LJUBLJANA, JUNIJ 2012
Vol. 20, št. 1: 65-72
ON THE CLASSIFICATION OF THE BEE GENUS MANUELIA (HYMENOPTERA: APIDAE)
Michael S. Engel
Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive - Suite 140, University of Kansas, Lawrence, Kansas 66045, USA
Abstract - The classification of the monogeneric and relic xylocopine tribe Manueli-ini is briefly addressed. Two subgenera are recognized in Manuelia Vachal - Manuelia s.str., consisting of Manuelia gayi (Spinola), and Uliamane subgen. n., comprising M. postica (Spinola) as type species and M. gayatina (Spinola). Diagnoses for the subgenera are provided along with a key to these groups and detailed images for M. pos-tica.
Key words: Apoidea, Anthophila, Xylocopinae, Manuelia, taxonomy, Chile, Argentina.
Izvleček - O RAZVRŠČANJU ČEBELJEGA RODU MANUELIA (HYMENOPTERA: APIDAE)
Razvrščanje enorodovnega in reliktnega plemena Manueliini poddružine Xylocopinae je predstavljeno na kratko. Prepoznana sta dva podrodova rodu Manuelia Vachal - Manuelia s.str., ki ga tvori vrsta Manuelia gayi (Spinola), in Uliamane subgen. n., ki vsebuje vrsti M. postica (Spinola) kot tipsko vrsto in M. gayatina (Spinola). Pripravljeni so opisi podrodov skupaj s ključem za ti skupini in podrobnimi slikami vrste M. postica.
Ključne besede: Apoidea, Anthophila, Xylocopinae, Manuelia, taksonomija, Čile, Argentina.
Considerable interest has developed in the relationships among the various lineages of Xylocopinae, encompassing the large carpenter bees (Xylocopini), the small car-
Introduction
penter bees (Ceratinini), the Old World allodapine bees (Allodapini), and the enigmatic, relic tribe Manueliini. A fifth tribe, the Boreallodapini, is known from mid-Eocene amber in Europe but was perhaps widespread (Engel, 2001). The boreallodapines were the closest relatives of the Allodapini (Engel, 2001). Phylogenetic considerations of the group have been presented by Daly et al. (1987), Sakagami and Michener (1987), Roig-Alsina and Michener (1993), Engel (2001), and Flores-Prado et al. (2010). In addition, Rozen (2010) recently documented numerous features from the mature larvae of the tribes of Xylocopinae, including remarks on one species of Manueliini [i.e., Manuelia gayatina (Spinola)].
As noted, the Manueliini are perhaps the basalmost lineage of the Xylocopinae and are relic in southern South America, predominantly Chile but also in Argentina. The tribe consists of three species classified in a single genus, Manuelia Vachal, and superficially resemble small carpenter bees of the genus Ceratina Latreille. Recently the biology of one species, M. postica (Spinola), has been studied in great detail (Flores-Prado et al., 2008a, 2008b; Flores-Prado & Niemeyer, 2010), while Claude-Joseph (1926) provided a general account of the nesting biology of the other two species. While there are few species of Manuelia they are as different from each other as is usually classified in separate genera or subgenera elsewhere in the Apoidea and most certainly within the Xylocopinae. Herein a subgeneric classification of Manuelia is proposed to highlight these significant morphological differences. Morphological terminology follows that of Engel (2001) and Michener (2007). In addition, given that Manuelia has rarely been figured, I provide detailed images of pertinent structures for M. postica (Figs. 1-13).
Systematics
Tribe Manueliini Sakagami & Michener Genus Manuelia Vachal
Diagnosis: Body small and slender (similar to Ceratina), 4.7-8.5 mm in length (Figs. 1-4); mandible tapering to apex, not abruptly narrowed; female labrum with elevated smooth triangular area; clypeus above anterior tentorial pits narrower than below (Figs. 5, 6), anterior tentorial pits situated above midheight of clypeus; apical meta-somal terga strongly convex (not depressed as in Allodapini); first flagellomere shorter than combined lengths of second and third flagellomeres; wings distally pubescent (Figs. 1-4), not papillate; forewing with pterostigma large, with three submarginal cells (Fig. 1, 2); hind wing jugal lobe ca. 0.2x length of vannal lobe; female pygidial fimbria present; female pygidial plate present as strong spine (Fig. 3); male metasomal sternum VIII large, sclerotized (Fig. 10), strongly convex ventrally, with robust apical process (apical process absent in Ceratinini) (Fig. 10), lateral arms short, robust (Fig. 10); male gonostylus not fused with gonocoxa (fused in Ceratinini except Euceratina Hirashima) (Figs. 12, 13).
Comments: The genus and its species were reviewed in detail by Daly et al. (1987).
Key to Subgenera of Manuelia
1. Bees dark metallic blue; lateral margin of female labral disc straight; male mandible bidentate; male with posterior ocelli in front of vertex summit; male basimetatibial plate present..........................................................Manuelia s.str.
— Bees black, non-metallic, sometimes with apical metasomal segments reddish (Figs. 1-4); lateral margin of female labral disc concave; male mandible simple; male with posterior ocelli on vertex summit (Fig. 6); male basimetatibial plate absent (Fig. 8)..............................................................Uliamane, subg. n.
Figs. 5-8: Photomicrographs of Manuelia postica (Spinola) from Chile; 5) Female face; 6) Male face; 7) Female basimetatibial plate; 8) Male outer base of metatibia (showing absence of plate).
Figs. 9-13: Photomicrographs of male terminalia of Manuelia postica (Spinola) from Chile; 9) Metasomal sternum VII; 10) Sternum VIII; 11) Genital capsule, lateral view; 12) Genital capsule, ventral view; 13) Genital capsule, dorsal view.
Subgenus Manuelia Vachal
Manuelia Vachal, 1905: 25. Type species: Halictus gayi Spinola, 1851, by designation of Daly et al. (1987: 104).
Diagnosis: Bees dark metallic blue; lateral margin of female labral disc straight; male mandible bidentate; male with posterior ocelli in front of vertex summit; male basimetatibial plate present; male penis valves robust; gonostylus short, shorter than penis valve apodemes.
Included species: Manuelia gayi (Spinola, 1851).
Uliamane subgen. n.
Type species: Halictus posticus Spinola, 1851.
Diagnosis: Bees black, non-metallic, sometimes with apical metasomal segments reddish (Figs. 1-4); lateral margin of female labral disc concave; male mandible simple; male with posterior ocelli on vertex summit (Fig. 6); male basimetatibial plate absent (Fig. 8); male penis valves relatively slender; gonostylus elongate, as long or longer than penis valve apodemes (Figs. 11-13).
Etymology: The new genus-group name is a euphonious combination of letters based on the generic name Manuelia, type genus of the tribe. The name is considered feminine.
Included species: Manuelia postica (Spinola, 1851) and M. gayatina (Spinola, 1851).
Acknowledgements
I am thankful to anonymous reviewers for comments on the manuscript; to Dr. Andrej Gogala for translating my abstract into Slovene and editorial expertise; and to Dr. Ismael A. Hinojosa-D^az for assistance with photomicrography. This is a contribution of the Division of Entomology, University of Kansas Natural History Museum.
References
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Received / Prejeto: 29. 11. 2011