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Discovery of the bee tribe Tarsaliini in Arabia (Hymenoptera, Apidae), with the description of a new species PDF

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AMERICAN MUSEUM NOVITATES Number 3877, 28 pp. March 20, 2017 Discovery of the Bee Tribe Tarsaliini in Arabia (Hymenoptera: Apidae), with the Description of a New Species MICHAEL S. ENGEL,1 ABDULAZIZ S. ALQARNI,2 AND MOHAMED A. SHEBL3 ABSTRACT The uncommonly encountered bee tribe Tarsaliini (Apinae) is recorded from the Arabian Peninsula for the first time, and based on a new species of the genus Tarsalia Morawitz. The tribes Ancylaini and Tarsaliini are diagnosed and their differences highlighted. Tarsalia kinda- hensis Engel, new species, is described and figured from the eastern portion of the Najd of central-eastern Saudi Arabia (Qassim and Riyadh regions). The new species is most similar to T. mimetes (Cockerell), known only from Egypt and Sudan, as well as the larger T. persica (Warncke) from Iran. These three species are morphologically and largely geographically dis¬ tinct from the remainder of the genus, and are segregated into a new subgenus, Astibomelissa Engel. An updated and corrected checklist of the genera and subgenera of bees recorded from Saudi Arabia is appended. 1 Division of Invertebrate Zoology (Entomology), American Museum of Natural History; Division of Ento¬ mology, Natural History Museum, and Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence. 2 Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia. 3 Department of Plant Protection, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia; Department of Plant Protection, Faculty of Agriculture, Suez Canal University, Ismailia, Egypt. Copyright © American Museum of Natural History 2017 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3877 INTRODUCTION The bee species of Tars alia Morawitz are some of the more uncommonly encountered and little studied among Eurasian and northeastern African Anthophila. The genus was first described by Morawitz (1895) from material collected by explorer naturalist Dmitry K. Glazu¬ nov (1869-1913) in Razavi Khorasan Province, Iran (at the time a southern part of “Greater Khorasan” of Persia), and near the present-day borders of Turkmenistan and Afghanistan. Subsequently, Popov (1935) described a related species from southern Tajikistan (eastern Khat- lon Province), and Baker (1971) a species from southern India (Karaikal), while subspecific forms of Morawitzs and Popov’s species were documented from the eastern Mediterranean (Pittioni, 1950; Mavromoustakis, 1952). Popov (1935) also noted a similarity between Tarsalia and the genus Ancyla Lepeletier de Saint Fargeau, considering that some species of the latter may eventually prove to belong within the former. Nearly a half century later, Warncke (1977) took this similarity a step further and united Ancyla and Tarsalia as subgenera within a single genus, and subsequently provided a revision for the combined group (Warncke, 1979). Baker (1998) corrected some errors of previous authors when describing another Indian species (Konkan), as well as transferring a further two overlooked taxa described in other genera, and thereby bringing the known fauna at the time to seven species (table 1). For nearly the last 20 years no further taxonomic treatments have considered species of the genus in any detail, nor has considerable material improved our impoverished understanding of the distribution or biology of Tarsalia. The general placement of Tarsalia among other groups of Apidae has also been challenging for several reasons, not the least of which is a dearth of material. Subsequent to Morawitzs (1895) description, Tarsalia were rarely considered by melittologists, usually overlooked (e.g., Ashmead, 1899) or mentioned only in passing as a genus of anthophorines (e.g., Friese, 1896: under his concept of “Podaliriinae”). Cockerell (1933) noted a general similarity between a species of Tarsalia and the New World tribe Exomalopsini, although he did not recognize his species as congeneric with Morawitzs genus and instead placed the Sudanese taxon in Tetralo- nia Spinola (Eucerini). This same author again noted a similarity between Tarsalia and the Exomalopsini, as well as Emphorini, when describing a species of Ancyla from Algeria, but again wrongly attributed its genus—this time to Ancyloscelis Latreille. It was Popov (1935) who first drew clearer attention to the affinities between Tarsalia and Ancyla, and documented the peculiar asymmetry in the male seventh metasomal sternum of the former. In his world clas¬ sification of bees, Michener (1944) brought Tarsalia and Ancyla together formally into a tribe Ancylaini4 (Michener, 1944), although the characters purportedly shared between these genera were problematic and applied only to one or the other of the two (Baker, 1998). Michener (1944) did not have material from which to base his character assessments, relying on generally inadequate published descriptions, and even excluded the tribe from his final phylogenetic scheme due to the dearth of substantive information. Popov (1949) subsequently suggested a relationship between Ancylaini and Exomalopsini (echoing the notion of Cockerell, 1933), as 4 The tribe was originally established as Ancylini (Michener, 1944), but was emended to Ancylaini to remove homonymy with a family group among the Gastropoda (Engel et al., 2008; ICZN, 2010). 2017 ENGEL ET AL.: ARABIAN TARSALIA 3 TABLE 1. Hierarchical classification of tribes Tarsaliini and Ancylaini, with summaries of species distribu¬ tions and synonyms (see also Baker, 1998). Tribe Tarsaliini Engel, 2015 Genus Tarsalia Morawitz, 1895 Subgenus Astibomelissa Engel, n. subgen. T. kindahensis Engel, n. sp. Saudi Arabia: Najd T. mimetes (Cockerell, 1933) Egypt, Sudan T. persica (Warncke, 1979) Iran: Khuzestan, Fars Subgenus Tarsalia s.str. T. ancyliformis Popov, 1935 Turkey, Israel, Turkmenistan, Uzbekistan, Tajikistan =T. ancyliformis mediterranea Pittioni, 1950 Cyprus, Sardinia T. cellularis (Cameron, 1898) India: Deccan Plateau T. deccana Baker, 1971 [1972] India: Eastern and Western ghats T. hirtipes Morawitz, 1895 Turkey, Uzbekistan, Iran: Razavi Khorasan, Fars, Khuzestan =T. hirtipes cyriaca Mavromoustakis, 1952 Cyprus T. strobilanthae Baker, 1998 India: Konkan Coast Tribe Ancylaini Michener, 1944 Genus Ancyla Lepeletier de Saint Fargeau, 1841 A. asiatica Friese, 1922 Lebanon, Turkey A. brevis Dours, 1873 Algeria = A. punica Friese, 1922 Tunisia A. cretensis Friese, 1902 Crete = A. cretensis kilikia Warncke, 1979 Turkey A. holtzi Friese, 1902 Bulgaria, Cyprus, Greece, Iraq, Iran: Fars =A. holtzi anatolica Warncke, 1979 Turkey A. nitida Friese, 1902 Armenia, Azerbaijan, Greece, Turkey = A. nitida nigricornis Friese, 1902 A. oraniensis Lepeletier de Saint Fargeau, 1841 Algeria, Morocco, Tunisia = A. flavilabris (Lucas, 1849) = A. heterodoxa (Cockerell, 1937) A. orientalica Warncke, 1979 Greece, Iraq, Syria, Turkey A. stolli Friese, 1922 Iran: Fars, Lebanon, Syria, Turkey well as Eucerini. The tribe has remained little understood (e.g., Michener and Moure, 1957), and attempts to place Ancylaini among other apine bees have given varied results but consis¬ tently place Ancyla and Tarsalia near Eucerini, though they are not always as a monophyletic group (Roig-Alsina and Michener, 1993; Silveira, 1993a, 1995; Baker, 1998; Praz and Packer, 2014; Plant and Paulus, 2016). Baker (1998) went so far as to constrain Ancylaini to only its type genus, removing Tarsalia as a basal branch within Eucerini and noting the birecurved gradulus on the female second metasomal sternum in both. Nonetheless, Michener (2000, 2007) retained Ancylaini in its traditional sense, correctly noting that several of Bakers (1998) characters were questionably defined or coded. Recently, the genus Tarsalia was again removed from Ancylaini, although this time placed within its own tribe, Tarsaliini, and as a putative sister group to Eucerini (Engel, 2015). Including a species established herein, the Tarsaliini and Ancylaini each consist of eight, uncommonly encountered species (table 1), the latter occurring in Mediterranean Europe, West¬ ern Asia, and northern Africa, while the former encompasses roughly the same area although 4 AMERICAN MUSEUM NOVITATES NO. 3877 more restricted in northeastern Africa and extending into Central Asia and India. The known records of Tarsalia reveal a rather disjunct distribution—the European/Western and Central Asian occurrences, the Indian species, and those species in northeastern Africa. Here we report the discovery of the tribe Tarsaliini for the first time from the Arabian Peninsula, expanding the distribution of this lineage into the region and closing the otherwise purported gap between those taxa in northeastern Africa and the Asiatic fauna. In addition, we provide revised diagnoses of the tribes Ancylaini and Tarsaliini, highlighting their distinctive traits. An initial survey of the supraspecific groups of bees occurring in Saudi Arabia was outlined by Engel et al. (2013) with the intention of encouraging further collecting throughout the peninsula. Subsequently, several genera have been newly recorded for the country, or even the entire peninsula, much as is done here with Tarsalia, and we accordingly append an updated list (appendix). MATERIAL AND METHODS Specimens of Tarsalia were taken at two localities in central Saudi Arabia during the 2011 and 2013 field seasons, while other attempts to locate individuals and nests were unsuccessful. The available material is deposited in the Division of Entomology (Snow Entomological Col¬ lections), University of Kansas Natural History Museum, Lawrence, Kansas (SEMC), and the King Saud University Museum of Arthropods, Plant Protection Department, College of Food and Agriculture Sciences, King Saud University, Riyadh, Saudi Arabia (KSMA). Comparative material of all other species of Tarsalia, with the exception of Tarsalia cellularis (Cameron), and diverse Ancyla and Eucerini were examined from the collections of SEMC. Descriptions are provided in the context of improving identifications of and species-level treatments of bees (Engel, 2011; Gonzalez et al., 2013). For the descriptive sections, the mor¬ phological terminology is largely adapted from Michener (2007) and Engel (2001), and from Snelling (1981) in terms of setal orientation. However, we have used a slightly revised terminol¬ ogy for tibial spurs, particularly the protibial calcar (= “fibula” sensu Snodgrass, 1956). Follow¬ ing Engel (2009) we use the term rachis for the main, longitudinal body of the spur along its entire length, and as commonly employed in biology for any spine or shaft forming the prin¬ ciple axis. For the protibial calcar, which forms a portion of the foreleg strigilis or “antenna cleaner,” we restrict the term malus5 (in Latin referring to a mast) to that apical portion of the rachis beyond the primary velum (and, when present, the secondary, anterior velum: Schonitzer and Renner, 1980; Schonitzer, 1986). The anterior velum is often lacking, but variably repre¬ sented among different lineages by a carina, thin lamella, or a reduced, lamellate lobe (Schonitzer, 1986). Microphotographs were prepared through the combination of consecutive images at suc¬ cessive focal plans, all shot using a Canon 7D digital camera attached to an Infinity K-2 long¬ distance microscope lens. Measurements of specimens relied upon an ocular micrometer and an Olympus SZX-12 stereomicroscope, those of the paratype male in parentheses. 5 The term malus as used by Michener (1944, 2007), Eickwort (1969), and Engel (2000, 2001) is equivalent to rachis, but the latter term is more broadly applicable to the axial length of all tibial spurs. 2017 ENGEL ET AL.: ARABIAN TARSALIA 5 SYSTEMATICS Subfamily Apinae Latreille, 1802 Tribe Ancylaini Michener, 1944 Ancylini Michener, 1944: 273. Type genus: Ancyla Lepeletier de Saint Fargeau, 1841. Ancylaini Michener, nomen emendatum; Engel et al., 2008: 199; ICZN, 2011: 116. Diagnosis: Bees of small to moderate size (5-10 mm in length); head about as wide as long to wider than long and face comparatively narrow, interocular distance less than com¬ pound eye length; vertex comparatively straight, not uniformly convex nor excavated between compound eyes and ocelli in facial view. Lower face often without maculation (see sexes below); clypeus flat to weakly protuberant in profile, scarcely bent back at side of labrum. Labrum broader than long. Malar space linear, nearly lacking with mandibular base abutting lower compound eye margin. Mandible simple, without distinct subapical tooth; articulations equi¬ distant from compound eye, posterior articulation behind mediolongitudinal axis of compound eye. Postpalpal portion of galea broad proximally, then attenuate and with weakly sclerotized strip extending to apex; maxillary palpus with six palpomeres. Glossa and labial palpus much shorter than prementum; submental spine present; paraglossa short; labial palpus with four palpomeres, first two palpomeres similar in shape to apical two palpomeres, not flattened; submental spine absent. Pronotum without transverse carina. Propodeum basally declivitous, basal area impunctate, not covered by setae. Protibial calcar of strigilis with anterior carina or thin lamella on rachis bordering primary velum but lacking anterior, secondary velum; primary velum rectangular with inner margin rounded, with inner apical corner projected as acutely pointed process; malus short, often shorter than velum, with apex straight, inner margin minutely ciliate. Mesotibial spur long, serrate, apex slightly incurved; mesobasitarsal comb absent. Metabasitibial plate short, wider than long, apically weakly and broadly rounded; metatibial spurs long, serrate, apices straight or slightly incurved. Pretarsal arolium present. Forewing with pterostigma small, longer than wide, scarcely wider than prestigma; pterostig- mal margin inside marginal cell convex, sometimes weakly so; marginal cell apex not truncate, bent away from anterior wing margin; three submarginal cells present; membrane with setae throughout. Hind wing with 2M+Cu about one-half as long as M, or slightly more; cu-a trans¬ verse, one-half length or more of M; jugal lobe short, slightly less than one-half length of vannal lobe. Metasomal tergum I without carina at angle of anterior- and dorsal-facing surfaces, at most sharply angulate in some males. Female: Face without maculation; probasitarsus without anterior or posterior combs; metati¬ bial and metabasitarsal scopa large, dense, composed of long, plumose setae, without oil-collect¬ ing structures; metabasitibial plate surface bare and smooth; pretarsal claws with small basal tooth; metasomal tergum VI with narrow pygidial plate; gradulus of metasomal sternum II sim¬ ple; sterna II-V with scattered long, subdecumbent setae apically, but not forming scopa. Male: Clypeus, supraclypeal area, paraocular area, and antennal scape with maculation; apical margin of clypeus with linear patch of distinctive setae; ventral surface of mandible and 6 AMERICAN MUSEUM NOVITATES NO. 3877 usually postgena near mandibular base with characteristic patch of dense setae; flagellum cren- ulate; mesobasitarsus, metatibia, metabasitarsus, and metatibial spurs frequently greatly modi¬ fied (e.g., mesobasitarsus swollen with dense setae; metatibia swollen, with modified setae; metabasitarsus thickened and arched, with concave inner surface and modified; metatibial spurs elongate and broadly curved); metabasitibial plate covered with appressed setae; pretarsal claws deeply cleft; tergum VII setose, with pygidial plate present; sternum V with subapical, paramedial setal patches and apical, paramedial, thumblike processes bordering medial concav¬ ity; sternum VI with medial tubercle extending between processes of sternum V, and variously with broadly concave areas laterally and medially, with mediolongitudinal ridge or carina and medioapical margin various modified; sternum VII with lateral and apical lobes; sternum VIII with apical lobes; gonostylus without setigerous parapenial lobe; spatha present. Comments: The presently included species, all in the genus Ancyla (= Plistotrichia Morawitz, 1874), and their distributions are summarized in table 1. The nesting biology and immature stages have been reported for two species of Ancyla (Straka and Rozen, 2012), and the species are generally considered specialists of Apiaceae and has mouthparts similar to short-tongued bees (Silveira, 1993b; Straka and Rozen, 2012). Tribe Tarsaliini Engel, 2015 Tarsaliini Engel, 2015: 4. Type genus: Tarsalia Morawitz, 1895. Diagnosis (modified and expanded from Engel, 2015): Bees of small to moderate size (5.5-13 mm in length); head wider than long, but face comparatively narrow, interocular distance less than compound eye length; vertex comparatively straight, not uniformly convex nor excavated between compound eyes and ocelli in facial view (fig. ID). Lower face often with maculation (see sexes below); clypeus weakly to moderately protuberant in profile, strongly bent back at side of labrum. Labrum broader than long. Malar space linear, nearly lacking with mandibular base abutting lower compound eye margin. Mandible simple, with¬ out distinct subapical tooth; articulations equidistant from compound eye, posterior articula¬ tion behind mediolongitudinal axis of compound eye. Postpalpal portion of galea broad proximally, then attenuate and with weakly sclerotized strip extending to apex; maxillary palpus with six palpomeres. Glossa and labial palpus longer than prementum; submental spine present; paraglossa short, not exceeding first labial palpomere; labial palpus with four palpomeres, first two palpomeres long, sheathlike (contrasting with condition in Ancylaini where first two palpomeres are similar in shape to apical two palpomeres); submental spine present. Pronotum without transverse carina. Propodeum basally with subhorizontal area, subhorizontal area punctate and setose. Protibial calcar of strigilis with anterior carina or thin lamella on rachis bordering primary velum but lacking anterior, secondary velum; pri¬ mary velum rectangular with inner margin blunt, straight, with inner apical corner projected as acutely pointed process (as in most Eucerini, Ancylaini, Exomalopsini, and Emphorini: Schonitzer, 1986); malus elongate, as long as or longer than velum, with apex straight or slightly incurved, inner margin minutely ciliate. Mesotibial spur long, serrate, apex slightly 2017 ENGEL ET AL.: ARABIAN TARSALIA 7 incurved; mesobasitarsal comb absent. Metabasitibial plate short, wider than long, apically weakly and broadly rounded; metatibial spurs long, serrate, apices slightly incurved. Pretarsal arolium present. Forewing with pterostigma small (fig. 1A), as long as wide, scarcely wider than prestigma; pterostigmal margin inside marginal cell weakly or not convex; marginal cell apex not truncate, bent away from anterior wing margin; three submarginal cells present; membrane with setae throughout. Hind wing with 2M+Cu one-half as long as M or less; cu-a transverse, more than one-half length of M; jugal lobe short, slightly less than one-half length of vannal lobe. Metasomal tergum I with or without carina at angle of anterior- and dorsal¬ facing surfaces (absent in two species). Female: Labrum, clypeus, and supraclypeal area usually with maculation (fig. ID) (absent in one species); paraocular area usually without maculation (present in one species); antennal scape usually without maculation (present in one species); probasitarsus without anterior or posterior combs; metatibial and metabasitarsal scopa large, dense, composed of long, plumose setae, without oil-collecting structures; metabasitibial plate surface bare and smooth; pretarsal claws with minute basal tooth; metasomal tergum VI with narrow pygidial plate; gradulus of metasomal sternum II weakly birecurved (similar to condition in some Thygater Holmberg); sterna II-V with scopa composed of long, dense setae, such setae often apically wavy or sinuous (similar to type IV of Pasteels and Pasteels, 1974). Male: Labrum with maculation (reduced to a medioapical spot in one species); clypeus and supraclypeal area usually with maculation (absent in one species); paraocular area with macula¬ tion; antennal scape usually without maculation (absent in one species), flagellum not crenulate nor elongate; metabasitibial plate often shorter than in female, covered with appressed setae; pretarsal claws deeply cleft; tergum VII setose, with true pygidial plate absent (no raised pygidial plate, instead surface continuous with disc and uniformly covered with setae), instead with dis¬ tinct lateral carina and apicomedially produced and truncate to weakly bilobate (broadly and shallowly emarginate), medial projected area rounded and deflexed in two species; apical margin sternum V straight to medially lob ate; sternum VI narrowed apically, apical margin usually char¬ acteristic (truncate, rounded, or medially emarginate), with lateral marginal areas broadly con¬ cave and typically smooth, producing narrow medial area usually bearing a medial tubercle or carina; sternum VII with lateral and apical lobes; sternum VIII with apical lobes; gonostylus with parapenial lobe bearing thickened setae; spatha present (absent in one species); hidden sterna and genitalia asymmetrical (sternum VII usually strongly asymmetrical; sternum VIII less dra¬ matically so; sometimes with penis valves strongly asymmetrical). Comments: Recently, the tribe was proposed as “new” again in Plant and Paulus (2016), despite having already been made available a year prior to this. The name as employed by these authors has no nomenclatural standing. Baker (1998) speculated that the characteristic sternal setae functioned as a scopa. However, conclusive evidence demonstrating the use of the sternal setae for pollen trans¬ port is lacking, and both Michener (2007) and Plant and Paulus (2016) indicated that specimens with copious pollen in the hind-leg scopae lacked pollen on the sterna. How¬ ever, we’ve followed Baker (1998) in considering the sternal setae as a metasomal scopa as AMERICAN MUSEUM NOVITATES NO. 3877 the female of the new species described below has appreciable pollen in only two places on the body: the hind-leg scopae and the sternal setae. It is hoped that biological studies will someday be possible to permit an evaluation of the true functional significance of the dense sternal setae of Tarsalia. The chiral form of the male terminalia is unique among bees. The asymmetry of the male sternum VII, and to a lesser degree sternum VIII, along with the sometimes hypertrophy of one penis valve, is immediately distinctive and peculiar. The purpose for this consistent chiral¬ ity is unclear and, as noted by Baker (1998), does not correspond to any asymmetry in the female terminal sclerites. Until such time as the mating behavior of these bees is discovered, the function of these asymmetries shall remain a mystery, but in the meantime they represent a strong synapomorphy for the tribe. We have considered the males of Tarsalia to lack a true pygidial plate. It is definitive that a pygidial plate as traditionally defined (Michener, 1944) is lacking in males of Tarsalia. How¬ ever, the seventh tergum is medioapically produced with the tergal apical margin carinate, suggestive of a plate, and leading some to speculate that the majority of the tergal apical dorsum is equivalent to a pygidial plate (Michener, 2007). However, this apical surface is continuous on all sides with the remainder of the tergal disc, as well as identically sculptured and uniformly covered with setae, in stark contrast to the distinctly raised surface of a pygidial plate which also almost invariably differently sculptured and either lacking in setae or with such setae distinctly differing from those of the surrounding disc. Thus, from a strictly morphological standpoint we do not consider a true pygidial plate to be present in Tarsalia, and the form of the seventh tergum in these bees should not be confused with those lineages in which such a plate is present (e.g., Ancylaini). Even if the apical protrusion of the seventh tergum of male Tarsalia should be shown later to be developmental^ homologous to the pygidial plate, it still remains a fundamentally different structure of uncertain function and such a character state should not be equated or confused with the pygidial plate in other bees. Referring to the pres¬ ent character state as a “pygidial plate” obscures the uniqueness of the condition found among Tarsalia, as well as misleads others, we believe, as to its morphological identity (i.e., that a true pygidial plate is lacking). Genus Tarsalia Morawitz, 1895 Tarsalia Morawitz, 1895: 9. Type species: Tarsalia hirtipes Morawitz, 1895, by monotypy. Friese, 1896: 211; Baker, 1998: 837; Michener, 2000: 666; Michener, 2007: 686. Ancyla (Tarsalia) Morawitz; Warncke, 1977: 58; Warncke, 1979: 192. Diagnosis: As for tribe (above). Comments: The genus currently includes two rather distinct groups, one with three species in northeastern Africa, Arabia, and Iran, while the other has five species in Western and Central Asia, south into India, and west on the Mediterranean islands of Cyprus and Sardinia (table 1). 2017 ENGEL ET AL.: ARABIAN TARSALIA 9 FIGURE 1. Representative species of Tarsalia Morawitz, subgenus Tarsalia s. str. A. Tarsalia (Tarsalia) ancy- liformis Popov, lateral habitus of female. B. T. (T.) hirtipes Morawitz, lateral habitus of female. C. T. (T.) dec- cana Baker, lateral habitus of male holotype. D. Facial view of female T. ancyliformis. E. Dorsal habitus of female T. ancyliformis. F. Dorsal habitus of male T. deccana. G. Dorsal habitus of female T. hirtipes. 10 AMERICAN MUSEUM NOVITATES NO. 3877 FIGURE 2. Posterior view of metatibial and metabasitarsal scopae of species of Tarsalia Morawitz. A. Open scopal form of Tarsalia (Tarsalia) ancyliformis Popov. B. Dense, closed scopal form of T. (Astibomelissa) per- sica (Warncke). Key to Subgenera of Tarsalia l.Metasoma and mesosoma dark reddish brown to black (fig. 1); metatibial scopa open (fig. 2A); subtriangular patch of distinctive, squamiform setae on outer surface of metatibia apical to metabasitibial plate (Sardinia, Cyprus, Western and Central Asia, India). .Tarsalia s. str. —Metasoma and majority of mesosoma light castaneous to testaceous (figs. 3, 4); metatibial scopa densely closed (fig. 2B); outer surface of metatibia apical to metabasitibial plate without patch of distinctive setae, area similar to remainder of scopal setae (northeastern Africa, Arabia, Iran).Astibomelissa, n. subgen. Astibomelissa Engel, new subgenus Type species: Tarsalia kindahensis Engel, new species. Diagnosis: Body length approximately 5.5-9.0 mm; integument pale castaneous to testa¬ ceous on metasoma and majority of mesosoma (figs. 3-5, 6). Female metatibial scopa densely

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