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Príomhchruthaitheoirí:	Jhuang, Wei-Cheng, Kusuma, Aradea Bujana, Liao, Te-Yu
Formáid:	Recurso digital
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Foilsithe / Cruthaithe:	Zenodo 2026
Ábhair:	Biodiversity Taxonomy Animalia Chordata Gobiiformes Gobiidae Stiphodon Stiphodon hadiatyae
Rochtain ar líne:	https://doi.org/10.5281/zenodo.18749366
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Stiphodon hadiatyae Nurjirana, Gustiano, Haryono & Wibowo, 2025Fig. 3 Common name: Renny’s stiphodon gobyMaterial examined.Seven specimens (four males and three females, 20.8–36.3 mm SL). Indonesia: • UNIPA-B 159 (male, 27.9 mm SL) • UNIPA-B 160 (male, 20.8 mm SL) • UNIPA-B 161 (female, 34.2 mm SL) • DOS 10029-3 (female, 26.8 mm SL) • DOS 10029-4 (male, 26.9 mm SL) • DOS 10029-5 (female, 36.3 mm SL) • DOS 10315 (male, 27.7 mm SL); Paco River, Malakoni Village, Enggano Island, Bengkulu, Southwestern Sumatra; 05°22'35.2S, 102°18'11.1E; coll. WCJ & ABK; 27 April 2025.Diagnosis.Stiphodon hadiatyae belongs to the Stiphodon sapphirinus group, diagnosed by a small body size (max size <40 mm) and usually 14 pectoral fin rays. It can be distinguished from congeners of the Stiphodon sapphirinus group by the following character combinations: first dorsal fin pointed with elongated and filamentous fourth spine, and a slightly pointed caudal fin in males; second dorsal fin rays 10; fewer premaxillary tricuspid teeth (21–29 in males; 21–31 in females); a belly entirely covered by cycloid scales. Coloration of males: head cobalt-blue with a brown snout; body greyish; median fins orange; distinct black blotch at caudal fin base; in females, first-, second-dorsal and anal fins orange to red; caudal fin proximally red to orange with four or five rows of brown spots.Description.Morphometric measurements and meristic counts are given in Tables 2–8. Body slender, cylindrical anteriorly, gradually becoming compressed posteriorly; dorsal and ventral body profiles nearly straight. Head small (20.5–25.5 % SL), approximately one-fifth to one-fourth of standard length. Snout short (3.7–6.4 % SL) and rounded, extending slightly beyond upper jaw, and upper jaw extending beyond lower jaw. Mouth inferior; upper lip thick and smooth, with a small medial cleft. Eyes large (5.6–6.8 % SL), positioned dorsolaterally. Interorbital width approximately equal to eye diameter (3.7–6.1 % SL). Anterior nostril short, tubular; posterior nostril without tubular structure.Dorsal fins VI and I, 10; in females, first dorsal fin semicircular, with the third or fourth spine longest; in males, first dorsal fin longer than female (22.1–33.0 % vs 16.4–19.0 % SL), with the fourth spine filamentous; in females, second ray of second dorsal fin longest; in males, second dorsal fin proportionally longer than female (34.1–40.0 % vs 32.1–37.7 % SL), with last ray longest. Anal fin I, 10, origin beneath origin of second dorsal fin; second ray longest in females; last ray longest in males. Pectoral fin with 14 (6) or 15 (1) rays, usually longer in males (longest ray 20.1–22.3 % SL vs 19.0–21.6 % SL in females; Table 2). Pelvic fins I, 5, origin directly below pectoral fin base, fused into a cup-like structure, supported by a folded fleshy frenulum. Caudal fin composed of 17 soft rays, including 13 principal caudal-fin rays; posterior margin slightly pointed in males, rounded or slightly truncated in females. Caudal fin longer in males (24.9–28.3 % SL vs 20.9–25.7 % SL in females; Table 2). Premaxillary teeth 21–31, fine and tricuspid (Table 3). Dentary with 1 (n = 1), 2 (n = 1), or 3 (n = 2) symphyseal teeth in males and 2 (n = 3) in females, curved and resembling canine-like shapes; dentary also with a single row of 24–34 unicuspid horizontal teeth enclosed in a fleshy sheath. Vertebral count 10 precaudal + 16 caudal vertebrae (including terminal centrum) (n = 7).Abbreviations: D 1 = first dorsal fin; D 2 = second dorsal fin; A = anal fin; P 1 = pectoral fin; P 2 = pelvic fin; C = caudal fin. * Type specimens of Stiphodon hadiatyae, data from Nurjirana et al. (2025).1 from Keith et al. (2009); 2 from Watson (1995); 3 from Watson et al. (2005); 4 from Keith et al. (2002); 5 from Keith et al. (2010); 6 from Watson et al. (1998).Scales in longitudinal row 30–39 (Table 4). Scales in transverse series divided into two regions, back 8 (n = 3) or 9 (n = 4) (Table 5); forward 10 (n = 2), 11 (n = 3), or 13 (n = 2) (Table 6). Scales in zigzag series 7 (n = 1), 8 (n = 5) or 9 (n = 1). Predorsal scales in midline 13 (n = 3) in females, and 1 (n = 1), 11 (n = 2), or 13 (n = 1) in males (Table 7 and Fig. 4). Ctenoid scales cover most of tail and lateral sides of body. Cycloid scales distributed on dorsum of head, first and second dorsal fin base, anterior trunk, belly, anal fin base, and caudal fin base. Pectoral-fin base, cheek, and operculum naked.1 from Keith et al. (2009); 2 from Watson (1995); 3 from Watson et al. (2005); 4 from Keith et al. (2002); 5 from Keith et al. (2010); 6 from Watson et al. (1998).1 from Keith et al. (2009); 2 from Watson (1995); 3 from Watson et al. (2005); 4 from Keith et al. (2002); 5 from Keith et al. (2010); 6 from Watson et al. (1998).1 from Keith et al. (2009); 2 from Watson (1995); 3 from Watson et al. (2005); 4 from Keith et al. (2002); 5 from Keith et al. (2010); 6 from Watson et al. (1998).1 from Keith et al. (2009); 2 from Watson (1995); 3 from Watson et al. (2005); 4 from Keith et al. (2002); 5 from Keith et al. (2010); 6 from Watson et al. (1998).Cephalic sensory pores small, comprising A’, B, C, D, F, H’, K’, L’, N’, and O’. Pore D singular, while others paired (Fig. 5). Pore A’ positioned at snout tip, slightly above anterior nostril; Pore B positioned laterally near posterior nostril. Pores C and D between eyes. Oculoscapular canal divided into anterior and posterior segments between pores H’ and K’. Cutaneous sensory papillae well-developed on head (Fig. 5). Infraorbital papillae short, in six transverse rows. Row c between nostril and upper lip, forming transverse and longitudinal papillae. Row d above posterior mouth edge. Rows e and i adjacent. Row f at tip of jaw. Rows g, m, and o on dorsum of head. Rows ot, oi, and os well-separated on preopercle. Row s anterior to anterior nostril. Row u 1 above pores F and H’. Row u 2 and x above pores H’, and K’. Row y above pore L’. Row z below pore H’ (Fig. 5). Urogenital papilla conical in males with a pointed posterior edge, and broader and rounded in females.Color in preservation. Sexual dichromatism well developed. In males, dorsal and side of body dark grey, ventral white; dorsum of head dark; occipital region to nape light grey. Pectoral fin base black; pectoral fin membranes translucent. First dorsal-, second dorsal- and anal-fin spines and rays white with black spots. First dorsal- and second dorsal-fin membrane light red. Anal fin light red, distal margin black. Caudal fin light red with pale-white and black spots with 8–10 transverse bars on middle of fin; a black blotch at caudal fin base. Pelvic fin rays and frenulum white; fin membrane transparent; fin rays and fin membrane with melanocytes, mainly on distal margin. In females, background of body and head cream; two black longitudinal bands extending through body and forming black blotches at end, one extending from upper lip to end of caudal peduncle through lateral midline; another extending from snout through eye to upper edge of caudal peduncle. Dorsum dusky; first and second dorsal-fin membranes almost red or transparent; usually two or three black spots along each spine and rays. Anal fin membrane transparent; black sub-margin distally; one or two black spots along each spine and rays. Caudal fin transparent with four or five transverse bands composed of black spots on middle of fin. Pectoral-fin membranes translucent. Pelvic fin translucent, usually with dense melanocyte at base and middle of rays, membranes, and frenulum.Color in life. Body and fin markings of male and female similar to those of preserved specimens. Males usually silver-grey in background of body; head grey or brown; ventral part of head to belly white; eyes cream color. First and second dorsal fins red to orange, fin-rays with 3–5 rows of black and white spots, membranes with a vertical brown stripe or 3–5 brown spots; pectoral fins transparent; anal fin white to light red with a broad black band along distal margin; caudal fin red to orange, fin-rays with 7–9 rows of brown spots, upper fin edge transparent (Figs 3 a, b, 6 a, b). In nuptial coloration, male’s cheeks metallic blue or metallic green. Females usually pale yellowish-brown in background of body and head; eyes cream color; first dorsal, second dorsal, and anal fins orange to red (small size transparent); pectoral fins transparent; caudal fin rays with four or five rows of brown spots, proximal red to orange, distal transparent; body markings similar to preserved specimens (Figs 3 c, d, 6 c).Distribution and habitat.Stiphodon hadiatyae was previously known only from its type locality, the Paco River in Malakoni Village, Enggano Island, Bengkulu, Southwestern Sumatra, Indonesia (Fig. 1). This study provides the first record of the species from outside Enggano Island, based on recent underwater photographs taken by aquarium hobbyists in the Pisang River in Padang, Sumatra (Fig. 6 d). The habitat substrate consists of gravel and sand (Fig. 2). In Enggano Island, it was found together with two other sicydiine goby species, Lentipes niasensis Harefa & Chen, 2022 and Sicyopterus micrurus (Bleeker, 1854).Molecular analysis.A total of 52 sequences (680 bp) from 25 species were analysed to reconstruct a Maximum Likelihood (ML) phylogenetic tree (Fig. 7) rooted with the outgroup taxa Awaous ocellaris and Sicyopterus lagocephalus. The tree topology revealed that sequences of Stiphodon hadiatyae form a monophyletic clade. Phylogenetically, S. hadiatyae is sister to a clade comprising S. hydoreibatus, S. sapphirinus, and S. tuivi. Together, these four species form a clade that is sister to another clade containing S. rubromaculatus, S. surrufus, and S. rutilaureus. The genetic distances between S. hadiatyae and other members of the S. sapphirinus group range from 9.4 % to 11 %. (Table 8).Remarks.The morphological description and meristic counts of Stiphodon hadiatyae in the present study are generally consistent with those reported by Nurjirana et al. (2025), but several differences are noted. These include the belly being fully covered with scales (vs always without scales in Nurjirana et al. (2025)); more second dorsal-fin rays (10 vs 9); longer snout length in males (5.9–6.3 % vs 3.4–4.6 % SL); smaller body depth at pelvic fin origin (11.1–13.3 % vs 13.1–15.2 % SL), and body depth at the anal fin origin (11.7–14.9 % vs 15.7–17.3 % SL). These discrepancies may be attributable to differences in measurement methods among researchers. In addition, prolonged storage of specimens (their material was collected by Renny K. Hadiaty and Sopian Sauri on 26 April 2015) or suboptimal preservation conditions may have resulted in scale loss or specimen shrinkage.Stiphodon hadiatyae can be distinguished from five other species from Sumatra, S. atropurpureus, S. carisa, S. maculidorsalis, S. ornatus, S. semoni, based on a combination of morphological characters. It differs from S. atropurpureus, S. carisa, and S. semoni by lacking fatty tissue behind pectoral fin base in males (vs with a white attachment), presence of an elongated and filamentous first dorsal fin spine in males (vs absence), and pectoral fin rays usually 14 (vs usually 15); from S. maculidorsalis and S. ornatus by the presence of an elongated and filamentous first dorsal fin spine in males (vs not filamentous), pectoral fin rays usually 14 (vs usually 15–16), and lacking spots on pectoral fin (vs 3–8 rows of black and white spots on pectoral fin rays in both sexes) (Maeda and Tan 2013). In addition, females of S. hadiatyae have the first and second dorsal, anal, and caudal fins orange to red, a condition not observed in females of those five Sumatran congeners (Maeda and Tan 2013; Keith et al. 2015 a).Stiphodon hadiatyae differs from S. alcedo, S. annieae, S. chlorestes, S. imperiorientis, S. larson, S. martenstyni, S. multisquamus, S. niraikanaiensis, S. pelewensis, and S. pulchellus in having typically 14 pectoral fin rays (vs 15–16) (Watson 1996, 1998; Maeda et al. 2011; Maeda 2013); from S. astilbos by presence of an elongated and filamentous first dorsal fin spine (vs absence), a black blotch on caudal fin base (vs absence) in males, more scales in longitudinal row in both sexes (30–39 vs 24–29) (Ryan 1986); from S. aureofuscus by a greater number of scales in the transverse forward series (10–13 vs 6–8) and presence of cycloid scales on belly (vs without scales) (Keith et al. 2015 b); from S. caeruleus by more scales in longitudinal row in both sexes (30–39 vs 18–21), presence of scales on belly (vs absence), and body silver-grey (vs blue) in males (Parenti and Maciolek 1993); from S. discotorquatus and S. julieni by absence of spots on pectoral fin rays (vs with alternating spots of black and white on each pectoral fin ray), and absence of spots on second dorsal fin rays (vs black and white spots on each second dorsal fin ray) in both sexes (Watson 1995; Keith et al. 2002); from S. kalfatak by lack of fatty tissue behind pectoral fin base in males (vs with a white attachment in males) and having an elongated and filamentous first dorsal fin spine (vs absence) (Keith et al. 2007); from S. mele, S. rubromaculatus, and S. surrufus by presence of an elongated and filamentous first dorsal fin spine in males (vs absence), scales on belly (vs absence), more scales in longitudinal row (30–39 vs <30), more scales in transverse forward series (10–13 vs <10) in both sexes, and body silver-grey in males (vs orange to red in males) (Keith and Marquet 2007; Keith et al. 2009); from S. percnopterygionus by fewer transverse back series (8–9 vs 10–11) and premaxillary tricuspid teeth (21–31 vs 33–46) in both sexes; from S. rutilaureus and S. zebrinus by fewer transverse back series (8–9 vs> 10) and premaxillary tricuspid teeth (21–31 vs> 40) in both sexes, and by coloration (males pale; females cream with two black longitudinal bands vs dusky transverse bars on side in both sexes; Watson et al. 1998).Stiphodon hadiatyae is morphologically similar to S. hydoreibatus, S. oatea, S. sapphirinus, and S. tuivi, as their differences in morphometric measurements and meristic counts are relatively minor (Watson 1995, 1999; Watson et al. 2005; Keith et al. 2010). However, S. hadiatyae can be distinguished from S. hydoreibatus, S. oatea, S. sapphirinus, and S. tuivi by fewer premaxillary tricuspid teeth (21–36 vs 35–52 in S. hydoreibatus, 40–44 in S. oatea, 31–51 in S. sapphirinus, and 34–53 in S. tuivi). Additionally, it differs from S. oatea, S. sapphirinus, and S. tuivi by fewer scales in the transverse back series (8–9 vs> 10). Furthermore, S. hadiatyae can be distinguished from all these species by differences in body coloration and distribution (Fig. 6) (Watson 1995, 1999; Watson et al. 2005; Keith et al. 2010).Since females of the Stiphodon sapphirinus group are superficially similar in color pattern, a comparative diagnosis based on female characters is provided. Females of S. hadiatyae can be distinguished from S. discotorquatus, S. julieni, and S. tuivi by pectoral-fin rays without alternating black and white spots (vs present); from S. caeruleus by belly with scales (vs absent); from S. astilbos, S. oatea, and S. percnopterygionus by black spots on first and second dorsal and proximal caudal fin rays (vs absent in those species); from S. hydoreibatus and S. sapphirinus by black spots on first dorsal fin rays (vs absent); from S. mele, S. rubromaculatus, and S. surrufus by more predorsal midline scales (vs absent or few) and black spots on proximal caudal fin (vs absent); from S. rutilaureus and S. zebrinus by sides without dusky transverse bars (vs present).The ML tree based on COI genes showed that the genus Stiphodon is monophyletic and can be divided into two groups, consistent with the results of Keith et al. (2011), Taillebois et al. (2014), and Jhuang et al. (2024). However, relationships within the “ S. sapphirinus group ” are poorly supported, resulting in uncertainty regarding the phylogenetic positions of some species within this group, which may be attributable to incomplete taxon sampling. In addition, several Stiphodon species are not recovered as monophyletic, potentially due to misidentifications or the limited resolution of single-gene analyses, suggesting that additional genetic markers are needed to clarify their taxonomic status.Based on both morphological and molecular evidence, the present study indicates that Stiphodon hadiatyae is sister to the clade comprising S. hydoreibatus, S. sapphirinus, and S. tuivi, and belongs to the “ S. sapphirinus group ” (Fig. 7). Stiphodon hadiatyae is distributed in western Sumatra and Enggano Island, whereas S. tuivi is currently known only from Marquesas, S. hydoreibatus from Futuna to Samoa, and S. sapphirinus from New Caledonia, Fiji, and Vanuatu (Fig. 1). Although phylogenetic relationships within the S. sapphirinus group remain weakly supported due to incomplete taxon sampling and the use of a single mitochondrial marker, the observed distribution pattern is broadly consistent with the biogeographic scenario proposed by Taillebois et al. (2014), which suggested that Stiphodon dispersed from the Indonesian shelf to Melanesia, followed by subsequent diversification.Taxonomic key to distinguish Stiphodon in SumatraAbbreviations: D 1 = first dorsal fin; D 2 = second dorsal fin; A = anal fin; P 1 = pectoral fin.Comparative material examinedStiphodon annieae: five specimens • DOS 07332, 22.1–24.8 mm SL, Maluku, aquarium trade, 15, January, 2020, coll. WC Jhuang. Stiphodon atropurpureus: 20 specimens • DOS 09876, 23.4–42.3 mm SL, Luzon, 23 June 2023, coll. AC Dimaquibo, CC Yu & NS Leung. Stiphodon carisa: six specimens • DOS 08610, 32.1–37.3 mm SL, Sumatra, aquarium trade, 14 March 2022, coll. WC Jhuang. Stiphodon chlorestes: one specimen • PNM 15728, holotype, 29.6 mm SL, Luzon, 12 January, 2023, coll. WC Jhuang, AC Dimaquibo, GC Liu & KH Chen; one specimen • ASIZP 0081725, paratype, 30.3 mm SL, Taiwan, 15 October, 2022, coll. WC Liang; five specimens • ASIZP 0081724 • DOS 09263 • NMMBA-P 038532–33 • PNM 15729, paratypes, 28.6–40.2 mm SL, Luzon, 12 January, 2023, coll. WC Jhuang, AC Dimaquibo, GC Liu & KH Chen. Stiphodon imperiorientis: one specimen • DOS 09250, 27.7 mm SL, Taiwan, 14 March, 2023, coll. WC Jhuang; one specimen • DOS 09258, 31.1 mm SL, Taiwan, 7 April, 2023, coll. W. C. Liao; one specimen • DOS 09259, 39.2 mm SL, Taiwan, 9 April, 2023, coll. WC Jhuang. Stiphodon multisquamus: two specimens • DOS 03177, 58.3–62.4 mm SL, Hainan, 7 September, 2020, coll. WC Jhuang; one specimen • DOS 09260, 26.7 mm SL, Taiwan, 18 July, 2021, coll. MT Zhou. Stiphodon palawanensis: five specimens • DOS 09268, 43.6–51.4 mm SL, Palawan, 24 April, 2023, coll. WC Jhuang, WC Huang & BA Balisco. Stiphodon pelewensis: ten specimens • DOS 10737, 36.6–45.6 mm SL, Papua, 13 September, 2024, coll. WC Jhuang. Stiphodon pulchellus: two specimens • DOS 09318, 51.1–55.6 mm SL, Palawan, 26 April, 2023, coll. WC Jhuang, WC Huang & BA Balisco. Stiphodon percnopterygionus: five specimens • DOS 09249, 21.1–31.0 mm SL, Luzon, 12 January 2023, coll. WC Jhuang, AC Dimaquibo, GC Liu & KH Chen. Stiphodon rutilaureus: 16 specimens • DOS 10740, 20.8–27.7 mm SL, Biak, aquarium trade, 20 February 2025, coll. WC Jhuang. Stiphodon semoni: six specimens • DOS 10736, 28.8–33.0 mm SL, Papua, 13 September 2024, coll. WC Jhuang. Stiphodon surrufus: 11 specimens • DOS 10735, 24.9–30.3 mm SL, Papua, 13 September 2024, coll. WC Jhuang.

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