A revision of the bee genus Ceratina, subgenus Rhysoceratina (Hymenoptera, Apidae, Xylocopinae)

the subgenus Rhysoceratina of Ceratina is restricted to South America, with known species in Argentina, Brazil, colombia, Paraguay, and uruguay. nine species are recognized, four of which are described as new: C. canaliculata, C. pusilla, C. mendozina, and C. nitidifrons. Lectotypes are designated for C. catamarcensis Schrottky, 1907, and C. volitans Schrottky, 1907. A phylogenetic analysis supports the monophyly of the subgenus and, within it, of two major species groups. A key to the species, descriptions, distributional data, and illustrations are provided.

the relationships within Ceratina have been studied by rehan et al. (2010) on the basis of one nuclear and two mitochondrial genes. these authors included representatives of 14 of the known subgenera, among them the new World Zadontomerus, Ceratinula, and Calloceratina. these three subgenera resulted closely related in the analysis, forming a clade together with the oriental subgenera Ceratinidia cockerell and Lioceratina Vecht. of the remaining new World subgenera, Crewella may well be suspected to belong in the same clade, based on its close morphological affinities with Calloceratina, but the relationships of Rhysoceratina remain unknown.
the subgenus Rhysoceratina was proposed by Michener (2000) to include a distinctive group of South American Ceratina which bear a transverse preapical carina on the sixth metasomal tergum of the female. the subgenus is also characterized by the strongly punctate body, the presence of a preoccipital carina, the basitibial plate of the female indicated by a strong tooth, the seventh tergum of the male with a midapical marginal projection, and by the presence of graduli on the second to fourth metasomal terga and second to third metasomal sterna of both sexes. According to Michener (2000), J. S. Moure had recognized the group, and "Rhysoceratina" was left as an unpublished name in collections. this name was used as a nomen nudum, for C. volitans Schrottky and C. stilbonota Moure, by Sakagami & Laroca (1971). the group was also recognized by Hirashima (1971), without naming it, as one of two new World groups with wax plates restricted to the female second sternum, other groups having wax plates on the second and third sterna. the other group of new World species with wax plates restricted to the second sternum is Neoclavicera, which has been recently revised by roig-Alsina (2013).
Most species of Rhysoceratina are known from Argentina and southeastern Brazil, but they also occur in uruguay and Paraguay, and one species is newly described here from colombia. Moure (2007), in his catalog of neotropical bees, recognized five species within Rhysoceratina; four of these are considered valid in the present contribution. Ceratina correntina Schrottky, unplaced to subgenus to date, is shown to be also a member of the subgenus, and four species are described as new, totaling the number of species to nine.
Information on the biology of species of Rhysoceratina is scarce. Besides a few notes on flower visitation (Holmberg, 1886;Jörgensen, 1909Jörgensen, , 1912aJörgensen, , 1912bSchlumpberger et al., 2009) the only available work is the study of Sakagami & Laroca (1971) on the nesting biology of Ceratina volitans Schrottky in southern Brazil. they described the nest structure and partitions, the pollen loaves and the eggs on them, and recorded duration of immature stages, phenological data, plants used as nest substrata, and flower visits.

MAtErIAL And MEtHodS
Specimens studied are deposited in the following institutions: california Academy of Sciences, San Francisco (cAS); Facultad de ciencias, universidad de la república, Montevideo (FcM); Instituto y Fundación Miguel Lillo, tucumán (IFML); Museo Argentino de ciencias naturales "Bernardino rivadavia," Buenos Aires (MAcn); Museo de La Plata, La Plata (MLP); natural History Museum, London (London). Acronyms are used to indicate depositories of the specimens. the following abbreviations are used. the maximum diameter of the median ocellus (Mod) is used as a reference to express the length of the pubescence and other structures. the metasomal terga (t) and sterna (S) are identified with Arabic numerals. the sex of the specimens is indicated by F, female, and M, male.
the phylogenetic analysis and the illustration of the cladogram were made with the aid of the programs tnt, v. 1.1 (Goloboff et al., 2003) and Mesquite (Maddison & Maddison, 2016). For outgroup comparison the following species were included in the analysis: C. (Ceratinula) oxalidis Schrottky, C. (Ceratinula) arizonensis cockerell, C.
(Zadontomerus) calcarata robertson, C. (Zadontomerus) acantha Provancher, C. (Neoclavicera) parvimacula roig Alsina, and C. (Neoclavicera) richardsoniae Schrottky. Rhysoceratina Michener, 2000: 599. type species: Ceratina montana, Holmberg, 1886. Diagnosis. Small species 4.8-8.5 mm long, with black to weakly metallic integument and yellow marks restricted to head, pronotal lobe and legs. Paraocular areas with punctures at least along the inner margin of eyes, but extensively punctate in most species; ocellocular areas always punctate. circumantennal area shallowly depressed. Preoccipital carina present. Mandible of male with two teeth. Pronotum with dorsolateral angle rounded to weakly carinate; lateral carina absent, or present below the transpronotal sulcus in a species group. Basal area of the metapostnotum slightly depressed, rugose or rugoso-striate, longer than metanotum. Anterior and middle tibiae with single dorsoapical tooth. Basitibial plate of female indicated by strong tooth. Sixth tergum of female with strong transverse preapical carina and weak median longitudinal impression. Metasomal graduli present on second to fourth terga and second to third sterna of both sexes. Wax plate present on female second sternum. Seventh tergum of male with midapical marginal projection.

Rhysoceratina
Phylogenetic analysis the analysis was based on 25 morphological characters, listed below. table 1 is a data matrix of character states for the nine species of Rhysoceratina and the six species of other subgenera selected as outgroups. trees were calculated with the implicit enumeration algorithm implemented in tnt, using unordered, unweighted characters. A single most parsimonious, fully resolved tree was obtained (length 48 steps, Fig. 1). Standard bootstrap (1000 replicates) values are shown below branches in figure 1. the resampling procedure collapses the branch leading to volitans and the montana species group, resulting in a tricotomy, and mendozina is shown as sister to montana and catamarcensis.
According to the analysis, two species groups can be recognized within the subgenus Rhysoceratina. the canaliculata species group, which also includes stilbonota and nitidifrons, is characterized by the peculiar channel-like, longitudinal depression of the paraocular area below the antennal socket, present in both sexes. the three species have a distinct lateral pronotal carina below the transpronotal sulcus, and an impunctate disc of the scutum behind the admedian lines, but these are plesiomorphic conditions for Rhysoceratina as a whole in the present analysis. the montana species group, which also includes mendozina, catamarcensis, pusilla, and correntina, is supported by two characteristics of the males: the dense sternal fringes on the apical margin of S4 and S5 (although lost in pusilla), and the sparsely punctate, depressed areas on S6. Ceratina volitans is intermediate between the two groups, sharing with canaliculata and allied species a peculiar depressed t5 in the female, but lacking a lateral pronotal carina and having an extended pilosity on the male hind femur as in the montana group.   5.-Yellow of paraocular area forming short spot at level of antennal insertion (Fig. 4A). Wings weakly infuscate, with dark brown veins. Gena usually black, though in some specimens a very dark brown spot can be seen. t4-t5 with slender setae (Fig. 2E)  Diagnosis. this species is closely related to C. stilbonota, from which it is separated by the yellow mark on the pronotal lobe in both sexes, the dark gena of the female, the broadened setae on the female t4-t5, and the short vestiture of the male, the ventral hairs in front of the mid coxa being 0.5-0.7 times Mod (these hairs in C. stilbonota being 1.0-1.7 times Mod). Ceratina canaliculata has been collected in the same places and at the same time as C. volitans, to which it is similar in size and color pattern. Besides the features mentioned in the key, canaliculata is separated by the sparser punctation the of ocellocular area and the nearly impunctate lower 2/3 of the gena.

Key to species of Ceratina
Female. Length, 5.0 mm (holo and paratype); length of forewing, 3.9-4.2 mm (holotype 3.9 mm). Color. Black, with dark bluish metallic tints, and coppery reflections on supraclypeal area. tegula brown. clypeus with dark brown apical band, and triangular yellow mark reaching above to level of tentorial pits. Paraocular yellow mark long (holotype, 0.75 times eye length; paratype, 0.72), reaching below lateral lower angle of clypeus and surpassing above antennal socket; mark narrower than diameter of antennal socket, not touching inner orbit of eye. Gena with dark Scutellum with punctures on disc leaving some shiny interspaces among them, which are larger than a puncture diameter.  Schrottky, 1907: 474, 480 (Lectotype female, catamarca, Argentina, Museo de La Plata, present designation). Schrottky, 1909: 271. Schrottky, 1913: 252. Schlumpberger et al., 2009: 1497 catamarcensis: Moure, 2007Moure, : 645. rasmussen et al., 2009 Schrottky's original description (1907) allows the identification of this species with certainty. He mentioned the dark clypeus, the yellowishbrown wing membranes and veins, and the long paraocular yellow band, all of which are characteristic of this Rhysoceratina. the species was already recognized as a Rhysoceratina by Moure (2007). A female specimen with a handwritten label of Schrottky "ceratina catamarcensis Schrottky" and a printed label "catamarca" is preserved in the Jörgensen collection in the Museo de La Plata, La Plata, Argentina. It undoubtedly belongs to the type series, and it is here designated as the lectotype. the specimen lacks the prothorax; the head, which lacks the antennae, has been glued to the mesothorax; the metasoma is glued onto a card on the same pin.
Schrottky did not mention in the original description any yellow on the gena, usually present in this species, but in some specimens, and also in the lectotype, the genal mark is dark brown, barely distinguishable.
Diagnosis. this species is readily distinguished by the color of its wings, with yellowish-brown membrane and veins. the female dark clypeus is only shared with C. montana, from which it is separated by the long yellow paraocular marks and the yellowish-brown wings. the male t7 has a strong, rounded, middle apical projection.
Female. Length, 5.7-8.5 mm; length of forewing 4.5-6.0 mm (lectotype, 4.7 mm). Color. Black, with dark olive green metallic tints, in some specimens with coppery reflections on head, scutum and scutellum. outer half of tegula and apical tarsomeres brown. clypeus without yellow, in some specimens with dark brown apical band. Paraocular yellow mark long (0.78-0.84 times eye length, lectotype, 0.82), reaching below lateral lower angle of clypeus and surpassing above antennal socket; mark wider than antennal socket diameter, touching in most of its length inner orbit of eye. Gena with yellow mark close to lower third of eye; in some specimens mark orange-brown to dark brown (as in lectotype). Foretibia with basal yellow spot followed by yellow stripe. Wings with yellowishbrown membrane; veins and pterostigma yellowish brown. Structure. Paraocular area punctate from eye to epistomal suture; next to subantennal suture punctate or, sometimes, with narrow smooth area as long as diameter of antennal socket. distance between lateral ocellus and preoccipital carina 1.55-1.85 times ocellar diameter. ocellocular area with dense, irregularly distributed punctures, close to ocellus separated by 0.1-0.5 puncture diameter, further apart close to eye. Gena close to upper half of eye with even punctures, separated approximately by a puncture diameter. Proportion between length of eye and length of second segment of labial palpus 1:0.70. dorsolateral angle of pronotum weakly carinate; lateral carina absent, except sometimes rudimentary on lower sector. Anterior surface of pronotum entirely punctate, punctures reaching dorsolateral angle. Scutum between admedian line and notaulus with punctures irregularly distributed, separated by 0.5-2.0 times their diameter, interspaces shiny; disc of scutum posteriorly to admedian lines with few scattered punctures. Scutellum with punctures sparse medially, leaving large shiny interspaces, but not forming transverse impunctate band. dorsal area of metapostnotum rugulose, basally with strong rugae, irregular medially and reaching posterior margin. Forecoxa with basal carina; lateral projection of forecoxa with apex squarish, mucronate, anterior margin concave and posterior margin sinuous. t4-t5 with broadened setae. t5 sculptured near to apical margin; apicalmost punctures separated from margin by approximately their diameter. t5 medially moderately flattened, base of tergum not abruptly elevated. t6 with weak longitudinal median impression.
Male. Length, 5.8-7.5 mm; length of forewing 4.8-5.9 mm. Color. Similar to that of female, with following parts yellow: long paraocular band, separated from subantennal suture by less than its width, extended from upper level of antennal socket to lateral lower angle of clypeus; inverted t-shaped mark on clypeus, pointed above, reaching upper margin of clypeus; triangular spot on base of mandible; basal spot on labrum; spot on pronotal lobe; forefemur with apical spot; foretibia with yellow stripe; small basal spot on middle and hind tibiae. Structure. Ventral surface of hind femur with long hairs on basal 3/5. t7 with median apical projection strong, rounded, approximately as broad as long. S4 and S5 with dense apical fimbriae formed by hairs 1.0-1.5 times Mod. S6 with median apical notch moderate and paramedian depressed area transverse, distinct.  In spite of the loss of the holotype, Schrottky's brief original description allows the identification of this species as a Rhysoceratina. Among the characters that he mentions , the strongly punctate head and the basal area of the metapostnotum with longitudinal rugae preclude the possibility that the species would belong to the subgenus Ceratinula. the dark body color, small size (6 mm), and the presence of a longitudinal impression on the t6, indicate that it is not a Calloceratina, the South American species of which are all rather large, bright metallic green, and with an unimpressed t6. the small size, the t1 finely punctate, and the long paraocular yellow band combined with a lack of yellow on the gena, differentiate C. correntina from all species of Crewella present in Argentina. the disc of the scutum with punctures, a characteristic stressed by Schrottky, indicates that it is not a species of Neoclavicera, the species of which have a characteristic broad, impunctate area posteriorly to the end of the admedian lines. the pattern of pale marks and the sculpture described by Schrottky fully agree with two morphospecies of Rhysoceratina, both with the scutum extensively punctate. one of them, with olive-green body and certain reddish tints, as is described by Schrottky, is here interpreted as C. correntina. the second morphospecies, with blue body, is described here as C. pusilla.
Diagnosis. Species closely related to C. pusilla, from which it is separated by the shape of the lateral projection of the forecoxa of the female, with rounded apex, by the yellow paraocular band of the female, truncate below, reaching the lateral lower margin of the clypeus, by the dense apical fringe of S5 of the male formed by several rows of hairs, and by the S6 of the male with deep median apical notch and drop-shaped paramedian depressed areas. this shape of the male S6 markedly differs from that of all other species of Rhysoceratina.
Female. Length, 5.6-6.5 mm; length of forewing, 4.1-4.5 mm. Color. Black, with dark olive green metallic tints, in some specimens with reddish reflections on head, scutum and scutellum. outer half of tegula and apical tarsomeres brown. Yellow spot on clypeus small, rounded. Paraocular yellow mark long (0.80-0.83 times eye length), reaching below lateral lower angle of clypeus and surpassing above antennal socket; mark wider than antennal socket diameter, touching in most of its length inner orbit of eye. Gena black. Pronotal lobe with yellow spot. Foretibia dorsally with basal yellow spot followed by longitudinal yellow band. Wings weakly infuscate; veins and pterostigma brown. Structure. Paraocular area punctate from eye to epistomal and subantennal sutures. distance between lateral ocellus and preoccipital carina 0.8-1.0 times ocellar diameter. ocellocular area densely punctate; punctures separated by less than a puncture diameter, coalescent in part. Gena close to upper half of eye with punctures dense, separated by less than a puncture diameter. Proportion between length of eye and length of second segment of labial palpus 1:0.60. dorsolateral angle of pronotum rounded, without carina; lateral carina absent. Anterior surface of pronotum entirely punctate, punctures reaching dorsolateral angle. Scutum between admedian line and notaulus with punctures dense, separated by less than their diameter, interspaces shiny; disc of scutum posteriorly to admedian lines with abundant punctures separated by 0.5-1.0 times their diameter; scutum with only two slim, longitudinal impunctate bands following notauli. Scutellum with punctures dense, without impunctate median areas. dorsal area of metapostnotum rugulose, basally with strong rugae, irregular medially and reaching posterior margin. Forecoxa without basal carina; lateral projection of forecoxa with apex rounded, anterior and posterior margins straight. t4-t5 with slender setae. t5 sculptured near to apical margin; apicalmost punctures separated from margin by approximately their diameter. t5 medially moderately flattened, base of tergum not abruptly elevated. t6 with weak longitudinal median impression.
Male. Length, 5.7-6.1 mm; length of forewing 4.3-4.4 mm. Color. Similar to that of female, with following parts yellow: narrow paraocular band, separated from subantennal suture by its own width, extended from upper level of antennal socket to lateral lower angle of clypeus; transverse apical mark on clypeus, in some specimens pointed above, but not reaching above half of clypeus; triangular spot on base of mandible; basal spot on labrum; spot on pronotal lobe; forefemur with apical spot; tibia with yellow stripe; in some specimens with small basal spot on middle and hind tibiae. Structure. Venter of male mesothorax with paramedian sparsely punctate swellings anterior to median depression. Ventral surface of hind femur with long hairs on basal half. t7 with median apical projection short, broad, approximately three times broader than long. S4 with few apical hairs, not forming fimbria, S5 with apical fimbria dense, formed by several rows of hairs, 0.8-1.0 times Mod. S6 with deep median apical notch and paramedian depressed area distinct, drop-shaped.  Diagnosis. this species can be recognized by the yellowish-brown anterior tibiae and tarsi, and by the sparse punctation of the head and thorax, being the anterior surface of the pronotum impunctate and polished close to the anterolateral angle. the clypeus of the female has a yellowish-brown apical band, and the small yellow spot is sometimes split in two; the yellow paraocular band is broad, above nearly touching the antennal socket. the t7 of the male has a broad apical projection, similar to that of C. correntina and C. pusilla, species from which it is separated by the sparse punctation. Holmberg's original description of this species leaves no doubts about its identity, in spite of the loss of the types. Holmberg (1886) mentioned for the female the presence of a transverse carina on the sixth tergum, as well as a median longitudinal impression. this kind of tergum is unique to the subgenus Rhysoceratina. other characters mentioned, as the black clypeus and the shape and position of the yellow paraocular spot, are diagnostic of the species.
Diagnosis. Females are distinguished by the black clypeus and gena, and by the oblong paraocular spot at level of the antennal socket, shorter than in any other Rhysoceratina, less than half of eye length. the male is distinguished by the small apical triangular projection of t7 and by the densely punctate scutellum, the punctures leaving no smooth areas between them.
Female. Length 5.5-8.2 mm; length of forewing 4.4-5.3 mm. Color. Black, with dark blue metallic tints. outer half of tegula and apical tarsomeres 2-4 light brown. clypeus without yellow, in some specimens with dark brown apical band. Paraocular yellow spot short (0.38-0.42 times eye length), oblong, slightly surpassing upper level of antennal socket above, and not reaching level of upper margin of clypeus below. Gena black, some specimens with difuse dark brown spot in the same position as yellow mark of other species. Foretibia with basal yellow spot. Wings weakly and evenly infuscate; veins and pterostigma dark brown. Structure. Paraocular area punctate from eye to epistomal suture; next to subantennal suture punctate or, sometimes, with narrow smooth area as long as diameter of antennal socket. distance between lateral ocellus and preoccipital carina 1.15-1.25 times ocellar diameter. ocellocular area densely punctate; punctures separated by less than a puncture diameter. Gena close to upper half of eye with punctures dense, separated by less than a puncture diameter. Proportion between length of eye and length of second segment of labial palpus 1:0.68. dorsolateral angle of pronotum rounded, without carina; lateral carina absent, except sometimes rudimentary on lower sector. Anterior surface of pronotum entirely punctate, punctures reaching dorsolateral angle. Scutum between admedian line and notaulus with punctures irregularly distributed, separated by 0.5-2.0 times their diameter, interspaces shiny; disc of scutum posteriorly to admedian lines with few scattered punctures. Scutellum with punctures dense, without impunctate median areas. dorsal area of metapostnotum rugulose, basally with strong rugae, irregular medially and reaching posterior margin. Forecoxa with basal carina; lateral projection of forecoxa with apex squarish, mucronate, anterior margin concave and posterior margin sinuous. t4-t5 with slender setae. t5 sculptured near to apical margin; apicalmost punctures separated from margin by approximately their diameter. t5 medially moderately flattened, base of tergum not abruptly elevated. t6 with weak longitudinal median impression.
Male. Length, 5.8-6.3 mm; length of forewing 4.0-5.1 mm. Color. Similar to that of female, with following parts yellow: narrow paraocular band, separated from subantennal suture by its own width, extended from upper level of antennal socket to lateral lower angle of clypeus; inverted t-shaped mark on clypeus, pointed above, reaching upper margin of clypeus; triangular spot on base of mandible; central spot on labrum; basal spot and dorsal band on foretibia; in some specimens with small basal spot on middle and hind tibiae; spot on pronotal lobe. Structure. Ventral surface of hind femur with long hairs on basal 3/5. t7 with median apical projection triangular, small. Apical fimbriae of S4-S5 dense, hairs 1.0-1.5 times Mod. S6 with median apical notch moderate and paramedian depressed area transverse, distinct. Female holotype. Length, 6.6 mm; length of forewing, 5.0 mm. Color. Black, with dark olive green metallic tints, and reddish reflections on supraclypeal area, scutum and metapostnotum. Antennal scape, inner basal part of foretibia, and tibial spurs reddish brown; tegula and apical tarsomeres dark brown. clypeus with trapezoidal yellow mark reaching above to level of tentorial pits. Paraocular yellow mark short (0.31 times eye length), reaching below lateral lower angle of clypeus and barely surpassing above upper level of clypeus. Gena with small yellow spot above middle of eye. Pronotal lobe with yellow spot. Forefemur with apical yellow spot; foretibia with basal yellow spot followed by yellow stripe. Wings infuscate, forewing darker along anterior margin of marginal cell; veins and pterostigma brown. Structure. Paraocular area next to subantennal and epistomal sutures depressed, impunctate, smooth, forming a trough reaching below to level of tentorial pit; such a trough 0.6-0.7 times as wide as paraocular area; this is the only species of Rhysoceratina described by Schrottky for which he mentions in the original desciption the lateral carina of the sixth tergum of the female, diagnostic of the subgenus.  mentions that he studied nine females and two males from Villa Encarnación, Paraguay. Moure (2007) andrasmussen et al. (2009) indicate that syntypes of this species are housed at the national Museum of natural History, Washington, and at the Museu de Zoologia, São Paulo. Specimens of C. volitans were not found in the former collection upon my request. two further syntype specimens have been located in the california Academy of Sciences, San Francisco, and another one in the Museo de La Plata, La Plata, in the P. Jörgensen collection. the specimen in La Plata is broken, only the mesosoma remaining on the pin; it bears the labels "Villa Encarnación 7-IX-1905" and "ceratina volitans Schrottky," both in Schrottky's handwriting; I have labeled it as paralectotype. the two syntypes housed at the california Academy of Sciences belonged, according to the labels, to Paul Herbst, another German student of bees, who was contemporary with Schrottky and lived in chile at that time. the specimens are as follows. one female in good condition with two labels in Schrottky's handwriting: "Villa Encarnación 14-I-1906" and "ceratina volitans Schrottky" plus a printed label "P. Herbst collection Ex reed." I designate hereby this specimen as the lectotype and label it accordingly. the second specimen is a female that lacks the metasoma, and the head is glued to one of the labels; it is designated as paralectotype. It also has three labels: "Villa Encarnación 4-IX-1905" and "ceratina volitans Schrottky," both in Schrottky's handwriting, and a printed label "P. Herbst collection Ex reed." Diagnosis. this species is distinguished, among species with punctate paraocular areas, by the disc of the scutum impunctate, or with a few punctures posteriorly to the admedian lines. the clypeus has a brown apical band, and the setae on t4-t6 are broadened, as in C. mendozina, species from which it is differentiated by the impunctate or scarcely punctate disc of the scutum, by the female t5 depressed medially on the apical 2/3, by the narrower yellow paraocular band of the female, and by the apical margin of the female t5 with a median area of sparse punctures. other features useful for distinguishing the species and mentioned by Schrottky in the original description are, for the female, the long yellow paraocular band narrowed below, the orangebrown mark on the gena, and the light brown under surface of the flagellum; for the male he mentions the short, sharply defined, middle apical projection of t7.
Female. Length, 5.6-6.5 mm (lectotype 5.7 mm); length of forewing, 4.0-4.5 mm (lectotype 4.2 mm). Color. Black, with dark olive green metallic tints, in some specimens with coppery reflections on head, scutum and scutellum. outer half of tegula and apical tarsomeres brown. clypeus with more or less developed brown apical band, and triangular yellow mark frequently reaching above to level of tentorial pits. Paraocular yellow mark long (0.70-0.75 times eye length, lectotype 0.73), reaching below lateral lower angle of clypeus and surpassing above antennal socket. Gena entirely dark, or in some specimens including lectotype with brown mark close to lower third of eye. Pronotal lobe with yellow spot. Foretibia with basal brown spot followed by brown stripe. Wings evenly infuscate; veins and pterostigma brown. Structure. Paraocular area punctate from eye to epistomal suture; below antennal socket, and next to subantennal suture, usually with triangular impunctate area tapering below, as long as 1.5 times diameter of antennal socket. distance between lateral ocellus and preoccipital carina 1.00-1.25 times ocellar diameter. ocellocular area with dense punctures, separated by 0.2-0.5 puncture diameter, further apart close to eye. Gena close to upper half of eye with even punctures, separated approximately by a puncture diameter. Proportion between length of eye and length of second segment of labial palpus 1:0.63. dorsolateral angle of pronotum carinate; lateral carina usually rudimentary on lower sector. Anterior surface of pronotum entirely punctate, punctures reaching dorsolateral angle. Scutum between admedian line and notaulus with punctures irregularly distributed, separated by 0.5-2.0 times their diameter, interspaces shiny; disc of scutum posteriorly to admedian lines impunctate, or with 2-5 grouped punctures inmediately following lines. Scutellum with punctures sparse medially, leaving paramedian large shiny interspaces (lectotype), in some specimens forming transverse impunctate band. dorsal area of metapostnotum rugulose, basally with strong rugae, irregular medially and reaching posterior margin. Forecoxa with basal carina; lateral projection of forecoxa with apex squarish, mucronate, anterior margin concave and posterior margin sinuous. t4-t5 with broadened setae. t5 sculptured near to apical margin, except a median area of sparse punctures. t5 medially flattened (smaller specimens) to depressed (larger specimens), at different level of elevated basal third of tergum. t6 with longitudinal median impression.
Male. Length, 5.2-6.0 mm; length of forewing 3.9-4.5 mm. Color. Similar to that of female, with following parts yellow: long paraocular band, separated from subantennal suture by a distance larger than its width, extended from upper level of antennal socket to lateral lower angle of clypeus; inverted t-shaped mark on clypeus, pointed above, reaching upper margin of clypeus; triangular spot on base of mandible; basal spot on labrum; spot on pronotal lobe; forefemur with apical spot; foretibia with yellow stripe. Structure. Ventral surface of hind femur with long hairs on basal 3/5. t7 with median apical projection short, triangular. S4 and S5 with apical fimbriae formed by sparse hairs 0.8-1.1 times Mod. S6 with median apical notch moderate, without paramedian depressed areas.