A NEW SPECIES OF CAPNIA PICTET, 1841 (PLECOPTERA: CAPNIIDAE) FROM KYUSHU, JAPAN, WITH DESCRIPTION OF MALE DRUMMING SIGNALS

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Hanada, Satoko. 2018. A new species of Capnia Pictet, 1841 (Plecoptera: Capniidae) from Kyushu, Japan, with description of male http://zoobank.org/ urn:lsid:zoobank.org:pub:3ff3d463-16a5-4255-bd16-16c30bf04d5c A NEW SPECIES OF CAPNIA PICTET, 1841 (PLECOPTERA: CAPNIIDAE) FROM KYUSHU, JAPAN, WITH DESCRIPTION OF MALE DRUMMING SIGNALS Satoko Hanada Nishijin, Sawara Ward, Fukuoka City, Fukuoka Prefecture 814-0002, Japan E-mail: s.hanada@email.plala.or.jp ABSTRACT A new species of the genus Capnia Pictet, 1841, Capnia mitsuseana sp. n. (Plecoptera: Capniidae), from Japan is described on the basis of male and female adults, and male drumming signal. Males of the new species are characterized by a modified abdominal tergum 7 with a pair of small tubercles on the anterior margin and with a large tubercle on the posterior margin; a long and rather narrow main epiproct sclerite with a pair of needle-like processes on either side of the tip. Females have abdominal sternum 7 protruding caudally and short subgenital plate that is nearly same length of sternum 8. The males produced monophasic call signals in succession by percussion involving up-down abdominal movements under solitary condition, and the male call signals show horizontal inter-beat interval patterns. Keywords: Capnia mitsuseana Hanada, stonefly, drumming INTRODUCTION The genus Capnia is known as a large genus in Plecoptera and contains more than 100 species (DeWalt et al. 2018). Thirteen species of Capnia have been recorded from Japan (DeWalt et al. 2018), and still more undescribed species are known (Shimizu & Negoro 2003). I report a new species of Capnia from northern Kyushu, western Japan resulting from studies in the region (Hanada 2014, 2016a, 2016b). I describe the new species based on morphological features of adults and from characteristics of the male vibrational communication signals (drumming), which are known within the infraorder Arctoperlaria as species-specific vibrational signals used for intersexual communication. MATERIALS AND METHODS Morphological observation. Adults were handcollected and preserved in 80% ethanol and studied using a Leica MZ12 stereomicroscope (magnification up to 100x). All illustrations were produced using a camera lucida attached to stereomicroscope. The epiproct and the fusion plate used for illustrations were removed from paratype specimens to facilitate illustrations. Female vaginal sclerites were observed and illustrated after abdomens were cleared in a solution of 10% potassium hydroxide (KOH). All examined materials were collected by Mr. H. Maruyama and the author, and therefore the collector s names were omitted in the materials examined list. The type series will be deposited in Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 155

1 2 3 4 5 Figs. 1-5. Capnia mitsuseana sp. n. 1. Habitus of Capnia mitsuseana sp. n. (female). 2. Right wings of Capnia mitsuseana sp. n. 3 5. Capnia mitsuseana sp. n. (male). 3. Terminalia, dorsal view. 4. Terminalia, lateral view. 5. Epiproct, lateral view (above), dorsal view (below). Ec = eversible crest, Ep-scl = main epiproct sclerite, arrow = needle-like process. Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 156

the Lake Biwa Museum, Shiga Prefecture, Japan, and additional materials are deposited in the author s collection. Recording of drumming signals. Four males collected from the type locality on 16 January and 7 February 2016 were used for recordings of male call signals under solitary condition. These males were kept individually in plastic cylinders with a piece of wet absorbent cotton and lichens from their habitat as food source. The male drumming signals were recorded and analyzed in accordance with the methodology conducted by Hanada (2018). The terminology of drumming signals follows that of Sandberg (2011) and Sandberg et al. (2015). DESCRIPTION OF SPECIES Capnia mitsuseana sp. n. http://lsid.speciesfile.org/urn:lsid:plecoptera.speciesfile.org: TaxonName:504342 (Figs. 1 10) Material examined. Holotype, Japan: Iiba, Sawara Ward, Fukuoka City, Fukuoka Prefecture, 33 28.73 N, 130 17.11 E, alt. 480 m, Muromi River, 1.II.2015. Paratypes: 1, type locality, 24.I.2015; 4, 1, type locality, 21.II.2015; 2, type locality, 3.I.2016; 4, type locality, 21.I.2018. Additional materials. Japan: Iiba, Sawara Ward, Fukuoka City, Fukuoka Prefecture (type locality), 16.II.2013, 1, 1 ; same locality, 23. III.2013, 1 ; same locality, 13.I.2014, 1, 1 ; same locality, 22.II.2014, 1, 1 (copulating pair); same locality, 10.I.2015, 1 ; same locality, 12.I.2015, 1 ; same locality, 18.I.2015, 1, 1 ; same locality, 24.I.2015, 4, 1 ; same locality, 22.II.2015, 1 ; same locality, 28.II.2015, 1 ; same locality, 3.I.2016, 1 ; same locality, 16.I.2016, 5 ; same locality, 2.II.2016, 2, 1 ; same locality, 7.II.2016, 1, 1 ; same locality, 28.II.2016, 1 ; same locality, 14.I.2017, 1 ; same locality, 29.I.2017, 1 ; same locality, 2.I.2018, 2, 2 ; same locality, 23.II.2018, 2, 1 ; same locality, 4.III.2018, 1 ; Magaribuchi, Sawara Ward, Fukuoka City, Fukuoka Prefecture, 33 28.97 N, 130 17.34 E, alt. 410 m, Muromi River, 24.I.2015, 9, 5 ; same locality, 31.I.2015, 1 ; same locality, 22.II.2015, 1 ; same locality, 21.II.2016, 1 ; Kumado, Hoshino Village, Yame City, Fukuoka Prefecture, 33 13.82 N, 130 50.88 E, alt. 716 m, Kumado River, 25.I.2015, 4 ; same locality, 16.I.2016, 1 ; same locality, 28.II.2016, 1 ; Kamiose, Fuji Town, Saga City, Saga Prefecture, 33 28.22 N, 130 16.21 E, alt. 515 m, Hatsuse River, 7.III.2015, 1 ; same locality, 3.I. 2016, 2 ; same locality, 9.I.2016, 2, 1 ; same locality, 30.I.2016, 1 ; same locality, 2.I.2018, 1 ; same locality, 7.I.2018, 1 ; Mitsuse, Mitsuse Town, Saga City, Saga Prefecture, 33 27.91 N, 130 16.47 E, alt. 448 m, Hatsuse River, 16.I.2016, 1 ; same locality, 2.I.2018, 1 ; same locality, 21.I.2018, 1 ; same locality, 23.II.2018, 1 ; Ikisa, Ouchi Town, Karatsu City, Saga Prefecture, 33 22.19 N, 130 02.95 E, alt. 320 m, Saikisa River, 17.III.2018, 1 ; Yoshida, Ureshino City, Saga Prefecture, 33 01.30 N, 130 02.09 E, alt. 460 m, Yoshida River, 29.I.2000, 2, 3 ; same locality, 8.III.2000, 1. Adult habitus. Macropterous (Fig. 1), but wing length somewhat variable (forewing length / body length = 0.68 1.17, n = 47 individuals). Wings subhyaline and slightly brownish; veins brown and partly pale. 1A of fore-wing typical to that of Capnia, bent abruptly caudad at its junction with cu-a and curved outward again; R1 of fore-wing typical to that of Capnia, bent upward at its origin; Cu1 and Cu2 of forewing originating from the different points on arculus (Fig. 2), but rarely from the same point on arculus (11%, n = 46 individuals). General color dark brown. Head, thoracic sclerites and basal segments of abdomen typical to those of Capnia. Cerci long, brownish, and composed of 16 26 segments (n = 10 individuals). Male. Body 4.8 7.0 mm (n = 23) long. Forewing 3.8 6.0 mm (n = 23) long. Tergum 7 modified (Figs. 3 & 4): the anterior margin rising and forming a pair of small and short tubercles with blunt tips; the posterior margin extending upward and forming a large tubercle with a rough, rounded apex bearing sensilla basiconia, the back of which covers the tip of epiproct; membranous field located on antero-middle of tergum 7. The anterior margin of tergum 8 slightly curved posteriorly. Main epiproct sclerite long and rather narrow with a taper, rounded tip in dorsal view; its bottom Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 157

6 7 8 9 10 Figs. 6 7. Capnia mitsuseana sp. n. (male). 6. Terminalia, ventral view. 7. Fusion plate, dorsal view (internal view). rp = retractoral plate, sbp = subgenital plate. Figs. 8 10. Capnia mitsuseana sp. n. (female). 8. Terminalia, ventral view. 9. Cleared terminalia and inner vaginal sclerites, ventral view. 10. Cleared terminalia and inner vaginal sclerites, ventro-lateral view. Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 158

11 Fig. 11. A male call signal sequence of Capnia mitsuseana sp. n. consisting of four signals. i = inter-beat interval. nearly straight in lateral view; a pair of needle-like processes located on either side of its tip (Fig. 5). Eversible crest of epiproct swollen in lateral view (Fig. 5). Sternum 9 without a vesicle on the basal margin (Fig. 6). Subgenital plate large and tonguelike shape with a small rounded quadrangular tip; indented basolaterally (Figs. 6 & 7). Fusion plate somewhat shield-shaped, and the anterior margin connected by membrane with a small, subtriangular, sclerotized retractoral plate (Fig. 7). Paraproct with a dilated apex (Fig. 7). Female. Body 6.0 8.9 mm (n = 24) long. Forewing 5.0 7.2 mm (n = 24) long. Abdominal terga 1 to 8 with broad longitudinal membranous fields along the midline; terga 9 and 10 fully sclerotized. Posterior margin of sternum 7 protruding backward (Fig. 8). Subgenital plate of sternum 8 nearly same length of sternum 8; rounded and having short posteromedial lobe (Fig. 8). A pair of longitudinal inner vaginal sclerites bent ventrally near the anterior ends; curved outward near the posterior ends; visible through the subgenital plate as two longitudinal stripes (Figs. 9 & 10). Table 1. Male call signal data of Capnia mitsuseana sp. n. under solitary condition at 16.8 17.0 C. The values are expressed as mean ± standard deviation (Mo = mode, n = number of samples) and range. Mean interbeat interval pattern was determined by mean ID (a maximum minimum interval difference): H = horizontal interval pattern determined by interval difference (ID) 10 (Sandberg et al. 2011). num. of beats mean interbeat interval (ms) inter-beat intervals (ms) i1 i2 i3 ID & mean interbeat interval pattern 3.7 ± 0.5 (Mo = 4, n = 43) 3 4 39.6 ± 2.5 (n = 115) 34.8 53.7 39.1 ± 2.1 (n = 43) 34.8 45.0 39.1 ± 2.0 (n = 43) 36.3 43.5 40.9 ± 3.1 (n = 29) 36.3 53.7 1.8 H Drumming signals. Two males (0-day-old and one-day-old after capture) produced signals by percussion involving up-down abdominal movements under solitary condition at 16.8 17.0 C. Because no female answers were recorded, the call of males is tentatively described as a monophasic signal (Fig. 11). The male call signal was composed of 3.7 ± 0.5 beats with 39.6 ± 2.5 ms Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 159

mean inter-beat interval (Fig. 11, Table 1). The signal durations averaged 105.8 ms (SD = 21.3, range = 71.1 132.1, n = 43). Interval difference was 1.8 ms (40.9 39.1 = 1.8 ms 10 ms ID) (Table 1), and therefore the mean inter-beat interval pattern was horizontal. The two males always produced series of monophasic signals (Fig. 11). The signal sequence consisted of 3.9 ± 0.5 signals (range = 3 5, mode = 4, n = 11) with 287.7 ± 30.4 ms mean signal interval (range = 220.6 364.3, n = 32). 12 Fig. 12. Type locality of Capnia mitsuseana sp. n. Ecological notes. This new species was collected from the upper reaches of rivers (Fig. 14). The emergence period is mainly during winter and early spring (I III). The adults were found on the parapets of bridges and the guard rails. Distribution. Japan (northern Kyushu). Etymology. The specific name mitsuseana refers to the Mitsuse Pass where the new species was found. Remarks. Males of this new species resemble Capnia asakawana Kohno, 1952, C. bituberculata Uéno, 1929, C. kibuneana Kawai, 1957 and C. fukushimana Kohno, 1952 in the structure of the modified abdominal tergum 7 where all members of the group displayed a pair of tubercles on the anterior margin and a large tubercle on the posterior margin (Uéno 1929, Kohno 1952, Kawai 1957). The new species differs from all the others by the anterior tubercles of tergum 7 being smaller and by the main epiproct sclerite being long and rather narrow with a taper, rounded tip in dorsal view and with nearly straight bottom in lateral view. Females of the new species can be distinguished from other Japanese Capnia species by abdominal sternum 7 extending caudally over the anterior margin of the 8 th sternum and by the short subgenital plate of sternum 8 not extending past the posterior edge of the sternum and by the distinct two longitudinal stripes seen on the Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 160

subgenital plate. ACKNOWLEDGMENTS Thanks are due to Mr. Hiroki Maruyama for his contribution collecting the specimens. I am grateful to two reviewers and the editors for their helpful comments and suggestions. REFERENCES DeWalt, R.E., M.D. Maehr, U. Neu-Becker & G. Stueber. 2018. Plecoptera Species File Online. Version 5.0/5.0. Accessed 25 September 2018. http://plecoptera.species file.org Hanada, S. 2014. Stoneflies (Insecta: Plecoptera) collected from Sefuri Mountains, Kyushu, western Japan. Biology of Inland Waters, 29:17 26. [Japanese with English Abstract] Hanada, S. 2016a. Stoneflies (Insecta: Plecoptera) collected from Tara Mountains in Kyushu, Japan. Biology of Inland Waters, 30:15 23. [Japanese with English Abstract] Hanada, S. 2016b. Stoneflies (Insecta: Plecoptera) collected from the Kumado River and the upper reaches of the Hoshino River in Fukuoka Prefecture, Japan. Biology of Inland Waters, 30:25 30. [Japanese with English Abstract] Hanada, S. 2018. A new species of Nemoura (Plecoptera: Nemouridae) from Kyushu, Japan, with notes on male vibrational signals. Illiesia, 14(07):135 143. Kawai, T. 1957. Studies on the holognathus stoneflies of Japan. III Descriptions of four new species of Capnid stoneflies. Mushi, 30(8):41 47. Kohno, M. 1952. Notes on Japanese Capniidae with descriptions of new species (Plecoptera). Mushi, 23(2):17 28. Sandberg, J.B. 2011. Vibrational communication of nine California stonefly (Plecoptera) species. Western North American Naturalist, 71:285 301. https://scholarsarchive.byu.edu/wnan/vol71/iss3/1 Sandberg, J.B, L.E. Serpa, & E.F. Drake. 2015. The drumming signals of three winter stonefly species (Capniidae, Leuctridae: Plecoptera); with discussion resolving two common interval patterns. Illiesia, 11(06):51 74. Shimizu, T. & H. Negoro. 2003. The stoneflies collected on snows in Toyama and its adjoining Prefectures: Stoneflies except apterous Capniidae. Bulletin of the Toyama Science Museum, 26:103 115. [Japanese with English Abstract] Uéno, M. 1929. Studies on the stoneflies of Japan. Memoirs of the College of Science, Kyoto Imperial University, Series B, 4(2):97 155 with plate XXIV. Submitted 28 September 2018, Accepted 1 November 2018, Published 3 December 2018 Hosted and published at the University of Illinois, Illinois Natural History Survey, Champaign, Illinois, U.S.A. Illiesia http://illiesia.speciesfile.org Volume 14 Number 9 Page 161