New species of the Eastern Hemisphere genera Afroheriades and Noteriades (Hymenoptera, Megachilidae), with keys to species of the former

Abstract New species of the rarely encountered megachilid genera Afroheriades Peters from South Africa, Afroheriades hyalinus sp. n., and Noteriades Cockerell from Myanmar and Thailand, Noteriades jenniferae sp. n. and Noteriades spinosus sp. n., are described and illustrated. The species are described to make their names available for forthcoming publications on phylogenetic studies of the family Megachilidae. Taxonomic notes and a comparative diagnosis for each genus are presented. Afroheriades hennigi (Peters) and Afroheriades reicherti (Brauns) are new junior synonyms of Afroheriades dolichocephalus (Friese). A key to the known species of Afroheriades is provided.


introduction
The purpose of this paper is to describe three new species of rarely encountered Eastern Hemisphere megachilid bees, one species of Afroheriades Peters from South Africa, and two species of Noteriades Cockerell from Southeast Asia. Specimens of these undescribed species have been used in recent molecular analyses (i.e., Praz et al. 2008;Cardinal et al. 2010;Litman et al. 2011); herein we describe them to make their names available for use in forthcoming publications on the phylogeny of Megachilidae. New synonyms and revised comparative diagnoses for both genera are also presented.
Afroheriades and Noteriades consist of small heriadiform or hoplitiform bees (< 10 mm in body length) whose biologies are largely unknown (a species of Noteriades was observed entering a hole in an upright log). Each genus contains a small number of species, although several new species have been recognized by one of us (TG) in the past few years. Afroheriades is known only from South Africa, whereas Noteriades is found in tropical and subtropical regions of sub-Saharan Africa, India, and Southeast Asia (Table 1). Both taxa are currently included in the Heriades group of genera of the tribe Osmiini (Michener 2007) but little is yet known about their phylogenetic relationships; Afroheriades has been thought to be related to Pseudoheri-ades Peters while Noteriades has been considered related to Heriades Spinola. In fact, Afroheriades was originally described as a distinctive monotypic subgenus of Pseudoheriades (Peters 1970) whereas Noteriades was described as a subgenus of Heriades (Cockerell 1931).
A recent comprehensive molecular analysis of the Osmiini (Praz et al. 2008) challenges these ideas regarding the taxonomy and relationships of Afroheriades and Noteriades. The molecular analysis supported the long suspected non-monophyly of Osmiini (e.g., Michener 2007), with Afroheriades, Noteriades, Pseudoheriades, and Ochreriades Mavromoustakis, excluded from the tribe (Praz et al. 2008). Afroheriades and Pseudoheriades formed a well-supported clade, but their phylogenetic position remained unclear; depending on the method employed in the analysis, they were either sister to the Anthidiini (parsimony analysis) or Megachilini + Noteriades (Bayesian analysis). Noteriades was consistently found to be the sister group of Megachilini (Praz et al. 2008). Such a relationship to Megachilini was first suggested by Griswold (1985) based on the presence of two apical spines on the outer surfaces of the fore and middle tibiae, and the modified hairs on the fifth sternum of the male. Except for the position of Noteriades, an on-going morphological phylogenetic analysis of Megachilidae (Gonzalez et al. in prep) is not concordant with the molecular analysis of Praz et al. (2008). In fact, Afroheriades, Pseudoheriades, and Ochreriades appear close to other "Osmiini" within a clade of most of the Heriades group (Gonzalez et al. in prep). Pending completion of that morphological analysis both genera are retained in the Osmiini.

Material and methods
Morphological terminology follows that of Michener (2007). Photomicrographs were taken using a Keyence® VHX-500F Digital Imaging System. Measurements were made with an ocular micrometer attached to a Leica® MZ12 stereomicroscope. Measurements in descriptions are for the holotype, with values for paratypes in parentheses. The abbreviations F, S, and T are used for antennal flagellomere, and metasomal sternum and tergum, respectively.
Institutional acronyms used herein are: Comments. Two species groups can be recognized in Afroheriades: one includes a rather robust form with the basal area of propodeum well below the level of the scutellum [A. primus (Peters)]; the other includes the remaining species, which have a more elongate mesosoma and the base of the propodeum at the same plane as the scutellum. If a phylogenetic analysis shows that these two groups are natural, they could be subgenerically separated as Afroheriades s.str. and Archeriades, as suggested by Michener (2007). For now, we treat them as species groups. Below are the species currently recognized in Afroheriades; the synonyms presented here are based on the study of the types by the senior author. Afroheriades capensis Griswold, in Michener (2007: 452), is a nomen nudum.

Afroheriades dolichocephalus (Friese)
Osmia dolichocephala Friese, 1925: 505   Diagnosis. The male of A. hyalinus is unique among Afroheriades in the S5 with erect, oval tuft of bristles (Figs. 4,6). In addition, males can be recognized by the combination of: antennal flagellum unmodified (Fig. 2); T7 without a lateral spine (Fig.  4); basal zone of the propodeum bounded posteriorly by a carina (Fig. 3); and anteriorfacing surface of T1 with fine, sparse, punctures smaller than those on dorsal face. The female can be easily recognized by the combination of: head about as long as broad (Fig. 12); clypeus sparsely punctate, not produced ventrally; hypostomal carina with anterior limb (i.e., portion of carina curving towards posterior margin of mandibular socket) longer than the straight, posterior limb (Fig. 13); basal zone of propodeum horizontal, scarcely longer than length of metanotum, broader laterally than medially; and anterior face of T1 very finely, sparsely punctate.
Integument smooth and shiny between punctures; punctation on interantennal area moderate, approximately one puncture width apart; on scutum moderate, approximately one puncture width apart; on disc of T3 sparse, more than one puncture width apart.
Body black except flagellomeres, tegula, legs dark brown. Wings hyaline with weak green and coppery highlights; veins, stigma, prestigma dark brown.
Etymology. The specific epithet makes reference to the thin, hyaline lamella of S2 that projects from the gradulus distally nearly to the posterior margin of the sternum. Basal zone of propodeum bounded posteriorly by carina (Fig. 3) Diagnosis. Noteriades can be easily separated from all other megachilid genera by the following combination of characters: small (4.5-10 mm), compact, hoplitiform bees with arolia present on all legs in both sexes; preoccipital carina complete; malar space linear; clypeus slightly projecting over clypeal-labral articulation; clypeus and usually supraclypeus with median longitudinal carina; pronotal lobe and omaulus carinate; scutellum carinate posteriorly; and propodeum vertical, without subhorizontal basal zone. Female. Mandible quadridentate, without differentiated cutting edges; T6 nearly vertical, except for apical hyaline flange. Male. Mandible bidentate; T1-T4 exposed dorsally, T5 and T6 curved ventrally, covering T7, S3 and remaining sterna; T6 with-out preapical carina; S1 subapically produced over its apical margin, forming a double carina; volsella distinct, with well-developed digitus and cuspis, and heavily sclerotized teeth resembling those of short-tongued bee families and within the Megachilidae, Pararhophites (Fideliinae).

Keys to species of Afroheriades
Comments. Griswold (1985;1994) provided the first synoptic list of species of Noteriades, transferring most of them from Heriades. The species list presented in Table  1 is provisional; an on-going revision of the genus by the authors has revealed several undescribed species as well as new synonyms. Diagnosis. This species is most similar to N. spinosus, sharing with it the following characters: clypeus with median longitudinal carina extending onto supraclypeal area; juxtantennal carina absent (Fig. 16); and scutum with unmodified posterolateral angle, not spined or sharply angled, without tomentum obscuring integument (Fig. 17). It can be easily separated from N. spinosus by: larger body size; head slightly broader than long (Fig. 16); and scutellum with posterior margin rounded, not projecting laterally into small, curved spine.

Noteriades jenniferae
Description. Female: Body length 9.0 mm (8.0 mm); forewing length 5.6 mm (5.4 mm). Head slightly broader than long (Fig. 16); compound eyes subparallel, about 2.4 times longer than broad, 1.3 times wider than gena in profile; interantennal distance 1.9 times median ocellar diameter, 1.6 times antennocular distance; interocellar distance 1.4 times median ocellar diameter, shorter than ocellocular distance; ocelloccipital distance slightly longer than median ocellar diameter; clypeus about twice as broad as long, flat in profile, distal margin crenulate, median longitudinal carina distinct, extending onto supraclypeal area; juxtantennal carina absent; scape 2.4 times longer than wide, pedicel 1.3 times longer than broad, about twice as long as F1, F1 broader than long, about as long as F2, remaining flagellomeres progressively increasing in length, apical flagellomere longer than broad. Scutum with posterolateral angle not spined or sharply angled (Fig. 17); scutellum with posterior margin broadly rounded, without lateral spine.
Mandible and labrum with outer surfaces dull, minutely roughened, irregularly punctate, punctures slightly coarser on basal half of mandible, condylar and outer ridges smooth and shiny; clypeus with small (one-sixth to one-fifth times median ocel-lar diameter), contiguous punctures, smaller, denser on disc, smooth, shiny between punctures; supraclypeus with punctures as on sides of clypeus, punctures becoming larger (one-fourth times median ocellus diameter), slightly sparser on frons and superior paraocular area; inferior paraocular area with dense punctures smaller than on clypeus; vertex and gena with large (one-third to one-half times median ocellar diameter), coarse punctures, largest on gena; hypostomal area with punctures shallow, faint, separated by two puncture widths or less, integument smooth and shiny between punctures as on face. Mesosoma somewhat dull, imbricate between punctures except smooth and shiny on anterior surface of mesepisternum, metepisternum and most segments of legs; scutum uniformly punctate, punctures nearly contiguous, small (onefourth to one-fifth times median ocellar diameter); axilla and scutellum with large (one-third to one-half times median ocellar diameter), coarse punctures as on gena (Figs. 15, 17); pronotum with punctures smaller than on scutum; anterior surface of mesepisternum minutely and densely punctate dorsally, impunctate ventrally; mesepisternum with large (one-fourth times median ocellar diameter) punctures separated by a puncture width or more, punctures becoming smaller and denser ventrally; metepisternum with small, contiguous punctures as on lateral surface of pronotum; metanotum and propodeal triangle shagreened, punctures faint, sparse on metanotum, remaining areas of propodeum with small, contiguous punctures; coxae finely, densely punctate, remaining areas of legs with large impunctate areas, punctures larger, sparser on hind legs. Metasomal terga and sterna smooth and shiny to finely imbricate, punctures smaller, denser than on scutum, punctures coarser, contiguous on distal terga (particularly T6), smaller, scattered on anterior face of T1.
Integument black throughout except tegula, legs, and metasomal sterna largely dark reddish brown. Wings light brownish with weak green or coppery highlights; veins, stigma, prestigma dark brown.
Pubescence in general short, sparse, white except yellowish on labrum, mandible, clypeal margin, and inner surfaces of tarsi; mandible and labrum with outer surfaces covered by dense, minute, erect hairs; paraocular area, dorsal surface of pronotum, pronotal lobe, lateral surfaces of coxae and propodeum with long, minutely branched hairs obscuring integument or nearly so; T1-T4 (also on T5 in paratype) with apical hair bands, medially interrupted on T1 and T2.
Male: As in female except longer and denser body pubescence, apical hair bands present on basal two terga only, coarser punctation, stronger median longitudinal carina of clypeus, and the following: Body length 8.0 mm; forewing length 5.1 mm. Head 1.2 times broader than long (Fig. 19); compound eyes converging below, 1.4 times wider than gena in profile; interantennal distance about twice as long as median ocellar diameter and antennocular distance; interocellar distance 1.6 times median ocellar diameter, subequal to ocellocular distance; clypeus 1.6 times broader than long, with distal margin not as crenulate as in female; antennal flagellum long, surpassing distal margin of scutellum, scape robust, 1.6 times longer than wide, pedicel broader than long, longer than F1, F1 about one-third length of F2, remaining flagellomeres much longer than broad. S5, S6 and S8, and genital capsule as in figures 20-23.
Etymology. This species is named after Mrs. Jennifer Lyman Strange, for her courage.

Noteriades spinosus
Diagnosis. This species is most similar to N. jenniferae (see above). It can be easily separated from that species and all other Noteriades by the combination of: relatively small body size, head shape (about as long as broad, Fig. 25), and posterior margin of scutellum with small, curved spine laterally (Fig. 27).
Description. Female. Body length 7.8 mm (7.1-7.8 mm); forewing length 4.8 mm (4.2-4.8 mm). Head about as long as broad (Fig. 25); compound eyes subparallel, about 2.4 times longer than broad, 1.3 times wider than gena in profile; interantennal distance about twice as long as median ocellar diameter, 1.5 times antennocular distance; interocellar distance 1.6 times median ocellar diameter, about as long as ocellocular distance; ocelloccipital distance 1.6 times median ocellar diameter; clypeus 1.7 times broader than long, flat in profile, distal margin crenulate, slightly projecting over clypeal-labral articulation, median longitudinal carina distinct on basal half, absent on distal half (thin but extending along clypeal length in some paratypes), extending onto supraclypeal area; juxtantennal carina absent; scape 2.6 times longer than wide, pedicel 1.4 times longer than broad, about twice as long as F1, F1 broader than long, about as long as F2, remaining flagellomeres progressively increasing in length, apical flagellomere longer than broad. Scutum with posterolateral angle not spined or sharply angled; scutellum with posterior margin rounded, laterally with small, curved spine (Fig. 27).
Mandible and labrum with outer surfaces dull, minutely roughened, irregularly punctate, punctures coarser on basal half of mandible, condylar and outer ridges smooth and shiny; clypeus with small (one-sixth to one-fifth times median ocellar diameter), rather shallow, contiguous punctures, smaller on disc, smooth, shiny between punctures; supraclypeus with coarser punctures than on clypeus, punc-tures becoming larger (one-third times median ocellus diameter) on frons, superior paraocular area, and vertex; inferior paraocular area with dense, shallow, smaller punctures than on clypeus; gena with large (one-half to two-thirds times median ocellar diameter), coarse punctures; hypostomal area with punctures small (about one-third times median ocellar diameter), shallower than on gena. Mesosoma weakly shiny, weakly imbricate between punctures except smooth and shiny on anterior surface of mesepisternum, metepisternum, propodeum and most segments of legs; scutum uniformly punctate (Fig. 26), punctures contiguous, small (one-fourth to one-third times median ocellar diameter); axilla as on scutum; scutellum with large (one-half times median ocellar diameter), coarse punctures; pronotum with smaller punctures than on scutum; anterior surface of mesepisternum minutely and densely punctate dorsally, impunctate ventrally; mesepisternum with small (one-third times median ocellar diameter) punctures separated by a puncture width or more, integument becoming shinier with smaller and denser punctures ventrally; metepisternum with coarse, contiguous punctures as on scutum; metanotum imbricate with sparse, coarse punctures; propodeal triangle smooth and shiny, impunctate, except imbricate and sparsely punctate basally; lateral and posterior surfaces of propodeum weakly shiny, with punctures contiguous, small (about-fifth times median ocellar diameter or less), faint; coxae finely, densely punctate, remaining areas of legs with large impunctate areas, punctures larger, sparser on hind legs. Metasomal terga and sterna weakly shiny, finely imbricate, punctures smaller, denser than on scutum, punctures coarser, contiguous on distal terga (particularly on T6), smaller, scattered on anterior face of T1.
Integument black throughout except tegula, legs, and metasomal sterna largely dark reddish brown. Wings light brownish with weak green or coppery highlights; veins, stigma, prestigma dark brown.
Pubescence in general short, sparse, white except yellowish on labrum, mandible, clypeal margin, and inner surfaces of tarsi; mandible and labrum with outer surfaces covered by dense, minute, erect hairs; paraocular area, dorsal surface of pronotum, pronotal lobe, mesepisternum near wing bases, lateral surfaces of coxae, and lateral surface of propodeum with long, minutely branched hairs obscuring integument or nearly so; T1-T4 with apical hair bands, medially interrupted on T1 and T2.
Male: Unknown. Etymology. The specific epithet makes reference to the curved posterolateral spine of the scutellum that distinguishes this species.
Comments. Specimens of this species were used in the molecular analysis of the Osmiini by Praz et al. (2008).