Halictidae

Mellitidia tomentifera, copyright mmpro.

Belongs within: Anthophila.
Contains: Halictinae, Nomia, Lipotriches.

The Halictidae are a family of short-tongued bees distinguished by a strongly arcuate basal vein in the fore wing (Engel 2001).

The Halictidae: short tongues and waxy chambers
Published 25 April 2020

In an earlier post, I introduced you to the diverse group of bees known as the Halictinae. In this post, I’m going to take a step back and consider the family of bees to which the halictines belong, the Halictidae.

Nomia sp. feeding at a flower, copyright Graham Wise.

The Halictidae are one of the families of what are known as ‘short-tongued bees’ (the other short-tongued families recognised by Michener, 2007, are the Andrenidae, Colletidae and Stenotritidae). Bees have their mouthparts modified compared to those of other wasps to form a mobile proboscis. The tongue works in three main sections from base to tip. The first two sections work like the upper and lower parts of your arm, or of the arm of a crane, to extend and fold back the proboscis against the underside of the head. The third section beyond these two includes a flexible structure, the glossa, that may be thought of as working like the tongue proper to collect nectar and pollen from the inside of flowers. Somewhat self-explanatorily, this glossa is extremely long and slender in the families of ‘long-tongued bees’ (the Apidae and Megachilidae) but relatively shorter and broader in short-tongued bees. Naturally, these differences in tongue structure may be reflected in differences in which types of flowers the different types of bees chose to visit. Just to confuse matters, some species of Halictidae may have relatively long proboscides overall, but in this case the extra length is achieved by extending the length of the middle ‘arm’ section rather than of the glossa itself. The primary features separating Halictidae from the other families of short-tongued bees relate to the structure of particular sclerites incorporated into the proboscis that I’m not going to go into here, but notable points include that the glossa of Halictidae is pointed at the tip and hairs on it are usually branched or bifid at the tips.

Male Halictus tetrazonianellus with proboscis extended (the glossa is the orange structure at the end of the proboscis), copyright Gideon Pisanty.

For the most part, halictids are moderately built bees: neither remarkably slender nor particularly robust. Halictids vary extensively in size: many are small, even minute, but some may be relatively large by bee standards. Coloration is similarly variable, with both metallic and non-metallic species belonging to the family. Members of the genus Nomia (which tend to be relatively large for halictids) often bear contrasting bright bands across the back of the metasoma. Michener (2007) recognised four subfamilies within the halictids: the Rophitinae, Nomiinae, Nomioidinae and Halictinae, with the Halictinae being considerably more diverse species-wise than the other three. Nomioidines have sometimes been included by other authors within the Halictinae but, as there is a general agreement that nomioidines form the sister lineage of the halictines in the strict sense, the question of whether to combine them or not is purely a matter of semantics. Rophitines differ from other halictids in having a relatively large labrum whose tip remains visible between the mandibles when they are closed (other subfamilies have the labrum hidden by the closed mandibles). Rophitines, as well as kleptoparasitic halictines, also have the tip of the labrum simply truncate or rounded; in other subfamilies, the tip of the labrum in females is produced into a distinct process. Rophitines also have the scopa (the array of long pollen-carrying hairs on the hind leg) less developed on the trochanter and femur than on the tibia whereas other subfamilies (excluding, again, kleptoparasitic forms in which the scopa is reduced) generally have the longest scopal hairs on the femur. Nomiines commonly have the third submarginal cell on the wing (if present) as long as the first submarginal cell or at least more than twice the length of the second. In nomioidines and halictines, the third submarginal cell is much shorter. Another notable feature of the last two subfamilies is that the basal vein (the upper of the three veins radiating from the basal midline of the wing) is much more strongly curved near the base than in other bee families; this feature may or may not be discernable in rophitines and nomiines.

Just to show that bees can sometimes get insane: a male of the Colombian species Chlerogella anchicaya, from Engel et al. (2014).

For the most part, halictids construct their nests in burrows in the ground (some halictines nest in rotting wood). Cells of the burrows are generally lined with a wax-like membrane secreted by the parent bee. The membrane is duller and less watertight in Rophitinae than in other subfamilies; one rophitine genus, the southwest North American Protodufourea, appears to not produce such a membrane. Most non-halictine halictids are solitary nesters though some nomiines are known to work communally, and may even show low levels of division of labour. Kleptoparasitism is not known outside the Halictinae.

Systematics of Halictidae

Characters (from Engel 2001): Subantennal suture meeting inner, lower margin of antennal socket, infrequently lower margin; one or (more rarely) two subantennal sutures. Facial foveae typically absent. Galea elongate prepalpally. Mentum and submentum absent; segments one and two of labial palpi similar to distal segments, cylindrical, rarely elongate; glossa acute, typically short, without flabellum. Preepisternal groove often complete (less commonly groove weak or absent below scrobe). Mesocoxa with exposed area shorter than distance from summit to metathoracic wing base; metabasibitial plate frequently present (except in parasitic forms); scopa developed on hind leg in non-parasitic females. Jugal lobe of hind wing much more than half as long as vannal lobe. Pygidial plate present in females. Volsellae present.

<==Halictidae
    |  i. s.: AcunomiaO01
    |         SystrophaPK17
    |           |--S. curvicornisPK17
    |           |--S. planidesR35
    |           `--S. spiralis [=Panurgus (Systropha) spiralis]G20
    |--+--DufoureaBD17 [DufoureinaeM65]
    |  |    |--D. dentiventrisPK17
    |  |    `--D. novaeangliaePK17
    |  `--RophitinaeE01
    |       |  i. s.: XeralictusE01
    |       |--RophitesBD17 [RophitiniE01]
    |       |    |--R. algirusBD17
    |       |    `--R. quinquespinosus [=Megachile (Rophites) quinquespinosa]G20
    |       `--PenapiniE01
    |            |--PenapisE01
    |            |--CeblurgusE01
    |            `--GoeletapisE01
    `--+--+--HalictinaeE01
       |  `--Nomioidinae [Nomioidini]H18
       |       |--Ceylalictus Strand 1913H18, M65
       |       |    |--*C. horni (Strand 1913) [=Halictus horni, Nomioides horni]M65
       |       |    `--C. perditellus (Cockerell 1905)H18, M65 [=Nomioides perditellaM65]
       |       `--Nomioides Schrenck 1866M65 (see below for synonymy)
       |            |--N. minutissima (Rossi 1790) (see below for synonymy)M65
       |            |--N. arnoldi Friese 1913 [=*Cellaria arnoldi, *Cellariella arnoldi]M65
       |            |--N. facilisR35
       |            |--N. obliqua (Friese 1924) [=Halictus obliquus]M65
       |            `--N. variegata (Olivier 1789) [=Andrena variegata, *Eunomioides variegata]M65
       `--NomiinaeE01
            |--NomiaM65
            |--LipotrichesMC13
            |--Epinomia trianguliferaR35
            |--DieunomiaE00
            |    |--D. heteropodaE00
            |    |--D. nevadensisBD17
            |    `--D. trianguliferaE00
            |--Reepenia Friese 1909H18, M65 [incl. Megaloptodes Moure 1958M65]
            |    |--*R. variabilis (Friese 1909) [=Nomia (*Reepenia) variabilis]M65
            |    |--R. bidentata (Smith 1863) [=Nomia (Reepenia) bidentata]M65
            |    |--R. bituberculata (Smith 1853)H18, M65 (see below for synonymy)
            |    |--R. brevicornisR35
            |    |--R. florea (Smith 1863) [=Nomia (Reepenia) florea]M65
            |    |--R. fuliginosa (Friese 1909) [=Nomia (Reepenia) fuliginosa]M65
            |    |--R. fulviventris (Cameron 1901) [=Nomia (Reepenia) fulviventris]M65
            |    |--R. ‘fulviventris’ (Friese 1909) [=Nomia (Reepenia) fulviventris non Cameron 1901]M65
            |    |--R. ‘triangulifera’ (Friese 1909) [=Nomia (Reepenia) triangulifera non Vachal 1897]M65
            |    `--R. zebrae (Friese 1909) [=Nomia (Reepenia) zebrae]M65
            `--Mellitidia Guérin-Méneville 1830H18, M65
                 |--*M. australis (Guérin-Méneville 1830) [=Andrena australis, Nomia (*Mellitidia) australis]M65
                 |--M. cincta (Smith 1859) [=Nomia (Mellitidia) cincta]M65
                 |--M. dentata (Smith 1859) [=Nomia (Mellitidia) dentata]M65
                 |--M. gressitti (Michener 1965) [=Nomia (Mellitidia) gressitti]M65
                 |--M. horvarthi (Friese 1909) [=Nomia (Mellitidia) horvarthi]M65
                 |--M. kochi (Friese 1909) [=Nomia (Mellitidia) kochi]M65
                 |--M. longicornis (Smith 1859) [=Nomia (Mellitidia) longicornis]M65
                 |--M. luteiventris (Friese 1909) [=Nomia (Mellitidia) luteiventris]M65
                 |--M. metallica (Smith 1863) [=Nomia (Mellitidia) metallica]M65
                 |--M. simillima (Smith 1863) [=Nomia (Mellitidia) simillima]M65
                 |--M. simplicinotum (Michener 1965) [=Nomia (Mellitidia) simplicinotum]M65
                 `--M. tomentifera (Friese 1909) [=Nomia (Mellitidia) tomentifera]M65

Nomioides Schrenck 1866 [incl. Cellaria Friese 1913 non Ellis & Solander 1786, Cellariella Strand 1926, Eunomioides Blüthgen 1937]M65

Nomioides minutissima (Rossi 1790) [=Apis minutissima; incl. Andrena pulchella Jurine 1807, *Nomioides pulchella]M65

Reepenia bituberculata (Smith 1853)H18, M65 [=Megalopta bituberculataM65, *Megaloptodes bituberculataM65, Nomia (Reepenia) bituberculataM65; incl. N. (R.) eboracina Cockerell 1912M65, Tetralonia testacea Smith 1854M65]

*Type species of generic name indicated

References

[BD17] Branstetter, M. G., B. N. Danforth, J. P. Pitts, B. C. Faircloth, P. S. Ward, M. L. Buffington, M. W. Gates, R. R. Kula & S. G. Brady. 2017. Phylogenomic insights into the evolution of stinging wasps and the origins of ants and bees. Current Biology 27: 1019–1025.

[E00] Engel, M. S. 2000. Classification of the bee tribe Augochlorini (Hymenoptera: Halictidae). Bulletin of the American Museum of Natural History 250: 1–89.

[E01] Engel, M. S. 2001. A monograph of the Baltic amber bees and evolution of the Apoidea (Hymenoptera). Bulletin of the American Museum of Natural History 259: 1–192.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

[H18] Houston, T. 2018. A Guide to Native Bees of Australia. CSIRO Publishing.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[M65] Michener, C. D. 1965. A classification of the bees of the Australian and South Pacific regions. Bulletin of the American Museum of Natural History 130: 1-362.

Michener, C. D. 2007. The Bees of the World 2nd ed. John Hopkins University Press: Baltimore.

[O01] OConnor, B. M. 2001. Historical ecology of the Acaridae (Acari): phylogenetic evidence for host and habitat shifts. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 76–82. CSIRO Publishing: Melbourne.

[PK17] Peters, R. S., L. Krogmann, C. Mayer, A. Donath, S. Gunkel, K. Meusemann, A. Kozlov, L. Podsiadlowski, M. Petersen, R. Lanfear, P. A. Diez, J. Heraty, K. M. Kjer, S. Klopfstein, R. Meier, C. Polidori, T. Schmitt, S. Liu, X. Zhou, T. Wappler, J. Rust, B. Misof & O. Niehuis. 2017. Evolutionary history of the Hymenoptera. Current Biology 27 (7): 1013–1018.

[R35] Rayment, T. 1935. A Cluster of Bees: Sixty essays on the life-histories of Australian bees, with specific descriptions of over 100 new species. Endeavour Press: Sydney.

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