A revision of Bittacomorphinae with a review of the monophyly of extant subfamilies of Ptychopteridae (Diptera) Author Fasbender, Andrew Author Courtney, Gregory W. text Zootaxa 2017 4309 1 1 69 journal article 32358 10.11646/zootaxa.4309.1.1 f5a96f72-1549-41e7-9cf7-f282c66d73b6 1175-5326 846180 C5C20BC9-B99C-47D0-970E-07E20D13CF72 Bittacomorpha clavipes Fabricius 1781 Tipula clavipes Fabricius, 1781 : 404 (original description) Tipula clavipes Fabricius, Fabricius 1787 : 323 (redescription) Ptychoptera clavipes Fabricius, Fabricius 1805 : 22 (catalog, redescription) Bittacomorpha clavipes Fabricius, Westwood 1835 : 282 (set as type species of Bittacomorpha ) Bittacomorpha clavipes Fabricius, 1890: 230 (key) Bittacomorpha clavipes Aldrich, 1895 : 200 (original description) Bittacomorpha clavipes Fabricius, Johnson 1905 : 75 (key) Bittacomorpha (Bittacomorpha) clavipes Fabricius, Alexander 1916 : 546 (key) Bittacomorpha clavipes Fabricius, Alexander 1920 : 784 –787 (redescription) Bittacomorpha clavipes Fabricius, Alexander 1927 : 9 (catalog) Bittacomorpha clavipes Fabricius, Peus 1968: 11 (catalog) Bittacomorpha clavipes Fabricius, Alexander 1965 : 98 (catalog) Bittacomorpha clavipes Fabricius, Alexander 1967 : 171 (illustration) Diagnosis. Larva ( Figs. 5 a, 6a–b). Genae round and abbreviate, cranium olive-colored and spotted with black; postgena with rounded anterolateral lobes. Abdominal segments with linear rows of tubercles transverse dorsally and ventrally, longitudinal laterally. Adult ( Figs. 7 a, 8, 9). Scutum with longitudinal white stripe. Male with paramere apex weakly furcate. FIGURE 8. Bittacomorpha clavipes thoracic sclerites, right lateral view. anclf: Anepisternal cleft, anest: Anepisternal suture, antbs: Anterior basalare, aprnt: Antepronotum; aspr: Anterior spiracle, epmpl: episternum of metapleuron, epmrn: Epimeron, epstr: Episternum, hltr: Halter, lttg: Laterotergite of postnotum; mdtg: Mediotergite of postnotum, mplst: Metapleural suture, mrmpl: Meron of metapleuron, mtn: Metanotum, phltr: Prehalter, pprnt: Postpronotum, prsct: Prescutum, pspr: Posterior spiracle, ptg: Paratergite, sbasl: Subalar sclerite, sct: Scutum, sctl: Scutellum wngbs: Wing base. Description. Larva. Dorsal surface of cranium domed, hemispherical, olive-colored, rows of dark spots posteriorly on frontoclypeal apotome and dorsal surface of genae. Genae rounded in dorsal view, not expanded posteriorly, ventroposterior margin weakly and broadly emarginate; subgenal sclerite narrowed at midpoint of length, apex with semicircular anterolateral lobes. Antenna two-segmented, basal segment cylindrical, apical segment weakly fusiform, apex rounded, apical sensilla cluster directed laterally. Abdominal segments with rows of tubercles transverse dorsally and ventrally, longitudinal laterally. Adult. Measurements (n=5). Overall length 10.82mm ( 10.51–11.16mm ) head width 1.06mm ( 1.02–1.12mm ); height 0.95mm (0.88–1.0mm), length 0.61mm ( 0.57–0.67mm ); antennal flagellum length 5.17mm , flagellomere proportions 1.00, 0.82, 0.89, 1.00, 1.00, 0.89, 0.94, 0.89, 0.76, 0.76, 0.64, 0.64, 0.64, 0.51, 0.43, 0.43, 0.38, 0.38, 0.17; maxillary palpus length 1.49mm , maxillary palpus segment proportions 0.24, 1.0, 1.3, 0.96, 1.0; thorax length 1.96mm ( 1.65–2.22mm ), height 1.78mm ( 1.65–1.91mm ); wing length 7.26mm ( 6.7–7.9mm ), width 1.42mm ( 1.31–1.6mm ), R3+4+5 stem 0.60mm , straight; R4+5 stem 2.00mm, R4 after fork 1.25mm , R5 after fork 1.23mm , R3+4+5:R4+5-stem: R4 1.00, 3.33, 2.08, r-m 0.30mm , M distal to r-m 2.97mm , Mdistal:R3+4+5 4.95, 1.00, R4:Mdistal 0.42, 1.00. CuA2 1.78mm ; leg femur 1.0, 1.12, 1.27, tibia 0.98, 0.95, 1.19, tarsomeres foreleg 0.53, 0.22, 0.14, 0.05, 0.04, midleg 0.54, 0.21, 0.12, 0.05, 0.03, hind leg 0.58, 0.25, 0.16, 0.04, 0.06; abdomen length 11.87mm ( 10.85–13.15mm ), male tergite proportions 1.00, 3.13, 2.20, 2.23, 1.88, 0.89, 0.28, male genitalia width 1.03mm , height 0.71mm . Head. Face narrow, incised by clypeus to dorsal margin. Clypeus subcylindrical, lateral margins somewhat pinched at junction with face, shorter than eye, inflated. Hypopharynx subconical triangular, directed anteroventrally. Labium beyond clypeus greater than ½ length; labella pad subequal width to labium. Scutum and prescutum pollinose black, presecutum with white line along medial suture. Wing membrane without macrotrichia, apex of veins R1 and R3, with macrotrichia, others bare. Posthalter palpiform. Male. Epandrium with posterolateral corners simple right angle, posterior margin transverse laterally, with semicircular emargination medially; posteriomedial stylus short, triangular. Internal fold extending medially with slight posterior deflection. Epandrial clasper pad width of subapical clasper. Subepandrial sclerite divided, broadly sickle-shaped, fused to medial margin of base of epandrial clasper, directed towards hypoproct (additional small crescent-shaped weak sclerotization exists between subepandrial sclerites in subepandrial membrane in some specimens). Paramere with lateral spine at 7/8 length supporting lateral membrane; apex subacute triangular spine, with prominent dorsal tooth subapically. Gonostylus apex with or without irregularly spaced trichoid sensilla. Hypandrium terminal division foliform, petiolate base, two flattened lateral lobes directed laterally, posterior margin variable. Lateral ejaculatory process lateral apex broadly triangular, medial apex acute fingerlike; ventral lobe dorsally subacute quadrilateral towards blunt anterior apex, apex with lightly sclerotized region directed laterally, deeply inserted into sperm sac. Aedeagal sclerites widely separated at base, ventral apical apodeme apex pointed, fused to subapical anterior portion of subaedeagal sclerite. Subaedeagal sclerite internal section anterolateral apex obtuse round point, medial margin emarginate. External portion of subaedeagal sclerite extending dorsally beyond apex of dorsal apical apodeme. Female. Epigynium posterior margin V-shaped, medial suture present, faint; hypoproct broad, tongue-like, with sclerotized triangular boat-like sclerite ventrally, cercus dorsal margin straight, apex rounded, ventral margin swollen, potbellied anteriorly, tapering posteriorly. Genital fork spatulate, posterior margin weakly emarginate. Hypogynium a hemi-ring, hypogynial valve a short triangular lobe medially, lateral margins straight and angled dorsally, medial notch square: 0.75x as deep as broad. Material examined. COSTA RICA . GUANACOSTE . Bijagua , 22.ii.1991 , coll. D.W. Webb & M.E. Irwin [pinned 1 ♂ ; INHS ] . UNITED STATES . ALABAMA . Hale Co. Payne Lake , 24 mi . SE. of Tuscaloosa , 25.v.1978 , coll. G.W. Byers , C.W. Young [pinned 1 ♂ , 1 ♀ on same pin; KU ] ; Lee Co. Chewacia State Park 4 mi . SW. of Auburn , 26.v.1978 , coll. G.W., Byers , C.W. Young [pinned 1 ♀ ; KU ] . ARKANSAS . Hot Spring Co. Malvern , City park, 22.v.78 , coll. G.W. Byers , C.W. Young [pinned 3 ♂ ; KU ] ; Montgomery Co. Gap Creek , 1.3mi . SE. of Joplin , 22.v.1978 , coll. G.W. Byers , C.W. Young [pinned 1 ♂ ; KU ] ; Gap Creek Picnic Grd. 1 mi SE Joplin , 1.vi.1984 , coll. S.A. Teale [pinned 1 ♂ ; KU ] ; 2 mi NW Mount Ida , 1.vi.1984 , coll. S.A. Teale [pinned 1 ♂ ; KU ] . COLORADO . Fremont Co. , Hayden Creek 4 mi . SW. Coaldale , 13.vii.1974 , coll. G.W. Byers , C.W. Young [pinned 8 ♂ , 1 ♀ ; KU ] ; Gunnison Co. Gothic 23.vii.1952 , coll. B. Pruitt [pinned 6 ♂ , 1 ♀ ; UMMZ ] ; Mineral Co. Wolf Creek Cpgr. San Juan Natl. Forest , 25.vii.1981 , coll. G.W. Byers , S.A. Teale [pinned 3 ♀ ; KU ] ; San Juan Co. 30 mi . N. Durango , 13.viii.1965 , coll. G.W. Byers [pinned 1 ♀ ; KU ] . Unknown locality, collector and date [pinned 1 ♂ ; WSU ] . CONNECTICUT . Fairfield Co. Redding , 2.viii.1928 , coll. A.L. Melander [pinned 1 ♂ ; USNM ] ; GEORGIA . Butts Co. Unknown locality and collector, 6.v.1973 [pinned 2 ♂ ; FMNH ] ; Fulton Co. East Point , 15.v.1965 , coll. B. Harrison [pinned 1 ♀ ; KU ] ; Lumpkin Co. #5 De Soto Falls , 23.vi.1967 , coll. G.W. Byers [pinned 2 ♂ ; KU ] ; Oconee Co. 2mi . N. Watkinsville , 3.x.1977 , coll. G.W. Byers [pinned 2 ♂ ; KU ] ; Union Co. Lake Winfield Scott , 23.vi.1967 , coll. G.W. Byers [pinned 1 ♂ ; KU ] . ILLINOIS . Cook Co. Palos Park , 20.vi.1909 , coll. W.J. Gerhard [pinned 1 ♂ ; FMNH ] ; Dupage Co. Argonne Natl. Lab , 11.ix.1974 , coll. D. Pearson [pinned 1 ♂ ; FMNH ] ; Kane Co. 23.vii.1972 , coll. D. Pearson [pinned 2 ♂ , 4 ♀ , additional dates July 23, 1972 – July 27, 1973 ; FMNH ] . INDIANA . Lagrange Co. Pigeon River , 4.ix.1928 B.E.M. [pinned 1 ♀ , UMMZ ] ; LaPorte Co. Michigan City , 2.ix.1955 , coll. H. Dybas [pinned 1 ♂ ; FMNH ] ; IOWA . Dickinson Co. Iowa Lakeside Lab , 18.vii.1959 , J.L. Laffoon [pinned 1 ♂ ; ISIC ] ; Iowa Lakeside Lab Hillside Seep near marsh, 17.v.2010 , 43.383374°N , 95.179974°W , coll. G.W. Courtney , A. Fasbender [pinned 2 ♀ ; ISIC ] ; Silver Lake Fen 17.v.2010 , 43.437472°N , 95.365009°W , coll. G.W. Courtney , A. Fasbender [pinned 11 ♂ , 9 ♀ ; alcohol: 4 ♂ , 3 ♀ , 3 larvae , adults reared; ISIC ] ; Guthrie Co. Springbrook State Park , 24.viii.1972 , A. Stewart Cott [pinned 1 ♂ , 1♀ ; ISIC ] , 19.vii.1960 , coll. J. Laffoon [pinned 1 ♂ ; ISIC ] ; Story Co. Ames , 17.v.1964 , coll. W.S. Craig [pinned 43 ♂ , 2 ♀ ; ISIC ] . KANSAS . Douglas Co. Lawrence , Snow Hall , 8.vii.1987 , J. Gelhaus [pinned 1 ♀ ; KU ] 19.v.1986 , coll. B.N. Danforth [pinned 1♀ ; KU ] . MAINE . Hancock Co. Bar Harbor , 5.viii.1936 , coll. A.E. Brower [pinned 2 ♂ ; USNM] ; Ellsworth , 12.ix.1967 , coll. P.J. Spangler [pinned 1 ♀ ; USNM ] ; Penobscot Co. Orono , 18.viii.1981 , coll. S. Struad [pinned 1 ♂ ; KU ] . MICHIGAN . Calhoun Co. Battle Creek , 5.vii.1979 , G.W. Byers [pinned 1 ♀ ; KU ] ; Battle Creek , date and collector unknown [pinned 3 ♂ , 5 ♀ ; WSU ] ; Cheboygan Co. Cochran Lake , 19.vii.1957 , coll. R.E. Beer [pinned 1 ♂ ; KU ] ; 4mi . N. of Topinbee , 8.vii.1961 , coll. H. Willis [pinned 1 ♀ ; KU ] ; No collector, vii.1931 [pinned 1 ♂ , KU ] ; Emmet Co. 4mi . W. Pellston , 15.vii.1969 [pinned 3 ♂ , 2 ♀ ; UMMZ ] ; Gratiot Co. Ithaca , 13.vii.1959 , coll. V. Potter [pinned 3 ♂ , 5 ♀ ; UMMZ ] ; Jackson Co. 7.v.1939 , coll. F. Awnerman [pinned 1 ♀ , UMMZ ] ; Kalamazoo Co. Gull Lake Biol. Sta., 4.vii.1965 , coll. B. Hlavac , T. Hlavac [pinned 1 ♀ ; FMNH ] ; Schoolcraft Co. 14.viii.1951 , coll. J.S. Rogers [pinned 1 ♂ ; UMMZ ] . MINNESOTA . Clearwater Co. Itasca State Park , 2.ix.1950 , coll. J.L. Laffoon [pinned 6♂ , 1 ♀ ; ISIC ] . MISSISSIPPI . Lafayette Co. Puskus Lake , 8mi . NE. Oxford , 23.v.1978 , G.W. Byers , C.W. Young [pinned 1 ♀ ; KU ] ; Warren Co. Vicksburg , 23.ix.1977 , coll. G.W. Byers [pinned 1 ♂ ; KU ] . NEBRASKA . Cherry Co. Valentine , 9.vi.1950 , coll. Hicks , Slater , J.L. Laffoon , multiple dates 9–11.vi.1950 [pinned 6 ♂ , 5 ♀ ; ISIC ] ; Sheridan Co. Rushville , Malaise Trap , coll. R.W. Dawson , 12.vii.1967 [pinned 1 ♂ ; WSU ] . NEW YORK . Chenango Co. Smyrna , 16.viii.25 , coll. A.C. Weed [pinned 1 ♀ ; FMNH ] ; Clinton Co. Saranac Inn , 1900 , coll. O.S. Wescott [pinned 5 ♂ ; KU ] ; Hamilton Co. 7 miles S . of Long Lake , 26.viii.1956 , coll. W.E. LaBerge [pinned 2 ♂ ] ; Wayne Co. North Rose , 17.vi.1927 , Rawson-MacMillan Exped ., coll. A.C. Weed [pinned 1 ♀ ; FMNH ] . NORTH CAROLINA . Buncombe Co. Asheville , on store window, 29.iv.62 , coll. R.D. Peterson [pinned 1 ♀ ; ISIC ] ; Swain Co. 1.2, mi. NE. Clingman’s Dome , GSMNP, 15.vi.1967 , coll. G.W. Byers [pinned 1 ♂ ; KU ] . OHIO . Portage Co. 4 mi . E Kent , 8.viii.1967 , coll. M. Duckenonod [pinned 1 ♀ ; USNM ] . PENNSYLVANIA . Berks Co. French Creek Park , 7.vii.1960 , coll. G.W. Byers , multiple dates 7–10.vii.1960 [pinned 30 ♂ , 2 ♀ ; KU ] ; 1mi . SE. Reading , 15.viii.1956 , G.W. Byers [pinned 3♂ , 1 ♀ ; KU ] ; Franklin Co. Caledonia St. Park , 2.vii.1960 , coll. G.W, Byers and party [pinned 2 ♂ , KU ] ; Luzerne Co. Hazelton , 8.vii.1979 , coll. G.W. Byers [pinned 1 ♂ ; KU ] . SOUTH CAROLINA . Abbeville Co. 5 mi . S. Abbeville , 29.v.1978 , coll. G.W. Byers , C.W. Young [pinned 2 ♂ , 1 ♀ ; KU ] ; Oconee Co. Westminster , 30.v.1978 , coll. G.W., Byers , C.W. Young [pinned 1 ♀ ; KU ] ; Oconee State Park , Small lake 20.viii.1939 , coll. B.E. Montgomery #3974 [pinned 1 ♂ ; UMMZ ] ; Pickens Co. Clemson U. Experimental Forest , Six-Mile Creek , 17.iii.2005 , 34.75°N , 82.85°W , coll. G.W. Courtney [alcohol 5 pupae ; ISIC ] coll. M.J. Petersen [ 1 ♂ , 2 pupae ; ISIC ] ; SOUTH DAKOTA Pennington Co. Black Hills Nat. For. , Stream nr. South Rochford Road , Sweeping , 44.109621°N 103.734564°W , A Fasbender 29.vi.2010 [alcohol 3 ♂ ; ISIC ] ; Custer Co. Custer State Park , French Creek Picnic , 30.vi.2008 , 43.718°N , 103.4891°W , coll. J.D. Petersen [alcohol: 1 ♀ , ISIC ] ; Custer State Park , Sylvan Lake , 30. vi.2 0 0 8, 43.8459°N , 103.5602°W , coll.: J.D. Petersen [alcohol 1 ♀ ; ISIC ] . TENNESSEE Fentress Co. Allardt , 2.vi.1978 , coll. G.W., Byers , C.W. Young [pinned 2 ♂ ; KU ] ; Sevier Co. Fighting Creek , Great Smoky Mts. 7.vii.1958 , 35.67833°N , 83.555°W , J.L. Laffoon [pinned 1 ♂ ; ISIC ] . TEXAS . Bastrop Co . Bastrop St. Park , 20.iii.1976 , E.M. May [pinned 1 ♂ ; KU ] ; Nacogdoches Co. Nacogdoches , 12.x.1960 , coll. R. Oliver , multiple dates 8.iv.1960 – 6.v.1962 [pinned 7 ♂ , 1 ♀ ; KU ] . VERMONT Orleans Co. 5 mi . NNE. Orleans , 19.vi.1982 , coll. G.W. Byers [pinned 1 ♀ ; KU ] . VIRGINIA . Arlington Co. Alexandria , 15.vi.1973 , coll. B.M. O’Connor [pinned 1 ♀ ; UMMZ ] ; Fairfax Co. Lake Accotink , 5.viii.1972 , coll. B.M. O’Connor [pinned 1 ♀ ; UMMZ ] ; Giles Co. Mt. Lake Biol. Stn. 6–12.viii.1979 , coll. G.W. Byers #163 [pinned 1 ♂ ; KU ] ; Prince William Co. Woodbridge , 28.viii.1946 , coll. L.D. Beamer [pinned 1 ♂ ; KU ] . WISCONSIN . Vilas Co. Squaw Lake , 16.viii.1954 , coll. J.L. Laffoon [pinned 2 ♂ ; ISIC ] . WYOMING . Albany Co. Medicine Bow Nat. For. North Laramie River at Sand Lake Rd. Beaver Dam , 21.vi.2012 41.351557°N , 106.166511°W , coll. A Fasbender [pinned 1 ♀ ; ISIC ] . Taxonomic notes. This species can be easily differentiated from its congener Bittacomorpha occidentalis by the medial white stripe on the scutum, shortened and narrowed clypeus, and paramere with lateral spur and apical dorsal tooth. Larvae can be distinguished in dorsal view, the head capsule of Bi. clavipes being ovoid and that of Bi. occidentalis triangular. The holotype of Bittacomorpha clavipes is not listed in the collection of Fabricius type material at ZMUC, and may be lost or destroyed. Fortunately, the distinctive character of the inflated 1 st tarsomere allows the determination of the genus without contention, and material from the range of Bittacomorpha occidentalis would not have been available to Fabricius at the time. Distribution. Fig. 18 a.