taxonID	type	description	language	source
0C4287800E75FFC6FE8DFADA06CD11C7.taxon	materials_examined	Material examined. In total six specimens were collected from rice paddy located to the south of Mt. Kuburadake, Yonaguni Island, Ryukyu Islands, Japan on 19 March 2014: KUZ Z 5159, Z 5160, 24.45064 ° N, 122.95425 ° E, by Yoshiko Yamane; KUZ Z 5161 – Z 5164, 24.44956 ° N, 122.95272 ° E, by Takafumi Nakano. Three specimens, KUZ Z 5159, Z 5162, Z 5164, were dissected.	en	Nakano, Takafumi, Lai, Yi-Te, Seo, Hong-Yul, Kambayashi, Chiaki (2024): First Record of the Predatory Leech Whitmania laevis (Hirudinea: Hirudiniformes: Hirudinidae) from the Ryukyu Islands, Japan, with New Insights into the Phylogenetic Relationships within Whitmania. Species Diversity 29 (2): 239-246, DOI: 10.12782/specdiv.29.239, URL: https://doi.org/10.12782/specdiv.29.239
0C4287800E75FFC6FE8DFADA06CD11C7.taxon	description	Description. Body firm, muscular, cephalic region small, from prostomium to somite XI with width increasing significantly in posterior direction, then from somite XII with constant width in posterior direction, ventrally flattened (Figs 1 A, B, 2 A, C). Oral sucker small, ovate, OL 1.4 – 1.8 mm, OW 1.3 – 1.8 mm (Figs 1 B, 2 C). Caudal sucker ventral elliptic, CL 2.9 – 4.6 mm, CW 3.8 – 5.3 mm (Figs 1 B, 2 G). Somite I completely merged with prostomium (Fig. 2 A). Somite II uniannulate (Fig. 2 A). Somite III uni- or biannulate, (a 1 + a 2) = a 3, a 3 sometimes with slight dorsal furrow (Fig. 2 A). Somites IV and V generally biannulate, (a 1 + a 2) = a 3, (a 1 + a 2) generally with slight dorsal furrow (Fig. 2 A, B); somite V forming posterior margin of oral sucker (Fig. 2 C). Somite VI dorsally triannulate, a 1> a 2 <a 3, or a 1 sometimes with obvious furrow, a 1 (b 1 = b 2)> a 2 <a 3; ventrally biannulate, (a 1 + a 2)> a 3 (Fig. 2 A – C). Somite VII quadrannulate, a 1> a 2 = b 5 = b 6 (Fig. 2 A – C), or sometimes a 1 (b 1 = b 2 dorsally)> a 2 = b 5 = b 6. Somites VIII – XXIII quinquannulate, b 1 = b 2 = a 2 = b 5 = b 6 (Fig. 2 A – E). Somite XXIV quinquannulate, b 1 = b 2 = a 2 = b 5 = b 6, or quadrannulate, b 1 = b 2 = a 2 <a 3 (b 5 = b 6) (Fig. 2 F, G). Somite XXV quadrannulate, b 1 = b 2 = a 2 = a 3 (Fig. 2 F, G). Somite XXVI biannulate, (a 1 + a 2)> a 3, (a 1 + a 2) sometimes with slight furrow (Fig. 2 F); (a 1 + a 2) being ventrally last complete annulus. Somite XXVII uniannulate (Fig. 2 F). Anus at posterior margin of XXVII (Fig. 2 F). Male gonopore in anterior margin of, or middle of somite XI b 6 (Fig. 2 D). Female gonopore in anterior margin of somite XII b 6 (Fig. 2 D). Gonopores separated by ≤ 5 annuli (Fig. 2 D). Eyespots in 5 pairs, in parabolic arc (Fig. 2 A, B); 1 st pair on II; 2 nd pair on III or III (a 1 + a 2); 3 rd pair on IV (a 1 + a 2); 4 th pair on V (a 1 + a 2); 5 th pair on VI a 2. Sensillae developed, 1 row on every annulus. Somital papillae developed, generally 6 papillae detected on a 2 of each somite in VII – XXVII (Fig. 2 A, B, E, F). Nephridiopores in 17 pairs, one each situated ventrally at posterior margin of b 2 of each somite in VIII – XXIV (Fig. 2 C, D, G). One median longitudinal furrow on ventral surface of oral sucker (Fig. 2 C). Three rudimental jaws in oral cavity, 1 dorsal and 2 ventrolateral; salivary papillae undetectable; distichodont. Pharynx reaching to IX b 2 – b 5. Crop reaching to XIX b 2, giving rise to 1 pair of crop ceca (post-crop ceca) in XIX b 1 – b 2 to XX b 5 – XXII b 2 (Fig. 3 A). Terminal end of intestine forming sphincter between intestine and rectum, in XXI a 2 to XXII b 1 – b 5. Testisacs in 9 or 10 pairs (Fig. 3 B): 1 st pair in XIII b 5 – b 6 to XIV b 2 (left testisac undetectable in KUZ Z 5159); 2 nd pair in XIV a 2 – b 5 to XV b 1; 3 rd pair in XV a 2 – b 5 to XVI b 1; 4 th pair in XVI a 2 – b 5 to XVI b 6 – XVII b 1; 5 th pair in XVII a 2 – b 6 to XVII b 6 – XVIII b 2; 6 th pair in XVIII a 2 – b 5 to XVIII b 6 – XIX b 1; 7 th pair in XIX a 2 – b 6 to XIX b 6 – XXb 1; 8 th pair in XX a 2 – b 5 to XX b 6 – XXI b 1; 9 th pair in XXI a 2 – b 6 to XXII b 1 – b 2 (left testisac undetectable in KUZ Z 5159); 10 th pair (detected in KUZ Z 5164 only) undeveloped, in XXII b 5 and b 6. Paired epididymides developed, globular or fusiform (Fig. 3 B, C). Ejaculatory bulbs developed, elongated, fusiform (Fig. 3 B – D); in large individuals (KUZ Z 5159 and Z 5162), in XI a 2 – XII b 1 to XII a 2 – b 6, occupying 6 or 7 annuli; in smaller individual (KUZ Z 5164), in XI b 5 – b 6 to XII b 1 – b 2, occupying 4 annuli. Ejaculatory ducts narrow, running toward male atrium in XI a 2 – b 5 to XII b 2 – XIII b 5 (Fig. 3 B – D); right duct crossing ventrally beneath nerve cord. Male atrium continuous with penis sheath (Fig. 3 B – D). Penis sheath hook-like, U- or L-shaped, reaching to XIII b 1 – b 6, then turning anteriorly to male gonopore (Fig. 3 B – D). Paired ovisacs ellipsoidal or globular, in XII b 5 – XIII b 1 to XIII b 1 – b 2 (Fig. 3 B, E, F). Oviducts short (Fig. 3 B, E, F); right oviduct crossing ventrally beneath nerve cord; both oviducts converging into common oviduct in XII b 6 or XIII b 2. Common oviduct thick, long, descending to female vagina in XII b 6 – XIII b 2 to XIII b 5 – XIV a 2 (Fig. 3 B, E, F). Female vagina continuous to vaginal duct, tubular, in XII b 6 to XIII b 5 – XIV a 2 (Fig. 3 B, E, F); vaginal cecum absent. Coloration. In life, dorsal surface olive green with 5 discontinuous whitish-green longitudinal stripes, all stripes bordered in brown, each stripe fully or slightly faded on a 2 of each mid-body somite (Figs 1 C, 2 E); several irregular black markings present on lateral margins; ventral surface tannish with black lateral margins, black markings present on lateral margins and caudal sucker, several irregular black markings also present on mid-ventral surface. Color faded in preservative, but longitudinal stripes and markings still present (Figs 1 A, B, 2 G). Natural history. All individuals were found crawling in water of a paddy field at night. Japanese name. The Japanese name “ Gohon-sesuzi-biru, ” which was proposed by Ohno (1998), is adopted herein. This name is likely to be derived from its five (= “ gohon- ”, Βψ) dorsal (= “ - se- ”, Ü) longitudinal stripes (= “ - suzi ”, ®); “ biru ” (or “ hiru ”) means a leech in Japanese. Phylogenetic position and genetic distances. The ML (ln L = − 7516.22; not shown) and BI (ln L = − 7559.22; Fig. 4) trees had congruent topologies. The monophyly of each of the four Whitmania species was well supported: W. acranulata (BS = 100 %, PP = 1.0), W. edentula (BS = 100 %, PP = 1.0), W. laevis (BS = 98 %, PP = 1.0), and W. pigra (BS = 86 %, PP = 0.98). Monophyly of three Whitmania species, i. e., W. pigra, W. laevis, and W. edentula, was well supported (BS = 94 %, PP = 1.0). Whitmania laevis formed a monophyletic lineage with W. edentula (BS = 98 %, PP = 1.0). The Taiwanese and Yonaguni W. laevis formed a distinct clade from the continental W. laevis sequences (BS = 89 %, PP = 0.99). The pairwise COI uncorrected p - distances between W. laevis and W. edentula were 10.5 % – 11.9 %. The intraspecific COI variations within W. laevis were as follows: 0.2 % – 0.7 % within the Taiwanese and Yonaguni populations; 2.1 % between the Korean and continental Chinese sequences; and 1.8 % – 3.5 % between the Taiwanese + Yonaguni and continental specimens.	en	Nakano, Takafumi, Lai, Yi-Te, Seo, Hong-Yul, Kambayashi, Chiaki (2024): First Record of the Predatory Leech Whitmania laevis (Hirudinea: Hirudiniformes: Hirudinidae) from the Ryukyu Islands, Japan, with New Insights into the Phylogenetic Relationships within Whitmania. Species Diversity 29 (2): 239-246, DOI: 10.12782/specdiv.29.239, URL: https://doi.org/10.12782/specdiv.29.239
0C4287800E75FFC6FE8DFADA06CD11C7.taxon	discussion	Remarks. The Whitmania leeches collected from Yonaguni Island were identified as W. laevis based on the presence of the following characteristics, which were also reported by the prior studies (Blanchard 1896; Moore 1927 a; Yang 1996; Lai and Chen 2010): dorsal surface with five discontinuous whitish-green longitudinal stripes and several irregular black markings, ventral surface with black markings on lateral margins and caudal sucker; male gonopore at XI b 6 and female gonopore at XII b 6; paired epididymides globular or fusiform; penis sheath hook-like; oviducts short; common oviduct long, descending to female vagina; vaginal duct tubular; vaginal cecum absent. The results of our molecular analyses supported the taxonomic account of the Yonaguni Whitmania specimens determined by those morphological features. The results showed close genetic relationships between the Yonaguni and Taiwanese populations. This occurrence report provides the first record of W. laevis from the Ryukyu Islands (see Itoh 2003). It remains uncertain whether W. laevis on Yonaguni Island was introduced from Taiwan or anywhere else in recent years, but nonetheless, it is noteworthy that the Yonaguni specimens have unique COI sequences. The large W. laevis specimens from Yonaguni Island possess nine pairs of testisacs. However, the Chinese and Taiwanese specimens had 11 pairs of testisacs (Yang 1996; Lai and Chen 2010) and those from eastern India had 10 pairs of testisacs (Moore 1927 a). Although the small specimen (KUZ Z 5164) possesses 10 pairs of testisacs, the last pair seemed to be rudimentary. The reduced number of testisac pairs may be a shared feature of the W. laevis population endemic to Yonaguni Island. The present phylogenies revealed a sister relationship between W. laevis and W. edentula for the first time. This relationship is consistent with the morphological similarities in female genital organs between the two species (Whitman 1886), both of which have a thick common oviduct directly descending to the vagina and a tubular vagina and vaginal duct. The calculated uncorrected p - distances between W. laevis and W. edentula were above 10 %, and thus met the average interspecific variations detected among the hirudinid predatory species of Haemopis Savigny, 1822 (Kvist et al. 2023). Therefore, W. laevis and W. edentula should both receive distinct species status within Whitmania. Three of the five Whitmania species, i. e., W. acranulata, W. edentula, and W. pigra, were originally described from Japan. Therefore, their topotypic DNA data and morphological features are essential to reveal the true species richness and evolutionary history of this predatory genus endemic to South and East Asia.	en	Nakano, Takafumi, Lai, Yi-Te, Seo, Hong-Yul, Kambayashi, Chiaki (2024): First Record of the Predatory Leech Whitmania laevis (Hirudinea: Hirudiniformes: Hirudinidae) from the Ryukyu Islands, Japan, with New Insights into the Phylogenetic Relationships within Whitmania. Species Diversity 29 (2): 239-246, DOI: 10.12782/specdiv.29.239, URL: https://doi.org/10.12782/specdiv.29.239
