Research Article |
Corresponding author: Michael S. Engel ( msengel@ku.edu ) Academic editor: Michael Ohl
© 2018 Michael S. Engel, Bo Wang, Abdulaziz S. Alqarni, Lin-Bo Jia, Tao Su, Zhe-kun Zhou, Torsten Wappler.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Engel MS, Wang B, Alqarni AS, Jia L-B, Su T, Zhou Z-k, Wappler T (2018) A primitive honey bee from the Middle Miocene deposits of southeastern Yunnan, China (Hymenoptera, Apidae). ZooKeys 775: 117-129. https://doi.org/10.3897/zookeys.775.24909
|
While fossils of honey bees (Apini: Apis Linnaeus) are comparatively abundant in European Oligocene and Miocene deposits, the available material from Asia is scant and represented by only a handful of localities. It is therefore significant to report a new deposit with a fossil honey bee from southern China. Apis (Synapis) dalica Engel & Wappler, sp. n., is described and figured from Middle Miocene sediments of Maguan County, southeastern Yunnan Province, China. This is the first fossil bee from the Cenozoic of southern China, and is distinguished from its close congeners present at the slightly older locality of Shanwang, Shandong in northeastern China. The species can be distinguished on the basis of wing venation differences from other Miocene Apis.
Aculeata , Apinae , Apis , Apoidea , Miocene, taxonomy
Honey bees (genus Apis Linnaeus) are iconic insects. The domesticated Western honey bee, Apis mellifera Linnaeus, is one of the most intensely studied animals (
As is the case for most bees, the fossil record of corbiculate Apinae is comparatively sparse and largely confined to the Cenozoic, with a heavy bias toward material of Eocene through Miocene ages (
Among the fossil Apini, there is apparently a gradation of taxa leading from the earliest Oligocene to the Miocene appearance of the first species of the clade comprising the surviving subgenera Micrapis Ashmead, Megapis Ashmead, and Apis s. str. (
Insect fossils were collected from the northwestern Maguan Basin, southeastern Yunnan, southwestern China (23°01'N, 104°23'E, 1320 m a.s.l.) (Figure
Fossil locality in Maguan County, southeastern Yunnan Province, China. 1 Outcrop overview, green arrow showing layers bearing the present fossil 2 Example of preservation, Acer cf. coriaceifolia H. Lév. (Sapindaceae) preserved together with a nematoceran fly (position indicated by white arrow) 3 Schematic cross section of the studied area.
The sediments bearing the present insect fossils are characterized by cyclic deposits of light-yellow or light-grey pelitic laminated mudstone and siltstone (Figure
Besides insect fossils, the sediments bear abundant fossils of fishes, birds, as well as plants in excellent preservation (Figure
For the description, morphological terminology is adapted from
Worker (Figure
Holotype worker of Apis (Synapis) dalica Engel and Wappler, sp. n., from Maguan County, southeastern Yunnan Province, China. 4 Entire holotype (NIGP154200) as preserved 5 Reconstruction of wing venation; forewing above, hind wing below 6 Detail of foreleg. 7 Detail of apical sterna. Abbreviations: ppl = propleuron, mcx = mesocoxa, tr = trochanter, fm = femur, tb = tibia.
The new species is most similar to those Miocene honey bees described from Shandong Province, China. Apis dalica differs from them in the gently arched basal vein (comparatively straight in the specimens from Shandong), which is also closer to 1cu-a (separated by about a vein width versus several vein widths and even up to 0.5–0.75 times crossvein length in material from Shandong: refer to figures presented by
Worker. Total length (as preserved) 17.06 mm; preserved in ventral orientation, with head thrust forward, wings extended obliquely away from body, and legs largely tucked underneath the body with most podites not preserved or indiscernible; coloration not preserved (appearing uniformly charcoal black). Head apparently slightly longer than wide as interpreted in ventral position; malar space elongate, longer than basal mandibular width; head narrower than mesosoma. Leg podites incompletely preserved. Metasoma typical for worker honey bee, length (as preserved) 9.03 mm, maximum width 4.36 mm; apical margins of sterna somewhat concave, those more basal sterna relatively straight, apical most sterna more strongly concave; sting not extended but slightly evident extending along midline of apical sterna (Figure
Forewing with venation typical of Apis and subgenus Synapis (Figs
The specific epithet refers to the Medieval Dali Kingdom which occupied the area of Yunnan from its founding in 937 AD at the close of the Nanzhao Kingdom and until its termination by Kublai Khan (1215–1294) and the Mongol invasion in 1253 AD.
Fossil honey bees are comparatively uncommon in Asia relative to the wealth of material available from a variety of European deposits of Oligocene and Miocene ages (e.g.,
The discovery of A. dalica expands the known localities with fossil honey bees southward in China and the presence of highly eusocial bees and critical pollinators within the Miocene of fauna of Yunnan. It is hoped that further exploration will recover larger numbers of workers from which the general morphometrics of the species can be determined and more precisely circumscribe the taxon among other Apini, as well as refine phylogenetic relationships among early honey bees. Phylogenetic studies on the modern species have demonstrated that open-nesting is ancestral for the genus (
This research was supported by the National Natural Science Foundation of China (41572010, 41622201, 41688103, U1502231) and the Chinese Academy of Sciences (XDPB05). The project was also supported by the International Scientific Partnership Program (ISPP) at King Saud University through ISPP #0083 (M.S.E. and A.S.A.). T.W. was supported by the German Research Foundation (WA 1496/6-1, Heisenberg grant WA 1496/8-1).