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

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Revision of the Heliomantis Giglio-Tos 1915 (Insecta: Mantodea: )

Evgeny Shcherbakov, Reinhard Ehrmann & Matthias Borer

To cite this article: Evgeny Shcherbakov, Reinhard Ehrmann & Matthias Borer (2016) Revision of the genus Heliomantis Giglio-Tos 1915 (Insecta: Mantodea: Hymenopodidae), Annales de la Société entomologique de France (N.S.), 52:3, 135-149, DOI: 10.1080/00379271.2016.1220264

To link to this article: http://dx.doi.org/10.1080/00379271.2016.1220264

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Download by: [Evgeny Shcherbakov] Date: 23 October 2016, At: 17:45 Annales de la Société entomologique de France (N.S.), 2016 Vol. 52, No. 3, 135–149, http://dx.doi.org/10.1080/00379271.2016.1220264

Revision of the genus Heliomantis Giglio-Tos 1915 (Insecta: Mantodea: Hymenopodidae) Evgeny Shcherbakova*, Reinhard Ehrmannb & Matthias Borerc aFaculty of Biology, Lomonosov Moscow State University, Moscow, 119991 Russia; bState Museum of Natural History Karlsruhe Division of Entomology, Erbprinzenstrasse 13, Karlsruhe, D – 76133, Germany; cNatural History Museum of Basel, Augustinergasse 2, Basel, CH – 4001 Switzerland (Accepté le 30 juillet 2016)

Summary. The little-known flower genus Heliomantis Giglio-Tos 1915 (Hymenopodidae) distributed in South and South-Eastern Asia is revised. All are redescribed. New data on the morphology, distribution and ecology are given, and all known specimens are listed. The lectotype of H. elegans (Navás 1904) is designated. Due to highly divergent morphology, H. latipennis Werner 1930 from Sarawak, Borneo, is transferred to Werneriana n. gen., leaving the genus Heliomantis with the only species, H. elegans (Navás, 1904).

Résumé. Révision du genre Heliomantis Giglio-Tos 1915 (Insecta: Mantodea: Hymenopodidae). Le genre peu connu de mante-fleur Heliomantis Giglio-Tos 1915 distribué en Asie du Sud et du Sud-Est est révisé. Toutes les espèces sont à nouveau décrites. De nouvelles données sur la morphologie, la distribution et l’écologie sont apportées, et tous les spécimens connus sont répertoriés. Le lectotype de H. elegans (Navas 1904) est désigné. En raison de sa morphologie très divergente, H. latipennis Werner 1930 de Sarawak, Bornéo, est transféré dans le nouveau genre Werneriana n. gen., laissant le genre Heliomantis avec une seule espèce, H. elegans. http://www.zoobank.org/urn:lsid:zoobank.org:pub:DDA3941B-1A65-4576-8E92-570B0C9B2FFE Keywords: ; Anaxarchini; taxonomy; revision; new genus; Oriental region Mots clés: Dictyoptera; Anaxarchini; taxonomie; révision; nouveau genre; région de l’oriental

the two species of the genus Heliomantis actually belong to Hymenopodidae (the flower ) is a diverse praying different genera. mantis family including at least 38 genera distributed in Africa and Asia (Svenson et al. 2015). The group is extremely heterogenous. While many of its genera are famous for sup- History of the genus posed plant mimicry, including flowers (O’Hanlon et al. Navás (1904) described the species Polyspilota elegans 2014), others do not exhibit such peculiar features – they Navás 1904 on the basis of a pair, a male and a female, have a very short or absent vertex process, possess a long collected in Kurseong (now a town in Darjeeling district prothorax, slender fore femora, very short middle and hind leg of West Bengal, India). The description was detailed, but lobes (if any) and are not particularly colourful; in other without figures. Later, the female was sent to I. Bolivar words, they resemble the “typical green mantis” ecomorph. and the male was sent to E. Giglio-Tos for further studies. Among such genera is Heliomantis Giglio-Tos 1919,a Bolivar (1914)erectedthegenusParaspilota Bolivar rarely collected genus, which has been placed in 1914 for P. elegans on the basis of the female sent to him. Anaxarchini in the recent revision of the In his opinion, the species is close to Polyspilota Burmeister flower mantises and allied families (Svenson et al. 2015). 1838, Prohierodula Bolivar 1908 and Sphodromantis Stål The genus includes two species with widely separated dis- 1871 (), with which he compared the new genus. tributions, H. elegans (Navás 1904)andH. latipennis Werner Giglio-Tos (1915) erected the genus Heliomantis Giglio- 1930. They also differ considerably by their morphology, Tos 1915 for Polyspilota elegans on the basis of the male sent making both the old key of Giglio-Tos (1927)andthenew to him, placing the new genus in . Giglio-Tos key by Svenson et al. (2015) insufficiently clear for identifica- provided no description and figures for the erected genus, only tion of the genus: the former was constructed before the noting that it “closely resembles” Anaxarcha Stål 1877 (“assai description of H. latipennis,whileonlyH. elegans was inves- affine a Anaxarcha”; Giglio-Tos 1915,p.2). tigated for the latter. By a thorough examination of the type Giglio-Tos (1919) placed the genus Heliomantis in material and the original descriptions, it became evident that Acromantinae Acromantes, noting, however, that it is almost

*Corresponding author. Email: [email protected]

© 2016 Société entomologique de France 136 E. Shcherbakov et al. intermediate between the groups Acromantes and Anaxarchae ICZN: International Code on Zoological Nomenclature (i.e. Anaxarcha, the only genus in the group at that time). He (ICZN 1999) mentioned the work of Bolivar and possible synonymy of MNCN: Museo Nacional de Ciencias Naturales, Madrid, Spain NHMW: Naturhistorisches Museum, Vienna, Austria Heliomantis and Paraspilota Bolívar 1914. He admitted that NMB: Natural History Museum of Basel, Basel, Switzerland he had no access to the work and no communication with I. NRM: Naturhistoriska Riksmuseet, Stockholm, Sweden Bolivar, so the true status of Paraspilota elegans was SMNK: State Museum of Natural History, Karlsruhe, Germany unknown to him. Apparently, both Giglio-Tos and Bolivar ZSI: Zoological Survey of India, Kolkata, India didn’t know that Bates (1888) described Paraspilota Bates ZSM: Zoologische Staatssammlung, München, Germany 1888 as a subgenus of the genus Anomala Samouelle 1819 (Coleoptera: Scarabaeidae: Rutelinae). According to Articles Examined material of Heliomantis elegans (Navás 1904) used 23, 52 of ICZN, that makes Paraspilota Bolívar 1914 a junior in this study is deposited in the collections of SMNK, NMB and ZSM. The male genitalia were prepared according to the standard homonym and invalid name. protocol for Mantodea (e.g. Shcherbakov & Savitsky 2015). Werner (1916) described the genus Deiroharpax Werner Photographs of specimens of the main material were taken with a 1916 with two species: D. viridis Werner 1916, based on a Canon PowerShot A630 digital camera (Tokyo, Japan) and pro- male from Sikkim, India and D. hyalina Werner 1916,based cessed using software Adobe Photoshop 8.0.1 (Adobe Systems, on a female from Sumatra. He provided figures of the San Jose, CA, USA). Photographs of the male genitalia were taken with Keyence VHX-2000 digital microscope (Osaka, Japan). pronotum and forewings along with the descriptions. Photographs of the lectotype of H. elegans were kindly provided Giglio-Tos (1927) treated the genus Heliomantis in the by Mercedes París (MNCN), made with a Nikon D700 camera group Acromantes and briefly described the male that was (Tokyo, Japan) and Tamron 90 mm macro lens (Saitama, Japan). sent to him, again with no figures. Photographs of the holotype of Deiroharpax viridis were kindly Werner (1930) described the second species of provided by Harald Bruckner (NHMW), made with Nikon D60 camera, equipped with Nikon AF-S Micro Nikkor 105 mm 1:2.9 G Heliomantis, H. latipennis Werner 1930, during his work ED lens and Sigma EM-140 DG flash (Kawasaki, Japan). on the Asian Mantodea from the collection of NRM. A brief High resolution photographs of the type of H. latipennis description on Latin was based on a female from Sarawak, a Werner 1930, made with Canon EOS Utility 5D and Zerene photograph of which was also in the paper. Werner noted its Stacker software (http://zerenesystems.com/cms/home) were stu- proximity to D. viridis, but provided no differential diagno- died, due to courtesy of Gunvi Lindberg (NRM). All figures were processed using Adobe Photoshop CS5 by sis. On the page 6 of the same paper Werner placed a note the first author. about a letter from Max Beier, who recognized that D. Terminology of the external features is given according to hyalina belongs to the genus Oligomantis Giglio-Tos 1915, Wieland (2013) and Svenson et al. (2015), of the male genitalia while D. viridis is identical to H. elegans. according to Klass (1997). Beier (1934), in his catalogue of Hymenopodidae, for- mally synonymized D. viridis with H. elegans (simulta- neously, the genus Deiroharpax became junior synonym of Heliomantis)andtransferredD. hyalina to the genus Results and discussion Oligomantis. While he left H. latipennis Werner 1930 Even at first glance, the two species of Heliomantis appear to under Heliomantis in the catalogue, he marked it with “?”, be very different from each other. H. latipennis differs from indicating his uncertainty. H. elegans in several important morphological characters. It Lombardo (1993) reported H. elegans from Nepal. lacks meso- and metafemoral preapical lobes (Figure 7C) Mukherjee et al. (1995) complemented the description and the fore coxal apical black band (Figure 5). The prono- of H. elegans, provided figures for the male and reported tum (Figure 6D) is shorter and bulkier, with weakly defined the species from additional locations in India. supracoxal extension, not strongly contracting in the meta- Mukherjee et al. (2014) listed additional location for zone. The forewings are lanceolate, barely exceeding the end H. elegans in Nepal. of abdomen, and the hind wings have partially pink colora- Ehrmann and Borer (2015) reported H. elegans from tion (Figure 5). In contrast, in H. elegans the pronotum is Bhutan. slender, with well-developed supracoxal extension and the Svenson et al. (2015) transferred H. elegans and the metazone noticeably constricting in the middle, while the whole genus from Hymenopodidae Acromantinae to forewings are oval and well exceed the end of the abdomen, Hymenopodidae Anaxarchini, placing it sister to the even in female. Other genera of Anaxarchini, Anaxarcha and genus Odondomantis Saussure 1871. Odontomantis, do not have meso- and metafemoral lobes. Species of Anaxarcha have slender pronotum similar to the pronotum of H. elegans (Figure 6A, B), while Odontomantis (except O. rhyssa Werner 1930) has very short pronotum Material and methods with (Figure 6E) or without (Figure 6F) strongly pronounced Abbreviations supracoxal extension; even in the latter case lateral edges of BMNH: The Natural History Museum, London, UK the prozone start to noticeably converge immediately Annales de la Société Entomologique de France (N.S.) 137 anteriad the point of maximum width, which is not the case metafemoral preapical lobes (Figure 7B), strongly curved in H. latipennis. main forewing veins and dense, irregular cross-vein Another notable character lies in female forewing vena- pattern (Figure 8A); from Odontomantis (Figures 6E–G, tion. Most mantises have a quite simple structure of cross- 9A, B) by elongated pronotum with pronounced veins pattern posteriad RP+M: straight or very slightly curved supracoxal extension (Figure 6C) and presence of ventral cross-veins form double rows of more or less rectangular cells meso- and metafemoral preapical lobes (Figure 7B); from between the main veins (Figure 9C;Wieland2013, fig. 334). Werneriana n. gen.(Figures 5, 6D, 7C, 8B) by elongated In other instances, the cross-veins, while still nearly perpendi- pronotum with pronounced supracoxal extension cular to the main veins at their base, form irregular cells, (Figure 6C), presence of ventral meso- and metafemoral which sometimes can be very dense, resembling archedictyon preapical lobes (Figure 7B), long wings and irregular cells (Figures 8A, 9A). The cells themselves are generally not in female forewing venation (Figure 8A); from elongated in any particular direction. The third state, resem- Oligomantis (Figure 9C) by dense forewing venation bling at first glance the second one, has in fact a totally (Figure 8A) and more dorsoventrally compressed different structure: dense cross-veins form elongated, mostly pronotum; from Psychomantis by absence of median longitudinal cells, or longitudinal themselves (Figure 8B, C). process on the head vertex (Figure 4A, B), more Females of most Hymenopodidae, including the majority dorsoventrally compressed pronotum, smooth surface of of Acromantinae genera, are characterized by the first state pronotum, straight dorsal edge of fore femora and only (e.g. Oligomantis, Figure 9C). The second state is rarer and one ventral lobe on middle and hind femora (Figure 7B); canbeobservedinmostHymenopodini,Psychomantis from Rhomantis by absence of median process on the head Giglio-Tos 1915 and in considerable number of species of vertex (Figure 4A, B), shorter and more dorsoventrally Acromantis Saussure 1870. Inside Anaxarchini it can be compressed pronotum and dense forewing venation observed in Heliomantis elegans (Figure 8A) and most spe- (Figure 8A); from Acromantis (Figure 7A) and cies of Odontomantis (e.g. O. foveafrons Zhang 1985, Citharomantis by rounded apeces of hind wings (Figures Figure 9A), the only two exceptions to our knowledge 1–3); from Ambivia by absence of median process on the being O. planiceps (De Haan 1842) possessing the first head vertex (Figure 4A, B), smooth surface of pronotum, state (the common “double rows” pattern), and O. rhyssa, straight dorsal edge of fore femora, only one ventral lobe which has only a single row of cells between each two main on middle and hind femora (Figure 7B) and dense veins (Figure 9B). This state is rarely encountered even out- forewing venation (Figure 8A); from Majangella by side Hymenopodidae. It should be noted that O. rhyssa has absence of median process on the head vertex many other interesting characters separating it from other (Figure 4A, B), smooth surface of pronotum and straight Odontomantis species (e.g. relatively elongated pronotum, dorsal edge of fore femora. densely covered by tubercles, Figure 6G) and probably deserves its own genus. The third state can be found in Systematic position. Giglio-Tos (1927) clearly separated Heliomantis latipennis (Figure 8B) and all species of Acromantes from the other tribes of his Acromantinae, Anaxarcha (Figure 8C) and appears to be unique at least including Anaxarchae, by a presence of preapical lobes among Hymenopodidae and allies, further stressing morpho- on middle and hind femora (Figure 7A), absence of a long logical differences between two species of Heliomantis. median head process, spherical eyes and smooth Based on these differences we suggest to erect a new pronotum. H. elegans (Figures 1–4, 6C, 7B, 8A) genus for H. latipennis, rendering the genus Heliomantis corresponds to this diagnosis of the tribe. Externally, it is monotypic. similar to the acromantine genera Psychomantis, Rhomantis Giglio-Tos 1915 and Oligomantis in slender pronotum (Figure 6C), with metazone strongly narrowing Genus Heliomantis Giglio-Tos 1915 in the middle, and fore coxae with contiguous apical lobes Heliomantis Giglio-Tos 1915: 2; Giglio-Tos 1919: 72, 77–79; and same coloration as in these genera, including a black Beier 1934: 11; Beier 1964: 939; Beier 1968: 6; Ehrmann preapical band (Figures 4A, B). However, it lacks vertex 2002: 169; Otte & Spearman 2005:75–76; Svenson et al. process; prozone of its pronotum is more gently convex, 2015: 23; Ehrmann & Borer 2015: 233. not strongly laterally compressed; its fore femora are – Paraspilota Bolívar 1914: 205 206 (nec Paraspilota Bates without a trace of dorsal lobes, and the meso- and 1888: 374–375). Deiroharpax Werner 1916: 283; Deiroharpax: Giglio-Tos 1927: metafemoral lobes are widely oval (Figure 7B) and not 524; Deiroharpax: Beier 1930: 435 (syn. by Beier 1934: 11). triangular like in the rest of Acromantes (e.g. Figure 7A). Type species. Polyspilota elegans Navás 1904 (by Nevertheless, the position of Heliomantis inside monotypy). Acromantes (= tribe Acromantini) was not disputed by later authors, until recently. Diagnosis. Heliomantis differs from Anaxarcha (Figures Svenson et al. (2015) conducted phylogenetic analyses 6A, B, 7D, 8C) by presence of ventral meso- and of all genera included in Hymenopodidae sensu Ehrmann 138 E. Shcherbakov et al.

Figure 1. Heliomantis elegans (Navás 1904), habitus. A, male, SMNK #12622, dorsally; B, female, SMNK #03625, dorsally; C, female, ventrally; D, male, ventrally. Photos: R. Ehrmann, © SMNK, published with permission. Annales de la Société Entomologique de France (N.S.) 139

2002, , Mantidae: and Mantidae: morphological analysis under maximum parsimony (see Phyllotheliinae, using molecular data from 10 gene frag- Svenson et al. 2015, fig. 6B) as well as in the total ments, as well as morphology. H. elegans was used as the evidence analysis (see Svenson et al. 2015, fig. 7) H. representative of the genus Heliomantis. Since a sample of elegans was recovered deep in Anaxarchini as the sister DNA of H. elegans was unavailable, only morphological taxon to Odontomantis, while Anaxarchini as a clade was data were used to determine its place on the phylogenetic very distant from Acromantinae sensu Svenson et al. tree. In the equally weighted analysis of morphology (2015) in both analyses. The only apomorphy uniting H. under maximum parsimony (see Svenson et al. 2015, fig. elegans and Odontomantis in the analysis is “the costal 6A) H. elegans was recovered in large basal polytomy. area width is 10–24% of the width of discoidal area of Next, Svenson et al. reconstructed ancestral states on the female fore wing” (state 1 of character 61; see Svenson total evidence topology (recovered through Bayesian et al. 2015, table S3), contrary to plesiomorphic state inference with Mk1 model for morphology), identified found in Anaxarcha, where the ratio is supposedly “synapomorphies” of the respective clades and increased 25–50%. However, this character is essentially continuous the weight of corresponding characters in morphological and no argumentation in favour of delineating these char- dataset up to 6, motivating it by poor resolution of the acter states in such a way was provided in the paper. strict consensus tree in case of equally weighted characters Therefore, there is no evidence of the clade uniting (Svenson et al. 2015, p. 12). In the upweighted Heliomantis and Odontomantis.

Figure 2. Heliomantis elegans (Navás 1904), lectotype female in dorsal and ventral perspective and its labels. Photos: M. Paris, © M. Paris, MNCN, published with permission. 140 E. Shcherbakov et al.

Figure 3. Deiroharpax viridis Werner 1916, holotype male in dorsal, lateral and ventral perspective and its labels. Photos: H. Bruckner, © NHMW, NOaS Image Collection, published with permission. Annales de la Société Entomologique de France (N.S.) 141

Figure 4. Heliomantis elegans (Navás 1904), details of morphology. A, anterior part of the male, ventrally; B, anterior part of the female, ventrally; C, asymmetrical variation of the subgenital plate, ventrally; D, male genitalia, preparation of SMNK #12622, ventrally; E, variation of the phalloid apophysis.

Though there is currently no counterevidence to the phylogenetically informative characters, usable for phyloge- monophyly of Anaxarchini including Heliomantis,we netic assessment of Heliomantis and its presumed sister taxa. believe that addition of molecular data from the fresh mate- rial to the analysis is important for reliable phylogenetic – placement of the genus. Moreover, it is desirable to reinves- Heliomantis elegans (Navás 1904)(Figures 1 4, 6C, tigate external morphological characters, as well as the male 7B, 8A) genitalia across Anaxarchini and possibly even Polyspilota elegans Navás 1904: 132–133 (male and female) Hymenopodidae, to ensure correct homologization of all (“Kurseong in Himalaya”; India-NE: West Bangal: Kurseong). 142 E. Shcherbakov et al.

Figure 5. Werneriana latipennis (Werner 1930), holotype female in dorsal and ventral perspectives and its labels. Photos: G. Lindberg, © NRM, published with permission.

Paraspilota elegans (Navás 1904): Bolívar 1914: 206 (female) Mukherjee et al. 2014: 8 (female) (Nepal: Bartār); Ehrmann (India-NE: West Bangal: Kurseong); Giglio-Tos 1919: & Borer 2015: 233 (male and female), fig. 28 (female) 77–79. (India-NE, Nepal, Bhutan); Svenson et al. 2015:5,8, Heliomantis elegans: Giglio-Tos 1915: 2 (male) (India-NE: West 12–14, 23, 26, 39, 43, fig. 17 C (female). Bangal: Kurseong); Giglio-Tos 1919:77–79; Giglio-Tos Deiroharpax viridis Werner 1916: 283–284, fig. 6 (male) (India- 1927: 522; Werner 1930: 6; Beier 1934: 11; Lombardo NE: Sikkim) (syn. by Beier 1934: 11); Giglio-Tos 1927: 524; 1993: 202 (male) (Nepal: Kathmandu Valley); Mukherjee Werner 1930:6. et al. 1995: 219–220, 345, figs 35–37 (males and female) (India-NE: Assam, Sikkim, West Bengal); Ehrmann 2002: Type material. Of two syntypes, male and female, the 169 (male) (India-NE, Nepal); Otte & Spearman 2005: 76; male could not be located. The second syntype, female, is Annales de la Société Entomologique de France (N.S.) 143

Figure 6. Pronotum of female. A, Anaxarcha limbata Giglio-Tos 1915; B, A. acuta Beier 1963; C, Heliomantis elegans (Navás 1904); D, Werneriana latipennis (Werner 1930); E, Odontomantis foveafrons Zhang 1985; F, O. planiceps (De Haan 1842); G, O. rhyssa Werner 1930. Scale bar 1 mm.

Figure 7. Apex of middle femur of female, dorsally, arrow marks the preapical lobe. A, Acromantis sp.; B, Heliomantis elegans (Navás 1904); C, Werneriana latipennis (Werner 1930); D, Anaxarcha limbata Giglio-Tos 1915. 144 E. Shcherbakov et al.

Figure 8. Proximal half of forewings in females, dorsally. A, Heliomantis elegans (Navás 1904); B, Werneriana latipennis (Werner 1930); C, Anaxarcha limbata Giglio-Tos 1915. Part of the cross-veins network highlighted. See the text for details. Annales de la Société Entomologique de France (N.S.) 145

Figure 9. Proximal half of forewings in females, dorsally. A, Odontomantis foveafrons Zhang 1985; B, O. rhyssa Werner 1930; C, Oligomantis mentaweiana Giglio-Tos 1915. Part of the cross-veins network highlighted. See the text for details. 146 E. Shcherbakov et al.

Table 1. Depositaries with known specimens of Heliomantis elegans (Navás 1904).

Depositary Number of specimens Country of origin Previous publications

BMNH 3 males, 1 female India Svenson et al. (2015) MNCN 1 female (lectotype) India Navás (1904) and Bolívar (1914) NHMW 2 males (including the India, India? Werner (1916) holotype of D. viridis) NMB 1 male Bhutan Ehrmann & Borer (2015) SMNK 1 male, 1 female Bhutan, Nepal Mukherjee et al. (2014) and Ehrmann & Borer (2015) ZSI 4 males, 1 female India Mukherjee et al. (1995) ZSM 3 males India, Nepal Lombardo (1993) and Ehrmann & Borer (2015)

deposited in the collection of MNCN under the code prosternum and the pin. Parts of mesothorax, middle and MNCN_Ent140643anddesignatedhereinasthe hind coxae destroyed; right middle leg from the femur lectotype (Figure 2). It is satisfactorily preserved. Head glued to right hind coxa apart from the right middle coxa. without both flagella, with eyes and frons partially Left pair of wings spread; anal fan of the hind wing destroyed. Prothorax with large longitudinal gap dorsally posteriorly torn. The lectotype has following labels: and ventrally. Distal parts of the fore coxae glued to the handwritten “Polyspilota elegans + Nav. Kurseong

Figure 10. Distribution of Heliomantis elegans (Navás 1904) and Werneriana latipennis (Werner 1930). Annales de la Société Entomologique de France (N.S.) 147

(Him.) Juno 1900”, handwritten “Paraspilota g. n.”, short spines. Dorsal margin of femora straight or very printed “Sintipo MNCN Tipos № 2614”,printed slightly concave in the middle; ventral margin with “MNCN_Ent 140643” (Figure 2). 13–14 alternating in length anteroventral spines, 4 poster- Re-examination of the holotype of Deiroharpax viridis oventral spines, 4 discoidal spines and small genicular – Werner 1916 (Figure 3) confirms its synonymy with H. spines on each side. Tibiae with 11 12 anteroventral – elegans. spines and 9 10 posteroventral. Posteroventral tibial spines strongly inclined, but their bases clearly separated Material examined. Nepal: 1 ♀, Bartār, 28.07°N 84.24°E, from each other. Middle and hind femora with small, very 2150 m, light trap, 7–29.VI.1988, leg. G. Lebisch & H. Probst elongated preapical lobe (Figure 7B). Middle and hind (SMNK #03625); 1 ♂, Kathmandu Valley, Godavari, tibiae shorter than femora, apically with triangular elonga- 1600–1800 m, 6.VI.1967, leg. W. Dierl & W. Schacht (ZSM). tion and two short spines. Hind metatarsus slightly longer Bhutan: 1 ♂, 21 km E Wangdi Phodrang, 1700–2000 m, 15. than all other tarsomeres combined. VI.1972, leg. C. Baroni Urbani, O. Stemmler, W. Wittmer & M. Both pair of wings far surpassing the end of abdomen Würmli [Nat.-Hist.Museum Basel Bhutan Expedition 1972] (Figure 1B, C). Forewings wide, gradually narrowing and (NMB #1); 1 ♂, same location and date (SMNK #12622) (ex narrowly rounded at apex. Costal field wide, main veins coll. NMB, #2). strongly curved posteriorly, venation between them very ♂ India: 2 , West Bengal, Darjeeling env. (ZSM). dense, irregular (Figure 8A). Stigma very elongated, not There are very few specimens in the collections worldwide: well defined. Hind wings slightly shorter than forewings, only 18 are known to the authors, including cited in various wide, more or less triangular. Anterior edge curved poster- literature (Table 1). iorly near apex, apex narrowly rounded. Redescription. Female (Figures 1B, C, 2, 4B, 6C, 7B, Abdomen wide, tergites and sternites simple, without 8A). Medium sized (body length 47–54 mm). lobes. Anal plate transverse, widely rounded. Cerci circu- lar in cross-section, simple. Head triangular. Vertex straight, smooth, without any Main life colour green. Anterior surface of fore coxae with process. Parietal furrows deep, but lateral tubercles absent. black band in apical third, not reaching the end of apical lobes. Ocelli small, with diameter less than diameter of scapus, Anteriorsurfaceofforefemorawith irregular black spot at the forming triangle, not elevated. Scutellum transverse, pen- base of first two anteroventral spines, almost reaching the tagonal, nearly smooth, anterior margin concave, posterior middle of femora’s width. Forewings subopaque with yellow- margin forming wide, triangular medial process projecting ish costal field. Wings transparent, hyaline with yellowish anteriorly. Clypeus rectangular. Labrum almost as wide as main veins, except yellowish-brownish costal field. clypeus, widely rounded at apex. Male (Figures 1A, D, 3, 4A, C–E) very similar to Pronotum (Figure 6C) moderately slender, with dis- female, but smaller (body length 32–37 mm). Comparing tinct oval, elongated supracoxal dilatation, dorsally to female, eyes bulging, pronotal and coxal spines rela- smooth. Prozone narrowing anteriorly, with rounded ante- tively shorter, femoral spot less distinct. Anal plate rior edge, dorsally slightly bulged, with shallow median slightly more triangular than in female, cerci with 13–14 groove. Prozone separated from metazone by distinct cercomeres. Genital plate wide, in some specimens more supracoxal sulcus, medially straight, laterally curved or less semicircular, in others unusually asymmetrical – toward the anterior. Metazone 2.6 2.7 times longer than (Figure 4C), with small and short styli. Male genitalia prozone, constricted posteriad supracoxal dilatation, with (Figure 4D) weakly sclerotized, similar in shape and struc- edges parallel and diverging again only near posterior ture to Anaxarcha. Sclerite L4A elongated, without any edge. Dorsal surface of metazone medially with very kind of process. Sclerite L4B very narrow and elongated. ’ shallow groove and low, short, less than 1/3 of metazone s Sclerite L2 mostly membranous, its titillator short and length, carina inside the groove. Lateral margins of pro- rounded. Sclerite L1 anteriorly curved, very long and notum slightly lamellar, in prozone covered by short, blunt narrow, posteriorly (phalloid apophysis) sclerotized, more spines, in supracoxal dilatation almost smooth, with only a or less rounded, with little variation of shape (Figure 4E). few barely noticeable low tubercles, in metazone covered Sclerite R1A triangular, its apex abruptly curved dorso- by very spaced, long, apically rounded tubercles. ventrally. Sclerite R3 in distal 2/3 widened, more or less Forelegs typical of praying mantises (Figure 4B). rectangular in shape. Coxae slightly shorter than metazone of pronotum, trian- gular in cross-section, with anterior apical lobes adjacent. Measurements (in mm). Total length male 32–37, female Dorsal margin with 7–9 relatively large denticles inclined 47–54; head width male 5.0, female 6.9; pronotum length toward the apex of coxa, with 1–3 smaller denticles male 11–12, female 16; metazone length male 7.2, female between them. Ventral margin with numerous small, very 11.0; pronotum maximum width male 3.6–4.0, female 5.2– 148 E. Shcherbakov et al.

5.5; fore coxa male 7.8, female 11.0; fore femur male 9.1–9.2, Redescription. Female (Figure 5). Medium sized (body female 12.0; fore wing length male 37–39, female 48.0–48.4; length 29 mm). – fore wing width male 10.0 10.1, female 14.3. Head triangular. Vertex slightly convex, without process. Posterior lateral tubercles between compound eyes and par- Distribution. India-NE, Nepal, Bhutan (Figure 10). ietal furrows small, but noticeable. Between vertex bump Apparently the species prefers mountain forests: there are and each parietal furrow noticeable depression, thus five no specimens with recorded elevation lower than 1600 m, small bumps in total between compound eyes. Ocelli small. – while some were collected as high as 2000 2150 m. Scutellum (without apical spine) strongly transverse, anterior border concave, posterior border very strongly angular, its middle extended into a narrow spine with rounded apex. Genus Werneriana n. gen. Prothorax (Figure 6D) elongated. Length of pronotum Type species. Heliomantis latipennis Werner 1930. 2.28 times its maximum width. Anterior edge of prozone widely rounded. Metazone of pronotum 2.2 times the Diagnosis. Werneriana n. gen. differs from Heliomantis length or prozone, separated from the latter by shallow (Figures 1–4, 6C, 7B, 8A) by shorter, much more robust supracoxal sulcus. Supracoxal dilatation weak, widely pronotum with much less pronounced supracoxal dilatation oval. Pronotum dorsally with a median groove, covered (Figure 6D), by absence of ventral lobes on middle and hind by low tubercles in prozone, otherwise smooth. Lateral femora (Figure 7C), by much shorter, lanceolate forewings, edges of pronotum covered by large obtuse tubercles, barely covering the apex of abdomen (Figure 5)andinfemale except anterior and posterior edges; in prozone tubercles by very elongated cells in forewing venation (Figure 8B); short, stout, densely arranged, in supracoxal dilatation from Anaxarcha (Figures 6A, B, 7D, 8C) by shorter, much very small and flat, in metazone longer, rounded and more robust pronotum with much less pronounced supracoxal more spaced than in prozone. dilatation (Figure 6D), by much shorter, lanceolate forewings, Forelegs typical of praying mantises. Coxa longer than barely covering the apex of abdomen (Figure 5) and by hind metazone of pronotum; anterior apical lobes contiguous; wings clearly shorter than forewings; from Odontomantis dorsal edge with numerous short, sharp, irregularly alternat- (Figures 6E–G, 9A, B) by almost parallel edges of ing larger and smaller spines; ventral edge with very short supracoxal extension in prozone (Figure 6D), by narrow sharpened tubercles. Femur wide, with dorsal edge straight. separation of ScP and RA and very elongated cells in female Anteroventral row with 14 spines total plus genicular spine, forewing venation (Figure 8B) and by complete absence of first 10 spines in the row long and short, alternating, penulti- dark pattern on hind wings of females (Figure 5). mate 3 spines short, last spine long; posteroventral row with 4 spines plus genicular spine, most distal spine a little shorter Systematic position. The genus fits the diagnosis of than others; discoidal row with 4 spines, the 1st much shorter Anaxarchini. Unique character of forewing cross-vein than the rest, the 2nd as the 4th, the 3rd more than twice the pattern shared with Anaxarcha suggest that the latter is a 2nd. Claw furrow in proximal half of fore femur. Tibia sister-group of Werneriana n. gen. However, as in the case approximately half the length of fore femur, anteroventral of Heliomantis, a thorough study of morphology row with 12 distally elongating spines, posteroventral row (including male genitalia) as well as obtaining DNA with 11 strongly inclined and packed spines plus very long sample is necessary to test this hypothesis. and curved tibial claw. Middle and hind legs short, simple. Ventral edges Etymology. The genus is named in honour of the author of middle and hind femora slightly lamellar of its type species, Austrian zoologist Franz Werner. The (Figure 7C), each apex of femur with very short geni- name gender is feminine. cular spine. Monotypic, therefore the generic characteristics are Forewings barely cover end of abdomen, wide, lan- those of the type species. ceolate, with narrowly rounded apex and very dense vena- tion (Figure 8B). Jugal area very small. Stigma small, very Werneriana latipennis (Werner 1930), n. comb. (Figures elongated, highly indistinct. Hind wings shorter than 5, 6D, 7C, 8B) forewings, anterior edge curved near apex. Heliomantis latipennis Werner 1930: 7, Taf. 2, fig. 2 (female) Abdomen wide. Cerci simple, circular, with 12 notice- (Borneo: Sarawak); Sjöstedt 1930: 13; Beier 1934:11; able cercomeres. Ehrmann 2002: 169; Otte & Spearman 2005: 76; Schwarz Coloration. Overall life colour green. Forelegs unicolor, & Konopik 2014: 142. except for completely black large femoral spines and small femoral and all tibial spines with dark-brown apices. Type material. Holotype: ♀, Malaysia: Borneo, Sarawak, Forewings greenish, semi-transparent, veins green, costal Murad Mt., 6500 f, 17.XI, Mjöberg leg. (NRM-MANT field and cells in proximal half of discoidal field light-brown 0002699) Annales de la Société Entomologique de France (N.S.) 149 in preserved specimen. Hind wings transparent, brownish Progreso de la Ciencias Congress. 5 (Sec. 4a). Madrid: AEPC; along anterior edge, bright rose centrally. p. 205–214. – Male not known. Ehrmann R. 2002. Mantodea Gottesanbeterinnen der Welt. Münster: Natur und Tier-Verlag; p. 519. Measurements (in mm). Total length 29, pronotum Ehrmann R, Borer M. 2015. Mantodea (Insecta) of Nepal: an anno- length 8, metazone length 5.5, maximum width 3.5, fore tated checklist. In: Hartmann M and Weipert J, editors. Biodiversität und Naturausstattung im Himalaya V. Erfurt: femur 8.5, forewing length 18, width 6.5. Verein der Freunde und Förderer des Naturkundemuseums Erfurt e.V.; p. 227–274. Distribution. The only known specimen is the holotype Giglio-Tos E. 1915. Mantidi esotici. VIII. – Acromantinae. female, collected by Eric Mjöberg 85 years ago on the Bollettino dei Musei di Zoologia ed Anatomia Comparata highest mountain of Sarawak, Mt Murud (Gunung Murud, della Reale Universita di Torino. 30:l–l6. 2423 m), at the elevation of 1830 m (Figure 10). No new Giglio-Tos E. 1919. Saggio di una nuova classificazione dei specimens were reported since then (Schwarz & Konopik mantidi. Bullettino della Società Entomologica Italiana. 49:50–87. 2014). Given the short wings and high elevation at which Giglio-Tos E. 1927. Das Tierreich. 50. Lfg. Orthoptera Mantidae. the specimen was collected, it may be hypothesized that this Berlin & Leipzig: Walter de Gruyter & Co.; p. XL + 707. species (and, probably, the genus as well) is endemic to [ICZN] International Commission on Zoological Nomenclature. Borneo.Presently,themountain is part of Pulong Tau 1999. International code of zoological nomenclature. 4th ed. National Park. London: The International Trust for Zoological Nomenclature; p. 306. Klass K-D. 1997. The external male genitalia and the phylogeny of Blattaria and Mantodea. Bonner Zoologische Monographien. Acknowledgements Bd. 42. Bonn: Zoologisches Forschungsinstitut und Museum We wish to sincerely thank Gunvi Lindberg (NRM) for providing the Alexander Koenig; p. 341. photographs of the holotype of H. latipennis, Mercedes París Lombardo F. 1993. Studies on the Mantodea of Nepal (Insecta). – (MNCN) for the photographs of the female syntype of H. elegans, Spixiana. 16:193 206. Mercedes París, Berta Caballero and Gloria Maso (Natural History Mukherjee TK, Ehrmann R, Chatterjee P. 2014. Checklist of Museum of Barcelona) for their efforts in search of the syntypes of H. Mantodea (Insecta) from India. Priamus (Serial Publication – elegans, Tushar Mukherjee (ZSI) for sending the pictures of H. of the Centre for Entomological Studies Ankara). 30:1 61. elegans specimens from the collection of ZSI, Harald Bruckner Mukherjee TK, Hazra A, Ghosh A. 1995. The mantid fauna of – (NHMW) for the photographs of D. viridis type and another male India (Insecta: Mantodea). Oriental . 29:185 358.

– of H. elegans. We also wish to thank Gavin Svenson (Cleveland Navás RPL. 1904. Notas Zoologicas. III. Algunos Insectos de Museum of Natural History) for sending us the morphological data- Kurseong en la cordillera del Himalaya. Boletin de la – set, Pavel Udovichenko (Moscow), Alexei Klimenko (Tver), Viktor Sociedad Aragonesa de Ciencias Naturales. 3:130 139. ’ Sinyaev (Moscow) and Mikhail Vasiliev (Moscow) for the additional O Hanlon JC, Holwell GI, Herberstein ME. 2014. Pollinator material used for comparison. We are grateful to Martin Stiewe deception in the Orchid Mantis. The American Naturalist. – (BMNH) for providing information about specimens of H. elegans 183:126 132. fi in BMNH, and to Christian J. Schwarz (Ruhr University Bochum, Otte D, Spearman L. 2005. Mantida species le. Catalog of the Bochum) for the valuable discussions regarding the taxonomy of Mantids of the world. Philadelphia (PA): Association of the Hymenopodidae. We thank Vladimir Savitsky (Lomonosov Moscow Diversity; 489 p. State University) and the two anonymous reviewers, who made Schwarz CJ, Konopik O. 2014. An annotated checklist of the valuable comments about earlier version of this work. praying mantises (Mantodea) of Borneo, including the results of the 2008 scientific expedition to Lanjak Entimau Wildlife Sanctuary, Sarawak. Zootaxa. 3797:130–168. References Shcherbakov EO, Savitsky V. 2015. New data on the fauna, taxon- Bates HW. 1888. On a collection of coleoptera from Korea omy and ecology of praying mantises (Dictyoptera, Mantodea) (Tribes Geodephaga, Lamcllicornia, and Longicornia), from Russia. Russian Journal of Zoology. 94:37–55. made by Mr. J. H. Leech, F.Z.S. Proceedings of the Sjöstedt Y. 1930. Orthopterentypen im Naturhistorischen Zoological Society of London. 56:367–380. Reichsmuseum zu Stockholm. Arkiv för Zoologi. Beier M. 1930. XLVII.—new and rare Mantodea (Orthoptera) in 21A:1–43. the British museum. Journal of Natural History Series 10. Svenson GJ, Hardy NB, Wightman H, Wieland F. 2015.Of 6:432–460. flowers and twigs: phylogenetic revision of the plant- Beier M. 1934. Mantodea, Fam. Mantidae. Subfam. mimicking praying mantises (Mantodea: Empusidae and Hymenopodinae. Genera Insectorum. Fascicule. 196:1–37. Hymenopodidae) with a new suprageneric classification. Beier M. 1964. Blattopteroidea. Mantodea. In: Bronn HG, editor. Dr. Systematic Entomology. 40:789–834. H. G. Bronns Klassen und Ordnungen des Tierreichs. Werner F. 1916. Zur Kenntnis afrikanischer und indischer Arthropoda. III. Insecta, 6 (5). Leipzig: Geest & Portig K.-G; Mantodeen. Verhandlungen der Zoologisch-Botanischen p. 850–970. Gesellschaft in Wien. 66:254–296. Beier M. 1968. Mantodea (Fangheuschrecken). In: Helmcke J-G, Werner F. 1930. Über asiatische Mantidae aus dem naturhistor- Stark D, Wermuth H, editors. Handbuch der Zoologie, eine ischen Reichsmuseum in Stockholm. Arkiv för Zoologi. aturgeschichte der Stämme des Tierreiches, 4 (2), Teil 2/12. 21A:1–10. Berlin: W. de Gruyter; p. 1–47. Wieland F. 2013. The phylogenetic system of Mantodea (Insecta: Bolívar I. 1914. Examen de un pequeno lote de Ortópteros de Dictyoptera). Species, Phylogeny and Evolution. 3:3–222. Mindanao y del Himalaya. In: Asociacion Espanola para el