J. HYM. RES. Vol. 18(1), 2009, pp. 74–79

A New Cleptoparasitic Lasioglossum (Hymenoptera, Halictidae) from Africa

JASON GIBBS Department of Biology, York University, 4700 Keele St., Toronto, ON, Canada, M3J1P3, email: [email protected] ______

Abstract.—Lasioglossum (Dialictus) ereptor Gibbs, new species, is described and illustrated. The sole known specimen, a female, is from Mount Kilimanjaro, Tanzania. The species has several characteristics typical of cleptoparasitic and socially parasitic Halictidae. A brief summary of the independent cleptoparasitic lineages of Lasioglossum is given. ______

Cleptoparasitism and social parasitism (2006). Terminology follows that of Mich- (for simplicity called ‘parasitism’ hereafter) ener (2007), Engel (2001) and Harris (1979). have arisen multiple times among halictid The diameter of the median ocellus (OD) is bees (reviewed in Michener 1978; see also used as a relative measure of hair length. Pauly 1984, 1997; Engel et al. 1997; Biani and Puncture density is given in relation to Wcislo 2007). The widespread genus Lasio- interspace size (i) and puncture diameter glossum Curtis sensu lato displays a wide (d). Flagellomeres and metasomal terga range of behavioural systems (reviewed in and sterna are abbreviated F, T and S, Michener 1974, 1990; Packer 1993, Yanega respectively, followed by the appropriate 1997) including multiple parasitic lineages number. (Table 1; Michener 1978, 2007). Parasitic Lasioglossum are known only from eastern RESULTS North America (Paralictus Robertson5Dia- lictus Robertson), central Africa (Paradialic- Lasioglossum (Dialictus) ereptor Gibbs, tus Pauly) and Samoa (Echthralictus Perkins new species and Cheesman) (Michener 2007). Each Figs 1–3. lineage is believed to be a recent derivative Diagnosis.—Female. Slender halictine of its non-parasitic host taxon (Michener with deep blue metallic reflections on the 1978; Pauly 1984). Halictine host species head and mesosoma; metasoma pale may be solitary or have eusocial societies orange; distal wing veins (1rs-m, 2rs-m, (e.g. Packer 1990; Eickwort et al. 1996) 2m-cu) weak; scopa absent, vestiture ex- which might invalidate the distinction tremely sparse and entirely pale; mandi- between cleptoparasitism and social para- bles not reaching opposing mandibular sitism for some parasitic halictines (Biani base, subapical tooth present; labrum with and Wcislo 2007). A new African species of medial tubercle and apical process; macro- parasitic Lasioglossum believed to be a sculpture and punctation largely absent. derivative of the subgenus Dialictus (sensu Differential Diagnosis.—Females of L. Michener 2007) is described. ereptor can be distinguished from other African Dialictus (5Afrodialictus Pauly) by METHODS the absence of scopal hairs and reduced The descriptive format and abbrevia- penicillus, basitibial plate and median tions follow that of Gibbs and Packer longitudinal specialized area of T5. Fe- VOLUME 18, NUMBER 1, 2009 75

males of L. ereptor may be distinguished

3 from the parasitic species L. (Paradialictus) 4 4 Dialictus Dialictus

5 5 synavei Pauly by the metallic colouration and pale abdomen of the former, and the black colouration, enlarged head, elongate mandibles with deep preapical incision L. (D.) zephyrum Afrodialictus Homalictus Afrodialictus Dialictus L. (D.) imitatum ? Dialictus L. (D.) versatum and lunate propodeal flange of the latter. Description.—Female.Bodylength:5.0 mm. Head width: 1.3 mm. Head length: 1.2 mm. Fore wing length: 3.5 mm. Inter- tegular distance: 0.9 mm. OD: 0.01 mm. Colouration: Head and thorax deep metallic blue. Labrum, distal margin of

and MN clypeus, basal half of mandible brown, MA to NC, west to KS remainder of mandible testaceous, apex red; scape and pedicel dark brown; flagel- lomeres brown, faintly ferruginous; post-

. gena faintly metallic. Tegula dark brown, infused with ferruginous; wing venation brown, pterostigma and distal veins testa- Canada, USA ceous; wing dusky throughout; legs brown, Lasioglossum tarsi infused with testaceous. Propodeum faintly metallic with purplish tinge. Meta- soma pale orange. Pubescence: Sparse and pale through- out. Basal area of labrum with apicolateral ctine bee genus bristles; mandibles and distal margin of ial. clypeus with sparse fringe of stiff bristles ’’ (Gibbs, in prep.) ($3OD). Pronotal lobe posterior margin Dalle Torre 1896: 57 Canada, USAGibbs 2008: 74Kohl 1908: ON 306 to NC,Perkins west & to Cheesman MN 1928: 23 Pauly 1984: ? 691 Tanzania Samoa DR Congo (Zaire) Virungas Mt. N. Kilimanjaro P. Samoa ? ? ? Mitchell 1960: 447 ON to AL, west to KS Robertson 1895: 117 ON to GA, west to LA Mitchell 1960: 448 ON to NC Robertson 1901: 230 USA with dense tomentum; remainder of meso-

Paralictus soma with sparse, erect hairs (2OD), longest along posterior margin of mesos-

2 cutellum (3OD); mid femoral and tibial combs present but weak; scopa absent; cephalotes ereptor extraordinarium latro synavei platyparium asteris michiganense simplex penicillus reduced; wing setae evenly distributed, brown; erect hairs on posterior surface of propodeum with slightly longer branches than other hairs. Terga with sparse, erect hairs on ventrally reflexed portions and premarginal line, more nu- merous on T4–T5 dorsal surface (2–3OD), 1928: 14

Robertson 1901: 229 Robertson 1902: 48 Perkins and Cheesman Pauly 1984: 691 fine setae visible on dorsal surface from

1 certain angles; sterna posterior portions with sparse, faintly sinuate, branched may belong to a distinct lineage from other ‘‘ hairs, oriented posteriorly (3–4OD). Dialictus also contains hundreds of non-parasitic species. 5 Surface sculpture: Microsculpture weak 1 Subgenus Author Speciesthroughout. Author Clypeus Country and supraclypeal Distribution Host area faintly imbricate, shining, with few Table 1. Summary of cleptoparasitic or socially parasitic taxa in the hali Dialictus L. (D.) asteris Parasite collected at nestingParasite site collected of inside putative nest host of species eusocial which host is species. eusoc Paralictus Dialictus 1 2 3 4 Echthralictus Paradialictus more or less uniformly spaced punctures 76 JOURNAL OF HYMENOPTERA RESEARCH

Fig. 1. Lateral habitus of Lasioglossum ereptor, n. sp. A) left side B) right side. Scale bar equals 0.5 mm. Inset: Holotype locality label.

(i51–2d), paraclypeal, parasupraclypeal, gitudinal striae on basal portion of dorsal and upper paraocular areas more densely surface. Metasoma smooth, sculpture very punctate (i5d), remainder of face faintly obscure, surface weakly coriarious, terga and closely scabriculous; genal area weak- with sparse (i52–5d), very fine punctures ly strigate. Mesoscutum and mesoscutel- on disc; sterna with piliferous punctures lum faintly imbricate, shining, punctures posteriorly. sparse (i52–4d) and fine, posterior mar- Structure: Head slightly broader than gins of mesoscutum and depressed medial long (1.3 : 1.2); inner orbits nearly parallel area of mesoscutellum with dense puncta- (slightly closer below at level of clypeus). tion (i#d); mesepisternum faintly scabri- Labrum short and broad with basal eleva- culous, tessellate below, hypoepimeral area tion, distal process present, over half imbricate. Propodeum tessellate, faint lon- length of remainder of labrum; mandible VOLUME 18, NUMBER 1, 2009 77

Fig. 3. Dorsal view of mesosoma of Lasioglossum ereptor, n. sp. Scale bar equals 0.5 mm.

cubital (1rs-m and 2rs-m) and 2nd recurrent (2m-cu) veins and 3rd abscissa of vein M weakened; marginal cell with apex pointed on wing margin, pterostigma more than 4 times as long as broad; basitibial plate Fig. 2. Frontal view of head of Lasioglossum ereptor, small, not enclosed by carinae; hind tibial n. sp. Scale bar equals 0.5 mm. spur pectinate with 4 teeth (not including apex of rachis); metapostnotum long and somewhat enlarged, not reaching opposing wide, constituting majority of dorsal pro- mandible base, preapical tooth present. podeal surface, declivous portions of basal Clypeus broader than long (2.5 : 1.0); area small, rounded onto lateral surface; clypeal length greater than distance from lateral carina short, reaching less than upper margin of clypeus to lower margin halfway to dorsal surface. Specialized of antennal socket; subantennal sutures median area of T5 reduced relative to faint; supraclypeal area convex, more non-parasitic species. Pygidial plate not protuberant than rest of face; median visible in holotype. ocellus above upper ocular tangent; dis- Male. Unknown. tance between orbit and lateral ocellus Etymology. The specific epithet means nearly double distance between lateral ‘thief’ in Latin and refers to the species’ ocelli (1.7 : 1.0). Hypostomal carinae presumed parasitic lifestyle. subparallel; genal area slightly wider than Type Material. Holotype label reads eye in lateral view, preoccipital carina (typed): ‘‘TANZANIA: Mt. Kilimanjaro, absent. Antenna rather long, nearly reach- Quadrant 2 Giant Senecio zone, area burnt ing mesoscutellum; scape over 6 times as in 1978, Sheffield U. 6.viii.1993 Meru Galdi’’ long as maximum width; pedicel about 1.5 The type specimen is in fair condition times as long as broad, longer than F1; except for three missing legs. The left front flagellomeres slightly longer than broad, leg is broken near the base of the femur succeeding flagellomeres each slightly lon- and the right mid and hind legs are ger and broader than preceding one. missing beyond the coxae. The pin was Dorsolateral angles of pronotum obtuse; placed through the posterior part of the mesoscutum convex with declivous anteri- mesoscutum and exits the left mesepister- or and lateral margins; mesoscutellum num near the ventral side. The mesoscu- with median longitudinal depression; te- tum and mesepisternum have both split as gula narrow ovoid; 2nd and 3rd transverse a result. 78 JOURNAL OF HYMENOPTERA RESEARCH

Type Depository. The holotype is current- Lasioglossum (Paradialictus) synavei is also ly deposited in the bee collection (PCYU) at believed to be a parasitic derivative of York University, Toronto, Canada. Afrodialictus (Pauly 1984; Arduser and Michener 1987). Several characters com- DISCUSSION mon in cleptoparasites are shared by L. No behavioural data are available for L. ereptor and L. synavei. The unique and ereptor but based on several convergent highly derived mandibles, the lunate characters common in parasitic halictines, flange of the propodeum, large head and such as the lack of scopa, weak basitibial black colouration of L. synavei are not plate and reduced penicillus, it seems shared by L. ereptor and suggest separate evident that L. ereptor is also a cleptoparasite origins of parasitism in these two species. or social parasite. Although many parasitic However, both species are of African halictids have thickened cuticles, coarse origin and their type localities are only sculpture and punctation (e.g. many Sphec. separated by approximately 900 km on a codes Latreille and Temnosoma Smith), which nearly east-west axis. Lasioglossum ereptor likely function in defence against hosts may represent a fourth independent line- (Michener 2007), these characteristics are age of cleptoparasitism in the genus. It is not present in L. ereptor nor in the related also probable that North American species parasitic groups of Dialictus and Paradialic- of parasitic Dialictus are not monophyletic tus. This lack of coarse sculpturing may be (Michener 1978; Gibbs in prep.). Based on an indication of recent origin or a mode of the low-level of differentiation from related parasitism that does not require aggressive nest-building species seen in parasitic encounters. The social parasite L. (D.) asteris Lasioglossum relative to other parasitic (Mitchell) is capable of entering guarded halictines, the origins of parasitic behav- nests without combat (Wcislo 1997) but iour are relatively recent (Michener 1978; nevertheless has an enlarged head and Arduser and Michener 1987). The existence mandibles that could be used for aggressive of multiple and recently derived parasitic encounters. Enlarged mandibles are also lineages make Lasioglossum an ideal group seen in Paradialictus and other parasitic for studying the origin of cleptoparasitism. Dialictus. The mandibles of L. ereptor are Unfortunately, parasitic Lasioglossum are larger than those of related non-parasitic not commonly collected. Additional sam- species but smaller than those of North pling, behavioural studies and systematic American parasitic Dialictus. treatment of these bees are needed. Lasioglossum ereptor is morphologically In the past, new generic names have been similar to the Afrodialictus group of the given to parasitic halictines believed to be large subgenus Dialictus (sensu Michener derived from non-parasitic hosts. For exam- 2007). The genus-group name Afrodialictus ple, the North American parasitic Dialictus refers to tropical African members of were formerly called Paralictus but are Lasioglossum s. l. with weak, often tessellate clearly derived from its host Dialictus (Dan- sculpturing, widely spaced punctures, an forth 1999; Danforth et al. 2003). Likewise, ecarinate and tessellate propodeum and the parasitic Echthralictus and Paradialictus lacking tomentum on the metasomal terga are believed to render their presumed hosts, (Pauly 1999). Afrodialictus has been classi- Homalictus and Afrodialictus, respectively, fied as a synonym of Dialictus (Michener paraphyletic. Both the names Echthralictus 2007; but see Pauly et al. 2008) and that and Paradialictus have been maintained to classification is followed here. Lasioglossum simplify the diagnostic characters of their ereptor shares the characteristic traits of non-parasitic host taxa (Arduser and Mich- Afrodialictus but may be differentiated by ener 1987; Michener 2007). For Paradialictus, its suite of parasitic characters. at least, this seems hardly necessary because VOLUME 18, NUMBER 1, 2009 79 it is likely derived from Dialictus s. l. which Natural History Museum, University of Kansas 5: includes both parasitic and non-parasitic 1–21. species. Lasioglossum ereptor is classified in Gibbs, J. and L. Packer. 2006. Revision and phyloge- netic analysis of Chilicola sensu stricto (Hymenop- the subgenus Dialictus to prevent an addi- tera: Colletidae) with the description of a new tional genus-group name in the already species. Zootaxa 1355: 1–37. complex nomenclature of Lasioglossum s. l. Harris, R. 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