Zootaxa 4341 (4): 528–538 ISSN 1175-5326 (print edition) http://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2017 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4341.4.5 http://zoobank.org/urn:lsid:zoobank.org:pub:0DA889AB-FD1A-403F-932D-BD0298A78251 (Diaulaconia) artemis sp. n., a new termitophilous species from Taiwan (Coleoptera: Staphylinidae: : )

WEI-REN LIANG1, MUNETOSHI MARUYAMA2,3 & HOU-FENG LI1,4 1Department of Entomology, National Chung Hsing University, 145 Xingda Rd., Taichung, 40227, Taiwan. E-mail: [email protected] 2The Kyushu University Museum, Fukuoka, Japan. 3Corresponding author. E-mail: [email protected] 4Corresponding author. E-mail: [email protected]

Abstract

The staphylinid tribe Lomechusini is known for the diverse associations with ants or termites. A new species of rove belonging to Zyras (Diaulaconia) is found to be associated with the fungus-growing termite, Odontotermes formosanus (Shiraki, 1909), and is showing meticulous predatory behavior. In the present study, this species is described as Zyras (Di- aulaconia) artemis Liang, Maruyama & Li, sp. n. Its assignment to the subgenus Diaulaconia is based on the comparison of female spermatheca, male aedeagus, and male secondary characters. The habitus photographs and the line drawings of diagnostic characters and their variations are provided. The current knowledge about the , phylogeny, and ecol- ogy of Diaulaconia is discussed. The checklist of Diaulaconia species of Taiwan is updated.

Key words: , spermathecal variation, synechthrans, termitophagous, Odontotermes formosanus

Introduction

Members of the tribe Lomechusini Fleming, 1821 (Staphylinidae: Aleocharinae) have been subject of biological interest for a long time because of their diverse associations with ants or termites (Hlaváč et al., 2011). A , Zyras species belonging to Lomechusini, was observed in the wood and litter of foraging sites of the fungus-growing termite, Odontotermes formosanus (Shiraki, 1909), and was occasionally accompanied with the termite dead bodies. This species immediately aroused our interest, and several behavioral experiments were conducted. The beetle shows an extraordinary predatory behavior and unique life history (Liang et al., in prep.). This species is a member of the subgenus Diaulaconia. Bernhauer (1928) established the subgenus Diaulaconia of the Zyras, without type designation, and included four species, Zyras biseriatus Bernhauer, 1915, Z. compressicornis Fauvel, 1905, Z. diversiventris Bernhauer, 1915, and Z. roepkei Bernhauer, 1914. According to Bernhauer (1928), Diaulaconia was distinguished from the subgenus Parophthalmonia Bernhauer, 1928 by its more arched body shape, the lack of two strong longitudinal furrows beside the middle zone of the pronotum, and the lack of lateral longitudinal furrows on the seventh tergite and the presence of punctures behind the frontal hunch at the head in males. Diaulaconia can also be separated from other subgenera of Zyras by the laterally compressed antennae. After that, Bernhauer (1929, 1939) described two species from China. Cameron (1946) described one species from Thailand. Later, Blackwelder (1952) designated Z. biseriatus as the type species of the subgenus Diaulaconia. Pace (1993, 1998, 2001, 2003, 2004, 2005, 2008, 2009, 2010, 2013) added 15 species to Diaulaconia and gave the key to the oriental species of compressicornis group (Pace 2003). Hlaváč (2005) redescribed Z. orientalis and discussed the taxonomic status of Diaulaconia. So far, the subgenus Diaulaconia contains 23 species distributed in Southeast Asia except one species described by Pace (2009) from Bolivia, South America (probably mis-affiliation).

528 Accepted by J. Klimaszewski: 28 Aug. 2017; published: 2 Nov. 2017 In the present study, the taxonomic status of the Zyras (Diaulaconia) sp. n. from Taiwan was examined based on the comparison of morphology among subgenus Diaulaconia. Taxonomy of the subgenus Diaulaconia and distribution of two species, Z. compressicornis and Z. roepkei, in Taiwan, are discussed.

Materials and methods

The specimens were examined under Leica® M205 C stereomicroscope and Leica® DM750 microscope with the drawing tube and measured by using Leica MC170 HD digital camera with the software LAS (Leica Application Suite Version 4.4.0, Leica, Wetzlar, Germany) on the computer monitor. The photographs were taken by Leica MC170 HD digital camera or Canon EOS 760D with a micro lens (Canon Inc., Tokyo, Japan), and then combined by the automontage software CombineZP (Hadley 2010). The methods for dissection and making permanent slides were followed Maruyama (2006). The following abbreviations are used for measurements: AL: antennal length; BL: body length (approximate whole length); EL: eye length (major axis); ELL: elytra length; ELW: elytra width; FBL: fore body length (apex of head to apices of elytra); HL: head length (apex of head to apices of head capsule, except for preocciput); HTL: hind tibial length; HW: head width (maximal, including eyes); PL: pronotal length; PW: pronotal width. All measurements are in millimeters.

Abbreviations for depositories:

KUM Maruyama collection at the Kyushu University Museum, Fukuoka, Japan; NCHU NCHU Termite collection, Department of Entomology, National Chung Hsing University, Taichung, Taiwan; NMNS National Museum of Natural Science, Taichung, Taiwan; NTU Museum of National Taiwan University, Taipei, Taiwan; TARI Taiwan Agriculture Research Institute, Taichung, Taiwan.

Taxonomy

Genus Zyras Stephens

Zyras Stephens, 1835: 430. Type species: Aleochara haworthi Stephens, established by decision of the International Commission on Zoological Nomenclature (ICZN., 1961).

Subgenus Diaulaconia Bernhauer

Zyras (Diaulaconia) Bernhauer, 1928: 73. Type species: Zyras biseriatus Bernhauer, designated by Blackwelder (1952).

Zyras (Diaulaconia) artemis Liang, Maruyama & Li, sp. n. (Figs 1–6)

Type materials (163 specimens). Holotype: male, Taiwan: Pingtung Co.: Sheding: 21.9646˚N; 120.8270˚E, 165m, 18.XI.2015, by high-intensity discharge lamp (HID light trap), W.-R. Liang & Y.-C. Lan, deposited in NMNS. Paratypes: Hualien Co.: Danongdafu: 23.6185˚N; 121.4009˚E, 160m, 20.X.2016, by ultraviolet light trap (UV light trap), W.-R. Liang & Y.-Z. Huang, 1♂ (dissected) (NMNS); Hsilin trail: 23.8275˚N, 121.4250˚E, 270m, 19.X.2016, by UV light trap, W.-R. Liang & Y.-Z. Huang, 1♀ (dissected) (NCHU). Nantou Co.: Xiaping: 23.7759˚N, 120.6720˚E, 150m, 25.IV–30.V.2015, by flight interception trap (FIT), W.-R. Liang, 1♂, 1♀ (all dissected) (KUM); 23.VI–15.VII.2015, by FIT, 3♂♂ (NMNS); 28.I.2016, near the foraging site of O. formosanus in the rotten wood, 1♂, 1♀ (NCHU); 28.VIII.2016, 1♂ (NMNS); 23.7760˚N, 120.6723˚E, 147m, 15.VII–

NEW SPECIES OF TERMITOPHILOUS ROVE BEETLE Zootaxa 4341 (4) © 2017 Magnolia Press · 529 2.VIII.2015, by UV light trap, 4♂♂, 5♀♀ (NCHU); 23.7748˚N, 120.6722˚E, 149m, 15.VII–2.VIII.2015, by UV light trap, 10♂♂, 11♀♀ (KUM); Shueili, 23.7661˚N, 120.8560˚E, 404m, 29.IV–16.V.2016, by Malaise trap, H.-T. Yeh, 1♂, 3♀♀ (NTU); 16.V–1.VI.2016, 1♂, 2♀♀ (KUM); 1.VI–15.VI.2016, 2♀♀ (NTU); Qingshuigou: 23.7615˚N, 120.7927˚E, 434m, 29.IV–16.V.2016, by Malaise trap, H.-T. Yeh, 5♀♀ (KUM); 16.V–1.VI.2016, 1♂ (KUM); 1–15.VI.2016, 2♂♂, 1♀ (KUM); 30.VI–1.VII.2016, 1♂ (NTU); 2–17.VIII.2016, 1♀ (NTU). Penghu Co.: Penghu Islands [Peng Hwu Isls.], 4–13.VI.1948, C.-W. Chen & L-T. Chen, 8♂♂, 7♀♀ (2♂♂, 2♀♀ dissected) (TARI). Pingtung Co.: same data as holotype, 7♂♂, 6♀♀ (4♂♂, 4♀♀ dissected) (KUM, NCHU, NMNS); Sheding: 21.9613˚N; 120.8236˚E, 193m, 24.VIII.2015, by sifting from the foraging site of O. formosanus in the leaf litter, Naomichi Tsuji & W.-R. Liang, 9♂♂, 8♀♀ (1♂, 3♀♀ dissected) (KUM, NCHU, NMNS); Dahanshan: 22.4073˚N; 120.6644˚E, 463m, 21.XI.2015, by UV light trap, W.-R. Liang, 1♀ (dissected) (NCHU); Formosa: Kuraru [Kueitzuchiao], 21–25.VIII.1932, Y. Miwa, 1♀ (TARI). Taichung City: Dakeng: 24.1758˚N; 120.7776˚E, 378m, 19.VIII.2016, by UV light trap, W.-R. Liang, 1♂, 3♀♀ (all dissected) (NCHU); 27.VIII.2016, 3♂♂, 1♀ (all dissected) (NCHU); Wanfeng Hill, X.1984, K.-S. Lin & K.-C. Chou, by Malaise trap, 9♂♂, 9♀♀ (TARI); Hsinshe, 5–8.XII.1955, 1♂, 2♀♀ (TARI). Taitung Co.: Basian Cave: 23.3995˚N, 121.4788˚E, 43m, 21.X.2016, near the foraging site of O. formosanus in the rotten wood, W.-R. Liang & Y.-Z. Huang, 3♂♂ (1 dissected) (NCHU); Taitung [Formosa: Taito], 25.II–27.III.1919, S. Inamura, J. Sonan, and M. Yoshino, 1♂ (dissected), Zyras compressicornis (det. T. Shiraki), Zyras artemis (det. W.-R. Liang, 2017) (TARI); Luanshan: 22.9139˚N, 121.1839˚E, 605m, 14.XI.2015, by UV light trap, W.-R. Liang, 8♂♂, 10♀♀ (2♂♂, 2♀♀ dissected) (KUM, NCHU, NMNS). Taoyuan Co.: Gaupo: 24.7634˚N, 121.3486˚E, 514m, 12.IV.2016, near the foraging site of O. formosanus in the rotten wood, W.-R. Liang & Y.-Z. Huang, 3♂♂, 1♀ (1♂, 1♀ dissected) (NCHU). Description. Body (Fig. 1) slender. Head dark blackish-brown. Antennae, pronotum, elytra, and legs orange to reddish-brown. Abdomen blackish-brown with posterior margins of 3rd to 7th segments dark reddish-brown. Head capsule widest behind eyes, surface finely wrinkled, moderately punctate with short setae. Eyes large and laterally prominent, length of eyes 2.08–2.17 times as long as head width. Antennae (Fig. 2B, C) slightly exceeding combined length of head and pronotum; 1st antennomere longest, longer than 2nd and 3rd combined; 2nd antennomere shorter than 3rd and narrowest, 3rd antennomere longer and slightly narrower than 4th; 4th to 10th antennomeres similar in shape and length; 11th pointed and approximately as long as 3rd; relative lengths of antennomeres: 26:10:15:10:10:10:10:10:11:11:18. Pronotum (Fig. 2A) subtrapeziform with anterior and posterior margin smoothly convex; 1.2 times as wide as long, widest just behind anterior margin, narrowed posteriorly; surface moderately punctate with short and light colour setae, with 4 black macrosetae on pronotal disc and with 40-50 black macrosetae along the whole margin of disc. Scutellum slightly wrinkled and hairy. Elytra (Fig. 2A) slightly widened apically, surface densely punctate with short and light coloured setae. Elytron with about 20 black macrosetae on main surface and 15 black macrosetae on the lateral margin. Legs hairy, hind tibia approximately as long as elytra; approximate relative lengths of tarsomeres: 14:14:12:29 in fore tarsi; 24:17:13:13:27 in mid tarsi; 30:19:16:15:28 in hind tarsi. Abdomen subparallel-sided, nearly as wide as elytra; 3rd to 7th tergites glabrous except for 1–2 rows of setae along each posterior area; 9th tergite with 3 macrosetae; 10th tergite with 6–8 macrosetae. Male. Frons of head with minute granulations (Fig. 2E). 4th tergite with a pair of projections on posterior margin (Fig. 3A, C); these projections apically bilobed in lateral view in large individuals (Fig. 3B), while in small individuals can be much reduced, not bilobed apically (Fig. 3D). 7th tergite with apical median tubercle (Fig. 3E, F). 8th tergite (Fig. 4A) with posterior margin emarginate and forming pair of projections, with about 20–30 macrosetae; posterior to posterolateral margin fringed with thick and light colour sensory setae. 8th sternite (Fig. 4B) with posterior margin slightly emarginate or rounded, with about 50 macrosetae; thick and light colour sensory setae fringed densely on posterior to posterolateral margin. Aedeagus (Fig. 4C, D) elongate pear-shaped in ventral view, pointed at apex; apical lobe with two lateral projections of internal sac, these projections are slender and curved ventrally (Fig. 4E, F); apical sclerite and copulatory piece of inner sac as in Fig. 4C–F. Female. Frons of head smooth, lacking granulation (Fig. 2D). 3rd to 7th tergite simple, lacking projection or small median tubercle on posterior margin (Fig. 3G). 8th tergite (Fig. 5A) similar to male, slightly shorter and wider than in male. 8th sternite (Fig. 5B) with posterior margin rounded, with about 50 macrosetae; thick and light colour sensory setae fringed sparsely on posterior and posterolateral margin. Spermatheca (Fig. 5C–F, 6) anfractuous, basal part highly variable in shape, at least 10 times as long as apical part, rounded and slightly emarginate at base (Fig. 5C, F), widest at base and suddenly constricted at basal 1/3, inner wall smooth; approximate median 1/3 of

530 · Zootaxa 4341 (4) © 2017 Magnolia Press LIANG ET AL. basal part thinnest, anfractuous and varied in shape (Fig. 6), inner wall smooth; apical 1/3 of basal part slightly curved, the inner wall unsmooth (Fig. 5C, E); connection of basal part and apical part bended about 90 degrees; apical part short, its inner wall curved apicad (Fig. 5C, D).

FIGURE 1. Habitus of Zyras (Diaulaconia) artemis sp. n., male, dorsal view.

NEW SPECIES OF TERMITOPHILOUS ROVE BEETLE Zootaxa 4341 (4) © 2017 Magnolia Press · 531 FIGURE 2. Zyras (Diaulaconia) artemis sp. n. A, forebody, dorsal view; B, right antenna, lateral view; C, ditto, dorsal view; D, frons of the female without granulations; E, frons of the male with granulations.

Measurements. BL ≈ 6.9–8.8; FBL ≈ 1.7–2.1; HL: 0.84–0.93; HW: 1.18–1.35; EL: 0.55–0.63; AL: 2.20–2.59; PL: 1.18–1.30; PW: 1.38–1.57; ELL: 1.43–1.61; ELW: 1.72–2.07; HTL: 1.23–1.49 (mm). Diagnosis. Zyras artemis sp. n. is distinguishable from all other Zyras (Diaulaconia) species by its remarkable shape of the spermatheca, which basal part anfractuous diversely and at least 10 times as long as apical part, as well as by the shape of the aedeagus and unique projections of 4th tergite in males. Etymology. Artemis is the ancient Greek goddess of hunting, wildlife, and childbirth. She is currently seen as the archetype of feminists (Bolen, 2014). The new species is named after Artemis in reference to its predatory behavior, extraordinary reproductive strategy, and in honor of Taiwan’s great progress in gender equality, being the first Asian country legalizing same-sex marriage. Biological notes. Zyras artemis sp. n. is found to be the obligatory predator of O. formosanus in Taiwan. We also observed its complete life cycle in the laboratory, and confirm its non-feeding larval development. The detail information will be published in other subsequent studies.

532 · Zootaxa 4341 (4) © 2017 Magnolia Press LIANG ET AL. FIGURE 3. Zyras (Diaulaconia) artemis sp. n. A, male 4th tergite with a pair of projections on posterior margin, large individual, dorsal view; B, ditto, projection bilobed apically, lateral view; C, ditto, small individual, dorsal view; D, ditto, projection with no bilobed apically, lateral view; E, male 7th tergite with apical median tubercle, dorsal view; F, ditto, lateral view; G, female 7th tergite with no small median tubercle, dorsal view.

Distribution. Zyras artemis sp. n. is commonly distributed in low altitude from 0 to 600m around Taiwan, where it is matching the main habitat of its host O. formosanus.This distribution pattern can be referred to their obligatory relationship, which O. formosanus as the only or main food resource for Z. artemis. There are several small islands around Taiwan, e.g., Lanyu Island, Green Island, and Penghu Islands. Up to the present, Z. artemis has been only recorded in Penghu Islands, where O. formosanus is commonly found, but so far it has not been found on the other two islands, where O. formosanus has been also absent.

NEW SPECIES OF TERMITOPHILOUS ROVE BEETLE Zootaxa 4341 (4) © 2017 Magnolia Press · 533 FIGURE 4. Zyras (Diaulaconia) artemis sp. n., male. A, eight abdominal tergite, dorsal view; B, 8th abdominal sternite, ventral view; C, median lobe of aedeagus, ventral view; D, ditto, the photograph; E, ditto, lateral view; F, ditto, the photograph.

Discussion

Taxonomy, systematic position, and ecology of the subgenus Diaulaconia. Bernhauer (1928) established the subgenus Diaulaconia and compared it with subgenus Parophthalmonia Bernhauer, 1928, which is well-defined by having two special bristle-bearing pores, named “setiferous craters” by Last (1958), on the head between the antennae in males. In addition to that, Diaulaconia can only be distinguished from other subgenera of Zyras by its laterally compressed antennae (Bernhauer, 1928). However, Diaulaconia is presumed to be related to genera Pedinopleurus Cameron, 1939 and Termitodonia Cameron, 1936, and subgenera Camonia Bernhauer, 1928, Glossacantha Gemminger & Harold, 1868, and Termidonia Motschulsky, 1860 by sharing with them numerous morphological characters, containing also laterally compressed antennae (Hlaváč, 2005; Hlaváč et al., 2011; Maruyama, 2006; Maruyama & Kanao, 2014). It is obvious that Diaulaconia was not well-defined by Bernhauer

534 · Zootaxa 4341 (4) © 2017 Magnolia Press LIANG ET AL. (1928), nevertheless, there are several species assigned to the subgenus Diaulaconia based on their similarity with the first described species, Zyras compressicornis. As a consequence, members of Diaulaconia are practically the Zyras compressicornis species complex. Some of the species, such as Z. ibanorum Pace, 1993, Z. lacunarumbeneolentium Pace, 1993, and Z. minangkabauorum Pace, 1993 can only be separated from Z. compressicornis by the morphology of their spermatheca because males were not described (Pace, 1993). The geographical variation of spermatheca had been recorded in Pella Stephens, 1835 (Maruyama, 2006). The present study reports that the spermatheca of Z. artemis sp. n. is variable not only allopatrically but also sympatrically (Fig. 6). Therefore, the species described only based on one character of one specimen would be insufficient for species delimitation.

FIGURE 5. Zyras (Diaulaconia) artemis sp. n., female. A, eight abdominal tergite, dorsal view; B 8th abdominal sternite, ventral view; C, spermatheca; D, the photograph of the apical part and spermathecal gland; E, the photograph of the inner wall of apical 1/3 of the basal part; F, the base of the basal part and the spermathecal duct.

The genus Zyras is considered to be a non-monophyletic group. Several subgenera of Zyras were reported as distantly related to Zyras s. str. and should be probably elevate to generic rank or synonymized with allied genera, e.g., Diaulaconia, Glossacantha, and Termidonia (Hlaváč, 2005; Hlaváč & Jászay, 2009; Maruyama, 2006;

NEW SPECIES OF TERMITOPHILOUS ROVE BEETLE Zootaxa 4341 (4) © 2017 Magnolia Press · 535 Maruyama & Kanao, 2014). One of them had already been recently assigned to genus Termitodonia (Maruyama & Kanao, 2014). Diaulaconia is apparently a distinct lineage rather than the subgenus of Zyras s. str. according to the various differences between their morphology such as the aedeagus, spermatheca, and the shape of pronotum (Hlaváč, 2005 and Hlaváč & Jászay, 2009). The comprehensive revision of the subgenus Diaulaconia is recommended for confirming its systematic position which is unsettled within the genus Zyras and other allied taxa. Termitophily of Diaulaconia is for the first time confirmed in this study, which can reveal the close relationship of this taxon with other termitophilous genera or subgenera. Note on the distribution of allied species in Taiwan. Two species, Z. compressicornis and Z. roepkei, belonging to the subgenus Diaulaconia were recorded in Taiwan (Hlaváč, 2005; Hlaváč et al., 2011; Smetana, 2004). In the original description of Z. compressicornis, Fauvel (1905) recorded this species in the "Chine méridionale: Hong-Kong". Smetana (2004) recorded Z. compressicornis in Taiwan, but we did not findYsufficient evidence (material examination or literature) to support this record. After the examination of specimens from the collection of TARI, we found one specimen of Z. compressicornis from Taiwan, which was determined by Tokuichi Shiraki. However, this specimen was misidentified and it belongs to the new species described in this paper.

FIGURE 6. Variations of the spermatheca of Zyras (Diaulaconia) artemis sp. n. Locality of each specimens: A, Gaupo (Taoyuan Co.); B, Hsilin trail (Hualien Co.); C, Dahashan (Pingtung Co.); D, E, Dakeng (Taichung City); F, Xiaping (Nantou Co.); G, H, Luanshan (Taitung Co.); I, Sheding (Pingtung Co.); J, K, Penghu Islands (Penghu Co.).

536 · Zootaxa 4341 (4) © 2017 Magnolia Press LIANG ET AL. Likewise, Bernhauer (1914) described Z. roepkei from Java (Telaga Patengan), but Smetana (2004) misreported it from Taiwan instead of Java, probably because of the typing error. Subsequent studies and catalogue followed Smetana (2004) and reported Z. compressicornis (Hlaváč, 2005; Hlaváč et al., 2011) and Z. roepkei (Assing, 2009; Hlaváč et al., 2011) to be distributed in Taiwan. Here, we exclude both species from the fauna of Taiwan.

Acknowledgments

The authors thank Yen-Chiu Lan (University of Kang Ning) and Naomichi Tsuji (Kyushu University) for assisting field collection. This study was supported by grants from the Experimental Forest, National Taiwan University (105-A03-4 and 106-B11) and the Ministry of Science and Technology, Taiwan (MOST 104-2311-B-005-002 and MOST 105-2628-B-005-003-MY3).

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