taxonID	type	description	language	source
03B887F1CD36FF94FF5BFC4A0AC8F91B.taxon	description	Perhaps the most immediately distinctive features of Glyptapis are the craterlike punctures of the mesosoma (figs. 2 B, 3, 4), the areolate propodeum (figs. 2 B, 3, 4), the hirsute compound eyes (fig. 2 A), and the large edentate margin to the mandible (fig. 5 I – J). These characters certainly make it possible to easily recognize the bees among fossil Apoidea, but there are numerous other interesting combinations of traits present in the groups, beyond just the aforementioned metatibial scopa. Continued study of these bees over the past 20 years and the availability of the scans reproduced here permit a more thorough and corrected diagnosis for the tribe.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD36FF94FF5BFC4A0AC8F91B.taxon	diagnosis	DIAGNOSIS: Integument apparently without maculation and nonmetallic (integumental coloration poorly preserved in available fossils).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD36FF94FF5BFC4A0AC8F91B.taxon	description	FEMALE. Mandible with single apical tooth and long edentate margin apically composed of expanded trimmal extension (fig. 5 I – J); fimbrial ridge present on mesal surface and paralleling apical edentate margin (fig. 5 I); interdental laminae lacking (fig. 5 I, K). Malar space linear (fig. 5 A – C). Maxillary palpus tetramerous. Labial palpus tetramerous (fig. 5 H); palpomeres I and II flattened, elongate, palpomere I about 0.5 × length of palpomere II; palpomere II with several erect stiff setae ventrally; palpomere III projecting obliquely from axis of palpomere II; ventral (posterior) surface of glossa with longitudinal groove, forming a glossal canal with inner longitudinal ridge and bordered by annulate, ectal surfaces of glossa (fig. 5 G). Clypeus somewhat flattened, extending below lower tangent of compound eyes (fig. 5 A – C); clypeus apical margin covering labral base, thus labroclypeal articulation obscured (fig. 5 A – C). Single subantennal sulcus directed to outer margin of antennal torulus (fig. 5 A). Juxtantennal carina absent (fig. 5 A). Compound eyes hirsute. Posterior margin of vertex gently concave; preoccipital area sharply angled above but not carinate, otherwise rounded (fig. 5 E, F). Pronotal collar virtually absent, with posterior border blending uninterrupted onto dorsal surface without transverse ridge medially (in profile slope continuous and without anterior, transverse ridge demarcating dorsal-facing surface comprising collar relative to lower anterior neck), with transverse dorsal ridge carinate laterally (effaced medially) and extending across pronotal lobe without dorsolateral angle (fig. 4 B) (dorsolateral angle present in Dioxyini except Prodioxys Friese), without dorsolateral ridge extending vertically across lateral surface (fig. 4 B) (present in Dioxyini). Mesoscutum broadly rounded anteriorly (figs. 3 C, 4 C), raised above pronotal posterior margin, with deep craterlike punctures / foveae (figs. 2 B, 3 B, C, 4 B – E) (also present on mesoscutellum and upper mesepisternum); parapsidal lines linear; preaxilla vertical and asetose (as in Dioxyini and Anthidiini); mesoscutalmesoscutellar sulcus deeply impressed, with small, medial, V-shaped notch on mesoscutellum; mesoscutellum low, rounded, posterior margin narrowly vertical with transverse row of small areolae, posterior margin not extending over metanotum; axillae simple (figs. 2 B, 3 D, 4 E); metanotum sloping, without medial tubercles or spines (metanotum tuberculate or spinous in Dioxyini); basal area of propodeum sloping, asetose, areolate (figs. 2 B, 3 D, 4 B, D, E); omaular ridge carinate (fig. 4 B); scrobal sulcus absent; preepisternal sulcus absent; mesepisternum areolate (fig. 4 B); dorsal lamella of metepisternum absent. Forewing (refer to Engel, 2001) with pterostigma proximal to r-rs as long as or slightly longer than pterostigmal width; pterostigmal length 2 × or more its basal width; prestigma more than 2 × as long as broad; basal vein (1 M) arched, thus orthogonal to 1 Cu (as in Ctenoplectrellini and the new tribe described below); marginal cell apex acutely rounded on anterior wing margin; two submarginal cells; 1 m-cu and 2 m-cu entering second submarginal cell, i. e., 2 rs-m proximal to 2 rs-m; hind wing with six distal hamuli. Protibial calcar (fig. 6) with anterior ridge on rachis bordering velum (fig. 6 D) (as in Anthidiini), malus simple, short (fig. 6 D), less than 0.5 × length of velum, apical margin of velum simple (rachis and apex of velum with serrations in Anthidiini and Osmiini); tibiae not spiculate; pro- and mesotibiae with apical outer spine (fig. 6 A), spine fainter on protibia; mesotibial spur long, slender, minutely ciliate (fig. 6); metabasitibial plate absent; metatibial scopa present (fig. 7) (as in Ctenoplectrellini and the new tribe described below; absent in other megachiline tribes); metatibial spurs long, slender, minutely serrate to minutely ciliate (fig. 7); pretarsal claws with short inner ramus (figs. 6, 7 D); arolium present (figs. 6, 7). Metasomal scopa present; sting and associated structures well developed (vestigial in Dioxyini). MALE. Unknown.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD36FF94FF5BFC4A0AC8F91B.taxon	discussion	REMARKS: It is worth noting here a clarification regarding the publication and date from which the name Glyptapis and that of its type species, Glyptapis mirabilis Cockerell, were first made nomenclaturally available. More than 20 years ago, one of us (M. S. E.) followed the assertion of Zeuner and Manning (1976) that the first usage of the names Glyptapis and G. mirabilis were in Cockerell’s (1909 b) paper in The Entomologist, and his more extensive descriptions of the genus, its type species, and the remaining species of the genus appeared subsequently in Schriften der physikalischökonomischen Gesellschaft zu Königsberg (Cockerell, 1909 a). However, examination of the prefatory material to volume 50 of the Schriften reveals at the end of the table of contents an outline of the dates of publication for the various numbers constituting the volume. Cockerell’s (1909 a) article was published 20 September 1909, while the article that Zeuner and Manning (1976) considered as the taxon name’s first usage — erroneously it turns out — was not released until December 1909. Thus, the genus Glyptapis and its type species were made available in the article Cockerell intended (Cockerell, 1909 a) and his usage of the names later that year in The Entomologist was merely a subsequent mention rather than the place from which the names were made available. Why Zeuner and Manning (1976) made such a reversal of precedence for the names is unclear but wrong based on currently available evidence. The same order of precedence applies to the name Ctenoplectrella.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2DFF8FFF6FFF620D9AFE3E.taxon	diagnosis	DIAGNOSIS: Integument with or without maculation (maculation present on face of males of Aspidosmia) and either nonmetallic or weakly dark metallic green (fig. 8).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2DFF8FFF6FFF620D9AFE3E.taxon	description	FEMALE. Mandible tridentate; frimbrial ridge absent (known for Ctenoplectrella and Aspidosmia); interdental laminae lacking. Malar space linear (figs. 8 A, 9 B, C, 10 A, C). Maxillary palpus tetramerous. Labial palpus tetramerous; palpomeres I and II flattened, elongate, palpomere I 0.5 – 0.9 × length of palpomere II; palpomere III projecting obliquely from axis of palpomere II. Clypeus somewhat flattened, not or only slightly extending below lower tangent of compound eyes (figs. 8 A, 9 C, 10 A); clypeus apical margin covering (figs. 8 A, 10 A) or not convering labral base (Aspidosmia), thus labroclypeal articulation either obscured or not (variable among genera, refer to key, below). Single subantennal sulcus directed to outer margin of antennal torulus (fig. 10 A). Juxtantennal carina absent. Compound eyes bare (fig. 8 A). Posterior margin of vertex gently concave; preoccipital ridge rounded, sometimes sharply angled dorsally. Pronotal collar virtually absent, with posterior border blending uninterrupted onto dorsal surface without ridge medially (in profile slope continuous and without anterior ridge demarcating dorsal-facing surface comprising collar relative to lower anterior neck), with dorsal transverse ridge effaced or rounded, without dorsolateral angle, without dorsolateral ridge extending vertically across lateral surface. Mesoscutum broadly rounded anteriorly (fig. 8 B, 9 A), raised above pronotal posterior margin, sculpturing variable, generally smooth to imbricate with scattered small to minute punctures (fig. 9 A); parapsidal lines linear; preaxilla sloping with setae (as in Megachilini); mesoscutal-mesoscutellar sulcus impressed, without V-shaped notch on mesoscutellum; mesoscutellum low, rounded, posterior margin meeting metanotum and not extending over metanotum (fig. 9); axillae simple (figs. 8 B, 9); metanotum sloping, without medial tubercles or spines; basal area of propodeum sloping, asetose, smooth to faintly imbricate (fig. 9); omaular ridge rounded (fig. 9 B, D); scrobal sulcus absent; preepisternal sulcus absent; mesepisternum smooth to faintly imbricate, sometimes with scattered punctures (fig. 9 B, D); dorsal lamella of metepisternum absent. Forewing (fig. 8 B; refer also to Engel, 2001; Gonzalez and Engel, 2011) with pterostigma proximal to r-rs about as long as or slightly longer than pterostigmal width; pterostigmal length 1.75 × or more its basal width; prestigma more than 2 × as long as broad; basal vein (1 M) arched, thus orthogonal to 1 Cu (as in Glyptapini and the new tribe described below); marginal cell apex rounded, slightly offset from anterior wing margin; two submarginal cells; 1 m-cu and 2 m-cu entering second submarginal cell, i. e., 2 rs-m proximal to 2 rs-m; hind wing with 6 – 13 distal hamuli (6 – 7 hamuli in Ctenoplectrella, 7 in Aspidosmiella, 10 in Glaesosmia, 11 – 13 in Aspidosmia). Protibial calcar with or without anterior ridge on rachis bordering velum, malus simple, short, less than 0.5 × length of velum, apical margin of velum simple; tibiae not spiculate; pro- and mesotibiae with apical outer spine, spine less prominent on protibia; mesotibial spur long, slender, minutely ciliate to serrate; metabasitibial plate absent; metatibial scopa present (fig. 8 A) (as in Glyptapini and the new tribe described below; absent in other megachiline tribes); metatibial spurs long, slender (fig. 8 A), minutely serrate to minutely ciliate; pretarsal claws with short inner ramus; arolium present. Metasomal scopa present; sting and associated structures well developed (vestigial in Dioxyini). MALE (known only for Aspidosmia and Ctenoplectrella). Mandible bidentate (fig. 9 C, 10 A) to quadridentate. Metasomal tergum VI without preapical carina; tergum VII exposed, directed posteriorly, without pygidial plate (fig. 11 B – D); sternum VI with apical margin simple (Aspidosmia) or with lateral notches for reception of gonostyli (fig. 11 F) (Ctenoplectrella); sternum VII well sclerotized, not divided medially, apical margin trilobate (fig. 11 G) (medial lobe large in Ctenoplectrella: fig. 11 G; medial lobe formed as short point in Aspidosmia: refer to Peters, 1972); volsella present (fig. 12) (particularly large in Ctenoplectrella), with or without differentiated digitus and cuspis (differentiated in Ctenoplectrella: fig. 12 A; not differentiated in Aspidosmia).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2DFF8FFF6FFF620D9AFE3E.taxon	discussion	REMARKS: A new key is provided to the genera of living and fossil Ctenoplectrellini. The subtribe Glaesosmiina Engel (new subtribe; type genus: Glaesosmia Engel, 2001; diagnosed by combination of subantennal sulcus as long as torular diameter, interocular distance greater than compound eye length, and forewing 2 Rs oblique to 2 M, forming acute apical angle) is here recognized to remove paraphyly of Ctenoplectrellina relative to Aspidosmiina and allow for retaining a group for Aspidosmia and its visible labroclypeal articulation and shorter pterostigmal form relative to the fossils. In addition, Ctenoplectrella phaeton Gonzalez and Engel is removed to a separate genus given its peculiarly sparse pubescence and rather discomfited inclusion in Ctenoplectrella.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2AFF8DFF7AFD9B0EADFBE8.taxon	materials_examined	TYPE SPECIES: Ctenoplectrella phaeton Gonzalez and Engel, 2011.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2AFF8DFF7AFD9B0EADFBE8.taxon	diagnosis	DIAGNOSIS: This genus is generally as described for Ctenoplectrella: tridentate female mandible, pronotal collar virtually absent, and 2 Rs orthogonal to 2 M but is more robust and with a noticeably sparse pubescence, especially on the mesoscutum and mesoscutellum where the surfaces are nearly asetose and the metasomal scopa composed of sparse bands of setae (refer to Gonzalez and Engel, 2011).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2AFF8DFF7AFD9B0EADFBE8.taxon	etymology	ETYMOLOGY: The new generic name is a combination of Aspidosmia and the Latin suffix - ellus, indicating a diminutive. The gender of the name is feminine.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD2AFF83FF75FBEF0F1BFDB9.taxon	type_taxon	TYPE GENUS: Glyptosmia Engel, new genus. DIAGNOSIS: This tribe possesses a unique intermingling of features of Glyptapini, Dioxyini, and Ctenoplectrellini as well as its own distinctive apomorphies. Like Glyptapini and Ctenoplectrellini the metatibia bears a scopa composed of abundant, long setae (fig. 13), although in this tribe many of the setae on the posterior surface are characteristically hooked apically, and some of those on the anterior surface have short apical branches. Likewise, the pretarsal claws have a short, subapical ramus, arolia on all legs, the basal vein is arched, and 2 m-cu is slightly basal to 2 rsm, and thus both 1 m-cu and 2 m-cu enter the second submarginal cell. Like Glyptapini the basal area and lateral surface of the propodeum are areolate and the mandible is long, slender, and has a large edentate margin above the apical tooth. Unlike glyptapines the compound eyes are bare, the omaular ridge is not carinate, and the mesosoma lacks large craterlike punctures and is instead covered with dense, strong, small punctures. Similar to Dioxyini the metanotum is medially beset with tubercles, albeit paired in the fossil versus the single tubercle or spine in the former tribe. Unlike Dioxyini, the sting and its associated structures are well developed. Quite distinctively, the mesoscutellum is flattened and projected posteriorly over the metanotum and the basal edge of the propodeum, and the apical margin is sinuate, and the metatibial spurs are noticeably thickened. FEMALE. Integument without maculation and shiny black (fig. 13). Mandible with single apical tooth and edentate margin apical to expansion of trimmal extension; interdental laminae lacking. Malar space linear. Maxillary palpus tetramerous. Labial palpus tetramerous; palpomeres I and II flattened, elongate, palpomere I subequal to length of palpomere II; palpomere III projecting obliquely from axis of palpomere II. Clypeus somewhat flattened, not extending below lower tangent of compound eyes; clypeus apical margin not covering labral base, thus labroclypeal articulation not obscured. Single subantennal sulcus directed to outer margin of antennal torulus. Juxtantennal carina absent. Compound eyes bare (fig. 14). Posterior margin of vertex gently concave; preoccipital area sharply angled above but not carinate, otherwise rounded. Pronotal collar virtually absent, posterior bor- der blending uninterrupted onto dorsal surface without transverse ridge medially to demarcate collar from dorsal surface of neck, with dorsal ridge rounded laterally without dorsolateral angle (carinate laterally in Glyptapini), without dorsolateral ridge extending vertically across lateral surface. Mesoscutum broadly rounded anteriorly (fig. 13 A, 15), raised above pronotal posterior margin, with dense, small punctures (without the deep craterlike punctures of Glyptapini); parapsidal lines linear; mesoscutalmesoscutellar sulcus deeply impressed and wide (fig. 15), without V-shaped notch on mesoscutellum; mesoscutellum flat (figs. 15, 16 A), posterior margin extending over metanotum and basal edge of propodeum (fig. 16 A), apical margin with broad medial concavity (fig. 15); axillae simple (fig. 15); metanotum vertical, with pair of medial tubercles on either side of midline (fig. 16 A); basal area of propodeum horizontal, asetose, areolate (as in Glyptapini); omaular ridge angled but not carinate (carinate in Glyptapini); scrobal sulcus absent; preepisternal sulcus absent; mesepisternum with dense, small punctures (areolate in Glyptapini); dorsal lamella of metepisternum absent. Forewing (fig. 17) with pterostigma proximal to r-rs longer than pterostigmal width; pterostigmal length more than 2 × its basal width; prestigma more than 2 × as long as broad; basal vein (1 M) arched, thus orthogonal to 1 Cu (as in Glyptapini and Ctenoplectrellini); marginal cell apex rounded, slightly offset from anterior wing margin; two submarginal cells; 1 m-cu and 2 m-cu entering second submarginal cell, i. e., 2 rs-m proximal to 2 rs-m; hind wing with six distal hamuli. Protibial calcar with anterior ridge on rachis bordering velum (as in Anthidiini and Glyptapini), malus simple, short, less than 0.5 × length of velum, apical margin of velum simple; tibiae not spiculate; pro- and mesotibiae with apical outer spine, spine fainter on protibia; mesotibial spur long, slender, minutely serrate; metabasitibial plate absent; metatibial scopa present (figs. 13 B, 18 A) (as in Glyptapini and Ctenoplectrellini; absent in other megachiline tribes); metatibial spurs long, distinctly thickened (fig. 18 B), minutely serrate (not crescentic, ciliate, nor comblike); pretarsal claws with short inner ramus (fig. 18 C, D); arolium present (fig. 18 C). Metasomal scopa present; sting and associated structures well developed (fig. 19). MALE. Unknown.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF83FF68FD230F89FC09.taxon	materials_examined	TYPE SPECIES: Glyptosmia hemiaspis Engel, new species.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF83FF68FD230F89FC09.taxon	diagnosis	DIAGNOSIS: As for the tribe (above).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF83FF68FD230F89FC09.taxon	etymology	ETYMOLOGY: The new genus-group name is a combination of ancient Greek adjective γλυπτός (glyptos, meaning “ engraved ”) and Osmia Panzer (derived from ὀσμή / osme, meaning “ odor ”). The gender of the name is feminine.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF87FF36FB9B0AFFF914.taxon	description	Figures 13 – 19	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF87FF36FB9B0AFFF914.taxon	diagnosis	DIAGNOSIS: As for the genus (above).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF87FF36FB9B0AFFF914.taxon	description	DESCRIPTION: FEMALE. Total body length (as preserved) 4.36 mm; forewing length (based on the left forewing as the right wing is taphonomically stretched) 2.27 mm. Integument black, shining (fig. 13); wing veins dark brown, membranes hyaline and clear (fig. 17). Head slightly longer than wide, length 1.03 mm, width 0.81 mm; labrum about as long as wide, with truncate apical margin; mandible elongate, with distinct apical tooth and larger preapical edentate margin (apex similar to Glyptapis except mandible more elongate); malar space linear; maxillary palpus tetramerous; labial palpomeres I and II long and flattened, albeit each shorter than prementum; clypeus flat, not extending below lower tangent of compound eyes; single subantennal sulcus straight to lower outer angle of torulus (visible faintly from right side; base of sulcus faint on left side), sulcus slightly longer than torular diameter; compound eye bare, slightly broader than gena in profile, inner margins straight and slightly converging below; upper interorbital distance greater than lower interorbital distance; scape short, length 0.18 mm, shorter than torulocellar distance; flagellomere I shorter than combined lengths of flagellomeres II and III (fig. 14 B), flagellomeres II and III of equal lengths; preoccipital ridge sharply angled dorsally, but not carinate, rounded laterally. Mesoscutum length 0.88 mm, mesoscutellum length 0.30 mm; intertegular distance 0.85 mm; parapsidal lines short, linear; transverse mesoscutal-mesoscutellar sulcus deeply impressed and wide, seemingly weakly foveolate within depression (challenging to observe as it is difficult to illuminate properly); tegula oval, brown and semitranslucent, with weak punctures. Forewing (fig. 17) with basal vein (1 M) noticeably arched, orthogonal to 1 Cu; hind wing with six distal hamuli arranged in a single series; jugal lobe more than 0.5 × but less than 0.65 × length of anal lobe (challenging to see precise measurement). Labrum, clypeus, and supraclypeal area with dense, small punctures, punctures separated by much less than a puncture width, nearly contiguous in some places, integument between punctures apparently smooth and shining. Lower face punctured as on clypeus, frons with similar punctures but more spaced (fig. 14), separated by 0.5 – 1.0 × a puncture width, although those on central frons denser, integument between punctures smooth and shining; ocellocular area (fig. 14 A) and vertex with punctures separated by 0.5 – 1.2 × a puncture width; gena apparently with punctures as on vertex. Mesosoma with dense punctures (figs. 13 A, 15), punctures coarser and slightly larger than those of head, punctures nearly contiguous on mesoscutum, mesoscutellum, and pleura, those of pleura seemingly slightly larger than those of mesoscutum (challenging to see clearly given poor state of preservation), integument between punctures smooth and shining. Metanotum irregularly roughened with two rounded tubercles on either side of midline. Basal area of propodeum areolate, with broad seemingly impunctate triangular area on upper posterior surface. Metasomal terga II – V with pregradular surfaces depressed and polished relative to disc and apical marginal zones, pregradular areas smooth to finely imbricate with some scattered, shallow, small punctures apically; tergal discs imbricate with small punctures separated by a puncture width (fig. 16 B), although laterally punctures denser, anterior-facing surface of tergum I with punctures much sparser and integument between smooth (fig. 16 B). Head (fig. 14) with numerous, short, fine, white, erect to suberect, minutely branched setae, such setae not obscuring integument, setae denser in antennal basin and supraantennal area; such setae present on labrum, labrum without distinct patches of setae; clypeus and supraclypeal area with sparsely scattered short, simple setae; frons with scattered, short, simple to minutely branched setae, such setae denser in antennal basins and becoming sparser in ocellocular area and vertex. Mesoscutum with scattered, short, simple, erect, pale setae, such setae slightly longer on mesoscutellar surface except setae along margin of axilla and mesoscutellum denser, long to elongate, lightly fuscous, and minutely branched; pleura with scattered, short, erect, largely simple setae, such setae slightly longer posteriorly and on lateral surface of propodeum; basal area of propodeum without setae. Legs with short, simple or minutely branched, pale, suberect setae except mesotibia with such setae more numerous, slightly longer, and intermixed with some thicker setae, setae on inner surface distinctly longer; mesobasitarsus with similar setae except noticeably longer than those of mesotibial outer surface; metatibia with scopa composed of abundant elongate erect setae, those of outer surface minutely branched apically or some simple, those of inner surface as on outer surface except intermixed with numerous elongate, simple, characteristically hooked setae; metabasitarsus with abundant erect to suberect, elongate, minutely branched, pale setae; meso- and metafemora with sparse, short, largely simple to minutely branched setae, such setae especially minute and simple on ventral and posterior apical surfaces; meso- and metabasitarsi with setae similar to those of corresponding tibiae. Metasomal terga with scattered, short, fine, decumbent, pale setae, such setae sparse on anterior-facing surface of tergum I, intermixed with slightly longer, erect to suberect setae, some with minute branches, laterally; tergum II onward with short discal setae intermixed with somewhat more erect setae and slightly denser apically (but not forming setal bands); narrow apical marginal zones without setae, imbricate; tergum VI with setae denser medioapically; metasomal scopa on sterna II – V composed of bands of long simple erect setae; sternum VI without scopal setae, with dense brush of minute plumose fuscous setae apically, apical margin narrowly truncate; sting simple, sting sheaths thick, short, with some apical setae. MALE. Unknown.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD24FF87FF36FB9B0AFFF914.taxon	materials_examined	HOLOTYPE: Female, CNU-HYM-LF- 2023 - 002 (fig. 13), in a single piece of amber without syninclusions, Fushun coalfield, Liaoning Province, northeastern China, Guchengzi Formation, Ypresian (Eocene); deposited in the fossil insect collection of the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China. The holotype of Glyptosmia hemiaspis is challenging to interpret given its state of preservation (fig. 13). The bee is preserved with the head slightly pulled forward and slightly detached from meso-	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD23FFBBFF5AFA7E0ADCFB8F.taxon	description	There are four extant tribes of corbiculate bees, along with an additional three tribes known only from fossils. These are some of the most familiar of all bees (Michener, 2007; Engel and Rasmussen, 2021): orchid bees (Euglossini), bumble bees (Bombini), stingless bees (Meliponini), and honey bees (Apini). Currently available evidence indicates that the three extinct tribes — Electrobombini, Electrapini, and Melikertini — disappeared during the Eocene-Oligocene transition (Engel, 2001). Some assert that the formal name Corbiculata is attributable to Shuckard (1866). In fact, Shuckard (1866) never established such a name and instead classified corbiculate bees as his Section Cenobites Shuckard, 1866 (160, 302); he also renamed all bees as Mellicolligerae Shuckard [= Anthophila Latreille]. In discussing the systematic arrangement of Apidae, Shuckard (1866: 165) wrote about the character uniting the Cenobites being, “ the glabrous surface of the posterior tibiae, with their lateral edges fringed with bristles slightly curved inwards ... a sort of natural basket for the conveyance of pollen or other stores to the nest. ” He continues, “ This, however, has not been made use of as a main feature for scientific distribution, although they might follow the Dasygasters [Megachilidae], as corbiculated bees, or little basket bearers. ” Thus, Shuckard introduces the term corbicula for this structure and uses the term as an adjectival description of the bees he actually classifies as Cenobites. Accordingly, Shuckard never proposed a “ Corbiculata ” and the formal name, rather than an adjective or new morphological term, cannot be said to derive from his work.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD1CFFBAFC8DFB0E0FFBF916.taxon	description	Melikertines were eusocial, likely living in colonies similar to those of modern stingless bees (Engel, 1998 b; Barden and Engel, 2021; Engel and Davis, 2021). For all but one of the species we know only the worker caste and therefore lack data for variations between workers and the other castes. Accordingly, it remains unknown whether some of the exaggerated morphologies observed in workers may also be present in queens or males. One would initially hypothesize that the queens lack these structures (e. g., facial protuberances), since they would seem to be associated with resin collection (Engel and Davis, 2021), although it cannot be ruled out that they may also be deployed in nest construction or less likely in defense (given the morphologies known, none seem suitable as a defensive weapon). An extensive account of the tribe and its diversity and putative paleobiology was recently provided by Engel and Davis (2021) and is therefore not repeated here except to update their key to genera.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD1FFFBFFF0BFAB50D7FFEA2.taxon	materials_examined	TYPE SPECIES: Thyreomelikertes electrosinicus, new species.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD1FFFBFFF0BFAB50D7FFEA2.taxon	diagnosis	DIAGNOSIS: This is a distinctive genus (figs. 20 – 31), noteworthy for the flattened, trapezoidal mesoscutellum that projects posteriorly well over the metanotum and wholly declivitous propodeum (figs. 23 A, 27). In addition, the mesosoma is densely setose, giving the bees a shaggy appearance (figs. 20, 21, 23 A, 26, 27). The clypeus lacks the protrusions and other modifications that characterize several genera of Melikertini. Among other genera, the new genus is similar to that of Mochlomelikertes Engel, Breitkreuz, and Ohl in Baltic amber as both have elongate and projecting mesoscutella, although that of the latter is vastly more extreme and covers portions of the metasoma (refer to Engel et al., 2014). Mochlomelikertes is noticeably larger at 8 mm in total length (forewing length 6.75 mm), while species of Thyreomelikertes are less than 6 mm in length. A full diagnosis for the genus is as follows: Small bees, less than 6 mm in length; integument seemingly dark brown to black, without maculation (figs. 20 – 31). Mandible with outer mandibular grooves largely reduced, outer ridge prominent, faint indication of auxiliary ridge, clear acetabular groove and pollicar basin, latter divided into auxiliary basin, rutellar basin evidence, condylar ridge forming dorsal ramus apically (refer to Discussion), basins vanishing preapically and proximally, apical margin oblique, with shallow, broad incision demarcating short first preapical tooth, above which a broad concave margin between first preapical tooth and second preapical tooth at upper edge of mandible (upper distal angle), condylar groove in apical third with line of 3 – 4 long, ventrally directed, suberect (obliquely angled toward apex of mandible), thickened, flat setae (fig. 22); malar space linear (fig. 22); labrum flat, slightly broader than long, lateral margins parallel (not converging apically), without lateral fringe setae, apical margin with broad, shallow concavity medially, weakly and broadly rounded lateral to concavity (resulting in a weakly and broadly bilobed appearance to apical margin), surface with sparsely scattered long, fine, erect, simple setae, apical margin with numerous long, simple, fuscous setae, such setae thicker than those of disc; clypeus weakly convex, unmodified (i. e., without clypeal specializations or protrusions); epistomal sulcus laterally forming obtuse angle; upper torular tangent slightly below head midlength; scape elongate, longer than torulocellar distance; flagellomere I longer than wide, longer than flagellomere II; ocelli high on vertex, situated above upper tangent of compound eyes; vertex rounded in facial view, unmodified (no depressions or ridges); preoccipital area rounded; gena narrower than compound eye in profile. \ Mesotibia elongate; mesotibial spur present, simple, elongate; mesotibia and mesobasitarsus densely setose (fig. 24), setae intermixed with bristles, some setae minutely branched and sometimes capitate (fig. 24) (in T. electrosinicus); metatibia slender, elongate (fig. 25 A), posterior margin gently convex and slightly widening medially; surface of corbicula not depressed; posterior margin with fringe of long plumose setae (branches minute and along length of setal rachis) (fig. 25 B), anterior margin with simple or minutely branched setae and bristles, corbicular surface with scattered, erect bristles and long setae; inner surface with keirotrichiate zone field covering most of surface except posterior, narrow, slightly depressed (i. e., a weak clivulus present) subglabrate zone and a broad, squarish apical subglabrate zone (sensu Rasmussen et al., 2017), latter with some scattered short, suberect bristles, apical subglabrate zone slightly longer than apical width of metatibia; rastellum composed of stiff bristles along entire inner apical width of metatibia; single metatibial spur present, spur minutely ciliate along inner margin; metabasitarsus with auricle present on proximal surface facing apex of metatibia; metabasitarsus rectangular (fig. 25 A) to squarish, longer or slightly longer than wide, margins roughly parallel, apical margin comparatively straight to concave, inner surface with abundant, elongate, suberect, simple bristles arranged in loose comb rows; pretarsal claws with minute inner subapical ramus; arolium present (figs. 20, 21, 25 A). Metasoma broad, ovoid (figs. 20, 21), with scattered, short, erect to suberect setae, such setae more abundant laterally; metasomal sterna II – V with abundant, long, erect, fine, simple setae, those setae of sterna IV and V shorter and more suberect to decumbent; sting present (difficult to observe in type species, although with care it can be found, but easily seen in the second species where the sting is exserted), simple (i. e., without proximally directed barbs).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD1FFFBFFF0BFAB50D7FFEA2.taxon	etymology	ETYMOLOGY: The new genus-group name is combination of the ancient Greek noun θυρεός (thyreos, meaning, “ long shield ”) and Melikertes, type genus of the tribe. The gender of the name is masculine.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD18FFB3FD06FAE50CBDFA1D.taxon	description	Figures 20 – 25	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD18FFB3FD06FAE50CBDFA1D.taxon	diagnosis	DIAGNOSIS: This species differs from its only congener (see below for alternative character states) in the that the metatibial fringe, fundal, and corbicular setae are as long as than the maximum width of the metatibia (figs. 20, 21, 25), and similarly, the setae of mesotibia are about as long as the maximum mesotibial width and are intermixed with capitate setae (fig. 24). The species is also larger, with a forewing length over 5 mm. It also seems that the facial setae are sparser than those of the other species (fig. 22).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD18FFB3FD06FAE50CBDFA1D.taxon	description	DESCRIPTION: WORKER. Total body length (as preserved) 5.44 mm; forewing length (as preserved) 5.30 mm. Head slightly longer than wide, length (summit of vertex to clypeal apical margin) 1.47 mm, width (maximum width across compound eyes, eyes partially collapsed inward thereby shortening the value) 1.29 mm; upper interorbital distance 1.26 mm, lower interorbital distance 1.22 mm. Scape elongate, length 0.58 mm, longer than torulocellar distance; flagellomere I longer than flagellomere II, about as long as combined lengths of flagellomeres II and III, flagellomere II shorter than flagellomere III. Gena distinctly narrower than compound eye in profile. Pronotum short, declivitous; mesoscutum anterior border broadly rounded, anterior lip gently curving to meet posterior pronotal margin; mesoscutum medial length 1.04 mm; intertegular distance 1.07 mm; mesoscutellum medial length 0.39 mm. Metatibia slender, length 2.09 mm, maximum width 0.41 mm; metabasitarsus longer than wide, length 0.81 mm, maximum width 0.34 mm, apical margin comparatively straight. Forewing with basal vein (1 M) straight, confluent with 1 cu-a, pterostigma longer than wide, maximum width slightly proximal to midlength, margin inside marginal cell sloping to costal margin; marginal cell broadly rounded apically, well separated from anterior wing margin; 2 Rs arched posteriorly; 3 Rs shorter than r-rs and slightly shorter than 4 Rs; 1 rs-m straight, 2 rs-m strongly arched apically in posterior half; 2 m-cu proximal 2 rs-m by about vein width; wing membranes hyaline clear; veins dark brown. Labrum faintly imbricate and impunctate; clypeus seemingly smooth with some weak, shallow punctures laterally; integument of much of face difficult to discern owing to collapse of face, but vertex smooth and shining, seemingly impunctate. Mesoscutum and mesoscutellum apparently smooth with sparse faint punctures (integument exceptionally challenging to observe); metasomal terga smooth to faintly imbricate and shining, with minute punctures at bases of setae. Pubescence generally fuscous and rather abundant; labrum with sparse, erect, simple setae on surface, apical margin with fringe of elongate setae, such setae about 0.75 × labral length; clypeus with scattered, simple, erect, short setae, such setae more numerous on remainder of head and becoming quite elongate on vertex. Scape with scattered short, erect to decumbent, simple, fine setae, without bristles. Mesosoma rather densely pubescent but not obscuring integument, setae elongate, erect, and simple or with minute branches along rachis, many setae noticeably thick on all surfaces except metanotum and basal area of propodeum; plumose setae more numerous on posterior of mesoscutellum, with branches slightly longer and more numerous, thus appearing brushy. Wing membranes with abundant microtrichia. Profemur and protibia with abundant, short setae, those of dorsal surfaces longer than those of prolateral, retrolateral, and ventral surfaces; probasitarsus with long, erect bristles on dorsal surface, similar bristles on other surfaces except slightly shorter; mesofemur with abundant short, erect setae, intermixed with short bristles on outer surface, ventrally setae shorter and bristles lacking; mesotibia with dense, erect setae and bristles, on dorsal surface, such setae and bristles progressively longer from base to near apex, setae with some minute branches, retrolateral surface with bristles sparse and setae longer, about as long as mesotibial width, such setae with abundant minute branches and some with thickened apices, thus appearing somewhat clavate apically, ventral surface with setae shorter; mesobasitarsus similar to mesotibia except bristles longer and more numerous on dorsal surface and more evident on retrolateral surface, setae of retrolateral surface not appear clavate; metafemur with short, erect setae on dorsal and prolateral surfaces, retrolateral and ventral surfaces with setae exceedingly sparse, with a patch of keirotrichia on retrolateral surface apically; metatibia with abundant, long, erect bristles along anterior edge of proventral surface and inferiorly on prolateral surface, such setae intermixed with long, minutely branched setae, proventral surface largely asetose, prolateral surface with sparsely scattered long, erect, simple setae and bristles, such setae similar on profundal and corbicular surfaces, retromarginal fringe composed of dense, long, minutely branched setae, such setae about as long as maximum metatibial width, retrolateral surface with keirotrichiate zone covering majority of surface except narrow, slightly depressed, superior, subglabrate zone and squarish, apical subglabrate zone; rastellum composed of dense series of thick bristles tapering in length; metabasitarsus with abundant erect bristles on proventral and prolateral surfaces, bristles intermixed with some short setae, retromarginal edge with bristles less prominent, with dense minutely branched setae, such setae slightly shorter than bristles and rounding onto superior apical angle, retrolateral surface with loose comb rows composed of long, suberect bristles; auricle bordered by fringe of short, branched setae. Metasomal setation as described for genus (above). QUEEN AND MALE. Unknown.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD18FFB3FD06FAE50CBDFA1D.taxon	materials_examined	HOLOTYPE: Female worker, CNU-HYM- LF- 2023 - 001 (figs. 20, 21), in a single piece of amber without syninclusions, Fushun coalfield, Liaoning Province, northeastern China, Guchengzi Formation, Ypresian (Eocene); deposited in the fossil insect collection of the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China. The holotype is the best preserved of the available bees from Fushun amber and is virtually complete, and observation is not hampered by prominent syninclusions, bubbles, or other imperfections in the amber. The bee is rather close to some curved surfaces that prevent preparation of the amber surface from some angles — the only hindrance to a perfect view of certain features. The wings are extended posteriorly over the body, although the left forewing is slightly more askew. The forelegs are raised near the head, although the left foreleg is more extended into the amber beneath the bee. The mid and hind legs are extended below and, with the exception of the left midleg, project out from the body. The worst defects concern the head, which is somewhat obscured by an oblique internal fracture. More critically, the integument of the face is partially collapsed, rendering interpretation a challenge. The top of the head and the mesosomal nota are partially obscured by encrusted material entangled in the setae, somewhat obscuring the integument. The mesosoma and metasoma are partially compressed but inconsequentially. The wings are excellent although the left forewing was slightly stretched during preservation, resulting in slight fractures along some of the fenestrae, bullae, and across the marginal cell. The right forewing is unmolested.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD18FFB3FD06FAE50CBDFA1D.taxon	etymology	ETYMOLOGY: The specific epithet is a combination of the Latin adjective electricus (from electrum), meaning, “ of amber ” and the Medieval Latin adjective sinicus, meaning “ of or related to China. ”	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD14FFB4FD52FA6A0F07FB06.taxon	description	Figures 26 – 31	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD14FFB4FD52FA6A0F07FB06.taxon	diagnosis	DIAGNOSIS: This species, which is known from three individuals (figs. 26, 27, 31 A), can be distinguished by the distinctly more elongate fringe, fundal, and corbicular setae on the metatibia (figs. 27, 29, 30, 31 A, B), the setae much longer than the maximum metatibal width. Likewise, the setae of the mesotibia are much longer than the maximum mesotibial width (up to about 1.5 × mesotibial width) and none of the setae are capitate. The species is overall smaller than that of the type species (refer to metrics) and the apical margin of the metabasitarsus is concave (fig. 30 A) (rather than straight). The facial setae, particularly on the clypeus, are seemingly denser than in T. electrosinicus (fig. 31 B).	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD14FFB4FD52FA6A0F07FB06.taxon	description	DESCRIPTION: As described for Thyreomelikertes electrosinicus (above), with the following exceptions: WORKER. Total body length (as preserved) 3.51 mm; forewing length (as preserved) 3.78 mm. Head slightly wider than long (fig. 31 B), length (summit of vertex to clypeal apical margin) 1.16 mm (paratype 1.21 mm), width (maximum width across compound eyes) 1.66 mm (paratypes 1.32 – 1.46 mm). Scape elongate, length 0.47 mm (artificially stretched) (paratypes 0.23 – 0.38 mm, both damaged), longer than torulocellar distance; flagellomere I longer than flagellomere II, shorter than combined lengths of flagellomeres II and III, flagellomere II shorter than flagellomere III. Mesoscutum medial length 0.57 mm; intertegular distance 0.67 mm (paratype 0.55 mm); mesoscutellum medial length 0.36 mm. Metatibia slender, length 1.50 mm (paratypes 1.11 – 1.61 mm), maximum width 0.45 mm (paratypes 0.22 – 0.37 mm); metabasitarsus longer than wide, length 0.50 mm (paratype 0.34 mm), maximum width 0.37 mm (paratype 0.22 mm), apical margin concave. Forewing (fig. 28) with basal vein (1 M) slightly proximad 1 cu-a, pterostigma maximum width at midlength; marginal cell apex well separated from anterior wing margin, with nebulous appendiculate vein; 3 Rs much shorter than both r-rs and 4 Rs; 1 rs-m arched; 2 m-cu proximal 2 rs-m by about 2 – 3 × vein width. Mesotibia with dense, erect setae and bristles (fig. 29 A), on dorsal surface, such setae and bristles progressively longer from base to apex, setae with some minute branches, retrolateral surface with bristles sparse and setae longer, longer than mesotibial width, such setae with abundant minute branches, no capitate setae; metatibia with dense elongate (distinctly longer than metatibial width), minutely branched setae forming retromarginal fringe (figs. 29 B, 30, 31 A), similar but shorter setae along proventral margin, retromarginal fringe sparsely intermixed with long, erect bristles and such bristles sparse along anterior edge of proventral surface and inferiorly on prolateral surface, proventral surface largely asetose, prolateral surface with sparsely scattered long, erect, simple setae and bristles, such setae similar on profundal and corbicular surfaces. QUEEN AND MALE. Unknown.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD14FFB4FD52FA6A0F07FB06.taxon	materials_examined	HOLOTYPE: Female worker, CNU-HYM- LF- 2023 - 003 (figs. 26, 27), in a single piece of amber with two further bees as syninclusions, Fushun coalfield, Liaoning Province, northeastern China, Guchengzi Formation, Ypresian (Eocene); deposited in the fossil insect collection of the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China. The holotype is preserved in a relatively clear piece of amber and has two syninclusions of the same species as well as a tiny chalcidoid wasp near the wing apex. The holotype is located at one end of the piece, while the two other bees are clustered together and overlapping each other. The holotype is well preserved and can be observed in all orientations except ventral. The wings are obliquely folded over the body and the legs tucked up alongside the body. PARATYPES: Female workers, CNU-HYM- LF- 2023 - 004 and CNU-HYM-LF- 2023 - 005, in the same piece of amber with the holotype (fig. 26), Fushun coalfield, Liaoning Province, northeastern China, Guchengzi Formation, Ypresian (Eocene); deposited in the fossil insect collection of the Key Laboratory of Insect Evolution and Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China. The paratypes are distinctly less well preserved relative to the holotype and, given that they are partially entwined as well as damaged through compression or missing sclerites, it was not possible to see each structure as well as in the holotype. It was also not possible to get all of the same measurements from the paratypes as was possible from the holotype. From observable characters, the paratypes agree perfectly with the holotype and so we are confident in their conspecificity with the holotype.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
03B887F1CD14FFB4FD52FA6A0F07FB06.taxon	etymology	ETYMOLOGY: The specific epithet honors Chuijin Kong, who graciously donated the specimens for this study.	en	Engel, Michael S., Xie, Jiaying (2024): The Bee Fauna Of Eocene Fushun Amber (Hymenoptera: Apoidea). Bulletin of the American Museum of Natural History 2024 (469): 1-81, DOI: 10.1206/0003-0090.469.1.1, URL: https://bioone.org/journals/bulletin-of-the-american-museum-of-natural-history/volume-2024/issue-469/0003-0090.469.1.1/The-Bee-Fauna-of-Eocene-Fushun-Amber-Hymenoptera-Apoidea/10.1206/0003-0090.469.1.1.full
