taxonID	type	description	language	source
530687B03854224723FBFCB7823BFAA9.taxon	vernacular_names	[New Japanese name: kubire-kugabiru]	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	description	(Figs 1 – 3)	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	diagnosis	Diagnosis. Body length of pre-mature individuals exceeding 5 cm; possibly middle-type. Somite IV uniannulate, somites VIII – XXV quadrannulate. Male gonopore in middle of somite XI b 6, female gonopore slightly anterior to middle of somite XIII a 1, behind gastropore, gonopores separated by 1 / 2 + 4 + 1 / 3 annuli. Pharynx reaching to somite XIV a 1 / a 2 – a 2. Gastropore conspicuous, slightly anterior to middle of somite XIII a 1. Gastroporal duct bulbous. Paired epididymides in somites XVIII – XX, occupying 6 – 7 annuli. Paired ejaculatory ducts thick. Atrial cornua developed, hyperboloidal, i. e., each middle part constricted, then expanding at respective junction with ejaculatory duct.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	materials_examined	Material examined. Holotype: KUZ Z 1439, dissected, collected from Mt. Gomadanzan, Ryujinmura, Tanabe, Wakayama Prefecture (Kii Peninsula, Honshu island), Japan (34.06082 ° N, 135.564722 ° E; elev. 1285 m) by Naoki Koike on 10 October 2012. Paratypes (3 specimens): KUZ Z 1440 – Z 1442, data same as for holotype (KUZ Z 1440 was dissected) Description. Body firm and muscular, elongate, with constant width in caudal direction, dorsoventrally compressed, BL 54.3 mm, BW 5.5 mm (Fig. 1 A, B). Caudal sucker ventral, elliptic, CL 2.6 mm, CW 3.2 mm (Figs 1 B, 2 D).	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	description	Somite I completely merged with prostomium (Fig. 2 A). Somites II (= peristomium), III and IV uniannulate (Fig. 2 A); somite II not separated from somite I. Somite V biannulate, (a 1 + a 2) = a 3; a 3 forming posterior margin of oral sucker (Fig. 2 A, B). Somites VI and VII triannulate, a 1 = a 2 = a 3 (Fig. 2 A, B). Somites VIII – XXV quadrannulate, a 1 = a 2 = b 5 = b 6 (Fig. 2 A – E). Somite XXVI dorsally quadrannulate, a 1> a 2> b 5 = b 6, ventrally triannulate, a 1> a 2 <a 3; a 3 (= dorsally b 5 and b 6) being ventrally last complete annulus (Fig. 2 C, D). Somite XXVII biannulate; a 3 being dorsally last annulus with slight furrow (Fig. 2 C). Anus behind somite XXVII; post-anal annulus absent (Fig. 2 C). Clitellum undeveloped. Male gonopore in middle of somite XI b 6 (Fig. 2 E). Female gonopore slightly anterior to middle of somite XIII a 1, inconspicuous, located posterior to gastropore (Fig. 2 E, F). Gonopores separated by 1 / 2 + 4 + 1 / 3 annuli (Fig. 2 E). Anterior ganglionic mass in somite VI a 2 and a 3. Ganglion VII in a 1 and a 2. Ganglia VIII – XIII, of each somite, in a 2 (Fig. 3 A). Ganglia XIV – XV, of each somite, in a 1 and a 2 (Fig. 3 A). Ganglion XVI in a 2 (Fig. 3 A). Ganglia XVII – XVIII, of each somite, in a 1 and a 2 (Fig. 3 A). Ganglion XIX in a 2 (Fig. 3 A). Ganglia XX – XXIV, of each somite, in a 1 and a 2 (Fig. 3 A). Ganglion XXV in somite XXIV b 6 and somite XXV a 1. Ganglion XXVI in somite XXV b 6. Posterior ganglionic mass in somite XXVI a 2 and a 3 (= dorsally b 5 and b 6). Eyes in 3 pairs, 1 st pair dorsally on anterior margin of somite III, 2 nd and 3 rd pairs dorsolaterally on posterior margin of somite V (a 1 + a 2) (Fig. 2 A). Papillae numerous, minute, hardly visible, on row on every annulus. Nephridiopores in 17 pairs, each situated ventrally at posterior margin of a 1 of each somite in somites VIII – XXIV (Fig. 2 B, D, E). Pharynx agnathous, euthylaematous, reaching to somite XIV a 1 / a 2 (Fig. 2 G). Crop tubular, acecate, reaching to somite XX a 1 / a 2. Gastropore conspicuous, ventral, slightly anterior to middle of somite XIII a 1 (Fig. 2 E, F). Gastroporal duct bulbous, winding at junction with gastropore, reaching to somite XIV a 2 (Fig. 2 G). Intestine tubular, acecate, reaching to somite XXIV a 1 / a 2. Rectum tubular, thin-walled, straight. Testisacs multiple, in somite XX b 5 to somite XXV b 6 (Fig. 3 A); on right side, in total ~ 22 testisacs, 1 in XX, 4 in XXI, 4 in XXII, 6 in XXIII, 4 in XXIV, 3 in XXV; on lefl side, in total ~ 22 testisacs, 2 in XX, 4 in XXI, 4 in XXII, 5 in XXIII, 4 in XXIV, 3 in XXV. Paired epididymides in somite XVIII / XIX to somite XX b 5, occupying 7 annuli (Fig. 3 A). Paired ejaculatory ducts thick, in somite XI b 5 to somite XVIII / XIX (Fig. 3 A); loosely coiled in position posterior to ovisacs; each duct crossing ventrally beneath each ovisac, then nearly straight in position anterior to ovisacs; each noticeably winding from respective junction with epididymis, narrowing at junction with atrial cornua, then turning proximally toward atrial cornua without pre-atrial loop (Fig. 3 A, B). Pair of muscular atrial cornua developed, hyperboloidal, in somite XI b 5 and b 6 (Fig. 3 A – D); each middle part constricted, then expanding at respective junction with ejaculatory duct. Atrium short, muscular, globular, in somite XI b 5 and b 6 (Fig. 3 B – D). Penis sheath and penis absent. Paired ovisacs globular; right ovisac in somite XIII a 2 and b 5; lefl ovisac in somite XIII a 2 (Fig. 3 A, E). Oviducts thin-walled, lefl oviduct crossing ventrally beneath nerve cord (Fig. 3 A, E); both oviducts converging in to common oviduct in somite XIII a 1 / a 2. Common oviduct thin-walled, short, directly descending to female gonopore (Fig. 3 E). Variation. Measurements (mean ± 1 SD, followed by ranges in parentheses; n = 4, including holotype): BL 51.9 ± 2.7 mm (48.9 – 54.3 mm), BW 5.1 ± 0.28 mm (4.8 – 5.5 mm), CL 2.4 ± 0.14 mm (2.4 – 2.6 mm), CW 2.8 ± 0.45 mm (2.3 – 3.2 mm). Somite XXVI generally triannulate, a 1> a 2 <a 3. Pharynx reaching to somite XIV a 2. Crop reaching to somite XX a 2 / b 5. Gastroporal duct reaching to somite XIV a 2 / b 5. Intestine reaching to somite XXIV a 2 / b 5. Testisacs in somite XIX a 2 to somite XXV b 5; both sides, ~ 29 sacs, respectively. Paired epididymides occupying 6 annuli, respectively; right side in somite XVII / XVIII to somite XIX a 2 / b 5; lefl side in somite XVIII a 1 to somite XIX a 2 / b 5. Paired ejaculatory ducts; right side in somite XI b 5 to somite XVII / XVIII; lefl side in somite XI b 5 to somite XVIII a 1; coiled in position posterior to ovisacs. Paired ovisacs in somite XIII a 2 and b 5. Coloration. In life, dorsal surface bluish black; ventral surface bluish white. Color faded in preservative.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	distribution	Distribution. This species was collected only from its type locality.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	biology_ecology	Natural history. Although both dissected individuals possessed developed testisacs, their clitella were undeveloped, and hardly detected. The reproductive season of the new species thus remains unclear.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	etymology	Etymology. The specific name is a participle in nominative singular derived from the Latin word angustatus (narrowed) referring to the fact that the male atrial cornua of this new species are constricted in their upper-middle parts.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
530687B03854224723FBFCB7823BFAA9.taxon	discussion	Molecular analyses results. The BI tree (mean ln L = − 28673.07; Fig. 4) for estimating the phylogenetic position of the new species had an identical topology to that of the ML tree (ln L = − 29119.04; not shown). Orobdella angustata belonged to a well-supported clade (BS = 99 %, PP = 1.0) containing the other four quadrannulate species inhabiting the western part of Honshu, Shikoku and northern Kyushu. Orobdella esulcata Nakano, 2010 formed a sister clade to the other four species including O. angustata. The monophyly of O. angustata and the well-supported clade consisting of O. brachyepididymis Nakano, 2016, O. kanaekoikeae Nakano, 2017 and O. naraharaetmagarum Nakano, 2016 (BS = 100 %, PP = 1.0) was also fully supported (BS = 100 %, PP = 1.0). Remarks. Orobdella angustata unquestionably belongs to Orobdella because it possesses the generic diagnostic characteristics defined by Nakano (2016 a); its molecular phylogenetic position also recovers the taxonomic position of the new species. According to previous studies (Nakano 2010, 2011, 2012; Nakano and Lai 2012; Nakano 2014; Nakano and Seo 2014; Nakano 2016 a, b, 2017 b; Nakano and Lai 2017), the new species clearly differs from the other 11 quadrannulate congeners in having short epididymides in somite XVIII – XIX to somite XIX – XX that are occupying 6 – 7 annuli, thick ejaculatory ducts, and hyperboloidal developed atrial cornua (see Table 2). The new species differs from the six sexannulate species and two octannulate species by its midbody somites that are quadrannulate. Because all the present individuals of O. angustata bear undeveloped clitella, it remains uncertain whether this new species is assigned to the small or middle types. The obtained phylogenies showed that O. angustata is phylogenetically close to the three “ small-type ” species, O. brachyepididymis, O. kanaekoikeae and O. naraharaetmagarum. However, the dissected specimens possessed fully developed genital organs, and thus they could be identified at least as pre-mature leeches. Because the body lengths of the dissected individuals exceed 5 cm, it is highly possible that O. angustata can be designated as a “ middle-type ” species. Future field surveys should collect individuals of O. angustata with obvious clitella, and reveal the natural history as well as the body length group of this species. The male atrial cornua of Orobdella angustata show quite unique characteristics among those of the other Orobdella species. However, in this new species, the gastroporal duct, which is a spermatophore receptor of Orobdella leeches (Nakano 2017 b), bears the bulbous feature that is common in the other congeners whose gastroporal ducts are developed. The characteristics of the gastroporal duct of O. angustata imply that its spermatophore may not be distinctive despite the noteworthy atrial cornua of the new species.	en	Nakano, Takafumi (2018): A New Quadrannulate Species of Orobdella (Hirudinida: Arhynchobdellida: Orobdellidae) from Kii Peninsula, Japan. Species Diversity 23 (1): 43-49, DOI: 10.12782/specdiv.23.43, URL: http://dx.doi.org/10.12782/specdiv.23.43
