidiusculum group species by 32 fixed substitutions: 79(T), 81(A), 117(C), 120(C), 213(C), 246(C), 276(T), 309(T), 315(A), 345(C), 354(C), 375(C), 384(C), 393(T), 396(C), 399(C), 402(A), 405(C), 423(A), 432(A), 456(C), 465(G), 474(T), 486(C), 535(C), 537(T), 546(A), 553(A), 591(A), 600(T), 603(T), and 645(C) (see Table 2). Two fixed substitutions distinguish L. inconditum from all other species included herein: positions 456(C) and 465(G).

Range. Fig. 23 .

Material examined. 645 (524 females, 121 males). Deposited in ACNS, AMNH, CAS, CNC, CUIC, EMEC, MCZ, MSUC, PCYU, PMAE, PMNH , ROMent, and UCMC.

CANADA. ALBERTA: Division 1: Elkwater Lake; Onefour; Division 11: Edmonton; Division 15: Banff; Division 16: McMurray; BRITISH COLUMBIA: Central Kootenay Dist.: Howser; Columbia-Shuswap Dist.: Revelstoke, Selkirk Mts.; Cowichan Valley Reg. Dist: Upper Carmanah Valley; East Kootenay: Moyie; Kitimat- Stikine Dist.: Terrace; Terrace, 18 mi. SW; Terrace, 32 mi. SW; Terrace, 6 mi. W, Gagnon Rd.; Terrace, Spring Creek; Lakelse, nr. Terrace; Lakelse Lake bog, S. of Terrace; Kleanza Creek, 14 mi. E Terrace; Kootenay Boundary Dist.: Rock Creek; Northern Rockies Dist.: Summit Lake; Okanagan-Similkameen Dist.: Oliver, White Lake ; Oliver, Orofino Mt.; Osoyoos, Richter Pass; Stikine Dist.: Atlin; MANITOBA: Division 7: Aweme; Division 13: Balmoral; Division 21: Cedar Lake; Division 23: Pearson Is.; NEW BRUNSWICK: Northumberland Co.: Trout Brook; Queens Co.: North Minto; Saint John Co.: St. Martins; York Sunbury Co.: Fredericton; NEWFOUNDLAND: Goose Bay; Harmond Field; Nicholsville; NORTHWEST TERRITORIES: Norman Wells; Tulita (Fort Norman), MacKenzie R.; Yellowknife; NOVA SCOTIA: Cape Breton, Highlands N.P., French Mt.; Colchester Co.: Truro; Digby Co.: Smiths Cove; Richmond Co.: Cape Breton; Victoria Co. : Baddeck; NUNAVUT: Resolution Is.; ONTARIO: Cochrane Dist.: Hearst; Iroquois Falls; Haliburton Co.: Algonquin P.P., by Crossbar Lk.; Algonquin P.P., by Madawaska Lk.; Kenora Dist.: Red Lake; Muskoka Reg.: Dorset; National Capital Reg: Ottawa; Nipissing Dist.: Timagami; Peel Reg.: Forks of Credit; Rainy River Dist.: Fort Frances; Sudbury Dist.: Penage Lake; Thunder Bay Dist.: Upsala; Timiskaming Dist.: Kenabeek; Kirkland Lk.; Swastika; PRINCE EDWARD ISLAND: Kings Co.: Albion Cross; Saint Georges; Woodville Mills; Prince Co.: Arlington; Grand River; Queens Co.: Belle River; Fredricton; Glenroy; Winsloe; QUEBEC: Abitibi-Témiscamingue Reg.: Laniel; Jamésie Reg.: Mistassini Post; Minganie Reg.: Anticosti Is., Ellis Bay; YUKON TERRITORY: Rampart House; Ross River; Shag.

USA. ALASKA: Fairbanks North Star Bor.: Fairbanks; Matanuska-Susitna Bor.: Palmer; CALIFORNIA: Alpine Co.: Winnemucca Lake, nr. Carson Pass; Fresno Co.: Frypan Meadow, 1 mi. E; Pioneer Basin; Humboldt Co.: Eureka; Marin Co.: Inverness; Mill Valley; Mono Co.: Duck Lake; Sonora Pass; Santa Cruz Co. : Felton; Siskiyou Co.: Devil’s Punchbowl; COLORADO: Boulder Co.: Boulder; Nederland; Pinecliff; Ward, 2 mi. E; Clear Creek Co.: Chicago Creek; Mt. Evans, Doolittle Ranch; Mt. Evans, Echo Lake; Gilpin Co.: Lump Gulch; Grand Co.: Kremmling, 2 mi. S; Gunnison Co.: Brush Creek; Gothic; Rocky Mtn. Biol. Lab, Gothic; Mt. Avery, Gunnison N.F.; Mt. Bellview, base, Gunnison N.F.; Washington Gulch; Larimer Co.: Red Feather Lakes, 2 mi. SW; Park Co.: Fairplay; Pennsylvania Mt., nr. Fairplay; Routt Co.: Clark, 5 mi. E; Steamboat Springs; Steamboat Springs, 5 mi. E; San Juan Co.: Silverton; Summit Co.: Dillon, 5 mi. SE; Teller Co.: Pikes Peak, nr. Crystal Creek Dam; CONNECTICUT: Litchfield Co.: Colebrook; IDAHO: Bear Lake Co.: Bloomington Lake; Emigration Canyon; Ovid, 5 mi. W; Blaine Co.: Oregon Gulch; Custer Co.: Sawtooth Lake; Franklin Co.: Willow Flat; MAINE: Hancock Co.: Acadia N.P.; Knox Co.: Glenmere; Piscataquis Co.; Washington Co.: Moosehead Wildlife Pres., SE of Calais; MASSACHUSETTS: Berkshire Co.: Mt. Greylock; Worcester Co.: Petersham; MICHIGAN: Alger Co.: Beaver Lk.; Pictured Rocks N.L.; Chippewa Co.: Drummond Is., nr. Maxton; Dickinson Co.: Channing; Sagola; Iron Co. ; Marquette Co.: Huron Mountain Club; Schoolcraft Co.: Seney; MINNESOTA: Clearwater Co.: Lake Itasca; MONTANA: Gallatin Co.; NEVADA: Elko Co.: Island Lake, Ruby Mtns.; NEW HAMPSHIRE: Belknap Co.: Meredith; Coos Co.: Gorham; Randolph; NEW YORK: Albany Co.: Huyck Preserve, nr. Rensselaerville; Cattaraugus Co.: Rock City; Clinton Co.: Black Brook; Delaware Co.; Essex Co.: Newcomb; Newcomb to Goodnow Mt. peak; Franklin Co.: Upper Saranak Lake, Adirondack Mts. Pk.; Genesee Co.; Hamilton Co.: 8 th Lake, 3 mi. E of Inlet; Lewis Co.: Tug Hill; Onondaga Co. ; Otsego Co.: East Worcester; Schuyler Co.: Hector Land Use Area, W of Reynoldsville; Texas Hollow; Watkins Glen S.P.; Tompkins Co.: Dawes Hill, 2 mi. SSW of West Danby; Dryden; Ellis Hollow, 6 mi. E of Ithaca; Enfield, Teeter Rd.; Ithaca, Bool’s Backwater; Ithaca, Carter Creek Rd.; Ithaca, Cornell U.; Ithaca, Monkey Run; Ithaca, Six-Mile Creek; Michigan Hollow, 5 mi. S of Danby; Mineak Rd., 7 mi. E of Ithaca; Slatervillle, wildflower pres.; Yellow Barn Hill, 7 mi. ENE of Ithaca; OREGON: Klamath Co.: Crater Lake; Merrit Cr., 1 mi. E; Lincoln Co.: Lincoln, 6 mi. W; UTAH: Box Elder Co. : Willard Peak; Cache Co.: Blacksmith Fork; Franklin Basin; Hodges Canyon, W.; Logan Canyon; Mantua, 10 mi. S; Tony Grove Jct.; Tony Grove Lake; Rich Co.: Allen Canyon; Monte Cristo; San Juan Co.: Geyser Pass, LaSal Mtns.; Uintah Co.: Vernal, 8 mi. NW; Weber Co.: Willard Bas., NE; VERMONT: Addison Co.; Bennington Co.; WASHINGTON: Island Co.: Whidbey Is.; King Co.: Seattle; Thurston Co.: Olympia; Whitman Co.: Pullman; WYOMING: Albany Co.: Tie Siding, 4 mi. SW; Park Co.: Cody, Sunlight Basin.

Figures

FIGURE 4. Faces of female Lasioglossum subgenus Hemihalictus in frontal view. (A) L. birkmanni. (B) L. fedorense. (C) L. foxii. (D) L. inconditum. (E) L. lustrans. (F) L. macoupinense. (G) L. nelumbonis. (H) L. pectinatum. (I) L. pectorale. (J) L. sopinci. (K) L. swenki. Scale bar = 1 mm.

FIGURE 5. Drawings of inner metatibial spurs of female Lasioglossum subgenera Hemihalictus (A-K), Evylaeus (L), and Sphecodogastra (M-Q). (A) L. birkmanni. (B) L. fedorense. (C) L. foxii. (D) L. inconditum. (E) L. lustrans. (F) L. macoupinense. (G) L. nelumbonis. (H) L. pectinatum. (I) L. pectorale. (J) L. sopinci. (K) L. swenki. (L) L. cinctipes. (M) L. boreale. (N) L. comagenense. (O) L. quebecense. (P) L. seillean. (Q) L. truncatum. See McGinley (2003) figures 61, 62, and 65 for images of inner metatibial spurs of L. lusorium, L. oenotherae, and L. texanum.

FIGURE 6. Faces of male Lasioglossum subgenus Hemihalictus in frontal view. (A) L. birkmanni. (B) L. fedorense. (C) L. foxii. (D) L. inconditum. (E) L. lustrans. (F) L. macoupinense. (G) L. nelumbonis. (H) L. pectinatum. (I) L. pectorale. (J) L. sopinci. (K) L. swenki. Scale bar = 1 mm.

FIGURE 7. Genital capsule of male Lasioglossum subgenus Hemihalictus in dorsal view. (A) L. birkmanni. (B) L. fedorense. (C) L. foxii. (D) L. inconditum. (E) L. lustrans. (F) L. macoupinense. (G) L. nelumbonis. (H) L. pectinatum. (I) L. pectorale. (J) L. sopinci. (K) L. swenki. Scale bar = 1 mm.

FIGURE 8. Genital capsule of male Lasioglossum subgenus Hemihalictus in ventral view. (A) L. birkmanni. (B) L. fedorense. (C) L. foxii. (D) L. inconditum. (E) L. lustrans. (F) L. macoupinense. (G) L. nelumbonis. (H) L. pectinatum. (I) L. pectorale. (J) L. sopinci. (K) L. swenki. Scale bar = 1 mm.

FIGURE 21. Lateral habitus of female Lasioglossum (Hemihalictus) inconditum (Cockerell). Scale bar = 1 mm.

FIGURE 22. Lateral habitus of male Lasioglossum (Hemihalictus) inconditum (Cockerell). Scale bar = 1 mm.

FIGURE 23. Distribution map of Lasioglossum (Hemihalictus) inconditum (Cockerell). Heat map indicates predicted suitable range based on maximum entropy niche-modelling. Black circles indicate actual geo-referenced specimen records.

FIGURE 54. Lateral habitus of male Lasioglossum (Sphecodogastra) boreale (Svensson, Ebmer & Sakagami). Scale bar = 1 mm.

FIGURE 57. Lateral habitus of male Lasioglossum (Sphecodogastra) comagenense (Knerer & Atwood). Scale bar = 1 mm.

FIGURE 66. Lateral habitus of male Lasioglossum (Sphecodogastra) quebecense (Crawford). Scale bar = 1 mm.

FIGURE 76. Mesepisternum of female Lasioglossum. (A) L. inconditum showing weak, rugulose sculpture. (B) L. pectinatum showing punctation. (C) L. pectorale showing coarse, rugose sculpture throughout. (D) L. comagenense showing weak, rugulose sculpture. (E) L. lusorium showing appressed pubescense. (F) L. truncatum showing rugose sculpture becoming rugulose anterior to mesocoxa.

FIGURE 79. Propodeum and metapostnotum of female Lasioglossum (Evylaeus) and L. (Sphecodogastra). (A) L. cinctipes showing coarse, rugose sculpture. (B) L. truncatum showing coarse, rugose sculpture. (C) L. boreale showing fine sculpture and weak propodeal carinae. (D) L. comagenense showing fine sculpture and distinct propodeal carinae. (E) L. quebecense showing fine sculpture and distinct propodeal carinae. (F) L. seillean showing fine sculpture and distinct propodeal carinae.

FIGURE 82. Propodeum, metapostnotum and anterior surface of T1 of female Lasioglossum (Hemihalictus). (A) L. lustrans showing fine metapostnotal sculpture and subappressed hairs on T1. (B) L. foxii showing metapostnotal rugae not reaching posterior dorsal margin. (C) L. inconditum showing metapostnotal rugae reaching posterior dorsal margin.

FIGURE 84. Clypeus and mandible of male Lasioglossum (Hemihalictus). (A) L. sopinci showing large clypeal maculation and elongate mandible. (B) L. lustrans showing entirely black clypeus and elongate mandible. (C) L. inconditum showing narrow, distal, clypeal maculation and short mandible.

FIGURE 87. Lateral view of gonostylus and retrorse lobe of Lasioglossum (Sphecodogastra) in the fulvicorne-fratellum species-group. (A) L. boreale. (B) L. comagenense. (C) L. quebecense. (D) L. seillean. Ventral view of retrorse lobe. (E) L. quebecense. (F) L. seillean. Arrows indicate apex of retrorse lobe.