Triepeolus segregatus ( Cockerell, 1900 )

Onuferko, Thomas M. & Rightmyer, Molly G., 2024, A revision of the simplex species group of the cleptoparasitic bee genus Triepeolus Robertson, 1901 (Hymenoptera: Apidae), European Journal of Taxonomy 950 (1), pp. 1-106 : 75-87

Triepeolus segregatus ( Cockerell, 1900)

Figs 12N , 15E, 20B, 29

Epeolus occidentalis var. segregatus Cockerell, 1900: 361 (♀, ♂).

Epeolus permixtus Cockerell, 1923: 94 (♀, ♂), syn. nov.

Triepeolus brunnescens Cockerell & Sandhouse, 1924: 313 (♂), syn. nov.

Triepeolus pacis Cockerell, 1925b: 201 (♂), syn. nov.

Epeolus sarothrinus Cockerell, 1929: 103 (♀, ♂), syn. nov.

Epeolus sarothrinus var. confluens Cockerell, 1929: 103 (♂), syn. nov.

Triepeolus segregatus – Cockerell 1904: 38; 1907: 63 (redescription).

Doeringiella (Triepeolus) sarothrinus – Vergara & Ayala 2002: 24 (in regional checklist).

Proposed common name

Segregated triepeolus.

Diagnosis

The following morphological features in combination tell T. segregatus apart from all other Triepeolus in the simplex species group: the axillae do not have pale pubescence all along their margins and are small to intermediate in size, each with the tip extending little if at all beyond the midlength of the mesoscutellum and the free portion distinctly pointed but mesally unattached to the mesoscutellum for less than 2/5 the medial length of the axilla ( Figs 20B , 29B ); the mesopleura laterally do not have erect/suberect, simple setae or have only sparse, short (<¼ MOD), erect/suberect, simple setae in addition to the usual appressed, branched setae ( Fig. 29A, C ); the T1 discal patch is trapezoidal with an inverted U- or horseshoe-shaped anteromedial projection, with the basal band so deeply invaginated that it appears to be completely interrupted medially in dorsal view ( Figs 15E , 29B ); and the T2 apical transverse band has a pair of well-defined basomedially convergent anterolateral extensions ( Figs 15E , 29A–C ). Specimens from Texas and adjacent Mexico may be confused with T. parahirsutus sp. nov. and specimens from California with T. saturninus due to their similar size and trapezoidal discal patch with an inverted U- or horseshoe-shaped anteromedial projection, but in the two similar-looking species the mesopleura have sparse, pale-yellow, erect/suberect, simple setae (reaching about ½ MOD in length) in addition to the usual appressed, branched setae.

Etymology

The specific epithet is a Latin adjective meaning ‘set apart’, presumably in reference to Cockerell’s (1900) recognition of the holotype as a separate variety that “[d]iffers from typical [E.] occidentalis from Colorado ”, with which it was mistakenly considered to be conspecific.

Material examined

Primary type material

MEXICO – Baja California • ♂, E. permixtus holotype; Isla Pond bay, Isla Ángel de la Guarda , Gulf of California ; 1 Jul. 1921; E.P. Van Duzee leg.; CAS 954 . – Baja California Sur • ♂, T. pacis holotype; La Paz; 29 Jun. 1919; G.F. Ferris leg.; CAS 1661 .

USA – California • ♂, T. brunnescens holotype; Poway , San Diego County; 10 Sep. 1884; F.E. Blaisdell leg.; CAS 1606 • ♀, E. sarothrinus holotype; Riverside , Riverside County; 26 May 1928; P.H. Timberlake leg.; AMNH AMNH_IZC 00323550 • ♂, E. sarothrinus var. confluens holotype; same data as for preceding; 26 May 1928; P.H. Timberlake leg.; USNM 534623 . – New Mexico • ♂, E. occidentalis var. segregatus holotype; Las Vegas Hot Springs ; 11 Jul.; USNM 534868 .

Secondary type material

USA • 1 ♀, E. sarothrinus paratype; California, Riverside , Riverside County; 6 Jul. 1928; P.H. Timberlake leg.; AMNH M.G.R. Database No. 1516 .

DNA barcoded material

Available. Three BINs in two clusters:

Cluster: BOLD:AAG7909+BOLD:AFP1028. BIN: BOLD:AAG7909. Specimen examined and sequenced:

USA • 1 ♀; California, Chollas Creek Park (San Diego), San Diego County; 32.7201° N, 117.0780° W; 15 Jul. 2011; K.L.J. Hung leg.; BOLD sample ID: KJH.4621 ; UCR .

Cluster: BOLD:AAG7909+BOLD:AFP1028. BIN: BOLD:AFP1028. Specimens examined and sequenced:

USA – Arizona • 1 ♀ (previously assigned to the BIN BOLD: AAG7909 ); Sierra Vista , Cochise County; 31.5100° N, 110.2590° W; 4 Sep. 2009; C.S. Sheffield and S. Dumesh leg.; BOLD sample ID: CMNTO_025 ; PCYU PCYU-SD09:631 • 1 ♂, T. cf. segregatus (previously assigned to the BIN BOLD: AAG7909 ); Southwestern Research Station (5 mi W of Portal ), Cochise County; 22 Aug. 2007; H.T. Ngo and J. Gibbs leg.; BOLD sample ID: CMNTO_485 ; PCYU • 2 ♂♂, T. cf. segregatus ( BIN previously unassigned); same data as for preceding; 22 Aug. 2007; H.T. Ngo and J. Gibbs leg.; BOLD sample IDs: CMNTO_487 , CMNTO_488 ; PCYU .

Cluster: BOLD:AAG7909+BOLD:AFP1028. BIN unassigned:

USA – Arizona • 1 ♂, T. cf. segregatus ; Southwestern Research Station (5 mi W of Portal ), Cochise County; 22 Aug. 2007; H.T. Ngo and J. Gibbs leg.; BOLD sample ID: CMNTO_489 ; PCYU • 1 ♂; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8830° N, 109.2070° W; 21 Aug. 2019; J. Gibbs leg.; BOLD sample ID: WRME0501356 ; WRME WRME0501356 .

Cluster and BIN: BOLD:AEI6013. Specimens examined and sequenced:

USA – Arizona • 1 ♂; 19 mi SE of Willcox , Cochise County; 18–25 Aug. 2002; BOLD sample ID: CCDB-38770 F09 ; RSKM RSKM_ENT_E-168248 • 1 ♂; Catalina Hwy ( Santa Catalina Mountains ), Coronado National Forest ; 32.3631° N, 110.7137° W; 16 Aug. 2015; T.M. Onuferko leg.; BOLD sample ID: CMNTO_047 ; TOPC • 1 ♂; Catalina Hwy ( Santa Catalina Mountains ), Coronado National Forest ; 32.3728° N, 110.7068° W; 16–26 Aug. 2015; T.M. Onuferko leg.; BOLD sample ID: CMNTO_109 ; PCYU • 1 ♀; Southwestern Research Station (5 mi W of Portal ), Cochise County; 1 Sep. 2013; BOLD sample ID: CMNTO_017 ; PCYU • 1 ♀; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8840° N, 109.2060° W; 25 Aug. 2011; N. de Silva leg.; BOLD sample ID: 14511G09- AZ ; PCYU 0003776 • 1 ♀; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8852° N, 109.2051° W; 24 Aug. 2017; R. Oram leg.; BOLD sample ID: CCDB-38770 F11 ; RSKM RSKM_ENT_E-167432 . – Colorado • 1 ♀; Escalante State Wildlife Area , Sawmill Mesa Road , Delta County; 38.7361° N, 108.1556° W; 31 Aug. 2013; M. Buck leg.; BOLD sample ID: CMNTO_314 ; RAM pmae00122773 . – New Mexico • 1 ♀; 7 km NE of Silver City, Grant County; 32.8270° N, 108.2410° W; 2 Sep. 2009; C.S. Sheffield and S. Dumesh leg.; BOLD sample ID: CMNTO_019 ; PCYU PCYU-SD09:289 • 1 ♀; 28 mi S of Animas , Hidalgo County; 31.5550° N, 108.8760° W; 27 Aug. 2019; J. Gibbs leg.; BOLD sample ID: CMNTO_162 ; WRME WRME0500765 • 1 ♀; NM-152 (along Iron Creek ), Gila National Forest , Grant County; 32.9078° N, 107.8160° W; 18 Aug. 2015; T.M. Onuferko leg.; BOLD sample ID: CMNTO_050 ; TOPC.

Non-barcoded material

MEXICO – Baja California • 1 ♀; 1 mi E of Santa Inés ; 6 Apr. 1985; Bloomfield and Faulkner leg.; INHS 387489 • 1 ♂; 3.3 mi S of El Cóndor ; 1 Sep. 1981; D.K. Faulkner leg.; SDNHM • 1 ♀; 4 mi S of El Cóndor ; 28 May 1981; Faulkner and Brown leg.; SDNHM • 1 ♀; 8 mi E of Tecate on Hwy 2 ; 30 Jun. 1980; Faulkner and Brown leg.; SDNHM • 1 ♀; 19 mi E of Ojos Negros ; 21 Jul. 1980; Faulkner and Brown leg.; SDNHM • 9 ♀♀; Agua Caliente ( San Carlos ), 18.5 km E of Maneadero ; 6 Jul. 1973; P.H. Arnaud, Jr. leg.; CAS • 2 ♀♀; El Pabellón ; 2 Jul. 1968; J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU • 1 ♀; Los Mártires ; 24 Jun. 1968; N. Leppla, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU • 1 ♀; Río San Salvador at Hwy 3; 31.8733° N, 116.0908° W; 7–12 Jun. 1997; M.E. Irwin and E.I. Schlinger leg.; INHS 387491 • 19 ♀♀; San Ángel ; 28 Jun. 1968; N. Leppla, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU • 1 ♀; same data as for preceding; 28 Jun. 1968; N. Leppla, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU M.G.R. Database No. 1215 • 1 ♀; Santa Ana ; 21 Jun. 1968; N. Leppla, J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU • 1 ♀; Sierra de Juárez (6 mi N of Laguna Hanson ); 21 Jul. 1980; Faulkner and Brown leg.; SDNHM . – Baja California Sur • 1 ♀; 2 mi S of La Paz ; 2 Aug. 1966; J.A. Chemsak leg.; EMEC • 1 ♀; 11 mi N of San Isidro ( La Purísima ); 3 Apr. 1985; Bloomfield and Faulkner leg.; SDNHM • 1 ♀; 36 km NE of San José del Cabo ; 8 Mar. 1988; W.J. Pulawski leg.; CAS • 1 ♂; Arroyo San Bartolo ; 28 Aug. 1982; D.K.F. leg.; SDNHM • 1 ♀; Loreto 21 Jun. 1983; R. Miller and L. Stange leg.; FSCA • 1 ♀; Playa Los Cerritos (11.2 mi S of Todos Santos); 28 Sep. 1981; F. Andrews and D. Faulkner leg.; SDNHM • 3 ♀♀; Puerto Escondido ; 15 Jun. 1992; R. Shaver leg.; UCBME • 1 ♀; S of El Rosario on Hwy 1 at km 115 marker; 24–25 Aug. 1992; D.E. Russell leg.; UCBME • 1 ♀; San Ignacio ; 26 Jun. 1938; Michelbacher and Ross leg.; CAS • 2 ♀♀; San José Viejo ; 17 Jul. 1968; J. Davidson, J. Bigelow, M. Bentzien, W. Fox, S. Williams, and M. Cazier leg.; ASU . – Chihuahua • 1 ♀; 10–15 km S of Maycoba ; 29 Aug. 1984; W.J. Pulawski leg.; CAS . – Coahuila • 2 ♀♀; 61 mi N of Saltillo ; 11 Aug. 1959; A.S. Menke and L.A. Stange leg.; UCBME • 1 ♀; Guadalupe ; 24 May 1952; M. Cazier, W. Gertsch, and R. Schrammel leg.; AMNH • 1 ♀; Rancho Las Lilas ( Reserva de la Biosfera Mapimí ); 26.7372° N, 103.7542° W; 12 Jun. 1995; L. Godínez leg.; UNAM LG-1095 7400 . – Durango • 1 ♂; Navajos (20 mi E of El Salto ); 23 Jul. 1964; L.A. Kelton leg.; CNC 808392 . – Guanajuato • 2 ♀♀; León ; 16 Aug. 1953; C. and P. Vaurie leg.; AMNH . – Jalisco • 1 ♀; Guadalajara ; Crawford leg.; USNM . – Michoacán • 1 ♀; Cotija ; 6 Sep. 1970; B.L. Villegas leg.; UCBME . – Nuevo León • 2 ♀♀; 50 mi SE of Monterrey ; 13 Oct. 1957; H.A. Scullen leg.; OSAC . – Sonora • 1 ♀; Bahia San Pedro, Gulf of California ; 7 Jul. 1921; E.P. Van Duzee leg.; CAS .

USA – Arizona • 1 ♀; 0.5 mi W of Hwy 92 on Ash Canyon Road ( Huachuca Mountains ), Cochise County; 13–22 Sep. 1993; N. McFarland leg.; CNC 1801913 • 1 ♀; 1 mi S of Portal ( Chiricahua Mountains ), Cochise County; 26 Aug. 1966; M.A. Cazier leg.; ASU • 1 ♀; 11 mi SW of Eloy , Pinal County; 19 Jun. 1953; T.R. Haig leg.; UCBME • 1 ♀; 12 km NE of Tombstone , Cochise County; 31.8460° N, 110.0030° W; 7 Sep. 2009; C.S. Sheffield and S. Dumesh leg.; PCYU PCYU-SD09:1064 • 1 ♀; 12 mi N of Yuma ; 8 Oct. 1958; T.R. Haig leg.; UCBME • 2 ♀♀; 15 mi E of Nogales ; 22 Sep. 1963; V.L. Vesterby leg.; UCBME • 7 ♀♀; 15 mi W of Chandler ; 9 Jun. 1961; G.D. Butler leg.; UAIC • 1 ♀; 15 mi W of Flagstaff on Hwy 66; 8 Sep. 1964; F.M. and M.C. Hull leg.; CNC 753870 • 1 ♀; 2 mi E of Apache , Cochise County; 3 Sep. 2003; J.G. Rozen and J.S. Ascher leg.; AMNH • 1 ♀; 2 mi N of Paradise , Cochise County; 9 Aug. 1967; F.G. Andrews leg.; UCR UCRC ENT 57283 • 1 ♀; 2.5 mi NE of Portal , Cochise County; 9 Aug. 1959; M. Statham leg.; AMNH • 1 ♀; 21 mi W of Portal , Cochise County; 15 Aug. 1970; R.F. Denno leg.; UCBME • 1 ♀; 2–10 mi S of Williams , Coconino County; 11 Aug. 1994; J.G. and B.L. Rozen leg.; AMNH • 1 ♂; 23 mi SE of Sonoita , Santa Cruz County; 27 Aug. 1968; R.W. Thorp leg.; UCBME • 1 ♀; 3 mi N of Portal , Cochise County; Aug. 2005; L. Packer leg.; PCYU PCYU-NA: 823 • 1 ♀; 3 mi S of Portal , Cochise County ( Chiricahua Mountains ); 13 Aug. 1965; C.A. Saario leg.; UCR UCRC ENT 57290 • 4 ♀♀; 3 mi W of Salome , La Paz County; 21 Jun. 1971; G. Bohart and P. Torchio leg.; BBSL BBSL522493 to BBSL522496 • 1 ♀; 4 mi N of Grand Canyon Junction ; 12 Jul. 1956; F.G. Werner leg.; UAIC • 1 ♀; 4 mi SW of Portal ( Chiricahua Mountains ), Cochise County; 16 Aug. 1973; Cazier family leg.; ASU • 1 ♀; 4 mi W of Paradise ( Chiricahua Mountains ), Cochise County; 9 Sep. 1950; W. Gertsch and M. Cazier leg.; AMNH • 1 ♀; 5 mi E of Fort Apache ; 28 Aug. 1964; E.I. Schlinger leg.; UCR UCRC ENT 57378 • 4 ♀♀; same data as for preceding; 28 Aug. 1964; M.E. Irwin leg.; UCR UCRC ENT 57397 to UCRC ENT 57400 • 1 ♀; 5.5 mi W of Peña Blanca , Santa Cruz County; 17 Aug. 1961; Werner and Bequaert leg.; UAIC • 1 ♀; 5803 North US Hwy 89, Flagstaff , Coconino County; 25 Jul. 1988; P.H. Arnaud, Jr. leg.; CAS • 1 ♀; 6 mi N of Pearce ; 6 Aug. 1955; G.D. Butler and Z. Noon leg.; UAIC • 1 ♀; 6 mi SW of Apache , Cochise County; 1 Sep. 2003; J.G. Rozen and J.S. Ascher leg.; AMNH • 1 ♀; 8 mi N of Flagstaff ; 15 Sep. 1955; G.D. Butler leg.; UAIC • 1 ♀; 8 mi N of Onion Saddle ( Chiricahua Mountains ), Cochise County; 22 Aug. 1986; McGinley and Norden leg.; USNM • 1 ♀; 8 mi S of Toltec , Pinal County; 24 Jun. 1953; T.R. Haig leg.; UCBME • 1 ♀; 9 mi N of Elfrida , Cochise County; 5 Aug. 1961; J.G. Rozen leg.; AMNH M.G.R. Database No. 1210 • 1 ♀; Benson ; 4 Jul. 1961; P.H. Johnson leg.; UAIC • 1 ♀; Boyce Thompson Arboretum ( Superior ); 15 May 1954; G.D. Butler leg.; UAIC • 2 ♀♀; Buckeye ; 11 Jun. 1954; G.D. Butler leg.; UAIC • 1 ♀; same data as for preceding; 12 Aug. 1954; G.D. Butler leg.; UAIC • 1 ♀; same data as for preceding; 27 May 1955; G.D. Butler leg.; UAIC • 1 ♀; Carr Canyon summit campground, Cochise County; 1–8 Aug. 1993; M. Sharkey leg.; CNC PCYU:CNC-Bee: 509 • 4 ♀♀; Carrizo , Gila County; 1 Sep. 1986; A.S. Menke leg.; USNM • 1 ♀; Casa Grande ; 20 Jun. 1956; C. Williams leg.; UAIC • 1 ♀; Arizona, Cave Creek Canyon ; 14 Aug. 1981; A.J. Gilbert and N.J. Smith leg.; UCBME • 1 ♀; Chiricahua Mountains , Cochise County; 31 Jul. 1946; H.A. Scullen leg.; OSAC • 1 ♂; Cienega Spring (near Portal ), Cochise County; 21 Aug. 1993; L. Packer, T. Pare, and M. Sharkey leg.; PCYU • 1 ♀; Cochise County; Nov. 1969; D. Schuh leg.; FSCA • 1 ♀; Double Adobe , Cochise County; 20 Aug. 1989; R.M. Bohart leg.; UCBME • 1 ♀; E of Box Canyon , Pima County; 31.7913° N, 110.7457° W; 15 Sep. 1994; T. Griswold leg.; BBSL BBSL582211 • 1 ♀; Eloy ; 14 Jun. 1955; G.D. Butler leg.; UAIC • 1 ♀; Flagstaff ; 22 Jul. 1934; F.E. Lutz leg.; AMNH • 1 ♀; Garden Canyon , Cochise County; 17 Aug. 2014; R. Parks leg.; CSUC • 1 ♀; Grand Canyon ; 18 Aug. 1939; E.C. Van Dyke leg.; CAS • 1 ♀; Grand Canyon ( South Rim ); 31 Jul. 1934; E.L. Bell leg.; AMNH • 1 ♀; Jhus Canyon , Cochise County; 11 Aug. 1967; F.G. Andrews leg.; UCR UCRC ENT 57406 • 2 ♀♀; Kansas Settlement; 23 Jun. 1954; G.D. Butler leg.; UAIC • 1 ♀; Madera Canyon ( Santa Rita Mountains ); “ 9-11-11 ”; J. Cowles leg.; SDNHM • 1 ♀; Madera Canyon ( Santa Rita Mountains ), Pima County; 13 Aug. 1952; M. Cazier, R. Schrammel, and C. and P. Vaurie leg.; AMNH • 1 ♀; Madera Canyon ( Santa Rita Mountains ), Santa Cruz County; 10 Aug. 1965; M.R. Irwin leg.; UCR UCRC ENT 57285 • 1 ♀; Marana ; 28 Sep. 1955; G.D. Butler leg.; UAIC • 2 ♀♀; same data as for preceding; 7 Jun. 1962; M. Wargo leg.; UAIC • 3 ♀♀; Maricopa ; 20 Jun. 1956; G.D. Butler leg.; UAIC • 1 ♀; same data as for preceding; 12 Jun. 1962; G.D. Butler leg.; UAIC • 3 ♀♀; Mesa ; 9 Jun. 1955; G.D. Butler leg.; UAIC • 1 ♂; Mount Wrightson ( Santa Rita Mountains ), Santa Cruz County; 28 Aug. 1962; H.E. Milliron leg.; CNC 808389 • 1 ♀; Oak Creek Canyon , Coconino County; 2 Sep. 1968; D.R. Miller and J.E. Lauck leg.; UCBME • 1 ♀; Onion Saddle ( Chiricahua Mountains ), Cochise County; 15 Aug. 1986; McGinley and Norden leg.; USNM • 2 ♀♀; Onion Saddle , Cochise County; 31.9334° N, 109.2631° W; 26 Aug. 2016; L. Packer leg.; PCYU • 1 ♂; Paradise , Cochise County; 31.9333° N, 109.2167° W; 11–16 Aug. 2008; W. Steiner and J.M. Swearingen leg.; BIML USGS-DRO 147147 • 1 ♀; Pinery Canyon (10 mi NW of Onion Saddle ) ( Chiricahua Mountains ), Cochise County; 16 Aug. 1965; K. Brown leg.; UCR UCRC ENT 57272 • 1 ♀; Ramsey ( Huachuca Mountains ); 30 Aug. 1953; G.D. Butler leg.; UAIC • 1 ♀; Ramsey Canyon (15 mi S of Sierra Vista ), Huachuca Mountains ; 15 Oct. 1969; R.F. Sternitzky leg.; CNC 753862 • 1 ♀; Roll ; 2 Jun. 1955; G.D. Butler leg.; UAIC • 1 ♀; Salt River Canyon at Hwy 60; 11 Jul. 1965; S.A. Gorodenski leg.; ASU • 1 ♀; Sierra Vista , Cochise County; 31.5100° N, 110.2590° W; 4 Sep. 2009; C.S. Sheffield and S. Dumesh leg.; PCYU PCYU-SD09:614 • 2 ♀♀; Skeleton Canyon ( Peloncillo Mountains ), Cochise County; 24 Aug. 1962; H.V. Weems, Jr. leg.; FSCA • 1 ♀; same data as for preceding; 8 Sep. 1958; H.V. Weems, Jr. leg.; FSCA • 1 ♀; Sonoita, Santa Cruz County; 19 Aug. 1986; R.M. Bohart leg.; UCBME • 3 ♀♀; Southwestern Research Station (5 mi W of Portal ), Cochise County; 1 Aug. 1956; C. and M. Cazier leg.; AMNH M.G.R. Database Nos 3763 , 3765 , 3771 • 1 ♀; same data as for preceding; 1 Aug. 1956; E. Ordway leg.; AMNH M.G.R. Database No. 3772 • 1 ♀; same data as for preceding; 2 Aug. 1956; C. and M. Cazier leg.; AMNH M.G.R. Database No. 3770 • 1 ♀; same data as for preceding; 2 Aug. 1956; E. Ordway leg.; AMNH M.G.R. Database No. 3767 • 1 ♀; same data as for preceding; 4 Aug. 1956; E. Ordway leg.; AMNH M.G.R. Database No. 3762 • 1 ♀; same data as for preceding; 10 Aug. 1956; C. and M. Cazier leg.; AMNH M.G.R. Database No. 3768 • 1 ♀; same data as for preceding; 21 Aug. 1956; C. and M. Cazier leg.; AMNH M.G.R. Database No. 3774 • 1 ♀; same data as for preceding; 17 Sep. 1956; E. Ordway leg.; AMNH M.G.R. Database No. 3766 • 2 ♀♀; same data as for preceding; 26 Sep. 1956; E. Ordway leg.; AMNH M.G.R. Database Nos 3764 , 3769 • 1 ♀; same data as for preceding; 13 Aug. 1958; P.D. Hurd leg.; UCBME • 1 ♀; same data as for preceding; 17 Aug. 1958; E.G. Linsley leg.; UCBME • 1 ♀; same data as for preceding; 19 Aug. 1958; E.G. Linsley leg.; UCBME • 2 ♀♀; same data as for preceding; 19 Aug. 1958; P.D. Hurd leg.; UCBME • 1 ♀; same data as for preceding; 26 Aug. 1958; E.G. Linsley leg.; UCBME • 1 ♀; same data as for preceding; 27 Aug. 1958; H.V. Weems, Jr. leg.; FSCA • 1 ♀; same data as for preceding; 25 Aug. 1960; P.H. Arnaud, Jr. and D.C. Rentz leg.; CAS • 1 ♀; same data as for preceding; 3 Sep. 1962; J.G. Rozen and S.J. Hessel leg.; AMNH • 1 ♀; same data as for preceding; 26 Aug. 1964; J.G. and B.L. Rozen leg.; AMNH M.G.R. Database No. 3773 • 1 ♀; same data as for preceding; 18 Sep. 1966; P.H. Arnaud, Jr. leg.; CAS • 1 ♀; same data as for preceding; 20 Sep. 1966; P.H. Arnaud, Jr. leg.; CAS • 2 ♀♀; same data as for preceding; 21 Aug. 1973; M. Favreau leg.; AMNH • 4 ♀♀; same data as for preceding; 22 Aug. 1973; M. Favreau leg.; AMNH • 2 ♀♀; same data as for preceding; 24 Aug. 1973; M. Favreau leg.; AMNH • 2 ♀♀; same data as for preceding; 26 Aug. 1973; Rozen and Favreau leg.; AMNH • 1 ♀; same data as for preceding; 27 Aug. 1973; J.G. Rozen leg.; AMNH • 1 ♀; same data as for preceding; 1 Sep. 1973; M. Favreau and R. McGinley leg.; AMNH • 1 ♀; same data as for preceding; 21 Aug. 1985; J.G. Rozen leg.; AMNH • 1 ♀; same data as for preceding; 18 Aug. 1995; J.G. Rozen and S.A. Budick leg.; AMNH • 1 ♂; same data as for preceding; 2003; L. Packer leg.; PCYU • 1 ♀; same data as for preceding; 17 Aug. 2004; J.S. Ascher leg.; AMNH • 1 ♂; same data as for preceding; 27 Aug. 2004; PCYU • 1 ♀, 1 ♂; same data as for preceding; 24 Aug. 2007; PCYU • 1 ♂; same data as for preceding; 25 Aug. 2007; PCYU • 2 ♀♀; same data as for preceding; 23 Aug. 2011; K.L.J. Hung leg.; SDNHM • 4 ♀♀; same data as for preceding; 25 Aug. 2011; K.L.J. Hung leg.; SDNHM • 1 ♀; same data as for preceding; 26 Aug. 2013; PCYU • 1 ♀; same data as for preceding; 27 Aug. 2013; PCYU • 3 ♀♀; same data as for preceding; 1 Sep. 2013; PCYU • 4 ♂♂; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8830° N, 109.2070° W; 26 Aug. 2019; J. Gibbs leg.; WRME WRME0500630 , WRME0500632 to WRME0500634 • 3 ♂♂; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8840° N, 109.2060° W; 22 Aug. 2011; N. de Silva leg.; PCYU 0005889 to 0005891 • 1 ♂; same data as for preceding; 25 Aug. 2011; N. de Silva leg.; PCYU 0003777 • 4 ♂♂; Southwestern Research Station (5 mi W of Portal ), Cochise County; 31.8852° N, 109.2051° W; 22 Aug. 2017; L. Graham leg.; PCYU • 2 ♂♂; same data as for preceding; 23 Aug. 2017; R. Oram leg.; RSKM RSKM_ENT_E-167429 , RSKM_ENT_E-167430 • 2 ♂♂; same data as for preceding; 24 Aug. 2017; R. Oram leg.; RSKM RSKM_ENT_E-167427 , RSKM_ENT_E-167431 • 3 ♂♂; same data as for preceding; 25 Aug. 2017; L. Graham leg.; PCYU • 1 ♀; Sunnyslope ; 26 May 1954; G.D. Butler leg.; UAIC • 1 ♀; Arizona, Sycamore Canyon (near Ruby); 16–17 Aug. 1961; Werner and Bequaert leg.; UAIC • 2 ♀♀; Tucson , Pima County; 19 May 1954; G.D. Butler leg.; UAIC • 1 ♀; same data as for preceding; 17 Jun. 1955; G.D. Butler leg.; UAIC • 1 ♀; same data as for preceding; 31 May 1962; G.D. Butler leg.; UAIC • 1 ♀; vicinity of Portal , Cochise County; 25 Aug. 1960; P.H. Arnaud, Jr. and D.C. Rentz leg.; CAS • 1 ♀; West Turkey Creek , Cochise County; 2 Sep. 2003; J.G. Rozen, J.S. Ascher, R.L. Staff, and R.E. Edwards leg.; AMNH • 1 ♂; Whiteriver , Navajo County; 9 Jul. 1977; R.C. Miller leg.; UCBME • 1 ♀; Willcox , Cochise County; 28 Aug. 1974; R.M. Bohart leg.; UCBME. – California • 1 ♀; 1 mi N of Hodge , San Bernardino County; 13 Jun. 1979; J.C. Hall leg.; UCR UCRC ENT 57190 • 1 ♀; 1.7 mi S of Hwy 223 along Hwy 99, Kern County; 8 Oct. 1999; G.R. Ballmer leg.; UCR UCRC ENT 00037952 • 2 ♀♀; 10 mi NE of Artois , Glenn County; 25 Jul. 1967; R.W. Thorp leg.; UCBME • 1 ♀; 6 mi N of Borrego Springs Rd and Di Giorgio Rd junction ( Coyote Creek ), Anza Borrego State Park , San Diego County; 5 Apr. 1968; S. and S. Frommer leg.; UCR UCRC ENT 57191 • 1 ♀; 7 mi N of Scissors Crossing , San Diego County; 15 May 1968; F.G. Andrews leg.; UCR UCRC ENT 57165 • 2 ♂♂; Alpine Ranch , San Diego County; 19 Aug. 1998; D.K. Faulkner leg.; SDNHM • 1 ♂; Anza Borrego State Park , San Diego County; 6 Jul. 1993; R.B. Parks leg.; SDNHM • 1 ♀; Anza Borrego State Park , San Diego County; 32.9695° N, 116.3088° W; 17 Aug. 2013; K.L.J. Hung leg.; SDNHM SDC.1995 • 3 ♀♀; Anza Borrego State Park , San Diego County; 33.1356° N, 116.3163° W; 30 Aug. 2014; K.L.J. Hung leg.; SDNHM SDC.11195 , SDC.11196 , SDC.11202 • 1 ♀; Davis ; 14 Sep. 1961; M.E. Irwin leg.; UCBME • 2 ♀♀, 1 ♂; E of Barrett Honor Camp , Cleveland National Forest , San Diego County; 25 Jun. 2003; D.K. Faulkner leg.; SDNHM • 1 ♂; Grapevine Canyon , Anza Borrego State Park , San Diego County; 12 Jul. 1985; R.B. Parks leg.; SDNHM • 1 ♀, 1 ♂; same data as for preceding; 20 Jul. 1985; R.B. Parks leg.; SDNHM • 1 ♀; same data as for preceding; 1 Sep. 1985; R.B. Parks leg.; SDNHM • 1 ♀; Imperial Valley ; 10 Jul. 1981; G.D. Butler leg.; UAIC • 1 ♀; Independence , Inyo County; 19 Jul. 1965; J. Birchim leg.; CAS • 2 ♂♂; Jamacha , San Diego County; 12 Aug. ??20; SDNHM • 1 ♀; Keeler , Inyo County; 12 Jul. 1982; R.M. Bohart leg.; UCBME • 1 ♀; Keosegan Ranch ( Indio ), Riverside County; 14–16 Jul. 1970; M.E. Irwin leg.; UCR UCRC ENT 57178 /Cotton Insect Survey M.E. Irwin Project Specimen Number 730 • 1 ♀; Lone Pine , Inyo County; 7 Jun. 1937; E.C. Van Dyke leg.; CAS • 1 ♀; Merced County; 27 Sep. 1959; W. Cothran leg.; UCBME P 0219947 • 1 ♀; Mercey Hot Springs , Fresno County; 21 Sep. 1963; R.L. Westcott leg.; UCBME • 2 ♂♂; Mission Gorge , San Diego County; 5 Sep. 1983; R.B. Parks leg.; SDNHM • 1 ♀; near La Mesa , San Diego County; 5 Sep. 1953; F.X. Williams leg.; CAS • 1 ♀; near Wendel , Lassen County; Jul. 1934; R.D. Murphy leg.; CAS • 1 ♀; Northridge , Los Angeles County; 2 Jul. 1940; SDNHM • 2 ♀♀; NW corner of UC Riverside Botanic Gardens ( Riverside ), Riverside County; 29 Apr. 1992; D.C. Hawks leg.; UCR UCRC ENT 57114 , UCRC ENT 57115 • 1 ♀; Pope Valley (⅓ km S of Aetna Springs ), Napa County; 3–14 Oct. 1993; L.S. Kimsey leg.; UCBME • 1 ♀; Potrero , San Diego County; 27 Jun. 1958; F.X. Williams leg.; SDNHM • 1 ♂; Ramona , San Diego County; 19 May 1992; R.B. Parks leg.; SDNHM • 1 ♀; Riverside , Riverside County; P.H. Timberlake leg.; USNM • 1 ♀; same data as for preceding; 18 Jul. 1928; P.H. Timberlake leg.; UCMC • 1 ♀; same data as for preceding; 21 Jul. 1933; P.H.Timberlake leg.; UCBME P 0219953 • 1 ♀; same data as for preceding; 10 Aug. 1957; E.I. Schlinger leg.; UCBME P 0218818 • 1 ♀; Saline Valley Salt Lake , Inyo County; 36.6967° N, 117.8328° W; 19 May 1995; T.L. Griswold leg.; BBSL BBSL Faunal Survey No. 000 102 989 • 2 ♀♀; San Diego , San Diego County; 20 Jul. 1920; W.S. Wright leg.; SDNHM • 1 ♀; same data as for preceding; 5 Jul. 1921; W.S. Wright leg.; SDNHM • 1 ♀; San Diego Country Estates , Ramona; 15 Jun. 1992; R.B. Parks leg.; SDNHM • 2 ♀♀; Scissors Crossing , Valle de San Felipe , San Diego County; 4 May 1969; E.I. Schlinger leg.; UCR UCRC ENT 57163 , UCRC ENT 57164 • 1 ♀; SWS3 Zircon Street , San Diego County; 32.7202° N, 117.0779° W; 28 Jun. 2022; J.L. Mullins leg.; UCR UCSD.JLM.4140 • 1 ♀; Tracy , San Joaquin County; 13 Jun. 1949; J.W. MacSwain leg.; USNM • 1 ♀; California, White Slough , San Joaquin County; 22 Aug. 1984; L.G. Bezark leg.; UCBME P 0219928 . – Colorado • 2 ♀♀; “ El Paso County Solar Trails County Park T14S R67W Sec 15 SW¼ ”; F.M. Brown leg.; UCMC • 1 ♀; 2 mi W of Walsenburg ; 1 Jul. 1975; B. Vogel leg.; UCMC • 1 ♀; 3 mi W of Arriola , Montezuma County; 4–24 Aug. 1975; T. Marquardt leg.; CSUC • 1 ♀; same data as for preceding; Sep. 1975; T. Marquardt leg.; CSUC • 2 ♀♀; 4 mi E of Steamboat Springs ; 28 Aug. 1979; U.N. Lanham leg.; UCMC • 1 ♀; Boulder , Boulder County; 13 Jul. 1925; C.H. Hicks leg.; UCMC • 2 ♀♀; same data as for preceding; 2 Aug. 1925; C.P. Custer leg.; UCMC • 1 ♀; Clear Creek ; 3 Aug. ??98; USNM • 2 ♀♀; County Road 42 (3 mi E of County Road 41.7 ), Las Animas County; 17 Jul. 2012; W. Cranshaw, B. Kondratieff, and D. Leatherman leg.; CSUC • 1 ♀; Evergreen ; 16 Jul. 1952; R.S. Beal leg.; ASU • 1 ♀; Glacier view Meadows , Larimer County; 27 Jul. 1996; H.E. Evans leg.; CSUC • 1 ♀; Jim Creek (near Boulder ); 24 Jul. 1922; AMNH • 1 ♀; Phantom Canyon , Larimer County; 30 Jul. 1992; S. Fitzgerald leg.; CSUC. – Nevada • 1 ♀; 7 mi N of Dyer , Esmeralda County; 2 Jul. 1958; R.C. Bechtel leg.; FSCA • 2 ♀♀; Alamo , Lincoln County; 13 Jul. 1958; F.D. Parker leg.; FSCA. – New Mexico • 1 ♀; 1 mi N of Rodeo , Hidalgo County; 22 Aug. 1996; J.G. and B.L. Rozen and A. Pence leg.; AMNH • 1 ♀; 3 km S of Seven Rivers , Eddy County; 3 Sep. 1990; T. Griswold leg.; BBSL BBSL522617 • 5 ♀♀; 5 mi NE of San Lorenzo , Grant County; 32.8197° N, 107.8967° W; 28 Aug. 2000; J.L. Neff leg.; CTMI 20245 to 20247 , 20259 , 20347 • 1 ♀; 8.3 km SW of Silver City , Grant County; 25 Aug. 1992; B.N. Danforth leg.; USNM • 1 ♂; 10 mi N of Mimbres , Grant County; 32.9778° N, 108.0528° W; 6 Aug. 2007; J.H. Kits leg.; DEBU debu00293461 • 1 ♀; 15 mi E of Santa Fe; 27 Jun. 1931; H.A. Scullen leg.; OSAC • 1 ♀; 20 mi W of Jal , Lea County; 3 May 1961; J.C. Bequaert and K. Roever leg.; UAIC • 1 ♀; 25–28 mi S of Animas , Hidalgo County; 16 Aug. 1994; J.G. Rozen and J.S. Ascher leg.; AMNH • 2 ♂♂; Animas , Hidalgo County; 31.9500° N, 108.8070° W; 26 Aug. 2015; L. Packer leg.; PCYU • 1 ♀; Iron Creek Campground , Gila National Forest , Grant County; 32.9089° N, 107.8058° W; 16 Aug. 2007; M. Buck leg.; DEBU debu00292521 • 1 ♀; Las Vegas; 20 Jul. ??02; Oslar leg.; USNM • 1 ♀; Loving ; 8 May 1945; J.W. MacSwain leg.; CAS • 1 ♀; same data as for preceding; 26 May 1945; J.W. MacSwain leg.; CAS • 2 ♀♀; same data as for preceding; 29 May 1945; J.W. MacSwain leg.; CAS • 1 ♀; Mesilla; 28 Jun.; A.P. Morse leg.; USNM • 1 ♀; New Canyon Campground (8 mi W of Manzano ), Manzano Mountains ; CNC PCYU: CNC-Bee: 449 • 1 ♀; Santa Fe; M. Boyle leg.; USNM • 1 ♀; Wrights Cabin Campground , Gila National Forest , Grant County; 32.9131° N, 107.7806° W; 16 Aug. 2007; J.H. Kits leg.; DEBU debu00293007 . – Oklahoma • 1 ♀; Sutton Wilderness Trail Park ( Norman ), Cleveland County; 35.2454° N, 97.4286° W; 21 Oct. 2023; K.L.J. Hung leg.; OKBS OKBS.POL.14137 . – Texas • 2 ♀♀; Belfrage leg.; USNM • 1 ♀; same data as for preceding; Morrison leg.; USNM • 1 ♀; 1 mi E of Eagle Pass , Maverick County; 9 Apr. 1999; J.L. Neff leg.; CTMI 09853 • 1 ♀; 1 mi E of Rio Grande City , Starr County; 15 Mar. 1999; J.L. Neff, A. Hook, and C.R. Riley leg.; CTMI 09421 • 1 ♀; 10 mi SE of Dryden , Terrell County; 22 Sep. 1972; W. Hanson and J. Poff leg.; BBSL • 1 ♀; 100 mi E of El Paso ; 13 Jul. 1942; E.C. Van Dyke leg.; CAS • 5 ♀♀; 13 mi E of Snyder , Scurry County; 4 Jun. 1965; R.M. Bohart leg.; UCBME • 1 ♀; 14 mi W of Grandfalls , Ward County; 9 Sep. 1989; J.L. Neff leg.; CTMI K07334 • 2 ♀♀; 2 mi S of El Sauz , Starr County; 29 May 1980; D.W. Webb leg.; INHS 7835 , 7836 • 3 ♀♀; 20 mi N of Van Horn ; 28 Sep. 1957; W. Nutting and F. Werner leg.; UAIC • 4 ♀♀; 8 mi NW of Girvin , Pecos County; 9 Sep. 1989; J.L. Neff leg.; CTMI K07226 , K07227 , K07231 , K07233 • 1 ♀; Alpine ; 10 Jun. 1930; G. Linsley leg.; CAS • 1 ♀; same data as for preceding; 1 Jul. 1942; E.C. Van Dyke leg.; CAS • 7 ♀♀; same data as for preceding; 1 Jul. 1942; H.A. Scullen leg.; OSAC • 1 ♀; same data as for preceding; 2 Jul. 1942; E.C. Van Dyke leg.; CAS • 2 ♀♀; same data as for preceding; 8 Jul. 1942; E.C. Van Dyke leg.; CAS • 3 ♀♀; same data as for preceding; 8 Jul. 1942; H.A. Scullen leg.; OSAC • 1 ♀; Big Bend National Park , Nine Point Draw ; 15 May 1959; W.R.M. Mason leg.; CNC 808404 • 2 ♀♀; Brackenridge Field Laboratory ( Austin ), Travis County; 22 Oct. 1985; J.L. Neff leg.; CTMI K02556 , K02557 • 1 ♀; same data as for preceding; 11.xi.1989; A. Hook leg.; CTMI • 1 ♀; Cotulla ; 18 Apr. 1906; F.C. Pratt leg.; USNM • 1 ♀; Cuevas Amarillas Springs (24 km ENE of Presidio ), Big Bend Ranch State Natural Area , Presidio County; 30–31 May 1992; C.R. Nelson, J. Gelhaus, and D. Koenig leg.; CTMI • 3 ♀♀; Davis Mountains ; 28 Jun. 1942; E.C. Van Dyke leg.; CAS • 1 ♀; Davis Mountains , Jeff Davis County; 26 Jun. 1942; H.A. Scullen leg.; OSAC • 2 ♀♀; same data as for preceding; 27 Jun. 1942; H.A. Scullen leg.; OSAC • 5 ♀♀; same data as for preceding; 28 Jun. 1942; H.A. Scullen leg.; OSAC • 1 ♀; Del Rio ; 8 May 1907; F.C. Bishopp leg.; USNM • 2 ♀♀; Devils River ; 3 May 1907; F.C. Bishopp leg.; USNM • 1 ♀; same data as for preceding; 6 May 1907; F.C. Bishopp leg.; USNM • 2 ♀♀; same data as for preceding; 7 May 1907; F.C. Bishopp leg.; USNM • 1 ♀; Dolan Creek above Devils River , Val Verde County; 18 Sep. 1993; C.R. Nelson, S.M. Stringer, and S. Thomas leg.; CTMI • 3 ♀♀; Dolan Falls ( Devils River ), Val Verde County; 26–29 May 1994; A. Hook and O. Hernandez leg.; CTMI • 1 ♀; same data as for preceding; 24 Jun. 1994; Kondratieff and Kippenhan leg.; CSUC • 1 ♀; same data as for preceding; 5–7 Aug. 1994; A. Hook and O. Hernandez leg.; CTMI • 1 ♀; same data as for preceding; 4–6 Nov. 1994; A. Hook and O. Hernandez leg.; CTMI • 1 ♀; Dripping Springs ; 10 Aug.; USNM • 1 ♀; Eagle Pass , Maverick County; 25 May 1952; M. Cazier, W. Gertsch, and R. Schrammel leg.; AMNH • 1 ♀; El Paso ; 23 Jun. 1942; E.C. Van Dyke leg.; CAS • 1 ♀; Fedor , Lee County; 23 Sep. 1897; Birkmann leg.; ZMB • 1 ♀; same data as for preceding; 15 Jun. 1898; Birkmann leg.; ZMB • 1 ♀; same data as for preceding; 3 May 1909; AMNH • 4 ♀♀; Texas, Fort Davis ; 27 Jun. 1942; E.C. Van Dyke leg.; CAS • 1 ♀; Gail , Borden County; 4 Jun. 1965; R.M. Bohart leg.; UCBME • 1 ♀; Green Gulch, Big Bend National Park ; 2 May 1959; W.R.M. Mason leg.; CNC 753901 • 1 ♀; Horsehead Crossing , Pecos County; 31.2333° N, 102.5167° W; 10 Sep. 1996; J.L. Neff leg.; CTMI 08075 • 1 ♀; same data as for preceding; 10 Sep. 1996; J.L. Neff leg.; CTMI 08076 /M.G.R. Database No. 1213 • 1 ♀; Hwy 163 (near Sterling City ), Sterling County; 23 Apr. 1998; S. Fitzgerald, B. Kondratieff, and D. Leatherman leg.; CSUC • 1 ♀; Hwy 84, Double Mountain Fork Brazos River , Justiceburg , Garza County; 1 Jul. 1993; A.W. Hook leg.; CTMI • 1 ♀; James River (20 mi NNW of Harper ), Kimble County; 2 Jun. 1996; J.L. Neff leg.; CTMI 07972 • 1 ♀; Kermit Sand Hills , County Road 302 , Winkler County; 31.9022° N, 103.0467° W; 12 May 2019; M. Buck leg.; RAM pmae00151961 • 4 ♀♀; Kerrville; Apr. ??07; P. Durham leg.; USNM • 2 ♀♀; same data as for preceding; 11 Apr. ??07; F.C. Pratt leg.; USNM • 2 ♀♀; same data as for preceding; 12 Apr. ??07; F.C. Pratt leg.; USNM • 1 ♀; same data as for preceding; 14 Apr. ??07; P. Durham leg.; USNM • 1 ♀; same data as for preceding; 22 Jun. 1942; E.S. Ross leg.; CAS • 1 ♀; Lake Childress , Childress County; 24 Aug. 1948; P.-C. Vaurie leg.; AMNH • 1 ♀; McAllen , Hidalgo County; 13 May 1952; M. Cazier, W. Gertsch, and R. Schrammel leg.; AMNH • 1 ♀; Palo Duro Canyon (22 mi S of Claude ), Armstrong County; 4 Apr. 1979; R.J. McGinley leg.; USNM M.G.R. Database No. 529 • 2 ♀♀; Pecos , Reeves County; 15 May 1927; J.O. Martin leg.; CAS • 1 ♀; same data as for preceding; 11 Jul. 1942; H.A. Scullen leg.; OSAC • 1 ♀; Pine Canyon , Big Bend National Park ; 4 May 1959; J.F. McAlpine leg.; CNC 753874 • 2 ♀♀; Quemado , Maverick County; 25 May 1952; M. Cazier, W. Gertsch, and R. Schrammel leg.; AMNH • 1 ♀; same data as for preceding; 3 Sep. 1960; P.M. Marsh leg.; UCBME • 1 ♀; Reagan Wells ; 25 May 1916; F.C. Bishopp leg.; USNM • 1 ♀; Rio Grande Village , Big Bend National Park , Brewster County; 23 Jun. 1970; M. Cazier, O. Francke, and L. Welch leg.; ASU • 1 ♀; Sanderson , Terrell County; 12 Jun. 1948; M. Cazier leg.; AMNH • 1 ♀; Sandy , Blanco County; 12 Jun. 1994; J.L. Neff and A. Hook leg.; CTMI • 1 ♀; Shumla , Val Verde County; 26 May 1952; M. Cazier, W. Gertsch, and R. Schrammel leg.; AMNH • 1 ♀; Texas Tech University , Junction , Kimble County; 22 May 1983; J.L. Neff leg.; CTMI K13693 • 1 ♀; The Basin (near base of Lost Mine Trail ) ( Chisos Mountains ), Big Bend National Park ; 14 Aug. 1962; H.V. Weems, Jr. leg.; FSCA • 1 ♀; The Rockpile (30 mi NW of Fort Davis ); 28 May 1959; W.R.M. Mason leg.; CNC 753875 • 1 ♀; Uvalde ; 3 Jun. 1916; F.C. Bishopp leg.; USNM • 1 ♀; Texas, Valentine ; 27 Sep. 1957; W. Nutting and F. Werner leg.; UAIC • 1 ♀; Van Horn ; 24 Jun. 1942; E.C. Van Dyke leg.; CAS • 1 ♀; same data as for preceding; 24 Jun. 1942; E.C. Van Dyke leg.; CAS M.G.R. Database No. 1212 • 1 ♀; Zapata, San Ygnacio 15 S; 13 May 1974; J.L. Neff leg.; CTMI K11139 . – Utah • 1 ♀; Beaver Canyon; Jun.; USNM • 1 ♀; St. George , Washington County; 8 Jun. 1964; W.J. Hanson leg.; BBSL BBSL522488 • 1 ♀; Wildcat Valley , Beaver County; Aug.; USNM .

Non-preserved material

USA – California • 1 ♀; Charles St , San Diego, San Diego County; 32.7180° N, 117.2443° W; 13 Jul. 2020; B. Nickel obs.; iNaturalist record #52965246 • 1 ♀; Mission Trails Regional Park , San Diego; 32.8347° N, 117.0758° W; 22 Aug. 2023; M.K. James obs.; iNaturalist record #179755886 • 1 ♀; Santee , San Diego County; 14 Aug. 2020; B.J. Stacey obs.; iNaturalist record #69844452 . – Texas • 1 ♀; Ector County; May 2021; S.M. McCoshum obs.; iNaturalist record #147628875 • 1 ♀; Mills County; 9 Apr. 2022; R. Pfau obs.; iNaturalist record #111471058 • 1 ♀; Road to Dry Creek Lake , Briscoe County; 3 Sep. 2020; C.C. Galley obs.; iNaturalist record #59724169 .

Redescription

MEASUREMENTS OF HOLOTYPE. Body length 8.6 mm; ITW 2.1 mm; head length 2.6 mm; head width 3.5 mm; fore wing length 8.2 mm.

Both sexes

INTEGUMENT COLORATION. Dark brown to black except as follows. Mandible with apical third golden yellow. Mandible with basal two-thirds, labrum along apical and lateral margins, scape and pedicel to some extent, F1 extensively, pronotal lobe in some non-type specimens, tegula, coxae to some extent, trochanters and femora predominantly to entirely, and tibiae and tarsi entirely orange. F2 with orange spot basally. Fore wing membrane subhyaline, apically dusky. Hind wing membrane dusky subhyaline to hyaline. Metasomal terga laterally, pygidial plate to some extent, and metasomal sterna to some extent reddish brown.

PUBESCENCE. Face with tomentum densest around antennal socket. Tomentum slightly sparser on clypeus; upper paraocular and frontal areas and vertexal area mostly exposed. Pronotal collar with tomentum uniformly pale yellow (off-white in some non-type specimens). Mesoscutum with well-defined paramedian band of pale-yellow tomentum (off-white in some non-type specimens), tapering slightly toward but not attaining anterior margin; pale tomentum otherwise mostly restricted to lateral and posterior margins. Mesopleuron with off-white to pale-yellow, appressed, branched setae; densely setose except for two sparsely setose circular patches (one beneath base of fore wing (hypoepimeral area) and slightly larger one in ventrolateral half of mesopleuron). Metanotum with tomentum uninterrupted, uniformly pale yellow (off-white in some non-type specimens). Propodeal triangle mostly glabrous, with (pale) setae restricted to small lateral patches. T1 with basal and apical transverse bands of off-white to pale-yellow tomentum interrupted medially (former more widely than latter) and subparallel, discal patch trapezoidal. T2–T4 with complete apical transverse bands of off-white to pale-yellow tomentum, those of T2–T3 medially somewhat removed from apical margins of terga, that of T2 with pair of basomedially convergent anterolateral extensions. S2–S3 with apical transverse bands of white tomentum.

SURFACE SCULPTURE. Labrum coarsely and densely (most i <1d) rugose-punctate. Clypeus densely punctate (most i <1d) but interspaces well defined, shining. Vertexal area somewhat sparsely punctate (some i>2d), especially around ocelli. Mesoscutum, mesoscutellum, and axilla with punctures more or less equally dense and nearly contiguous (most i <1d). Mesopleuron with punctures in upper half nearly contiguous and denser (i <1d) than in ventrolateral half (i ≤ 2d); interspaces shining where punctures not contiguous; punctures similar in size throughout. Discs of metasomal terga with punctures very fine, dense (i ≈1d), and evenly distributed; interspaces shining somewhat.

STRUCTURE. Labrum with pair of small subapical denticles, each preceded by discrete longitudinal ridge. Pronotal collar rather short (medial length ~ 3/5 MOD). Mesoscutellum moderately bigibbous. Axilla extending little if at all beyond midlength of mesoscutellum; tip distinctly pointed, but mesally unattached to mesoscutellum for less than 2/5 medial length of axilla; lateral margin relatively straight. Fore wing with three submarginal cells.

Female

T5 with concave apical margin and large patch of off-white to pale-yellow tomentum on each side lateral to pseudopygidial area. Pseudopygidial area with underlying integument to some extent reddish orange in some non-type specimens, with triangular region of posteriorly directed setae with three subregions (basal patch of dense, golden setae; darker subapical band of sparser, coppery brown setae; and apical row of dense, suberect, silvery setae; poorly differentiated in some non-type specimens) within larger trapezoidal space of posteromedially directed coppery brown setae. Pygidial plate apically truncate. S4 with apical transverse band of white tomentum. S5 straight in lateral view, with apical fimbria of coppery bristle-like setae and brown tomentum laterally if not throughout; S5 otherwise covered in off-white tomentum.

Male

T5 with complete apical transverse band of pale-yellow tomentum. T6 with small posteromedial patch of off-white tomentum. Pygidial plate relatively flat and apically rounded. S4–S5 each with apical/ subapical fringe of dense, long (> 1 MOD), curved setae, those of S4 coppery to silvery, those of S5 coppery and contrasting with apical transverse bands of white tomentum of preceding sterna.

Distribution

Southern Great Plains to California and western Mexico ( Fig. 12N ).

Ecology

Host records

Unknown.

Floral records

Cockerell (1900, 1929), images on iNaturalist, and labels of examined voucher specimens indicate that this species has been collected from the following flowering plant species: in Apocynaceae , Asclepias sp. ; in Asteraceae , Aster sp. , Baccharis sp. , B. salicina , Bidens sp. , Carthamus tinctorius , Encelia palmeri Vasey & Rose , Gaillardia pulchella , Grindelia pulchella Dunal , Gutierrezia californica (DC.) Torr.&A.Gray , G. sarothrae , Haplopappus sp. or spp., Helenium amarum (Raf.) H.Rock , Heterotheca sp. , H. subaxillaris (Lam.) Britton & Rusby , Isocoma menziesii (Hook. & Arn.) G.L.Nesom , I. pluriflora , Pectis papposa Harv. & Gray , Pluchea sericea (Nutt.) Coville , Ratibida columnifera (Nutt.) Wooton & Standl. , Rudbeckia sp. , Solidago sp. , Symphyotrichum sp. , Tetraneuris linearifolia Greene , Verbesina sp. or spp., Xanthisma gracile (Nutt.) D.R.Morgan & R.L.Hartm. , and X. spinulosum (Pursh) D.R.Morgan & R.L.Hartm. ; in Boraginaceae , Heliotropium sp. or spp.; in Brassicaceae , Lepidium sp. or spp.; in Cleomaceae Bercht. & J.Presl , Wislizenia sp. ; in Fabaceae , Acacia sp. , Dalea candida Willd. , Medicago sativa L., Melilotus albus , Phaseolus sp. , Pithecellobium sp. , and Trifolium repens ; in Lamiaceae , Marrubium vulgare L., Monarda sp. , and M. citriodora Cerv. ex Lag. ; in Malvaceae , Sphaeralcea sp. ; in Polygonaceae , Eriogonum fasciculatum Benth. ; in Tamaricaceae , Tamarix sp. ; in Verbenaceae , Lippia sp. ; and in Zygophyllaceae , Kallstroemia grandiflora .

Remarks

In the original description of E. permixtus, Cockerell (1923) did not mention T. segregatus , having misidentified the two female and two male representatives as an Epeolus based on the presence of two (as opposed to three) maxillary palpomeres, which is not diagnostic for either genus ( Rightmyer 2004), and instead compared them to Triepeolus dacotensis (as E. dacotensis ), which is a visibly different bee, and T. concinnus Cockerell, 1917 , a name that has since been synonymized under T. townsendi Cockerell, 1907 ( Rightmyer 2008), which he described as superficially similar but with a much larger pseudopygidial area.

In the original description of T. brunnescens, Cockerell & Sandhouse (1924) did not mention T. segregatus and instead compared the (male) holotype to T. brunneus Cockerell, 1921 , a name that has since been synonymized under T. balteatus Cockerell, 1921 ( Rightmyer 2008), which the authors described as easily distinguishable “by the pleura and pattern of abdomen”, Triepeolus balteatus is more easily separated from the holotype of T. brunnescens (and all other Triepeolus ) by the presence of a pair of curved, clasper-like structures near the bases of the metacoxae.

In the original description of T. pacis, Cockerell (1925b) did not mention T. segregatus or T. brunnescens and instead compared the (male) holotype to T. blaisdelli Cockerell & Sandhouse, 1924 , to which T. pacis was said to be related, T. mensae Cockerell, 1924 , and T. norae Cockerell, 1907 . The (male) holotypes of T. blaisdelli (CAS 1603) and T. mensae (CAS 1607) were examined and recognized as members of the verbesinae species group; as in most Nearctic species in the T. verbesinae group, in both specimens the propodeal triangle is mostly covered in dense, appressed pale setae (as opposed to mostly glabrous, with (pale) setae restricted to small lateral patches). The pseudopygidial area of the (female) holotype of T. norae is subtriangular (with the apical margin of T5 slightly convex), confirming the placement of this species outside the T. simplex group.

In the original description of E. sarothrinus, Cockerell (1929) compared the (female) holotype to E. lectiformis , which is herein established as a junior synonym of T. rhododontus (vide supra), and E. permixtus , with which E. sarothrinus was said to have evident affinity but differing “by [ E. permixtus ] being distinctly larger and more robust, [with] the eyes much larger and very much darker, the antennae stouter and the apical plate broader”. However, Cockerell (1929) added that “it may later seem expedient to treat [ E. sarothrinus ] as a northern race of E. permixtus ”. Cockerell (1929) described a variety of E. sarothrinus under the name confluens based on a male (the holotype) in which the paramedian bands are anteriorly joined to lateral transverse bands of pale tomentum along the anterior margin of the mesoscutum, the pro- and metafemora are partially brown, the T1–T2 apical transverse bands are complete, and the pygidial plate is less obtuse.

Barcoded specimens resembling the types of E. occidentalis var. segregatus , E. permixtus , T.brunnescens , T.pacis , E.sarothrinus , and E. sarothrinus var. confluens were assigned to three BINs—BOLD:AAG7909, BOLD:AEI6013, and BOLD:AFP1028—which appear in two widely separated clusters (with a minimum distance of 3.9%) in a neighbor-joining tree that includes all available sequences of species in the T. simplex group (Supp. file 4). The BIN BOLD:AFP1028 includes specimens that were previously assigned to the BIN BOLD:AAG7909 (its nearest neighbor), from which it was split in a subsequent iteration of the RESL clustering algorithm despite exhibiting a barcode sequence divergence of only 1.0%. The sequences in these two BINs cluster together closely (Supp. file 4). The BIN BOLD:AEI6013 exhibits a larger barcode sequence divergence (of 5.3% (p-dist)) from its nearest neighbor BIN (BOLD:AFP1028) in the aforementioned cluster. The sequences in both clusters lack in-frame stop codons, have a similar mean GC content (25.0%, SE ± 0.2 for BINs BOLD:AAG7909+BOLD:AFP1028; 23.3%, SE ± 0.2 for BIN BOLD:AEI6013), and strong AT-bias in the third codon position (95.3%, SE ± 0.2 for BINs BOLD:AAG7909+BOLD:AFP1028; 94.7%, SE ± 0.2 for BIN BOLD:AEI6013). However, there are four specimens for which barcoding attempts at the CBG failed (BOLD sample IDs: CCDB-24584 D10, CCDB-24584 D12, CCDB-30384 E02, and CCDB-30384 E10; see Supp. file 1 for specimen and vouchering information). Examination of the associated trace files and their reassembly into consensus sequences, which were aligned, revealed stop codons in every reading frame in all four sequences and indels in one. Although these are clearly non-target sequences, they were matched with 97.7–98.9% similarity to the same sequence (BOLD sample ID: CMNTO_025) in BIN BOLD:AFP1028 and cluster more closely to the sequences in this cluster than any other. The sequences of another four (male) specimens that closely resemble the allotype of T. apache (BOLD sample IDs: CMNTO_485, CMNTO_487, CMNTO_488, and CMNTO_489) unexpectedly appeared in this cluster and are thus listed here as T. cf. segregatus . If these specimens are heterospecific as suggested by their morphology, the observed low sequence divergence from T. segregatus in this cluster could be explained by the possible sequencing of conserved nuclear mitochondrial pseudogenes (numts) mistaken for COI sequences. Françoso et al. (2019) found very similar numts (1.2% divergence) that coincided with highly divergent COI sequences (16.5% divergence) in two species of Tetragonula Moure, 1961 ( Hymenoptera : Apidae : Apinae ) stingless bees. Thus, of the three BINs available for T. segregatus , it is possible if not probable that BINs BOLD:AAG7909 and BOLD:AFP1028 are comprised of COI paralogs (numts) despite those sequences having been designated as barcode compliant. In any event, the placement of any sequences in this cluster should be treated cautiously for identification purposes.

Further complicating the treatment of these MOTUs as separate species is that barcoded specimens from the two groups cannot reliably be told apart morphologically. For example, although there is pronounced variation among specimens in mesopleural puncture density, with punctures ranging from nearly contiguous throughout to separated by>2d ventrolaterally, specimens whose sequences comprise separate MOTUs exhibit considerable overlap with respect to this feature. Specifically, those with sequences in the cluster to which the BINs BOLD:AAG7909 and BOLD:AFP1028 were assigned exhibit continuous variation in mesopleural puncture density, with punctures ranging from nearly contiguous throughout to separated by>2d ventrolaterally, whereas in those with sequences corresponding to BIN BOLD:AEI6013 the mesopleura are sparsely punctate ventrolaterally (many i>2d). In the holotypes of E. permixtus , T. brunnescens , T. pacis , E. sarothrinus , and E. sarothrinus var. confluens , the mesopleura are finely rugose-punctate, each with the punctures more or less equally dense throughout (most i <1d), which is typical of (but not exclusive to) specimens from the Californias. Specimens from Arizona to Texas exhibit greater variation in mesopleural puncture density, even locally. For example, in the AMNH collection there are non-barcoded specimens taken from the same locality (the Southwestern Research Station near Portal, Arizona) in which the mesopleura range from finely and densely to coarsely and sparsely punctate. In the holotype of E. occidentalis var. segregatus , the mesopleura are somewhat more coarsely and sparsely punctate (most i ≤ 1d) than in the aforementioned types, but the specimen does not more obviously resemble barcoded specimens of one haplotype than the other in this or any other regard.

In specimens from Texas and adjacent Mexico, for which DNA barcode sequences are presently unavailable, the pronotal lobes are more commonly orange (as opposed to entirely black) and the metasomal tergal bands are typically (but not always) off-white (as opposed to yellow). Although these and the aforementioned examples of pronounced morphological variation as well as available DNA barcode data suggest the possibility of a cryptic species complex, it is not clear if or where species limits should be drawn. Thus, given the apparent lack of morphological or geographic support for each of the MOTUs, the unusual clustering of available barcode sequences (see Supp. file 4), and that it is possible for real species to have split (i.e., multiple) BINs ( Ratnasingham & Hebert 2013), they are herein regarded as representing a single, highly variable species in which intraspecific barcode sequence divergence is greater than might normally be expected among conspecifics, assuming that none of the sequences are of nuclear paralogous copies of the COI gene as discussed above.Additionally, as there are no discernable morphological differences among the holotypes of E. occidentalis var. segregatus , E. permixtus , T. brunnescens , T. pacis , E. sarothrinus , and E. sarothrinus var. confluens that fall outside the range of variation observed among both barcoded and non-barcoded specimens, E. permixtus , T. brunnescens , T. pacis , E. sarothrinus , and E. sarothrinus var. confluens are herein synonymized under T. segregatus .

Figures

Fig. 12A–D (continued on next page). Distribution maps. A. All species in the Triepeolus simplex group. B. T. apache sp. nov. C. T. eumeniformis sp. nov. D. T. hirsutus sp. nov. Approximate geographic ranges are indicated in orange and were estimated from occurrence records known to the authors (yellow and blue circles, with the latter indicating localities of DNA barcoded specimens).

Fig. 20. Axillae and mesoscutellum of female, dorsal view. A. Triepeolus kathrynae Rozen, 1989 (AMNH M.G.R. Database No. 3711). B. T. segregatus (Cockerell, 1900) (SDNHM SDC.11202). Blue lines indicate the posterior extent of the axilla relative to the length of the mesoscutellum; red lines indicate the extent of the free portion of the axilla relative to its entire medial length.

Fig. 29. Triepeolus segregatus (Cockerell, 1900). A. Female (CNC PCYU:CNC-Bee: 509), habitus, lateral view. B. Female (CNC PCYU:CNC-Bee: 449), habitus, dorsal view. C. Male (PCYU 0005890), habitus, lateral view. D. Female (BOLD sample ID: CMNTO_017; PCYU) pseudopygidial area, dorsal view.

Fig. 15. T1 of female, dorsal view. A. Triepeolus apache sp. nov., paratype (AMNH). B. T. hirsutus sp. nov., paratype (AMNH). C. T. oblongimacula sp. nov., paratype (AMNH). D. T. rhododontus Cockerell, 1921 (AMNH). E. T. segregatus (Cockerell, 1900) (PCYU). F. T. shoshone sp. nov., paratype (PCYU PCYU-GS-07:2171).