Larvae and juveniles of the plunderfishes <Emphasis Type="Italic">Artedidraco shackletoni</Emphasis> and <Emphasis Type="Italic">A.</Emphasis><Emphasis Type="Italic">skottsbergi</Emphasis> (Notothenioidei,Artedidraconidae) from the Indian sector of the Southern Ocean

Early life stages of Artedidraco skottsbergi and A. shackletoni were collected off Adélie Land. The morphology and pigmentation pattern of nine larvae and juveniles of A. skottsbergi between 17.2 and 21.4 mm in standard length (SL), and of two juveniles of A. shackletoni measuring 25.1 mm SL were described. A. skottsbergi was characterized by a heavily pigmented body, except for the caudal peduncle, with distinctively dense pigmentation on the ventrolateral half of the body and caudal section (17.2–17.9 mm SL). Furthermore, they had no pigmentation on the pectoral fin base until they attained 21.4 mm SL. Juvenile A. shackletoni had a heavily pigmented body except for the ventral side of the abdomen and the anal fin base. The proximal part of the dorsal fin and most of the anal fin were covered with melanophores. Although knowledge of larval and juvenile Artedidraco species is limited, the distribution of melanophores on the fins, pectoral fin base and caudal peduncle at each developmental stage may be useful for species identification.     

Embryonic, larval and juvenile development of the roughskin sculpin,Trachidermus fasciatus (Scorpaeniformes: Cottidae)

Embryonic, larval and juvenile development of the catadromous roughskin sculpin,Trachidermus fasciatus, were described using eggs spawned in an aquarium. The eggs, measuring 1.98–2.21 mm in diameter, were light reddish-yellow and had many oil globules, 0.05–0.18 mm in diameter. Hatching occurred 30 days after spawning at 2.3–11.3°C. The newly-hatched larvae, measuring 6.9–7.3 mm BL, had a single oil globule, 9–10+25–26=34–36 myomeres and 6 or 7 large stellate melanophores dorsally along the gut. The yolk was almost resorbed, number of pectoral-fin rays attained 16–17, and two parietal, one nuchal and four preopercular spines were formed, 5 days after hatching, at 8.2–8.4 mm BL. The oil globule disappeared, and one supracleithral spine was formed, 11 days after hatching, at 8.9–9.5 mm BL. Notochord flexion began 15 days after hatching, at 9.7–10.3 mm BL. A posttemporal spine was formed 20 days after hatching, at 10.7–10.9 mm BL. The first dorsal fin spines (VII–VIII), second dorsal fin and anal fin rays (18–19, 16–18, respectively) appeared 23 days after hatching, at 12.0–13.7 mm BL. The pelvic fin spine and rays (I, 4) were formed and black bands on the head and sides of the body began to develop 27 days after hatching, at 13.8–15.8 mm BL. Newly-hatched larvae swam just below the surface in the aquaria. Preflexion larvae (8.9–9.5 mm BL), in which the oil globule had disappeared, swam in the middle layer, while juveniles (13.8–15.8 mm BL) began swimming on the bottom of the aquaria. Swimming behavior observed in the aquaria suggested that the fish started to change to a demersal existence at the juvenile stage.     

Juveniles of two sillaginids,Sillago aeolus andS. sihama,occurring in a surf zone in the Philippines

Sillaginid juveniles collected from the surf zone at Tigbauan, Iloilo, Philippines, between May 1986 and September 1987 were identified asSillago aeolus (n=702, 8.9–26.0 mm SL) andS. sihama (n=3414, 8.6–22.9 mm SL), based on the numbers of dorsal and anal soft fin rays and vertebrae. The two species were easily distinguishable by the pattern of melanophores distributed on the caudal fin base,S. aeolus having a triangular-shaped cluster, whereas the melanophores formed a vertical line inS. sihama. The ratios of pre-anal and caudal peduncle lengths to SL also differed between the species, both being higher inS. aeolus. The occurrence ofS. aeolus was limited to the dry season, from January to March. On the other hand,S. sihama occurred year-round, although a peak was observed in the dry season, from November to April.     

Description and ecology of larvae and juveniles of three native cypriniforms of coastal southern California

Larval series of the Santa Ana sucker, Catostomus santaanae (Federally Threatened), arroyo chub, Gila orcutti (California Species of Special Concern), and Santa Ana speckled dace, Rhinichthys osculus (California Species of Special Concern) are described from wild-caught specimens from the Los Angeles and Santa Ana river drainages. Santa Ana sucker larvae are elongate, having 41–46 myomeres and a distinctive paired-triangle patch of melanophores over the midbrain. Melanophores present on the snout, dorsal body, lateral midline, dorsal gut, postanal ventral body, and caudal fin. Preanal length 74–79% in body length (BL), typical of catostomids. Arroyo chub larvae relatively deep-bodied, 36–39 myomeres, and a heart-shaped patch of melanophores over the midbrain with a line of melanophores trailing posteriorly. Heavy pigment present on the snout, lower jaw, dorsal body, lateral midline, gill arches, dorsal gut, postanal ventral body, and caudal fin; short preanal length of 65–72% BL, typical of native North American cyprinids. Santa Ana speckled dace are similar to arroyo chub except for having less pigment on the ventral gut, large distinct melanophores on the ventrolateral caudal peduncle, a wedge-shaped patch of midbrain melanophores with no distinct line trailing posteriorly, and lateral midline melanophores that do not extend anteriorly. These three species often occur together and with nonnative cyprinids. Characters distinguishing them from other local larvae, including southern fathead minnow, Pimephales promelas confertus and red shiner, Cyprinella lutrensis, are discussed with their habitat preferences.     

Osteological development in the Japanese sardine,Sardinops melanostictus

The development of all osteological elements, except scales, of the Japanese sardine,Sardinops melanostictus, is described from newly-hatched larvae to adult fishes. Newly-hatched larvae lacked osteological elements. Part of the head skeleton began to develop in 53 hour old larvae (4.2 mm in notochord length [NL]). Larvae at the first-feeding stage (77 hours, 5.5 mm NL) possessed several elements of the head skeleton and pectoral fin supports. In a 10.5 mm NL specimen, part of the caudal and dorsal fin supports were apparent. The centra appeared in specimens 18–22.7 mm in standard length (SL). Gill rakers were first observed in the lower branchial arches at 13 mm NL and spine-like processes with spiny nodules from about 25 mm SL. The distance between the predorsal and first dorsal proximal radial relative to SL rapidly decreased with forward translocation of the dorsal fin and became constant beyond approximately 34 mm SL. At this stage, most basic osteological elements were established. Completion of the osteological structure was characterized by the disappearance of the dentary teeth at 60–70 mm SL. Based on the osteological development, ontogenetic intervals consisting of four periods and eight phases were recognized.     

The Saurida undosquamis group (Aulopiformes: Synodontidae), with description of a new species from southern Japan

Four lizardfishes of Saurida (family Synodontidae), S. undosquamis, S. umeyoshii sp. nov., S. macrolepis, and S. longimanus, are described. All are recognized here as the Saurida undosquamis group, characterized by having dark dots on the upper margin of the caudal fin, pectoral fin exceeding origin to pelvic fin, anterior rays of dorsal fin neither elongate nor filamentous, predorsal length greater than distance between dorsal-fin and adipose-fin origins, 46–55 pored lateral-line scales, and vomer with 0–8 teeth. Saurida undosquamis, from northern West India and West Pacific, excluding East Asia, differs from others in having lateral-line scales ridged on the caudal peduncle, conspicuously concave posterior margin of the pectoral fin, 51–55 pored lateral-line scales, and 50–53 vertebrae. Saurida umeyoshii sp. nov., from southern Japan and the East China Sea, is defined by three rows of indistinct dark blotches on, above, and below the lateral line, distribution of scale pockets with melanophores on their posterior part extending over the entire abdominal region from the lateral line in specimens over ca. 130 mm SL, lateral-line scales not ridged on caudal peduncle, 49–52 pored lateral-line scales, and 48–50 vertebrae. Saurida macrolepis, from the Indo-West Pacific, is characterized by 46–49 pored lateral-line scales and 45–48 vertebrae. Saurida longimanus, from northern West India, northwest Australia, and southern Indonesia, differs from the others in having a long pectoral fin extending past the origin of the dorsal fin. Some geographic variations are found in S. macrolepis. Saurida grandisquamis is confirmed as a junior synonym of S. undosquamis, based on examination of the type specimens. A key to species in the S. undosquamis group is included.     

Induction of ovarian maturation and development of eggs,Larvae and Juveniles of the Puffer,Takifugu exascurus,Reared in the Laboratory

Ovarian maturation of the pufferTakifugu exascurus (Jordan et Snyder) was induced, and embryonic, larval and juvenile development was observed. The brood fish were collected in Tassha Bay, Sado Island (38°05′N, 138°15′E), during the spawning season in 1986 which seemed to extend from late June to mid-July. To each female 3 mg acetone-dried pituitary ofHypophthalmichthys molitrix was injected to induce ovarian maturation, which took place in about 77 hours at a water temperature of 19.5–21.0°C. The eggs obtained by hormone injection were artificially fertilized with the milt from a collected male. The hatched larvae were fed successively on rotifersBrachionus plicatilis, Artemia nauplii and minced fish meat, and reared for a period of about one year. The eggs were spherical, 1.24±0.04mm in diameter, demersal and adhesive. The egg-membrane Was transparent and yolk was orange in color, containing a cluster of small oil-globules. The incubation period was about 160 hours at a water temperature of 18.5–21.0°C. The newly-hatched larvae, measuring 2.9– 3.1 mm TL, had 8+15 = 23 myomeres. Absorption of the yolk was completed 3 days after hatching, by which time the larvae had attained 3.5–3.6 mm TL. Larval finfolds disappeared and rudimentary dorsal, anal and caudal fins formed at 4.1–4.4 mm TL, in 6 days after hatching. In 9-day old larva (5.4 mm TL), fin ray rudiments appeared on the dosai, anal and caudal fins and spine-like scale formed on the belly. In 16-day old specimens, 9.1–10.2 mm TL, the full complements of fin rays were completed on all the fins and the fish reached the juvenile stage. The growth of larvae and juveniles reared in 1986–1987 is expressed by the following equations, where y is total length (mm) and x is days after hatching. y₁ = 2.9420· 1.0639^x 0 ≦ x ≦ 19 (r = 0.998) y₂ = 4.0286· 1.0464^x 19≦ x ≦ 33 (r = 0.998) y₃ = 9.8854· 1.0180^x 33 ≦ x ≦ 72 (r = 0.996) y₄ = 20.1555· 1.0080^x 72 ≦ x ≦ 115 (r = 0.998) y₅ = 28.0610· 1.0049^x 115 ≦ x ≦ 202 (r = 0.995)     

Larval and juvenile Polymetme elongata (Stomiiformes: Phosichthyidae) collected from Suruga Bay and offshore waters, Japan

Two larvae [17.4 mm standard length: SL (postflexion stage)] and 26.1 mm SL (transformation stage)] and a juvenile (31.7 mm SL) of a phosichthyid, Polymetme elongata, from Suruga Bay and offshore waters, central Japan, are described. These specimens had an elongate body with relatively short preanal length (53–63% SL), long anal fin base (2.6–3.4 times dorsal fin base length), and anal fin origin below dorsal fin base, and were further characterized by a blackish flap on each eye and internal clusters of melanophores (e.g., along caudal myosepta around midlateral line and on ventral margin of caudal peduncle). The short preanal length and larval melanophore pattern were very similar to those of another phosichthyid, Yarrella blackfordi, from the Atlantic Ocean.     

Embryonic and morphological development of larvae and juveniles of the Amberjack,Seriola dumerili

Embryonic and morphological development of larvae and juveniles of the amberjack,Seriola dumerili Risso, are described using specimens raised at Yaeyama Station (Ishigaki Island, Okinawa Pref.), Japan Sea Farming Association. The specimens obtained from brood fish (3 females, 3 males) were treated with gonadotropin and spawned on 6th of April 1987. The eggs of amberjack are pelagic, spherical in shape and 1.01–1.17 mm in diameter. The yolk is roughly segmented and has a single oil globule 0.22–0.24 mm in diameter. The perivitelline space is narrow. During development, a few melanophores and no xanthophores were observed on yolk. Hatching took place 35 hrs. 15 min. after spawning out at temperatures 23.1–23.7°C. The newly hatched larvae were 2.84–3.04mm in TL with 27 (13+14) myomeres and an oil globule anteriorly situated beyond the head. 3 days after hatching 4.00 mm TL, the mouth opened. 10 days after hatching 4.26 mm TL, small denticles appeared on the margin of the upper jaw and there were 1 anterior and 2 posterior preopecular spines. At 5.96mm TL, notochord was slightly flexed. Caudal, dorsal and anal fins with rudiments of rays appeared at 8.00 mm TL. The specific numbers of all fin rays and spines were obtained in a juvenile 9.60 mm TL. In a juvenile 34.25 mm TL, 54 days after hatching, the characteristic brown band of amberjack had appeared on head. Some notable changes in relative growth were observed at 5 mm and 15 mm in TL.     

Pelagic larvae of Ventrifossa garmani (Gadiformes: Macrouridae) from Suruga Bay and offshore waters of Japan

Three pelagic larvae [5.1–5.9 mm in head length (80+ to 101+ mm in total length)] of a macrourid fish, Ventrifossa garmani, from Suruga Bay and offshore waters of central Japan are described. The specimens were characterized by a remarkably elongate caudal region (caudal region length >15.6 times head length), the longest known to date among macrourid larvae and juveniles. Other characteristics included a short snout, first dorsal and pelvic fin rays not elongated, external melanophores on most of the body and posteriorly on the anal fin membrane, and six or seven rectangular clusters of internal melanophores laterally on the anterior caudal region.     

Development of eggs,larvae and juveniles of the labrid fish,Halichoeres poecilopterus,reared in the laboratory

Embryonic, larval and juvenile development of the labrid fish,Halichoeres poecilopterus, is described using a laboratory-reared series. The eggs, measuring 0.60–0.72 mm in diameter, were pelagic and spherical with a single oil globule (0.12–0.16 mm in diameter). Hatching occurred 18 h 48 min after spawning. The newly-hatched larvae, measuring 1.46–1.70 mm TL, had 8–114 + 16–18 myomeres. A conspicuous melanophore appeared on the dorsal finfold 8 h after hatching, at ca. 2 mm TL. The yolk was completely absorbed 3 days after hatching, at 2.52–2.72 mm TL. Flexion of the notochord started at ca. 6 mm TL and was finished at ca. 8 mm TL. Aggregate numbers of all fin rays were completed at ca. 14 mm TL. Squamation was almost completed at ca. 20 mm TL.     

Larvae of <Emphasis Type="Italic">Lamprogrammus shcherbachevi</Emphasis> (Ophidiiformes: Ophidiidae) from the western North Pacific Ocean

Three exterilium larvae (18.2 mm notochord length to 113.3 mm standard length) of an ophidiid, Lamprogrammus shcherbachevi, from the western North Pacific are described. The specimens had a highly specialized morph with a remarkably elongate trailing gut and ventral coracoid process, and many elongate anterior dorsal fin rays, as occur in other exterilium larvae, but were characterized by unique melanophore patterns (a cluster of melanophores on the back of the stalked pectoral fin base, a row of clusters midlaterally on the trunk and caudal region, and further clusters on the trailing gut). Although the largest specimen (113.3 mm standard length, much larger than the previously recorded maximum size of exterilium larvae) retained typical features of the exterilium stage, the ventral coracoid process was significantly reduced in size compared with that of a smaller specimen (37.8 mm standard length). Comparison of the largest specimen with an adult suggests that the anterior dorsal fin rays would disappear during the transformation stage.     

Larval development of Asemichthys taylori (Cottidae)

Larvae of Asemichthys taylori were reared in the laboratory to identifiable juveniles. A preserved series of larvae was characterized in part by fin meristics (X–XI, 14–15 dorsal fin rays; 15 anal fin rays), as well as a combination of the alignment of the jaw tip with the ventral margin of the gut, heavy lateral melanistic pigment except on the caudal peduncle, and a series of postanal ventral melanophores. An internal, horizontal band of melanin extends through the head from the snout to the gut. Hypural plates do not align vertically even after settlement, giving the appearance that the larvae do not complete notochord flexion during their planktonic stage. Larval A. taylori resemble known larvae of Radulinus spp., a genus under which some authors have synonymized the monotypic genus Asemichthys.     

Pelagic eggs and larvae of Coelorinchus kishinouyei (Gadiformes: Macrouridae) collected from Suruga Bay, Japan

Pelagic eggs and larvae of the macrourid fish Coelorinchus kishinouyei, collected from Suruga Bay, southern Japan and subsequently identified by 16S rRNA gene nucleotide sequences, are described. The spherical eggs, 1.18–1.31 mm in diameter, contained a single oil globule, 0.28–0.33 mm in diameter, and had hexagonally patterned ornamentation on the chorion, 0.017–0.022 mm in width. Melanophores were present on the embryo, yolk and oil globule after the blastopore had closed. Within 1 day after hatching, the body axis of the yolk-sac larvae was bent slightly at the anterior trunk region. During this stage many melanophores formed on the head, trunk, tail, yolk and oil globule, along with small irregular wrinkles on the dorsal and ventral finfolds. Pelagic eggs (after the caudal end of the embryo had detached from the yolk) and yolk-sac larvae also developed xanthophores on the embryo and yolk, and head, trunk, dorsal and ventral finfolds just before tail tip, and yolk, respectively. The pelagic larvae had a short tail, stalked pectoral-fin base and no elongate first dorsal and pelvic-fin rays. Three clusters of melanophores were present on the tail (anterior two embedded to muscle and one just before tail tip subsequently lost with development) and a cluster around the anus (beyond 3.9 mm head length). Nucleotide sequence analyses of comparative adult specimens appeared to confirm a previous proposal that C. productus is a junior synonym of C. anatirostris.     

Review of the genusBembras Cuvier, 1829 (Scorpaeniformes: Bembridae) with description of three new species collected from Australia and Indonesia

The bembrid genusBembras Cuvier is reviewed. Five species,B. japonica Cuvier,B. adenensis Imamura & Knapp and three undescribed species, were assigned to the genus. Type species of the genus,Bembras japonica is redescribed on the basis of 36 specimens including the holotype, and three new species,B. macrolepis, B. longipinnis andB. megacephala, previously misidentified asB. japonicus, are also described on the basis of specimens collected from Australia and Indonesia.Bembras macrolepis differs from its congeners by having large body scales, a long pectoral fin with 17–19 rays and a dark blotch on slightly upper portion to middle of margin, 14–15 anal-fin rays, small head and orbit, and caudal fin with a broad vertical dark band near posterior margin.Bembras longipinnis is distinguished from other members of the genus by having a slightly long pectoral fin with 17–19 rays and lacking a small black blotch near tip of upper rays, caudal fin with a large dark spot most intense in lower lobe, 1–2 gill rakers on upper gill arch, 13–14 anal-fin rays, slightly elong ated head and small orbit.Bembras megacephala is characterized by the following combination of characters: caudal fin with several irregular narrow vertical dark bands, small orbit, pectoral fin with 19–20 rays and lacking a small black blotch near tip of upper rays, head elongate, 2–4 gill rakers on upper gill arch, 15 anal-fin rays and small body scales. A key separating the five species ofBembras is given.     

A new loach, Cobitis shikokuensis (Teleostei: Cobitidae), from Shikoku Island, Japan

A new species of spinous loach, Cobitis shikokuensis, is described based on 297 specimens from Shikoku Island, Japan. The new species was formerly known as the Shikoku group of Cobitis takatsuensis. It can be distinguished from other species of Cobitis and closely related genera by a combination of the following characters: dorsal fin with 6 branched soft rays; anal fin with 5 branched soft rays; one brownish streak across eye from the tip of nose, no streak on cheek; a black spot smaller than eye diameter near the dorsal corner of the caudal fin base; 3–5 small brownish speckles on ventral side of caudal peduncle; high caudal peduncle with well-developed fleshy keels on dorsal and ventral side; a lamina circularis at base of dorsal part of pectoral fin absent; first branched soft ray of pectoral fin broad in males; pectoral soft rays widely branched from the approximate midpoint; last anal fin ray with 2 elements; interorbital width 11.2–17.1% of head length.     

Pelagic eggs and larvae of Coryphaenoides marginatus (Gadiformes: Macrouridae) collected from Suruga Bay, Japan

The pelagic eggs, yolk-sac and pelagic larvae of the macrourid fish, Coryphaenoides marginatus, from Suruga Bay in southern Japan, are described. The identification of the pelagic eggs based on 16S rRNA gene nucleotide sequences agreed with that obtained from morphological analyses. The spherical eggs, 1.14–1.30 mm in diameter, contained a single oil globule 0.30–0.38 mm in diameter, and had hexagonally patterned ornamentation on the chorion, 0.025–0.033 mm in width. Many melanophores were present on the anterodorsal region of the embryo after the caudal end had detached from the yolk. Within a day after hatching, each of the yolk-sac larvae had a body axis that was bent slightly at the anterior trunk region, many dorsal and lateral melanophores on the trunk plus several on the gut, and small irregular wrinkles on the dorsal and anal fin membranes. The pelagic larvae had a short caudal region in comparison to other known congeners (length 2.0–3.2+ times head length vs. 4–7, respectively), a short stalked pectoral fin base, and no elongate first dorsal and pelvic fin rays. They were further characterized by the presence of numerous very dense melanophores from just behind the eye to the anterior part of the caudal region at 5.1 mm head length (25.8+ mm total length). The significant difference in vertical distribution between the pelagic eggs and larvae (dominant depths ca. 200–350 m vs. ca. 10–100 m, respectively), with no subsequent collection of pelagic larvae with greater than 6 mm head length, indicate two stages (rising and falling) of ontogenic vertical migration.     

Developmental morphology of the cyprinid fish Inlecypris auropurpureus

Embryonic, larval, and juvenile development of a Myanmarese cyprinid fish, Inlecypris auropurpureus, is described from laboratory-reared specimens. The eggs, measuring 0.9–1.0 mm in diameter, were demersal, almost spherical in shape, transparent and unpigmented, with a pale yellow yolk without oil globules. Hatching occurred 49–56 h after fertilization at 26.2°–27.3°C. The newly hatched larvae, measuring 2.9–3.1 mm in body length (BL) with 17 + 19–20 = 36–37 myomeres, had melanophores on the head and body. A cement organ on the forehead for adhering to objects during the yolk sac and early preflexion larval stages was distinctive. The yolk was completely absorbed at 3.6–4.0 mm BL. Notochord flexion was initiated at 5.1–5.6 mm BL and finished at 7.1 mm BL. Aggregate numbers of all fin rays were completed at 14 mm BL. Squamation was initiated midlaterally on the anterior trunk at 14 mm BL and completed at 27 mm BL. Although the eggs of I. auropurpureus resembled those of the closely related species Chela dadiburjori, Danio rerio, and Devario malabaricus, they differed from those of Danio rerio and Devario malabaricus in having a narrower perivitelline space. The larvae and juveniles of I. auropurpureus were also similar to those of C. dadiburjori, Danio rerio, and Devario malabaricus in general morphology, but they differed from the latter three species in having a series of dark blotches laterally on the body in the juvenile stage. Moreover, I. auropurpureus differed from C. dadiburjori in having more myomeres and a near-single row of melanophores on the body along the dorsal midline from the yolk-sac to early postflexion larval stages, from Danio rerio in having a cement organ on the forehead during the yolk-sac and early preflexion larvae, and a single melanophore on the lower eye margin in the early yolk-sac larvae, and from Devario malabaricus in having a single melanophore on the lower eye margin in the early yolk-sac larvae. The presence of a cement organ on the forehead indicates a close relationship among the genera Inlecypris, Chela, and Devario.     

Growth and morphological development of larval and juvenileepinephelus bruneus (perciformes: Serranidae)

The growth and morphological development of larval and juvenileEpinephelus bruneus were examined in a hatchery-reared series. Average body length (BL) of newly-hatched larvae was 1.99 mm, the larvae growing to an average of 3.96 mm by day 10, 6.97 mm by day 20, 12.8 mm by day 30, 22.1 mm by day 40 and 24.7 mm by day 45 after hatching. Newly-hatched larvae had many mucous cells in the entire body epidermis. By about 4 mm BL, the larvae had developed pigment patterns peculiar to epinepheline fishes, including melanophores on the dorsal part of the gut, on the tips of the second dorsal and pelvic fin spines, and in a cluster on the ventral surface of the tail. Spinelets on the second dorsal and pelvic fin spines, the preopercular angle spine and the supraocular spine, had started to develop by about 6 mm BL. The notochord tip was in the process of flexion in larvae of 6–8 mm BL, by which time major spines, pigments and jaw teeth had started to appear. Fin ray counts had attained the adult complement at 10 mm BL. After larvae reached 17 mm BL, elements of juvenile coloration in the form of more or less densely-pigmented patches started to appear on the body. Squamation started at 20 mm BL. Major head spines had disappeared or became relatively smaller and lost their serrations by 20–25 mm BL.