Sphecodes

Sphecodes is a cosmopolitan genus with species represented on every continent except Antarctica.^[2] The genus is also very species rich, with 21 species described from Siberia,^[3] 33 species from Central Europe,^[2] 17 species from the Indian region,^[4] 26 from the Arabian Peninsula and surrounding region,^[5] and 21 from Southeast Asia.^[6] In the Americas, there are 11 species described from the Neotropics^[7] and 63 from the Nearctic.^[8] Australia has only two native species, S. manskii and S. profugus, both of which are restricted to the northeast.^[9] The species Sphecodes albilabris is thought to have been introduced to both Australia and the United States by accident.^[2]

Summarize Timeline Fact Check

The genus was erected in 1804 by Pierre Latreille for the species Sphecodes gibbus. That species was initially described by Carl Linnaeus as Sphex gibba.^[10][11] The fact that the type specimen was initially described as a wasp was not lost on Latreille as the name he chose for the genus, Sphecodes, means "like a wasp".^[12] In the late 18th and early 19th-centuries, the taxonomy of Apoidea was very unclear, and at various points Sphecodes species were described as the following genera: Sphex Linnaeus, 1758; Nomada Latrielle, 1802; Halictus Latrielle, 1804; Apis Linnaeus, 1758; Melitta Kirby, 1802; Andrena Fabricius, 1775; and Tiphia Fabricius, 1775.^[10][11]

The following genera are now considered synonyms of Sphecodes: Dichroa Illiger, 1806; Sabulicola Verhoeff 1890; Thrausmus Buysson 1900; Dialonia Robertson, 1903; Machaeris Robertson, 1903; Stelidium Robertson, 1903; and Sphegodes Mavromoustakis, 1949.^[13]

Charles Robertson was particularly prolific in describing genera that would be synonomized with Sphecodes later on, although the differences he noticed are now reflected in the subgenera. The following genera are now considered subgenera: Drepanium Robertson, 1903; Proteraner Robertson, 1903; and Sphecodium Robertson, 1903.^[13][14] Subgeneric classifications are currently debated, and whether they are truly monophyletic groups remains unknown.

Callosphecodes Friese, 1909 may either be a synonym or subgenus, but specimens that match its description are exceedingly rare.^[15]

Relatively few phylogenetic studies have been completed for the genus, and none that are exhaustive or analyze biogeography. Sphecodes is a member of the family Halictidae and in the tribe Sphecodini. Sphecodini is sister to Halictini sensu stricto.^[16] Genus-level relationships within Sphecodini are still being determined,^[17] but recent phylogenies suggest that Sphecodes is sister to the genus Eupetersia.^[18] It is unknown exactly how many species there are worldwide, and even well-studied regions of the world such as the United States and Europe likely have many species to be described or synonymized.

A note on Halictinae tribes

There are two diverging approaches to tribe-level phylogenies within Halictinae. The first, used by taxonomists like Danforth,^[19] Gibbs,^[16] and Brady^[16] recognizes five distinct tribes: Augochlorini, Thrinchostomini, Caenohalictini, Halictini sensu stricto, and Sphecodini. This is also the approach that Wikipedia uses. The second, used by Ascher,^[8] Michener,^[20] Straka,^[18] and Engel^[14] among others recognizes only two: Augochlorini and Halictini sensu lato, with the rest of the tribes relegated to subtribes within Halictini s. l. This approach is also used by resources such as iNaturalist and BugGuide. Both are valid and represent the same evolutionary relationships.

Sphecodes is in the family Halictidae, a group distinguished from other bee families by a highly arcuate basal vein. In North America, Sphecodes is the only genus of Sphecodini present.^[8]

Sphecodes are small-to-medium bees, usually with distinctive red abdomens although males are sometimes entirely black.^[21] The genus has somewhat oval, brachycephalic heads and thicker antennae compared to other bees. Most species have three submarginal cells, but some consistently have two, and others have either two or three.^[21] Their body size is correlated with that of their host: large Sphecodes parasitize larger hosts like Andrena, while small Sphecodes are more often associated with small hosts like Perdita. Like other kleptoparasitic bees, they are largely hairless and females lack scopae on their hind tibia.^[22] Both sexes have thickened and highly rugose cuticles. This adaptation allows them to deflect and block stings when fighting their hosts. In many other cuckoo bee genera, the larvae sport large mandibles used to kill the host. These are absent in the young of Sphecodes, where it is the adult that dispatches the host egg or larva.^[19]

Identification of these bees is difficult, in part due to lack of research and published keys. Important characters include the antennae, mandibles, propodeum, and male genitalia.^[11]