Annales de la Société entomologique de France (N.S.) International Journal of Entomology

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Revision of the genera Nemotha Wood-Mason, 1884 and Tricondylomimus Chopard, 1930 stat. rev., with description of a new (: Mantodea)

Martin B. Stiewe & Evgeny Shcherbakov

To cite this article: Martin B. Stiewe & Evgeny Shcherbakov (2017) Revision of the genera Nemotha Wood-Mason, 1884 and Tricondylomimus Chopard, 1930 stat. rev., with description of a new species (Dictyoptera: Mantodea), Annales de la Société entomologique de France (N.S.), 53:3, 175-196, DOI: 10.1080/00379271.2017.1327330 To link to this article: http://dx.doi.org/10.1080/00379271.2017.1327330

Published online: 07 Jun 2017.

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Download by: [Evgeny Shcherbakov] Date: 07 June 2017, At: 09:07 Annales de la Société entomologique de France (N.S.), 2017 Vol. 53, No. 3, 175–196, https://doi.org/10.1080/00379271.2017.1327330

Revision of the genera Nemotha Wood-Mason, 1884 and Tricondylomimus Chopard, 1930 stat. rev., with description of a new species (Dictyoptera: Mantodea) Martin B. Stiewea* & Evgeny Shcherbakovb aThe Natural History Museum, London, Department of Entomology, Cromwell Road, SW7 5BD, London, UK; bFaculty of Biology, Lomonosov Moscow State University, Moscow 119991, Russia (Accepté le 26 avril 2017)

Summary. The oriental species previously included in the rarely collected Nemotha Wood-Mason, 1884 (Mantodea: ) are revised. N. coomani (Chopard, 1930) and N. mirabilis Beier, 1933 strongly differ from the type species N. metallica (Westwood, 1845) by family-level morphological features, but share a unique type of forefemoral armament with the genera Hapalopeza Stål, 1877 and Amantis Giglio-Tos, 1915, suggesting Nemotha is of a polyphyletic nature. The genus Tricondylomimus Chopard, 1930 is resurrected from synonymy to accommodate T. coomani, T. mirabilis n. comb. and the species which is newly described in this article, T. intermedius n. sp.The monotypic genus Pseudogousa Tinkham, 1937 and its species P. s inens is Tinkham, 1937 are synonymized with Tricondylomimus and T. mirabilis, respectively. The genus Tricondylomimus is placed in Iridopterygidae, while Nemotha is transferred to : Anaxarchini.

Résumé. Révision des genres Nemotha Wood-Mason, 1884 et Tricondylomimus Chopard, 1930 stat. rev., avec description d’une nouvelle espèce (Dictyoptera : Mantodea). Les espèces orientales précédemment incluses dans le genre rarement collecté Nemotha Wood-Mason, 1884 (Mantodea : Iridopterygidae) sont révisées. N. coomani (Chopard, 1930) et N. mirabilis Beier, 1933 diffèrent fortement de l’espèces type N. metallica (Westwood, 1845) par des caractères morphologiques du niveau familial, mais partagent avec les genres Hapalopeza Stål, 1877 et Amantis Giglio-Tos, 1915 un ’

type unique de l armature des fémurs antérieurs, suggérant la nature polyphylétique de Nemotha dans sa compréhension actuelle. Le genre Tricondylomimus Chopard, 1930 est ressuscité de la synonymie pour inclure T. coomani, T. mirabilis n. comb. et une espèce nouvellement décrite, T. intermedius n. sp. Le genre monotypique Pseudogousa Tinkham, 1937 et son espèce P. sinensis Tinkham, 1937 sont synonymes respectivement de Tricondylomimus et de T. mirabilis. Le genre Tricondylomimus est placé dans les Iridopterygidae, tandis que Nemotha est transféré aux Hymenopodidae : Anaxarchini. http://zoobank.org/urn:lsid:zoobank.org:pub:EA2C7A1A-DB81-4C9F-959A-4F629736D1BE Keywords: Hymenopodidae; Iridopterygidae; ; revision; new species; oriental region

History of the genera Nemotha, Tricondylomimus and The genus Nemotha Wood-Mason, 1884 is one of the least Pseudogousa known Oriental genera of praying (Mantodea). It currently includes three species known almost exclusively Westwood (1845) described metallica based on a by their type specimens: Nemotha metallica (Westwood, single female from Sylhet, modern Bangladesh. He pro- 1845) known from India, Bangladesh, Myanmar and vided a drawing and a description of the coloration. China, N. mirabilis Beier, 1933 from China and N. coo- Saussure (1871)includedM. metallica in his genus mani (Chopard, 1930) from Vietnam. At present, the Micromantis Saussure, 1870. He divided the genus into genus is classified in the family Iridopterygidae three groups or subgenera: the nominative subgenus with (Ehrmann 2002). However, recent advances in Mantodea the type species, M. glauca Saussure, 1870, and M. planiceps (Yager & Svenson 2008; Svenson & (De Haan, 1842), the subgenus Odontomantis Saussure, Whiting 2009) show this family to be polyphyletic, leav- 1871 with its type species, M.(O.) javana Saussure, 1871, ing uncertainty about the correct place of the genus. The and the third group, unnamed subgenus, with M. metallica. principal aim of this work is to revise the genera Nemotha Today, M. planiceps and M. javana are considered synonyms and Tricondylomimus stat. rev., bring clarity to its taxo- (Ehrmann 2002; Otte & Spearman 2005) and placed in the nomic status and volume, describe a new species and elevated genus Odontomantis of the family Hymenopodidae discuss the taxonomic position of all species in the system (tribe Anaxarchini), while the genus Micromantis,withthe of Mantodea. species M. glauca, belongs to Iridopterygidae.

*Corresponding author. Email: [email protected]

© 2017 Société entomologique de France 176 M.B. Stiewe & E. Shcherbakov

Wood-Mason (1884) created a new subgenus, and the second to Iridopterygidae, just like Tinkham Nemotha, for Mantis metallica and provided a description (1937a, 1937b). Wang did not examine the specimens. of the second female collected on Nemotha Peak, Assam. Mukherjee et al. (1995) provided additional details on In his opinion, the species is very closely related to the morphology of N. metallica, based on the specimens Odontomantis javana and Antissa pulchra (Olivier, collected earlier in Arunachal Pradesh, while treating the 1792) (today Odontomantis pulchra: Otte & Spearman species as a member of Hymenopodidae. 2005). Ehrmann & Roy (in Ehrmann 2002) transferred Giglio-Tos (1919, 1927) placed Nemotha in his tribe Pseudogousa from Iridopterygidae to and Euantissae (a new name for Antissae), of the subfamily left Nemotha (with Tricondylomimus as a synonym) in Hymenopodinae, together with Odontomantis and Iridoptergygidae: Iridopterygini. Euantissa Giglio-Tos, 1927 (= Odontomantis). Otte & Spearman (2005) used the same arrangement. Chopard (1930) described the genus Tricondylomimus Yager & Svenson (2008) used a female of Nemotha Chopard, 1930 and species T. coomani based on a male and a male of Pseudogousa as part of their study on the and female from Tonkin, now North Vietnam. He sug- mantodean cyclopean ear morphology and evolution, and gested the placement of this new genus in his tribe coded them as having DO and MSMT types, correspond- Metallytici of the subfamily Eremiaphilinae, though he ingly. Unfortunately, they did not indicate the species. noted considerable differences from the only genus of Thinh (2010) reported a male of N. coomani from Kon this tribe, Westwood, 1835. Tum, Vietnam. Beier (1933) described a second species of the genus Zhu et al. (2012) provided new data for P. sinensis: its Nemotha, N. mirabilis, based on a single female from distribution in China, photos of the male, female and Lung Tao Shan, northern Guangdong, China. He assigned ootheca, as well as life history and breeding data. They this species to Nemotha with some doubt, as he knew N. followed Ehrmann (2002) in arranging Pseudogousa metallica by Westwood’s picture and descriptions only. under Liturgusidae and Nemotha (with N. coomani and Based on his generic placement of N. mirabilis and mor- N. mirabilis listed) under Iridopterygidae. No specimens phology of the latter, he transferred the genus from of the latter two species were examined. Hymenopodinae to Iridopteryginae. Mukherjee et al. (2014) considered N. metallica in Beier (1935) synonymized Tricondylomimus with Iridopterygidae in a new checklist of the mantid fauna of Nemotha, without any comments, and listed N. coomani India. as a valid species. Most recently, Wu & Liu (2017) reported N. metallica Tinkham (1937a) described the genus Pseudogousa and from China for the first time. species P. sinensis on the basis of two adult males and a male nymph from Hainan and Kwangtung, respectively. He also listed an additional female of T. coomani from Tonkin, Material and methods close to the border of China, treating the genus Abbreviations Tricondylomimus as valid (probably being unaware of the BMNH: British Museum of Natural History, London, UK. ’ synonymization). He placed both genera in Iridopteryginae MNHN: Muséum national d Histoire naturelle, Paris, France. OXUM: Oxford University Museum of Natural History, because, in his opinion, their pronotal and leg features point Oxford, UK. to relationships with Spilomantis Giglio-Tos, 1915 and ZMBH: Museum für Naturkunde der Humboldt-Universität, Muscimantis Henry, 1931. He also noted a close morpho- Berlin, Germany. logical similarity between Pseudogousa and Liturgusa,but ZIN: The Museum of the Zoological Institute of the Russian felt that the arrangement of the discoidal spines in the Academy of Sciences, Saint-Petersburg, Russia. ZSI: Zoological Survey of India, Kolkata, India. former precludes the membership in Liturgusinae. Tinkham (1937b) mentioned the occurrence of N. mir- The examined material used in this study is deposited in the abilis in China and moved the genus Nemotha to the tribe collections of BMNH, MNHN, ZMBH and ZIN. Odontomantes, of the subfamily . He never External male genitalia were removed and placed in a hot saw any specimens of N. mirabilis. 10% KOH solution for approximately 30 min and then placed in Beier (1964, 1968) consistently placed Nemotha in water for approximately 90 min. After the external genitalia were dissected in parts, all were placed in 96% ethanol solution for Iridopteryginae, but never mentioned Pseudogousa in 1 h. The parts were then embedded in Euparal on a glass slide for any of his works. preservation. Mukherjee & Hazra (1985) reported new specimens of Pinned specimens and male genitalia were studied using N. metallica, including the first male, from Arunachal MBS-10 (Lomo-Eltem) and Leica MZ6 stereomicroscopes. Pradesh, India, placing the species in Hymenopodidae: Drawings were based on the photographs made with a Canon EOS 1000D camera, connected to the MBS-10 stereo- Acromantinae: Acromantini. microscope through an MFU adapter and with a Nikon D90 Wang (1993)listedN. mirabilis and P. sinensis as camera body, mounted to a Carl Zeiss Jena Technival 2 occurring in China, allocating the first to Hymenopodidae stereomicroscope. Annales de la Société entomologique de France (N.S.) 177

The terminology for the general morphology follows Giglio-Tos, 1915 and Amantis Giglio-Tos, 1915. The Wieland (2013), for the wing nomenclature it follows Béthoux only difference is that in both subgenera of Hapalopeza and Wieland (2009) and for the male genitalia it follows Klass the second discoidal spine is noticeably smaller than the (1997). fi All measurements are based on the specimens studied by the third, and in Hapalopeza (Hapalopeza) the rst discoidal authors, as well as the measurements reported in literature. spine is reduced to a tubercle. In all three taxa the largest The map was created in SimpleMappr (Shorthouse 2010). anteroventral spine is the second one, while the first is very minute, just like in N. mirabilis (Figure 28, i1). Interestingly, Amantis has always been classified in Results Gonypetini (: Amelinae), but recently molecular Upon detailed investigation of the morphology of all phylogenetics (Svenson & Whiting 2009) has shown the species included in Nemotha, the genus appears to be polyphyly of this group, with Amantis forming a clade very heterogenic. In particular, the type species, N. metal- with Hapalopeza (Nemotha was not sampled). Polyphyly lica, differs most significantly from the other two species, of both Iridopterygidae and Iridopteryginae has also been N. coomani and N. mirabilis, by features at the family- demonstrated but the groups are yet to be revised. It seems level. Posteroventral foretibial spines of N. metallica are reasonable to retain N. coomani and N. mirabilis in decumbent (Figure 24), while in N. coomani and N. mir- Iridopteryginae – at least until a proper revision of the abilis they are inclined at no less than 45° (Figure 25). subfamily has taken place. Since N. metallica is the type Among the four discoidal spines of N. metallica the third species of the genus Nemotha, the name Tricondylomimus spine is the longest, and the first the shortest (Figure 26). must be restored from its synonyms. On the forewing, the main veins are elongated and very Our study of the holotype of Nemotha mirabilis and gently curved around the middle of the forewing, while in our discovery of a new species related to N. mirabilis and its distal half the cross-veins are aligned parallel to the T. coomani has allowed the picture to be clarified even forewing and form dense elongated cells, much like in further. Firstly, the new species described below represents Anaxarcha Stål, 1877 and Werneriana Shcherbakov, a morphological link between the other two and demon- Ehrmann & Borer, 2016 (Shcherbakov et al. 2016). N. strates that they, in fact, belong to one genus. Secondly, metallica is also different from its supposed congenerics our study of high resolution photographs, together with by having two ventral cervical sclerites. None of the the original description of Pseudogousa sinensis, revealed described features of N. metallica correspond to the diag- the latter to be conspecific with Nemotha mirabilis, which nosis of Iridopteryginae, but instead are very similar to is why it consequently has to be treated as a new Odontomantis and must be transferred to Hymenopodidae: synonym. Anaxarchini. At the same time, N. coomani and N. mirabilis have a very special type of forefemoral armament, described by Family Hymenopodidae Giglio-Tos, 1915 the following features (Figures 28 and 29). The first and Subfamily Hymenopodinae Giglio-Tos, 1915 second discoidal spines are shifted laterally so that the first Tribe Anaxarchini Giglio-Tos, 1919 one falls in line with the anteroventral row and the second Genus Nemotha Wood-Mason, 1884 is only slightly more basal than the third discoidal spine Nemotha Wood-Mason 1884: 35; Giglio-Tos 1919: 73; Giglio- (Figure 29, d1–d2). The second and third discoidal spines Tos 1927: 543; Beier 1935: 45; Beier 1964: 947; Beier 1968: are large and almost of the same size (Figure 27, d2–d3). 10; Wang 1993: 50; Mukherjee et al. 1995: 224; Ehrmann – The anteroventral spine is immediately opposite the third 2002: 238 239, 376; Otte & Spearman 2005: 113; Yager & fi Svenson 2008: 565; Thinh 2010: 20; Zhu et al. 2012: 30; discoidal spine (the rst in row in N. coomani and the Mukherjee et al. 2014: 20; Wu & Liu 2017. second in N. mirabilis), is just as large and is much larger than the other anteroventral spines (Figures 27, i1 and 28, Type species. Mantis metallica Westwood, 1845 (by i2). The smallest discoidal spine is the fourth one monotypy). (Figure 29, d4). The first spine is much smaller than the second and third (Figures 27 and 29; d1). Between the two Diagnosis. Closely related to Odontomantis, but differs in distal-most large spines of the anteroventral series there is the following features: (1) metazone noticeably narrowed a large gap with 1–2 very small spines (Figures 27 and in the middle; in Odontomantis, the metazone is not, or 28). The first posteroventral spine is slightly shifted medi- only slightly, medially narrowed; (2) abdomen, forelegs ally (Figure 29). Finally, the whole ventral surface of the and hind legs completely metallic black; in Odontomantis, forefemur between the anterior and posterior rows of adults with completely green abdomen, forelegs and hind spines is covered by multiple tubercles (Figure 29, tbr). legs – only early instar nymphs are showing a shiny We found that only three other mantodean taxa have a coloration; (3) female foretibia narrow, longer than nearly identical forefemoral armament: Hapalopeza foretarsi; in Odontomantis, the female foretibia is shorter (Hapalopeza) Stål, 1877, Hapalopeza (Spilomantis) or as long as the foretarsi; (4) forewings yellowish, hind 178 M.B. Stiewe & E. Shcherbakov wings yellowish in costal area and dark brownish in forming an angle of about 140° and connected anteriorly discoidal area, rest of the body black with blueish by a feeble elevation with respect to the area between the metallic shine; in Odontomantis the main body color is antennae. Median ocellus slightly raised posteriorly above green, the forewings are always green, while the hind the surrounding area, and partially covered by the cuticle wings are reddish in costal and discoidal areas, distally flush with surrounding surface, lateral ocelli much less so. infumated. Vertex between ocelli and frontal suture feebly rugose. Frontal suture raised medially into a very low carina. Parietal sutures well developed, moderately deep. Juxta- Nemotha metallica (Westwood 1845) ocular bulges very prominent, protruding beyond com- (Figures 1, 6, 7, 11, 16, 24, 26, 33, and 34) pound eyes and median part of the head, making whole posterior line of the head weakly U-shaped. Area between parietal sutures posteriorly with a pair of very feeble, wide Mantis metallica Westwood 1845:51–52, pl. 62, fig. 3. Micromantis metallica: Saussure 1871: 33. paramedian longitudinal depressions, and a short, shallow Nemotha metallica: Wood-Mason 1884:34–37; Giglio-Tos median groove. Antennae filiform, reaching at least 1927: 543; Beier 1935: 45; Mukherjee & Hazra 1985: 38; beyond posterior edge of the pronotum (in all known Mukherjee et al. 1995: 224–225; Ehrmann 2002: 239; Otte & preserved specimens the antennae partially or completely Spearman 2005: 114; Mukherjee et al. 2014: 20; Wu & Liu broken). Scapus cylindrical, of almost uniform width. 2017: 18, figs 7, 21–23. Pedicellus with median constriction, of approximately Type material. Holotype: ♀, Silhet (today Sylhet in the same length and 2 times as narrow as the scapus. Bangladesh) (OXUM) (Figure 6). The type is in Remaining antennomeres cylindrical, basally very short fi satisfactory condition: the antennae (except scapi and and signi cantly narrower than the pedicellus. pedicelli), distal tarsomeres of foretarsi, right middle leg Pronotum (Figure 16) relatively short, robust, roughly from the coxa, most of the hind wings and the left cercus semi-oval in cross section. Prozone parabolic-trapezoidal, are missing; some parts of the hind wings are glued to the with lateral edges possessing a small angular ledge on each paper rectangle; most of the prosternum and parts of the side at about half of their lengths. Prozone dorsally bulging forecoxae, trochanters and femora are covered with glue. and slightly compressed laterally in its anterior most part; a pair of very low, smooth carinae runs diagonally across its

Other material. 1 ♀, Upper Burma, Nam Tamai Valley, lateral sides, starting dorsally near the middle and diverging 12.IX.1938, leg. R. Kaulback, B.M.1938-741 (BMNH) anteriorly; area between each of these carinae and trans- (Figure 1). 1 ♂, India, Arunachal Pradesh, Tirap, verse groove covered by rough wrinkles. Transverse groove Hornbill, 30 km. NE of Miao, 13.XII.1983, coll. S. very shallow medially, noticeably deepening laterad, Biswas (ZSI) (Figure 7a, b) slightly angular. Supracoxal dilatation short, oval, approxi- mately the same width as the head and 1.3–1.4 times as wide as the prozone. Metazone 1.6–1.7 times as long as the Redescription prozone, more or less strongly constricted in the middle, Female. Figures 1, 6, 11, 16, 24, 26, and 33. Medium with semi-oval posterior edge; its maximum posterior width sized (body length 26.5–28 mm). only slightly less than the maximum width of supracoxal Head (Figure 11) pentagonal, slightly flattened dorso- extension. Dorsal profile of metazone slightly bulged at the ventrally, 1.4 times as wide as long. Labrum pentagonal, point of maximum width. Dorsal surface of metazone at medially with a strong bulge. Clypeus trapezoidal, its this point with a pair of wide depressions preceding the dorsal margin slightly convex. Anterior surface with a bulge, otherwise with thin median carina and a pair of pair of lateral tubercles placed at some distance from roughly oval low paramedian elevations right before poster- dorsal angles of clypeus; medially with very strongly ior edge. Angle between posterior parts of pronotum and and sharply raised, obtuse-angular transverse carina reach- prosternum about 50°. Lateral edges slightly lamellar, with ing lateral sides of clypeus, with feeble median longitudi- small rounded tubercles. Cervicalia with two ventral cervi- nal carina; posteriorly more or less flat, with median cal sclerites. Intercervicalia very elongated, triangular, triangular groove. Scutellum pentagonal, transverse, its shorter than lateral cervical sclerites. Postcervical plate width 2.6 times as large as its length at the middle, ventral extremely short, as is T-shaped sclerite. Furcasternum with margin concave, dorsal margin obtuse-angular, raised, low medial carina. with very small dorsal projection at the apex. Anterior Forelegs typical of praying mantises. Coxa triangular surface ventrally with a pair of paramedian transversely in cross-section, longer than metazone of pronotum, nearly elongated tubercles mirroring those on clypeus, otherwise smooth. Anterior apical lobes short, divergent. Trochanter with a few very thin transverse creases curved anteriorly at bucket-shaped, strongly narrowed distally. Femur the middle. Compound eyes large, oval, bulbous, with (Figure 26) elongated, roughly triangular in cross-section, very narrow semi-oval eye sockets. Ocelli small, oval, its length 4.5 times as large as maximum dorsoventral Annales de la Société entomologique de France (N.S.) 179

Figures 1–5. Genera Nemotha and Tricondylomimus, habitus. 1, N. metallica, ♀; 2, T. coomani, ♂; 3, T. coomani, ♀; 4, T. mirabilis, ♀ holotype; 5, T. intermedius n. sp., ♀ holotype. width. Dorsal edge very slightly lamellar, smooth, basal discoidal, 4 posteroventral, 1 anteroventral genicular and 1 half a little convex. Anterior surface with shallow claw posteroventral genicular spines. Anteroventral spines elon- groove situated at basal 2/5 of the length and with oval gated, sharp, curved distally at the base, of alternating femoral brush. The femur armed with 11 anteroventral, 4 length, with a gap between the two distal-most spines. 180 M.B. Stiewe & E. Shcherbakov

Figure 6. Nemotha metallica, ♀ holotype and its labels. © Oxford University Museum of Natural History. All rights reserved. Published with permission.

First discoidal spine the smallest, straight, sharp, smaller spine, and a larger but even more shallow depression than small anteroventral spines, shifted laterally and between second and third posteroventral spine. Both geni- almost in line with them. Second discoidal spine 2.5 cular spines of similar structure, shorter than smaller ante- times as long as the first. Third discoidal spine the long- roventral spines, very robust and strongly curved distally. est, 2.6 times as long as the second and much thicker at Tibia (Figure 24) thin, straight, 1.4 times as short as the base, straight, sharp. Fourth discoidal spine a little femur, armed with 11–13 anteroventral spines (without shorter than the second, strongly curved distal at the tibial claw) and 11 posteroventral spines. Anteroventral base. Posteroventral spines thin and elongated, the distal spines equally spaced, gradually elongating distally, three placed at equal distances from each other, while the inclined at approximately 45°, slightly curved. Tibial first one is much closer to the second. First posteroventral claw very long and sharp, overall inclined at 70° with spine the smallest, straight, slightly shifted medially, sec- respect to the tibia, slightly curved along its length. ond and third posteroventral spines identical, 1.3 times as Posteroventral spines extremely decumbent. Tarsi a little long as the first, slightly curved medially, fourth poster- longer than the tibia. Basitarsus slightly longer than the oventral spine a little shorter and thicker than preceding remaining tarsomeres combined. two. Posteroventral edge with a small circular shallow pit- Metathorax with a cyclopean ear of the DK type, with like depression between first and second posteroventral comparatively large opening. Annales de la Société entomologique de France (N.S.) 181

Figure 7. Nemotha metallica, ♂. A, dorsal and B, ventral view, from Arunachal Pradesh, India. © T. K. Mukherjee, published with permission. C, dorsal view, from Yunnan, China. D, genitalia ventral view, from Yunnan, China; C and D © Chao Wu, published with permission.

Middle and hind legs cursorial. Coxae short, triangu- plate semi-oval, cerci formed by at least 10 cercomeres, lar, mesocoxa with a very low ventral carina. Femora and reaching beyond the apex of subgenital plate and ovipositor; tibia straight. Mesofemora shorter than metafemora, cercomeres similar to each other, circular in cross-section, smooth, both with a short apical spine. Tibia equal to basally wider than long, very slightly elongating distally. femora in length but thinner, straight, apically with dorsal Coloration. Main body color, including wide margin triangular extension and a ventral spine. Tarsi shorter than along the edges of the pronotum (except small area between tibia, basitarsus shorter than the remaining tarsomeres disconnection of pronotum and prosternum and posterior combined. edge) as well as wide bat-shaped band along transverse Wings. Both pairs fully developed, extending beyond supracoxal groove shiny black with metallic blue reflections the end of the abdomen by about 1/6 of their length. (Figure 33). Most of the prozone and metazone of the pro- Forewings 3.7–4 times as long as wide, elongated, oval. notum are yellow, except the posterior of the head, the ocelli Main veins gently curved around the middle. Costal field and the areas between the parietal sutures. Forefemur poster- widest at the end of basal third, gradually narrowing and iorly with a medially elongated pale patch. Forewings yel- not reaching the apex. RA usually simple, RP+M with two low, precostal field black. Hind wings opaque red basally and to four branches. CuA forked basally, anterior branch with anteriorly, the apex and an irregular spot in the basal third two to three, posterior branch with four simple branches; yellowish, the rest heavily smoky and lighter cross-veins; all all branches of CuA gently curved medially anteriorly, the articular membranes, two streaks on each eye, the middle anterior ones more than the others. Pseudoveins absent. half of fore basitarsus, thin dorsal marks on mesotarsus, The cross-veins in the basal half forms irregularly shaped small dorsal apical mark on the four posterior femora, a cells usually situated in three rows, while in the distal half }-shaped band along the posterior edge of sternites II–VI they align almost longitudinally and form dense elongated and the posterior half of subgenital plate pale turquoise-blue cells. Stigma highly elongated along RP + M, indistinct. in live (Figure 33) and yellowish in dried specimens. Hind wings shorter than forewings, triangular with semi- oval apex. Male. Figures 7a–dand34. Very similar to female, but Abdomen oval, about 1.5 times as long as wide. All significantly smaller (body length 18 mm). Compared to tergites and sternites transverse, with fifth tergite and third female, its pronotum is more rhomboidal, the metazonal con- visible sternite being the widest. Posterior edges of sternite I striction weaker and the bat-shaped transverse band on pro- convex, sternites II–IV straight, sternite V concave. Anal notum wider. Posteriorly, the forefemur has a much bigger 182 M.B. Stiewe & E. Shcherbakov elongated pale patch and the forewings are brownish-yellow, Type species. Tricondylomimus coomani Chopard, 1930 hind wings uniformly infumate, much lighter than in female (by monotypy). and dorsally with reddish coloration in the discoidal field (Figure 7c). Diagnosis. Similar to Hapalopeza and Amantis,withthe In the male genitalia (Figure 7d), sclerite L4A elongated, following differences: (1) scutellum transverse, more than 3 noticeably sclerotized on the anterior half; covered with dots times as wide as long; in Hapalopeza and most species of with setae and without a distal process pda. Heavily scler- Amantis the scutellum is less transverse, about twice as otized round apex. Sclerite L4B of complex structure, with wide as long; (2) first discoidal spine longer than fourth the anterior half very strongly sclerotized. Sclerite L2 well discoidal spine; in Hapalopeza and Amantis the first developed and trilobate, with a short, strongly sclerotized discoidal spine is shorter than both the fourth discoidal spoon-like apical process paa. Sclerite L1 with a large and spine and the larger forefemoral spines of the strongly sclerotized, globular phalloid apophysis afa. anteroventral row, sometimes it is even reduced to a Sclerite R1 rectangular, round at the apex; sclerotized tubercle; (3) wings moderately reduced in females, with ventral plate pia, smooth ventral tooth pva, strongly forewings at rest reaching any point from the posterior curved by 180°, same length as ventral plate pia. Sclerite edge of tergite IV to the base of the anal plate; in fi R3 very elongated, nger-like. Hapalopeza the wings in females reach beyond the apex of the abdomen, while in Amantis they may be moderately Measurements (in mm). Body length, male 18.0, female to almost completely reduced; (4) at least tergites and – 26.5 28.0; head width, female 3.8; pronotum length, male sternites with blue metallic shine; some species of – 6.5, female 7.3 8; maximum pronotum width, female 4.5; Hapalopeza are shiny, but without a true metallic shine; (5) forecoxa length, male 5.0, female 6.0; forefemur length, while females of Hapalopeza and Amantis have a male 6.0, female 7.5; hind femur length, female 7.5; metathoracical ear of the DK type (Yager & Svenson – forewing length, male 14.0, female 19.5 22. 2008), females of Tricondylomimus have an ear of the POT type (D. Yager, pers. comm. unpublished) (Figure 23), an Remarks. This species occurs in a variety of geographical extreme deviation from DK type with walls tilted posteriorly locations (Figure 36); however, the different localities do not like in the MSMT type and a deep, wide-open, triangular pit with hardly developed knobs, like in the DO type. have a high density of specimens. So far, only 9 specimens are known worldwide. Of these 9 known specimens, 4 were just recently discovered in the province Yunnan in China (Gaolinggong Mt., 1400 m, leg. Chao Wu, 15.VII–10. Key to the species of Tricondylomimus IX.2015, 3 ♂,1♀)(Figures 33 and 34). This is the first 1. Metazone of pronotum strongly bulging posteriorly the record of N. metallica from China (Wu & Liu 2017). Despite supracoxal dilatation and sharply angled up near pos- the fact that many of the recorded localities are now terior edge of the pronotum, while concave at its mid- frequently visited by scientific researchers, nothing is dle (Figures 19 and 20). Network of cross-veins in the known about the behavior and life cycle of N. metallica. distal half of the forewing highly irregular (Figure 32). The limited number of specimens contributing to this species Shiny brown (Figures 2, 3, 8, and 9)...... have been taxonomically misplaced in the past...... Tricondylomimus coomani – Metazone of pronotum flat or only slightly Ƨ-shaped, fl Distribution. Bangladesh: Sylhet. India: Assam, its posterior part at (Figures 17 and 18)...... 2 Arunachal Pradesh. Myanmar: Tamai Valley. China: 2. CuA with three, M with two branches in the forew- Yunnan (Figure 36). ing. Cross-veins in the distal half of the forewing form mostly regular, rectangular cells (Figure 31). Olive-brown, with reddish, monochrome middle Family Iridopterygidae Giglio-Tos, 1915 and hind femora; forewings with monochrome Subfamily Iridopteryginae Giglio-Tos, 1915 veins and without bright spots (Figure 5)...... Tribe Iridopterygini Giglio-Tos, 1915 ...... T. intermedius n. sp. Genus Tricondylomimus Chopard, 1930 stat. rev. – CuA with two branches, M simple for most of its length in the forewing. Network of cross-veins in the Tricondylomimus Chopard 1930: 229; Naviaux 2002: 19. distal half of the forewing irregular (Figure 30). Olive- Syn. Pseudogousa Tinkham 1937a: 491, n. syn. Wang 1993: 97; green, with bright-green spots on the body and forew- Ehrmann 2002: 294, 375; Otte & Spearman, 2005: 135; ings, and whitish stripes on the legs, including middle Yager & Svenson 2008: 565; Zhu et al. 2012: 30, 135. and hind femora (Figure 4)...... T. mirabilis Annales de la Société entomologique de France (N.S.) 183

Tricondylomimus coomani Chopard, 1930 Other material. 1 ♀, Tonkin occ., Reg. de Hoa Binh, 1919 (Figures 2, 3, 8, 9, 12, 13, 19, 20, 22, and 32) (MNHN); 1 ♂, Tonkin, Reg de Hoa-Binh, A de Cooman 1926 (MNHN); 2 ♂, Tonkin, Rég de Hoa-Binh, De Cooman 1927 Tricondylomimus coomani Chopard 1930: 229–232, Pl. 3, figs. (genital slide: R. Roy 4601, MNHN (EP) 2800), (MNHN (EP) 3–4; Tinkham 1937a: 489–490, pl. 16, fig. 5. 2798); 3 ♀, Tonkin, Rég de Hoa-Binh, A. De Cooman 1926 Nemotha coomani: Beier 1935: 45; Ehrmann 2002: 239; Otte & (MNHN (EP) 2802, 2805, 2806); 1 ♀, Vietnam, NE Region, Spearman 2005: 114; Thinh 2010: 20; Zhu et al. 2012: 30. Vin Phuc Prov., Me Linh Distr., Station for Biodiversity, – Type material. The species was described by Chopard 24 27.VIII.2014, leg. T. McCabe (Cleveland). based on one male (the holotype) and one female (the allotype), collected by A. de Cooman in Tonkin, Hoa-Binh Redescription (present day Vietnam, Hoa Binh province, Hoa Binh). Female. Figures 3, 9, 12, 20, and 32. Small (body length Holotype: ♂ (Figure 8), “Tonkin, Lac Toh, Hoa-Binh, 23 mm). A de Cooman” (MNHN (EP) 2796); allotype: ♀ Head (Figure 12) large, pentagonal. Labrum pentago- (Figure 9), “Tonkin, rég. de Hoa-Binh, A de Cooman nal, medially bulging, dorsally fused with clypeus. 1926” (MNHN (EP) 2804). The specimens are in satisfac- Clypeus massive, with bulging transverse carina, the tory condition, the male lacks both hind legs and most of latter gradually lowering ventrally, with hyperbolic pro- the antennae, the female lacks antennae and distal parts of file. Anterior wall of the transverse carina with a thin the left fore and right hind tarsi. Both specimens are longitudinal carina running ventrally. Posterolateral provided with corresponding red labels (Figures 8 and 9). angles of clypeus flat, rounded. Scutellum transverse, of

Figures 8–10. Types of Tricodylomimus coomani and habitus of Tricondyla tiger beetle. 8, Tricondylomimus coomani, ♂ holotype and its labels, MNHN-EP-EP2796; 9, ♀ allotype and its labels, MNHN-EP-EP2804; 10, Tricondyla mellyi Chaudoir. Figures 8 and 9 © MNHN, M. Depraetere, published with permission, Figure 10 © E. Shcherbakov. 184 M.B. Stiewe & E. Shcherbakov

Figures 11–15. Genera Nemotha and Tricondylomimus, head. 11, N. metallica, ♀; 12, T. coomani, ♀; 13, T. coomani, ♂; 14, 15, T. intermedius n. sp., ♀. 11–14, anterior view; 15, dorsal view. nearly constant length across its width, except its ante- shorter than long, distally gradually elongating and nar- rolateral lobes very slightly projecting anteriorly. Width rowing; about 4 basal-most antennomeres of the flagel- of scutellum 3.7 times as large as its length at the middle. lum completely fused. Ventral margin between lateral lobes nearly straight. Pronotum about 2 times as long as wide, in cross Dorsal margin obtuse-angular, carinate in the middle, section nearly semicircular. Prozone semicircular, strongly smooth laterally. Surface of scutellum slightly rugose. bulging dorsally, with converging lateral depressions in its Compound eyes large, oval, bulbous, with narrow semi- anterior half. Transverse groove deep and very slightly oval eye sockets. Ocelli very small, nearly circular, lat- curved. Supracoxal dilatation short but well developed, eral ones form a slight angle to the surrounding cuticle. limited anteriorly by transverse groove, about 0.6 times Frontal suture slightly depressed, inconspicuous. The the width of the head. Metazone 2.0–2.3 times as long as posterior vertex looks like it has a suture that is bulging, prozone, with strongly Ƨ-shaped lateral profile, very bul- with posterior boundary very convex when viewed dor- ging in its anterior half, then strongly concave and rising sally, protruding far beyond the compound eyes; juxta- again near posterior edge. Dorsal surface of pronotum in ocular bulges moderate, more or less leveled with the the prozone, lateral parts of supracoxal dilatation, and vertex. Antennae filiform, almost as long as the body. posterior half of metazone, as well as along the medial Scapus cylindrical, with a slight medial depression on line densely covered by small tubercles, each possessing a one side. Pedicellus hourglass shaped, as long as and 1.5 single rather long seta. Additionally, posterior half of timesasnarrowasscapus.Remaining antennomeres supracoxal dilatation with a short, transverse, very shallow cylindrical, basally slightly narrower than pedicellus, and smooth groove on each side; metazone with a pair of Annales de la Société entomologique de France (N.S.) 185

Figure 22. Tricondylomimus coomani, ♂ genitalia in ventral view. afa = phalloid apophysis, paa = apical process, pda = distal process, pia = ventral plate, pva = ventral process.

armed with 8–9 very small, rounded tubercles decreasing in size distally, each accompanied by a long seta, with even smaller spinules in between. Anterior surface of the coxa

loosely covered with weak tubercles, each with a long seta, the latter becoming longer in the basal half of the coxa. Posterior edge with spines similar to the spines on the dorsal edge. Anterior apical anterior lobe short, wide, strongly diverging from the base; the dorsal one extended and visible from posterior as a small lobe. Trochanter only slightly narrower than coxa, sharply curved. Femur comparatively wide in lateral aspect; 3.7–3.9 times as long as maximum dorsoventral width. Dorsal edge wide and smooth, basally slightly convex, otherwise straight. Anterior surface with a very wide and deep claw groove, situated close to the distal end of the femur’s basal third, the femoral brush, formed by Figures 16–21. Genera Nemotha, Tricondylomimus and long thick setae, and several setae between the claw groove Tricondyla, pronotum in left lateral view. 16, N. metallica; 17, and the femoral brush. Femur armed with 10–11, rarely 9, Tricondylomimus mirabilis; 18, T. intermedius n. sp.; 19, T. anteroventral, 4 discoidal, 4 posteroventral, 1 anteroventral coomani, ♂; 20, T. coomani, ♀; 21, Tricondyla mellyi. Scales genicular and 1 posteroventral genicular spines. Ventral sur- 1 mm. face between basal 8 anteroventral spines and basal 3 poster- oventral spines covered by multiple small tubercles, and the strong pyramidal tubercles near its posterior edge. Lateral surface between the base and the second discoidal spine with edges narrowly lamellar, with very small rounded tuber- a double row of flattened tubercles. First anteroventral spine cles, smaller and sparser in the prozone and at supracoxal greatly enlarged, as long as the first posteroventral spine and dilatation, bigger and denser along most of the metazone. about 0.8 times as long as the second discoidal spine. Second Postcervical plate extremely short, similar to the T-shaped to ninth anteroventral spines of alternating sizes, usually sclerite, the latter with its posterior part very thin and starting with a small spine, so that the third spine is about partly reduced. Furcasternum anteriorly angular, raised one third of the length of the first one, and the second spine is into a carina. about two thirds of the third spine in length; both, the spines Forelegs overall typical for a praying mantis. Coxa rela- and the spaces between them, are slightly elongated distally. tively wide, longer than metazone, with smooth dorsal edge The shorter spines are slightly shifted medially comparing to 186 M.B. Stiewe & E. Shcherbakov larger spines. Gap between the ninth (long) and the distal- triangular, with very low ventral carina. Femora straight, most (long) spines is much larger than between the rest, wide, narrowing distally, with an apical spine. Tibia usually with a small tenth spine inside, situated closer to straight, roughly as long as the femora, thin, straight, the ninth. Distal-most spine half the length of the first ante- covered by short setae, apically with a dorsal triangular roventral spine, thinner, sharper and more curved than the dilatation and two ventral spines. Tarsi 0.8 times as long longer anteroventral spines. Anteroventral genicular spine as femora, hind basitarsus as long as remaining tarsome- very similar to the distal-most anteroventral spine. All poster- res combined. oventral spines emerging from short sockets. First poster- Wings shortened, of variable length among the exam- oventral spine straight, a little smaller than the second ined specimens, covering the first 6–8 tergites. Forewings discoidal spine, slightly shifted medial; second to fourth roughly rectangular, slightly widening distally, with para- spine approximately of equal length, becoming more curved, bolic apex. Costal field widest near base, gradually nar- thinner and sharper distally; space between the first and the rowing and nearly reaching the apex. All main veins second spines equals the length of the first spine, between the curved, except a nearly straight RA; Sc very irregular in third and the fourth spines slightly longer than that, and the basal half. CuA with three branches. Cross-veins very between the second and the third even more so. prominent, carinate (Figure 32); their network with very Posteroventral genicular spine similar to but slightly shorter narrow more or less rectangular cells in the anal area, than the fourth posteroventral spine. First discoidal spine otherwise generally irregular, especially in the costal very slightly curved, about two fifths of the length of second, field and in the distal half of the forewing, sometimes shifted on the same line as the anteroventral row, immedi- forming a honeycomb-like structure. Stigma situated on ately before claw groove; second discoidal spine very mas- basal-most fork of RP+M, extremely indistinct, noticeable sive, almost straight; third spine identical to the second; the only by its much smoother surface when compared with second and third discoidal spines are in line with the first the remaining forewing. Hind wings significantly shorter posteroventral spine; fourth discoidal spine the smallest, than forewings with rounded apex. shorter than smaller anteroventral spines, strongly inclined Abdomen ovoid, all tergites and sternites wider than distally. Tibia straight, expanding dorsoventrally toward its long, with fifth tergite and fourth visible sternite being the apex, armed with 10–12 anteroventral spines (without tibial widest. Anterior and posterior edges of tergites and ster- claw) and 6 posteroventral spines. Basal seven anteroventral nites straight. Supra-anal plate semicircular, with short spines almost straight, inclined, very sharp, first to fifth transverse fold in the middle, transitioning into a very spines gradually elongating, the sixth and seventh spines short medial longitudinal fold posteriorly. Cerci formed very slightly shorter than fifth; the four distal spines notice- by 11–12 cercomeres, gradually elongating distally, the ably thicker and less inclined than the previous ones. Tibial last cercomere more than twice as long as penultimate claw about 0.4 times as long as the rest of the tibia, strongly one, flattened laterally, other cercomeres circular in curved at the base at about 70° with respect to the tibia, very cross-section. thick, with a longitudinal, anterodorsal, very narrow groove Coloration. Main body color dark-brown with strong and obtuse apex. First posteroventral spine situated at about bronze metallic shine. Scutellum with blue or green 1.5 of its length from the base; second spine the longest, reflections. Compound eye sockets with narrow yellow situated close to the first; third spine shorter than the first, stripe along the edge. Posterior-most part of the vertex with a gap between it and the second spine which is slightly with a pair of yellow-brown spots. Palpi yellow with longer than the length of the second spine; fourth spine dark apices. Antennae yellow, except blackish scapus slightly smaller than the third, with the gap between the and pedicellus. Pro-, meso- and metasternum with blue third and the fourth spines smaller than the gap between or green shine. Anterodorsal part of forecoxae, anterior the second and the third spine; the fifth spine is the smallest, part of fore trochanters, and forefemora brown, with a situated as far from the fourth as the fourth from the third light-yellow stripe along the dorsal edge and largely spine; sixth spine almost like the second, placed very closely orange anterior surface, foretibia brown with various to the fifth. All posteroventral spines sharp, slightly curved, shades of dark yellow in some parts. Middle and hind inclined at about 45°. Tarsus substantially longer than the coxae black with blue reflections, femora reddish- femur, about the same length as the pronotum. Basitarsus brown, tibia with tarsi a little darker. Forewings very longer than the remaining tarsomeres combined. The distal smoky, all veins black; hind wings smoky at apex and part of the basitarsus and the remaining tarsomeres are cov- less so along the anterior edge, otherwise transparent ered by very dense setae. with brown veins. Abdomen with tergites and lateral Metathoracical cyclopean ear of POT type, with deep parts of sternites brown. triangle pit limited anteriorly by the basisternum wall, posteriorly by fused walls. Male. Figures 2, 8, 13, 19, and 22. Very similar to female, Middle and hind legs long, cursorial, with middle but smaller (body length 14.5 mm). Clypeus less bulging, legs shorter than hind legs. Coxae very short and wide, while the dorsal margin of the scutellum is more convex Annales de la Société entomologique de France (N.S.) 187

Figures 23–32. Genera Nemotha and Tricondylomimus, morphological details. 24, 26, N. metallica; 32, T. coomani; 28, 30, T. mirabilis; 23, 25, 27, 29, 31, T. intermedius n. sp. 23, metathoracic ear; 24, 25, foretibia in posterior view; 26–28, forefemora in anterior view (only anterior row of spines shown for T. mirabilis); 29, forefemur in ventral view; 30–32, posterodistal part of forewing venation. Scales for 24–29 1 mm; 23, 30–32 not to scale. i1–i2 = first to second anteroventral spines; d1–d4, first to fourth discoidal spines; tbr = field of tubercles. and the ventral margin more concave (Figure 13). Ocelli ocular bulges comparatively shorter. Pronotum (Figure 19) slightly larger. Frontal suture raised into a very low carina. covered by tubercles across the whole surface. Forelegs Posterior boundary of vertex only slightly arcuate, juxta- substantially more slender, all femoral and tibial spines 188 M.B. Stiewe & E. Shcherbakov thinner. Tarsomeres of meso- and metatarsi white, with Binglan, pers. comm.). However, the species has been black apices. Wings longer, reaching the end of abdomen. collected in China since its description, most recently Supra-anal plate more transverse, subgenital plate with during a field trip to Ledong in Hainan (C. Wu pers. trapezoidal posterior half and short styli. In the male comm., see Figure 35), and there have also been attempts genitalia (Figure 22), sclerite L4A strongly elongated, to breed it in captivity (Zhu et al. 2012). with noticeably sclerotized posterior half. Basal lobe short, with ragged posterior edge and membranous con- tinuation of its right edge; this membrane and large adja- Redescription cent dextro-posterior area of L4A covered by dots with Female. Figures 4, 17, 28, 30, and 35. Medium sized setae. Distal process pda short, robust, situated on and (body length 21–24 mm). directed to the left, contrary to the situation in most Head comparatively large, pentagonal, very slightly Mantodea. Sclerite L4B large, twisted along its axis ante- flattened dorsoventrally. Labrum pentagonal, mediodor- riorly and expanded posteriorly. Sclerite L2 elongated, sally fused with clypeus. Clypeus with a low transverse curved, with apical process paa strongly curved dorsally, carina, and an adjacent longitudinal, very low and thin rounded at apex; left edge of L2 immediately anterior to medial carina. Scutellum transverse, short, ventral margin the apical process folded inside. Sclerite L1 with small, straight except lateral lobes slightly protruding anteriorly, hatchet-like phalloid apophysis afa and long, weakly dorsal margin a little convex, medially elevated into a low curved anterior process wider than the phalloid apophysis. transverse carina. Width of scutellum 3 times its height at Sclerite R1 with three “arms”: the left arm long, rectan- the middle. Compound eyes large, globular, bulbous, with gular, twisted and blade-like at the apex; the right arm narrow semi-oval eye sockets. Ocelli very small, nearly protrudes anteriorly and folds ventrally, with small but circular. Medial ocellus almost flat, lateral ocelli slightly heavily sclerotized ventral plate pia and simple, smooth larger than the medial one and forming a slight angle to ventral tooth pva, strongly curved by 180°, with ventral the surrounding cuticle. Vertex almost flat, with only very lobe of the latter longer than its dorsal lobe; the posterior low elevations near each ocellus, and fontal suture raised arm of R1 parabolic, finger-like. Sclerite R3 spoon-like, medially into a very low carina. Posterior edge of the head with rectangular and flat expanded part. weakly arcuate, protruding a little beyond the compound eyes. Parietal sutures shallow, with a pair of short, even

Measurements (in mm). Body length, male 14.5, female more shallow grooves medial of them; juxta-ocular bulges 23; head width, male 3.6, female 4.8; pronotum length, male rounded, almost at the same level as the posterior bound- 4.5, female 6; pronotum maximum width, male 2.0, female ary of the vertex. Antennae filiform, reaching beyond half 2.7; forecoxa length, male 3.2, female 4.0, forefemur length, of body length. Scapus cylindrical, thickened near the male 3.5, female 5.0; hind femur length, male 7.2, female base; pedicellus hourglass shaped, a little shorter and 8.5, forewing length, male 8.5, female 11.0. half as narrow as the scapus. Remaining antennomeres Distribution. Vietnam: Lào Cai, Hòa Bình, Vin Phuc, cylindrical, basally slightly narrower than the pedicellus, Kon Tum (Figure 36). distally gradually elongating and narrowing; about 4 basal-most antennomeres of the flagellum completely fused. Tricondylomimus mirabilis (Beier, 1933) n. comb. Pronotum (Figure 17) robust, 1.7–1.8 times as long as (Figures 4, 17, 28, 30, and 35) its maximum width. Prozone parabolic; transverse groove deep laterally but almost flat dorsally, weakly Nemotha mirabilis Beier 1933: 323–325; Beier 1935: 45; arcuate; supra-coxal dilatation short, limited anteriorly Tinkham 1937b: 567; Wang 1993:50–51; Ehrmann 2002: by the transverse groove, 0.6 times the width of the 239; Otte & Spearman: 114; Zhu et al. 2012: 30. head; metazone 1.7–1.9 times as long as the prozone, Syn. Pseudogousa sinensis Tinkham 1937a: 491–492, Pl. 16, fig. 9, n. syn.; Wang 1993:97–98, fig. 97; Ehrmann 2002: 294; with the most narrow point posteriorly in the middle of it. Otte & Spearman 2005: 135; Zhu et al. 2012: 135, figs Dorsal surface of pronotum overall relatively flat, very therein. slightly bulging in the prozone and at the supracoxal dilatation, with a very thin medial carina along the meta- Type material. Holotype: ♀, China, Himalaya, Lung-tao- zone and a pair of transverse shallow grooves posteriorly. shan, 7.IX.1916, leg. Mell (ZMBH). The type (Figure 4)is The middle point of the supracoxal dilatation is inclined in good condition, except missing distal parts of antennae a little anteriorly. Prozone, supracoxal extension, middle and significant discoloration. line and the lateral pars of the posterior half of the The type of its synonym, Pseudogousa sinensis,is metazone covered by very small tubercles. Lateral likely lost, as it could not be found in the Lingnan edges very finely denticulated in the narrow part of the Natural History Survey and Museum materials, presently metazone, otherwise smooth. Furcasternum anteriorly deposited at Sun Yat-sen University (H. Pang & Zh. with low medial carina. Annales de la Société entomologique de France (N.S.) 189

Figures 33–35. Genera Nemotha and Tricondylomimus, life photographs. 33, N. metallica, ♀ from Yunnan, China, 34, N. metallica, ♂ from Yunnan, China; 35, T. mirabilis, ♀ from Hainan, China. Photographs © Chao Wu, published with permission.

Forelegs typical for praying mantises. Coxa triangular in loosely covered by tubercles with attached setae; ventral cross-section, substantially longer than the metazone of the surface covered by similar tubercles. Anterior apical lobe pronotum, armed dorsally with multiple very small spines, short, wide, strongly diverging from the base; the dorsal one seven of which are larger than the rest; a long seta is attached extended and visible from posterior as a small lobe. to each spine. Dorsal half of the anterior surface of the coxa Trochanters only slightly narrower than coxa, sharply 190 M.B. Stiewe & E. Shcherbakov

Figure 36. Distribution of Nemotha metallica, Tricondylomimus coomani, T. intermedius n. sp. and T. mirabilis. Only provinces are shown for T. mirabilis. curved. Femur widened ventrally, its length 3.7 times the medially; space between first and second spine equals the dorsoventral width. Dorsal edge smooth, a little convex near length of the first spine; second to fourth spine approximately the base. Anterior surface with a wide and deep claw groove, of equal size, becoming sharper and more curved distally; situated at the end of basal third of the femur, elongated spaces between second to fourth spines equal, each about 1.6 femoral brush, formed by straight setae, and several longer times as long as the space between the first and the second setae between the apex of the femur and the claw groove, as posteroventral spine. Posteroventral genicular spine similar well as around clawal groove. The femur (Figure 28)is in size and sharpness to the fourth posteroventral spine, but armed with 12–13 anteroventral, 4 discoidal, 4 posteroven- nearly straight. First discoidal spine very similar to the tenth tral, 1 anteroventral genicular and 1 posteroventral genicular anteroventral spine, but straight, shifted to the same line as spines. Ventral surface between basal 11 anteroventral spines the anteroventral row, immediately before claw groove; sec- and posteroventral spines covered by multiple small tuber- ond spine more than twice the length of the first, almost cles, especially numerous near the basal half of the antero- straight; third spine identical to the second, situated right at ventral row of spines, while the surface between the base of the median line of the femur, with the second and third the femur and the second discoidal spine bearing a row of discoidal spines on the same straight line with the first poster- flattened tubercles, each accompanied by a long seta. First oventral spine; fourth discoidal spine the smallest, situated anteroventral spine small, situated on the boundary of the near the beginning of the claw groove, a little anterior to the claw groove and directed ventro-basad. Second anteroventral median line of the femur, shorter than the smaller anteroven- spine greatly enlarged, only slightly smaller than the second tral spines, but wider, with very obtuse apex and inclined discoidal spine, situated at a distance from the first spine distally. Tibia straight, armed with 11–12 anteroventral equal to the length of the latter. Third to tenth anteroventral spines (without tibial claw) and 7 posteroventral spines. spine of alternating sizes, so that the third spine is less than a Basal seven anteroventral spines straight, inclined at about third of the length than the first one, and the second spine is a 30°, very sharp, first to fifth spine gradually elongating, sixth little shorter than the third; both the spines and the spaces and seventh spine very slightly shorter than fifth; eighth between them slightly elongating distally. Shorter spines spine as long as the fifth, thicker and less inclined, next shifted slightly medially compared to larger spines. three to four spines gradually becoming thicker, more curved Eleventh to penultimate spine approximately half as long and less inclined, with the spaces between them decreasing. the preceding uneven spines. Distal-most spine half as long Tibial claw large and wide, only half the length of the tibia, the second anteroventral spine, thinner, sharper and more with an anterodorsal, longitudinal and very narrow groove curved, situated at great distance from the tenth spine. and with rounded apex, its apex forms an 80° angle with the Anteroventral genicular spine very similar to the tenth ante- tibia. First posteroventral spine situated at about twice its roventral spine. All posteroventral spines emerging from length from the base; second spine the longest, 1.5 times as short sockets. First posteroventral spine very weakly curved, long as the first, its distance from the first equals its length; a little longer than the largest anteroventral spine, shifted third spine shorter than the first, as far away from the second Annales de la Société entomologique de France (N.S.) 191 as the length of the latter; fourth to fifth spine like the third, at the latter, not reaching the apical lobes; ventral surface dark- about the same distance from each other; sixth spine the olive; surface between dorsal and posterior edges with two shortest, about 0.7 times as long as the fifth and removed more or less fuzzy dark transverse bands. Anterior side of from the latter at a distance equaling the distance between the forefemora with a longitudinal dark brown streak in the distal second and the third spine; seventh spine of a size in between 2/3, the band being widened near its beginning and end; the third and the fourth spine, situated close to the sixth dorsal of this band there are two very short dark-brown spine. All posteroventral spines inclined at about 45°, basal transverse bands continuing its counterparts on the posterior four spines very sharp and very slightly curved, distal three side of the femur; the latter dark-olive, with one (basal) blunt and straight. Tarsi longer than femur and pronotum. incomplete and one (distal) complete transverse green Basitarsus as long as remaining tarsomeres combined, cov- band; dorsal edge of the femur green. Foretibia pale green; ered by straight rows of short setae. anterior side with a dark-brown, medial longitudinal narrow Metathoracical cyclopean ear of POT type, with a deep streak and a medial transverse wide band, running dorsally triangle pit limited anteriorly by the basisternum wall, till the median line of the posterior side of the tibia; together, posteriorly by fused walls. these two bands form an extremely stretched upside-down T. Middle and hind legs cursorial, very long. Coxae short Apex of foretibia brown. Fore basitarsus black dorsally, pale and wide, triangular, with a low ventral carina. Femora with dark base and apex ventrally, other tarsomeres pale with straight, a little thickened in basal 2/3, with a posterior row dark apices. All spines dark-brown to black. Middle and hind of small, spaced cuticular spines, each with a seta, and coxae pale yellow, basally mottled with olive. Middle and with a long apical spine. Tibia thin, straight, covered by hind femora with three transverse dark-brown bands, absent setae, apically with a dorsal triangular extension and two on the ventral side. On middle femur, starting from the base, ventral spines. Tarsi slightly shorter than femora, hind the first band is situated at the end of the basal third, with basitarsus longer than remaining tarsomeres combined. diffuse basal and sharper distal boundaries, dorsally sending Wings shortened, covering first six tergites. Forewings well-defined thin line towards base; the second band with more or less rectangular, 2.7 times as long as wide, with light gaps separating it from the first and the third band widely parabolic apex. Costal field widest at the end of the occupies roughly the middle third of the femur, with uneven basal third, gradually narrowing and almost reaching the basal and distal boundaries, posteriorly connected to adjacent apex. All main veins weakly curved. CuAwith two branches. bands via dark medial lines; the third band almost reaching Cross-veins form irregular cells (Figure 30), except in the the apex of the femur, with sharp boundaries; all bands anal area. Pseudo-veins absent, except the short ones in the sharply and significantly shrinking on the anterior side. On most distal part of discoidal area. Stigma large, longer than hind femur, the bands are similar to those on the middle wide, roughly triangular, situated on RP+M and the anterior femur, except the dark dorsal line directed basal from the first branch of CuA, with fuzzy boundaries. Hind wings shorter band expands basally and sometimes forms posteriorly than forewings, semi-circular with widely rounded apex. another, very small transverse band. Apical spines dark- Abdomen relatively short, all tergites and sternites wider brown. Middle and hind tibia with three transverse bands, than long, with the fifth tergite and the fourth visible sternite both their length and the distance between them elongating being the widest. Anterior and posterior edges of tergites and distally; at least the first and the second band connected by a sternites straight. Anal plate wider than long, semicircular, thin dorsal line; the third band reaches the apex of tibia. with a feeble longitudinal medial folding. Cerci formed by Middle and hind basitarsi with brown base and apex, remain- 10 cercomeres, gradually elongating distally, the last cerco- ing tarsomeres completely dark-brown. Forewings blackish, mere more than twice as long as the penultimate one, flattened main veins brown, except Sc, proximal halves of R, M and laterally, remaining cercomeres circular in cross-section. CuA; the latter and all cross-veins pale green; an irregular Coloration. Main body color dark-olive (Figure 35). spot in the distal third of the costal field, the stigma and a Labrum and scutellum blackish, clypeus pale yellow with basal spot between A1 and A2 bright to pale green. Wings anterior triangular and posterior roughly rectangular medial infumate, especially near the anterior edge. Tergites I–VI marks. Eye sockets green dorsally, pale yellow with an oval each with large, metallic blue dark spot narrowing ante- dark spot ventrally. Scapi and area between them pale yel- riorly, several very small lateral brown spots and a bright low. Vertex with a pair of reverse arrowhead-shaped green green lateral line above spiracle. Tergites VII–IX each marks, continuing posteriorly as a pair of parallel lines. with a medial, pale green and triangular mark directed Metazone of pronotum with three pairs of paramedian, posteriorly, emarginated by diffuse brown spots, and with fuzzy, elongated, light strokes, all marginal and lateral parts an irregular lateral brown line on each side. Each sternite of the prozone and of the supracoxal dilatation green. Pro-, with a large, central metallic and blue dark spot, flanked meso- and anterior part of metasternum blackish with an on each side by a very small dark spot. Cerci black. intense blue metallic shine. Forecoxae green on and around Subgenital plate completely metallic black. Preserved spe- posterior edge, pale yellow on dorsal edge and anterior sur- cimens may experience discoloration, especially in the face, with a dark narrow dorsal streak in the apical third of dark pattern. 192 M.B. Stiewe & E. Shcherbakov

Male. Similar to female, but smaller and more slender pentagonal, medially bulging, dorsally fused with clypeus. (body length 19.0). Antennae longer than body. Clypeus very thick and bulging, medially strongly elevated, Forefemora with 14 anteroventral spines. Fore and hind forming a transverse carina, centrally with a very short and wings narrower; forewings 3.4–3.6 times as long as wide, low medial transverse carina. Ventral carina of clypeus reaching the end of abdomen. Coloration overall lighter, interrupted in the middle by a rhomboidal relatively flat light elements on head and pronotum enlarged. Metazone area with a very thin longitudinal carina running from the of pronotum with a pair of large paramedian, arcuate lines, central transverse carina to the place of the clypeus-labrum homological to the three pairs of light strokes in females; fusion. Scutellum short, transverse, arcuate, of constant additionally, metazone with medial pale triangle adjacent length across its width. Width of scutellum 3.7 times as to the posterior edge, directed anteriorly. Forecoxae pale large as its length at the middle. Surface of scutellum green with an indistinct basal dark spot on the posterior rugose, much less so in the lateral quarter of the width, side. The longitudinal dark line on the anterior side of the with carina vanishing in the same lateral fourths. forefemora shifted slightly dorsally, while an additional Compound eyes large, oval, bulbous, with narrow semi- narrow darkening is present along the anteroventral series oval eye sockets. Ocelli very small, nearly circular, covered of spines. Anterior side of foretibia almost completely with thick rough cuticle flush with surrounding surface. blackish. Anterior side of middle and hind femora without Lateral ocelli slightly larger than the medial one and form- dark bands. Forewings semi-opaque, cross-veins very pale ing a slight angle to the surrounding cuticle. Vertex almost with light areas along them. Tergites darker. flat, with only very low elevations near each ocellus and fontal suture raised medially into very low carina. Posterior Measurements (in mm). Body length, male 19.0, female edge of the head weakly arcuate, protrudes far beyond the 21.0–24.0; head width, male 4.3–4.7, female 5.5; compound eyes (Figure 15). Parietal sutures well devel- pronotum length, male 5.0–5.5, female 5.6; pronotum oped, juxta-ocular bulges large, reaching slightly beyond maximum width, male 2.7, female 3.2; forecoxa length, the posterior edge of the head. Antennae filiform, nearly as male 4.0, female 4.7; forefemur length, male 5.5–6.0, long as the body. Scapus cylindrical, thickened near the female 6.0; hind femur length, male 11.0, female 10.0; middle, 2.5 times as wide as the lateral ocellus. Pedicellus forewing length male 12.5–13.0, female 13.5. hourglass shaped, slightly shorter and half as wide as the scapus. Remaining antennomeres cylindrical, basally

fi Remarks. Zhu et al. (2012) published the rst and only slightly narrower than the pedicellus, distally gradually life history data known for this species. They observed the elongating and narrowing; about 4 basal-most antenno- “ species at 250 and 850 m. Ecologically, it is a bark meres of the flagellum completely fused. ” mantis , a fast trunk dweller like members of Pronotum (Figure 18) relatively short, dorsally bul- Liturgusidae. Last instar nymphs were observed at the ging, in cross section nearly semicircular. Prozone para- end of July, moving quickly in spurts on the surface of bolic; transverse groove deep, only weakly angular; trunks of Schima sp. and Ficus pumila L. Several adult supracoxal dilatation short, but well developed, limited females were collected by the end of August. The ootheca anteriorly by the transverse groove, 0.6 times as wide is small, containing very few eggs, laterally rhomboidal, the head; metazone twice the length of the prozone, with higher than long, with the anterior sprout almost as long as most narrow point on the posterior in the middle of it. the height of the ootheca. Dorsal profile of metazone slightly Ƨ-shaped (Figure 18), with highest point in the posterior part of the supracoxal Distribution. China: Guangxi, Guangdong, Fujian, dilatation and with a pair of very weak elevations adjacent Yunnan, Hainan (Figure 36). to the posterior edge of the pronotum. Dorsal surface of pronotum with a pair of shallow incomplete transverse Tricondylomimus intermedius n. sp. grooves in the posterolateral parts of the supracoxal dila- (Figures 5, 14, 15, 18, 23, 25, 27, 29, and 31) tation and very small sharp tubercles in the lateral parts of metazone, otherwise with extremely fine honeycomb microstructure or smooth. Anterior, posterior and lateral Type material. Holotype (Figure 5): ♀, Vietnam, Lâm Đồng edges at the widest point of the supracoxal dilatation province, BảoLộc pass, 11°26ʹ59.6ʺN107°42ʹ43.4ʺE, smooth, the rest of lateral edges with small, sparse spines. III.2015, leg. Văn Đặng. (ZIN). Each of the lateral spines and dorsal metazonal tubercles with a short seta. Ventral cervicalia completely absent. Description Intercervicalia medially strongly narrowed, not fused, Female. Figure 5, 14, 15, 18, 23, 25, 27, 29, and 31. longer than lateral cervicalia. Postcervical plate extremely Medium sized (body length 21.7 mm). short, similarly to the T-shaped sclerite, the latter with its Head (Figures 14 and 15) pentagonal, slightly flattened posterior part very thin and partly reduced. Furcasternum dorsoventrally, 1.3 times as wide as long. Labrum anteriorly strongly angular, medially raised into a carina. Annales de la Société entomologique de France (N.S.) 193

Forelegs relatively wide, with smoothed dorsal edges, the median line of the femur, with the second and third contrary to most Mantodea. Coxa almost circular in cross- discoidal spine on the same straight line with the first poster- section, longer than metazone of pronotum, armed dorsally oventral spine; fourth discoidal spine the smallest, situated with six very small spines decreasing in size distally, each near the beginning of the claw groove, a little anterior to the accompanied by a long seta, with even smaller spinules in median line of the femur, shorter than the smaller anteroven- between. Anterior surface of the coxa with a loose, irregular tral spines, but wider, with very obtuse apex. Tibia row of very low, weak tubercles. Posterior edge with spines (Figure 25) straight, its ventral surface with an anterior row similar to the spines on the dorsal edge, but much smaller. of tubercles near the apex, armed with 11 anteroventral Ventral surface smooth. Anterior apical lobes short, wide, spines (without tibial claw) and 6 posteroventral spines. strongly diverging from the base; the dorsal lobe extended The seven proximal anteroventral spines straight, inclined, and visible from posterior as a small lobe. Trochanter only very sharp, the first to fifth spine gradually elongating, the slightly narrower than coxa, sharply curved. Femur (Figures sixth and seventh spine a little shorter than the fifth; the sixth 27 and 29) widened ventrally, its length 3.8 times the max- spine thicker and less inclined, almost three spines becoming imum dorsoventral width. Dorsal edge wide and smooth, gradually thicker, more curved and less inclined, with the more abrupt posteriorly, convex in the basal. Anterior surface space between them decreasing. Tibial claw wide, 1/3 the of femur with a very wide and deep claw groove, situated in length of the tibia, with a longitudinal, anterodorsal, very the middle of the basal half of the femur, femoral brush narrow groove and a posteroventral ridge, as well as with an formed by thick, long and curved setae, and with several obtuse apex forming a straight angle with the tibia. First longer and straighter setae before and especially after the posteroventral spine situated at about 1.3 times its length claw groove. The femur is armed with 11–12 anteroventral, 4 from the base; the second spine is the longest, 1.5 times as discoidal, 4 posteroventral, 1 anteroventral genicular and 1 long as the first, its distance from the first equals the length of posteroventral genicular spines. Ventral surface between the latter; third spine as the first, as far from the second as the basal 10 anteroventral spines and basal 3 posteroventral length of the latter; fourth spine half the length of the third, its spines covered with multiple small tubercles, more numer- distance from the third slightly smaller than the distance ous near the anteroventral row of spines (Figure 29, tbr), between the preceding two spines; fifth spine the shortest, while the surface between the base and the second discoidal less than 0.7 times the length of the fourth and removed from spine bears a row of flattened tubercles, each accompanied the latter at a distance equaling the distance between the by a long seta. First anteroventral spine greatly enlarged, second and the third spine; sixth spine longer than the fourth only slightly smaller than the second discoidal spine. but shorter than the third, situated close to the fifth spine. All Second to ninth anteroventral spines of alternating sizes, so posteroventral spines inclined at around 45°, the basal three that the third spine is less than a third as long the first one, spines very sharp and very slightly curved, distal three blunt and the second spine a little shorter than the third; both the and straight, the sixth spine particularly thick. Tarsus longer spines and the spaces between them slightly elongating dis- than femur and pronotum. Basitarsus as long as remaining tally. Shorter spines shifted slightly medially compared to tarsomeres combined, covered by straight rows of short larger spines. Tenth to penultimate spine approximately half setae. Euplantulae of the proximal three tarsomeres oval, as long as the preceding even spines. Distal-most spine 0.7 with a median separation; each euplantula of fourth tar- times as long as the first anteroventral spine, thinner, sharper, somere rhomboidal, enlarged; in the dried specimen the and more curved, with a large gap between it and the penul- cuticle of the fourth tarsomere forms a sheath around euplan- timate spine. Anteroventral genicular spine similar in size to tulae, with an anteromedial U-shaped window. the ninth anteroventral spine, but thinner and with sharper Metathoracical cyclopean ear (Figure 23) of POT type, apex. All posteroventral spines emerging from short sockets. with a deep triangular pit limited anteriorly by the basi- First posteroventral spine straight, with rounded apex, 0.7 sternum wall, posteriorly by fused walls. times as long as the third discoidal spine, shifted medially; Middle and hind legs cursorial, very long, though the space between the first and the second spine equals the middle legs noticeably shorter than hind legs. Coxae length of the first spine; second to fourth spine approximately short and wide, triangular, metacoxa with a very low of equal size, becoming more curved and sharper distally; ventral carina. Femora straight, thickened basally, distally spaces between the second and third, and third to fourth with a posterior row of small, spaced cuticular spines, spine equal, each slightly larger than the space between the each with a seta, and with an apical spine. Tibia thin, first and the second posteroventral spine. Posteroventral straight, covered by setae, apically with a dorsal triangular genicular spine similar in size and sharpness to the fourth extension and two ventral spines. Tarsi slightly shorter posteroventral spine, but straight. First discoidal spine iden- than femora, hind basitarsus longer than remaining tar- tical to the posteroventral genicular spine, shifted to the same someres combined. line as the anteroventral row, immediately before claw Wings shortened, covering first four tergites. groove; second spine twice the size of the first, almost Forewings more or less rectangular, with widely rounded straight; third spine identical to the second, situated right at apex. Costal field widest near the base, gradually 194 M.B. Stiewe & E. Shcherbakov narrowing and almost reaching the apex. All main veins length 10.1, hind tarsus length 7.6, forewing length 8.7, straight or very weakly curved. CuA with three branches. forewing width 3.9, hind wing length 8.0. Cross-veins irregular in costal field and forming rectangu- lar cells in most of the discoidal and anal area. Pseudo- Diagnosis. This elegant and unusual species can be veins absent, except the short ones in the most distal part distinguished from T. coomani by the following characters: of discoidal area. Stigma longer than wide, roughly trian- (1) the vertex when viewed from the anterior perspective is fi gular, situated on RP+M and the rst branch of CuA, with much less bulging in T. intermedius (Figure 14)thaninT. irregular boundaries. Hind wings shorter than forewings, coomani (Figure 12); (2) the prozone is more gently sloping semi-circular with widely rounded apex. dorsally in T. intermedius (Figure 18)thaninT. coomani Abdomen relatively short, all tergites and sternites (Figure 20); (3) the anterior part of the metazone is only fi wider than long, with the fth tergite and the fourth visible weakly convex dorsally in T. intermedius (Figure 18), while sternite being the widest. Anterior and posterior edges of it is very strongly bulging in T. coomani (Figure 20); (4) the tergites and sternites straight. Supra-anal plate semicircu- posterior part of the metazone is almost flat in T. intermedius lar. Cerci formed by 10 cercomeres, gradually elongating (Figure 18), contrary to T. coomani (Figure 20); (5) the distally, the last cercomere more than twice as long as the forewings of T. coomani aremuchlongerthanthoseofT. fl penultimate one, attened laterally, other cercomeres cir- intermedius, reaching almost the end of the abdomen cular in cross-section. (Figures 3 and 9); (6) the cross-veins in the distal half of Coloration. Head brown with a weak bronze shine; the forewing form almost regular rows of large and lateral parts of scutellum with a metallic greenish shine, rectangular cells in T. intermedius (Figure 31), unlike the cuticle near lateral ocelli with metallic blue shine. irregular net of polygonal cells in T. coomani (Figure 32); the Pronotum brown with weak metallic shine. T-shaped scler- cross-veins are also much more raised in T. coomani; (7) the ite and furcasternum metallic blue. Anterior surface and stigma is large, triangular and brown in T. intermedius most of the posterior surface of the forecoxae dorsally of (Figure 31), but reduced, rectangular and whitish in T. the posterior edge are reddish-brown, lighter distally; ven- coomani (Figure 32). tral surface, all posterior surface ventral the posterior edge T. intermedius differs from T. mirabilis by the follow- and a small basal part dorsally black. Anterior side of fore ing characters: (1) the metazone of T. intermedius is trochanters anteriorly basally yellow at base, distally

slightly Ƨ-shaped (Figure 18), while in T. mirabilis it is white; posterior side dark-brown. Forefemora light red- almost flat, with only a weak concave part around the dish-brown with white dorsal, anteroventral and poster- middle of its length (Figure 17); (2) the largest anteroven- oventral edges, and yellow ventral surface; all femoral fi spines black. Foretibia anteriorly whitish, posteriorly tral spine of the forefemur is the rst in T. intermedius (Figure 27, i1), while it is the second in T. mirabilis brownish, with two black stripes, an anterior and a poster- (Figure 28, i2); (3) the foretibia has 6 posteroventral ior one, not reaching the apex of tibia; all edges white, all spines in T. intermedius, but 7 in T. mirabilis; (4) the tibial spines black. Fore basitarsus black, all other tar- forewings of T. mirabilis are noticeable longer, reaching someres grey to brown. Meso- and metathorax dorsally at least tergite VII, while in T. intermedius they are reach- light-brown, ventrally metallic blue. Meso- and metafe- ing only tergite V; (5) RP + M is bifidinT. intermedius, mora dark reddish-yellow. Meso- and metatibia and tarsi but simple for the most or all of its length in T. mirabilis; dark-brown to black. Forewings olive, transparent except semi-opaque costal field, with brown veins and stigma. (6) CuA in the forewing with 3 branches in T. intermedius and 2 in T. mirabilis; (7) A1 on the forewing with 1 Hind wings transparent, very slightly darkened, except branch in T. intermedius and 2 branches in T. mirabilis; brownish costal edge and apex; main veins dark-brown. (8) the cross-veins in the distal half of forewing form Tergites olive-brown, with a narrow, metallic dark blue almost regular rows of large and rectangular cells in area along posterior edge, which gradually widens caud- T. intermedius (Figure 31), unlike the highly irregular net ally, attaining the whole tergite’s length at tergite V; tergite in T. mirabilis, especially between CuA1 and RP + M VI only with a slight bronze shine, while tergites VII to IX (Figure 30); (9) the coloration is strongly different: and supra-anal are all matt. All sternites with a slight metallic green shine. T. intermedius is olive-brown with reddish legs and areas of metallic blue or green, while T. mirabilis is olive-green with stripes and spots of bright green, white, and dark- Male. Not known. brown; only parts of the tergites are metallic blue. Measurements (in mm). Body length 21.7, head width 4.6, pronotum length 5.2, prozone length 1.7, maximum Remarks. This species likely represents an intermediate width of pronotum 2.7, minimum width of pronotum 1.6, ecological form between the other two species. It forecoxa length 3.9, forefemur length 4.8, foretibia length resembles T. mirabilis in the shape of most body parts, 3.3, foretarsus length 6.3, hind femur length 9.3, hind tibia however in the number of the branches of CuA and RP+M Annales de la Société entomologique de France (N.S.) 195 and its coloration it is similar to T. coomani. The shape of all major groups, preventing superficial similarities from the pronotum is also intermediate between these species. obscuring relationships. It has to be mentioned that both genera revised here, Distribution. Vietnam: Lâm Đồng (Figure 36). Nemotha and Tricondylomimus, have species with charac- teristics unique among Mantodea. Etymology. The species was named on the basis of its The bright body coloration of N. metallica can be features showing similarities with both of the other two considered highly unusual for a praying mantis. While species of the genus. the genera Metallyticus Westwood, 1835 and Tricondylomimus Chopard, 1930 show a similar metallic coloration (but are taxonomically distant from Nemotha), Discussion only N. metallica features a striking yellow and black Analyses of the literature on the genera Nemotha, bi-coloration in adult specimens (Figures 1, 6, and 7). Tricondylomimus and Pseudogousa show that most The vertex part of the head is yellow and the rest of authors who mentioned these species in their works did head is metallic black. The middle part of the prozone is not examine actual specimens but relied instead on the yellow and the surrounding margin is metallic black. The original descriptions. A chain of assumptions, combined metazone is similarly colored as the prozone, with bright with the fact that many boundaries between the taxa in the yellow coloration and a metallic black lateral margin, and traditional Mantodea system are highly subjective, led to the wings are yellowish. In other orders (other than the acceptance of a polyphyletic genus and the two cryptic Hymenoptera), such a bright, contrasting yellow-and- synonyms, which sometimes coexist within one book. black coloration is usually indicative of Batesian wasp Despite not seeing the actual specimens of Nemotha mimicry. However, the potential hymenopteran model in metallica, Beier (1933) tentatively assumed that his new case of N. metallica is unclear. species mirabilis belongs to the same genus, referring to On the other hand, T. coomani possesses a similarities in pronotal features and rare metallic colora- surprising similarity to the tiger beetle genus Tricondyla tion. On this basis, he moved the whole genus Nemotha to Latreille, 1822 (Coleoptera: Cicindelidae), which gave Iridopteryginae. His doubts about the generic affinity of Tricondylomimus its name (Figure 10). Besides general mirabilis with metallica were left unnoticed by subsequent habitual resemblance (Figures 8–10), the profile of the authors. The holotype of mirabilis had not been studied by pronotum shows the same number of swellings and con- anyone since, and no new specimens had been collected. strictions (Figures 19–21), the cells of forewings are Tinkham (1937b) did not take into account Beier’s transfer noticeably deepened, forming a honeycomb structure of Nemotha to Iridopterygidae. Although he was uncertain similar to the punctuation of the elytra in Tricondyla about the systematic position of Nemotha due to his unfa- (Figure 32), and the coloration is the same in all major miliarity with the species, he moved mirabilis and metal- aspects (Figures 8–10). Chopard (1930) was puzzled by lica back to Odontomantes, following the placement of N. this resemblance. Despite its accuracy, he expressed metallica by Giglio-Tos (1927). At the same time, doubts that this can be a case of Batesian mimicry, since Tinkham (1937a) described Pseudogousa sinensis. it is difficult to see what advantages such mimicry can Though he recognized similarities between Spilomantis, give to T. coomani, supposedly a fierce predator just like Tricondylomimus coomani and P. sinensis, he considered Tricondyla. As an explanation, Chopard suggested an the morphological separation between the latter two sig- influence of similar living conditions, i.e. a simple nificant enough to justify a new genus. Unfortunately, a convergence. number of similarities between P. sinensis and Beier’s Aside from Tricondylomimus, only nymphs of the description of N. mirabilis, despite them relying on differ- katydid genus Leptoderes Serville, 1838 (= Condylodera ent sexes, have escaped his attention. Beier (1935) syno- Westwood, 1841; Orthoptera: Tettigoniidae) bear even nymized Nemotha and Tricondylomimus without stronger resemblance to Tricondyla and are long regarded comments. It is unknown if he ever saw T. coomani as mimics of the latter (Shelford 1902; Naviaux 2002). specimens. This synonymy was considered valid in all The lifestyle of T. coomani is unknown, but it may be subsequent works. Later authors who personally saw similar to T. mirabilis, which is a trunk dweller like N. coomani were apparently unfamiliar with N. metallica Tricondyla (Zhu et al. 2012; see above). If T. coomani is and vice versa. indeed a mimic of Tricondyla, then T. intermedius might Overall, this resembles the confusion about identity represent an initial stage of the evolutionary transition to and systematic placement of the genera Majangella or from this state, bearing general resemblance to Giglio-Tos, 1915 and Ephippiomantis Werner, 1922 Tricondyla in habitus and coloration (Figure 5), yet lack- (Svenson & Vollmer 2014). There is a strong need for a ing the specialized features of T. coomani and being more phylogenetic high level classification of the whole similar in morphology to T. mirabilis. However, more Mantodea system, with clearly stated synapomorphies for studies are required to confirm whether the similarity of 196 M.B. Stiewe & E. Shcherbakov

Tricondyla and Tricondylomimus represents a case of Giglio-Tos E. 1927. Das Tierreich. 50. Lfg. Orthoptera Mantidae. mimicry at all. Berlin: Walter de Gruyter & Co.; XL + 707 p. These questions posed by the morphology make the Klass K-D. 1997. The external male genitalia and the phylogeny of Blattaria and Mantodea. Bonner Zoologische Monographien. genera Nemotha and Tricondylomimus highly interesting Bd. 42. Bonn: Zoologisches Forschunginstitut und Museum targets for a life history study. Alexander Koenig; 341 p. Mukherjee TK, Ehrmann R, Chatterjee P. 2014. Checklist of Mantodea (Insecta) from India. Priamus (Serial Publication Acknowledgements of the Centre for Entomological Studies Ankara). 30:1–61. Mukherjee TK, Hazra A. 1985. Insecta: Mantodea. Records of The authors sincerely thank Roger Roy (MNHN), Christian the Zoological Survey of India. 82:33–39. Schwarz (University Bochum, Germany), Reinhard Ehrmann Mukherjee TK, Hazra A, Ghosh A. 1995. The mantid fauna of (SMNK), Michael Ohl (ZMB), Gavin Svenson (CMNH), George India (Insecta: Mantodea). Oriental . 29:185–358. Beccaloni (BMNH) and James Hogan (OXUM), without whom Naviaux R. 2002. Les Tricondylina (Coleoptera, Cicindelidae). this publication in its present form would not have been possible. Révision des genres Tricondyla Latreille et Derocrania We are thankful to Amoret Spooner (OXUM) for providing us Chaudoir et descriptions de nouveaux taxona. Mémoires de with high resolution photographs of the holotype of Mantis metal- la Société entomologique de France. 5:1–106. lica and to Roger Roy and Marion Depraetere (MNHN) for Otte D, Spearman L. 2005. Mantida species file. Catalog of the permission to use the photographs of the T. coomani type speci- mantids of the world. Diversity Association, Philadelphia; 489 p. mens. We are indebted to Tushar Mukherjee, Kailash Chandra and Saussure H. de 1871. Mélanges orthoptérologiques. Supplément au the staff of the Orthoptera Section of Zoological Survey of India IIIme Fascicule. Mantides. Mémoires de la Société de Physique (Kolkata) for sending us additional N. metallica photos and infor- et d’Histoire naturelle de Genève. 21:239–337, pl. 7. mation, We thank Hong Pang, Zhang Binglan (Sun Yat-sen Shcherbakov E, Ehrmann R, Borer M. 2016. Revision of the University, Guangzhou) and Mark A. McGinley (Lingnan genus Heliomantis Giglio-Tos, 1915 (Insecta: Mantodea: University, Hong Kong) for their efforts to locate Pseudogousa Hymenopodidae). Annales de la Société entomologique de sinensis holotype. We also thank David Yager (University of France (N.S.). 52:135–149. Maryland) for the information on metathoracical ear of Nemotha Shelford R. 1902. Observations on some mimetic insects and and Pseudogousa, Andrey Matalin and Kirill Makarov (Moscow spiders from Borneo and Singapore. Proceedings of the State Pedagogical University, Moscow) for lending us a specimen Zoological Society of London. 2:230–284. of Tricondyla mellyi Chaudoir 1850, Alexander Prosvirov Shorthouse DP 2010. SimpleMappr, an online tool to produce (Lomonosov Moscow State University) for help with a macro- publication-quality point maps. [cited 2017 Jan 18]. photography system and Sergey Dementiev (Moscow) for the Available from: http://www.simplemappr.net literature on the genus Tricondyla. Furthermore, we would also

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