CHIROTEUTHID and RELATED PARALARVAE from HAW Allan WATERS Richard Edward Young

BULLETIN OF MARINE SCIENCE, 49(1-2): 162-185, 1991

CHIROTEUTHID AND RELATED PARALARVAE FROM HAW AllAN WATERS Richard Edward Young

ABSTRACT Eleven species of paralarvae belonging to the Chiroteuthidae and related families are described from Hawaiian waters, The doratopsis stage is shown to be a diagnostic feature of the family Chiroteuthidae. This family, as now defined, includes the genera Chiroteuthis, Asperoteuthis, Grimalditeuthis and Planktoteuthis. Doratopsis sagitta, Valbyteuthis, Tankaia, Echinoteuthis and E'noptroteuthis spinicauda are placed asjunior synonyms ofGrimalditeuthis bonplandi. Planktoteuthis. Chiroteuthis, Mastigoteuthis and Lepidoteuthis grimaldii respectively. An unknown type of paralarva referred to as "big-fin" is described. Members of the "chiroteuthid lineage" which apparently includes the Chiroteuthidae, Mastigoteuthidae, Joubiniteuthidae, Batoteuthidae, Promachoteuthidae and "big-fin" may all have secondarily derived tentacular clubs,

This paper is dedicated to the memory of Gil Voss and to the advancement of passion and controversy in science. Passion and controversy, in my opinion, are the life-blood of science. Gil was not only a prolific researcher, but a leader and a strong and passionate advocate for his field and he frequently stirred emotion and debate. Indeed, the considerable force of his personality may well have had a greater effect in advancing the field of cephalopod systematics and biology than did his substantial contributions to the literature. Gil Voss was not fond of the paralarval stages of the Chiroteuthidae. These stages, because of their large size and diverse forms, spawned a variety of generic and specific names that, for the most part, could not be merged with the nomenclature for older stages. He considered these names to be a time bomb (many had priority over subadultladult names) that someday would produce an ugly explosion and disrupt the systematics of the family. He was only partly correct. Un- fortunately, he can't be here to see that the explosion (at least its beginning) is not ugly at all. Surprisingly, the effect is providing order where confusion was growing. The family Chiroteuthidae was established by Gray (1849). Chun (1910) divided the family into three subfamilies: Chiroteuthinae, Mastigoteuthinae and Grimal- diteuthinae. Pfeffer (1912), however, considered the latter group to be a separate family. Naef (1923) fi)llowed Chun's arrangement as did Thiele (1935) and Voss (1956). Clarke (1966), however, followed Pfeffer, while Roper, Young and Voss (1969), Young (1972), Voss (1977), Nesis (1982/87) and Clarke and Trueman (1988) considered all three groups as separate families. Chiroteuth is, the oldest and best known genus in the Chiroteuthidae, is characterized by a distinctive paralarval stage that can reach large size. For many years these paralarvae were placed in the genus Doratopsis even after Ficalbi (1899) presented strong evidence that Doratopsis vermicularis was the young of Chiroteuthis veranyi. Naef (1923) clearly demonstrated that this relationship was valid. Pfeffer (1912), however, had rejected Ficalbi's conclusion and although Chun (1910) believed Ficalbi to be correct, he retained the genus. Chun's reluc- tance to synonomize Doratopsis is understandable today when we realize that he was actually dealing with the young stages of three different genera. The name "doratopsis" now is used to designate a developmental stage. Two other peculiar paralarvae have been placed in the "chiroteuthid" families. Chiroteuthoides hastula Berry, 1920, was considered to be a possible mastigoteuthid by Naef (1923) but a valid genus in the Chiroteuthidae by Nesis (1982/87).

162 YOUNG: CHIROTEUTHID PARA LARVAE FROM HAWAIIAN WATERS 163

Enoptroteuthis spinicauda Berry, 1920, was considered a young grimalditeuthid by Naef(1921; 1923) and Grimpe (1925). Nesis (1982/87), however, considered this species to be a probable member of the Cycloteuthidae. The family Mastigoteuthidae contains two genera, Mastigoteuthis and Echino- teuthis. The latter was described by Joubin (1933) and is based on two early juvenile/late paralarval specimens. This genus has been maintained by most sub- sequent workers (Voss, 1977; Nesis, 1982/87; Clarke and Trueman, 1988) even though subadults have never been described. The sole species in the family Joubiniteuthidae, Joubiniteuthis portieri, was originally placed by Joubin, 1912, in the genus Chiroteuthis. It was soon elevated to generic (Berry, 1920), then familial status (Naef, 1922). The paralarval stage has never been described. In this paper we will describe the paralarval stages of the Chiroteuthidae and related families from Hawaiian waters. We will demonstrate that Grimalditeuthis is one of four genera in the Chiroteuthidae, and that the primary feature relating these genera is the presence of a doratopsis stage in their development. We will, also, show that Berry's two species do not belong in this family and that Echinoteu- this is not a valid genus. We will describe a paralarva from an unknown genus/ family that shows some similarities to the chiroteuthid line. Finally, we will suggest that the "chiroteuthid families" are characterized by the loss of their "primary" tentacle clubs.

MATERIALS AND METHODS

Specimens were obtained from plankton tows taken from waters around the high Hawaiian Islands. Tows were taken in all seasons and primarily between 1985 and 1987. Most tows were made with a 4 m2 ring net with mesh sizes of 0.333 mm or 0.505 mm. Most tows were oblique between the surface and 300 m. The largest paralarvae were taken by a 3 m Isaacs-Kidd midwater trawl or a 3 m Tucker trawl (Young, 1978). Plankton samples were fixed in 4-5% seawater formalin and transferred to 50% isopropyl alcohol buffered with calcium carbonate chips after 24 h. After several years in preservation, most specimens exhibit some fading of chromatophore pigment. Scanning electron micrographs were taken with an I.S.I.-SS40 SEM. Voucher specimens are deposited in the National Museum of Natural History, Washington, D.C. Definitions. -Arm formula: One convention has been added to that of the standard arm formula: here the relative size between arms is indicated in the formula in addition to the size sequence. For example: Arm II 2 x I means that arm II is two times the length of arm I. Brachial pillar: Head between the anterior end of the eyes and the base of the brachial crown. Brachial pillar chambers and vesicles: Transverse septa from pillar surface to the digestive tract define chambers; septa forming chambers of lesser extent, often small and spherical spaces, define vesicles. CML: Mantle length measured to the conus of the giadius. Eyes (ventral projection): Percent of eye length that projects ventral to ventral surface of head. FL: Fin length. ML: Mantle length measured along the dorsal midline to the posterior end of the fins or for attenuate fins, to the point of maximum curvature on the posteroconcave fin margin. Neck: Region between posterior end of the optic lobes and the anterior end of the collar. N umber of neck septa: Number between anterior edge of collar and cephalic cartilage. Position of gill base: Distance, measured laterally, from anterior margin of collar to posterior end of gill. Collar: Muscular flaps lateral to the funnel that form a one-way valve for water flow into the mantle cavity. Shoulder of funnel: Region of junction between tubular funnel and collar. Caution must be used when relying on chromatophore patterns for species identification. While each species examined has a distinctive pattern, the patterns can be easily modified by damage during capture or differential fading. Patterns described here are not complete in all cases.

Chiroteuthis imperator Chun, 1910 Figures lE-H, M, N; 2; 4A

Material Examined.-68 specimens, 2-45 mm ML. 2-3 mm ML. Morphology. -Arms I and II: 2 and 3 (rarely 4) suckers respectively; basal sucker enlarged; arm II > I. Arm III: Not detectable. Arm IV: Ridge only. 164 BULLETIN OF MARINE SCIENCE. VOL. 49. NO. 1-2,199]

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Figure 1. A, B, I, J: Chiroteuthis sp, nov.; C, D, K, L: Asperoteuthis sp nov.; E-G, M, N: Chiroteuthis imperator. A, B: ventral and dorsal views, 5.3 mm ML. C, D: ventral and dorsal views, 4.2 mm ML. YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 165

Figure 2. Chiroteuthis imperator, 5.2 mm ML, scanning electron micrographs. A: tip of tentacle showing numerous buds. Scale bar = 20 ILm; B: lower magnification of same tentacle with fully-formed suckers in more than 4 series distally. Scale bar = 100 ILm.

Tentacle: 2-3 x eye diameter, robust with 14-18 suckers in two series over length of tentacle; 2-3 enlarged basal suckers. Brachial pillar: Virtually absent, eyes nearly abut buccal crown; buccal mass occupies entire region. Head: Eyes together (rather than individually) bulge ventrally. Neck: Length 112 ofML; about 7 septa. Funnel: Funnel cartilage with straight, shallow groove. Gladius: Terminates in a point just posterior to fins. Viscera: Digestive gland oblique, spindle-shaped; near midpoint ofML. Chromatophores. - Tentacle: One, at base on oral surface (3 mm ML). Head (ventral): One, large, centrally between eyes; one, elongate, posterior to each eye; one, median, just posterior to arms IV. Head (dorsal): One, posterior to or just over each eye (3 mm ML). 5 mm ML. Morphology (changes).-Arm IV: Small papilla. Tentacle: Two large suckers near base; scattered suckers along stalk in two series increasing to 4-6 irregular series in distal 113; tapering tip with numerous buds. Brachial pillar: Short, 1f4 of eye diam; esophagus well ventral of dorsal surface; one septum ventral to esophagus; dorsal to esophagus two well separated single series of several vesicles each. Head: Eyes approximately circular in lateral view; width across statocysts = or > width across optic lobes. Gladius: Extends well posterior to fins; invariably broken off. Viscera: Digestive gland at anterior 1/4 ofML; gill base at anterior 30% ofML. Chromatophores (additions).-Head (dorsal): One, large, posterior to each eye. Funnel: One, large, on each shoulder. 8 mm ML. Morphology (changes).-Arm III: Small papilla. Arm IV: Just> arm II; about 8-10 suckers in two series. Tentacle: Two large suckers at base followed by 2-3 of intermediate size and a short bare region, then a region of scattered suckers; on distal 1/4, suckers more densely packed into 6 irregular series which

~ E, F: ventral and dorsal views, 2.8 mm ML. G, H: ventral and dorsal views, 7.7 mm ML. I, J: ventral and dorsal views, 9.6 mm ML. K, L: ventral and dorsal views, chromatophore patterns missing. M, N: ventral and dorsal views, 12 mm ML. Scale bars-l.O mm. 166 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991 diminish in number near tip. Brachial pillar: Length Ih eye diam; esophagus a bit dorsal to center; dorsally (to esophagus) two adjacent single series of6-8 chambers, each series bordered laterally and proximally by three small vesicles in a single series. Ventrally, 3-4 major chambers on either side of medial septum and a single series of small vesicles along each dorsolateral margin. On ventral surface, a patch of small vesicles on proximal half of pillar. Funnel: Locking cartilage with deep, elongate oval groove. Mantle: Small vesiculate area at posterior tip of muscular mantle and anterior end of fins. CML overlaps fins by 15% of FL. Chromatophores (additions). -Arms: One on arm IV. Tentacle: Single series along aboral surface on distal3f4. Neck: At least 2 on dorsal midline. Mantle: One, large, ventrally on either side at level of anterior margin of fin. At least 2 more on dorsal midline. 12 mm ML. Morphology {changes).-Arms I and II: 3-4 and 6-7 suckers respectively. Arm III: Arm III nearly = I; few small suckers and two series of buds. Tentacle: One large sucker at base, otherwise, proximal 213of tentacle bare; distal IIJ divided into two portions, a proximal portion with suckers in 4 series (at least 30 rows) decreasing to 3 series at base of club, a distal portion that forms a distinct club; club with dorsal keel and protective membranes extending full length of club, suckers in 4 series (about 15 rows); club suckers larger in size than proximal suckers; club suckers of ventral series the largest, distal few with enlarged teeth. Brachial pillar: Length slightly> eye diam; dorsally two series of chambers bounded laterally on either side of pillar by a single series of small vesicles, becoming 2-3 series proximally. Head: Eye with small, silvery rostrum. Olfactory organ a small nipple ventral to each optic lobe. Funnel: Locking cartilage with tragus and antitragus. Viscera: Digestive gland in anterior 15-20% of ML. Gill base at 20% ofML. Chromatophores {additions).-Arms: Series on aboral surface of arm IV. Ten- tacle: One, at base on oral surface. Head (dorsal): One, posteromedial. Neck: At least 7 (total) in dorsal midline. Funnel: Second pair. Fins: 4 on dorsal midline. 18 mm ML. Morphology {changes).-Arms I and II: 4 and 8 suckers plus buds respectively. Arm III: Arm III = I < II. Arm IV: Length 6-7 x arm II. Tentacle: Scattered suckers proximal to club; distal toothed suckers smaller than large suckers of ventral series. Brachial pillar: Length 1.5 x eye diam. Head: Olfactory papilla on short stalk. Mantle: Vesiculate area extends anteriorly from fins about twice as far dorsally as ventrally. Chromatophores {additions).-Some increase in numbers but pattern unchanged. 35 mm ML. Morphology (changes). - Tentacle: Distal third oflong tentacle bearing four series of suckers in regular arrangement on stalk and bordered by low protective membranes. 45 mm ML. Morphology (changes). - Tentacle: Stalk with more numerous suckers. Brachial pillar: Length 2 x eye diam. Neck: Length 30% of ML. Comments. - The younger stages are unusual in the virtual lack of a brachial pillar, the central position of the esophagus within the pillar as the latter develops, the lack of protruding eyes and a median chromatophore ventrally between the eyes. These features are shared only with Asperoteuthis sp. See comments on this latter species for characters needed to distinguish these two. Although considerable changes occur during metamorphosis, the advanced paralarvae can be identified with one of the two appropriate subadult types by (1) the less complex arrangement YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 167 of vesicles in the dorsal region of the brachial pillar, (2) the less pronounced tragus and antitragus in the funnel locking-cartilage and (3) the somewhat more centrally located esophagus within the brachial pillar.

Chiroteuthis new species Figures lA, B, I, J; 4D

Material Examined.-28 specimens, 2-80 mm ML. 2-3 mm ML. Morphology. -Arms I and II: 1 and 2 suckers respectively; arm II > I. Arm III: Not detectable. Arm IV: Ridge only. Tentacle: Length 2 x eye diam; 2 irregular series of large suckers throughout; basal sucker is largest. Brachial pillar: Length 1.3 x eye diam; esophagus runs along dorsal wall; approx. 6 pairs of large chambers. Httad: Eyes irregular (including small bump on posterior margin) but approximately circular in outline. Neck: Length 112 of ML; number of septa uncertain. Gladius: Terminates in a point just posterior to fins. Viscera: Digestive gland spindle-shaped, oblique. Chromatophores. - Tentacle: One, large, at base on oral surface; one, small on aboral tip. Brachial pillar: One, median, near base of arms IV. Head (ventral): One, median, between anterior region of eyes; one, small, posterior to each eye. Head (dorsal): One, large, posteromedial to each eye; one, median line near statocyst. Neck: One, small, on dorsal midline. Mantle: Several on dorsal midline. 5 mm ML. Morphology (changes).-Arm IV: Papilla. Tentacle: Length 3x eye diam; large basal sucker followed by few scattered suckers in two series in proximal half; distal halfwith suckers more closely packed and in 4 series becoming more numerous and irregularly arranged near tapering tip where they are replaced by numerous buds. Brachial pillar: Length 2 x eye diam; 6-7 pairs of chambers. Head: Eyes together (rather than individually) bulge ventrally; olfactory organ just detectable as a pad immediately posterior to each eye. Neck: 6-8 septa. Funnel: Locking cartilage with straight groove. Mantle: Vesiculate area represented by a few small vesicles ventrally. CML overlaps fin by 20% of FL. Gladius: Extends well posterior to fins; invariably broken off. Viscera: Digestive gland at anterior 1/4 of ML; gill base at anterior 25% of ML. Chromatophores (additions). - Tentacle: Two on aboral surface near tip. Neck: At least one in dorsal midline. Funnel: One on each dorsolateral surface near shoulder. Mantle: Series on dorsal midline. 8-10 mm ML. Morphology (changes). - Arms I and II: 2-3 suckers and 3-5 suckers respectiveiy. Arm III: Papilla. Arm IV: Arm IV 1-2 x II; up to 20 suckers in two series. Tentacle: Length 40-50% of ML; proximal 30-50% bare except for large basal sucker; distal portion with suckers in 4 irregular series becoming more compact and in 5 series near tapering tip; tip covered with buds. Brachial pillar: Length 2.5 x eye diam; 4 slender chambers anterior to 6 major chambers. A single incomplete series of vesicles on either side of dorsal surface extends anteriorly 113 of pillar length. Head: Eyes with short, blunt, ventral rostrum. Funnel: Locking cartilage with elongate oval depression. Mantle: Vesiculate area extends around mantle. Chromatophores (additions).-Arm IV: Aboral series present. Tentacle: Trans- verse chromatophores over much of aboral surface of distal half; one basal. Bra- chial pillar: One, dorsal midline, anterior. Mantle: At least 9 along dorsal midline; 6-8 scattered over sides and ventral surface. Fins: At least 5-6 dorsally along midline between fins. 168 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

13-15 mm ML. Morphology (changes). -Arm III: From greatly to slightly shorter than arm I. Arm IV: 6 times length of arm II, about 112 tentacle length; numerous suckers. Tentacle: Except for large basal sucker, suckers only on distal half and in 4-5 irregular series; club not clearly defined but keel present. Brachial pillar: Length 3 x eye diam. Funnel: Locking apparatus with tragus and antitragus just detectable. 18 mm ML. Morphology (changes).-Arms I and II: Approx. 4 and 6 suckers respectively. Arm III: Arm II > III > I. Arm IV: Arm IV lOx II. Tentacle: Stalk suckers cover distal quarter of which the distal third is a distinct club; suckers on club in ventral series are largest, distal 3 suckers in both ventral rows with enlarged teeth; dorsal keel and protective membranes extend full length of club; stalk suckers in 4 irregular series and flanked by protective membranes. Brachial pillar: Sheet of small vesicles (7-8 vesicles wide at mid-level) covers dorsal surface above esophagus. Mantle: Very slender, width < 10% oflength. Viscera: Gill base at anterior 15% of ML. 45 mm ML. Morphology (changes).-Arms: All vesiculate and with numerous suckers; arm III ~ II > I. Tentacle: Club with slender tip and terminal pad with suckers. Stalk with more numerous suckers in 4 regular series, protective membranes more developed. Brachial pillar: Length 4 x eye diam; esophagus does not quite reach dorsal wall; dorsal vesiculate region with 10-13 vesicles across; ventral surface with layer of small vesicles (6 across); 9 major chambers with several oblique half-chambers posteriorly. Head: Olfactory papillae present ventral to optic lobes. Mantle: Vesiculate area extends anteriorly from fins along dorsal midline for 30% of FL. 65 mm ML. Morphology (changes).-Brachial pillar: Length about 60% of neck length; esophagus about 20% of pillar diameter below dorsal surface. Head: Ol- factory papillae slightly posterior to optic lobes. Neck: 1;j of ML. Comments. - The distinctive dorsal position ofthe esophagus within the brachial pillar is shared only with Planktoteuthis danae. These two species differ in eye shape, ventral eye protrusion, number of brachial chambers and numerous other characters. The smallest metamorphosed subadult seen is 65 mm ML while the largest paralarva is 80 mm ML. This suggests considerable variation in the size at metamorphosis. The undescribed adult is closely related to the species described by Nesis and Nikitina (1984) and incorrectly identified as Chiroteuthis joubini.

Asperoteuthis new species Figure 1C, D, K, L

Material Examined.-16 specimens, 2-25 mm ML. 4 mm ML. Morphology. -Arms I and II: 5 suckers each; arm I = II. Arm III: Papilla. Arm IV: IV ~ I, with one sucker. Tentacle: Long, thick; enlarged basal sucker; suckers scattered in 2 irregular series becoming 4 series in distal third; buds at tip. Brachial pillar: Length 1J2 long axis of eye; esophagus in center; dorsal to esophagus numerous vesicles (about 6 across); number of chambers uncertain. Head: Eyes slightly elliptical, bulge ventrally slightly individually. Neck: Length 112 of ML. Funnel: Straight, broad cartilage lacking groove. Gladius: Extends well posterior to fins, invariably broken off. Viscera: Digestive gland oblique, spindle- shaped, located mid-mantle; gill base at anterior 40-45% of ML. Chromatophores. -Anns: One, at base of arm IV. Tentacle: One, at base on oral surface; series on distal aboral surface. Brachial pillar: One, near base of arm IV. YOUNG: CHIROTEUTHID PARA LARVAE FROM HAWAIIAN WATERS 169

Head (ventral): One, central between eyes; one, posterior to each eye; one, small, posterolateral to each eye. Head (dorsal): One, posterior to each eye; one, small, posterolateral to each eye. Neck: Several on dorsal midline. Funnel: One, dorsally at each shoulder. Mantle: Two on dorsal midline; one pair ventrally near fins. 6 mm ML. Morphology (changes). -Arms: Arm II > I = IV, III a papilla. Mantle: Vesiculate area present surrounding mantle; extends dorsally from fins for 112 FL; ventrally, vesiculate area does not extend anterior to fins. 19 mm ML. Morphology (changes).-Arms: Arm IV 5 x II > I = III; vesiculate; arms IV with numerous well-spaced suckers in 2 series become very small distally. Tentacle: Muscular; length?; arm IV. Funnel: Locking cartilage with straight groove. Mantle: Length dorsal vesiculate area from fin = FL. Fin: FL = 34% of ML. Viscera: Digestive gland horizontal; gill base at anterior 30% of ML. 25 mm ML. Morphology (changes).-Arms I-III: Arm II = III slightly> I. Arm IV: Arm IV 2 x III; numerous suckers in two closely-aligned series. Tentacle: Muscular; about 1.5 times ML; distal portion divided into carpal area (unex- panded) with 4 series of suckers (8-12 rows) becoming two series where they merge with expanded club; remainder of stalk bare; club with suckers in 4 series, decreasing to 2 at tip; ventral series with largest suckers; dorsal keel and protective membranes along entire club; carpal region bounded by protective membranes which extend further down stalk for a distance equal to the carpal length. Brachial pillar: Length uncertain; numerous vesicles posteriorly; pattern of chambers cannot be determined. Head: Olfactory organ papilla beneath posterior margin of optic lobe; optic lobes not separate anteriorly. Neck: Length 25% ofML. Funnel: Locking cartilage with broad, straight groove. Mantle: Vesiculate area extends anteriorly along dorsal mantle to anterior quarter ofML. Fins: 40% ofML. Viscera: Digestive gland elongate, horizontal; gill base at anterior lki of ML. Comments.-Large specimens were badly damaged. Young stages are easily con- fused with Chiroteuthis imperator. Young stages of Asperoteuthis sp. differ from this latter species by the chromatophore on the oral surface of its tentacle base, its greater number of arm suckers, more numerous dorsal brachial pillar vesicles, the dorsally elongate vesiculate area (6 mm ML) and by the lack of a groove in the funnel cartilage. At larger sizes the shape of the mantle vesiculate area, the more circular shape ofthe combined fins and the straight funnel locking-cartilage is diagnostic. The larger paralarvae can be related to subadults on the basis of the shape of the funnel locking-cartilage, the shape of the dorsal mantle vesiculate region and by default. The undescribed adult differs from the only described species, Asperoteuthis acanthoderma, in the lack of dermal denticles and the lack of an antitragus in the funnel locking-cartilage among other features, but retains the most diagnostic feature of the genus: tentacular clubs with suckers restricted to the distal half.

Planktoteuthls danae (Joubin, 1931) Figures 3D, E, I, J; 4E

Material Examined. -32 specimens, 2.0-21 mm ML.

2.5 mm ML. Morphology. -Arms I and II: 2 and 3 suckers resp., arm II = I. Arm III: Not detectable. Arm IV: Nipple. Tentacle: Short (1.0-1.5 x long axis of eye), tapering; about 8 suckers. Brachial pillar: Length 112 long axis of eye; esophagus along dorsal surface, muscle bundles along dorsal and ventral surfaces are strong 170 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. \-2, 199\

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Figure 3. A-C, F-H: Planktoteuthis sp.; D, E, I, J: Planktoteuthis danae. A, B: ventral and dorsal views, 3.7 mm ML. C: distal tentacle with club (2.3 mm), 19 mm ML. D, E: ventral and dorsal (chromatophore pattern missing) views, 8.4 mm ML. F-H: ventral view, side view of head showing eye shape and ventral eye protrusion, dorsal view, 14.4 mm ML. I, J: ventral and dorsal views, 22 mm ML. Scale bars-I.O mm. c

Figure 4. A-E: side views of heads showing position of esophagus in brachial pillar. A: Chiroteuthis imperator. 2.4 mm ML. B: Grimalditeuthis bonplandi, about 2.7 mm ML. C: Planktoteuthis sp., about 2.5 mm ML. D: Chiroteuthis sp. nov., 2.4 mm ML. E: Planktoteuthis danae, 2.6 mm ML. Asperoteuthis sp. nov. not illustrated but appears very similar to Chiroteuthis imperator. F: Aspero- teuthis sp. nov., distal tentacle with club, 27 mm ML. G-M: Grimalditeuthis bonplandi. G, H: ventral and dorsal views, 3.9 mm ML. I, J: ventral and dorsal views, 7.2 mm ML. K: distal tentacle with club, 19 mm ML. L, M: ventral and dorsal views, 17 mm ML. Scale bars: 0.5 mm for side views of heads and tentacles, 1.0 mm for whole views. 172 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991 and opaque; 2 chambers. Head: Eyes elongate (width just > lens diam) with rostra, each protrudes anteroventrally (112 eye length). Neck: Length 112 ofML; number of septa uncertain (at least 1). Gladius: Terminates in a broad point at posterior end of short fins. Viscera: Digestive gland spindle-shaped, obliquely oriented in midmantle. Chromatophores. - Buccal mass: One small on aboral surface at base in midventral line. Tentacle: One small on aboral tip. Head: One small, between eyes on midventral surface. Mantle: Two on lateral side. 5-6 mm ML. Morphology (changes).-Arms I and II: 3 and 5-6 suckers resp. Arm III: Nipple. Arm IV: Arm IV > II; many small suckers. Tentacle: Small, compact club with suckers in 4 series; scattered suckers along stalk, large basal sucker. Brachial pillar: Length = long axis of eye; 3 major chambers. Funnel: Locking cartilage with straight, broad groove. Viscera: Digestive gland at anterior 1f3 ofML; gill base at anterior 30% ofML. Gladius: Extends well posterior to fins, invariably broken off. Chrornatophores (additions).-Buccal mass: One small, aboral surface at base in dorsal midline. Arm IV: Two aborally. Tentacle: Two, aboral to club; one aborally at base of stalk. Head: One pair dorsally; one lateral, posterior to eye. Mantle: Two on ventral midline near tail. 12 mm ML. Morphology (changes).-Arms I-III: Arm II > I slightly> III; numerous suckers; Arm II vesiculate. Arm IV: Arms IV 10 x II; extremely small suckers in virtual single row along most of arm length; vesiculate. Tentacle: Length = arm IV; robust; compact club with keel and only few carpal suckers; stalk robust, without suckers. Brachial pillar: Length 2 x long axis of eye; thin layer of vesicles dorsal to esophagus. Funnel: Antitragus on funnel cartilage just detectable. Mantle: Small vesicular area near fins. Viscera: Gill base at anterior 25% of ML. Chromatophores (additions).-Arm IV: Group at aboral base. Tentacle: 3 series on aboral surface of cluo. Funnel: One dorsally at junction with collar. Mantle: Group on ventral mantle near fins. 21 mm ML. Morphology (changes).-Arms: Arm IV ~ III = II> I; all vesiculate; arm IV suckers forming nearly single series, broadly spaced throughout arm length. Tentacle: Length 1.5 x arm IV; clubs with few slightly enlarged suckers distally in ventral series but without enlarged teeth. Neck: Length 1/4 of ML. Comments. - The most distinctive feature in the smaller stages is the dorsal position of the esophagus in the brachial pillar and the small number of pillar chambers. The midventral chromatophore between the eyes separates this species from its congener. Most specimens were badly damaged. The broad fins suggest that the adult is Planktoteuthis danae; however, subadults and intermediate stages have not been taken here.

Planktoteuthis sp. Figures3A~,F-H;4C

Material Examined.-28 specimens, 2-19 mm ML. 2.5-3.0 mm ML. Morphology. -Arms I and II: Subequal, each with 2 suckers. Arm III: Undetectable. Arm IV: Small ridge. Tentacle: Short (1.0-1.5 x long axis of eye), and tapering; 4-10 suckers depending on size; basal sucker just slightly larger. Brachial pillar: Length 112 x long axis of eye; esophagus passes approximately YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 173 through center; approximately 4 chambers dorsally and ventrally. Head: Eyes elongate (width about equal to lens diameter) and each project anteroventrally (If2 of eye length with midpoint of lens level with ventral surface of head). Neck: Length 2-3 times length of optic lobe; probably 2 septa. Gladius: Does not extend past tiny fins. Viscera: Digestive gland spindle-shaped, oblique, midmantle; gill base at anterior 45% of ML. Chromatophores. - Buccal mass: One, posteroventral midline. Brachial pillar: One, ventral midline. Head: One pair on dorsal surface. 4.0 mm ML. Morphology (changes).-Arms I and II: Arm II with 3 suckers; arm I uncertain. Arm III: Nipple. Arm IV: Arm I = II > IV. Tentacles: Suckers throughout; at least 12 suckers plus terminal buds; basal sucker largest. Funnel: Locking cartilage oval with oval depression. Gladius: Terminates in a point just posterior to fins. Chromatophores (additions).-Funnel: One dorsally at junction with collar. 5.0 mm ML. Morphology (changes).-Arms I and II: 3 suckers and 4 suckers respectively. Arm III: Much the smallest. Arm IV: Arm IV slightly> II; 3 suckers. Tentacle: Length 4 x long axis of eye, tapering; scattered suckers along most of length with cluster of small suckers near tip; large basal sucker. Brachial pillar: Small vesicles at anterodorsal end. Head: Width across statocysts > across optic lobes. Funnel: Locking cartilage with antitragus. Chromatophores (additions).-Arm IV: Two on aboral surface. Tentacle: 3 on aboral surface. Head: One on lateral surface posterior to eye. 11 mm ML. Morphology (changes). -Arms I and II: 7 and 10 suckers respectively. Arm III: 3 suckers and numerous buds; arm II > I = III. Arm IV: Arm IV 5 x II; vesiculate. Tentacle: Stalk robust, without suckers except near club; club damaged. Brachial pillar: Additional layer of vesicles ventrally. Head: Eyes with posteroventrally protruding, pointed rostra; olfactory papillae ventral to optic lobe. Mantle: Vesicular tissue near fins; more extensive dorsally where it extends anteriorly from fins 40% ofFL. Gladius: Extends well posterior to fins; invariably broken off. Viscera: Digestive gland horizontal; gill base at anterior 30% of ML. 15 mm ML. Morphology (changes).-Arms: All with numerous suckers; arm IV suckers very small, in nearly single series throughout arm. Tentacle: 8-10 rows of small suckers in 4 series on carpus. Suckers on club much larger, in 4 series, grade gradually in size with largest in ventral series, size disparity accentuated near tip; 2 series at tip; protective membranes extend to tip; dorsal keel present. Brachial pillar: Numerous vesicles dorsally and ventrally. Neck: At least 4 septa; length If2 of ML. Mantle: Dorsal vesicular region extends anteriorly in narrow region the length of the fins from anterior end of fins. 19 mm ML. Morphology (changes).-Arms: All vesiculate; IV :> III slightly> II slightly > I. Tentacle: Carpal region consisting of 4 series of small suckers; length of carpal region about 75% oflength of remainder of club. Comments. -Species of Planktoteuthis are characterized in the early stages by the strong ventral projection of the eyes, possession of only a few suckers on the arms and tentacles, relatively small, tapering tentacles and a reduced vesicular region on the mantle. Planktoteuthis sp. is most easily separated from P. danae by the central position of the esophagus in the brachial pillar, two (vs. one) chromatophores on the ventral midline near the arms (one is on the ventral base of the 174 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991 buccal mass) and the lack of a chromatophore between the eyes. This species is similar to P. lippula Chun, 1908, but differs in the structure of the tentacle club and ventral arms.

Grimalditeuthis bonplandi (Verany, 1837) Figure 4B, G-M

Material Examined.-37 specimens, 2-24 mm ML. 2-3 mm ML. Morphology. -Arms I and II: 3 and 4 suckers respectively, arm I = II. Arm III: Notd(~tectable. Arm IV: Ridge only. Tentacle: Two series of suckers; basal 2-3 largest. Brachial pillar: Length = long axis of eye; esophagus passes through center; 4 pairs of chambers dorsally, 3 or 4 ventrally. Head: Eye elliptical with ventral rostrum; each eye protrudes ventrally. Neck: Length 112 of ML; 6 septa; width at collar about twice head width across optic lobes. Gladius: Pointed tip reaches just poste:rior to fins. Viscera: Digestive gland spindle-shaped, oblique, in midmantle region. Chromatophores. - Tentacle: Two, at base on oral surface. Brachial pillar: One, ventral midline near base of arms IV. Head: Two, posterior to each eye on ventrolateral surface. 4-5 mm ML. Morphology (changes).-Arms I and II: 3-4 suckers plus buds and 5-6 suckers plus buds respectively; basal suckers enlarged. Arm III: Papilla. Arm IV: Arm I > IV; approx. 6 sucker buds in 2 series. Tentacle: 1-3 large suckers at base followed by scattered suckers in 2 irregular series becoming more compact in arrangement in distal 113. Head: Eye slightly elongate; silvery rostrum present. Neck: Length 113 of ML. Funnel: Locking cartilage with straight groove. Mantle: Vesiculate region surrounding posterior end of mantle; length equals FL. Gladius: Extends well posterior to fins; invariably broken off. Viscera: Digestive gland at anterior 40% of ML; gill base at anterior 50% of ML. Chromatophores (additions). -Arms: 2 on arm IV. Tentacle: About 4, transversely elongate, aboral surface of stalk. Brachial pillar: 2, anteroventral midline; one, anterodorsal midline. Head (ventral): 2, posterior to each eye. Funnel: 2-4, small, dorsal surface on each shoulder. Mantle: Up to 4 on ventral surface of vesiculate area. 6-7 mm ML. Morphology (changes).-Arms I and II: 5 and 6 suckers respectively, with buds. Arm III: Arm II > I ~ III. Arm IV: Length 2 x arm II; 6 sucker buds in two series. Tentacle: 2 or 3 large basal suckers; broadly scattered small suckers along stalk in two irregular series; distal 20-25% forms club; carpal area with suckers in 2 irregular series merging into expanded club with suckers in 4 series; dorsal keel present. Head: Eyes project ventrally only slightly; olfactory organ a small nipple just anterior to the posterior wall of the statocyst. Mantle: Vesiculate area separates muscular mantle from fins by 213 ofFL, anterior extension dorsally does not exceed that ventrally. 9 mm ML. Morphology (changes).-Arm III: II > I = III. Arm IV: Length about 3 times arm II; numerous suckers. Tentacle: 0-1 basal suckers, otherwise, stalk bare; carpal region with about 10 suckers in 2 irregular series becoming 3 series (3 rows) then 4 series (about 10 rows) on expanded club but decreasing in number near tip; suckers subequal in size. Brachial pillar: Length 2 x long axis of eye. Head: Eyes nearly circular, do not protrude ventrally; width across statocyst greater than across eyes; optic lobes slightly separated anteriorly; superior buccal and brachial lobes adjacent to optic lobes. Neck: Length 1/4 ofML; olfactory papilla YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 175 at 40% of distance between optic lobes and collar. Funnel: Shoulder region with numerous vesicles. Viscera: Gill base at 30% of ML. Chromatophores (additions).-Head: Numerous small chromatophores over dorsal and ventral surfaces just posterior to eyes. Neck: Scattered small chromatophores on dorsal surface. Funnel: At least 8 on each shoulder. Mantle: Few, small, near dorsal midline; one large pair ventrally near fins.

15-17 mm ML. Morphology (changes).-Arms I-III: Arm II = III > I; all with numerous suckers. Arm IV: Arm IV 2 x II; numerous suckers in two series. Tentacle: Club suckers equal in size except for 4 suckers near tip which are enlarged and bear enlarged teeth. Brachial pillar: Length 3.5 x eye diam (without rostrum); esophagus passes just dorsal of center; arrangement of chambers uncertain. Head: Eyes small and nearly circular with blunt rostrum; optic lobes broadly separated anteriorly; superior buccal and brachial lobes well separated anteriorly from optic lobes and each other. Neck: Olfactory papilla lateral to base of funnel (85% of distance from optic lobes to collar). Funnel: Locking cartilage with broad, straight groove that ends abruptly posteriorly; mantle component nose-like. Fins: Com- bined outline round. Viscera: Digestive gland at anterior 1/4 of ML. Chromatophores (additions). - Brachial pillar: Distinctive series of transversely oriented chromatophores across ventral surface at level of anterior edge of eye, incomplete in midline. Row of small chromatophores on lateral surface from anterior edge of each eyelid to base of each arm III. Head and Neck: Series of small elongate chromatophores along lateral side of head and neck to olfactory organ (marks path of olfactory nerve for most of its length); other lines of small chromatophores approximately follow posterior and near-anterolateral edges of each optic lobe; small, scattered chromatophores on dorsal surface of head, funnel, etc. but pattern uncertain. 22 mm ML. Morphology (changes).-Funnel: Locking cartilage with groove in anterior 213 only; slightly undercuts posterior shelf. Comments. - Young squids are characterized by a long brachial pillar with a central esophagus. This feature is shared with Planktoteuthis sp. Other features that, in combination, separate small Grimalditeuthis from this and the other species are: number of chromatophores on the funnel shoulders and ventral head posterior to the eyes, eye shape and shape of vesiculate area on the mantle. At sizes larger than 9 mm ML numerous features distinguish this species: anterior separation of optic lobes, position of superior buccal and brachial lobes, eye size, position of olfactory papillae and numerous features of the chromatophore pattern. Between 9 and 15 mm ML considerable changes occur which caused some dif- ficulty in linking the stages of this species. The critical features that allowed the gap to be bridged are the shape of the vesiculate area on the mantle (i.e., equally developed dorsally and ventrally) and the more posterior position of the olfactory organ. The larger specimens described here appear to be identical to Doratopsis sagitta Chun, 1910.

Family Joubiniteuthidae

Joubiniteuthis portieri (Joubin, 1912)

Material Examined.-6 specimens, 5.8-18 mm ML. 6-7 mm ML. Morphology. -Arms I and II: Arm I = II; suckers in two series (9- 10/arm); largest suckers are most proximal; attenuate tips bare; vesiculate. Arms 176 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

III and IV: Papillae. Tentacle: Short (2.5-3.0 x long axis of eye) and thick (width 20% oflength excluding slender tip). Suckers (approx. 25) over nearly full length; approximately uniform in size except for few small suckers near tip. Two irregular series proximately increasing to 4 irregular series distally (7 mm ML). Tips attenuate and bare. Brachial pillar: Short; 1;5 long axis of eye. Head: Eyes elliptical, project ventrally slightly. Small vesicles present on dorsal and ventral surfaces between eyes and arm bases. Head widest across eyes; width across statocysts greater than width across optic lobes. Short olfactory papilla just posterior to each eye (7 mm ML). Neck: Present; anterior edge of collar at level of posterior end of statocysts. Funnel: Locking cartilage elongate-oval with similarly shaped depression. Mantle: Muscular mantle terminates near anterior edge of fin. Fins: Small, length about 5% of ML. Gladius: Length posterior to fins slightly greater than ML. Viscera: Digestive gland cylindrical, horizontal, abuts cephalic cartilage. Chromatophores. - T,entac1e: Numerous scattered chromatophores over surfaces. Head: Scattered chromatophores over dorsal surface, above brain; two large chromatophores on each ventrolateral surface of head posterior to eyes. Mantle: Scat- tered on dorsal surface and on tail. 9 mm ML. Morphology (changes).-Arms: Arm II slightly> I > 3x III = IV; arms III and IV with sucker buds. Tentacle: Suckers in two series proximally becoming 4-5 series distally; numerous buds near tip; large proximal suckers; tip abruptly attenuate. 18 mm ML. Morphology (changes).-Arms: Suckers in 5-6 series; arm II = III > I > IV. Tentacle: Suckers in 8 or more series; club with protective membrane but no dorsal keel. Brachial pillar: Virtually absent. Head: Long olfactory papilla at level of posterior margin of optic lobe. Neck: Present; elongate, anterior margin of collar does not quite reach posterior end of cephalic cartilage. Funnel: No medial depression between adductor muscles; funnel adductor muscles arise from head over midpoint of statocysts. Mantle: CML overlaps fin by 35-40% of FL. Fin: Anterior and posterior lobes present. Viscera: Gill base at 15% of ML. Comments. -Paralarvae are very distinctive and easily identified to this species by their small fins and long tails.

Family Mastigoteuthidae

Mastigoteuthis famelica (Berry, 1909) Figure 5A, B, F, G

Material Examined. -30 specimens, 3.5-25 mm ML. 4-5 mm ML. Morphology. -Arms I and II: Arm I with 4-5 suckers in two series; arm II with about 6 suckers in two series. Basal sucker not especially enlarged. Arm III: Papillae. Arms IV: Arms IV <: II (4 mm ML); arm IV about = II (5 mm ML). Tentacle: Length 112 ML; suckers throughout but more numerous in distal 1/3; arranged in two series proximally becoming four irregular series distally; approximately 45 suckers; basal sucker enlarged; numerous buds at tip. Brachial pillar: Short (length 1f4 long axis of eye); vesicles and chambers absent. Head: Head narrow; width across statocysts greater than width across region of optic lobes; eyes elliptical and project ventrally. Neck: Short; collar just overlapping statocysts ventrally but more extensively dorsally but does not reach optic lobes. Funnel: Locking cartilage with straight, shallow depression. Mantle: CML overlaps 1f4 of FL. Gladius: Ends in a point a short distance posterior to fins (1f4-1f3 of YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 177

Figure 5. A, B, F, G: Mastigoteuthisfamelica; C, H: loubiniteuthis portieri; D, E, I, J: Mastigoteuthis inermis. A, B: ventral and dorsal views, 4.8 mm ML. C: dorsal view, 6.9 mm ML. D, E: ventral and dorsal views, 4.6 mm ML. F, G: ventral and dorsal views, 8.5 mm ML. H: ventral view, 6.9 mm ML. I, J: ventral and dorsal views, 7.2 mm ML. Scale bars-l.O mm. 178 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

FL). Viscera: Digestive gland (5 mm ML) with bent spindle shape; oblique only at anterior end. Gill base at anterior 25% of ML. Chromatophores. -Arms: Single series on aboral surface of arm IV. Tentacle: Single series on distal 213 of aboral surface. Head: One in midventralline of head near base of arms IV; one posterior to each eye on ventral surface and on dorsolateral surface of head; additional pair on dorsal surface of head appears in slightly older paralarvae. 7-9 mm ML. Morphology. -Arms I and II: Arm I with 5 suckers, arm II with 7 suckers (7 mm ML). Arm III: Papillae. Arm IV: Arms IV ~ II (7 mm ML); arm IV 2 x II (9 mm ML). Tentacle: Buds at tip; largest specimen with intact tentacles was 11 mm ML. Brachial pillar: Length 213 long axis of eye. Head: Silvery rostrum projects ventrally or anteriorly depending on eye rotation. Funnel: Locking cartilage with broad, straight groove. Mantle: Slender; CML overlaps 15% of FL. Fin: Length 1/3 of ML; anterior and posterior lobes lacking. Gladius: Gladius extends beyond fins by 25% of FL. Viscera: Gill base at 17% of ML. 17 mm ML. Morphology (changes).-Arms: Arm IV 4x II; arm II > I > III; not vesiculate. Tubercles: Small multicuspid tubercles are present over the surface of the mantle, head and aboral surfaces of arms (bases only of dorsal arms). 25 mm ML. Morphology (changes).-Funnel: Locking cartilage oval and bearing tragus and antitragus. Head: Eye abuts arm bases when rotated with rostrum directed anteriorly. Olfactory papillae not detectable. Tubercles: Well developed, also on funnel. Comments. -Among paralarvae, tubercles were found only in large M. famelica. In subadults of 40 mm ML they are still present but more difficult to detect. These paralarvae are easily identified to this species by their very elongate shape and narrow elongate fins.

Afastigoteuthis inermis Rancurel, 1972 Figure 5D, E, I, J

Material Examined.-25 specimens, 3.5-12.8 mm ML. 4-5 mm ML. Morphology. -Arms I and II: Arm I with 4 suckers, arm II with 8 suckers; proximal suckers are the largest; arm II > I. Arm III: Barely detectable. Arm IV: Arm IV nearly = I. Tentacle: At 4 mm ML, suckers scattered (number uncertain) along entirt: stalk with large basal sucker; at 5 mm ML, length 1/2 of ML or 4 x arm II; suckers only in distal 11J; at least lOin two series. Brachial pillar: Short (length 11.1 long axis of eye), apparently to accommodate forward rotation of eye; chambers and vesicles absent. Head: Eyes elliptical and protrude ventrally. Width across statolith> across optic lobes. Neck: Short; anterior margin of collar just overlaps statocysts ventrally but more strongly dorsally but doesn't reach optic lobes. Funnel: Locking cartilage with just detectable, elongate depression, funnel adductors arise from head over posterior IIJ of statocyst. Mantle: CML overlaps 1/4-IIJ of FL. Gladius: ends in a point a short distance (about 1/3 of FL) posterior to fins. Viscera: Digestive gland spindle-shaped, oblique, situated at anterior 11J of ML. Chromatophores. - Tentacle: Single series on aboral surface in region of club. Arms: Single series on aboral surface of arm IV; one at tip of each arm II; one YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 179

in midventralline of head near base of arm IV. Head: One behind each eye on ventral surface; four on dorsal surface posterior to eyes. Mantle: One ventral pair just anterior to fins; one on dorsal midline at posterior end of fins. 7-9 mm ML. Morphology (changes).-Arms: Arm IV > II > I; arms III are papillae. Tentacles: Distal suckers in 2-4 irregular series (number of apparent series may be affected by coiling of the tentacle); buds absent from tip; largest specimen with intact tentacles was 7 mm ML. Funnel: Locking cartilage with straight groove. Fin: Length lis of ML with anterior and posterior lobes. Viscera: Digestive gland at anterior 28% of ML; base at anterior 1/3 of ML. 12.8 mm ML. Morphology (changes).-Arms: Arm IV 2.5-3 x II. Arms III nearly = I. Head: Eyes abut arm bases; olfactory organ a small pad half way between each eye and posterior end of optic lobe. Mantle: CML overlaps 112 FL; no tubercles in skin. Fin: Slightly longer than combined width. Comments. - These paralarvae are identified as M. inermis by default: this is the only other species of Mastigoteuthis known from Hawaiian waters. At 5 mm ML M. famelica is more slender, has smaller eyes, a more anterior digestive gland, a more extended coverage of tentacular suckers and a more extended arrangement of chromatophores on the tentacle than does M. inermis. The shape differences, including fin shape, are accentuated with increasing size. The specific differences between these two species diminish as size decreases below 4.5 mm ML. However, the position of the digestive gland is a useful character to at least 3.5 mm ML. Smallest paralarvae captured were 2.5 mm ML and contained yolk indicating that they hatch from rather large eggs.

Family Lepidoteuthidae

"Enoptroteuthis spinicauda" Berry, 1920 Figure 6D-F

Material Examined.-One specimen, 9.7 mm ML.

Morphology. -Arms: Arm II > I 2: III = IV; arms I and II bear two series of tightly packed globular suckers; sucker size less than half the diameter of the largest club suckers; suckers of arms III and IV also globular but smaller. Tentacle: Length> arm II with thick, bare stalk and short terminal clubs. Club with 5 large suckers (largest of these is most proximal) and 3 very small suckers (2 at club base and one at tip); club with protective membranes but no keel. Brachial pillar: Present (length 112 eye diam); eyes well posterior to base of arm crown; arm crown is supported only by a narrow stalk due to loss of delicate outer tissues during capture. Head: Eyes circular, bulge laterally from head; loss of head integument gives false appearance of stalked eyes (eyes abut optic lobes which in tum abut central brain); width across statocysts < width across optic lobes; olfactory organ not seen. Neck: Present; anterior margin of collar overlaps the statocysts but doesn't reach optic lobes. Funnel: Adductor muscles arise from head at anterior 113 of statocyst; the funnel cartilage elongate with straight, long but shallow central depression. Mantle: Mantle muscle thin; CML overlaps fins by 5% of FL; FL > CML. Tubercles: Absent. Gladius: Gladius extends posterior to fin '12 of FL in a slender tail. Fin: Anterior and posterior lobes lacking. Viscera: Digestive gland rounded cone-shape with base anterior, gland nearly abuts cephalic cartilage; gills base at 39% of the CML or 19% of the ML; large ink sac present. 180 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

Chromatophores. - The chromatophore pattern is mostly obscured by damage except for the presence ofa few chromatophores on the aboral surface of the club, the aboral surface of arm IV and a few on the ventral surface of the head. Comments. - This specimen differs from Berry's primarily in the more robust tentacles which carry more suckers and in a slightly different shape of the funnel cartilage. These apparent differences may be due simply to damage and interpre- tation. The identity of this paralarva is examined in the Discussion.

Family, genus, species unknown ("big-fin") Figure 6A-C

Material Examined.-One specimen, 19.1 mm ML. Morphology. - Arms I and II: Suckers in two rows; proximal 4 suckers large, distally they progressively decrease in size (total count about 10/arm); tips bare, tips attenuate; arm I = U. Arms III and IV: Several large suckers proximally, sharp decrease in size distally (total count, including buds, about 12 and 20 respectively). Suckers in 3 (arm III) or 4 series (arm IV) at point of maximum density; tips attenuate; I = II > IV > III. Tentacle: Short and broad; width nearly 50% of length; suckers throughout, 2 large basal suckers, suckers in 2 series at base, shortly grading into 7-8 series with numerous buds near tip. Suckers in region of greatest density larger than basal suckers. Protective membranes present; dorsal keel absent. Brachial pillar: Absent, eyes abut arm bases. Head: Eyes large, hemispherical, protrude ventrally; small olfactory knob on posterolateral surface of each eye; width across statocyst nearly = width across optic lobes. Neck: Absent, anterior edge of collar dorsally at level of posterior margin of optic lobes. Funnel: Funnel cartilage not well d(~fined but with broad, shallow, straight depression; funnel adductors fused, without medial depression; adductor muscles arise from head at anterior end of statocyst. Mantle: Mantle wall thin; CML overlaps fins by 15% of FL. Fins: Length equals width (10.9 mm); anterior and posterior lobes lacking. Gladius: Extends posteriorly beyond fins by nearly IJ2 FL. Viscera: Gill base at 27% of the CML and 11% of ML. Digestive gland broadly rounded but spindle- shaped and oblique; nearly touches cephalic cartilage. Chromatophores. - Three series of chromatophores lie on the aboral surface of the tentacle and a few scattered chromatophores are found near the base on the oral surface. One chromatophore is present at the base of arm IV and one in the ventral midline of the head. One chromatophore lies on the aboral base of arm III. The dorsal surfaces of the head and mantle have scattered chromatophores and a patch of chromatophores lies at the ventral termination of the muscular mantle. Comments. -Site of .attachment of ventral buccal connectives cannot be determined. This squid cannot be placed in any known family at present. It is included in this paper because of its similarities (i.e., the position of the fins virtually posterior to the mantle and the unusual tentacle clubs) to the chiroteuthid lineage.

DISCUSSION Chun (1910) described several chiroteuthid paralarvae. One of these he named Doratopsis sagitta. It has several peculiar features: 1) The olfactory papilla arises at the base of the neck lateral to the funnel; 2) A transverse simple band of chromatophores, each transversely elongate, lies on the ventral surface of the head just anterior to the eyes; 3) Eyes are small, round; 4) Eyes do not project ventrally. YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 181

Figure 6. A-C: "big-fin," 19.1 mm ML; D-F: "Enoptroteuthis spinicauda" (=Lepidoteuthis grimaldii), 9.7 mm ML. A: ventral view. B: entire tentacle, sucker arrangement not exact. C: dorsal view. D: ventral view. E: tentacle club. F: dorsal view. Scale bars-I.O mm for whole views, 0.5 mm for tentacles.

All four of these features are characteristic ofsubadult Grimalditeuthis bonplandi; the first two are diagnostic of it. D. sagitta is the young of G. bonplandi and the former thereby becomes a junior synonym of the latter. Several features of Gri- malditeuthis have kept it from being included in the Chiroteuthidae: the fused funnel-mantle locking cartilages, subequal arms and lack of tentacles. The early paralarval stages, however, are nearly identical to those of Chiroteuthis with typical funnel-locking cartilages, elongate ventral arms and tentacles. In subadult Gri- malditeuthis bonplandi the tentacle is actually present, though invariably lost during capture, and it bears a club similar to that of Asperoteuthis except that suckers are absent from the distal half in addition to the proximal half (i.e., the tentacle lacks suckers; L. Burgess, pers. comm.). Grimalditeuthis bonplandi, therefore, belongs within the Chiroteuthidae. Chun (1910) described, in addition to D. sagitta, two other species of Doratopsis: 182 BULLETIN OF MARINE SCIENCE, VOL. 49, NO. 1-2, 1991

D. exopthalmica and D. lippula. In 1912 Pfeffer created Planktoteuthis as a new subgenus of Chiroteuthis and included these species along with his own Chiro- teuthis planctonica. He hesitated to include the latter species which was much smaller than the others. The distinctiveness of Chun's two species is apparent in the original descriptions which report the presence of an antitragus, aberrant armature of arms IV, projecting eyes and loss of most suckers of the tentacular stalks. Nesis (1982/87) recognized that Planktoteuthis and Valbyteuthis Joubin, 1931, which was originally described from subadult specimens, were synonyms but he did not apply the rule of priority. Since Planktoteuthis has clear priority it must be elevated to generic status and Valbyteuthis becomes its junior synonym. In 1929, Sasaki described a peculiar paralarva from Japan that he called Tankaia borealis. This species was listed as a subgenus of Chiroteuthis by Clarke (1966) although he suggested that it may be a young of Mastigoteuthis sp. Neither Voss (1977) nor Clarke and Trueman (1988) listed the genus in their classification of the Cephalopoda (presumably because they did not wish to recognize genera based on para1arvae). Nesis (1982/87), however, maintained it as a valid genus in the Chiroteuthidae. The most distinctive feature of this chiroteuthid is the presence of 6-8 sucker rows on the tentacular clubs and the short brachial pillar. As shown here, Chiroteuthis imperator paralarvae from Hawaii pass through a stage that exhibits these features. This same species is common off Japan (Kubota et aI., 1981). A second species, C. calyx is known from more northern Japanese waters (Okutani and Satake, 1978). Presumably, Tankaia borealis is the young of one of these species. Therefore, we place Tankaia in the synonomy of Chiroteuthis. In 1977 Lu described a peculiar chiroteuthid, Chiroteuthis acanthoderma, from the tropical North Pacific. Nesis (1980) incorrectly placed Lu's species in the synonomy of the Hawaiian species Chiroteuthis famelica Berry, 1909 (now M as- tigoteuthis famelica) and then elevated it to a new genus, Asperoteuthis. The confusion on the generic designation of Berry's species is due, in part, to Berry's description of a small specimen (39 mm ML) that had lost its tentacles during capture. Subadult specimens of Mastigoteuthisfamelica in our collections possess all the diagnostic features of the genus Mastigoteuthis. C. acanthoderma Lu, 1977 must be considered the type species of Asperoteuthis. This generic name, however, requires approval by the International Commission on Zoological Nomenclature. Chiroteuthoides hastula clearly is not a doratopsis. The presence of elongate arm IV and the retarded development of arm III indicate that it is the young of some species of Mastigoteuthis as suggested by Naef (1923). Indeed, Berry's illustration is quite similar to Figure 51 of Mastigoteuthis inermis from Hawaiian waters. Echinoteuthis is distinguished primarily by the possession of multicuspid tubercles on the body. Similar tubercles occur in the paralarvae of M. famelica which, as indicated above, is a typical member of the genus Mastigoteuthis. Other features ofJoubin's specimens (arrangement of suckers on arms IV and tentacles) are less distinctive and will probably fall within normal developmental patterns when these are better understood. Joubin originally included Berry's Chiroteuthis famelica (now Mastigoteuthis famelica) in his new genus. Tubercles are now known to occur in subadults of at least two additional species of Mastigoteuthis (M. cordiformis and Ai. hjorti; Roper and Lu, 1990). While future phylogenetic studies may show that tubercle-bearing and non-tubercle-bearing species of Mas- tigoteuthis fall into separate lineages, there is no evidence for making this as- sumption at present. Echinoteuthis, therefore, should be considered a junior synonym of Mastigoteuthis. Enoptroteuthis spin.icauda, also, does not belong in the Chiroteuthidae. Its primary similarity to this family is the termination of its mantle at the anterior YOUNG: CHIROTEUTHID PARALARVAE FROM HAWAIIAN WATERS 183 end of the fins. The only other squid to exhibit this feature outside the "chiroteuthid families" is Lepidoteuthis grimaldii Joubin, 1895. The advanced paralarva examined here has a very small, compact club with only a few suckers in two rows. This club is strikingly similar to that of young octopoteuthids. In this latter family the tentacles are lost during ontogeny. This similarity suggests that the tentacles of Enoptroteuthis spinicauda will also be lost. The only other squid besides the octopoteuthids and E. spinicauda to lose its tentacles during growth is Lepidoteuthis grimaldii. The youngest specimens of L. grimaldii described (60 mm ML) have reduced tentacles that are lost in the adult (Clarke, 1964). Although our paralarva lacks scales or tubercles characteristic of Lepidoteuthis grimaldii, slightly larger specimens in the U.S. National Museum of Natural History do have scales. As a result, there is little doubt that E. spinicauda belongs in the synonomy of Lepidoteuthis grimaldii. The family Chiroteuthidae, as defined here, is composed of the genera Chiro- teuthis, Planktoteuthis, Asperoteuthis and Grimalditeuthis. Other nominal genera in the family (i.e., Chiropsis, Bigelowia) are based on individuals belonging to valid species of Chiroteuthis (Roper and Young, unpublished). The family Chi- roteuthidae as now constituted is not easily defined on the basis of adult morphology. The most distinctive feature of the family is the unique presence of a doratopsis paralarva which occurs in all genera and presumably all species. The doratopsis stage is unlike the paralarval stage of any other cephalopod and is characterized by: 1) an elongate, chambered neck; 2) a chambered brachial pillar at some point in development; 3) a gladius that extends posteriorly beyond the fins and presumably supports a flotation device ("secondary fin" in some) and! or other "ornamentation" although this is usually lost during capture; 4) a posterior region of the mantle just anterior to the fins that contains vesicular tissue (absent in the youngest stages); 5) vesiculate arms in advanced stages; 6) greatly elongate ventral arms in advanced stages; and 7) tentacular clubs, in advanced stages, bounded over most of their lengths by parallel dorsal protective membranes and keels. The changes that occur at metamorphosis are poorly known and vary with the genus. Basically, in Chiroteuthis there is a shift in body proportions, loss of most of the "tail," loss of paralarval clubs (see below), increase in pigmentation and development of photophores. In Grimalditeuthis the tail is maintained, the funnel-mantle locking cartilages presumably fuse at this time, the relative lengths of the arms apparently changes and the paralarval club is lost. Changes in As- peroteuthis probably are similar to those of Chiroteuthis except that most photophores are absent and greater changes in relative arm lengths probably occur. Planktoteuthis shows the fewest changes and these mostly concern morphometrics.

PHYLOGENETIC RELATIONSHIPS A relationship between the Chiroteuthidae, Mastigoteuthidae and Joubiniteu- thidae has long been suspected (Naef, 1923) and to this grouping the Promach- oteuthidae and Batoteuthidae have been added more recently (Roper and Young, 1967; Young and Roper, 1968). We can now add to this assemblage "big-fin" described here. These "chiroteuthid families" have yet to be examined with a careful phylogenetic analysis and such an analysis is beyond the scope of the present paper. One feature, however, that all these families share in common that has not been clearly recognized is the apparent loss of the decapod tentacle club. One of the characteristics of the decapod club is the presence of a dorsal keel. None of the species in the chiroteuthid families possesses a club as an adult with a keel with the probable exception of Planktoteuthis. A keel is present, however, 184 BULLETINOFMARINESCIENCE,VOL.49, NO. 1-2. 1991 on the clubs of nearly all other species of decapods that have tentacles. (The exceptions are the small species of the Idiosepiidae and Pyroteuthidae.) The development of the club in Chiroteuthis provides a clue to what may have happened in the chiroteuthid :families.At about 8 mm ML in C. imperator the tentacle bears about 6 irregular rows of suckers distally prior to the development of the dora- topsid club. If evolution truncated the development of the tentacle at this point, a club with strong similarities to the mastigoteuthid-type club would result. Indeed, Clarke (1966) suspt:cted'that Tankaia borealis was a mastigoteuthid due to this stage oftentacle development. As the tentacle develops beyond this point, a club is formed with membranes and keel. Eventually the adult club forms on the tentacle stalk and the paralarval club is lost (presumably resorbed) (Naef, 1923; Roper and Young, 1967). This bizarre developmental sequence is unique within modem cephalopods. This sequence does not occur in all genera of chiroteuthids since the paralarval club is retained in adult members of Planktoteuthis. Presumably the chiroteuthid paralarval club represents the only true decapod club within the chiroteuthid families. Most likely, all other "clubs" are simply tentacle stalks covered with stalk suckers.

ACKNOWLEDGMENTS

I thank the numerous people that assisted in collecting and sorting the paralarvae, especially, R. Harman, K. Bigelow, N. Markel and D. Stevens. I also thank the officers and crews ofthe R/Vs KANA KEOKl,KILA and MOANAWAVEfor their able assistance. I am especially indebted to C. F. E. Roper for providing the opportunity to examine specimens in the collections of the National Museum of Natural History. This study is based on research supported in part by the National Science Foundation under grant number OCE 85-0064.

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DATEACCEPTED: December 3, 1990.

ADDRESS: Department of Oceanography. University of Hawaii. 1000 Pope Road. Honolulu. Hawaii 96822.