Revision of the Neotropical Laelius (Hymenoptera: Bethylidae) with notes on some Nearctic species

. Laelius Ashmead includes 54 known nominal species worldwide. We present a revision of the Neotropical species of Laelius and provide keys for the identification of males and females of these species. We propose fourteen nomenclatural changes and additions. For the Neotropics, seven new species from Brazil are described and illustrated: Laelius arryni sp. nov. which has clypeus bearing median lobe with apical margin straight; propodeal disc with inner discal carina complete; aedeagus with apical lobes touching each other and inner margin with hook-shaped process, L. baratheoni sp. nov. which has propodeal disc with second pair of discal carinae incomplete; genitalia with paramere wing-shaped; with long setae around the margin and cuspis wide, L. lannisteri sp. nov. which has median lobe of clypeus with apical margin straight; mesopleuron with mesopleural suture distinct; genitalia with paramere slender and angled, cuspis wide, L. martelli sp. nov. which has pronotum with series of foveae posteriorly; genitalia with paramere slender and placed dorsally, aedeagus not reaching apex of paramere, L. targaryeni sp. nov. which has head with clypeus wide and with apical margin rounded; propodeal disc without longitudinal ridge between median and inner discal carinae; forewing with r-rs & Rs longer than Rs+M, L. tullyi sp. nov. which has head densely punctate; pronotal disc, mesoscutum and mesoscutellum rugulose; mesopleuron with mesopleural suture not fused with posterior fovea, L. starki sp. nov. which has mesoscutellar sulcus straight; mesopleuron with all foveae closed; genitalia with paramere club-shaped and with aedeagus wide. For the Nearctic region, Laelius areolatus (Rosmann & Azevedo, 2005) is a new junior synonym of Laelius pedatus (Say, 1836) and lectotypes for L. rufipes Ashmead, 1893, L. nigripilosus Ashmead, 1893, L. tricarinatus Ashmead, 1893, L. trogodermatis Ashmaed, 1893, and Mesitius nigripilosus Ashmead, 1895 are designated. We also propose the name Laelius billi Barbosa & Azevedo nom. nov. for Mesitius nigripilosus Ashmead, 1895 and re-erect Laelius huachucae (Evans, 1965) from L. nigripilosus (Ashmead, 1895). As result, Laelius now includes 61 species, 15 of which are from the Neotropical region. fol-lows: body length, measured from apex of clypeus to posterior margin of last metasomal segment, excluding male genitalia or female sting; (LH) length of head, measured in frontal view from vertex crest to median apical margin of clypeus; (WH) maximum width of head, measured in frontal view, including eyes; (WF) minimum width of frons, measured in frontal view, usually at the level of the base of eye bases; (HE) maximum height (length) of eye, measured in lateral view; (OOL) ocello-ocular line, measured in laterodorsal view, the shortest distance from eye top to posterior ocellus; (WOT) width of ocellar triangle, measured in frontal view, maximum width, including ocelli; (DAO) diameter of anterior ocellus, measured in frontal view; distance of ocellar triangle to vertex, measured in dorsoposterior view; distance from posterior ocellus to vertex crest; (VOL) vertex-ocular line, measured in dorsal view, distance eye top to vertex crest.

Measurements and indices used in this study are as follows: body length, measured from apex of clypeus to posterior margin of last metasomal segment, excluding male genitalia or female sting; (LH) length of head, measured in frontal view from vertex crest to median apical margin of clypeus; (WH) maximum width of head, measured in frontal view, including eyes; (WF) minimum width of frons, measured in frontal view, usually at the level of the base of eye bases; (HE) maximum height (length) of eye, measured in lateral view; (OOL) ocelloocular line, measured in laterodorsal view, the shortest distance from eye top to posterior ocellus; (WOT) width of ocellar triangle, measured in frontal view, maximum width, including ocelli; (DAO) diameter of anterior ocellus, measured in frontal view; distance of ocellar triangle to vertex, measured in dorsoposterior view; distance from posterior ocellus to vertex crest; (VOL) vertex-ocular line, measured in dorsal view, distance from eye top to vertex crest.
The descriptions were elaborated with DELTA (Descriptive Language for Taxonomy) as proposed by DALLWITZ et al. (1993). All species were described based on the holotype, and the characteristics that are not present in the holotype are mentioned in the topic "Variations" of the species' description.
In the material examined section, additional information is in brackets. Label information is reproduced in the original language, except for the country.
The male genitalia of L. gracilis, L. multilineatus, and L. muesebecki are described and illustrated for the first time. As we had the opportunity to study types of several New World species, we also present comments on some Nearctic species.
Material Variation. Body length 2.23-3.86 mm; mesosoma and metasoma castaneous; mesoscutum with notauli incomplete; propodeal disc without longitudinal ridge between median and inner discal carina, without longitudinal ridge between inner and second pair of discal carina, and mesopleuron with mesopleural suture with inner margin not striate.
Remarks. This species is similar to L. gracilis ) by having the frontal angle of the ocellar triangle obtuse; ocelli large; pronotal disc wider than long and with series of foveae posteriorly; mesoscutum with notauli complete; propodeal disc with five carinae; mesopleuron with all foveae closed; genitalia with paramere with apex rounded, cuspis slender and reaching beyond the apex of digitus, aedeagus dilated at median region. However, L. arryni has five apical teeth on the manidible, with the two basal ones wider than the others; clypeus bearing median lobe with apical margin straight; propodeal disc with inner discal carina complete; mesopleural suture distinct; paramere uniform, aedeagus with apical lobes touching each other and inner margin with hook-shaped process, and base of apodeme slightly curved outwards, whereas L. gracilis has five equally wide apical teeth on the mandible; clypeus bearing median lobe with apical margin angulate; propodeal disc with inner discal carina incomplete; mesopleural suture indis-tinct; paramere S-shaped, aedeagus with apical lobes not touching each other and inner margin without hook-shaped process, and apodemes parallel to each other. [92][93][94] Description. Holotype (Fig. 5) (Figs 7,8). Pronotal disc 0.4 x as long as wide, coriaceous, trapezoidal, side straight, with series of foveae posteriorly, with few setae, dorsal setae long and thick. Mesoscutum coriaceous, with long and thick setae between notaulus and parapsidal signum; notaulus conspicuous, complete, convergent posteriorly. Parapsidal signum narrow, as conspicuous as notaulus, shorter than notaulus, not reaching anterior margin of mesoscutum, convergent posteriorly. Mesoscutellum coriaceous, with post-sulcal, fore, mesal and hind setae; mesoscutellar sulcus narrow. Propodeal disc 0.7 x as long as wide, with five carinae; median carina complete; space between median and inner discal carinae with longitudinal ridge; inner discal carinae incomplete; space between inner and second pair of discal carinae without ridge; second pair of discal carinae incomplete; sublateral carina present and conspicuous; lateral carina present and conspicuous; followed by short striae; posterior carina complete. Declivity of propodeum not strigate, with median carina. Mesopleuron with acropleural fovea elongate and closed; anterior fovea closed; lower fovea closed; posterior fovea closed; mesopleural suture with inner margin foveolate. Variation. Body length 2.81-3.28 mm; propodeal disc with space between median and inner discal carinae without longitudinal ridge.

Laelius baratheoni
Remarks. This species is similar to L. muesebecki  by having the median lobe of clypeus with apical margin angulate; pronotum wider than long and with a series of foveae posteriorly; mesoscutum with notauli complete; propodeal disc with median carina complete, and inner and second pair of discal carinae incomplete; mesopleuron with all foveae closed and mesopleural suture distinct; aedeagus elliptical. However, L. baratheoni has the frontal angle of the ocellar triangle obtuse; propodeal disc with second pair of discal carinae incomplete; genitalia with paramere wing-shaped and with long setae around the margin; cuspis wide and apodemes parallel to each other, whereas L. muesebecki has the frontal angle of the ocellar triangle right; propodeal disc with second pair of discal carinae complete; genitalia with apex of paramere rounded and without long setae on the margin; cuspis slender and apodemes diverging from each other.
Remarks. This species is similar to L. gracilis  by having the mandible with equally wide apical teeth; pronotum wider than long with a series of foveae posteriorly; mesoscutum with notauli complete; propodeal disc with five carinae, median carina complete, inner and second pair of discal carinae incomplete; mesopleuron with all foveae closed. However, L. lannisteri has the median lobe of clypeus with apical margin straight; frontal angle of ocellar triangle right; mesopleuron with mesopleural suture distinct; genitalia with paramere slender and angled, cuspis wide and aedeagus with apical lobes touching each other, whereas L. gracilis has the median lobe of clypeus with apical margin angulate; frontal angle of ocellar triangle obtuse; mesopleuron with mesopleural suture not distinct; genitalia with paramere S-shaped, cuspis slender and aedeagus with apical lobes not touching each other.

Figs 17-20
Description. Holotype (Fig. 17). Female. Body 3.08 mm. LFW 1.88 mm. Color. Wings hyaline; palpi, wing venation, trochanters, protibia, mesotibia, metatibia and tarsi light castaneous; scape, pedicel and mandible castaneous; flagellum, tegula, procoxa, mesocoxa, metacoxa, profemur, mesofemur, metafemur and metasoma dark castaneous; head and mesosoma dark castaneous to nearly black. Head (Fig. 18 Remarks. This species is similar to L. centratus (Say, 1836) by having the head longer than wide, with eyes large; mesoscutum with notauli incomplete and as wide as parapsidal signum; mesoscutellum with mesoscutellar sulcus narrow; propodeal disc with five carinae, inner and second discal carinae incomplete; mesopleuron with only anterior fovea. However, L. targaryeni has the head with clypeus wide and with apical margin rounded; ocellar triangle compact, with frontal angle right and with ocelli large; propodeal disc without longitudinal ridge between median and inner discal carinae; forewing with r-rs & Rs longer than Rs+M, whereas L. centratus has the head with clypeus not wide and with apical margin straight; ocellar triangle not compact, with frontal angle obtuse and with ocelli small; propodeal disc with longitudinal ridge between median and inner discal carinae; forewing with r-rs & Rs shorter than Rs+M.
Remarks. This species was known from the U.S.A. (Arizona and California) and Mexico (Durango) and is now recorded for the first time from the Dominican Republic (San Cristóbal), Nicaragua (Granada and San Juan del Sur), Trinidad and Tobago (Arima), and Brazil (Amazonas, Bahia, Distrito Federal, Espírito Santo, Minas Gerais, Pará, Rio Grande do Norte and São Paulo).
Although the list of studied material is fairly large, we did not observe significant intraspecific variation. Only the proportion between LH x WH varies among the specimens analyzed. This species is promptly recognized by having the head remarkably longer than wide, LH is usually at least 1.2 x WH, whereas in other species it is 1.0 x WH or less.
Material examined. Remarks. The elongate process on the inner margin of the dorsal portion of the basiparamere is unique among Bethylidae.
Laelius mellipes   Remarks. This species was known from the U.S.A. (southeastern states) and it is now recorded for the first time from Brazil (Bahia, Minas Gerais, and Espírito Santo).
Laelius minutulus  Figs 57-60 Rhabdepyris (Rhabdepyris) minutulus Evans, 1965: 72, 77-76. Laelius minutulus  Remarks. This species was known from Peru and now it is recorded for the first time from Brazil (Minas Gerais and São Paulo). The small number of specimens and the narrow distribution of the species indicate that L. minutulus is rare. After EVANS (1965) described it, no further specimens were found or analyzed. This is the second record of L. minutulus, but it corresponds to only two specimens.
Diagnosis. Female. Body 3.8 mm. Dark castaneous to nearly black. Mandible with five sharpened apical teeth, two lowermost teeth wider than others. Clypeus with median lobe trapezoidal. Frons coriaceous. WH 0.98 x LH; WF 0.57 x WH; WF 1.12 x HE; VOL 0.54 x HE; OOL 1.21 x WOT; posterior ocellus distant from vertex crest 0.5 x DAO. Frontal angle of ocellar triangle right. Vertex slightly convex. Mesoscutum with notaulus complete, very wide and convergent posteriorly; parapsidal signum incomplete. Propodeal disc 1.15 x as long as wide; with five discal carinae; median discal carina complete; with longitudinal ridge between median and inner discal carina; inner discal carina complete; with longitudinal ridge between inner and second discal carina; second pair of discal carinae incomplete. Declivity of propodeum with median carina. Mesopleuron with lower and posterior fovea fused; with mesopleural suture and posterior fovea fused.
Diagnosis. Male. Body 3.0 mm. Black. Mandible with five sharpened apical teeth, two lowermost teeth wider than others. Clypeus with median lobe trapezoidal, with apical margin angulate. Frons coriaceous. WH 1.0 x LH; WF 0.62 x WH; WF 1.29 x HE; VOL 0.5 x HE; OOL 1.15 x WOT; posterior ocellus distant from vertex crest 0.4 x DAO. Frontal angle of ocellar triangle almost right. Vertex slightly convex. Pronotal disc with series of foveae posteriorly. Mesoscutum with notaulus complete, very wide and convergent posteriorly; parapsidal signum incomplete. Propodeal disc 1.14 x as long as wide; with five discal carinae, median discal carina complete; with longitudinal ridge between median and inner discal carina; inner discal carina complete; with longitudinal ridge between inner and second discal carina; second discal carina incomplete. Declivity of porpodeum with median carina. Mesopleuron with lower and posterior fovea fused; with mesopleural suture and posterior fovea fused. Genitalia (Figs 108-110). Paramere much longer than basiparamere, wide, apex rounded, base narrow, dorsal margin convex, ventral margin concave, dorsal and apical margins strongly setose; volsella with cuspis wide basally progressively narrowing to apex in lateral view, apex as high as paramere apex; aedeagus narrow and straight, apex nearly as high as paramere apex, base of apodeme curved outwards. Remarks. This species was known from Canada, U.S.A., and Mexico. The known distribution indicates that L. centratus probably is restricted to the Nearctic region. It is promptly recognized by its broad ocellar triangle.
The type of this species, which is supposed to be at USNM, has apparently been lost. Laelius trogodermatis was described based on a couple, but both syntypes deposited at USNM are females. Here we considered the specimen with code USNM #2187, deposited at the primary type collection, as the lectotype.
Diagnosis. Female. Body 4.65 mm. Dark castaneous to nearly black, forewing hyaline. Mandible with five apical sharpened teeth, two lowermost teeth wider than others. Clypeus with median lobe trapezoidal with apical margin angulate. Frons strongly coriaceous. WH 0.92 x LH; WF 0.59 x WH; WF 1.57 x HE; VOL 0.7 x HE; OOL 1.1 x WOT, frontal angle of ocellar triangle right, posterior ocellus distant from vertex crest 0.5 x DAO. Vertex slightly convex. Pronotal disc without series of foveae posteriorly. Mesocutum with notaulus complete, wide, and converging posteriorly; parapsidal signum incomplete. Propodeal disc 1.38 x as long as wide; with five discal carinae, median discal carina complete; with longitudinal ridge between median and inner discal carina; inner discal carina complete; with longitudinal ridge between inner and second pair of discal carinae; second discal carina incomplete; sublateral and lateral carinae conspicuous. Declivity areolate with median carina. Mesopleuron lower and posterior fovea fused; posterior fovea not fused with mesopleural suture.