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. Bau- delot's ligament from basioccipital passing through this notch, attaching to medial surface of supracleithrum. Vertical limb of cleithrum developed, not extending to ventral border of cleithrum. Lower part of horizontal limb of cleithrum broadly expanded. Scapula foramen not enclosed in scapula. Posterodorsal margin of scap- ula with marginal keel supporting upper four pectoral-fin rays. Four small plate- like radials; upper two supported by scapula, lower two by coracoid. Four rounded slits present between scapula and uppermost radial, and between adjacent radials. Lower postcleithrum free from upper postcleithrum. Pelvic bones suture on mid- line, with a fossa on posterodorsal margin where muscle of infracarinalis medius inserts. A strong ligament extending between ventrolateral corner of pelvis and posteroventral tip of coracoid.

Postcranial Axial Skeleton (Fig. 5).- There are 15 abdominal vertebrae and 22 caudal vertebrae including urostylar vertebra (on the radiograph's observation; 14 a.v. + 23 c.V. (six specimens), 14 + 22 (two), 13 + 24 (one)). Anteriormost centrum modified to form three facets for articulation with basioccipital and both exoccipitals. Neural arch of anteriormost centrum closed with a short strong neural spine. Parapophyses present on abdominal vertebrae except for anteriormost two, well developed extending horizontally on posterior half of abdominal vertebrae. Parapophysis on last abdominal vertebra closed to form a haemal arch without haemal spine. 16 pairs of epipleurals associated with first to 16th vertebrae, lying in lateral septum between epaxial and hypaxial muscles; distal tip of each epi- pleural extending above lateral line nerve; anteriormost two epipleurals attaching to lateral sides of neural arches; successive epipleurals attaching to parapophyses, but last a few rudimentary, free from vertebrae. Pleural rib absent. Urostylar vertebra separated from parhypural and hypurals. Parhypural with a hypurapo- physis. First and second hypurals fused to each other, forming lower caudal lobe with parhypural. Preural centrum 2 with a low neural crest and an autogenous haemal spine. Other haemal spines fused with centra. Upper lobe of caudal skel- eton supporting two upper unbranched rays and five branched rays; lower lobe 498 BULLETIN OF MARINE SCIENCE. VOL. 58. NO.2. 1996

ceratohyal A hypobranchial

pharyngobranchial2nd - 4th epibranchial2nd -4th

pharyngobranchial epibranchial

o ceratohyal

branchiostegal rays

Figure 3. Hyoid and branchial arches of Normanichthys crockeri. A, dorsal view (dorsal elements were removed); B, ventral view of right dorsal elements of branchial arch; C, dorsal view of right dorsal elements of branchial arch (first epibranchial was removed); D, hyoid arch (lateral view). Scale bars = 2 mrn. YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 499

Figure 4. Pectoral (A; lateral view, supracleithrum was removed), and pelvic girdle (B; dorsal (above), ventral (middle) and lateral views (below)) of Normanichthys crockeri. Scale bars = 2 rnm. supporting five branched rays and a lower unbranched ray. 11-13 dorsal and 12- 13 ventral procurrent caudal rays. First dorsal fin comprising of 9-11 spines, separated from second dorsal fin of 10-11 soft rays. First proximal pterygiophore of dorsal series inserting between second and third vertebrae, articulating to sec- ond proximal pterygiophore posterodorsally; second proximal pterygiophore sup- porting first dorsal-fin spine without distal pterygiophore; successive proximal pterygiophores of first dorsal fin supporting spiny rays with a single plate-like distal pterygiophore; fifth and sixth (or 4th and 5th) proximal pterygiophores inserting to same interneural space. Four slender proximal pterygiophores present

A proximal distal pterygiophore plerygiophore epipleuralrib 1st abdominal vertebra

Figure 5. Postcranial axial skeletons of Normanicthys crockeri. A, lateral view of anterior 21 ver- tebrae and associated bones; B, lateral view of caudal skeleton; C, ventral view of posterior four abdominal vertebrae. Scale bars = 2 mm. 500 BULLETIN OF MARINE SCIENCE, VOL. 58. NO, 2. 1996

dilatator operculi

ligamentum primordium adductor arcus palatini adductor mandibu lae adductor mandibu lae section A1 section A 2

Figure 6. Cheek muscle of Normanichthys crockeri. Lateral view. Scale bar = 2 mm. between first and second dorsal fins. Each proximal pterygiophore of second dor- sal fin supporting a soft ray with a paired distal pterygiophore; last proximal pterygiophore supporting two small soft rays. Anal-fin comprising of 14-15 soft rays. Anteriormost two proximal pterygiophores of anal series inserting before first haemal spine; each proximal pterygiophore except for the last supporting a soft ray of anal-fin with a paired distal pterygiophore; the last supporting two small soft rays. Cheek Muscle (Fig. 6).-Adductor mandibular section Al subdivided into anterior and posterior muscular masses; fibers of both masses converged on middle part with a slender tendinous connection; some fibers of anterior mass connecting with ligamentum primordium. Adductor mandibular section A2 originating from lower angle of suspensorium, tendinously connecting with adductor mandibular section Aw; connecting with section Al by a slender tendon anterodorsally. A part of adductor mandibular section A3 connecting with posterior mass of section Al by a thin tendon. Levator arcus palatini originating from posterolateral part of sus- pensory region, broadly inserting to sphenotic. Adductor arcus palatini filling in floor of orbital cavity. Levator operculi divided into superficial and profound layers; superficial layer originating from dorsomedial margin of opercle, inserting to ventrolateral margin of pterotic and posttemporal; profound layer originating from medial surface of posterodorsal process of hyomandibular and anterodorsal part of opercle, inserting to pterotic. Dilatator operculi originating from ventro- lateral margin of pterotic, inserting to anterodorsal corner of opercle. Muscles of Hyoid Arch (Fig. 7A).-Protractor hyoidei originating from antero- medial surfaces of both dentaries of immediately behind symphysis, running pos- teriorly to form a common cylindrical muscular mass on ventral mid-line, then branched off either sides, inserting to ventrolateral side of ceratohyal between bases of first and second branchiostegal rays. Some elements of hyohyoidei ab- ductores supporting between hyoid arch and branchiostegal rays. Anteriormost YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 501

B basihyal c

pharyngoclavlcularis externus

Figure 7. Muscles of hyoid and branchial regions in Normanichthys crockeri. A, ventral view of chin region; B, ventral view of branchial arch; C, dorsal view of branchial arch. Scale bars = 2 mm. element of hyohyoidei abductores, a developed cylindrical muscular mass, ex- tending between anteroventral hypohyal and first branchiostegal ray. Slender mus- cular masses of hyohyoidei abductores present between first to fourth branchio- stegal rays and ventromedial surface of ceratohyal. Two thin muscular sheets of hyohyoidei abductores extending between first branchiostegal rays of both sides with a medial raphe attaching to isthmus. Hyohyoidei adductores present as a sheet between branchiostegals and medial surface of opercle. Hyohyoides infer- ioris absent. Rectus communis originating from upper margin of urohyal, taper- ingly inserting to lateral margin of fifth ceratobranchial. Sternohyoideus originat- ing from posteroventral surface of urohyal, extending to ventral tip of cIeithrum where it assimilating obliquus inferior of hypaxialis. Muscles of Branchial Arch (Figs. 7-8).-Obliquus ventralis present between ven- tral surfaces of hypobranchial and ceratobranchial on first and third branchial arches. Rectus ventralis interconnecting between ventral surface of third hypo- branchial and fourth ceratobranchial; medially some fibers connecting with its antimere. Transversus ventralis divided into anterior and posterior sections; an- terior section extending between anteroventral parts of both fourth ceratobran- chials; posterior section between entirely ventromedial margins of fifth cerato- branchials; posteriormost fibers of posterior section assimilating to sphincter oes- ophagi. Adductores present between ceratobranchial and epibranchial of each gill arch and between fifth ceratobranchial and fourth epibranchial. Pharyngoclavi- cularis externus, a broad muscular mass, extending between ventrolateral margin of fifth ceratobranchial and ventral part of cleithrum. Pharyngoclavicularis inter- nus, a slender muscle, interconnecting between ventromedial surface of fifth ce- ratobranchial and anteromedial part of middle of cleithrum. Levator externus and levator internus originating from lateral side of cranium immediately below facets of craniohyomandibular articulation; the former passing superficial to the latter. Four levator externus elements; the first a cylindrical mass, inserting on the dorsal process of first epibranchial; the second to fourth forming broad converged mass, 502 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

epaxialis levatorposterior

pharyngoclavicularis infracarinalisanterior externus Figure 8. Muscles between pectoral girdle, skull and branchial arch in Normanichthys crockeri. Lateral view (suspensorium and hyoid arch were removed). Scale bars = 2 mm.

taperingly inserting to the second to fourth epibranchials. Two levator internus elements; the anterior inserting to anterior margin of second pharyngobranchial; the posterior inserting to dorsolateral margin of third pharyngobranchial. Levator posterior, a cylindrical muscular mass, extending between fourth epibranchial and posterolateral side of otic region of cranium. Obliquus dorsalis originating from dorsal surface of third pharyngobranchial, broadly inserting to fourth epibranchial; some fibers inserting to dorsal process of third epibranchial. Anterior element of transversus dorsalis developed, extending between second epibranchials of both sides; posterior element, a thin muscular sheet, extending below obliquus dorsalis, Retractor dorsalis originating from ventrolateral side of second and third centra, inserting to dorsomedial margin of third pharyngobranchial. A slender muscular branch of sphincter oesophagi passing on oral surface along pharyngobranchials; it originating from posterolateral corner of second pharyngobranchial, running along lateral margins of third and fourth pharyngobranchials, assimilating to sphincter oesophagi posteriorly. Muscles of Occipital Region (Fig. 8).-Epaxial muscle broadly inserting to pos- teromedial margin of the cranium dorsomedially, but not extending to dorsal sur- face of parietal nor supraoccipital. Lateral branch of epaxialis inserting to post- temporal fossa. Protractor pectoralis originating from lateral surface of exoccipital as a cylindrical muscular mass, gradually changed to a thin muscular sheet, in- serting to anterior margin of c1eithrum. Levator pectoralis originating from lateral surface of exoccipital immediately medial to origin of protractor pectoralis, in- serting to anterodorsal process of cleithrum. Obliquus superior of hypaxial muscle broadly inserting to lateral surface of basioccipital. Another specialized branch of hypaxial is not observed on this region. Muscles of Pectoral, Pelvic and Caudal Fins (Fig. 9).-Pectoral fin supported by abductor superficialis, abductor profundus and arrector ventralis on lateral side, by adductor superficialis, adductor profundus and arrector dorsalis on medial side. YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 503

8

Infracarinalls medius A

ligament

Infracarinalls anterior abductor superflclalle

obliquus inferlorls 01hypaxlalle

Figure 9. Muscles of pectoral, pelvic and caudal fins in Normanichrhys crockeri. A, medial view of pectoral girdle; B, dorsal view of pelvic girdle (above), and ventral view of thorax region (below); C, lateral view of caudal muscles (epaxialis, hypaxialis, and upper part of hypochordal longitudinalis were removed). Scale bars = 2 mm.

Coracoradialis, a small muscular mass, present between lowermost radial and posteroventral projection of coracoid on medial side. Pelvic fin supported by ar- rector ventralis pelvicus, abductor superficialis pelvicus and abductor profundus pelvicus on ventral side, by adductor superficialis pelvicus and adductor profundus pelvicus on dorsal side, and by arrector dorsalis on lateral side. Extensor proprius absent. Interradialis serve upper seven and lower seven principal caudal rays on proximal part of caudal fin. Hypochordal longitudinalis inserting to upper four branched rays. Flexor ventralis externus tendinously inserting to upper two rays of lower caudal lob. Adductor dorsalis of caudal fin absent. Alimentary Canal and Swimbladder (Fig. 1O).-Stomach thick-walled, V-shaped. Six finger-like pyloric caeca present. Intestine short, simply coiling on right side of stomach. A physoclistous swimbladder present behind stomach, located pos- terior two-third of abdominal cavity. Swimbladder thin-walled, rounded anteriorly, tapered posteriorly; it loosely attached to parapophyses of sixth to 14th vertebrae by connective tissue. Eight retia mirabilia radiated on anteroventral wall of swim- bladder.

DISCUSSION Yabe (1985) established the superfamily Cottoidea as a monophyletic group based on a combination of 22 synapomorphies. He also suggested that the cy- clopteroid group including the Cyclopteridae and Liparidae is the sister group of the Cottoidea because it shares 20 of the cottoid synapomorphies. The monophy- letic group comprising the cottoid and cyclopteroid fishes conforms closely to the suborder Cottoidei presented by Nelson (1984). To reconsider the relationships 504 BULLETIN Of MARINE SCIENCE. VOL. 58, NO.2, 1996

Figure 10, Photomicrograph of abdominal cavity in Normanichthys crockeri, HUMZ 113858, 57.7 mm SL, lateral view (a part of liver was removed). between Normanichthys and cottoid fishes, we use the concept of the suborder Cottoidei sensu Nelson (1984) except we exclude the Normanichthyidae, and characterize this suborder by the combination of 20 synapomorphies presented by Yabe (1985). As the following comparisons, Normanichthys crocked has different character states in the following 10 synapomorphies of the Cottoidei. Among them, the first five synapomorphies are uniquely shared within the Cottoidei among the Scor- paeniformes, and some of them were regarded as the peculiar of this species by Mandrytza (1991). 1) Fourth pharyngobranchial absent: the numbers of pharyn- gobranchials and tooth plates except for the first (suspensory pharyngeal) are one or two in the cottoids. Although Norman (1938) stated that Normanichthys had two pairs of upper pharyngeals in which the third and fourth were united with each other, Mandrytza (1991) described that this species has three pharyngobran- chials with tooth plates. Our specimens have the second to the fourth pharyngo- branchials with three tooth plates. 2) Basihyal rudiment or absent: most cottoids lack the basihyal. Only some species have a rudimentary or small cartilaginous basihyal (Yabe, 1985; Kido, 1988; Kanayama, 1991). Normanichthys has a long rod-like basihyal. 3) Posttemporal fossa reduced: in the cottoid fishes, the post- temporal fossa is very shallow and does not receive the epaxial muscle. Norman- ichthys has a developed posttemporal fossa into which the anterolateral branch of the epaxial muscle inserts. This difference was also suggested by Mandrytza (1991).4) Extrinsic muscle of swimbladder extending between cranium and c1ei- thrum: all cottoids share this unique muscular structure associating with loss of the swimbladder (Hallacher, 1974; Yabe, 1985). Normanichthys does not have the YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMAN1CHTHYS 505 extrinsic muscle, although it has the swimbladder. 5) Pleural ribs absent at least on anterior five vertebrae: many cottoids have pleural ribs on posterior abdominal vertebrae. Among the Cottoidei, most of agonid and liparid fishes lack pleural ribs, but a few primitive members of each family have the pleural ribs (Kido, 1988; Kanayama, 1991). These reductions could be occurred in each lineage in- dependently. The absence of pleural ribs has been regarded as a peculiar feature of Normanichthys since Clark (1937) described, but Mandrytza (1991) described that this species has rudimental pleural ribs on 7th to 11th vertebrae, each of them is composed of two or three short fragments freeing far from corresponding cen- trum. We could not observe such rudimental bones on our materials. The absence or reduction of pleural ribs in Normanichthys is peculiar, and should probably be considered homoplasy. 6) Lachrymal-palatine articulation present: in the cottoid fishes, the first infraorbital (lachrymal) has a dorsal facet for the lachrymal-lateral ethmoid articulation, and a medial condyle for the lachrymal-palatine articulation. This condition is also known in hexagrammids and zaniolepidids among Scor- paeniformes (Yabe, 1985). In Normanichthys, the first infraorbital has an articular socket for the lachrymal-lateral ethmoid articulation anteromedially, but does not have a medial condyle for the lachrymal-palatine articulation. 7) Scapula sepa- rated from coracoid: in most cottoids, the scapula is separated from the coracoid by a cartilage. Among the Cottoidei, an exception in the specialized agonid As- pidophoroides was illustrated by Kanayama (1991); it is probably a secondary condition. A separate scapula is also known in hexagrammids and triglids (Starks, 1930; Quast, 1965). In Normanichthys, the scapula is firmly attached to the cor- acoid. 8) Hypurapophysis absent: this condition is also known in Erisphex (Al- poactinidae) and hexagrammids among the Scorpaeniformes (Yabe, 1985; Fujita, 1990). Normanichthys has a normally developed hypurapophysis on the parhy- pural. 9) Caudal skeleton composed of one or two plate-like bones: in the cottoids, the hypurals, parhypural and urostyle are fused into one or two bony plates. In Normanichthys, the caudal skeleton comprises four hypurals and a parhypural. It could be more primitive than those of cottoids, but the condition in Normanichthys is unique in having only a fusion of hypural I and 2. Such a configuration in the caudal skeleton has been reported only in some Acanthopterygii: Odonthestes bonariensis (Atherinidae), Parapercis cylindrica (Pinguipedidae), Ruvettus pre- tiosus (Gempylidae), Notothenia tessellata (Nototheniidae) and Parapterois het- erurus (Scorpaenidae) (Fujita, 1990). 10) Levator externus III of branchial muscle absent: same condition is known in synanceiids among the Scorpaeniformes (Yabe, 1985). Normanichthys has the third levator externus inserting on the third epibranchial as in many of non-cottoid scorpaeniforms. On the other hand, Normanichthys shares the following nine character states with the cottoids. Some were presented as evidence of Norman's (1938) hypoth- esis, but all of these character states are known to occur homoplasiously (or synapomorphic) in non-cottoid fishes (Table 1): 11) suborbital stay present, 12) basisphenoid absent, 13) intercalar bone small, 14) metapterygoid lamina absent, 15) scapular foramen open, 16) anal-fin spines absent, 17) hyohyoidei abductores section 2 of hyoid muscle fused to its antimere, 18) extensor proprius of pelvic muscle absent, 19) adductor dorsalis of caudal muscle absent. The remaining synapomorphy of the cottoids, 20) epipleural ribs attach to vertebrae not to pleural ribs, is shared by Normanichthys. But it could not be estimated as a homologous condition, because the pleural rib is absent in Normanichthys. A close relationship between Normanichthys and cottoids has little support because the former lacks a set of 20 synapomorphies limiting the suborder Cot- toidei. Normanichthys shares nine derived conditions with the cottoids, but all of 506 BULLETIN OF MARINE SCIENCE, VOL. 58, NO, 2. 1996

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them also occur homoplasiously among the percomorpha. In the five characters (nos. 1 to 5) which are uniquely shared by the Cottoidei at least within the Scor- paeniformes, Normanichthys has more primitive conditions or has lost the com- ponents of the characters. Quast (1965) suggested the parasphenoid-pterosphenoid junction is an impor- tant evidence of the hexagrammids-cottids lineage. Although Normanichthys has the junction, its relation to the hexagrammids-cottids lineage could not be tested, because some families of the Cottoidei lack the junction; e.g., Cyclopteridae, Ereuniidae, Psychrolutidae, Hemitripteridae (Deno, 1970; Yabe, 1981, 1983, 1985; Nelson, 1989). More recently, Shinohara (1994) presented nine synapo- morphies of his zaniolepidoid-hexagrammoid-cottoid clade, which corresponds to the hexagrammids-cottids lineage of Quast (1965). Among the synapomorphies, Normanichthys lacks three; presence of lachryopalatine articulation, absence of predorsal bone, and dorsal pterygiophores arranged singly in each interneural space. We also considered the swimbladder of Normanichthys. Although Clark (1937), in the original description of Normanichthys, reported the swimbladder to be absent, Balbontin and Perez (1980) described its presence in the larvae of Normanichthys (4.6-16.3 mm TL). Our adult specimens have a thin-walled blad- der placed on the posterior half of the abdominal cavity. We regard it as a phy- soclistous swimbladder, because it has the rete mirabile on the anteroventral wall. Although the lack of the swimbladder was not regarded as a synapomorphy of cottoids by Yabe (1985), it can be regarded as an additional synapomorphy of cottoids, and also hexagrammoids and zaniolepidoids, because they lack the swim- bladder. Moreover, cottoids share a unique modification of muscles associating with the loss of the swimbladder. In this sense, Normanichthys does not share this cottoid synapomorphy. On the other hand, Mandrytza (1991) described the seismosensory system of Normanichthys, which is characterized the presence of three neuromasts in the lachrymal and four in the dentary as same as many of scorpaenoids not as cottoids. On the basis of this condition and other osteological features, Mandrytza (1991) doubted the former systematic position of Norman- ichthys, and modestly suggested that if following to traditional determination of Cottoid fishes (Regan, 1913; Berg, 1940; Taranetz, 1941), Normanichthys could be a member of the suborder Cottoidei, but it may be necessary to rank up to the superfamily-level. But as discussing above, Normanichthys cannot be placed in the suborder Cottoidei. We cannot present any reasonable sister group for Normanichthys, but we con- sider that Normanichthys is a member of the order Scorpaeniformes, because it has the suborbital stay (Fig. 11). The suborbital stay could be probably regarded as a unique synapomorphy for the Scorpaeniformes, however its polyphyletic origin has been often suggested but never verified (Quast, 1965; Greenwood et aI., 1966; Gosline, 1971; Lauder and Liem, 1983; Washington et a\., 1984; John- son, 1993). In addition to the suborbital stay, Johnson (1993) suggested that dis- tinct parietal spines in the larvae associated with a bone-enclosed passage of the supratemporal sensory canal through the parietal in adult may be the synapo- morphy of the Scorpaeniformes. Shinohara (1994) also regarded the parietal sup- porting a sensory canal as a synapomorphy of the Scorpaeniformes. Balbontin and Perez (1980) reported that the larvae of Normanichthys have no spines on the head. The adults of Normanichthys have a tabular bone for passage of sensory canal on the parietal, which is similar to the condition of some psychrolutid- cottoids reported by Nelson (1982) and Yabe (1985). Thus we do not deny the present placement of order-level category for Normanichthys. We also consider the monotypic family Normanichthyidae to be valid, because its unique set of 508 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

Figure 11, Photomicrograph of right side of head in Normanichthys crockeri, HUMZ 113858. 57.7 mm SL, after staining with alizarin res-So

morphological features (e.g., poorly ossified olfactory and posttemporal regions, elongate nasal bones, first infraorbital bone with medially situated socket for the lachrymal-lateral ethmoid articulation, no ascending process of angular, slender metapterygoid, five branchiostegals, horizontally developed parapophysis) does not fit to any other available families of the Scorpaeniformes; especially the con- dition of the adductor mandibular AI, its uniqueness among the Scorpaeniformes was also suggested by Mandrytza (1990), and a branch of the sphincter oesophagi should be regarded as autapomorphies of this taxon. In the Scorpaeniformes, Nelson (1984) recognized five suborders: Scorpaenoidei, Platycephaloidei, Ano- plopomatoidei, Hexagrammoidei, and Cottoidei. At least the Normanichthyidae is not a member of the suborder Cottoidei as discussed above, but its suborder rank is still uncertain. Therefore it is better to place Normanichthys crockeri in the Scorpaeniformes, incertae sedis: Family Normanichthyidae, until its reasonable sister group will be clarified by further morphological comparison with perco- morphs.

ACKNOWLEDGMENTS

We wish to express out thanks to K. Amaoka and K. Nakaya (HUMZ) for their valuable advice and criticisms of a draft manuscript, and to J. S. Nelson (UAMZ), G. D. Johnson (USNM) and T. W. Pietsch (UW) for their review of a draft of the manuscript. We are also grateful to T. Iwamoto (CAS) for the loan of study materials. Our special thanks go to A. Zama of Miyagi Fisheries High School and F. Balbontfn of Univ. de Valparaiso for supplying us with materials and valuable information about fishes of Chile. YABE AND UYENO: ANATOMICAL DESCRIPTION OF NORMANICHTHYS 509

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DATE ACCEPTED: August 24, 1994.

ADDRESSES: (M.Y.) Laboratory of Marine Zoology, Faculty of Fisheries, Hokkaido University, 3-1- ] Minato-cho, Hakodate, Hokkaido 04], Japan, email: [email protected]; (T.U,) Department of Paleontology, National Science Museum (Nat. Hist.), 3-23-/ Hyakunin-cho, Shinjuku-ku, Tokyo 169, Japan.