Morphological Comparisons of Lefua Costata (Teleostei: Nemacheilidae), L
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Biogeography 22. 9–15. Sep. 20, 2020 Morphological comparisons of Lefua costata (Teleostei: Nemacheilidae), L. pleskei and L. nikkonis, with notes on morphological variations in L. costata introduced to Japan Shotaro Ohgita 1 and Mizuki Matsunuma 2* 1 Graduate School of Agriculture, Kindai University, 3327-204 Nakamachi, Nara 631-8505, Japan 2 Department of Environmental Management, Faculty of Agriculture, Kindai University, 3327-204 Nakamachi, Nara 631-8505, Japan Abstract: Morphological comparisons of Lefua costata (Kessler, 1876) with related congeners, L. pleskei (Herzenstein in Warpachowski & Herzenstein, 1888) and L. nikkonis (Jordan & Fowler, 1903), resulted in a revision of diagnostic features for L. costata, including: a rhomboidal or triangular dark blotch in the middle of the caudal-fin base; a dark lateral stripe along the body in mature males; pre-dorsal vertebrae 16–18 (modally 17); abdominal vertebrae 18–22 (21); longitudinal scales 101–136; body depth 12.3–21.7% SL (mean 16.5% SL); body width 12.0–15.3% SL (13.8% SL); head depth 9.1–12.4% SL (10.9% SL); and interorbital width 7.3–9.6% SL (8.5% SL). Morphological differences between native populations of L. costata and introduced Japanese populations are also discussed. Key words: Lefua nikkonis, Lefua pleskei, distribution, morphology, diagnosis of pre-dorsal vertebrae (Naseka & Bogutskaya, 2004), and Introduction from L. nikkonis in having a greater number of longitudinal The genus Lefua Herzenstein, 1888 is characterized by scale rows and a smaller and narrower body (Nakajima & a rounded caudal fin, the anterior nostril with a long nasal Uchiyama, 2017). However, careful comparisons of L. costa- barbel, and the absence of a folding spine under the eyes and ta with the latter species resulted in a revision of diagnostic postcleithrum and epural bones (Nichols, 1943; Sawada, 1982; features of the former. Since specimen-based description of L. Prokofiev, 2005; Ito et al., 2019). The genus is distributed costata was never given in the previous studies, a description throughout East Asia, including Mongolia, Russia, China, of L. costata is given, together with detailed comparisons with Korea and Japan (Kottelat, 2012; Hosoya, 2013). According congeners, and comments on intraspecific morphometric vari- to Kottelat (2013), Hosoya et al. (2018) and Ito et al. (2019), ations evident in introduced Japanese populations. the following seven nominal species are currently recognized Materials and Methods as valid: Lefua costata (Kessler, 1876), L. pleskei (Herzenstein in Warpachowski & Herzenstein, 1888), L. nikkonis (Jordan Counts and measurements followed Hubbs & Lagler (2004), & Fowler, 1903), L. echigonia Jordan & Richardson, 1907, L. except for vertebral counts which followed Hosoya (1983), sayu (Herre & Lin, 1936), L. torrentis Hosoya, Ito & Miyaza- with the following additions: pre-dorsal-fin trunk length - the ki, 2018 and L. tokaiensis Ito, Hosoya & Miyazaki, 2019. distance between the upper origin of the gill slit and the dor- Among the above species, L. costata is naturally distribut- sal-fin origin; and post-dorsal-fin length - the distance between ed (hereafter referred to as “native distribution”) in the Amur the dorsal-fin origin and mid-point of the caudal-fin base. River drainage, Korea [given the Japanese name “Hime-dojo” Barbel length was taken as the distance from the base to the by Mori (1928) in his checklist of Korean fishes] and north- extreme tip. Fin rays and vertebrae were counted from radio- eastern China, but is also known as an invasive species in graphs of the examined specimens (except for FAKU 51061, Japan (Naseka & Bogutskaya, 2004; Kottelat, 2012; Hosoya, FAKU 116365, FAKU 116366 and FKUN 31994–31998). The 2013; Nakajima & Uchiyama, 2017). Similar to L. pleskei and last two rays of the dorsal and anal fins were counted as a sin- L. nikkonis in having a rhomboidal or triangular dark blotch gle ray, each pair being associated with a single pterygiophore. in the middle of the caudal-fin base and a dark lateral stripe For morphometric comparisons, data from “narrow-body along the body in mature males (Ito et al., 2019; this study), form” L. costata specimens were excluded (see Table 1). L. costata can be distinguished from L. pleskei by the position Standard length is abbreviated SL. Institutional codes follow of the dorsal fin, snout length (or orbit diameter) and number Sabaj (2019). Identifications of L. costata, L. pleskei and L. −−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−−− nikkonis are based on Naseka & Bogutskaya (2004), Shedko *Corresponding author: [email protected] et al. (2008), Hosoya (2013), Nakajima & Uchiyama (2017), – 9 – Morphological comparisons of Lefua costata (Teleostei: Nemacheilidae), L. pleskei and L. nikkonis, with notes on morphological variations in L. costata introduced to Japan Hosoya et al. (2018) and Ito et al. (2019). male, KUN-P 52870 41.9 mm SL, female, KUN-P 52871 43.0 mm SL, male, Kurobe River, Hakanoki Nature Park, Nyuzen, Lefua costata (Kessler, 1876) Shimonikawa, Toyama Pref., 36°52’20”N, 137°32’35”E, 0.3 [Standard Japanese name: Hime-dojo] m depth, hand net, K. Mochizuki, 26 June 2019; robust-body (Figs. 1, 2, 4, 5, Tables 1, 2) form: KUN-P 48180, 67.3 mm SL, female, KUN-P 48181, Materials examined. Fifty eight specimens, 28.5–67.3 mm 62.5 mm SL, female, KUN-P 48182, 40.7 mm SL, male, SL. Japan: narrow-body form: HUMZ 228716, 34.8 mm SL, KUN-P 48183, 45.1 mm SL, male, Lake Midori, Hokuto, female, HUMZ 228717, 43.1 mm SL, male, HUMZ 228718, Yamanashi Pref., 35°51’29”N, 138°21’52”E, collection date 12 specimens, 28.5–52.8 mm SL, 2 males, 9 females and 1 unknown. North Korea: CAS-SU 25611, 3 specimens, 32.0– sex undetermined juvenile, Lake Midori, Hokuto, Yamanashi 43.8 mm SL, 2 males and 1 female, Seikoshin, J. Thompson, Prefecture (Pref.), 19 Mar. 2017; KUN-P 46649, 57.1 mm SL, 13 May 1911; CAS-SU 28151, 2 specimens, 37.5–39.3 mm female, KUN-P 46650, 47.0 mm SL, female, KUN-P 46651, SL, females, Nampo, Pyeongannam-do, D. Jordan, 1911; 49.2 mm SL, female, KUN-P 46652, 64.6 mm SL, female, CAS-SU 29531, 53.2 mm SL, female, Wonsan, J. Thompson; KUN-P 46653, 48.5 mm SL, female, KUN-P 46654, 39.5 CAS-SU 34741, 3 specimens, 42.8–48.8 mm SL, 2 males mm SL, female, KUN-P 46655, 42.8 mm SL, male, KUN-P and 1 female, Hoeryong, J. Thompson, 21 May 1911. South 48166, 41.3 mm SL, female, Kurobe River, Hakanoki Nature Korea: FAKU 51061, 41.0 mm SL, female, precise locality Park, Nyuzen, Shimonikawa, Toyama Pref., 3 Nov. 2011; unknown; FKUN 31994, 55.4 mm SL, female, FKUN 31995, KUN-P 46737, 53.6 mm SL, female, KUN-P 46738, 51.7 mm 48.1 mm SL, female, FKUN 31996, 42.8 mm SL, male, SL, female, KUN-P 46739, 46.6 mm SL, male, Lake Midori, FKUN 31997, 42.8 mm SL, male, FKUN 31998, 31.6 mm SL, Hokuto, Yamanashi Pref., 35°51’29”N, 138°21’52”E, 2018; male, Chodo, Goseng, Gangwon. China: FAKU 116365, 79.0 KUN-P 51148, 42.4 mm SL, male, Lake Midori, Hokuto, mm SL, female, FAKU 116366, 60.1 mm SL, female, precise Yamanashi Pref., 35°51’29”N, 138°21’52”E, hand net, K. locality unknown, 21 Jan. 1995. Mochizuki, 21 Aug. 2018; KUN-P 51153, 43.2 mm SL, male, Diagnosis. Lefua costata is distinguished from congeners KUN-P 51154, 38.2 mm SL, female, KUN-P 51164, 32.9 by the following combination of characters: rhomboidal or mm SL, female, KUN-P 51165, 36.8 mm SL, female, KUN-P triangular dark blotch in middle of caudal-fin base; dark lateral 51166, 31.6 mm SL, female, KUN-P 51167, 36.6 mm SL, stripe along body in mature males; pre-dorsal vertebrae 16–18 sex undetermined juvenile, KUN-P 51168, 32.1 mm SL, sex (modally 17); abdominal vertebrae 18–22 (21); longitudinal undetermined juvenile, KUN-P 51169, 29.3 mm SL, sex un- scales 101–136; body depth 12.3–21.7% SL (mean 16.5% SL); determined juvenile, Lake Midori, Hokuto, Yamanashi Pref., body width 12.0–15.3% SL (13.8% SL); head depth 9.1–12.4% 35°51’29”N, 138°21’52”E, 0.2 m depth, hand net, M. Fujita SL (10.9% SL); and interorbital width 7.3–9.6% SL (8.5% and Y. Yashima, 3 May 2019; KUN-P 52869, 45.3 mm SL, fe- SL). Fig. 1. Fresh specimens of Lefua costata from Japan. (A) KUN-P 51153, 43.2 mm SL, male, Yamanashi; (B) KUN-P 52871, 43.0 mm SL, male, Toyama; (C) KUN-P 46737, 53.6 mm SL, female, Yamanashi; (D) KUN-P 52869, 45.3 mm SL, female, Toyama; (E) KUN-P 51168, 32.1 mm SL, sex unknown, Yamanashi; (F) KUN-P 51169, 29.3 mm SL, sex unknown, Yamanashi. – 10 – Shotaro Ohgita and Mizuki Matsunuma Fig. 2. Preserved specimens of Lefua costata from native populations (A–D) and Japan, [“robust-body form” (E and F) and “narrow-body form” (G and H)]. (A) FAKU 116366, 60.1 mm SL, male, China; (B) FAKU 116365, 79.0 mm SL, female, China; (C) CAS-SU 29531, 53.2 mm SL, fe- male, North Korea; (D) FKUN 31998, 31.6 mm SL, male, South Korea; (E) KUN-P 48182, 40.7 mm SL, male, Yamanashi; (F) KUN-P 48181, 62.5 mm SL, female, Yamanashi; (G) KUN-P 51153, 43.2 mm SL, male, Yamanashi; (H) KUN-P 51165, 36.8 mm SL, female, Yamanashi.