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<I>Normanichthys Crockeri</I>

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<I>Normanichthys Crockeri</I> BULLETIN OF MARINE SCIENCE. 58(2): 494-510.1996 ANATOMICAL DESCRIPTION OF NORMANICHTHYS CROCKERI (SCORPAENIFORMES, INCERTAE SEDIS: FAMILY NORMANICHTHYIDAE) Mamoru Yabe and Teruya Uyeno ABSTRACT To reconsider the doubtful systematic position of Normanichthys crockeri in the suborder Cottoidci, its osteology, myology and other morphology are described. Normanichthys does not share 10 of the 20 synapomorphies that limit the Cottoidei (4th pharyngobranchial absent; basihyal absent or rudiment; posttemporal fossa reduced; extrinsic muscle of swimbladdcr between cranium and cleithrum; pleural ribs absent on anterior vertebrae; lachrymal-palatine articulation present; scapula separated from coracoid, hypurapophysis absent; one or two hypurals; levator externus III absent). Other 10 synapomorphies of Cottoidci arc shared by Normanichthys. but each of them is known to occur homoplasiously (or synapomorphic) in non-colloid fishes (no basisphenoid absent; small interealar; no metapterygoid lamina; scap- ular foramen open; no anal-fin spine; hyohyoidei abductores II fused to its antimere; no extensor proprius; no adductor dorsalis; epipleural attached to vertebra). Moreover, Norman- ichthys has the swimbladder which is not present in all colloid fishes. The placement of the monotypic family Norrnanichthyidae in the suborder Cottoidei is rejected. Normanicllthys is regarded as a member of the order Scorpaeniformes because it has the suborbital stay which is the only synapomorphy of this order. At least Normanichthys is not a member of the monophyletic taxon Cottoidei, but its suborder rank is still uncertain because its reasonable sister group could not been clarified. Normanichthys crockeri is a small pelagic fish endemic to the coastal region of Chile, which has been classified as a perciform or scorpaeniform fish (Norman, 1938; Tortonese, 1941; Pequeno, 1978; Balbontfn and Perez, 1980). In the original description, Clark (1937) established a new family Normanichthyidae but did not place it in any order-level taxon, although he quoted Carl L. Hubbs who thought the fish was a modified percoid. Norman (1938) stated that several osteological features of Normanichthys fit the osteological diagnosis of the division Cottifor- mes defined by Regan (1913); e.g., small opisthotic, no basisphenoid, reduced mesopterygoid, two pairs of upper pharyngeals, plate-like pectoral radials, no ribs. He regarded Normanichthys as a primitive member of the family Cottidae sensu Regan (1913). Based on Norman's conclusion, Berg (1940) added the genus to his superfamily Cottoidea, but placed it in the family Normanichthyidae because of its complete squamation. This placement has remained in subsequent classifi- cations of cottoid fishes without additional evidence (Quast, 1965; Greenwood et aI., 1966; Nelson, 1976, 1984; Yabe, 1985). Doubt has often been expressed about the relationship of Normanichthys to cottoids (Tortonese, 1941; Balbontfn and Perez, 1980; Nelson, 1984; Washington et aI., 1984). The relationships remain poorly known because of the poor morphological description of Normanichthys, and because of ambiguity of the limit of the higher taxa of cottoid fishes. Recently, Yabe (1985) established the monophyly of the superfamily Cottoidea. And more recently, Mandrytza (1991) documented the condition of the cephalic sensory system and some osteological features of Normanichthys. The objectives of this study are to describe the anatomical features of Normanichthys crockeri in detail and to reconsider the relationships between this species and the Cottoidei through the comparison with cottoid synapomorphies. 494 YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 495 supratemporal prevomer o. ," f exoccipital ethmoid cartilage otolith chamber Figure]. Neurocranium of Nonnanichthys crockeri. Above; dorsal view. below; lateral view. Scale bar = 5 mm. MATERIALS AND METHODS The specimens examined for bone and muscle were dissected after staining with alizarin red-S, and observed with a binocular microscope. Some anatomical information was determined from radio- graphs. The morphological terminology chiefly follows Winterbottom (1974) and Yabe (1985); that for ligaments of the jaws follows Stiassny (1986). Institutional abbreviations are as listed by Leviton et al. (1985). Anatomical description of Normanichthys crocker; is based on the following specimens. Dissected observation: HUMZ 113858, one specimen, 57.7 mm SL; NSMT-P 45649-45650, 2 spec- imens, 4 I. 7-66.7 mm SL; CAS 5520-552 I, 2 paratypes, 51.0-54.0 mm SL. Radiographic observation: NSMT-P 45648,2 specimens, 32.1-35.0 mm SL; CAS 5522-5524, 3 paratypes, 41.0-47.0 mm SL; HUMZ 113859-113863, 5 specimens, 47.5-58.1 mm SL. ANATOMICAL DESCRIPTION Neurocranium (Fig. l).-Olfactory region poorly ossified; ethmoid cartilage ex- posed dorsolaterally. Prevo mer attached to mesethmoid dorsally, to parasphenoid ventrally. Dorsal mid-line of prevomer unossified. Anterior half of mesethmoid elongate forming median keel; posterior half deeply notched on mid-line to form laminar regions connecting with frontal. Nasal an elongate tabular bone. Lateral ethmoid with a small condyle for lachryocranial articulation on ventrolateral cor- ner, and a blunt process for palatocranial articulation anteroventrally. Tabular bone 496 BULLETIN OF MARINE SCIENCE. VOL. 58. NO.2. 1996 Figure 2. Oromandibular region of Normanichthys crockeri. A, medial view of the first infraorbital (above), and lateral view of five infraorbital bones (below); B, suspensorium (lateral view); C, upper and lower jaws (lateral views); D, maxillary (dorsal view). Scale bars = 2 mm. for occipital sensory canal present on parietal. Posttemporal fossa moderately developed; its floor less ossified. Supraoccipital crest not expanded beyond hor- izontal level of cranial roof. Epiotic with a shallow groove for temporal sensory canal anterodorsally. Ascending process of parasphenoid connected to pterosphen- oid anterodistally. Posterior region of parasphenoid notched to form posterior opening of myodome. Basisphenoid absent. Trigeminofacialis chamber on prootic covered with a wide vertical bridge. Moderate gap (foramen) present between prootic and pterosphenoid. Intercalar not extending to prootic. A T-shaped tabular supratemporal with three openings. Oromandibular Region (Fig. 2).-Five infraorbital bones. First infraorbital (lach- rymal) with a small socket anteromedially for receiving articular condyle of lateral ethmoid. Third infraorbital with a blunt projection posteriorly, the suborbital stay, its posterior margin not extending to preopercle. Fourth and fifth infraorbitals small, gutter-shaped. Palatine connected to ascending process of premaxilla by palato-premaxillary ligament, to prevomer by anterior and posterior palato-vomer ligaments. No teeth on palatines. Entopterygoid extending to dorsal part of quad- rate, attached to metapterygoid. A slender metapterygoid without lateral lamina. Preopercle with a series of incomplete tubular structures for operculo-mandibular sensory canal. Cranial condyle of maxilla well developed, articulating with pre- vomer through a meniscus medially. Lateral process of maxillary head weakly projected, connected to mesethmoid by ethmo-maxillary ligament. Medial process of maxillary head, a slender projection, articulating with articular process of pre- maxilla through a meniscus on lateroproximal position (premaxillary condyle); a small process just below cranial condyle receiving a tendon of adductor mandib- ular section AI. Maxillary prominence, a blunt process immediately behind max- illo-palatine articulation, receiving ligamentum primordium. A strong ligament extending between posteromedial surface of maxilla and ascending process of dentary. Supramaxillary absent. Dentary with a narrow band of fine conical teeth on dorsal margin except for posterior region where a blunt triangular process YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 497 present. Dorsal margin of angular straight without ascending process. Angular with a lateral keel attached by ligamentum primordium, an articular fossa receiv- ing condyle of quadrate on posterior tip, a broad groove for mandibular sensory canal on posterolateral surface. Hyoid and Branchial Arches (Fig. 3).-An elongate basihyal with a spatulate cartilage on anterior tip. Anteroventral hypohyal connecting with urohyal by a strong ligament on medial side. Five branchiostegal rays. Epihyal ligamentously connecting with medial surface of interopercle. Urohyal with two processes; a short anterior process ligamentously connecting with anteroventral hypohyals of both sides; a long dorsal process articulating with ventral side of first basibran- chial. Third basibranchial elongated. First and second hypobranchials with small processes on anterolateral margins where each process connecting with successive arch by a thin ligament. Third hypobranchial extending below second hypobran- chial anteriorly, articulating with third and fourth ceratobranchials posteriorly. Fifth ceratobranchial with an elongate tooth-plate. No tooth plate on third epi- branchial. First pharyngobranchial bone (suspensory pharyngeal) and interarcual cartilage absent. Second to fourth pharyngobranchials present with tooth-plates; tooth-plate on third largest, that on second smallest. Pectoral and Pelvic Girdles (Fig. 4).-Supracleithrum with an incomplete tube for the sensory canal on its dorsal margin. Dorsal margin of cleithrum deeply notched with a spiny-like anterior process and a laminar posterior process.
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