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Kunishima, Taiga, Palla, Herminie P., Tachihara, Katsunori, Maeda, Ken (2022): First Records of an Estuarine Goby Acentrogobius ocyurus (Gobiiformes: Gobiidae) from Japan and the Sulu Sea in the Philippines. Species Diversity 27 (1): 129-138, DOI: 10.12782/specdiv.27.129, URL: http://dx.doi.org/10.12782/specdiv.27.129

Rhinogobius estrellae Maeda, Kunishima & Palla, new species [New English name: Estrella goby] (Figs. 1���5; Tables 2���6) Material examined. Eight males and 16 females from Narra, Palawan Island in the Philippines. Holotype. NSMT-P 140091, male (40.6 mm SL), Estrella Falls, 13 May 2016, coll. K. Maeda, T. Kunishima, and H. P. Palla. Paratypes. NSMT-P 140092, female (38.9 mm SL), same data as holotype; URM-P 49295���49301, 1 male (40.7 mm SL) and 6 females (36.1���40.9 mm SL), same data as holotype; URM-P 49302, female (43.0 mm SL), creek in front of the Estrella Village Barangay Hall, 28 May 2018, coll. K. Maeda, H. Kobayashi, and H. P. Palla; URM-P 49303���48306, 2 males (37.8���39.6 mm SL) and 2 females (36.4���45.3 mm SL), Estrella Falls, 28 May 2018, coll. K. Maeda, H. Kobayashi, and H. P. Palla; WPU-PPC-P 50���54, 1 male (37.5 mm SL) and 4 females (35.5���41.1 mm SL), same data as holotype; WPU-PPC-P 55���59, 3 males (37.3���38.4 mm SL) and 2 females (33.2���37.8 mm SL), Estrella Falls, 28 May 2018, coll. K. Maeda, H. Kobayashi, and H. P. Palla. Diagnosis. Pectoral fin with 14���16 rays (usually 15 rays). Nape and posterior part of occipital region covered by cycloid scales; scaled area extending anteriorly to around a vertical line through posterior margin of preopercle or a little posterior to this line. Longitudinal scales 27���31, predorsal scales 3���11. Number of vertebrae 26. Cephalic sensory pore system usually with B��, C, D(S), E, F, H��, K��, L��, M��, N, and O��, but often lacking one or both side(s) of pore E. Transverse rows of sensory papillae on cheek. In preservative, lateral and dorsal sides of body with dark brown mesh-like markings, snout and cheek with three dark brown stripes, and pectoral fin with dark brown band vertically across proximal part of the upper and middle rays except lower 2���6 rays. In life, upper and middle parts of pectoral-fin base with bright white vertical band. Description. Body nearly cylindrical anteriorly and somewhat compressed posteriorly. Head depressed and larger in male than female (head length 33.5���36.0 vs 30.3���32.9% of SL; Fig. 3). Eyes located dorsolaterally. Mouth terminal and oblique with thick upper and lower lips. Anterior tips of upper and lower jaws almost even or upper jaw slightly protruding beyond lower jaw. Posterior end of upper jaw always exceeding a vertical line through anterior margin of eye. Mouth larger in male than female (upper-jaw length 14.1���16.1 vs 10.5���11.9% of SL; Fig. 3). Anterior nostril short tubular, posterior nostril a pore. Cephalic sensory pore system usually with B��, C, D(S), E, F, H��, K��, and L�� in oculoscapular canal and M��, N, and O�� in preopercular canal, but 5/ 24 specimens lacking one pore C, 2/ 24 specimens lacking both pores C, 7/ 24 specimens lacking one pore E, 5/ 24 specimens lacking both pores E, 3/ 24 specimens having pore G on one side, and 1/ 24 specimens lacking one pore N. One specimen having an additional single pore anterior to another single pore D. Arrangement of cutaneous sensory papillae of head shown in Fig. 4. Cheek having two longitudinal rows of papillae and 4���6 transverse rows both between eye and upper longitudinal row and between upper and lower longitudinal rows. Vertebrae 10+16=26 (n=9) or 11+15=26 (n=1), P-V 3/II II I I 0/9 (n=9) or 3/II I II II 0/9 (n=1). First dorsal fin usually with six spines supported by six pterygiophores, but 1/ 24 specimen with seven spines supported by seven pterygiophores. Second dorsal fin usually with one spine and eight soft rays, but 3/ 24 specimens with one spine and seven soft rays. First- and second-dorsal-fin bases separated each other by a small interval (0.3���3.9% of SL). First dorsal fin rounded, usually almost semi-circular; posterior tips of fin (usually tips of fourth to sixth spines) not reaching second dorsal fin origin (n=13), just touching base of second-dorsal-fin spine (n=6), or exceeding base of second-dorsal-fin spine but not reaching base of first soft ray of second dorsal fin (n=5). Anal fin usually with one spine and eight soft rays, but 3/ 24 specimens with one spine and seven soft rays and 2/ 24 specimens with one spine and nine soft rays. Caudal fin with 17 segmented rays, including 12 (n=3), 13 (n=9), or 14 (n=12) branched rays; posterior margin rounded. Male having larger second dorsal, anal, and caudal fins than female (second-dorsal-fin length 34.6���38.7 vs 27.7���30.2% of SL, anal-fin length 31.3���33.7 vs 24.2���29.3% of SL, and caudal-fin length 28.6���31.7 vs 24.4���28.0% of SL; Fig. 3). Pectoral fin with 14���16 rays (usually 15 rays) (Table 2). Pelvic fin with one spine and five soft rays; pelvic fins joined together to form a cuplike disk with fleshy bilobed frenum. Posterior tip of pelvic fin located below middle of first-dorsal-fin base. Ctenoid scales covering lateral side of body and dorsal and ventral sides of caudal peduncle. Nape and posterior part of occipital region covered by cycloid scales; scaled area extending anteriorly around vertical line through posterior margin of preopercle (above pore H��) or to a little posterior to this line (above area between pores H�� and K��). Other regions in head naked. Belly and first-dorsal-fin base covered with cycloid scales. A few cycloid scales also occurring on trunk behind pectoral fin, along bases of second dorsal, anal, and caudal fins, and on proximal part of caudal fin. Pectoral-fin base and breast (prepelvic area) probably naked (at least invisible without staining by arizarin red; see Suzuki et al., 2016). Longitudinal scales 27���31 (usually 28���30), transverse scales 8���10 (usually 9), transverse scales in caudal peduncle always 7, and predorsal scales 3���11 (Tables 3���5). Colour in preservative (Fig. 1): Background of head and body cream. Head with three dark brown stripes; upper one running on snout and connecting upper lip and eye; lower two running diagonally across cheek and connecting upper lip and posterior margin of preopercle. These stripes often broken-up with gaps, branches, and additional dots. Posterior margin of opercle fringed by narrow dark brown vertical band. Pectoral-fin base dark brown with thin, whitish vertical line along bases of middle and ventral rays. Scales on dorsal and lateral sides of body having dark brown margins forming mesh-like patterns. Darkness of scale margins variable and body often showing four or five dark brown saddles from trunk to caudal peduncle. First and second dorsal fins greyish brown with multiple transparent or pale grey spots on proximal and middle parts. Anal fin pale grey or greyish brown, often darker in males. Caudal fin pale grey or greyish brown with 6���10 dark brown vertical bands, but ventral part lacking these bands. Pectoral fin pale grey or greyish brown with a dark brown vertical band proximally on upper and middle parts. Pelvic fin pale grey or greyish brown. Colour in life (Fig. 5): Background of head and body pale reddish- or yellowish grey but more whitish on ventral side; ventral surface of head and belly often tinged with orange. Dark brown markings on head and body described in ���colour in preservative��� present but more reddish. Background of cheek and opercular region often tinged with greyish blue. Pectoral-fin base black with a bright white vertical band posteriorly on upper to middle parts, and often tinged with greyish blue anteriorly. Background of body tinged with sky blue. First and second dorsal fins reddish brown or orange with many white spots on proximal and middle parts. Anal fin reddish brown or orange but proximal and posterior parts greyish blue. Caudal fin reddish brown or orange with 6���10 translucent or white transverse bands on middle part. Pectoral fin translucent often lightly tinged with orange, and proximal parts of some ventral rays white. Pelvic fin pale orange proximally and light grey or greyish blue distally. Distribution. Rhinogobius estrellae is endemic to Palawan Island, Philippines. Most of the specimens examined were collected from a pool and the lower reaches of a stream below the Estrella Falls (9��21'26"N 118��23'50"E), located where a steep stream from Mt. Victoria (1,726 m) comes out to a large plain in Narra, Sulu Sea-side of the island. This new species was abundant in the pool below the first major fall with many cyprinids and gobies but absent in the reaches above it which are occupied by sicydiine gobies including Sicyopus zosterophorus (Bleeker) and Lentipes palawanirufus Maeda & Palla, and eels of the genus Anguilla Schrank (unidentified to species level). The new species was also found in a creek in front of the Barangay Hall of Estrella Village (9��21'28"N 118��24'58"E), drawing from the stream below Estrella Falls. The stream flows into Malatgao River. When we surveyed a site in the middle reaches of the Malatgao River (9��20'16.7"N 118��27'27.0"E), no Rhinogobius were found there. The water at Estrella Falls was cool even during the hottest dry season (May) in this region (water temperature was 26.5��C when measured on 13 May, 2016), while water in the Malatgao River was warmer (appeared to be nearly 30��C although we did not measure). Etymology. The type locality of the new species is Estrella Falls in Barangay Estrella Village. Therefore, the new species is named as Rhinogobius estrellae, derived from Estrella, and the Latin suffix -e., Published as part of Maeda, Ken, Shinzato, Chuya, Koyanagi, Ryo, Kunishima, Taiga, Kobayashi, Hirozumi, Satoh, Noriyuki & Palla, Herminie P., 2021, Two new species of Rhinogobius (Gobiiformes: Oxudercidae) from Palawan, Philippines, with their phylogenetic placement, pp. 81-98 in Zootaxa 5068 (1) on pages 84-89, DOI: 10.11646/zootaxa.5068.1.3, http://zenodo.org/record/5702104, {"references":["Suzuki, T., Shibukawa, K., Senou, H. & Chen, I. - S. (2016) Redescription of Rhinogobius similis Gill 1859 (Gobiidae: Gobionellinae), the type species of the genus Rhinogobius Gill 1859, with designation of the neotype. Ichthyological Research, 63 (2), 227 - 238. https: // doi. org / 10.1007 / s 10228 - 015 - 0494 - 3"]}

Rhinogobius tandikan Maeda, Kobayashi & Palla, new species [New English name: Tandikan goby] (Figs. 1, 3, 6���8; Tables 2���6) Material examined. Thirteen males and 16 females from Cayulo River, Bahile, Puerto Princesa City, Palawan Island in the Philippines. Holotype. NSMT-P 140093, male (43.6 mm SL), 1 June 2018, coll. K. Maeda, H. Kobayashi, and H. P. Palla. Paratypes. NSMT-P 140094, female (45.7 mm SL), 31 May 2018 coll. K. Maeda, H. Kobayashi, and H. P. Palla; URM-P 49307���49315, 7 males (36.1���46.9 mm SL) and 2 females (37.3���38.9 mm SL), 31 May 2018 coll. K. Maeda, H. Kobayashi, and H. P. Palla; URM-P 49316���49323, 2 males (40.8���44.8 mm SL) and 6 females (35.6���45.2 mm SL), same data as holotype; WPU-PPC-P 60���64, 5 females (30.8���36.5 mm SL), 31 May 2018 coll. K. Maeda, H. Kobayashi, and H. P. Palla; WPU-PPC-P 65���69, 3 males (43.6���46.1 mm SL) and 2 females (39.6���41.3 mm SL), same data as holotype. Diagnosis. Pectoral fin with 16 or 17 rays. Nape and posterior part of occipital region covered by cycloid scales; scaled area extending anteriorly to around a vertical line through posterior margin of preopercle or a little posterior to this line. Longitudinal scales 29���34, predorsal scales 5���13. Number of vertebrae 26. Cephalic sensory pore system usually with B��, C, D(S), E, F, H��, K��, L��, M��, N, and O��, but often lacking one or both side(s) of pore C. Transverse rows of sensory papillae on cheek. In preservative, lateral and dorsal sides of body without clear meshlike markings, obscure markings on cheek, and upper half of pectoral fin with dark brown band vertically across proximal part. In life, pectoral-fin base with a bright white vertical band reaching near bottom of fin. Description. Body nearly cylindrical anteriorly and somewhat compressed posteriorly. Abdomen of male often thin. Head depressed and larger in male than female (head length 32.1���37.4 vs 30.8���33.4% of SL; Fig. 3). Eyes located dorsolaterally. Mouth terminal and oblique with thick upper and lower lips. Anterior tips of upper and lower jaws almost even or upper jaw slightly protruding beyond lower jaw. Posterior end of upper jaw always exceeding a vertical line through anterior margin of eye. Mouth larger in male than female (upper-jaw length 13.3���20.9 vs 10.1���12.7% of SL; Fig. 3). Anterior nostril short tubular, posterior nostril a pore. Cephalic sensory pore system usually with B��, C, D(S), E, F, H��, K��, and L�� in oculoscapular canal and M��, N, and O�� in preopercular canal, but 9/ 29 specimens lacking one pore C, 7/ 29 specimens lacking both pores C, 4/ 29 specimens lacking one pore E, 1/ 29 specimen having two pores E on one side, 4/ 29 specimens having pore G on one side, 1/ 29 specimen having pores G on both sides, and 5/ 29 specimens lacking one pore N. One specimen having combined pores K�� and L�� on one side (to be a groove without roof). Arrangement of cutaneous sensory papillae of head shown in Fig. 7. Cheek having two longitudinal rows of papillae and 3���6 transverse rows both between eye and upper longitudinal row and between upper and lower longitudinal rows. Vertebrae 10+16=26 (n=10), P-V 3/II II I I 0/9 (n=10). All males had damage to the caudal fin, except holotype; 10/ 16 female specimens with damaged caudal fin (Figs. 1, 3). Other fins of male and female sometimes damaged also. First dorsal fin with six spines supported by six pterygiophores. Second dorsal fin usually with one spine and eight soft rays. First and second dorsal-fin bases separated each other by a small interval (0.2���4.4% of SL). First dorsal fin rounded, usually almost semi-circular; posterior tips of fin (tips of fourth to sixth spines) not reaching second dorsal fin origin (N=9), just touching base of second dorsal-fin spine (n=12), or exceeding base of second dorsal-fin spine but not reaching base of first soft ray of second dorsal fin (n=8). Anal fin usually with one spine and eight soft rays, but 4/ 29 specimens with one spine and seven soft rays. Caudal fin usually with 17 segmented rays but 3/ 29 specimens with 16 segmented rays, including 11 (n=1), 12 (n=6), 13 (n=15), or 14 (n=22) branched rays (unknown in four specimens due to fin damage); posterior margin rounded in non-damaged individuals. Male having larger second dorsal and anal fins than female (second dorsal-fin length 33.8���43.6 vs 29.5���35.3% of SL and anal-fin length 30.2���38.3 vs 26.7���32.7% of SL), and nondamaged male with larger caudal fin than female (Fig. 3). Pectoral fin with 16 or 17 rays (Table 2). Pelvic fin with one spine and five soft rays; pelvic fins joined together to form a cuplike disk with fleshy bilobed frenum. Posterior tip of pelvic fin located below middle of first-dorsal-fin base. Ctenoid scales covering lateral side of body and dorsal and ventral sides of caudal peduncle. Nape and posterior part of occipital region covered by cycloid scales; scaled area extending anteriorly around vertical line through posterior margin of preopercle (above pore H��) or to a little posterior to this line (above area between pores H�� and K��). Other regions in head naked. Belly and first dorsal-fin base covered with cycloid scales. A few cycloid scales also present on trunk behind pectoral fin, along bases of second dorsal, anal, and caudal fins, and on proximal part of caudal fin. Pectoral-fin base and breast (prepelvic area) probably naked (at least invisible without staining by arizarin red; see Suzuki et al. 2016). Longitudinal scales 29���34 (usually 30 or 31), transverse scales 8���10 (usually 9 or 10), transverse scales in caudal peduncle always 7, and predorsal scales 5���13 (Tables 3���5). Colour in preservative (Fig. 1): Background of head and body brown dorsally and laterally and cream ventrally. Snout with obscure dark brown stripe connecting upper lip and eye. Cheek often with two obscure, dark brown intermittent lines. Posterior margin of opercle fringed by dark brown vertical band. Pectoral-fin base blackish except cream or pale brown bottom. Four to eight dark brown transverse bars or round blotches aligned along lateral midline of body. First and second dorsal fins pale grey with 3���7 brown horizontal stripes on proximal and middle parts, but anterior half of first dorsal fin lacking this marking. Anal fin greyish brown but light brown proximally. Caudal fin pale grey or greyish brown with 6���11 dark brown vertical bands, but dorsal and ventral parts lacking these bands. Pectoral fin pale greyish brown with a dark brown vertical band proximally on upper half. Pelvic fin greyish brown. Colour in life (Fig. 8): Background of head and body olive green or yellowish grey. Snout with reddish brown stripe connecting upper lip and eye. Cheek with two reddish brown longitudinal intermittent lines and several reddish brown spots. Posterior margin of opercle fringed by reddish-brown vertical band. Pectoral-fin base black or dusky with a large greyish blue blotch anteriorly, and with a bright white vertical band posteriorly. Four to eight, dark brown transverse bars or round blotches aligned along lateral midline of body. Irregularly arranged dark brown markings dorsally on body. Centre of lateral scales tinged with sky blue. First and second dorsal fins light yellowish brown with 3���7 dark brown horizontal stripes on proximal and middle parts and reddish brown on distal part, but anterior half of first dorsal fin reddish brown without clear marking. Anal fin reddish brown or orange but yellowish proximally. Caudal fin reddish brown dorsally and ventrally and whitish with 6���11 dark reddish-brown transverse bands on middle part. Pectoral fin translucent but proximal parts of some ventral rays white. Pelvic fin reddish brown proximally and light grey distally. Distribution. Rhinogobius tandikan is endemic to Palawan Island. It was found only in the Cayulo River, Bahile, northern part of Puerto Princesa City. The Cayulo River is a small stream (length approximately 4 km including 1 km of an estuary) flowing into Ulugan Bay on South China Sea-side of the island. The stream system is surrounded by mountains of about 200���400 m above sea level. Cayulo Falls, composed of a series of small waterfalls, is located at the middle eaches of this stream (10��02'12"N 118��45'03"E). Rhinogobius tandikan was found at both the lower and upper reaches of this waterfall, as well as pools in the middle of the falls. The new species was abundant in this site with many other goby species, cyprinids, and a jungle perch species. Etymology. Tandikan is a local name of Palawan peacock-pheasant, Polyplectron napoleonis Lesson, known as a symbol of Puerto Princesa City. The new goby species is named Rhinogobius tandikan, as it has blue markings on the body, suggestive of the plumage of the Palawan peacock-pheasant, and as the type locality is located in the northern part of Puerto Princesa City. The new specific name is a noun in apposition. Comparison. Rhinogobius similis Gill, 1859 is the only species previously known in the genus having transverse rows of sensory papillae on cheek (Suzuki et al., 2016), and our two new species share this unique feature with R. similis. However, the two species in Palawan are distinguished from R. similis in having fewer rays in the pectoral fin (14���17 vs 18���19), predorsal squamation composed of cycloid scales (vs ctenoid scales), anterior-most dorsal scale located around a vertical line through posterior margin of the preopercle or more posteriorly (vs dorsal squamation reaching anteriorly near the posterior margin of the eye), and proximal part of the caudal fin covered by cycloid scales (vs by ctenoid scales). Rhinogobius estrellae shares many characters with R. tandikan, but the former differs from the latter by having fewer pectoral-fin rays (usually 15 vs 16���17; Table 2), fewer scale counts (longitudinal scales 27���31 vs 29���34, predorsal scales 3���11 vs 5���13; Tables 3, 5), and colourations of the body. In preservative, R. estrellae has dark brown mesh-like markings on the lateral and dorsal sides of the body (vs rather uniformly brown in R. tandikan), three dark brown stripes on the snout and cheek (vs often obscure), and a dark brown vertical band on the proximal part of the upper and middle pectoral-fin rays except lower 2���6 rays (vs this band restricted to upper half of the fin). In life, R. estrellae has a more reddish body than the yellowish or olive green R. tandikan, and the former has a bright white vertical band restricted to the upper and middle parts of the pectoral-fin base while this band reaches nearly the bottom of the fin in R. tandikan. Arrangements of the cephalic sensory pore system are basically the same between the two species, but R. estrellae more often lacks pore E on one or both sides than in R. tandikan (12/24 vs 5/29) while R. tandikan more often lacks pore C on one or both sides than in R. estrellae (16/29 vs 7/24). In addition to the transverse rows of sensory papillae on cheek (present vs absent), there are many differences between the two new species from Palawan and three other Philippine species, R. carpenteri Seale, 1910, R. bucculentus (Herre, 1927), and R. philippinus (Herre, 1927), such as numbers of the vertebrae (26 vs 27���29), first-dorsalfin pterygiophores (usually 6 vs usually 7), first-dorsal-fin spines (usually 6 vs usually 7 in R. bucculentus and 6 or 7 in R. carpenteri and R. philippinus), pectoral-fin rays (14���17 vs 17���20), having fewer scales in the longitudinal scales (27���34 vs 35���43), in the transverse scales (8���10 vs 12���18), and in the transverse scales in caudal peduncle (7 vs 9���11), and presence of scales on belly behind the pelvic fins (vs absent in R. carpenteri and R. bucculentus). The latter three species are distributed in Luzon, northern Philippines (see Discussion). Molecular phylogenetic analysis. In the ML phylogenetic tree using the aligned 16,409 bp of mitochondrial genomes (Fig. 9), the species in Rhinogobius were divided into two robust clades. One clade included R. similis and the two new species from Palawan, while all other species belonged to another clade. Six specimens of R. estrellae and four specimens of R. tandikan formed each well-supported monophyletic clade and were sister species to each other. Rhinogobius similis was a sister to this pair., Published as part of Maeda, Ken, Shinzato, Chuya, Koyanagi, Ryo, Kunishima, Taiga, Kobayashi, Hirozumi, Satoh, Noriyuki & Palla, Herminie P., 2021, Two new species of Rhinogobius (Gobiiformes: Oxudercidae) from Palawan, Philippines, with their phylogenetic placement, pp. 81-98 in Zootaxa 5068 (1) on pages 89-94, DOI: 10.11646/zootaxa.5068.1.3, http://zenodo.org/record/5702104, {"references":["Suzuki, T., Shibukawa, K., Senou, H. & Chen, I. - S. (2016) Redescription of Rhinogobius similis Gill 1859 (Gobiidae: Gobionellinae), the type species of the genus Rhinogobius Gill 1859, with designation of the neotype. Ichthyological Research, 63 (2), 227 - 238. https: // doi. org / 10.1007 / s 10228 - 015 - 0494 - 3","Gill, T. N. (1859) Notes on a collection of Japanese fishes, made by Dr. J. Morrow. Proceedings of the Academy of Natural Sciences of Philadelphia, 11, 144 - 150.","Seale, A. (1910) New species of Philippine fishes. The Philippine Journal of Science, Section A, 4, 491 - 543, pls. 1 - 13.","Herre, A. W. C. T. (1927) Gobies of the Philippines and the China Sea. The Philippine Bureau of Science Monographic Publications of Fishes, 23, 1 - 352, pls. 1 - 30."]}

Kunishima, Taiga, Maeda, Ken, Inui, Ryutei, Hibino, Yusuke (2021): First Japanese Record of Muraenichthys gymnopterus (Anguilliformes, Ophichthidae) from Ishigaki-jima Island, Ryukyu Archipelago. Species Diversity 26: 343-349, DOI: 10.12782/specdiv.26.343

Kunishima, Taiga, Tanaka, Ryosuke, Hirashima, Kentarou, Maeda, Ken (2021): Northernmost Record of Stiphodon multisquamus (Gobiiformes: Oxudercidae) Based on a Specimen from Wakayama, Japan. Species Diversity 26 (1): 37-41, DOI: 10.12782/specdiv.26.37, URL: http://dx.doi.org/10.12782/specdiv.26.37

Acanthogobius insularis Shibukawa and Taki (1996) is a threatened goby species, previously known to be restricted to estuaries on Amami-oshima Island and northern part of Okinawa-jima Island. However, we found this species in two estuaries in the southern part of Okinawa-jima Island (Toyosaki and Manko), which are separated from their habitats in the northern part of the island by approx. 60 km of coral reefs. Morphologies of four specimens collected in Toyosaki and Manko examined in the present study correspond to the original description of A. insularis. This study reports that Toyosaki is the southernmost habitat of this species based on specimens. Because it is thought that A. insularis move to unknown spawning habitats in winter and because only one individual has been found in Manko, it is necessary to understand the habitats that these newly discovered small populations use, in their entirety.

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