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Electronic Supplement File 2: liu&al-myrmeciathorax-asp2019-electronicsupplement-2.xls — Table S2. Muscular homology chart of Hymenoptera (present with “+” or muscular name in green, absent with “–” in pink, uncertain with “?” or “/” in yellow. In Formicidae, the muscles only present in workers are labeled in dark green; muscles only occurring in alate castes labeled in dark blue.)

Electronic Supplement File 1: liu&al-myrmeciathorax-asp2019-electronicsupplement-1.xls — Table S1. Character list and matrix.

The mesosomal skeletomuscular system of workers of Myrmecia nigrocincta was examined. A broad spectrum of methods was used, including micro-computed tomography combined with computer-based 3D reconstruction. An optimized combination of advanced techniques not only accelerates the acquisition of high quality anatomical data, but also facilitates a very detailed documentation and visualization. This includes fine surface details, complex configurations of sclerites, and also internal soft parts, for instance muscles with their precise insertion sites. Myrmeciinae have arguably retained a number of plesiomorphic mesosomal features, even though recent molecular phylogenies do not place them close to the root of ants. Our mapping analyses based on previous morphological studies and recent phylogenies revealed few mesosomal apomorphies linking formicid subgroups. Only five apomorphies were retrieved for the family, and interestingly three of them are missing in Myrmeciinae. Nevertheless, it is apparent that profound mesosomal transformations took place in the early evolution of ants, especially in the flightless workers. The modified mesosoma is characterized by four character complexes: a) an enlarged prothorax with elongate procoxae, a large plate-like pronotum, strongly developed muscles of the forelegs and especially of the neck region; b) highly differentiated legs with complex cleaning and attachment devices; c) a reduced flight apparatus with greatly simplified pterothoracic musculature and mechanically reinforced exoskeleton and d) strongly developed specialized muscles inserted on the base of the metasoma. Structural modifications of the prothorax and neck region allow ant workers to transport items efficiently with a highly movable head with strongly developed cervical muscles. Their differentiated legs enable them to move efficiently on various surfaces and to maintain their complex apparatus of sensilla. The mechanically reinforced mesosoma provides protection against predators and likely against detrimental environmental agents. The enhanced movability of the metasoma increases the defensive capacity with a sting or other mechanisms.

A series of traditional and modern morphological techniques were used to examine and document the thoracic skeletomuscular structures in detail of the flightless insects from BIG4 orders: Lepidoptera, Coleoptera, Diptera and Hymenoptera. The flightlessness of female Nyssiodes lefuarius (Lepidoptera: Geometridae) with an independent evolution background is linked not only with the modifications of flight related skeletomuscular structures, but also with the enlarged ovaries and the reduced digestive system. The evolutionary scenario of flight ability enhancement in Lepidoptera is demonstrated using the combined morphological characters mapping on the phylogenometric topology. The extremely simplified thoracic skeletomuscular structures and a well-developed swimming apparatus of Orectochilus villosus (Coleoptera: Gyrinidae) were clearly demonstrated. The limited skeletomuscular elements take over more functions to support the flight capacity. The strongly flattened thorax and the specialized leg structures of Crataerina pallida (Diptera: Hippoboscidae) adapt to the ectoparasitism. The flight related skeletomuscular structures are strongly modified as the reason of the flightlessness of this species. The monophyly of Hippoboscoidea is supported based on morphological parsimony analysis. The thoracic skeletomuscular system of workers of Myrmecia nigrocincta (Hymenoptera: Formicidae) was examined. The primitive status of Myrmeciinae was challenged based on recent researches from both morphology and phylogenomics. The strongly modified thoracic structures of the worker ants support the entire family as an ecologically dominant group. Parsimony analyses support a sister group between Strepsiptera and monophyletic Coloptera. Archostermata is recovered as the sistergroup of the remaining suborders of Coleoptera, and Polyphaga as the sister group of Myxophaga. The Bayesian analyses based on the same morphological dataset provide a different topological pattern.

Ammothereva Lyneborg Ammothereva Lyneborg, 1984: 206 (including revision and illustration: numerous, e.g. antennae, heads, male genitalias.). Type species, Psilocephala gessakovskyi Zaitzev, 1973, by original designation. Gaimari & Irwin, 2000: 163 (diagnosis, phylogenetics, biogeography, illustrations, in key). Holston et al., 2005 (phylogenetics)., Published as part of Liu, Si-Pei, Gaimari, Stephen D. & Yang, Ding, 2012, Species of Ammothereva Lyneborg, 1984 (Diptera: Therevidae: Therevinae: Cyclotelini) from China, pp. 1-13 in Zootaxa 3566 on page 4, DOI: 10.5281/zenodo.213838, {"references":["Lyneborg, L. (1984) Ammothereva, a new Palaearctic genus of Therevidae, with a review of the 13 included species (Insecta, Diptera). Steenstrupia, 10 (7), 205 - 222.","Zaitzev, V. F. (1973) New species of stiletto flies of the genus Psilocephala Zett. (Diptera, Therevidae) from Palaearctic. Entomologicheskoe Obozrenie, 52, 703 - 713. [in Russian; English translation, 1974, Entomological Review, 52, 471 - 477]","Gaimari, S. D. & Irwin, M. E. (2000) Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Therevidae). Zoological Journal of the Linnean Society, 129 (2), 129 - 240."]}

Ammothereva brevis sp. nov. (Figs. 23���35) Diagnosis. Eyes separated on frons by a distance about 3 times width of anterior ocellus. Frons yellowish orange in anterior third. Antennal scape strongly elongated and yellowish orange basally. Upper occiput covered with uniform scale-like white pile, without postocular setae. Outer gonocoxal process large, as high as gonocoxite, and bifurcated apically. In aedeagus, lateral ejaculatory process distinct and separated from dorsal and ventral apodemes. Description. Male. Body length 8.5 mm, wing length 6.2 mm. Head (Fig. 23) with dense pale pruinescence over black ground color except lower frons with brown ground color; anterior third of frons with yellowish orange pruinescence. Scale-like white pile on ocellar tubercle, frons, gena and occiput; upper occiput without postocular setae; parafacial bare. Eyes separated on frons by a distance about 3 times width of anterior ocellus. Antenna with dense pale pruinescence; scale-like white pile on scape and pedicel; scape strongly elongated, yellowish orange basally and dark brown apically; pedicel barrel-shaped, dark brown; flagellum broken off so no further information available; antennal ratio (as length ratio of scape: pedicel) 4.1: 1.0. Proboscis yellow with large labellum and covered with brown pile; palpus with scale-like yellow pile. Thorax with dense pale pruinescence; notum with black ground color; pleuron with brown ground color; prosternum yellow; mesonotum with 2 brown vittae separated by narrow grey stripe. Mesonotum with short scalelike white pile; prosternum and pleuron with white pile; macrosetae on thorax pale yellow. Scutal chaetotaxy (pairs): np 5, sa 2, pa 1, dc 2, sc 2. Legs yellowish brown, except tips of tibiae and tarsi dark brown, pale pruinescence on coxae to femora, pulvilli yellow. White pile present on coxae and femora, setae of coxae through basal tibiae pale yellow, setae of apical tibiae to tarsi dark brown. Fore coxa with a 1, av 1; mid coxa with a 2; hind coxa with a 3���4, d 1. Fore femur with av 4; mid femur with av 3. Fore tibia with ad 4, pv 5; mid tibia with ad 4, pd 3, av 5, pv 5, apically with 5 setae. Hind legs broken off so no further information is available. Wing hyaline, tinged brownish yellow; pterosigma very narrow, yellow, at end of R 1; veins yellow except apices brown; cell m 3 closed with short petiole apically. Halter stalk and knob yellow. Abdomen yellow with pale pruinescence, except basal parts of tergites dark brown. Dense scale-like pile on tergites recumbent, sparse yellow pile on sternites. Terminalia with dense yellowish brown pile. Male genitalia: Epandrium (Fig. 27) elongated, 1.2 times longer than wide, with posterior edge emarginate, with broadly rounded posterolateral flange. Subepandrial sclerite with semicircular medial invagination; nearly as long as cercus distally. Cercus ovoid. Hypandrium short. Gonocoxite (Fig. 28) with broad, apically bifurcated outer gonocoxal process. Gonostylus much elongated, nearly 9 times longer than wide. Aedeagus (Fig. 29) with dorsal apodeme wide; ventral apodeme short and narrow; lateral ejaculatory process clearly separated from dorsal and ventral apodeme; distiphallus (Fig. 30) short and ventrally curved. Female. Body length 11.7 mm, wing length 7.0 mm. Most characters of female similar to male, with following exceptions. Eyes separated on frons by distance about 5 times width of anterior ocellus. Antennal setae shorter than in male; flagellum broken off so further information unavailable; antennal ratio (as length ratio of scape: pedicel) 4.8: 1.0. Proboscis yellow with yellow pile; palpus yellow with long yellow pile. Setae on legs from coxae through femora and on ventral tibiae pale yellow; setae from dorsal tibiae through tarsi dark brown. Fore coxa with a 1, av 1; mid coxa with a 3; hind coxa with a 3, d 1. Fore femur with av 4; mid femur with av 2; hind femur with av 4. Fore tibia with ad 4, pd 3, pv 4���5, apically with 2���3 setae; mid tibia with ad 5, pd 3, av 4, pv 5, apically with 5 setae; hind tibia with ad 3, pd 2, av 4, pv 4, apically with 7 setae. Sternites of abdomen black, with pale pruinescence. Scale-like white pile on tergites shorter and less dense than in male; sparse white pile on sternites. Female terminalia: Sternite 8 (Fig. 32) rather large with apical incision. Acanthophorite A 2 setae on tergite 10 sharp apically. Cercus semicircular in lateral view, but slightly elongated in dorsal and ventral views. Subepandrial sclerite rounded basally and distinctly narrow apically. Furca (Fig. 33) 1.3 times longer than wide, with furcal bulla anteriorly. Accessory glands with separated ducts. Spermathecal sac relatively small and spherical; 2 spherical spermathecae present. Type material. Holotype male, CHINA: Ningxia, Jingheyuan (35 �� 24 'N, 106 �� 24 'E, 1940 m), 13. VII. 1980, Chi-Kun Yang. Paratype, 1 ��, same data as holotype. Distribution. Palaearctic: China (Ningxia) (Fig. 35). This is biogeographically part of the North China Region (Zhang 1999). Remarks. This new species is similar to A. salentioides Lyneborg in the following features: eyes separated on frons by a distance about 3 times width of the anterior ocellus; scape of antenna strongly elongated; upper occiput covered with uniform pile without postocular setae; epandrium with broadly rounded posterolateral flange; outer gonocoxal process large, bifurcated apically. But the new species can be separated from it by the following features: anterior frons and basal scape yellowish orange; epandrium elongated, 1.2 times longer than wide; aedeagus with dorsal apodeme distinctly shorter than ejaculatory apodeme; lateral ejaculatory process distinct and separated from dorsal and ventral apodemes. In A. salentioides, the anterior frons is dark and the antennal color is entirely blackish; the epandrium is about as long as wide; the dorsal apodeme of the aedeagus (Lyneborg 1984: 220, fig. 48) is slightly longer than the ejaculatory apodeme; the lateral ejaculatory process (Lyneborg 1984: 220, fig. 47) is completely subsumed into the dorsal and ventral apodemes. Etymology. The specific name refers to the short dorsal apodeme of the aedeagus, from the Latin adjective ��� brevis ��� meaning short., Published as part of Liu, Si-Pei, Gaimari, Stephen D. & Yang, Ding, 2012, Species of Ammothereva Lyneborg, 1984 (Diptera: Therevidae: Therevinae: Cyclotelini) from China, pp. 1-13 in Zootaxa 3566 on pages 8-12, DOI: 10.5281/zenodo.213838, {"references":["Zhang, R. - Z. (1999) Zoogeography of China. Science Press, Beijing, 487 pp.","Lyneborg, L. (1984) Ammothereva, a new Palaearctic genus of Therevidae, with a review of the 13 included species (Insecta, Diptera). Steenstrupia, 10 (7), 205 - 222."]}

Ammothereva nuda sp. nov. (Figs. 1 ���11, 35) Diagnosis. Frons densely silver pruinose, but without any pile or setae. Antenna entirely yellow. Macrosetae on thorax black. Hind femur yellow. Abdominal tergites 2���3 yellow, each with a dark brown central spot. Epandrium as long as wide with distinct posterolateral flange. Outer gonocoxal process long and narrow. Aedeagus with dorsal apodeme wide. Description. Male. Body length 9.2 mm, wing length 6.4 mm. Head (Fig. 1) with dense pale pruinescence over black ground color; frons covered with dense silver pruinescence, upper frons light brown. White pile from gena to occiput, upper occiput with some black postocular setae; ocellar tubercle, frons and parafacial bare. Eyes nearly contiguous on upper frons. Antenna (Fig. 2) yellow with pale pruinescence; black setae on scape thick, several being very strong, but those on pedicel and basal first flagellomere short and thin; scape conical; pedicel ovoid; first flagellomere swollen in basal part and wider than scape and pedicel; style apical with tiny distal spine; antennal ratio 1.3: 1.0: 5.0: 0.9. Proboscis brownish yellow with brown pile; palpus yellow with yellow pile. Thorax with dense pale pruinescence over black ground color; mesonotum with 2 pale yellow vittae. Notum and pleuron nearly bare, with only sparse short white pile; prosternum bare; macrosetae on thorax black, scutal chaetotaxy (pairs): np 3, sa 2, pa 1, dc 1, sc 2. Coxae and trochanters yellow with pale pruinescence; hind femur yellow, hind tibia yellow with dark brown tip, hind tarsomere 1 yellow with dark brown apex, other hind tarsomeres dark brown, hind pulvilli dark brown (fore and mid legs broken off, so further information not available). Coxae and trochanters with white pile and yellow setae, posterior surface of mid coxa bare; hind femur, tibia and tarsus with black setae. Fore coxa with a 1, av 1; mid coxa with a 2; hind coxa with a 3���4, d 1. Hind femur with av 5, pv 3. Hind tibia with ad 14, pd 9, av 8, pv 6, apically with 5 setae. Wing (Fig. 4) hyaline, tinged yellow; pterostigma very narrow, brown, at end of R 1; veins yellow in basal half of wing, becoming brown in distal half; cell m 3 closed with a short petiole apically. Halter stalk brown in basal half, becoming yellow distally; knob yellow. Abdomen yellow with pale pruinescence, except basal parts of segments 3���7 dark brown; covered with sparse short yellow pile mixed with short black recumbent pile; tergites 2���3 each with dark brown median spot. Terminalia yellow. Male genitalia: Epandrium (Fig. 8) as long as wide, with distinct posterolateral flange, a dark brown bell-shaped spot in central area. Subepandrial sclerite with a shallow medial invagination, slightly shorter than cercus distally. Hypandrium short. Gonocoxite (Fig. 9) with long and narrow outer gonocoxal process; ventral lobe triangular. Gonostylus elongated, nearly 3 times longer than wide. Dorsal apodeme of parameral sheath (Fig. 10) wide, nearly 5 times wider than ventral apodeme; ventral apodeme short and narrow; distiphallus (Fig. 11) long and curved ventrally. Female. Unknown. Type material. Holotype male, CHINA: Inner Mongolia, Helan Mountain, Yaoba, Xiazigou (39 ��00'N, 105 �� 50 'E), 30. VII. 2010, Li-Hua Wang. Distribution. Palaearctic: China (Inner Mongolia) (Fig. 35). This is biogeographically part of the Mongolia- Xinjiang Region (Zhang 1999). Remarks. This new species is similar to A. splendida (Kr��ber 1912) in having the black antennal setae, the black mesonotal macrosetae and the relatively wide dorsal apodeme of the aedeagus. But it can be separated from it by the following features: frons bare without any pile or setae; hind femur totally yellow; tergites 2���3 each with a dark brown central spot; epandrium as long as wide; outer gonocoxal process distinctly narrow; distiphallus directly downcurved. In A. splendida, the frons has the blackish pile as long as width of the scape; the femora are brownish black; tergite 2���5 each have the large brownish black triangular areas in the middle; the epandrium (Lyneborg 1984: 210, fig. 12) is 1.2 times longer than wide; the outer gonocoxal process (Lyneborg 1984: 210, fig. 11) is wide basally and gradually tapers distally; the distiphallus (Lyneborg 1984: 210, fig. 13) is dorsally recurved basally before turning ventrally. The species Ammothereva mongolica (Zaitzev, 1970) is not in the key, because only the female is known. Despite this species sharing the characteristic of having black mesonotal setae, the new species differs in having the coxae (Fig. 3) and hind femur yellow. In A. mongolica, the coxae and hind femur are black. Etymology. The specific name refers to the bare frons; from the Latin adjective ���nudus��� meaning bare or naked; a feminine adjective., Published as part of Liu, Si-Pei, Gaimari, Stephen D. & Yang, Ding, 2012, Species of Ammothereva Lyneborg, 1984 (Diptera: Therevidae: Therevinae: Cyclotelini) from China, pp. 1-13 in Zootaxa 3566 on pages 4-5, DOI: 10.5281/zenodo.213838, {"references":["Zhang, R. - Z. (1999) Zoogeography of China. Science Press, Beijing, 487 pp.","Krober, O. (1912) Monographie der palaarktischen und afrikanischen Thereviden (Dipt.). Deutsche Entomologische Zeitschrift, 1912, 109 - 140.","Lyneborg, L. (1984) Ammothereva, a new Palaearctic genus of Therevidae, with a review of the 13 included species (Insecta, Diptera). Steenstrupia, 10 (7), 205 - 222.","Zaitzev, V. F. (1970) New species of genus Neothereva (Diptera, Therevidae) from Palearctics. Zoologicheskii Zhurnal, 49, 795 - 797. [in Russian]"]}

Ammothereva flavifemorata sp. nov. (Figs. 12 –22, 35) Diagnosis. Antenna with apical part of first flagellomere and style dark brown. Macrosetae on thorax pale yellow. Legs entirely yellow. Tibiae with mixed pale and black setae. Abdomen mostly black with pale pruinescence. Aedeagus with dorsal apodeme narrow, lateral ejaculatory apodeme narrow and not extending beyond edge of dorsal apodeme, distiphallus long and slender. Description. Male. Body length 4.2–6.5 mm, wing length 3.8–5.5 mm. Head (Fig. 12) with dense pale pruinescence over black ground color; frons with light brown pruinescence. White pile from gena to occiput, upper occiput with some pale yellow postocular setae; ocellar tubercle, frons and parafacial bare. Eyes nearly contiguous on upper frons. Antenna (Fig. 13) yellow with pale pruinescence, except apex of first flagellomere and entire style dark brown; scape and pedicel with pale yellow pile, several strong black setae on apical scape and basal first flagellomere; scape conical; pedicel ovoid; first flagellomere swollen basally, wider than scape and pedicel; style apical with tiny distal spine; antennal ratio 2.3: 1.0: 4.3: 1.1. Proboscis pale yellow with short yellow pile; palpus pale yellow with white pile. Thorax with dense pale pruinescence over black ground color. White pile on marginal notum and pleuron; prosternum bare; macrosetae on thorax pale yellow. Scutal chaetotaxy (pairs): np 3, sa 2, pc 1, dc 1, sc 2. Legs entirely yellow; pulvilli yellow. Coxae and femora with white pile and pale yellow setae, but several setae on apical femora black; posterior surface of mid coxa bare; tibiae with pale yellow setae mixed with several black setae; tarsi with black setae. Fore coxa with a 1, av 1; mid coxa with a 2; hind coxa with a 1–3, d 1. Fore femur with av 1; mid femur with av 1, pv 2; hind femur with av 3–4, pv 1. Fore tibia with ad 1–2, pd 0–3, pv 3–4, apically with 5–6 setae; mid tibia with ad 3, pd 2, av 2, pv 3–4, apically with 5–6 setae; hind tibia with ad 8, pd 5–7, av 6–8, pv 4–5, apically with 8–9 setae. Wing hyaline, tinged yellow; pterostigma very narrow, yellow, at the end of R 1; veins brown; cell m 3 closed with short petiole apically. Halter stalk yellow, knob pale yellow. Abdomen with pale pruinescence; covered with white pile; tergites black in ground color except posterior corners of each tergite yellow, sternites yellow except basal area of each sternite black, posterior margin of each segment pale yellow. Male genitalia: Epandrium (Fig. 19) nearly as long as wide, basal and distal margins each with triangular medial invagination. Subepandrial sclerite with small medial invagination; nearly as long as cercus distally. Hypandrium short. Gonocoxite (Fig. 20) tapering posteriorly into short outer gonocoxal process. Gonostylus much elongated, nearly 8 times longer than wide. Aedeagus (Fig. 21) with dorsal apodeme narrow; lateral ejaculatory apodeme narrow; distiphallus (Fig. 22) ventrally curved. Female. Unknown. Type material. Holotype male, CHINA: Gansu, Sunan, Lama Plain (38 ° 50 'N, 99 ° 37 'E, 2158 m), 6. VII. 2011, Xiao Zhang. Paratypes, CHINA: 1 3, Ningxia, Helan Mountain (38 ° 34 'N, 106 ° 20 'E), 21. VII. 1980, Chi-Kun Yang; 1 3, Gansu, Sunan, Lama Plain (38 ° 50 'N, 99 ° 37 'E, 2158 m), 6. VII. 2011, Ya-Jun Zhu. Distribution. Palaearctic: China (Gansu, Ningxia) (Fig. 35). This is biogeographically part of the Mongolia- Xinjiang and North China Regions (Zhang 1999). Remarks. This new species is very similar to A. laticornis (Loew, 1856), especially in the external habitus, the lateral view of the gonocoxite and even the aedeagus. But it can be separated from it by the following features: epandrium as long as wide, parallel-sided and markedly emarginated distally and basally; subepandrial sclerite as long as cercus; lateral ejaculatory apodeme narrow, not extending beyond margin of dorsal apodeme. In A. laticornis, the epandrium (Gaimari & Irwin 2000: 221, fig. 56) is 1.3 times longer than wide, tapering distally and not emarginated basally; the subepandrial sclerite is distinctly longer than the cercus; and the lateral ejaculatory apodeme (Lyneborg 1984: 214, fig. 29) extends laterally beyond the edge of the dorsal apodeme. Etymology. The specific name refers to the entirely yellow legs, from the Latin adjective “flavus” meaning yellow and the Latin derivation “femorata” referring to the legs.

Liu, Si-Pei, Gaimari, Stephen D., Yang, Ding (2012): Species of Ammothereva Lyneborg, 1984 (Diptera: Therevidae: Therevinae: Cyclotelini) from China. Zootaxa 3566: 1-13, DOI: 10.5281/zenodo.213838