Epeolus obscuripes Cockerell, 1917, 2019

Onuferko, Thomas M., 2019, A review of the cleptoparasitic bee genus Epeolus Latreille, 1802 (Hymenoptera: Apidae) in the Caribbean, Central America and Mexico, European Journal of Taxonomy 563, pp. 1-69 : 41-45

Epeolus obscuripes Cockerell, 1917 stat. nov.

Figs 1H , 4B , 7I , 12 C, 17

Epeolus bifasciatus obscuripes Cockerell, 1917: 298 (♂).

Epeolus schmidti Friese, 1925: 32 (♀, ♂), new lectotype designation, syn. nov.

Proposed common name

Dark-legged epeolus.

Diagnosis

Together with the morphological features that are diagnostic for the ‘ Trophocleptria group’, the following in combination can be used to tell E. obscuripes apart from all other Epeolus except E. bifasciatus : the mesoscutum lacks paramedian bands; the axillae are smooth (i.e., not crenulate) along the lateral margin ( Fig. 17D ); the mesoscutellum does not have a pair of posteriorly directed teeth ( Fig. 17D ); the mesopleura are coarsely punctate, each with smaller and sparser punctures ventrolaterally (many i≥1d) than in the upper half, with interspaces smooth and shining; T1 has only a broad, medially narrowed or interrupted, bright to pale yellow basal fascia ( Fig. 17 A–C); and T2 has a complete, bright to pale yellow apical fascia ( Fig. 17 A–C). Whereas in E. bifasciatus the frontal area has a pair of pronounced granulose protrusions, the pseudopygidial area of the female is very wide (the apex>2 × the medial length), only T1 and T2 of the male are distinctly fasciate, and the pronotal collar, pronotal lobes, axillae and mesoscutellum are entirely ferruginous, in E. obscuripes the frontal area typically has only a pair of weak protrusions, which are virtually lacking in some specimens, the pseudopygidial area of the female is more elongate medially (the apex ≤2 × the medial length), T3–T6 of the male typically have well-developed bright to pale yellow fasciae, and the pronotal collar, pronotal lobes, axillae and mesoscutellum range from entirely black to entirely ferruginous. Epeolus obscuripes is also similar to E. fumipennis in that in males of both species T1–T6 are typically fasciate. However, in E. fumipennis the mesoscutum has a pair of well-defined paramedian bands, the axillae are longer, extending as far back as or beyond the ridge overhanging the depressed posterior margin of the mesoscutellum, and T1 has a broad, medially narrowed bright to pale yellow submedial fascia. Despite the specific epithet ‘ obscuripes ’, meaning dark foot in Latin, the color of the legs does not reliably distinguish this species from E. bifasciatus , which may also have dark brown to black legs.

Material examined

Primary type material

COSTA RICA • ♂, E. schmidti lectotype; San José, San José; May 1922; Schmidt leg.; ZMB .

MEXICO • ♂, E. bifasciatus obscuripes holotype; Veracruz, Medellín; “ H. H. Hyde; Baker coll. 1785”; USNM 534041 .

Secondary type material

COSTA RICA • 1 ♀, E. schmidti lectoallotype; Alajuela, San Mateo ; May 1922; Schmidt leg.; ZMB .

DNA barcoded material with BIN-compliant sequences

Available. BOLD:ACW1534. Specimens examined and sequenced:

MEXICO • 1 ♂; Jalisco, San José del Carmen ; 10 Oct. 2008; L. Packer leg.; BOLD sample ID: CCDB- 28238 A08; PCYU • 1 ♀; Oaxaca, S of San Sebastián Frontera ; 18.2163° N, 97.6466° W; 25 Sep. 2008; L. Packer leg; BOLD sample ID: CCDB-22014 C06; PCYU • 1 ♀; Sonora, Rancho Fundición (30 km E of Álamos); 27.0183° N, 108.7483° W; 3 Oct. 2006; M.E. Irwin leg.; BOLD sample ID: CCDB- 28239 G08; BBSL FDP124693 .

Non-barcoded material

BELIZE • 1 ♂; Cayo, Central Farm ; 325 ft a.s.l.; 17.1773° N, 89.0053° W; 10–20 Feb. 2004; G. Steck and B. Sutton leg.; FSCA • 1 ♀; Cayo, Las Cuevas Access Rd and Caracol Rd ; 29 Apr. 2009; J.S. Ascher leg.; AMNH AMNH _ IZC 00290833 .

COSTA RICA • 1 ♀; Puntarenas, Las Cruces ; 25 Aug. 1977; T.P. Cogley leg.; FSCA .

EL SALVADOR • 1 ♂; La Libertad, Mount San Salvador; 8 Jul. 1963; M.E. Irwin and D.Q. Cavagnaro leg.; EMEC 1135879 .

GUATEMALA • 1 ♀; Alta Verapaz, 5 km W of Purulhá ; 11 Oct. 2005; J.B. Heppner leg.; FSCA .

HONDURAS • 1 ♀; Cortés, Estación Experimental Café (ca. Peña Blanca rainforest); 15 Aug. 1992; C. Porter and L. Stange leg.; FSCA • 1 ♂; Santa Bárbara, La Fé, Finca La Roca (5.3 km S of Peña Blanca); 14.9500° N, 88.0333° W; 21 Jun. 1994; Brooks and Ashe leg.; KUNHM SEMC1248337 .

MEXICO • 1 ♀; ANSP • 1 ♂; Chiapas, 3 km S of Palenque ( Nututun ); 25 Apr. 1993; W. LaBerge leg.; KUNHM SEMC1248323 • 1 ♂; Chiapas, 32 mi W of San Cristóbal ( Jct 190-195); 20 May 1969; H.J. Teskey leg.; CNC 754056 • 1 ♂; Chiapas, Palenque ruins; 22 Jun. 1969; B.V. Peterson leg.; CNC (754061) • 1 ♀; Jalisco, Chamela ; 7 Nov. 1986; J.G. and B.L. Rozen leg.; AMNH • 1 ♀; Jalisco, El Tuito; 6 Nov. 1987; L. Godinez leg.; KUNHM SEMC1248317 • 2 ♀♀; Jalisco, Estación de Biología - Chamela ; 30 Sep. 1985; J.G. Rozen leg.; AMNH • 1 ♂; same collection data as for preceding; 6 Oct. 1985; J.G. Rozen leg.; AMNH • 1 ♀; same collection data as for preceding; 4 Nov. 1987; L. Godinez leg.; KUNHM SEMC1248316 • 1 ♂; Jalisco, Guadalajara ; “ 8.1.03 ”; McClendon leg.; ANSP • 1 ♀; Jalisco, Tuxpan ; “ ix.4 ”; McClendon leg.; ANSP • 1 ♂; Morelos, 12 mi E of Cuernavaca ; 14 Aug. 1954; J.G. Chillcott leg.; CNC 754076 • 2 ♀♀; Nuevo León, Cola de Caballo ; 20 Jun. 1976; D. Weems leg.; FSCA • 1 ♀; Oaxaca, 3 mi S of El Camarón ; 2 Oct. 1986; R. Miller and L. Stange leg.; FSCA • 1 ♂; Puebla, Tepexco-Izúcar de Matamoros ( Carretera Federal 160); 18.6540° N, 98.6595° W; 4 Sep. 1998; T.L. Griswold leg.; BBSL BBSL 334999 • 1 ♀; Querétaro, Jalpan de Serra (along Río Jalpan ); 3 Sep. 1991; D. Yanega leg.; KUNHM SEMC1248286 • 1 ♀; San Luis Potosí, 14 mi W of Xilitla ; 22 Jul. 1954; J.G. Chillcott leg.; CNC 754075 • 1 ♀; San Luis Potosí, 9 km N of Tamazunchale ( Hwy 85); 9 Jul. 1990; W. Bell leg.; KUNHM SEMC1248283 • 1 ♀; San Luis Potosí, Cascada el Salto ( 12 km NW of El Naranjo ); KUNHM SEMC1248330 • 1 ♀; same collection data as for preceding; 20 May 1989; D. Yanega leg.; KUNHM SEMC1248284 • 1 ♀; same collection data as for preceding; 4 Jul. 1990; I. Yarom leg.; KUNHM SEMC1248288 • 1 ♂; San Luis Potosí, Col Salto del Agua ; 20 May 1989; D. Yanega leg.; KUNHM SEMC1248285 • 1 ♂; San Luis Potosí, El Salto; 19 Jun. 1973; H.V. Weems Jr. leg.; FSCA • 1 ♂; Tabasco, Teapa ; Mar. 1911 –1924; “Godman-Salvin Collection”; KUNHM SEMC0938794 • 1 ♂; Veracruz, 8 km S of Carrizal ; 5 Nov. 1991; T. Griswold leg.; KUNHM SEMC1248324 • 1 ♀; Veracruz, Catemaco ; 16–18 Jun. 1969; W.R.M. Mason leg.; CNC 754059 • 2 ♀♀; Yucatán, Chichén Itzá ; 24 Nov. 1981; L.A. Stange leg.; FSCA • 1 ♂; same collection data as for preceding; 5 Jan. 1992; J.R. Vockeroth leg.; CNC 754062 • 1 ♂; same collection data as for preceding; 17 Dec. 1993; L. Masner leg.; CNC 754060 .

Redescription

Male

MEASUREMENTS. Length 6.1 mm; head length 1.9 mm; head width 2.3 mm; fore wing length 5.6 mm.

INTEGUMENT COLORATION. Mostly black; notable exceptions as follows: at least partially ferruginous on mandible, labrum, antenna, pronotal collar, pronotal lobe, tegula, axilla, mesoscutellum, metanotum and legs. Mandible with apex darker than all but basal quarter; preapical tooth lighter than mandibular apex (difficult to see in E. bifasciatus obscuripes holotype because mandibles closed; described from nontype specimens). Antenna brown except scape, pedicel and F1 extensively orange. F2 with orange spot basally. Pronotal lobe and tegula pale ferruginous to amber. Wing membrane dusky subhyaline, slightly darker at apex. Legs with brown or black more extensive than reddish orange.

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 and dorsum of metasoma with bright yellow setae. Mesoscutum without pale tomentum, except for small patch between tegula and axilla. Mesopleuron nearly bare, except along margins. Metanotum with tomentum uninterrupted, uniformly pale yellow. T1 with broad, medially interrupted bright yellow basal fascia. T2 with narrower, complete, bright yellow apical fascia without anterolateral extensions. T3–T6 with tomentum sparse and partly rubbed off in E. bifasciatus obscuripes holotype, but with complete, bright yellow apical fasciae of moderately dense tomentum in E. schmidti lectotype and most non-type specimens. S3–S5 with long (>1 MOD), curved, coppery to silvery subapical hairs.

SURFACE SCULPTURE. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deeper and more distinct. Labrum with larger punctures than clypeus, but punctures of both equally dense (most i<1d). Small, impunctate, shiny spot lateral to lateral ocellus. Mesoscutum, mesoscutellum and axilla very coarsely and densely rugose-punctate. Tegula densely punctate anteriorly and mesally (i≤2d), sparsely punctate (i>2d) to impunctate posteriorly and along margins. Mesopleuron with larger and denser (i<1d) punctures in upper half than ventrolateral half (many i≥1d), interspaces shining. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc. Pygidial plate with large deep punctures closely clustered basomedially and sparser apically and laterally, with interspaces shining.

STRUCTURE. Preapical tooth acute. Labral apex with three small denticles, each preceded by small discrete longitudinal ridge. Frontal keel strongly raised. Frontal area with pair of sparsely punctate granulose protrusions, each located near upper mesal margin of compound eye. In E. schmidti lectotype and lectoallotype and multiple non-type specimens, protrusions discrete and more densely punctate, interspaces shining. Head dorsally with pair of protrusions, each located where upper genal area meets vertexal area. Vertexal area strongly convex in frontal view. Scape with greatest length 1.7 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.3). Preoccipital ridge separated from hypostomal carina by ≥2 MOD. Pronotal collar elongate (medial length ~1 MOD), expanded laterally to about 2 × medial length in dorsal view and relatively straight along anterior margin. Mesoscutellum moderately bigibbous, depressed along posterior margin beneath overhanging ridge. Axilla intermediate in size, its lateral margin nearly half as long as mesoscutellar width (AL/MSCW ratio = 0.4–0.5) and tip extending well beyond midlength of mesoscutellum but not as far back as its posterior margin; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion between ⅓ and 2 ∕ 5 its medial length; axilla with lateral margin relatively straight and carinate. Metanotum with blunt median process obscured by tomentum (process more pronounced in E. schmidti lectotype and multiple non-type specimens than in E. bifasciatus obscuripes holotype). Mesopleuron with carina delineating its anterior and lateral surfaces. Fore wing with three submarginal cells. Pygidial plate apically truncate.

Female

Description as for male except for usual secondary sexual characters and as follows: F2 even longer than wide (L/W ratio = 1.5); T3–T5 usually without fasciae; T5 laterally with long, erect simple setae; T5 with pseudopygidial area lunate, its apex twice as wide as medial length, indicated by silvery setae on impressed disc of apicomedial region elevated from rest of tergum; pygidial plate with smaller, denser punctures; S4 and S5 with straight and much shorter hairs (S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~⅓ MOD).

Distribution

Northwestern Mexico to Costa Rica ( Fig. 7I ).

Ecology

Host records

Unknown.

Floral records

The label of one examined voucher specimen indicates a floral association with Bidens pilosa .

Remarks

In Moure et al. (2007), E. bifasciatus obscuripes appears as a junior synonym of E. fulvopilosus , although it is not clear when and where this change in taxonomic status was first proposed, as no such change appears in any of the accompanying references. The holotype of E. bifasciatus obscuripes differs from that of E. fulvopilosus in many aspects, and the latter is structurally more similar to E. boliviensis , E. claripennis and E. nomadiformis sp. nov. In the holotype of E. bifasciatus obscuripes , the axillae are much shorter than in the holotype of E. fulvopilosus , the lateral margin of each axilla is smooth (not crenulate as in E. fulvopilosus ) and the axillae extend beyond the midlength of the mesoscutellum but not as far back as its posterior margin. By contrast, in the holotype of E. fulvopilosus the axillae extend as far back as the apex of the horizontal dorsal portion of the mesoscutellum, which is produced to two posteriorly directed teeth (not straight as in E. obscuripes ). Cockerell (1932) indicated that E. bifasciatus obscuripes is sufficiently distinct such that there may be justification to elevate its name to the taxonomic rank of species. In addition to the diagnostic morphological features that separate E. obscuripes from other similar species, its status as a separate species is supported by a separate BIN and large barcode sequence divergence (6.3%) from its nearest neighbor, E. bifasciatus (Supplementary File 3).

Friese (1925) described E. schmidti from both sexes, represented by two syntypes (one female and one male) deposited at the ZMB. Both specimens were examined, and the male is herein designated as the lectotype, the same sex as the primary types of E. bifasciatus obscuripes (herein recognized as belonging to the same species) and E. bifasciatus , the species to which it is most similar. The female syntype at the ZMB is herein designated as the lectoallotype. Below what is presumed to be Heinrich Friese’s original type label for the male specimen, which simply says “Type”, is a label that says “ LECTOTYPE ” and “desig. Melo, 2016”. The female bears a label that says “ PARALECTOTYPE ” and “desig. Melo, 2016”. Since Melo’s (2016) lectotype designations of Friese’s Neotropical Epeolus types cannot be traced to any publication, the designation for E. schmidti is made herein instead. Ferrari (2017) cited personal communication with G. Melo regarding the addition of the latter’s lectotype label to another of Friese’s syntype specimens (in this case Colletes nigritulus ), with 2015 given as the year of the designation, indicating that the designation (at that time) remained to be published.

The E. schmidti lectotype differs from the E. bifasciatus obscuripes holotype most notably in that the frontal protrusions are much less obvious and more densely punctate and that the frontal keel and metanotum are much more strongly protuberant. However, these differences fall within the range of variation observed among sequenced specimens that were assigned the same BIN. Also, in the E. schmidti lectotype T3–T6 have more pronounced bands of dense, bright yellow pubescence, which are less conspicuous and more sparsely hairy in the E. bifasciatus obscuripes holotype and multiple nontype specimens. These bands are absent altogether in most studied female specimens of E. obscuripes .

Figures

Fig. 1. Male genitalia of Epeolus spp. A. E. boliviensis Friese, 1908, in which the penis lacks lobes. B. E. claripennis Friese, 1908, in which the penis lacks lobes. C. E. fulvopilosus Cameron, 1902, in which the penis lacks lobes. D. E. fumipennis Say, 1837, in which the penis lacks lobes. E. E. hanusiae sp. nov., paratype (CNC 754086), showing the penis with a short, fleshy lateral lobe (blue arrow) on each side. F. E. luteipennis Friese, 1916, showing the penis with a long, fleshy lateral lobe (blue arrow) on each side. G. E. niger (Michener, 1954), in which the penis lacks lobes. H. E. obscuripes Cockerell, 1917 stat. nov., in which the penis lacks lobes. I. E. pulchellus Cresson, 1865, in which the penis lacks lobes. Scale bars = 0.5 mm.

Fig. 4. Head of female. A. E. bifasciatus Cresson, 1864 showing frontal area with impunctate, granulose protrusion (blue arrow) on each side. B. E. obscuripes Cockerell, 1917 stat. nov. showing frontal area with very weak/inconspicuous protrusion (blue arrow) on each side. C. E. scutellaris Say, 1824 showing dorsal protrusion (blue arrow) on each side where upper genal area meets vertexal area. D. E. pusillus Cresson, 1864 showing frontal and vertexal areas without protrusions. E. E. chamaesarachae Onuferko, 2018, paratype, showing vertexal area with two shiny, impunctate protrusions (blue arrows) on each side. F. E. flavofasciatus Smith, 1879 showing single (closely punctate) dorsal protrusion (blue arrow) on each side where upper genal area meets vertexal area. Scale bars = 1 mm.

Fig. 7 (opposite page). Occurrence records (yellow circles except where indicated otherwise) of: A. Epeolus boliviensis Friese, 1908. B. E. claripennis Friese, 1908. C. E. danieli (Genaro, 2014) comb. nov. (orange square) and E. luteipennis Friese, 1916. D. E. fulvopilosus Cameron, 1902. E. E. fumipennis Say, 1837. F. E. hanusiae sp. nov. G. E. niger (Michener, 1954). H. E. nomadiformis sp. nov. I. E. obscuripes Cockerell, 1917 stat. nov. J. E. odyneroides sp. nov. K. E. pulchellus Cresson, 1865.

Fig. 12. Pseudopygidial area of female, dorsal view. A. E. fumipennis Say, 1837, which consists of a basal impressed triangular or semicircular portion covered in silvery short, appressed setae differentiated from a transverse band of coppery short, appressed setae along the posterior-facing apical margin of T5. B. E. bifasciatus Cresson, 1864, which is more than twice as wide as long. C. E. obscuripes Cockerell, 1917 stat. nov., which is not more than twice as wide as long. The pseudopygidial area is the apical portion of T5 that changes slope from the rest of the tergum and is covered in short, coppery or silvery hairs uniform in length (posteromesad the light blue lines). Scale bars = 1 mm.

Fig. 17. Epeolus obscuripes Cockerell, 1917 stat. nov. A. Female, habitus, lateral view. B. Female, habitus, dorsal view. C. Male, habitus, lateral view. D. Female axillae and mesoscutellum, dorsal view (blue lines indicate the posterior extent of the axilla relative to the length of the horizontal dorsal portion of the mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm.