A review of the cleptoparasitic bee genus Epeolus Latreille, 1802 (Hymenoptera: Apidae) in the Caribbean, Central America and Mexico Author Onuferko, Thomas M. text European Journal of Taxonomy 2019 2019-10-07 563 1 69 journal article 25281 10.5852/ejt.2019.563 4de0fe4e-219b-432f-9d36-43d2f4a41835 3476748 6F6E082D-0675-49C1-A603-F7BABB546C46 Epeolus claripennis Friese, 1908 Figs 1B , 5A , 7B , 8 Epeolus claripennis Friese, 1908: 88 ( ♀ ). Trophocleptria odontothorax Michener, 1954: 126 ( ♀ ), syn. nov. Proposed common name Clear-winged epeolus. Diagnosis Together with the morphological features that are diagnostic for the ‘ Trophocleptria group’, the following in combination can be used to tell E. claripennis apart from all other Epeolus : the axillae are crenulate along the lateral margin, each with a large tooth near the base ( Fig. 8D ); the mesoscutellum has a pair of posteriorly directed teeth ( Fig. 8D ); the mesopleura are coarsely punctate, each with sparser punctures ventrolaterally (many i≥2d) than in the upper half, with interspaces dull due to tessellate surface microsculpture ( Fig. 5A ); the fore wings are deeply infuscate basally and (unusually for Epeolus ) clear apically ( Fig. 8 A–B); and the metasomal terga lack pale pubescence ( Fig. 8 A–C). Epeolus claripennis most closely resembles E. niger ( Michener, 1954 ) in that both species are almost entirely black and at least the fore wings of females are deeply infuscate basally and clear apically, but in E. niger there is no large tooth laterally near the base of each axilla, the mesoscutellum does not have a pair of posteriorly directed teeth and the mesopleura are more finely punctate, each with punctures more or less equally dense throughout (few i≥2d). Material examined Primary type material BOLIVIA • ♀ , E. claripennis holotype ; Cochabamba Department , Tarata ; 1900 ; ZMB . PANAMA • ♀ , Tro. odontothorax holotype ; Chiriquí , El Volcán Chiriquí ; 27 Feb. 1936 ; F.E. Lutz leg.; AMNH AMNH _ IZC 00324262 . Secondary type material PANAMA • 1 ♂ , Tro. odontothorax allotype ; Coclé , El Valle de Antón ; 1 Apr. 1945 ; C.D. Michener leg.; AMNH AMNH _ IZC 00324263 . DNA barcoded material with BIN-compliant sequences Available. BOLD:ADB 1637 . Specimens examined and sequenced: COSTA RICA • 1 ♂ ; San José , Montecarlo (MORA ref. site); 9.3419° N , 83.6041° W ; 14 Feb. 2005 ; H.T. Ngo leg.; BOLD sample ID: CCDB-28315 F06; PCYU . Non-barcoded material COSTA RICA • 1 ♀ ; Puntarenas , F. Las Cruces ( 6 km S of San Vito ); 21 or 25 Aug. 1976 ; E.M. Fisher leg.; LACM LACM ENT 363332 • 1 ♂ ; San José , Montecarlo (MORA ref. site); 9.3420° N , 83.6039° W ; 7 May 2005 ; H.T. Ngo leg.; PCYU • 1 ♀ ; same collection data as for preceding; 13 May 2005 ; H.T. Ngo leg.; PCYU • 1 ♂ ; same collection data as for preceding; 13 Jul. 2005 ; H.T. Ngo leg.; PCYU • 1 ♀ ; same collection data as for preceding; 2 Sep. 2005 ; H.T. Ngo leg.; PCYU • 1 ♀ ; same collection data as for preceding; 9 Jan. 2006 ; H.T. Ngo leg.; PCYU • 1 ♀ ; San José , Quizarra ( Sr. Rojas’ farm); 12 May 2005 ; H.T. Ngo leg.; PCYU • 2 ♀♀ ; San José , San Isidro de El General ; Feb. 1993 ; F.D. Parker leg.; BBSL . PANAMA • 1 ♂ ; Panamá , 15 km N of El Llano (Camino Cartí); 10 May 1981 ; R.W. Brooks leg.; KUNHM SM 0738700 • 1 ♀ ; same collection data as for preceding; KUNHM SM 0738701 . VENEZUELA • 1 ♀ ; Aragua , Estación Biológica de Rancho Grande ( Portachuelo Pass ); 10.3500° N , 67.6833° W ; 9 Mar. 1995 ; R. Brooks leg.; KUNHM SEMC 1248355 • 3 ♂♂ ; same collection data as for preceding; 4 Jun. 1998 ; J. Ashe , R. Brooks and R. Hanley leg.; KUNHM SM 0124175, SM0124178 , SM0124179 • 2 ♀♀ ; same collection data as for preceding; 4 Jun. 1998 ; J. Ashe , R. Brooks and R. Hanley leg.; KUNHM SM 0124176, SM0124177 • 1 ♂ ; Carabobo , Henri Pittier National Park ( Portachuelo Pass ); 10.3475° N , 67.6878° W ; 15 Sep. 2008 ; J. Skevington leg.; PCYU • 1 ♂ ; Lara , Parque Nacional Yacambú ( 16.1 km SE of Sanare ); 9.7000° N , 69.5850° W ; 2 Jun. 1998 ; J. Ashe , R. Brooks and R. Hanley leg.; KUNHM SM 0122036 . Redescription Female MEASUREMENTS. Length 8.5 mm ; head length 2.0 mm; head width 2.4 mm ; fore wing length 7.5 mm . INTEGUMENT COLORATION. Mostly dark brown to black; notable exceptions as follows: at least partially ferruginous on mandible, antenna, tegula and legs. Mandible with apex darker than rest of mandible; preapical tooth lighter than mandibular apex (difficult to see in E. claripennis holotype because mandibles closed; described from non-type specimens). Flagellum, except left F1 and F2, missing in E. claripennis holotype , but brown and (except F1) slightly lighter than partially dark brown (otherwise orange) scape, pedicel and F1, primarily due to extensive pilosity on flagellum, in Tro. odontothorax holotype and multiple non-type specimens. Wing membrane subhyaline, basally dusky. 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 with narrow band of ferruginous short, appressed setae along posterior margin. Mesoscutum without pale tomentum in E. claripennis holotype , but with some ferruginous short, appressed setae along posterior margin in multiple non-type specimens. Mesopleuron nearly bare, except along margins. Metanotum with tomentum discolored or rubbed off in E. claripennis holotype , but sparser medially and uniformly light brown/pale ferruginous in Tro. odontothorax holotype and multiple non-type specimens. Dorsum of metasoma without bands of pale tomentum. T5 laterally with long, erect simple setae. T5 with pseudopygidial area lunate, its apex twice as wide as medial length, with basal impressed triangular portion covered in gray-brown short, appressed setae differentiated from transverse band of coppery to silvery short, appressed setae along posterior-facing apical margin. S5 with apical fimbria of coppery to silvery hairs extending beyond apex of sternum by ~½ MOD. SURFACE SCULPTURE. Punctures dense, but those of head and mesosoma sparser in some areas, larger, deeper and more distinct. Labrum with larger and sparser punctures (most i>1d) than clypeus (i≤1d). Impunctate spot lateral to lateral ocellus absent in E. claripennis holotype , but shiny spot present in multiple non-type specimens. Mesoscutum, mesoscutellum and axilla very coarsely and densely rugosepunctate. Tegula densely punctate mesally (i=1–2d), sparsely punctate (i>2d) to impunctate along margins. Mesopleuron with denser (i<1d) punctures in upper half than ventrolateral half (many i≥1d), interspaces dull due to tessellate surface microsculpture; mesopleuron with punctures similar in size throughout. Metasomal terga with punctures very fine, dense (i≈1d), evenly distributed on disc. Fig. 8. Epeolus claripennis Friese, 1908 . A . Female, habitus, lateral view. B . Female, habitus, dorsal view. C . Male, habitus, lateral view. D . Female axillae and mesoscutellum, dorsal view (blue arrow indicates one of two posteriorly directed tooth-like projections of mesoscutellum). Scale bars: A–C = 3 mm; D = 0.5 mm. STRUCTURE. Preapical tooth acute. Labral apex with pair of small denticles (separated by shallow concavity), each preceded by longitudinal carina. Frontal keel strongly raised. Frontal area with pair of discrete, sparsely punctate granulose protrusions, each located near upper mesal margin of compound eye. 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 1.5 × greatest width. F2 noticeably longer than wide (L/W ratio = 1.5). Preoccipital ridge separated from hypostomal carina by about 1.5–2 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 weakly bigibbous, depressed along posterior margin beneath distinct overhanging ridge produced to pair of posteriorly directed teeth. Axilla large, its lateral margin more than half as long as mesoscutellar width (AL/MSCW ratio = 0.7) and tip extending beyond apex of horizontal dorsal portion of mesoscutellum; axilla with tip conspicuously diverging from side of mesoscutellum, distinctly hooked, and axilla with free portion approximately half its medial length; axilla with lateral margin crenulate, with large tooth near base, and carinate but relatively straight. Mesopleuron with carina delineating its anterior and lateral surfaces. Fore wing with three submarginal cells. Pygidial plate apically truncate. Male Description as for female except for usual secondary sexual characters and as follows: F2 shorter, but still longer than wide (L/W ratio = 1.2); pygidial plate apically rounded, with larger and deeper punctures, closely punctate throughout; S3–S5 with much longer (>1 MOD) coppery to silvery subapical hairs, those of S4 and S5 curved. Distribution Central America and northwestern to western-central South America ( Fig. 7B ). Ecology Host records Unknown. Floral records Labels of examined voucher specimens indicate floral associations with Coffea and Psychotria L. ( Rubiaceae ). Remarks The examined primary type specimen of E. claripennis bears a label that simply says “Type”, which is presumed to be Heinrich Friese’s original type label. Below it is a label that says “ LECTOTYPUS ” and “A. Roig Alsina, 1989”. The lectotype designation cannot be traced to any publication, nor does it seem warranted since Friese’s (1908) original description of the species gives no indication that it was described from more than one (female) specimen. Moure et al . (2007) refer to this specimen as the holotype , not the lectotype , and in the present study it is also recognized as such. Although the holotypes of E. claripennis and Tro. odontothorax are from widely separated localities, in Bolivia and Panama , respectively, there are no apparent differences between the two specimens that fall outside the range of variation observed among conspecifics that were, by contrast, collected at the same place and time. Although only a single BIN-compliant sequence is available for a specimen identified as E. claripennis , a partial sequence 297 bp in length obtained from a visibly different bee (see Figs 8 , 16 ), herein considered to be a separate species ( E. nomadiformis sp. nov. ), has been matched to it with 99.7% similarity. Thus, it seems unlikely that additional barcodes from specimens exhibiting the diagnostic features of E. claripennis should uncover any cryptic species. Epeolus claripennis exhibits very little intraspecific morphological variation and appears to be one of the more widespread and commonly collected species in the ‘ Trophocleptria group’. The extensive black coloration and apically clear fore wings of this species (presumably the latter is what inspired the epithet ‘ claripennis ’) and E. niger give both the distinctive appearance of Parachartergus R. von Ihering, 1904 ( Hymenoptera : Vespidae : Polistinae ). These features are also shared with various tropical stingless bees (e.g., Trigona Jurine, 1807 spp.), some of which are known to be in mimetic complexes that include a diversity of aculeates, including other bees ( Smith-Pardo 2005 ), and even nonhymenopterous insects such as flower flies ( Diptera : Syrphidae ) ( Reemer 2013 ).