Lasioglossum (Dialictus) coactus (Cresson, 1872)

Figs 9–11, 76B, 82B, 85A, 86A, 106A, 107A

Halictus coactus Cresson, 1872: 254 (lectotype designated in Cresson (1916), ♀, deposited in ANSP, type no. 2123, examined).

Halictus coactus – Cresson 1887: 292 (catalogue); 1916: 107 (lectotype designation). — Dalla Torre 1896: 58 (catalogue).

Halictus (Chloralictus) coactus – Ellis 1914: 100, 101 (comparison to jamaicae = gemmatum and tegulare), 103 (comparison to perparvum).

Lasioglossum (Chloralictus) coactum (!) – Michener 1951: 1112 (catalogue). — Scott et al. 2011: 29 (checklist).

Dialictus coactus – Hurd 1979: 1965 (catalogue). — Moure & Hurd 1987: 95 (catalogue).

Diagnosis

Females of L. coactus have T1–T3 distinctly convex in lateral view (although not as strongly so as in the male), with apical rims strongly depressed, shiny, and impunctate or very sparsely punctate, contrasting with densely punctate discs (IS ≤ 1 PD); T3 with dense and complete subapical band of tomentum; tegula relatively small (reaching but not exceeding posterior margin of mesoscutum) and densely punctate (IS ≤ 1 PD); metapostnotum usually shiny with strong anastomosing rugae reaching posterior margin (rarely dull with fine rugae); mesepisternum imbricate with punctures often dense (IS <1 PD) but distinctly separated and round (sometimes crowded (IS = 0 PD)); and inner metatibial spur with 3–4 branches.

Females of L. coactus are most similar to those of L. carlinvillense and L. tegulare, both of which have T1–T3 flat in lateral view. In addition, females of L. carlinvillense have the inner metatibial spur with only 2 branches and L. tegulare have the mesepisternum with crowded indistinct punctures (IS = 0 PD) or rugulose. Some species in the L. stictaspis species complex (particularly L. stictaspis s. str.) are also similar, but have T1–T3 flat in lateral view and more finely and sparsely punctate (IS = 1–3 PD), the mesepisternum shiny, and/or metapostnotum dull with fine, subparallel rugae.

Males of L. coactus have T1–T4 very strongly convex, with abruptly depressed, shiny, impunctate apical rims contrasting with densely punctate discs (IS <1 PD); tegula relatively small (reaching but not exceeding posterior margin of mesoscutum in dorsal view), with inner posterior margin weakly concave, and somewhat densely punctate (IS ≤1 PD); and mesepisternum densely punctate (IS <1 PD). They are not likely to be confused with any other species so long as T1–T4 are visible, but specimens with broken metasomas may be indistinguishable from L. tegulare .

Etymology

The specific epithet ʻ coactus ʼ could be either a Latin noun (compulsion, constraint) or a perfect participle (forced, compelled). Because Cresson (1872) did not specify the intended grammar, according to Article 31.2.2 of the International Code of Zoological Nomenclature (ICZN 1999), the epithet is treated as a noun and is therefore not declinable to ʻ coactum ʼ in combination with ʻ Lasioglossum ʼ, as has been done in the past.

Material examined

Holotype UNITED STATES – Texas • ♀; ANSP.

Other material

MEXICO – Chihuahua • 1 ♀; 83 km N of Chihuahua; [29.38° N, 106.07° W]; 29 Aug. 1991; Terry L. Griswold leg.; ex Coreopsis tinctoria; BBSL849296 . – Coahuila • 1 ♂; 61 mi. N of Saltillo; [26.31° N, 101° W]; elev. 853 m; 8 Sep. 1972; Byers, Thornhill leg.; SEMC • 1 ♀; San Juan de Sabinas; [27.93° N, 101.3° W]; elev. 354 m; 22 May 1952; P.S. Chrapliwy leg.; SEMC . – Hidalgo • 1 ♀; Grutas Tolantongo, 30 km NW of Cardonal (Barranco del Tolantongo); [20.65° N, 98.96° W]; elev. 1230 m; 13 Jul. 1990; Robert L. Minckley leg.; ex mimosoid legume; SEMC . – Nuevo León • 1 ♀; 12 mi. W of Linares; 24.8537° N, 99.7244° W; 21 Sep. 1974; George E. Bohart, W.J. Hanson leg.; BBSL1101070 • 1 ♀, 1 ♂; 15 mi. SW of China; [25.54° N, 99.41° W]; 19 Jul. 1954; SEMC • 1 ♀; 16 mi. S of Montemorelos; [24.96° N, 99.84° W]; 19 Jul. 1951; University of Kansas Mexico Expedition leg.; SEMC • 1 ♀; 4 km S of Linares; [24.81° N, 99.55° W]; elev. 400 m; 22 Mar. 1991; R.W. Brooks, R. Leschen leg.; ex Cercidium macrum; SEMC • 1 ♀; ibid.; 22 Mar. 1991; R.W. Brooks, R. Leschen leg.; ex yellow flower Opuntia; SEMC • 1 ♀; Ciudad Universitaria, San Nicolás de Garza, Jardin Desértico; [25.73° N, 100.31° W]; elev. 520 m; 4 Jun. 2009; Liliana Ramírez Freire leg.; LRF • 1 ♀; Ciudad Universitaria, San Nicolás de Garza, Jardin Desértico; [25.73° N, 100.31° W]; elev. 529 m; 1 Jun. 2009; Liliana Ramírez Freire leg.; ex Leucophyllum frutescens; LRF • 2 ♀♀; Parque Estatal El Cuchillo, China; [25.66° N, 99.27° W]; elev. 205 m; 27 Jun. 2009; Liliana Ramírez Freire leg.; LRF . – San Luis Potosi • 3 ♀♀, 3 ♂♂; 3.4 mi. NE of El Naranjo; [22.56° N, 99.3° W]; elev. 244 m; 5 Sep. 1962; University of Kansas Mexico Expedition leg.; SEMC • 1 ♀; Salinas; [22.63° N, 101.72° W]; Apr. 1982; R. Del Castillo leg.; SEMC • 1 ♂; Xilitla; [21.385° N, 98.975° W]; elev. 400 m; 7 Jul. 1990; Robert L. Minckley leg.; SEMC . – Tamaulipas • 1 ♂; 15 mi. N of Llera; [23.53° N, 99.03° W]; 24 Feb. 1972; F.D. Parker, D. Miller leg.; BBSL1101048 • 2 ♀♀; 3 mi. S of San Fernando; [24.8° N, 98.15° W]; 9 Jul. 1966; SEMC • 16 ♀♀; 8 mi. S of Jiménez; [24.1° N, 98.49° W]; 15 Jun. 1953; ex Parthenium hysterophorus; SEMC • 1 ♀; Antiguo Morelos; [22.55° N, 99.08° W]; elev. 300 m; 6 Jul. 1990; Ilan Yarom leg.; SEMC • 1 ♂; Forlon; [23.24° N, 98.8° W]; 30 Sep. 1938; L.J. Lipovsky leg.; SEMC • 1 ♀, 2 ♂♂; Jiménez; [24.22° N, 98.49° W]; 19 Jul. 1962; University of Kansas Mexico Expedition leg.; SEMC • 1 ♀; Matamoros; [25.87° N, 97.5° W]; 27 Sep. 1954; SEMC • 3 ♀♀, 1 ♂; Padilla; [24.05° N, 98.9° W]; 15 Jun. 1953; University of Kansas Mexico Expedition leg.; SEMC • 1 ♀; Playa Altamira; [22.4° N, 97.845° W]; 5 Jul 196?; M.S. Wasbaner, J.E. Slansky leg.; UCDC • 1 ♂; Rio Conchos, San Fernando; [24.84° N, 98.16° W]; 9 Jun. 1966; University of Kansas Mexico Expedition leg.; SEMC • 1 ♀; Tampico; [22.23° N, 97.86° W]; 27 Mar. 1951; W.P. Stephen leg.; SEMC . – Veracruz • 5 ♀♀; Llave Panuco; 22.0504° N, 98.185° W; 26 Feb. 1972; F.D. Parker, D. Miller leg.; BBSL1101039, BBSL1101040, BBSL1101041, BBSL1101042, BBSL1101043 .

UNITED STATES – Colorado • 1 ♀; Boulder Co., Palisades, “L. H. Can.” [Lefthand Canyon]; [40.11° N, 105.305° W]; 25 Jul. 1942; Hugo G. Rodeck leg.; ex Carduus; UCMC • 2 ♀♀; Jefferson Co.; 39.7433° N, 105.2365° W; 22 Aug. 2001; Sam W. Droege leg.; PCYU USGS-DRO 020468, USGS-DRO 020471 • 1 ♂; Las Animas Co., Model; 37.493° N, 103.8345° W; 23 Jul. 2009; J. Newton leg.; PCYU • 1 ♂; Grand Junction; [39.06° N, 108.55° W]; 4 Sep. 1938; D.E. Hardy leg.; SEMC . – Kansas • 2 ♀♀; Barber Co.; 37.0873° N, 98.379° W; 29 Apr. 2013; S. Griffin leg.; WRME . – New Mexico • 1 ♀; De Baca Co., Old Fort Sumner; [34.402° N, 104.194° W]; 17 Jun. 1999; W.F. Chamberlain leg.; ex Melilotus; TAMU-ENTO X1400068 • 1 ♀; Doña Ana Co., New Mexico State University campus; [32.28° N, 106.75° W]; elev. 1189 m; 11 Jun. 1979; G.S. Forbes leg.; ex Santolina; NMSU • 2 ♂♂; Eddy Co., Carlsbad; [32.42° N, 104.23° W]; 31 May 1979; Carrola, D.R. Delorme, Friedlander leg.; TAMU-ENTO X1365290, X1399323 • 1 ♀; Eddy Co., Carlsbad; [34.42° N, 104.23° W]; 20 May 1969; D.J. Brothers, Krueger, Charles D. Michener leg.; ex Lepidium integrifolium; SEMC • 1 ♂; Eddy Co., Pecos River at Highway 31; [32.31° N, 104.06° W]; 13 Jul. 1979; D.R. Delorme, C.P. McHugh leg.; TAMU-ENTO X1365429 • 1 ♀; Eddy Co., Rattlesnake Springs; 32.1095° N, 104.4716° W; elev. 1115 m; 18 Jun. 2010; J.D. Herndon leg.; BBSL CAVE2826 • 1 ♂; Otero Co., Alamogordo Dam; [32.93° N, 105.95° W]; 6 Jun. 1969; D. Liesnee leg.; ex Tamarix; NMSU • 1 ♀; Quay Co., Tucumcari; [35.17° N, 103.73° W]; 10 Jul. 1963; R.M. Bohart leg.; UCDC • 2 ♀♀; Sandoval Co., Corrales; [35.24° N, 106.61° W]; 23 Apr. 2000; R. Fagerlund leg.; PCYU • 2 ♀♀; 3 miles west of Cimarron, “Ovka? Kenton ”; [36.51° N, 104.97° W]; 23 Jun. 1970; D. Liesnee leg.; ex Tamarix; NMSU • 11 ♀♀; Carrizozo; [33.64° N, 105.88° W]; 9 Jun. 1950; L.D. Beamer leg.; ex Asclepias galloides; SEMC • 27 ♀♀; ibid.; 9 Jun. 1950; L.D. Beamer leg.; ex Dasylirion wheeleri; SEMC • 1 ♀; ibid.; 9 Jun. 1950; L.D. Beamer leg.; ex Marrubium vulgare; SEMC • 14 ♀♀, 5 ♂♂; ibid.; 9 Jun. 1950; L.D. Beamer leg.; ex Pyrrhopappus multicaulis; SEMC • 37 ♀♀, 1 ♂; ibid.; 9 Jun. 1950; L.D. Beamer leg.; SEMC . – Texas • 1 ♀; Anderson Co., 10 mi. SW of Elkhart; [31.52° N, 95.7° W]; 5–6 Jun. 1976; H.R. Burke leg.; TAMU-ENTO X1365914 • 1 ♀; Anderson Co., Salmon; [31.57° N, 95.5° W]; 17 May–6 Jun. 1975; H.R. Burke leg.; TAMU-ENTO X1365410 • 1 ♀; Bandera Co., 10 miles west of Medina; [29.8° N, 99.42° W]; 17 May 2004; W.F. Chamberlain leg.; TAMU-ENTO X1416743 • 1 ♀; Bastrop Co., Camp Swift; 30.2833° N, 97.3233° W; 2 May 2002; J.L. Neff leg.; ex Coreopsis basalis; CTMI 23406 • 1 ♀; Bastrop Co., Camp Swift; 30.284° N, 97.323° W; 2 Oct. 2003; J.L. Neff leg.; ex Heterotheca subaxillaris; CTMI 26042 • 1 ♀; ibid.; 2 Apr. 2004; J.L. Neff leg.; ex Ilex vomitoria; CTMI 26762 • 1 ♀; Bastrop Co., Camp Swift; 30.296° N, 97.31° W; 2 Oct. 2003; J.L. Neff leg.; ex Croptilon divaricatum; CTMI 26078 • 4 ♀♀, 1 ♂; Bastrop Co., Stengl Biological Station, 6 mi. N of Smithville; [30.09° N, 97.17° W]; 12 May 2000; J.C. Abbott, R. Caesar leg.; PCYU • 2 ♀♀; Bexar Co., Ebony Hill Research Station (Kendall residence); [29.488° N, 98.696° W]; 12 May 2003; Roy O. Kendall leg.; TAMU-ENTO X0408802.1, X0408802.2 • 1 ♀; Bexar Co., San Antonio; [29.42° N, 98.49° W]; 29 Aug. 1964; A.H. Alex leg.; ex Helianthus annuus; TAMU-ENTO X1400725 • 2 ♀♀; ibid.; 12 Jul. 2018; Richard S. Peigler leg.; ex Astrophytum myriostigma; TAMU-ENTO X1600774, X1602051 • 1 ♀; ibid.; 13 Jul. 2018; Richard S. Peigler leg.; ex Astrophytum ornatum; TAMU-ENTO X1597095 • 1 ♀; ibid.; 13 Jul. 2018; Richard S. Peigler leg.; ex Leuchtenbergia principis; TAMU-ENTO X1601105 • 1 ♀; ibid.; 14 Jul. 2018; Richard S. Peigler leg.; ex Leuchtenbergia principis; TAMU-ENTO X1599632 • 1 ♀; Bexar Co.; [29.4° N, 98.5° W]; 20 Oct. 1928; H.B. Parks leg.; TAMU-ENTO X1387356 • 1 ♀; ibid.; 3 Apr. 1929; H.B. Parks leg.; TAMU-ENTO X1418300 • 1 ♀; ibid.; 9 May 1930; H.B. Parks leg.; TAMU-ENTO X1370844 • 1 ♀; ibid.; 24 Aug. 1930; H.B. Parks leg.; TAMU-ENTO X1386235 • 1 ♀; ibid.; 30 Sep. 1931; H.B. Parks leg.; TAMU-ENTO X1370854 • 1 ♀; Bosque Co., 2 mi. W of Iredell; [31.98° N, 97.91° W]; 21 Apr. 1972; J.C. Schaffner leg.; TAMU-ENTO X1362257 • 1 ♂; Brazos Co., 0.3 mi. N of Jones Road & Highway 60 junction; [30.586° N, 96.391° W]; 12 Jul. 1975; S.J. Merritt leg.; TAMU-ENTO X1276417 • 1 ♀; Brazos Co., 9 km SSE of College Station, 15889 Woodlake Drive; 30.5314° N, 96.2817° W; elev. 90 m; 3 Sep. 2013; J.D. Oswald leg.; TAMU-ENTO X1003142 • 1 ♂; Brazos Co., Bryan; [30.67° N, 96.37° W]; 1–2 May 1976; H.R. Burke leg.; TAMU-ENTO X1405850 • 9 ♀♀; ibid.; 29–30 May 1976; H.R. Burke leg.; TAMU-ENTO X1364704, X1364839, X1365258, X1365688, X1398752, X1398831, X1399013, X1399406, X1399493 • 2 ♂♂; ibid.; 29–30 May 1976; H.R. Burke leg.; TAMU-ENTO X1365317, X1399226 • 1 ♀; Brazos Co., College Station; [30.63° N, 96.33° W]; 21 Oct. 1954; A.H. Alex leg.; ex Aster; TAMU-ENTO X1386948 • 3 ♀♀; ibid.; 5 Jun. 1955; A.H. Alex leg.; ex Leucophyllum frutescens; TAMU-ENTO X1389396, X1389399, X1389440 • 1 ♀; ibid.; 13 Jul. 1962; A.H. Alex leg.; ex Leucophyllum frutescens; TAMU-ENTO X1386731 • 1 ♀; ibid.; 15 Jun. 1963; A.H. Alex leg.; ex Lactuca sativa; TAMU-ENTO X1312803 • 3 ♀♀; ibid.; 16 Jun. 1963; A.H. Alex leg.; ex Melilotus alba; TAMU-ENTO X1312806, X1312916, X1313021 • 1 ♀; ibid.; 27 Jun. 1963; A.H. Alex leg.; ex Melilotus alba; TAMU-ENTO X1313095 • 1 ♂; ibid.; 29 Jun. 1963; A.H. Alex leg.; ex Mentha spicata; TAMU-ENTO X1399190 • 1 ♀; ibid.; 15 Jul. 1963; A.H. Alex leg.; ex Chrysopsis pilosa; TAMU-ENTO X1365181 • 1 ♀; ibid.; 28 Mar. 1964; J.C. Schaffner leg.; TAMU-ENTO X1393271 • 1 ♀; ibid.; 11 May 1964; A.H. Alex leg.; ex Opuntia macrorhiza; TAMU-ENTO X1313206 • 1 ♀; ibid.; 15–22 May 1977; S.J. Merritt leg.; TAMU-ENTO X1365222 • 1 ♀; ibid.; 27 Apr. 1979; R. Krenek leg.; TAMU-ENTO X1370547 • 1 ♀; ibid.; 9 Mar. 1985; S.J. Merritt leg.; ex Lamium; TAMU-ENTO X1404418 • 1 ♀; Brazos Co., College Station, Church Avenue; [30.62° N, 96.346° W]; 17 Mar. 2000; B. Rodriguez-Velez leg.; TAMU-ENTO X1404670 • 2 ♀♀; Brazos Co., College Station, Riley Estate; [30.5885° N, 96.2537° W]; 14 Apr. 2001; E.G. Riley leg.; TAMU-ENTO X1315282, X1419587 • 1 ♀; Brazos Co., College Station, Riley Estate; 30.5885° N, 96.2537° W; 1–3 Apr. 2011; E.G. Riley, M. T. leg.; TAMU-ENTO X1428008 • 1 ♀; ibid.; 9–10 Apr. 2011; E.G. Riley, M. T. leg.; TAMU-ENTO X1428075 • 2 ♀♀; ibid.; 18–19 May 2011; E.G. Riley, M. T. leg.; TAMU-ENTO X1428021, X1428040 • 2 ♀♀; ibid.; 4–21 Mar. 2013; E. Dinh, K. Henson, E. Riley leg.; TAMU-ENTO X1370402, X1414696 • 1 ♂; Brazos Co., Jones Road, 1.6 mi. N of Highway 60; [30.597° N, 96.408° W]; 1 Jun. 1975; S.J. Merritt leg.; TAMU-ENTO X1401125 • 1 ♀; Brazos Co., Lick Creek Park; [30.56° N, 96.21° W]; 9 Aug. 2018; Karen W. Wright leg.; TAMU-ENTO X1600920 • 2 ♀♀; Brazos Co., near College Station, TAMU Range Sci. Area; [30.58° N, 96.37° W]; 18 Jun. 1978; S.J. Merritt leg.; ex Cassia sp.; TAMU-ENTO X1393258, X1395177 • 8 ♀♀; ibid.; 30 Jul. 1978; S.J. Merritt leg.; ex Euphorbia; TAMU-ENTO X1391760, X1393557, X1395215, X1395805, X1395980, X1396015, X1397381, X1397397 • 1 ♀; Brazos Co., on FM 2154 6.9 miles south of Wellborn; [30.47° N, 96.22° W]; 12 Sep. 1981; S.J. Merritt leg.; ex Euphorbia sp.; TAMU-ENTO X1397470 • 3 ♀♀; Brazos Co., Southwest Parkway and Wellborn Road; [30.59° N, 96.33° W]; 10 Jun. 1979; S.J. Merritt leg.; ex Monarda sp.; TAMU-ENTO X1393434, X1395788, X1397468 • 1 ♀; Brazos Co., TAMU-Riverside Campus; 30.6428° N, 96.4935° W; elev. 80 m; 19 Sep. 2013; J.D. Oswald leg.; TAMU-ENTO X1000976 • 2 ♀♀; ibid.; 28 Sep. 2013; K. Fryer leg.; TAMU-ENTO X1014172, X1015505 • 1 ♀; Brazos Co.; [30.6° N, 96.3° W]; 23 Jul. 1955; A.H. Alex leg.; ex sesame; TAMU-ENTO X1312589 • 1 ♀; Brazos Co.; 30.5922° N, 96.2727° W; elev. 88 m; 8–26 Feb. 2017; Karen W. Wright leg.; TAMU-ENTO X1534944 • 1 ♀; ibid.; 26 Feb.–8 Mar. 2017; Karen W. Wright leg.; TAMU-ENTO X1533805 • 1 ♀; ibid.; 25–31 Mar. 2017; Karen W. Wright leg.; TAMU-ENTO X1534899 • 2 ♀♀; ibid.; 16–23 Apr. 2017; Karen W. Wright leg.; TAMU-ENTO X1534915, X1534949 • 3 ♀♀; ibid.; 23–30 Apr. 2017; Karen W. Wright leg.; TAMU-ENTO X1534926, X1534945, X1534950 • 12 ♀♀; ibid.; 30 Apr.–13 May 2017; Karen W. Wright leg.; TAMU-ENTO X1533803, X1533804, X1533808, X1533822, X1533827, X1534913, X1534922, X1534955, X1534958, X1534971, X1534974, X1534975 • 1 ♀; Burleson Co., Highway 50 1.8 mi. N of Highway 60 junction; [30.558° N, 96.47° W]; 30 May 1975; S.J. Merritt leg.; TAMU-ENTO X1400784 • 2 ♀♀; Burnet Co., 5 mi. SE of Marble Falls; [30.53° N, 98.33° W]; 9 Sep. 2017; Karen W. Wright leg.; ex Argemone albiflora; TAMU-ENTO X1520087, X1520473 • 1 ♀; ibid.; 9 Sep. 2017; Karen W. Wright leg.; ex Polanisia dodecandra; TAMU-ENTO X1519455 • 2 ♂♂; Burnet Co., Inks Lake State Park; [30.73° N, 98.36° W]; 2 May 1987; J.B. Woolley leg.; TAMU-ENTO X1314946, X1417609 • 1 ♀; Caldwell Co., 4 mi. W of Luling; [29.68° N, 97.71° W]; 27 Sep. 1965; George E. Bohart leg.; ex Chrysopsis viscida; BBSL700725 • 1 ♀; Cameron Co., Brownsville; [25.9° N, 97.5° W]; 10–13 Mar. 1979; T.P. Friedlander leg.; TAMU-ENTO X1365387 • 1 ♀; Comanche Co., DeLeon; [32.11° N, 98.54° W]; 7 Jun. 1978; F.L. Mitchell leg.; TAMU-ENTO X1364763 • 4 ♀♀; Denton Co.; 33.2043° N, 97.0816° W; 27 Apr.; H.W. Ikerd leg.; PCYU USGS-DRO 004828, USGS-DRO 004837, USGS-DRO 004913, USGS-DRO 004960 • 1 ♂; Eastland Co., 3 miles north of Desdemona; [32.31° N, 98.55° W]; 1 Jun. 1998; W.F. Chamberlain leg.; TAMU-ENTO X1398711 • 1 ♀; Edwards Co., 2 mi. NW of Camp Wood; 29.6826° N, 100.0277° W; 14 Apr. 2018; Karen W. Wright leg.; ex Salvia farinacea; TAMU-ENTO X1601179 • 1 ♂; Edwards Co., 23 mi. W of Rocksprings; [30.02° N, 100.59° W]; 6 Aug. 1982; J.C. Schaffner leg.; ex Rhus sp.; TAMU-ENTO X1393599 • 10 ♀♀; Erath Co., 3 mi. W of Bluff Dale, bank of Paluxy River; [32.33° N, 98.07° W]; 8–15 Jul. 1976; R.L. Sams leg.; TAMU-ENTO X1364558, X1365416, X1365671, X1365893, X1366146, X1400758, X1400803, X1400873, X1401168, X1401183 • 4 ♀♀; ibid.; 29 Jul.–4 Aug. 1976; R.L. Sams leg.; TAMU-ENTO X1398534, X1399137, X1401145, X1401178 • 1 ♀; Erath Co., 5 mi. N of Stephenville; [32.29° N, 98.2° W]; 21–23 May 1980; P.T. Riherd leg.; TAMU-ENTO X1365904 • 4 ♀♀; ibid.; 27– 30 May 1980; P.T. Riherd leg.; TAMU-ENTO X1312282, X1314928, X1386058, X1386231 • 1 ♂; ibid.; 30 May 1980; P.T. Riherd leg.; TAMU-ENTO X1366121 • 5 ♀♀; ibid.; 1–6 Jun. 1980; P.T. Riherd leg.; TAMU-ENTO X1312347, X1365493, X1399026, X1420881, X1422490 • 2 ♀♀; Erath Co., Bluff Dale; [32.35° N, 98.02° W]; 13–20 May 1978; R.L. Sams leg.; TAMU-ENTO X1395228, X1417326 • 2 ♀♀; Erath Co., Bluff Dale, 1 mi. N of Paluxy River; [32.37° N, 98.03° W]; 21–27 Apr. 1978; R.L. Sams leg.; TAMU-ENTO X1365448, X1366157 • 6 ♀♀; ibid.; 28 Apr.–4 May 1978; R.L. Sams leg.; TAMU-ENTO X1391859, X1393360, X1399245, X1399334, X1399353, X1399485 • 8 ♀♀; ibid.; 5–13 May 1978; R.L. Sams leg.; TAMU-ENTO X1313256, X1388076, X1388078, X1388210, X1391955, X1392376, X1397504, X1400009 • 2 ♂♂; ibid.; 5–13 May 1978; R.L. Sams leg.; TAMU-ENTO X1393257, X1395802 • 7 ♀♀; ibid.; 19–26 May 1978; R.L. Sams leg.; TAMU-ENTO X1392365, X1393152, X1393435, X1396001, X1396075, X1397414, X1397524 • 2 ♂♂; ibid.; 19–26 May 1978; R.L. Sams leg.; TAMU-ENTO X1397525, X1397529 • 1 ♀; ibid.; 13–20 Jun. 1978; R.L. Sams leg.; TAMU-ENTO X1362234 • 1 ♀; Erath Co., Bluff Dale, Paluxy River; [32.35° N, 98.02° W]; 25–29 Mar. 1977; R.L. Sams leg.; TAMU-ENTO X1400161 • 5 ♀♀; Erath Co., FM 1188 3 miles west of Bluff Dale; [32.36° N, 98.06° W]; 16–23 Jul. 1976; R.L. Sams leg.; TAMU-ENTO X1365188, X1365505, X1365517, X1398865, X1399298 • 2 ♂♂; ibid.; 16–23 Jul. 1976; R.L. Sams leg.; TAMU-ENTO X1365131, X1413145 • 5 ♀♀; ibid.; 22–29 Jul. 1976; R.L. Sams leg.; TAMU-ENTO X1365458, X1365579, X1398636, X1398887, X1399390 • 1 ♂; ibid.; 22–29 Jul. 1976; R.L. Sams leg.; TAMU-ENTO X1365178 • 1 ♀; Fayette Co., Schulenburg; [29.68° N, 96.9° W]; 28 Sep. 1965; George E. Bohart leg.; ex Baccharis; BBSL700732 • 1 ♀; Fayette Co.; 29.897° N, 98.8379° W; 23 Jul. 2016; Sam W. Droege leg.; WRME 376569 • 2 ♀♀; Frio Co., 6 mi. SE of Pearsall; [28.83° N, 99.02° W]; 7 Jul. 1972; E.E. Grissell, J. Smith leg.; TAMU-ENTO X1362222, X1396247 • 2 ♂♂; ibid.; 7 Jul. 1972; E.E. Grissell, J. Smith leg.; TAMU-ENTO X1362225, X1393179 • 1 ♀; Galveston Co., Texas A&M University at Galveston; [29.32° N, 94.82° W]; 14 May 2000; J.C. Abbott, R. Caesar leg.; PCYU • 1 ♀; Garza Co.; [33.2° N, 101.3° W]; 9 Jun. 1980; Thieband leg.; TAMU-ENTO X1388272 • 1 ♀; Gillespie Co., Morris Ranch; [30.22° N, 99.01° W]; 11 Sep. 1994; W.F. Chamberlain leg.; TAMU-ENTO X1416318 • 1 ♀; Harris Co., Houston, University of Houston; 29.7245° N, 95.3451° W; 27 Jul. 2017; Scott Gilb. leg.; ex Croton capitatus; TAMU-ENTO X1576029 • 2 ♀♀; Hidalgo Co., 3 mi. E of Hargill; [26.44° N, 97.96° W]; 4 May 1983; D.A. Dean leg.; ex cotton; TAMU-ENTO X1382605, X1386601 • 1 ♂; Hidalgo Co., Bentsen-Rio Grande Valley State Park; [26.175° N, 98.385° W]; 16 Jun. 1978; C.C. Porter leg.; TAMU-ENTO X1386607 • 1 ♂; ibid.; 25 Aug. 1982; C.C. Porter leg.; TAMU-ENTO X1388263 • 1 ♀; Hidalgo Co., McAllen Botanical Garden; [26.2062° N, 98.2663° W]; 21 Aug. 1982; C.C. Porter leg.; TAMU-ENTO X1387872 • 1 ♀; Hill Co., 6 mi. S of Hillsboro; [31.92° N, 97.13° W]; 9 Aug. 1983; D.A. Dean leg.; TAMU-ENTO X1395167 • 1 ♂; Hockley Co.; [33.6° N, 102.4° W]; 11 Jun. 1981; L. Berger leg.; ex sunflower; TAMU-ENTO X1388815 • 1 ♀; Kendall Co., Boerne City Park; [29.785° N, 98.71° W]; 19 Apr. 1984; J.C. Schaffner leg.; TAMU-ENTO X1417011 • 1 ♀; Kenedy Co., Rest Stop #2, 4 mi. N of Raymondville; [27.13° N, 97.79° W]; 2 Aug. 1978; M. Stiller leg.; TAMU-ENTO X1418448 • 4 ♂♂; Kerr Co., “ 3.3 mi. N ” [E] of Hunt on S. R. 39 Bumblebee Creek; [30.1° N, 99.3° W]; 22 Jul. 1974; H. Greenbaum leg.; TAMU-ENTO X0283941, X1386577, X1387035, X1387359 • 1 ♀; Kerr Co., 9 miles west of Mountain Home; [30.17° N, 99.52° W]; 16 Jun. 1997; W.F. Chamberlain leg.; TAMU-ENTO X1395186 • 2 ♀♀; Kerr Co., Center Point; [29.95° N, 99.04° W]; 20–26 May 1987; Wharton, Praetorius leg.; TAMU-ENTO X1388019, X1388477 • 4 ♂♂; ibid.; 20–26 May 1987; Wharton, Praetorius leg.; TAMU-ENTO X1312365, X1387784, X1388547, X1388732 • 1 ♀; ibid.; 5–11 Jun. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1386942 • 2 ♀♀; ibid.; 12–18 Jun. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1386333, X1386474 • 1 ♂; ibid.; 12–18 Jun. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1386275 • 8 ♀♀; ibid.; 25 Jun.–3 Jul. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1386045, X1386375, X1386585, X1387684, X1388548, X1388824, X1388972, X1389066 • 3 ♂♂; ibid.; 25 Jun.– 3 Jul. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1383867, X1387373, X1388916 • 1 ♂; ibid.; 24–30 Jul. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1388122 • 2 ♀♀; ibid.; 7–13 Aug. 1987; Wharton, Praetorius leg.; TAMU-ENTO X1312283, X1386811 • 1 ♀; Kerr Co., Kerrville; [30.05° N, 99.14° W]; 28 Apr. 1965; TAMU-ENTO X1405137 • 2 ♂♂; Kleberg Co., 20 miles southeast of Kingsville; [27.31° N, 97.62° W]; 4–5 Jun. 1983; W.J. Pulawski leg.; CAS • 1 ♀; ibid.; 6–8 Jun. 1983; W.J. Pulawski leg.; CAS • 4 ♂♂; ibid.; 6–8 Jun. 1983; W.J. Pulawski leg.; CAS • 1 ♀; Kleberg Co., Leo Kaufer Memorial Park; [27.315° N, 97.68° W]; 2 Aug. 1978; M. Stiller leg.; TAMU-ENTO X1416758 • 4 ♀♀; Lavaca Co., 15 mi. S of Hallettsville; [29.22° N, 96.97° W]; 28 Aug. 1965; George E. Bohart leg.; BBSL700728, BBSL700733, BBSL700774, BBSL700775 • 1 ♀; ibid.; 28 Sep. 1965; George E. Bohart leg.; ex Croton; BBSL700721 • 1 ♀; Leon Co., 2.8 mi. N of Normangee; [31.07° N, 96.12° W]; 12 Jun. 1976; S.J. Merritt leg.; ex Monarda; TAMU-ENTO X1398606 • 1 ♀; Live Oak Co., 8 miles south of George West; [28.22° N, 98.11° W]; 22 Aug. 1975; D.W. Plitt leg.; ex Garza pasture; TAMU-ENTO X1399068 • 1 ♂; ibid.; 25 Aug. 1975; D.W. Plitt leg.; TAMU-ENTO X1399237 • 1 ♀; Llano Co., 10 mi. W of Llano; 30.709° N, 98.824° W; 2 Mar. 2009; J.L. Neff leg.; ex Rhus trilobata; CTMI 33372 • 1 ♀; Milam Co., 14.6 miles NW of junction of Highway 79 and FM 487; [30.75° N, 97.18° W]; 10 Jun. 1978; S.J. Merritt leg.; ex Coreopsis sp.; TAMU-ENTO X1402200 • 1 ♂; Palo Pinto Co., Possum Kingdom Reservoir; [32.9° N, 98.5° W]; 31 Aug. 1990; J.L. Tracy leg.; TAMU-ENTO X1498264 • 9 ♀♀; Potter Co., Bushland; [35.19° N, 102.06° W]; 17–24 Jul. 1983; T.J. Kring leg.; TAMU-ENTO X1392217, X1392366, X1393210, X1393614, X1395180, X1395920, X1397321, X1397441, X1397460 • 1 ♂; ibid.; 17–24 Jul. 1983; T.J. Kring leg.; TAMU-ENTO X1395872 • 10 ♀♀; ibid.; 7–15 Aug. 1983; T.J. Kring leg.; TAMU-ENTO X1383647, X1383683, X1386832, X1386845, X1387976, X1388196, X1388344, X1388354, X1388721, X1388886 • 1 ♂; ibid.; 7–15 Aug. 1983; T.J. Kring leg.; TAMU-ENTO X1388693 • 1 ♀; Reeves Co., 11 mi. W of Balmorhea State Park (“ Route 290 ” [3078?]); [31° N, 103.94° W]; 18 Aug. 1980; T.P. Friedlander, T.A. Friedlander, P.W. Kovarik leg.; TAMU-ENTO X1386814 • 1 ♀; Reeves Co., TAMU Research Station at Pecos; [31.37° N, 103.62° W]; 17 Aug. 1976; S.J. Merritt leg.; ex alfalfa; TAMU-ENTO X1400891 • 1 ♀; Robertson Co., 3.4 miles east of junction of Highway 6 and OSR; [30.75° N, 96.4° W]; 14 Aug. 1976; S.J. Merritt leg.; ex Euphorbia; TAMU-ENTO X1365218 • 3 ♀♀; Robertson Co., Salter Research Farms; 31.0333° N, 96.7725° W; elev. 88 m; 23 Apr. 2017; Karen W. Wright leg.; TAMU-ENTO X1533744, X1533769, X1533775 • 3 ♀♀; San Patricio Co., Welder Wildlife Refuge; [28.11° N, 97.42° W]; 28 Jun. 1984; J.C. Schaffner leg.; ex hackberry motte; TAMU-ENTO X1391640, X1391716, X1391878 • 1 ♀; San Patricio Co., Welder Wildlife Refuge; 28.1122° N, 97.4167° W; elev. 20 m; Mar. 2004; Sty Reed leg.; PCYU • 1 ♀; Somervell Co., 3 mi. S of Glen Rose; [32.19° N, 97.76° W]; 4 May 1973; J.C. Schaffner leg.; TAMU-ENTO X1362253 • 3 ♀♀; Somervell Co., 6 mi. N of Glen Rose, Comanche Peak Electric Station; [32.3° N, 97.8° W]; 16–17 Jun. 1979; J.P. Cuda leg.; TAMU-ENTO X1393570, X1395851, X1396070 • 1 ♂; Starr Co., Falcon State Park; [26.58° N, 99.14° W]; 10 Jun. 1969; Veryl V. Board leg.; TAMU-ENTO X1395821 • 1 ♂; ibid.; 19 Jun. 1969; Veryl V. Board leg.; TAMU-ENTO X1396041 • 2 ♀♀; ibid.; 18 May 1978; C.C. Porter, A. Cerbone leg.; TAMU-ENTO X1313681, X1385482 • 1 ♂; ibid.; 18 May 1978; C.C. Porter, A. Cerbone leg.; TAMU-ENTO X1387384 • 3 ♂♂; ibid.; 20 Apr. 1985; J.B. Woolley leg.; TAMU-ENTO X1387994, X1388896, X1416975 • 2 ♀♀; Taylor Co., Dyess Air Force Base; 32.4072° N, 99.829° W; 12–13 Apr. 2001; H.W. Ikerd leg.; TAMU-ENTO X1387947, X1388030 • 1 ♀; Taylor Co., Dyess Air Force Base; 32.4415° N, 99.8111° W; 12–13 Apr. 2001; H.W. Ikerd leg.; TAMU-ENTO X1312644 • 1 ♀; ibid.; 9–10 May 2001; H.W. Ikerd leg.; TAMU-ENTO X1387955 • 2 ♀♀; Travis Co., Austin; [30.26° N, 97.74° W]; 28 Sep.–10 Oct. 1975; Bob Henderson leg.; TAMU-ENTO X1398967, X1399473 • 1 ♀; Travis Co., Austin, Barton Creek Park; [30.26° N, 97.82° W]; 8 Oct. 1983; L.E. Carroll leg.; TAMU-ENTO X1401177 • 5 ♂♂; ibid.; 8 Oct. 1983; L.E. Carroll leg.; TAMU-ENTO X1385439, X1386250, X1388246, X1400810, X1401157 • 2 ♀♀; Travis Co., Austin, Zilker Park; [30.27° N, 97.77° W]; 8 Oct. 1983; J.B. Woolley leg.; TAMU-ENTO X1415630, X1418344 • 2 ♂♂; ibid.; 8 Oct. 1983; J.B. Woolley leg.; TAMU-ENTO X1314578, X1316273 • 1 ♀; Travis Co., Heep Farm, 11 mi. S of Austin; [30.11° N, 97.8° W]; 27 Jul. 1972; E.E. Grissell leg.; TAMU-ENTO X1362335 • 1 ♀; Uvalde Co., Garner State Park; [29.59° N, 99.74° W]; 22 May 1979; W.P. Foerster leg.; TAMU-ENTO X1418161 • 1 ♀; Val Verde Co., Dolan Creek Camp Ground; 29.9° N, 100.8833° W; 14 Jun. 1975; J.S. Ashe leg.; TAMU-ENTO X1365716 • 1 ♀; Walker Co., Ellis Prison; [30.88° N, 95.46° W]; 23 Jul. 1977; W.L. Sterling leg.; TAMU-ENTO X1396331 • 1 ♀; ibid.; 25 Jul. 1977; W.L. Sterling leg.; TAMU-ENTO X1396334 • 5 ♀♀; Williamson Co., CR 278 3.5 mi. SE of Liberty Hill; [30.63° N, 97.89° W]; 9 Sep. 2017; Karen W. Wright leg.; TAMU-ENTO X1519447, X1519897, X1519918, X1519935, X1520149 • 1 ♀; Williamson Co., Highway 183 4 mi. N of Leander; [30.64° N, 97.87° W]; 9 Sep. 2017; Karen W. Wright leg.; TAMU-ENTO X1536161 • 1 ♂; 10 miles east of Sonora; [30.57° N, 100.48° W]; 25 Jul. 1986; W.F. Chamberlain leg.; ex Rhus; TAMU-ENTO X1387486 • 1 ♀; 10 miles NW of Ballinger; [31.84° N, 100.07° W]; 15 Jul. 1993; G.M. Chamberlain leg.; TAMU-ENTO X1388634 • 1 ♀; 10 miles S of Austin; [30.11° N, 97.74° W]; 28 Sep. 1965; George E. Bohart leg.; BBSL700645 • 1 ♂; ibid.; 28 Sep. 1965; George E. Bohart leg.; BBSL700808 • 1 ♀; 10 miles west of Port Mansfield; [26.55° N, 97.59° W]; 9 Mar. 2001; W.F. Chamberlain leg.; TAMU-ENTO X1488468 • 3 ♀♀; 11 miles SW of Kerrville; [29.93° N, 99.27° W]; 21 May 1989; W.F. Chamberlain leg.; TAMU-ENTO 1395661, X1313963, X1370712 • 1 ♂; ibid.; 21 May 1989; W.F. Chamberlain leg.; TAMU-ENTO X1409734 • 1 ♀; 13 miles NE of Fredricksburg; [30.41° N, 98.72° W]; 20 Apr. 1997; W.F. Chamberlain leg.; TAMU-ENTO X1400248 • 1 ♀; 5 miles east of Lampasas; [31.06° N, 98.1° W]; 21 Apr. 1997; W.E. Chamberlain leg.; TAMU-ENTO X1393218 • 1 ♀; 5 miles north of Hondo; [29.42° N, 99.14° W]; 8 May 1988; W.F. Chamberlain leg.; TAMU-ENTO X1417885 • 1 ♀; 5 miles north of Utopia; [29.69° N, 99.53° W]; 22 Apr. 1994; W.F. Chamberlain leg.; TAMU-ENTO X1403909 • 1 ♀; 6 miles east of Doss; [30.44° N, 99.03° W]; 27 Aug. 1995; W.F. Chamberlain leg.; TAMU-ENTO X1400060 • 1 ♀; 9 miles SE of Hondo; [29.25° N, 99.04° W]; 17 Mar. 1994; W.F. Chamberlain leg.; TAMU-ENTO X1420319 • 1 ♂; Austin; [30.26° N, 97.74° W]; 11 Sep. 1975; W.P. Foerster leg.; TAMU-ENTO X1365304 • 1 ♀; ibid.; 25 Mar. 1976; W.P. Foerster leg.; TAMU-ENTO X1401264 • 1 ♀; ibid.; 17 May 1976; W.P. Foerster leg.; TAMU-ENTO X1398517 • 1 ♀; ibid.; 14 Jun. 1976; W.P. Foerster leg.; ex dandelion; TAMU-ENTO X1365981 • 1 ♂; ibid.; 27 Jun. 1976; W.P. Foerster leg.; ex Gaillardia; TAMU-ENTO X1399165 • 1 ♀; ibid.; 1 Jul. 1976; W.P. Foerster leg.; ex cult. camphor; TAMU-ENTO X1365509 • 1 ♂; ibid.; 5 Jul. 1976; W.P. Foerster leg.; TAMU-ENTO X1365939 • 1 ♀; Enchanted Rock; 30.51° N, 98.82° W; 20 May 2015; Paul A. Rude leg.; EMEC • 1 ♀; Kerrville; [30.05° N, 99.14° W]; 18 Jul. 1981; W.F. Chamberlain leg.; TAMU-ENTO X1387184 • 1 ♀; ibid.; 13 Sep. 1996; G.M. Chamberlain leg.; TAMU-ENTO X1422788 • 1 ♀; ibid.; 27 May 1999; G.M. Chamberlain leg.; TAMU-ENTO X1400002 • 1 ♀; New Braunfels; [29.7° N, 98.12° W]; 23 Apr. 1977; W.F. Chamberlain leg.; TAMU-ENTO X1388564 • 1 ♂; Presidio; [29.56° N, 104.37° W]; 4 May 1979; W.F. Chamberlain leg.; TAMU-ENTO X1312628 • 1 ♀; Ratcliff Lake Recreation Area; [31.39° N, 95.16° W]; 6 Jun. 1994; W.F. Chamberlain leg.; TAMU-ENTO X1415799 • 1 ♂; Taylor; [30.57° N, 97.41° W]; 16 Sep. 1933; J.E. Gillaspy leg.; TAMU-ENTO X1387673 • 9 ♂♂; Texas A&M Campus, College Station; [30.62° N, 96.34° W]; 29 Jul. 1995; John W. Wenzel leg.; PCYU • 1 ♂; Waco; [31.55° N, 97.15° W]; 10 Aug. 1939; P.A. Glick leg.; ex cotton; TAMU-ENTO X1386262 .

Note: During final preparation of the manuscript some Montana specimens were discovered in MTEC.

Range

Chihuahuan Desert and western Great Plains from eastern Colorado to northern Veracruz, rarely occurring in western Colorado and New Mexico (Fig. 11).

Floral hosts

ANACARDIACEAE R.Br.: Rhus L.: R. trilobata Nutt. • APOCYNACEAE Juss.: Asclepias L.: A. verticillata L. • AQUIFOLIACEAE Bercht. & J.Presl: Ilex L.: I. vomitoria Aiton • ASPARAGACEAE Juss.: Dasylirion Zucc.: D. wheeleri S.Watson ex Rothr. • ASTERACEAE: Aster L. • Baccharis L. • Bradburia Torr. & A.Gray: B. pilosa (Nutt.) Semple • Carduus L. • Coreopsis L.: C. basalis (A.Dietr.) S.F.Blake • C. tinctoria Nutt. • Croptilon Raf.: C. divaricatum (Nutt.) Raf. • Gaillardia Foug. • Helianthus L.: H. annuus L. • Heterotheca Cass.: H. subaxillaris (Lam.) Britton & Rusby • H. viscida (A.Gray) V.L. Harms • Lactuca L.: L. sativa L. • Parthenium L.: P. hysterophorus L. • Pyrrhopappus DC.: P. pauciflorus (D.Don) DC. • Santolina L.• BRASSICACEAE Burnett: Lepidium L.: L. integrifolium Nutt. • CACTACEAE Juss.: Astrophytum Lem.: A. myriostigma Lem. • A. ornatum (DC.) Britton & Rose • Leuchtenbergia Hook.: L. principis Hook. • Opuntia Mill.: O. macrorhiza Engelm. • CLEOMACEAE Bercht. & J.Presl: Polanisia Raf.: P. dodecandra (L.) DC. • EUPHORBIACEAE Juss.: Croton L.: C. capitatus Michx. • Euphorbia L. • FABACEAE Lindl.: Cassia L. • Cercidium Tul.: C. macrum I.M.Johnst. • Medicago L.: M. sativa L. • Melilotus (L.) Mill.: M. officinalis (L.) Lam.: M. o. subsp. alba (Medik.) H.Ohashi & Tateishi • LAMIACEAE Martinov: Lamium L. • Marrubium L.: M. vulgare L. • Mentha L.: M. spicata L. • Monarda L. • Salvia L.: S. farinacea Benth. • MALVACEAE Juss.: Gossypium L. • PAPAVERACEAE Juss.: Argemone L.: A. albiflora Hornem. • PEDALIACEAE R.Br.: Sesamum L.: S. indicum L. • SCROPHULARIACEAE Juss.: Leucophyllum Bonpl.: L. frutescens (Berland.) I.M.Johnst. • TAMARICACEAE Link: Tamarix L.

Additional records from Moure & Hurd (1987): ASTERACEAE: Erigeron L.: E. philadelphicus L. • Gutierrezia Lag.: G. texana (DC.) Torr. & A.Gray: G. t. var. glutinosa (S.Schauer) M.A.Lane • BORAGINACEAE Juss.: Phacelia Juss.: P. congesta Hook. • PAPAVERACEAE: Argemone: A. sanguinea Greene.

DNA barcodes

Eight confirmed sequences available, two haplotypes (BOLD process IDs: BWTWO1314-10, DLII051- 06, LASNA016-08 (haplotype 1); DIAL1132-07, DLII307-07, DLII318-07, DLII338-07, DLII547- 07 (haplotype 2)). Haplotype 1 is very similar to L. carlinvillense, and haplotype 2 is very similar to L. tegulare . There is a large amount of divergence within these sequences (3.36% maximum intraspecific p-distance). They are closest in terms of p-distance to L. carlinvillense (0.15% minimum interspecific p-distance). No fixed nucleotide substitutions distinguish all L. coactus from all other Nearctic species of the L. gemmatum complex, but one fixed substitution distinguishes haplotype 1, and two fixed substitutions distinguish haplotype 2: 156(C) (haplotype 1); 429(G) and 441(C) (haplotype 2) (Supp. file 2).