Stingless bees in Miocene amber of southeastern China (Hymenoptera: Apidae)
Authors: Engel Michael S. , Herhold Hollister , Davis Steven , Wang Bo , Thomas Jennifer 
Published: August 2021

Among the many inclusions from the exceptionally rich fossiliferous amber of Zhangpu, China (Middle Miocene: Langhian), stingless bees (Apinae: Meliponini) are particularly common, analogous to the merely slightly older amber sites of Mexico and the Dominican Republic. While there is a large number of workers in Zhangpu amber, only two species are represented. The systematics and morphology of the tribe Meliponini is outlined, including a revision to terms of orientation and direction when discussing surfaces and features of appendages, all in order to better discuss the traits of the fossils as well as place them into a broader context in the global systematics of stingless bees. The two amber species are representative of two Old World genera of Meliponini: Tetragonula Moure and Austroplebeia Moure. While the former is widespread across southern Asia, Malesia, Papuasia, and Australia, the latter is today known only from New Guinea and Australia. Neither genus occurs in the environs of Zhangpu today. Tetragonula (Tetragonula) florilega Engel, new species, is a generally typical species of the subgenus although it intermingles traits otherwise found in two large species groups of non-Australian Tetragonula s.str. The species of Austroplebeia from Zhangpu is sufficiently distinct to place in a new subgenus, Anteplebeina Engel. Austroplebeia (Anteplebeina) fujianica Engel, new species, shares a long list of character states with Austroplebeia s.str., including the presence of yellow maculation on the face, mesoscutum, mesoscutellum, and metepisternum, found only in this clade among Asiatic Meliponini. Nonetheless, the species differs in the length of the malar space and various features of wing venation. Keys are provided to the genera in Zhangpu amber and the subgenera of both genera. The diversity of stingless bees in amber is discussed, as well as the presence of these two fossils in the Middle Miocene of mainland Asia in relation to prior divergence time estimates, phylogenetic relationships, as well as the paleogeography of the region and potential biogeographic hypotheses. The biological association of stingless bees with resins, particularly the collection of dipterocarp resins, is discussed as Zhangpu amber is a Class II resin likely produced by an extinct species of Dipterocarpaceae (Malvales). Also discussed are the ecological preferences of modern Tetragonula and Austroplebeia relative to the reconstructed paleoenvironment of the Zhangpu amber rainforest. Emendations to the classification of Meliponini are appended, with the following new taxa proposed: Atrichotrigona Engel, new subgenus of Axestotrigona Moure; Lispotrigona V.H. Gonzalez & Engel, new subgenus of Nannotrigona Cockerell; Asperplebeia Engel, new genus; Nanoplebeia Engel, new subgenus of Plebeia Schwarz; Aphaneuropsis Engel, Koilotrigona Engel, Necrotrigona Engel, Dichrotrigona Engel, Nostotrigona Engel, Ktinotrofia Engel, all new subgenera of Trigona Jurine; Chapadapis Engel, new subgenus of Schwarziana Moure. The following higher groups are also established: Hypotrigonina Engel, new sutribe; Heterotrigonitae Engel, new infratribe; Trigoniscitae Engel, new infratribe.