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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 97

The Systematics and Areal Distribution of Pelagic Cephalopods from the Seas off Southern California

Richard Edward Young

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972

A B S T R A C T

Young, Richard Edward. The Systematics and Areal Distribution of Pelagic Cephalopods from the Seas off Southern California. Smithsonian Contributions to Zoology, number 97, 159 pages, 15 figures, 38 plates, 1972.—The mid-water cephalopods from off the coast of southern California and adjacent areas are described and their areal distribution is discussed. Forty-two species of pelagic cephalopods are now known from this area including ten new species. Off southern California the fauna is part of the transitional and subarctic fauna to the north, while primarily tropical species are found off northern Baja California.

Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution’s annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE WASHINGTON : 1972

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Contents

Introduction

Material and Methods

Acknowledgments

Description of Study Area

Zoogeography

Material Examined

Key to the Species Described in this Report

Systematics

Records of Additional Pelagic Species from off Southern California

Checklist of Cephalopods Definitely Known to Occur in Southern Californian or Neighboring Waters

Summary

Systematics

Distribution

Literature Cited

Appendix 1

Appendix 2

Plates

Publication in Smithsonian Contributions to Zoology

The Systematics and Areal Distribution of Pelagic Cephalopods from the Seas off Southern California

Richard Edward Young

____________

Richard Edward Young, Department of Oceanography, University of Hawaii, Honolulu, Hawaii 96822.

Introduction

The pelagic cephalopod fauna of the eastern North Pacific is poorly known. The single major paper treating cephalopod systematics of this area (Berry 1912b) dealt with only a small portion of the pelagic species. Material for this study was collected by the University of Southern California over a six-year span from the fall of 1960 to the fall of 1966. During this period, 445 mid-water tows were taken with an Isaacs-Kidd mid-water trawl. This represents an estimated 911 hours of “fishing” at discrete depths from the surface to about 1200 meters. This program yielded over 5,000 specimens of pelagic cephalopods belonging to 40 species of which 33 are described in this report.

The present paper attempts to delineate the pelagic cephalopod fauna occurring in the waters off southern California and to discuss the zoogeographic relationships of this fauna. Subsequent papers will treat the vertical and seasonal distribution of the most abundant species.

Materials and Methods

Specimens were collected with an Isaacs-Kidd mid-water trawl with a mouth 10 feet in diameter. The net was operated from the University of Southern California’s R/V VELERO IV. Comparative material was available from the large collections gathered by G. L. Voss at the University of Miami’s Institute of Marine and Atmospheric Sciences. The University of Southern California trawling program initially studied the waters over the most accessible inshore basin, the San Pedro Basin, which lies between the mainland and Santa Catalina Island. The program subsequently was extended to include the area of the Santa Caltalina Basin to the west of Santa Catalina Island. The most intensive sampling (over 200 stations) has occurred in this area. While never completely abandoning these two basins, the program was extended to include San Nicolas Basin, San Clemente Basin, the waters just west of the continental slope, and the waters to the south as far as the vicinity of Guadalupe Island. The total area studied is well within a region extending from 28°N to 34°N. While the region sampled is rather small, it lies within a transitional faunal region and, as a result, distinct differences were noted in the species composition between the northern and southern ends of this area. To demonstrate such geographic differences, I have divided the area of study into seven zones (Figure 2) so that each includes a major area of sampling.

In this paper, the following abbreviations are used:

Sta. no. number of station where specimen was captured

M.L. mantle length

M.W. mantle width

P.L. pen length

P.W. greatest width of pen

F.L. fin length

H.W. head width

H.L. head length measured from the anterior end of the nuchal cartilage of the base of arms I

Tail L. tail length

Arm L. I length of arm I measured from the arm base, between arms I (where the muscular portion of the arms join) to the arm tip

Arm L. II length of arm II measured from the arm base between arms II and III to the arm tip

Arm L. III length of arm III measured from the arm base between arms II and III to the arm tip

Arm L. IV length of arm IV measured from the arm base between arms IV to the arm tip

A.C. I–IV arm count (Total number of suckers and/or hooks on the indicated arm)

½ A.C. I–IV total number of suckers and/or hooks on the proximal 50 percent of the arm length of the indicated arm measured from the basal sucker

A.C. Dist. S. I–IV number of suckers distal to arm hooks on the indicated arm

A.C. Prox. S. I–IV number of suckers proximal to arm hooks on the indicated arm

A.C. Hooks I–IV number of hooks on the indicated arm

Sucker D. diameter of largest arm sucker + accurate count or measurement greater than given figure due to damaged arm.

Club L. length of tentacular club

Club H. number of hooks on club

Club H. F. club hook formula (number of hooks in the dorsal medial row over the number in the ventral medial row)

Club S. number of suckers on club

Club S. D. diameter (long axis) of largest club sucker

Carp. S. number of suckers in carpal cluster

Carp. K. number of knobs in carpal cluster

Carp. F. carpal formula (number of carpal knobs over the number of carpal suckers)

Dact. S. number of suckers on the dactylus of the club

Dact. T.S. number of suckers in the sucker pad at the tip of the dactylus

Tent. L. tentacle length

Stalk S. number of suckers on the tentacular stalk

D.S.S. number of suckers in the dorsal row on the tentacular stalk

V.S.S. number of suckers in the ventral row on the tentacular stalk

M.S.S. number of suckers in median portion of tentacular stalk

R. right

L. left

Tub. L. length of cartilagenous strip bearing tubercules

Tubercles number of cartilagenous tubercules at funnel-mantle fusion

Gill L. gill length

G.C. gill count (total number of primary gill lamellae)

Gill W. gill width

Lam. L. length of longest lamella

Eye D. eye diameter

Egg D. egg diameter

Ov. G. Diam. diameter of oviducal gland.

FIGURE 1.—Localities of trawling stations. Dots correspond to the midpoints of the trawling transects.

FIGURE 2.—Breakdown of area studied into zones corresponding to major areas of sampling.

Unless otherwise indicated in the tables, measurements and counts of structures on the right and left sides have been averaged to give single figures. Measurements are in mm and have been rounded to the nearest whole number in most cases.

Acknowledgments

I am indebted to many people who assisted me in the present work. J. S. Savage, J. S. Garth, R. Lavenberg, D. Bright, and R. Given associated with the University of Southern California provided specimens and data. F. Hochberg kindly supplied specimens taken by the R/V SWAN. G. Y. Hendrix of the University of Miami provided a considerable amount of help in sorting and caring for the collection and in making preliminary identifications during the early stages of this work. Diane McGinnis helped with sorting when time was becoming a critical factor.

Drs. Garth and Savage also provided funds for a trip to the University of Southern California during the course of the study. I am also indebted to the personnel of the R/V VELERO IV and the students and staff at the University of Southern California that participated in the cruises.

This study was accomplished under the supervision of G. L. Voss, whom I thank for his assistance in innumerable ways and for his support, encouragement, and help with the manuscript. The comparative material, which was invaluable to this study, was drawn from the large collections gathered by Dr. Voss at the University of Miami’s Institute of Marine and Atmospheric Sciences.

I am grateful to D. DeSylva and R. Hurly for reading the manuscript and offering valuable suggestions.

I am also indebted to C. F. E. Roper for supplying type material and much valuable information concerning specimens in the collections of the National Museum of Natural History. Dr. Roper has added considerably to the present work through numerous discussions on systematic and other biological problems. E. S. McSweeny also provided aid through discussions of many of the problems encountered.

The great majority of the illustrations were done by Mrs. Constance McSweeny. Mrs. McSweeny, a biologist as well as an excellent artist, on many occasions detected important systematic characters that had previously gone unnoticed. I thank Mrs. McSweeny for her excellent illustrations and also for her help with the systematic aspects of the paper. Mrs. McSweeny deserves double thanks for her contribution. After all the plates were completed, they were destroyed in a fire at Miami and had to be redone.

During the course of this study, a number of visits were made to Redlands, California where S. Stillman Berry and I were able to have long discussions on many aspects of cephalopod biology. I thank Dr. Berry for his interest and enthusiasm.

Finally, I wish to thank my wife, Shelia, for assisting with the typing, compiling data, and for supplying ecouragement.

This research was partially supported by grants G–10691 and G–23467 from the National Science Foundation to the University of Southern California, and by grants GM–5729X, GA–0709, and GA–1493 from the National Science Foundation to the Univrsity of Miami, and by the National Geographic Society–University of Miami Deep-Sea Biology Program.

This work was done in partial fulfillment of the Ph.D. degree at the Institute of Marine and Atmospheric Sciences of the University of Miami.

Description of Study Area

The ocean floor off southern California between the shoreline and the continental slope, a distance of 50 to 160 miles, is unusually complex. Shepard and Emery (1941) designated this region the continental borderland to distinguish it from typical continental shelves. This area contains a dozen complete basins ranging from about 500 to 2,100 meters in depth, several troughs, numerous banks, and eight islands. At the western edge of the borderland, the continental slope drops rapidly to a depth of about 3,500 meters.

The waters over the continental borderland are derived from the complex interaction of several water masses. The California Current is the eastern component of the great anticyclonic circulation of the North Pacific and, as such, it is a continuation of the North Pacific Drift (sensu lato). The waters of the northern portion of the California Current are derived largely from the Subarctic water. As this cold current extends southward, it warms through insolation and by increased mixing with waters from Central water to the west. At about 25° N, it begins to turn westward and eventually merges into the North Equatorial Current. Off southern California a nearly permanent eddy is present, creating north-flowing currents between San Clemente and Santa Catalina islands.

The surface waters over the borderland are modified to some extent by upwelling. Prevailing winds along the southern California coast, which are from the north and northwest, are strongest in May and June. Surface waters, under the influence of these winds and the earth’s rotation, are driven offshore and replaced by nutrient-rich waters which upwell from below to a maximum depth of 200 to 300 meters. Entrainment of surface water by the California Current over the shallow Santa Rosa-Cortes Ridge may also play a role in the upwelling.

During the winter an inshore surface countercurrent is present along most of the coast from the southern tip of Baja California to at least 45° N. This current seems to be somewhat variable off Baja California and off southern California, but is well defined north of Point Conception where it is known as the Davidson Current.

Below 150 to 200 meters a deep countercurrent of Equatorial Pacific Water flows northward along the coast to at least 40° N. and progressively mixes with the lower layers of the California Current. Off southern California the major flow of the deep countercurrent, which has become transitional in composition at this latitude, occurs just seaward of the continental slope, although portions flow over the borderlands apparently filling the basins and forming the water of the intermediate depths.

Thus, over the borderland, the influence of the California Current, the surface countercurrent, and upwelling predominate in about the upper 200 meters while below 200 to 300 meters the influence of the southern water predominates. (The above discussion is based primarily on Reid, Roden, and Wyllie 1958, and Emery 1960.)

Surface temperature ranges are shown in Figures 3 and 4.

FIGURE 3.—Extremes of temperature regression curves in the study area based on data taken from 1950 to 1962. (Reproduced from R. J. Lynn, 1967.)

Zoogeography

The present study area lies within an oceanographic and faunistic transitional region. The faunal changes that occur within this area have been demonstrated by a number of authors. The faunal boundries seem to apply equally well to the littoral (Ekman 1953), deep benthic (Parker 1964), epipelagic (McGowan 1968), and mid-water (Brinton 1962, Lavenberg and Ebeling 1967) faunas.

Brinton (1962) gave an excellent account of the distribution of euphausiids in the northeastern Pacific. His chart showing the faunal relationships of euphausiids in this area for a 2-month period (August–September) is reproduced here (Figure 5). A more generalized chart (Figure 6) is given by McGowan (1968) based on the distribution of pteropods and heteropods.

From the limited information available, it appears that cephalopods in this area exhibit general faunal distributions similar to the euphausiids, pteropods, and heteropods. Many cephalopod species during the period studied were absent or only sparcely represented in the southern zones, but were abundant in the northern zones; the reverse trend was also apparent. In many cases species can be confidently related to the established faunal assemblages of the North Pacific, while for other species only indications of their faunal relationships can be determined.

FIGURE 4.—Extremes of temperature regression curves in the study area based on data taken from 1950 to 1962. (Reproduced from R. J. Lynn, 1967.)

Subarctic and Transitional species. Due to our present lack of information, it is difficult in many cases to determine whether a species occurs primarily in Subarctic, Transitional, or Subarctic and Transitional waters. Therefore, the placement of certain species within one category or the other must be considered tentative.

Moroteuthis robusta appears to be the species most closely associated with Subarctic water which has been reported off southern California. It has only rarely been collected from these waters (Phillips 1961). It is known from off northern California (Phillips 1966), Oregon (van Hyning and Magill 1964), Canada, the Aleutian Islands, and northern Japan (Clarke 1966).

Two other species, Berryteuthis magister and B. anonychus, probably are distributed primarily in Subarctic waters and are not known from California. Berryteuthis magister (Berry 1913) has been recorded from Japan, Kuril region, coastal waters of the Aleutian Islands, the Gulf of Alaska, Puget Sound, and off Oregon (Clarke 1966). Berryteuthis anonychus (Pearcy and Voss 1963) is known only from the coast of Oregon.

Gonatopsis borealis is also known from Subarctic waters, but is more abundant in the southern California waters than Moroteuthis robusta and has a southern limit of distribution off northern Baja California. It has been recorded from California, Oregon, the Bering Sea, Kamchatka, and nothern Japan.

Material Examined

Onychoteuthis borealijaponicus is known from off California, Oregon, Canada, and northern Japan. Probable records also occur in the Bering Sea.

If all northern Pacific Galiteuthis (except G. pacifica) belong to G. phyllura, then this species would be known from the Kurile Islands and the Bering Sea as well as off California and Oregon; it seems likely that this will prove to be the case.

Loligo opalescens may extend as far south as the tip of Baja California and extends north to the southern end of British Columbia (Fields 1965).

A number of additional species are believed to have the southern limit of their distribution off northern Baja California and are known to extend north to the middle Oregon coast. These are: Gonatus onyx, Chiroteuthis calyx, Octopoteuthis deletron, and Abraliopsis felis. It seems likely that these four species will prove to be distributed primarily in Transitional waters, although Abraliopsis felis has tentatively been recorded off Hawaii. Octopoteuthis deletron is also known from off northern Peru and may be an antitropical species.

Histioteuthis heteropsis is known from the southern California area north to Monterey Bay. Its southern limit is thought to lie off northern Baja California. It has also been reported off Chile and is also considered to be antitropical in distribution.

FIGURE 5.—Faunistic zones of the northeastern Pacific based on the distribution of euphausiids. (Reproduced from Brinton, 1962).

Other species which, at present, are thought to have southern limits off northern Baja California and are known only from Transitional waters are Gonatus berryi and Gonatus pyros. Gonatus californiensis extends somewhat farther south along Baja California, and there is a single record of a larva from the Bay of Panama that apparently belongs to this species. This is, however, considered tentatively to be outside the normal range for this species.

Central species. Two species, Leachia dislocata and Abraliopsis falco, show definite preference for the southern zones of the study area and have also been found off Hawaii and therefore probably belong to the Central Water Mass fauna.

FIGURE 6.—Faunistic zones off California and Baja California based primarily on the distribution of pteropods and heteropods. (Reproduced from McGowan, 1968.)

Equatorial species of the eastern Pacific. Two species, Galiteuthis pacifica and Valbyteuthis danae, are know from the eastern Pacific off Panama where they are thought to be common. Only a few specimens have been taken in the southern California area.

Occasionally, Dosidicus gigas is found off southern California. A major invasion of this species took place from 1934 to 1937. This squid first became abundant in 1935 when 12 tons were landed at San Pedro Harbor during August and September. The squid varied from 2 to 2½ feet in length. Many individuals were taken north of Monterey Bay during the fall and winter of 1935 and January of 1936. Most of these specimens were between 3 and 4 feet long. In January 1936, 400 pounds were taken in a purse seine. The individuals averaged close to 4 feet in length (Clark and Phillips 1936). Croker (1937) states that in 1934 and 1935 specimens weighed about 5 pounds, while those caught in the early part of 1937 were averaging over 20 pounds.

This information indicates that D. gigas was not breeding in the southern California waters but invaded, grew, and disappeared after 1937. Apparently this species has not been present in any numbers since 1937. Young D. gigas are common in the Bay of Panama (personal observation) while only large specimens are known from the coast of Chile, and these disappear during part of the year (P. Garcia-Tello, personal communication). It seems likely that this species is an equatorial form which migrates away from its spawning grounds into cold water to feed. If this is so, it is strange that this species does not occur more commonly off southern California.

Argonauta pacifica has been reported from the Galápagos Islands to Monterey Bay (Berry 1912b). Its occurrence off southern California seems to be sporadic. On occasion, it has been reported off southern California in considerable numbers (Berry 1912b).

Cosmopolitan tropical and subtropical species. Species known from California that have been recorded as having a cosmopolitan distribution in tropical and subtropical waters are: Pterygioteuthis gemmata, P. giardi, Ommastrephes bartramii, Alloposus mollis, Ocythoe tuberculata, Histioteuthis dofleini. Two additional species, Cranchia scabra and Vampyroteuthis infernalis, have a similar distribution but also occur in somewhat higher latitudes. Symplectoteuthis luminosa appears to be broadly distributed within warm waters of the Pacific Ocean only.

Species with insufficient distributional data. Mastigoteuthis pyrodes is found in greater abundance within the southern zones of the study area, but is not known from either the waters off Panama or off Hawaii. Valbyteuthis obligobessa and Bathyteuthis berryi are known only from the present trawling program and show no preference for the southern or northern latitudes within the study area. Pyroteuthis addolux also exhibits no latitudinal preference in the study area, but is clearly more abundant in the more oceanic zones. This species is also known from as far west as longitude 147° W. Systematic problems or insufficient data prevent discussion of the zoogeography of the following species: Neoteuthis sp., Japatella diaphana, Japetella sp., Grimalditeuthis bomplandii, and Heliocranchia pfefferi.

TABLE 1.—Distribution frequencies (specimens per hour of trawling) of most abundant species. Frequencies are corrected for estimated depth of capture and for contamination resulting from the use of open-net trawls.

Inshore and offshore species. Some species that occur along the coast show preference for inshore waters over the continental borderland, while others show a preference for the more offshore waters seaward of the continental slope. In the present study, Gonatus onyx, the most abundant species taken, seemed to prefer near-shore waters as do G. californiensis and G. pryos, while Onychoteuthis borealijaponicus and Pyroteuthis addolux showed preference for offshore waters. McGowan and Okutani (1968) found that larvae of Abraliopsis felis were taken most frequently in offshore waters. A similar trend was found in the present study in specimens 20 mm M.L. or less, but not in specimens larger than 20 mm. Valbyteuthis obligobessa is also more abundant in the offshore zones; this appears to be correlated with its deep-water habitat.

Faunal affinities. It is apparent that in the northern zones off Los Angeles the species with northern affinities (i.e., Transitional and Subarctic species) predominated during the study period from 1960–1966. The 14 more abundant species (disregarding Helicocranchia and Japetella due to systematic complications) taken in this survey give a total frequency of capture (specimens/hour) of 9.41 in zone 2 (Catalina Basin—the most heavily sampled area), of which 8.59 specimens per hour represent species with northern affinities. In zone 7 (the most southernly zone) these same 14 species have a total catch frequency of 3.92 specimens per hour, of which 3.08 specimens per hour represent species with southern affinities (i.e., species of tropical waters including the North Pacific Central waters). These figures also suggest a greater abundance of individuals in zone 2 compared to the more oceanic zone 7.

The predominant affinity with a more northern fauna can be further emphasized by the total counts of individuals. Almost 80% of the total specimens taken in the survey belong to species with northern affinities, while only slightly more than 11% of the total specimens belong to species with southern affiinities. (The remaining 9% represent species whose affinities could not be determined for various reasons.)

Of the 42 species of pelagic cephalopods recorded off southern California and nearby waters 8 are unidentified at the specific level as a result of their damaged condition, larval state, or merely insufficient numbers. Of the remaining 36 species, 10 also occur in the Atlantic Ocean. Available material was sufficient to allow a detailed comparison between Atlantic and Pacific populations of only four of these species: Pterygioteuthis gemmata, Ommastrephes bartramii, Cranchia scabra, and Vampyroteuthis infernalis. All four showed distinct differences between the two oceans. The differences in local specimens of P. gemmata are probably a result of geographical variation. The differences in the local specimens of O. bartramii, C. scabra, and V. infernalis are presently interpreted as resulting from geographical variation, but this will require confirmation. There was abundant material of a fifth species, Heliocranchia pfefferi, available, but comparative material was inadequate; however, a slight difference in eye size was noted between specimens from the different oceans. It is felt that mature specimens as well as more adequate comparative material must be examined before the identification of this species can be confirmed.

Although direct comparisons were made, material of Alloposus mollis, Ocythoe, tuberculata, and Grimalditeuthis bomplandii was insufficient to allow a careful evaluation of variations in specimens from different oceans.

Comparisons of populations of Histioteuthis dofleini have been made by N. Voss. She informs me that this widely distributed species shows considerable geographical variation.

I have not examined specimens of the final cosmopolitan species, Pterygioteuthis giardi, which was recently reported from southern Californian waters by McGowan.

It should be emphasized that all ten of these presumed cosmopolitan species are primarily distributed in tropical and subtropical waters and in those species that have been carefully examined, Pacific specimens exhibit distinct differences from those of the Atlantic Ocean.

Of the 26 species known only from the Pacific Ocean, four, Onychoteuthis borealijaponicus, Moroteuthis robusta, Gonatopsis borealis, and Symplectoteuthis luminosa, are know from the waters around Japan. It is possible, however, that Octopoteuthis deletron and Galiteuthis phyllura are identical to similar described forms from Japan (i.e., Octopodoteuthis sicula and Galiteuthis armata).

Only three species, Leachia dislocata, Abraliopsis falco, and Abraliopsis felis, of the 26 species restricted to the Pacific Ocean have been found off Hawaii, and it seems likely that the juvenile A. felis (if this is a correct identification) seen from these waters are outside their area of primary abundance.

Key to the Species Described in this Report

SYSTEMATICS

Class CEPHALOPODA

Order TEUTHOIDEA

Family ENOPLOTEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Arms with biserial armature (occasionally modified at arm tips) consisting both of hooks and suckers. Tetraserial armature on the tentacular clubs, although marginal rows may be lost with growth in some species. Hooks usually present on clubs. Eight buccal lappets; buccal connectives attach to the dorsal margins of arms IV. Photophores present.

Subfamily ENOPLOTEUTHINAE Pfeffer, 1912

DIAGNOSIS.—Nidamental glands absent. Numerous small photophores present on the surface of the mantle, head, and arms, but not in the tentacles or on the viscera. Posterior margins of fins concave.

Genus Abraliopsis Joubin, 1896

DIAGNOSIS.—Arms IV with 2–4 very large photophores at the arm tips. Tentacular clubs with 2 rows of hooks on manus.

TYPE SPECIES.—Enoploteuthis hoylei Pfeffer, 1884.

Abraliopsis falco, new species

PLATES 1A, B; 2A–G

DESCRIPTION.—The mantle is cylindrical anteriorly and tapers to a blunt point posteriorly. The mantle musculature terminates at the end of the pen which is in advance of the posterior tip of the body. This broad posterior tip appears to be composed of vacuolated connective tissue which continues laterally, for a short distance, up the sides of the muscular portion of the mantle. The anterior margin of the mantle projects slightly in the areas of the dorsal and ventrolateral cartilages. The mantle is moderately thick and muscular.

The large, muscular fins are terminal; they are about 70–80% of the mantle in length and about 90–100% in width. Their lateral apex forms a rounded point. The posterior margin is drawn-out along the mantle to its apex. Anterior lobes are free.

The funnel is short and muscular. Each funnel locking-cartilage has a straight groove which opposes a straight ridge on the mantle. The dorsal pad of the funnel organ has an inverted V-shape. A large ridge extends diagonally across each lateral limb of the organ and an anterior papilla is present. The ventral pads are oval. A funnel valve is present. The funnel is free laterally.

The head bears large eyes that occupy its entire lateral portions. A large sinus is present in the anterior margin of each eyelid, and distinct dorsal and ventral windows lie above and below each eye. A nuchal crest with 4 folds on either side of the head is present; the ventral fold on each side is thick and partially embrances the funnel; the second fold from the funnel bears the “olfactory” organ; the dorsalmost 2 folds join posteriorly on each side forming crescent-shape ridges.

The arms are large and muscular; they are considerably longer in the male. The arm formula is: 4=2>3>1. Arms I–III all have well developed aboral keels and arms IV have well-developed lateral keels. Protective membranes are nearly absent proximally on the dorsal borders of arms I where they are represented primarily by isolated trabeculae; the membranes are very low distally. The dorsal borders of arms II have a few trabeculae proximally, but otherwise lack trabeculae and protective membranes except near the sucker-bearing tip where a trabeculate protective membrane is present. The dorsal borders of arms III show only a trace of a few proximal trabeculae except for a small trabeculate protective membrane near the distal, sucker-bearing tip. The dorsal borders of arms IV have flat, rounded trabeculae with low membranes in females. Arms I–III all have large, trabeculate, protective membranes on their ventral borders. These are present, but not as well developed on arms IV in females.

Each arm I–III possesses a double series of hooks which is replaced by a double row of suckers at the arm tip. The number of distal suckers is difficult to count with certainty. One female had about 34 on each arm I, 23 on each arm II, and 15 on each arm III. The most proximal suckers in this series have broad outer rings; their inner rings have about 10–12 broad teeth which are rounded to truncate on the distal margin. The more distal suckers are globular; they lack outer rings and have reduced apertures without teeth. All suckers are globular and lack outer rings on the dorsal pair of arms in males. Arms IV possess hooks, but lack distal suckers.

The arms in male specimens are longer and have more hooks than do the arms of females. The increased number of hooks in males results from both the greater length of the arms and the smaller size of the hooks. The following chart comparing averages for 4 adult females and 5 adult males demonstrates these relationships.

Both arms IV are hectocotylized in the males; however, the left arm is much more highly modified. The trabeculate protective membrane is greatly enlarged on the ventral margin of this arm and is heavily pigmented with dark brownish purple chromatophores. Small conical knobs line the oral margins of the large trabeculae. On the dorsal margin, trabeculae are very prominent proximally but diminish in size distally, while the protective membrane is low and nearly absent proximally; these trabeculae are also lined with conical papillae on their oral margins. The proximal 3–4 hooks on the ventral margin are much larger than the proximal 2–3 hooks on the dorsal margin. The fourth or fifth hook on the ventral margin and the third or fourth hook on the dorsal margin are comparable in size. Distally the hooks on the dorsal margin increase in size, while those of the ventral margin abruptly decrease. Shortly, the hooks of the ventral margin begin to increase in size, while those of the dorsal margin begin a gradual decrease. Approximately the fifteenth hook of the ventral margin approaches in size the adjacent hooks of the dorsal margin. The hooks of the ventral margin then decrease more rapidly in size than those of the dorsal margin up to the end of the series. The fleshy bases from which the hooks arise in the dorsal row bear small conical papillae; there are also a few papillae on some of the hook bases of the ventral row. The oral face of the arm proximal to the first hook also possesses numerous papillae.

The right arm IV shows a lesser degree of modification. On the ventral border, from the second hook to about the middle of the arm, are 4–5 enlarged trabeculae, bearing conical papillae, with a low membrane between them; trabeculae are lacking on the distal half of the arm. Along the distal third of the ventral border, there are two semilunar flaps. The dorsal border has trabeculae that are less prominent, but also bear papillae and have very low protective membranes. A small flap, present distally on the dorsal border, opposes the distal flap on the ventral border. The proximal 3–4 hooks in the dorsal row are considerably smaller than the first 4–5 in the ventral row. Distally the hooks in the ventral row decrease abruptly in size, while those of the dorsal row increase abruptly in size so that the fifth or sixth hook of the ventral margin is much smaller than the fifth one of the dorsal margin. Each series terminates at the proximal end of the distal flaps on each margin. The tips of the ventral arms bear large photophores identical to those of the female. Both modified arms IV are about the same length.

In the male, arms I–III show a slight amount of dimorphism. The trabeculae on the dorsal borders of arms I are considerably larger than in the females. They lack a protective membrane proximally, but one is well developed distally. The oral margins of the trabeculae on both dorsal and ventral borders are lined by conical papillae. In males, papillae are also present to some extent on the trabeculae of arms III and to a lesser degree on arms II. All arms in males generally have 5–20 papillae proximal to the first pair of hooks. In males, the proximal hook on the ventral margin of each arm III is greatly enlarged over the succeeding hook and is approximately equal to (or larger than) the proximal hook of the dorsal margin. The second through fourth hooks of the ventral margin are distinctly smaller than those opposing them on the dorsal margin; farther distally, they are approximately equal in size. Hooks on arms I–III are smaller in males than in females.

The tentacles are short and robust. The tentacular club is short and not expanded. The manus bears hooks in 2 rows; those of the ventral row are about twice the height of those in the dorsal row in females, while the difference may be less in males. Hooks of the dorsal row vary from 3–4, while the smaller hooks of the ventral row vary from 2–4. Suckers alternate with the hooks in the ventral row. Only the proximal sucker is present in the dorsal row, and it is occasionally absent. The dactylus has 4 rows of suckers that extend to a circle of suckers at the tip of the tentacle. A random count of 8 clubs shows a variation in dactyl suckers between 48 and 60. A distinct carpal cluster is present which possesses 4 or occasionally 5 suckers and pads. No membrane or flap lies adjacent to the carpus or the manus. A low, thick protective membrane lines the ventral margin of the dactylus. There is no trace of a keel.

Numerous photophores of varying sizes are embedded in the mantle in a rather irregular arrangement, although some indication of a pattern is discernible. In the ventral midline, there is a narrow strip that is generally free of photophores; however, this strip is sometimes difficult to detect and often does not extend to the anterior margin or to the posterior end of the mantle. On either side of this strip is a diffuse longitudinal band of photophores. Lateral to these bands, the number of photophores gradually diminishes to only a few scattered organs situated dorsally. Superimposed on this pattern is a single line of photophores that extends from the anterior mantle margin at the projection of the mantle locking-cartilage, posteriorly for about half of the mantle length. Between this series and the midventral strip is another single line of photophores that can be traced about half of the mantle length from the anterior margin. Both series are not always readily apparent.

The funnel has 6 separate groups of photophores, none of which extends to the exhalant opening. One group is located on either side of a clear median strip on the ventral surface of the funnel. A second group is found along each lateral margin of the funnel, and a third group is located on either side of the dorsal surface of the funnel and extends onto the bases of the bridles.

A triangular patch of photophores is located on the ventral surface of the head dorsal to the funnel. This patch continues forward over the ventral surface of the head in the midline as a narrow, well-defined strip to the junction of arms IV where the series divides into a row that runs along the inner margin of each arm IV for about half of the arm length. A second narrow strip on each side of the head runs anteriorly from the first nuchal fold, passes medial to the ventral “window” of the eye, and continues onto the lateral margin of arm IV at the base of the lateral keel. This series continues to the tip of the arm, but is broken at 4–5 spots by short areas that lack photophores. Between these first 2 bands, near the base of the ventral arms, a few scattered photophores are occasionally present.

A third strip extends anteriorly from each second nuchal fold, passes on either side of the head lateral to the ventral “window” and joins an area of scattered photophores anterior to each “window.” Each patch continues onto the margin of the lateral keel of each arm IV as a single row of photophores which extends along the proximal third of the arm. A final series extends around the margin of each eyelid. In the 3 males which allowed accurate counts, the number of photophores in this series varied from 48–54, while in 4 females the number varied from 52–63. This series extends anteriorly on either side onto arm III at the base of the aboral keel and continues the entire length of the arm except for a few breaks in the series in the distal half of the arm.

On the ventral surface of each eye is a series of 5 light organs of which the first and last members are considerably larger than the middle 3.

Distally the tips of arms IV bear 3 large, dark-purple photophores and 2–3 small photophores.

Within the mantle cavity of females on the inner surface of the dorsal mantle wall, immediately anterior to the stellate ganglion, is a highly folded, apparently glandular, pocket. The pleats of this pocket also extend beneath the collar in the nuchal region. Several sperm reservoirs (discharged spermatophores) were found attached within the pocket. The pocket has a dark-purple pigmentation which can be seen through the dorsal surface of the mantle beginning about one-eighth of the mantle length from the anterior end of the pen.

The buccal membrane has 8 supports and bears numerous long fleshy papillae on its brownish purple oral surface. The buccal membrane connectives attach to the dorsal borders of arms I, II, and IV, and to the ventral borders of arms III.

The specimens are covered with numerous small, reddish brown chromatophores.

TYPE LOCALITY.—28°54′ N, 118°08′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod Type No. 1.

DISCUSSION.—The genus Abraliopsis has been in an extremely confused state due to many inadequate early descriptions, often based on larvae, followed by numerous misinterpretations by subsequent authors. Fortunately, the genus has been monographed by Voss (in manuscript) and most of the problems have been resolved.

The genus Abraliopsis is usually divided into two subgenera: Abraliopsis, with a diffuse pattern of photophores on the ventral surface of the head, and Micrabralia, with photophores on the ventral surface of the head arranged in distinct rows (Voss 1967). A. falco clearly falls into Micrabralia which, according to Voss (in manuscript), contains only two other named species: Abraliopsis affinis (Pfeffer 1912) and Abraliopsis gilchristi (Robson 1924). Pfeffer based A. affinis on Hoyle’s (1904) description of 4 specimens taken from the eastern tropical Pacific between 2°34′ N, 82°29′ W and 14°46′ N, 9840′ W. The specimens were only partially described; however, specimens of A. affinis in the collections of the Institute of Marine and Atmospheric Sciences which were captured at 7°19′ N, 79°44′ W have allowed adequate comparisons to be made with A. falco.

Abraliopsis falco differs most distinctly from A. affinis on 2 characters: A. affinis has a distinct membrane on the ventral margin of the tentacle that extends from the proximal end of the carpus to the base of the manus; no such membrane is present in A. falco. A. affinis has a large keel on the club that extends from the tip of the dactylus to about the midpoint of the manus; a keel is lacking in A. falco. The specific value of these features within the genus has been emphasized by Voss (in manuscript). Several other characters are also of value in separating these species. In A. falco, the first hook in the ventral row of each arm III in males is usually equal to or larger than the first hook in the dorsal row of the same arm (this feature was valid in 9 out of 10 cases). In A. affinis the first hook in this same position in males is distinctly smaller than the first hook in the dorsal row (this feature was valid in 35 out of 36 cases). The midventral strip on the mantle in A. affinis which lacks photophores is very broad and generally expands at its posterior termination less than two-thirds of the mantle length from the anterior end. In A. falco, this strip is very narrow throughout and seldom reaches either the anterior or posterior end of the mantle. The arms are considerably longer in A. affinis, but further study on more material is necessary to quantify this feature. There are a number of other features that seem to show significant differences such as the relative sizes of the club hooks and the number of proximal arm papillae; however, the characters listed are sufficient to separate the species.

TABLE 2.—Measurements in mm of Abraliopsis falco, new species.

Since Abraliopsis falco appears to be a warm-water species, it is possible that the ranges of A. falco and A. affinis overlap. Although the original description of A. affinis is incomplete, there can be no question of the true identity of this species since a club was illustrated by Hoyle (1904, plate 8, figure 1) that clearly shows the presence of a keel and another illustration (plate 10, figure 1) shows the midventral clear strip on the mantle with the appearance characteristic of specimens discussed in this paper as A. affinis.

Abraliopsis gilchristi was described by Robson (1924) from specimens taken off South Africa and has recently been redescribed by Voss (1967). The most obvious feature that separates this species from A. falco is the persence of 2 distinctly separate rows of photophores on the midventral surface of the head in A. gilchristi, each of which is continuous with the medial series of the nearest arm IV. In A. falco, there is only a single series in the midventral line of the head. There are many other features (including the presence of a semicircular membrane on the ventral border of the tentacle between the carpus and manus in A. gilchristi) which separates these 2 species, but need not be discussed here.

There is another but unnamed species in the Atlantic Ocean which is treated fully by Voss (in manuscript), and a comparison between this species and A. falco will be made in this paper. For the present, it is only necessary to state that they are distinct.

DISTRIBUTION.—The 10 specimens of A. falco taken by the VELERO all came from zone 7. An additional specimen (M.L., 20 mm) was taken by the SWAN near Hawaii at approximately 24° N and 151° W. These data indicate that this species is a warm-water form and probably a mid-oceanic species.

Abraliopsis felis McGowan and Okutani, 1968

PLATES 1C–D; 2H–N

Abraliopsis sp., Pearcy, 1965, p.261.

Abraliopsis felis McGowan and Okutani, 1968, p.72.

DESCRIPTION.—The mantle is long and slender; its width is about one-fourth of its length. It is cylindrical anteriorly, but tapers posteriorly to end in a blunt point. The muscular portion of the mantle terminates at the end of the pen which is in advance of the posterior tip of the body. This broad posterior tip appears to be composed of vacuolated connective tissue which continues for a short distance laterally along the sides of the muscular portion of the mantle. The rhachis of the pen is visible in the dorsal midline. The free margin of the mantle projects in the nuchal region and in the regions of the mantle locking-cartilages.

The fins are large, 62–68% of the mantle in length, and broad, 70–83% of the mantle in width. The lateral angles of the fins are rounded. Much of the posterior fin margin forms a nearly straight line at about 45 degrees to the body axis; however, as the edge approaches the mantle, it turns posteriorly and terminates at the tip of the mantle. Anterior lobes are large and free.

The funnel, which is free laterally, is small and muscular. The funnel locking-cartilage bears a straight groove. The dorsal pad of the funnel organ has an inverted V-shape and bears an anterior papilla and lateral ridges. The ventral pads are oval. A large funnel valve is present.

The head width is approximately equal to the mantle width. The eyes are large and occupy the entire lateral portions of the head. A small but distinct anterior sinus is present on each eyelid. Four large nuchal folds are present on each side of the head, the dorsal 2 of which are joined posteriorly. The second fold from the funnel bears the “olfactory” organ, and the fold nearest the funnel forms the posterolateral edge of the funnel groove and bears photophores on its ventral surface. The nuchal cartilage is elongate and has a prominent median ridge that bears a central groove.

The arms are relatively short (arms IV are 55–60% of the mantle in length), and in the order 4>3=2>1. All arms I–III have well-developed aboral keels; arms IV have large lateral keels. Well-developed trabeculate protective membranes are present on the ventral border of arms I–III, but are absent from the dorsal borders of these arms. Arms IV lack protective membranes on both borders.

Arms I–III each bear 2 alternating rows of hooks which give way abruptly to 2 rows of suckers near the arm tip. Each arm has approximately 15–20 hooks and about 20–26 distal suckers. These suckers have roughly 7–10 slender teeth along the distal half of their inner chitinous rings. Arms IV have 13–15 hooks and lack distal suckers; these hooks are smaller than those of the other arms. The tips of arms IV have usually 3 (sometimes 2 or 4) very large photophores encased in dark-purple pigment followed by 2–4 small photophores of similar appearance.

In the males the right arm IV is hectocotylized. It carries 10–12 small hooks and near the distal end of this hook-bearing portion a bilobed membrane, broadest in its proximal half, is present on the ventral border of the arm. Slightly more distal to the hooks, a second membrane arises on the dorsal border. The area beyond the membranes lacks hooks or suckers. McGowan and Okutani (1968) have shown that rarely both ventral arms are hectocotylized. The terminal photophores are the same as in the non-hectocotylized ventral arms.

The tentacles are robust and have short unexpanded clubs which lack protective membranes. The manus has from 3–4 hooks (rarely 2) in each of 2 rows, with those of the ventral row being much larger than those of the dorsal row. The latter generally alternate with small marginal suckers, although the full compliment is not always present. The dactylus consists of approximately 14 transverse rows of 4 suckers each. The inner chitinous ring of each of these suckers is nearly smooth, but may have a few irregular teeth on the distal margin of the first few suckers. A distinct carpal cluster is present which has 4–5 suckers and knobs. Membranes are lacking from the clubs and stalks except for a slight indication of a dorsal keel along the distal half of the dactylus. This keel, when detectable, is very low and appears as only a slender ridge.

The buccal membrane is purple and 8-parted. The buccal membrane connectives attach to the dorsal borders of arms I, II, and IV and to the ventral borders of arms III.

In females, there is a dark patch on either side of the anterior end of the nuchal cartilage which marks the position of the spermatophore receptacles. The inner side of the collar and the outer surface of the body wall adjacent to the nuchal cartilage form a glandular pleated pocket which receives sperm reservoirs. Mature eggs are roughly 1.5 mm in diameter. Nidamental glands are absent and the oviducal glands are greatly enlarged.

The mantle is covered with numerous photophores of varying sizes. The light organs have the greatest concentration on the ventral surface, considerably less on the lateral surfaces, and only a few scattered organs are present on the dorsal surface. These organs from a diffuse pattern over the mantle; mere is, however, a narrow strip extending the full length of the mantle along the midventral line which lacks photophores, and there is a somewhat regular line of photophores that extends around the ventral two-thirds of the mantle along its free margin. Photophores on either side of the mantle are not strictly bilaterally arranged.

The funnel has 6 separate patches of photophores, none of which extends to the exhalent opening. In fully mature specimens, one patch, located on either side of the midline on the ventral surface of the funnel, has between 16–23 photophores. A second patch is found along each lateral margin of the funnel and usually has 5 or occasionally 6 photophores. The other patches, each of which has about 13–15 photo phores, are located on either side of the dorsal surface of the funnel and extend onto the bases of the bridles.

Photophores are abundant on the ventral surface of the head, but have a diffuse pattern with the following exceptions: 2–3 photophores in a single line extend along the medial nuchal fold. From this series, extending dorsally along the posterolateral margin of the head, is a single row of usually 4 photophores (occasionally up to 6) that reaches to a point nearly level with the center of the lens. From usually the second photophore from the ventral end of this series, a single line of broadly separated photophores extends anteriorly along the head (occasionally one or two photophores fall above or below the main line). This series runs lateral to the ventral “window” of the eye. Between this line and the eye opening, there are no photophores except for a line of organs which lie along the margin of the eyelid. The number of photophores in the latter series is relatively constant. A count on 19 eyelids gave 2 with 29 photophores, 3 with 30, 11 with 31, one each with 32, 33, and 34 photophores. There are no photophores over the ventral “window” of the eye. Medial to this area the ventral surface of the head is covered with numerous scattered photophores, diffusely arranged but with a nearly bilaterally symmetrical pattern. Anterior to each “window” there are a few scattered photophores. In the ventral midline, the most posterior photophore lies dorsal to the funnel and immediately in front of the point of attachment of the converging bridles. Immediately anterior to this photophore is a distinct oval area that is usually devoid of photophores.

Arms IV each bear three series of photophores. One series runs along the ventromedial margin of the arm for about two-thirds of the arm length. The middle series runs along the ventrolateral edge of the arm. At the base of the arm this series is arranged in 2–3 irregular rows which shortly converge into a single row that continues to the tip of the arm. Near the arm tip, 2–4 photophores are greatly enlarged and covered with a dark-purple pigment. These are followed by 2–4 small photophores. The final series runs along the edge of the lateral membrane and terminates roughly at the same level as the ventromedial series.

A single row of photophores extends along each arm III at the base of the aboral keel for slightly more man half of the arm length. This series is aligned with 2 photophores of the eyelid series that are situated on the dorsal edge of the eyelid sinus; however, a distant gap separates these 2 series (Plate 1D).

The number of photophores increases with the size of the animal. Therefore, the above discussion applies primarily to mature specimens although many of the features mentioned can also be detected in young specimens.

This squid is covered with numerous reddish brown chromatophores.

TYPE LOCALITY.—31°59′ N, 122°24′ W. Eastern North Pacific Ocean.

LOCATION OF TYPE.—U.S. National Museum, catalog number 678792.

DISCUSSION.—A. felis belongs to the subgenus Abraliopsis which contains only 2 other species: Abraliopsis (Abraliopsis) pfefferi Joubin, 1896, from the Mediterranean; and Abraliopsis (Abraliopsis) hoylei (Pfeffer 1884), from the Mascarene Islands, Indian Ocean (Voss 1967).

A. pfefferi was originally taken off Villefranchesur-Mer, but has since been found to be distributed throughout much of the Atlantic Ocean and into the eastern Indian Ocean (Voss 1967). This species is easily distinguished from the Californian species by a number of characters. A. pfefferi has a semicircular membrane on the ventral margin of the tentacle that originates opposite the carpal cluster and extends distally just past the first hook on the manus. This membrane is lacking in A. felis. A low, but distinct keel on the club in A. pfefferi extends from about the middle of the manus to the dactylus. In A. felis an indistinct keel is present only on the distal half of the dactylus. There are usually between 10–12 hooks on the tentacular clubs in A. pfefferi, while A. felis has usually 6–8. In A. pfefferi there are roughly 44–46 photophores surrounding the eyelid, while A. felis has usually 30–31 photophores. In A. pfefferi, lateral to the ventral “window” of each eye, there is a patch of photophores consisting of at least 4 irregular longitudinal rows. In A. felis there is only a single row of photophores in this position. In A. pfefferi, the series of photophores surrounding the eyelid continues onto arms III. In A felis a distinct gap separates the 2 series. In A. felis there is an oval patch devoid of photophores near the midposterior portion of the ventral surface of the head which is lacking in A. pfefferi. There are other distinctive characters such as the fin length and the number of arm hooks (see also discussion of A. hoylei); however, the features mentioned should be sufficient for the present.

A. hoylei has been the subject of considerable discussion since its original discovery. Hoyle (1904) ascribed several specimens from the Pacific Ocean near Panama to this species. Pfeffer (1912), recognizing the distinctive features of Hoyle’s specimens, placed these forms in a new species A. affinis (a member of the subgenus Micrabralia). Robson (1948) rejected Pfeffer’s name and assigned specimens that he examined from Panama to A. hoylei which he considered to be a polymorphic species. There can be no doubt, however, that A. affinis and A. hoylei are distinct species and even belong in separate subgenera (Voss, in manuscript). Both Berry (1913, 1914) and Allan (1945) ascribed specimens from the South Pacific to A. hoylei, although with some hesitation. These specimens were either too small or too inadequately described for their identity to be ascertained with certainty; however, it is unlikely that they belong to A. hoylei. Therefore, the only valid descriptions of A. hoylei are those of the type specimen (Pfeffer 1884, 1912).

A. hoylei is much more difficult to separate from A. felis than is A. pfefferi. This is not unexpected since one must rely solely on Pfeffer’s (1884, 1912) descriptions of the type as the type specimen has been destroyed (Voss, in manscript), and there are no other specimens available. It is, however, possible to detect the following differences: in the single specimen of A. hoylei, the fins are 78% of the mantle in length and 95% of the mantle in width; the arms (apparently Pfeffer refers to arms IV which are the longest) are 75% of the mantle length. In A. felis the fins are 62–68% of the mantle in length and 70–83% of the mantle in width. Arms IV are 55–60% of the mantle length. The first few distal suckers in A. hoylei are stated to have pointed teeth on the inner rings while the rest have smooth rings. In A. felis the distal suckers have long slender teeth in at least the first 10 suckers. Pfeffer could not describe the tentacular membranes in A. hoyei since the tentacles had undergone some desiccation; however, his illustration (plate 17, figure 9) shows a keel that originates near the middle of the manus. This differs from the previously mentioned conditioned in A. felis.

A number of differences can be detected in the light-organ patterns. Pfeffer described a ventral field of photophores on the ventral surface of the mantle which has a median bare strip lacking photophores on the posterior two-thirds of the mantle. The illustration shows this clear band extending somewhat farther anteriorly, but clearly not approaching the anterior margin of the mantle. In A. felis this clear band extends to the anterior margin or sometimes is separated from it only by a single line of photophores. The ventral field in A. hoylei is bounded laterally by an area which “fast den Eindruck eines leeren Langstreifens macht.” Lateral to this zone is another area of more densely packed photophores which Pfeffer called the laterodorsal field. In A. felis a relatively clear band separating the laterodorsal field is lacking; instead, the number of photophores gradually diminishes laterally. On the ventral surface of the head, Pfeffer described a row of photophores that corresponds to the row immediately lateral to each ventral “window” of the eye of A. felis. Lateral to this row, he describes “eine weitere, unbedeutende kleine Reihe.” This additional series is lacking in A. felis.

One might be somewhat dubious of the small differences here indicated in the photophore pattern between A. felis and A. Hoylei; however, A. pfefferi differs from A. felis in exactly the same features and in the case of A. pfefferi there can be no question concerning the validity of these characters. In fact, the other characters of A. hoylei mentioned here, which separate it from A. felis, are also shared by A. pfefferi. This condition suggests that A. hoylei and A. pfefferi may be synonymous; however, characters such as the number of hooks on the tentacular clubs and other features make this relationship somewhat uncertain. At any rate, it is possible to separate A. felis from both A. hoylei and A. pfefferi and thus verify the specific status of A. felis.

McGowan and Okutani (1968) reduced Watasenia to the status of a subgenus of Abraliopsis and included in it Abraliopsis (Watasenia) scintillans and Abraliopsis (Watasenia) felis. In doing this, these authors relied greatly on the sexual modifications of the ventral arms in members of Watasenia and Abraliopsis.

Descriptions of the males of A. pfefferi and A. hoylei have never been published; therefore, McGowan and Okutani were unaware of the nature of the hectocotylus in these species. I have examined two males of A. pfefferi from the Indian Ocean. The right ventral arm is hectocotylized in each specimen and bears an elongate membrane distally on the ventral margin of the arm. A much smaller membrane is present on the dorsal margin in one specimen, but not the other. The left ventral arm in each specimen is not modified. It is assumed that the very similiar A. hoylei would have a comparable condition.

Therefore, within the genera Abraliopsis and Watasenia all male members seem to have similarly modified right ventral arms although varying slightly between species. In the genus Abraliopsis two species, A. affinis and A. falco, have highly modified trabeculate protective membranes on the left ventral arm in males; a third species, A. gilchristi has the left ventral arm slightly modified by the presence of large trabeculae, but with a reduced or absent protective membrane (Voss 1967). These three species are placed in the subgenus Micrabralia. In A. felis, A. pfefferi, presumably A. hoylei (all subgenus Abraliopsis), and W. scintillans the left ventral is not modified (rarely, in A. felis both ventral arms are similarly modified). Therefore, the sexual modification of the ventral arms in W. scintillans is simliar to these species of Abraliopsis; however, these should not be included in the same subgenus for the following reason: All species of Abraliopsis have two rows of hooks on each tentacular club. This is an extremely important feature which is of generic value (all species of Abralia have a single row of hooks, all species of Enoploteuthis have two rows of hooks). W. scintillans has a single row of hooks on each tentacular club and, therefore, is clearly separable from all members of the genus Abraliopsis.

The tips of both ventral arms in W. scintillans bear several greatly enlarged photophores. This is a highly characteristic feature which is shared by all members of Abraliopsis, but no other groups. Therefore, at present, the status of Watasenia as a separate genus or a subgenus of Abraliopsis rests purely on personal preference, but it is clear that Watasenia scintillans remains in an isolated position and that A. felis belongs in the subgenus Abraliopsis of the genus Abraliopsis.

TABLE 3.—Measurements in mm for Abraliopsis felis McGowan and Okutani, 1968

DISTRIBUTION.—Specimens of Abraliopsis felis over 20 mm M.L. are well represented from zones 1, 2, 5, and 6, having respective frequencies of 0.66, 1.13, 0.50, and 0.63 specimens/hour. In zone 7 the frequence of captures is significantly less, being 0.22 specimens/hour. Specimens less than 20 mm M.L. have a similar occurrence, but have a greater frequency of capture in the more oceanic zones 5 and 6.

McGowan & Okutani (1968) record A. felis from about 43° N to about 27° N along the coast of Oregon, California, and Baja California. Apparently, the southern limit for the species is off Baja California. Juvenile specimens taken by the SWAN, which appear to belong to this species, indicate that the species may extend westward to at least 147° W at 26° N.

Subfamily PYROTEUTHINAE Pfeffer, 1912

DIAGNOSIS.—Nidamental glands present. Mantle, head, and arms without photophores. Photophores present on viscera and embedded in tentacular stalks. Posterior margin of fins convex.

Genus Pyroteuthis Hoyle, 1904

DIAGNOSIS.—Left oviduct present, but reduced. Hooks present on tentacular clubs.

TYPE SPECIES.—Enoploteuthis margaritifera Rüppell, 1844.

Pyroteuthis addolux, new species

PLATES 3B; 4I–Q, S

DESCRIPTION.—The mantle is conical and tapers rapidly to a posterior point. The tip of the mantle consists of little more than an integumentary layer which covers the pointed conus of the pen. The mantle wall, for most of its length, is thick and muscular. The free margin projects in the area of the ventral locking-cartilages and in the nuchal region.

The fins are separate and each is nearly circular in outline, having broad, free anterior and posterior lobes. The fin length is about 40–50% of the mantle length, and the combined fin width is about 70–80% of the mantle length.

The funnel is short; it reaches anteriorly to approximately the level of the midpoint of the lenses. The funnel locking-cartilage has a straight groove. The dorsal pad of the funnel organ has in inverted V-shaped pad which bears median ridges on each of the lateral arms and a very small anterior papilla. The ventral pads are oval and extremely large. A funnel-valve is present.

The head is large; its width is greater than that of the mantle. The large eyes occupy the entire lateral sides of the head. A distinct anterior sinus on the eyelid is lacking. A thick “olfactory” papilla is present on either side of the head at the posterior end just lateral to the funnel; the papilla curves forward in an anteromedial direction. A nuchal crest and nuchal folds are lacking. The nuchal cartilage is oval and bears a median ridge with a central groove.

The arms are robust; they are about 50–60% of the mantle in length and in the order of 4=3=2>l. Large trabeculate protective membranes are present on the ventral borders of arms I–III, but are greatly reduced on the dorsal borders. Arms IV have low membranes on their dorsal borders, but none on their ventral borders. Arms I–III all have well-developed aboral keels; arms IV have large, lateral keels. The armature is biserial throughout and consists of hooks over most of the arm length, although there are suckers basally and distally. The hooks of arms IV are considerably smaller than those of the other arms. The hooks have no trace of an aperature and a small secondary cusp is present beneath the major one. This secondary cusp is greatly elaborated in the proximal hooks of the hectocotylus. There appears to be considerable sexual dimorphism in the armature; however, since only a single male was examined, this will need confirmation. The male lacks distal suckers on arms II and III, while females have approximately 4–9 suckers. The male also has a slightly greater number of hooks (21–23 hooks) on arms II and III than do females (14–17) and a lesser number of proximal suckers. The structure of the suckers appears to be similar on all arms except the dorsal pair. The first 2–6 suckers in the distal series of arms I are typical, but the following suckers have lost the outer chitinous ring and are more globular. The first few of these latter suckers are slightly enlarged over the adjacent typical suckers. The globular suckers have smooth rings. The proximal suckers from arms IV have approximately 10 teeth around the margin of the inner chitinous ring. The teeth of the distal margin are rounded and separate from one another, while those of the proximal margin are truncate and closely adjacent, almost fused to one another.

The right arm IV of the male is hectocotylized by the presence of a short semicircular membrane on the ventral border of the arm about one-third of the arm length from the tip of the arm. The 2 rows of hooks proximal to the membrane are more widely separated than in the other arms.

The tentacles are moderately long and bear short, unexpanded clubs. The club, on its proximal two-thirds, is bounded by very low, trabeculate, protective membranes. The manus has a single row of 3–5 hooks dorsal to which is a slightly irregular row of relatively large suckers, and ventral to it is a row of small suckers. The remainder of the club distally bears 4 rows of suckers that gradually diminish in size. There is a distinct carpal cluster mat consists of 3 suckers and an equal number of pads. The largest suckers of the club each have about 13 broadly separated, blunt teeth around the entire margin of the inner chitinous ring, with those of the proximal margin distinctly larger than those of the distal margin.

There are no photophores present on the surface of the mantle; however, a number of light organs are situated on the visceral sac. A large photophore is located on either side of the anus. A transverse series of 3 photophores is located on the midline between the gills; the median member is much the largest. A large round photophore is located at the base of each gill. Posterior to the median, ventral mantle artery lies a single round photophore that is followed farther posteriorly by another median photophore. The latter 2 photophores are often bilobed. There is a final median, round photophore located near the tip of the body at approximately the level of the most posterior edge of the fin.

Twelve photophores are present on the bulbus of the eye: 3 are situated on the anterior margin near the bases of arms II and III, another is immediately posterior to the lens. On the anteroventral margin of the bulbus, mere is a series of 5 large photophores that are somewhat staggered in position. The second photophore from the posterior end of the series is the largest. Anteriorly and medially from this photophore is an additional series of 3 very small light-organs.

A series of 7 photophores is imbedded in the tentacular stalk. Near the base of the tentacle, the stalk is bent sharply. This bend is easy to locate and is a permanent feature of all members of the subfamily. At the distal end of this area is a round photophore that is deeply buried in the stalk. Shortly distal to this is a smaller round photophore that lies more superficially near the aboral surface of the stalk. Near the proximal end of the carpal cluster is another round photophore located near the aboral surface of the stalk. Approximately evenly spaced between these proximal and distal photophores are 4 elongate light organs near the aboral surface of the stalk.

The dorsal surface of the mantle, head, and arms is covered with large, reddish brown chromatophores; smaller chromatophores of the same color are present on the ventral surfaces. Most chromatophores overlie a silvery, iridescent layer.

The buccal connectives attach to the dorsal borders of arm IV.

The eggs of the gravid females measure 1.0 mm in in diameter. The spermatophores’ of the male are short, compact, and lack coils in the ejaculatory apparatus.

TYPE LOCALITY.—31°39′ N, 133°16′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod Type No. 2.

DISCUSSION.—In addition to Pyroteuthis addolux, there is only one other species, P. margaritifera (Rüppell 1844), in the genus. Hoyle (1886) assigned 2 specimens captured in the South Pacific (4°21′S, 129°7′E, and 0°33′S, 151°34′W) to P. margaritifera. He did not describe the animal, but merely illustrated the club. Pfeffer (1912) felt mat these specimens belonged to a new subspecies, P. m. oceanica, since the illustration of die club showed only 3 hooks on the manus and only 2 suckers in the carpal cluster. I examined specimens from the South Pacific which agree with P. margaritifera from the Atlantic in the numbers of hooks on the manus and the number of suckers in the carpal cluster and in all other discernable features. The craracters Pfeffer referred to are unusual for P. margaritifera, but almost certainly are within die range of variation for this species. Therefore, it does not seem advisable to retain P. m. oceanica as a separate taxon.

Joubin (1924) described a form of P. margaritifera from the Atlantic Ocean which he named P. m. aurantiaca which supposedly had 2 more occular photophores than does the typical P. margaritifera. Joubin’s illustration, however, shows only typical photophores. Voss (personal communication) has examined the type which is deposited in the Monaco Museum and informs me that the eyes are indeed normal for the species and that P. m. aurantiaca has no taxonomic validity.

The specimens of P. addolux are very similar to P. margaritifera, but differ in a number of important features. The most distinctive character is the presence of an additional photophore on the tentacular stalk in P. addolux, giving a total of 7 photophores compared with 6 in P. margaritifera. The long fleshy pad situated on the hectocotylus of P. margaritifera is quite different from the shorter, more membranous pad of P. addolux. The hooks at the proximal end of the hectoctylus are much larger and of a different shape in P. margaritifera (see Plate 4R, S). These hooks are also more numerous in P. margaritifera. I have counted the hooks proximal to the pad on 12 males from the Atlantic and South Pacific oceans and find the following variation in P. margaritifera:

Number of hooks

13

14

15

16

Number of specimens

1

7

3

1

TABLE 4.—Measurements in mm for Pyroteuthis addolux, new species.

the single male of P. addolux has only 10 hooks. At the tip of the hectocbtylus, P. addolux has 6 small suckers while P. margaritifera usually has none, although a maximum of 2 small suckers was observed on the 12 males examined. The spermatophores in P. addolux also seem to be fairly distinctive, but this cannot be certain until more material can be studied since spermatophores are sometimes subject to considerable variation.

DISTRIBUTION.—Only 20 specimens of P. addolux have been taken during the VELERO program. Of these 20, however, only 1 specimen came from the inner 2 basins. Nine specimens were captured in zone 3, 1 specimen in zone 5, 7 specimens in zone 6, and 2 specimens in zone 7. In addition to these, 2 specimens, including the type, were captured by the R/V SWAN between southern California and the Hawaiian Islands at 147° W, 26° N and 133° W, 32° N.

The spotty occurrence off California, the nearly complete absence of P. addolux from the inner basins, and the capture of specimens from the Eastern North Pacific Central Water indicates that this species is probably mid-oceanic and is only rarely present in the inshore waters over the continental borderland.

Genus Pterygioteuthis Fischer, 1896

DIAGNOSIS.—Left oviduct absent. Hooks not present on tentacular clubs.

TYPE SPECIES.—Pterygioteuthis giardi Fischer, 1896.

Pterygioteuthis gemmata Chun, Fischer, 1908

PLATES 3A; 4A–H

Pterygioteuthis gemmata Chun, 1908, p.87; 1910 p.108, pl.13, fig.3; pl.14, figs.4, 5, 9; pl.15, figs.2, 3, 6–12; pl.16, figs. 1, 2, 5, 7–19.—Pfeffer, 1912, p. 194.—Thiele, 1921.—Voss, 1967.

DESCRIPTION.—The mantle is cylindrical anteriorly, but conical posteriorly. The mantle wall is thick and muscular. The pen extends posterior to the fins and terminates in a sharp point. The free margin of the mantle project distinctly in the nuchal region and at the mantle locking-cartilages.

Each fin is large and nearly circular in outline and has broad, free anterior and posterior lobes. The fin length is about 40–50% of the mantle length, and the combined width is about 70–80% of the mantle length.

The funnel is moderate in size, reaching to the level of the midpoint of the eyes. Each funnel locking-cartilages bears a straight median groove. The dorsal portion of funnel organ is an inverted V-shaped pad with a small anterior papilla and a low ridge on each lateral arm. The ventral pads are roughly oval in shape. A large funnel valve is present.

The head is large and has enormous eyes which occupy the entire lateral portions of the head. The eyelid has only a slight indication of an anterior sinus. An “olfactory” papilla is present at the posteroventral edge of each eye as a triangular flap which is swollen at the tip. The olfactory nerve passes along the posterior edge of the flap.

The arms are short and robust. The arm formula is 3>4=2>1. Extremely well-developed trabeculate protective membranes are present on the ventral borders of arms I–III and very weakly developed ones are present on the dorsal borders. Arms IV in females have low, fleshy, protective membranes that are particularly thick near the bases of the arms. Large aboral keels are present on arms I–III, but are best developed on arms III. Arms IV have large lateral keels. The arms have biserial armature consisting mostly of suckers, but with 3–7 hooks on the midportion of the ventral row of each arm I–III. Generally there are 2–4 hooks on each arm I, 4–5 on each arm II, and 5–7 on each arm III, although the maximum range is slightly greater than this (Figure 7). The hooks are considerably larger in males. The suckers of arms IV are much smaller than those of the other arms. The proximal suckers are very small but are gradually larger in size over the first two thirds of the arm, then are progressively smaller to the tip. The larger suckers on all arms possess approximately 6–8 long, slender, truncate teeth on the distal margin of the inner chitinous rings. The hooks are low with broad bases and possess a rather large but irregular secondary cusp beneath the primary one. No remnant of the sucker aperture remains.

The left arm IV of the male is hectocotylized. It bears 2 large fleshy longitudinal ridges (the distal one is somewhat bilobed) between which is located a pocket with pleated lateral sides that contains a toothed plate. The dentition of this plate is slightly variable, but usually has a proximal portion with 5 cusps, then a large single cusp, and a distal set of 2–3 cusps. The distal one-third of the hectocotylus bears low, protective membranes which embrace small, paired suckers in the last 15–20% of the arm. These suckers are fragile and easily lost during capture and preservation. A large lateral keel is present along the length of the arm.

FIGURE 7.—Comparison of numbers of arm hooks between specimens of Pterygioteuthis gemmata from the North Atlantic Ocean (white) and from the North Pacific Ocean off California (black). Crosshatched area represents overlap. Comparisons shown: a, Arms II; b, Arms III; c, Arms II and III.

The tentacles are short and the tentacular stalks are naked. The club is only slightly expanded, and possesses moderately developed trabeculate protective membranes; the dorsal membrane is slightly the larger. A distinct keel is lacking. Each carpal cluster consists typically of 3 suckers and 3 pads; occasionally only 2 suckers may be present. Suckers on the manus and dactylus are in 4 rows. The suckers, in a tranverse series on the manus, grade in size from the largest dorsally to the smallest ventrally. The suckers of the ventral row are roughly half the diameter of those of the dorsal row. The suckers of the dactylus are small and equal in size in a transverse series. Dentition of the inner chitinous ring of each of the club suckers consists of approximately 10–14 short, slender, pointed teeth on the distal margin. The proximal margin is smooth or slightly knobby.

The buccal membrane is very long, purple, and has 8 lappets. The inner lining of the membrane is produced into numerous, pale, globular papillae. The primary buccal membrane connectives attach to the dorsal borders of arms I, II, and IV, and to the ventral border of arms III. Secondary connectives attach to the ventral borders of arms I and II.

Each eye possesses 14 photophores on the surface of the bulbus. On the anterior margin of the eye there is a series of 3 large photophores followed posteriorly by a slightly smaller series of 4 photophores, the third of which has a distinctly different structure. Just posterior to the lens is a rather large, single photophore. Along the ventral medial margin of the eye are 2 large well-separated light organs and a more anterior medial series of 4 very small light organs. Most light organs on the eyes are spectacular even in preservation, reflecting any color of the rainbow depending on the angle of incident light. Each tentacle contains 4 light organs buried deep in the aboral tissue of the stalk. One of these photophores lies beside the carpal cluster; 2 lie close together about one-third of the tentacle length from its proximal end, and the other organ lies at the base of the tentacle. Several light organs of various sizes lie within the mantle cavity. A large photophore lies on either side of the rectum. A medium-size photophore lies in the midline on the viscera between the bases of the gills, while 2 very large ones lie just posterior to the branchial hearts. Farther posteriorly on the midline, beginning at approximately the level of the midpoint of the fins, there is a series of 3 small photophores of which the anterior 2 are sometimes completely divided into paired organs; however, more often the organs are simply bilobed. The most posterior organ lies near the tip of the pen approximately at the level of the most posterior point on the fins.

Large, paired nidamental glands are present. Only the right oviduct, which has a well-developed oviducal gland, is present.

The body, head, and arms are covered by large, reddish brown chromatophores. There are distinct silvery iridescent areas on the body and head, although these are rendered nearly invisible by preservation. The sides of the mantle anterior to the fins seem to have been silvery in life although overlain by chromatophores. Similar iridescent spots are present on the posterior-dorsal margin of the head, the posterior-ventral corners of the head, anterior-dorsal corners of the eyes, and the bases of arms II, III and IV and the tentacles.

TYPE LOCALITY.—South Atlantic Ocean.

LOCATION OF TYPE.—Berlin Museum.

DISCUSSION.—At present the following 3 species of Pterygioteuthis are established in the literature: P. giardi Fischer, 1896; P. gemmata Chun, 1908; and P. microlampas Berry, 1909. P. giardi is very distinctive and easily identified by the presence of hooks in both the dorsal and ventral rows of arms I–III (P. gemmata and P. microlampas have hooks only in the ventral rows). It can also be separated by the presence of 15 photophores on each eye compared to 14 in the other 2 species, and by the lack of suckers on arms IV.

Pterygioteuthis microlampas poses a problem. It is known from only 2 specimens captured near the Hawaiian Islands, both of which are apparently no longer extant.

In establishing P. microlampas, Berry (1914) listed 4 features that distinguished his species from P gemmata. The slight difference in the arrangement of the ocular photophores noted by Berry is probably only a result of the slightly damaged condition of his specimens. The small number of hooks on arms II and III (3 on each) which was previously thought to fall outside the range of variation of P. gemmata is now known to be well within the limits of the latter species, although it is somewhat unusual for a Pacific Ocean specimen. Berry could not locate suckers on the proximal half of arms IV and considered this a distinctive feature of P. microlampas; however, the proximal suckers are often very difficult to observe on P. gemmata since they are very small and are frequently covered by the fleshy protective membranes. It seems doubtful that there are any valid differences concerning this feature. The final character concerns the presence of only 2 suckers in the carpal cluster of P. microlampas. This feature is unusual, but is also occasionally shared by P. gemmata. Therefore, the laterature provides no basis for separating P. microlampas from P. gemmata. The examination of specimens from Hawaii, however, is necessary before synonymizing the two species.

I examined a number of specimens of P. gemmata from the North Atlantic and find that they have fewer hooks on arms II and III than do the California specimens. The total number of hooks on all 4 lateral arms shows almost no overlap (Figure 7c). This is the only feature, however, that seems to differ. I recently examined 17 specimens from the South Pacific and find the hook counts fall somewhat between the specimens from the North Atlantic and North Pacific. The mean number of hooks on arm II is 3.3 for specimens from the North Atlantic, 3.6 for specimens from the South Pacific, and 4.5 for the specimens from the North Pacific. The values for arm III are 4.2, 5.0, and 6.1 for the three areas, and the mean values for the total counts of all 4 lateral arms are 15.2, 17.3, and 21.1. The different hook counts, therefore, seem to reflect geographical variation.

DISTRIBUTION.—P. gemmata has previously been recorded from the equatorial Atlantic (Thiele 1921), from the South Atlantic (Chun 1910; Thiele 1921) from the Indian Ocean (Voss 1967) and off Hawaii (Berry 1914, P. microlampas). I have examined numerous specimens captured by the DANA and PILLSBURY from areas throughout much of the north Atlantic and specimens captured by the ELTANIN in the region of the subtropical convergence in the South Pacific. The species is worldwide in distribution, occurring primarily in warmer waters. For more details concerning the distribution of P. gemmata see Voss (in manuscript).

TABLE 5.—Measurements in mm for Pterygioteuthis gemmata Chun, 1908.

In the present study P. gemmata had relatively low frequencies of capture in the more northern zones of the area under study (i.e., 0.15, 0.08, and 0.30 specimens/hour for zones 1, 2, and 3, respectively) and relatively high frequencies of capture in the more southern zones (1.07 and 1.34 specimens/hour for zones 6 and 7, respectively). This species clearly shows a preference for warm water.

Family OMMASTREPHIDAE Steenstrup, 1857

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with tetraserial suckers on the manus and usually on the dactylus. Buccal connectives attach to the dorsal borders of arms IV. Funnel locking-cartilage with an inverted T-shaped groove. A muscular bridge anterior to the funnel locking-cartilage passes from the funnel to the ventral surface of the head.

Genus Ommastrephes Orbigny, 1835

DIAGNOSIS.—An enlarged tooth present in each quadrant of the largest tentacular suckers. Protective membranes on lateral arms greatly enlarged, particularly on arms III. Arm tips not attenuate; arms containing 50–70 suckers. Funnel and mantle cartilages not fused.

TYPE SPECIES.—Loligo Bartramii Lesueur, 1821

Ommastrephes bartramii (Lesueur, 1821)

PLATES 5B, 6C; 7J–R

Loligo Bartramii Lesueur, 1821, p.90, pl.7.

Ommastrephes Bartramii.—Orbigny, 1839, p.347, in Férussac and Orbigny, Loligo pl.2; pl.21, fig.5; Ommastrephes pl.2, figs. 11, 12.—Berry, 1912b, p.298; pl.47; pl.50, figs.4–5; text figs. 7–8.

Ommastrephes Bartrami.—Steenstrup, 1880, p.73, fig.2; p.79, fig.3; p.81.

Stenoteuthis bartrami.—Pfeffer, 1912, p.465; pls.35–36; pl.39, figs. 1, 2.—Sasaki, 1929, p.289, pl.1, fig.8; pl.24, figs. 1–3; text fig.139.

DESCRIPTION.—The mantle is long and slender (width about 20% of length). Anteriorly it is nearly cylindrical, but at the level of the anterior edge of the fins, it begins to taper rapidly and terminates in a blunt point. A median ventral keel extends from the level of the conus forward for about 20% of the mantle length. The free anterior margin of the mantle has no distinct projections. The mantle wall is extremely thick and muscular.

The fins are about 40%–45% of the mantle in length and about 70%–75% of the mantle in width. They are rather sharply angled laterally and have free anterior lobes.

The funnel is very muscular and rather short. The funnel locking-cartilage is complex. The groove has roughly the shape of an inverted T. The dorsal pad of the funnel organ is very large and has an inverted V-shaped appearance. A small anterior papilla can sometimes be detected. The ventral pads are large and oval. A large funnel valve is present. The funnel is fused to the head forward of each locking-cartilage by a muscular bridge.

The head, approximately the same width as the mantle, has large eyes which occupy almost its entire lateral sides. Each eyelid has a large anterior sinus. A nuchal crest with 4 folds on each side is present; the most medial folds embrance the funnel. Posteriorly the folds all converge. A low oblique ridge on the posterior extension of the second fold from the funnel is apparently the “olfactory” organ. A small “window” is situated ventral to each eye, but there is only a slight trace of a dorsal “window.” The funnel groove contains a large foveola at its anterior end. Seven folds are present within the foveola, and 2–4 side pockets are located lateral to it on either side.

The arms are short (30% to 40% of the M.L.) and powerful. Arms I–III all have well-developed aboral swimming keels. In addition, arms I possess accessory lateral keels. Large lateral keels are present on arms IV. All arms have well-developed trabeculate protective membranes. The membranes on the ventral margins of arms I–III are greatly enlarged, but are most extensive on arms III. The elongate trabeculae extend to the margins of the membranes.

The arms have biserial suckers. Arms I–III bear from 50–55 suckers while arms IV (excepting the hectocotylus) have about 65 suckers. The dentition of the suckers is somewhat variable. The basal suckers from arms III have about 16–18 pointed teeth (some teeth have irregular shapes) on the distal three-fourths of the ring; the proximal margin is smooth or finely toothed. By the fifth transverse row of suckers from the base of the arm, the teeth are more regularly shaped and the entire ring is clearly lined with about 28–32 teeth. The largest teeth occur on the distal margin; however, these teeth frequently alternate with smaller teeth. The outer ring is made up of roughly 75 elongate slats; each, with only a few exceptions, extends unbroken from the inner to the outer margin of the ring. The slats on the distal portion of the ring bear small knobs at their bases. At the fourteenth row the proximal portion of the inner ring is smooth, and the distal portion bears about 7 large, pointed teeth that may or may not alternate with very small teeth. The largest suckers of arms IV are just slightly smaller than those of the first arms which, in turn, are slightly smaller than those of arms II and III.

The left arm IV is usually hectocotylized in males, although occasionally the right arm is modified instead. The largest specimen available (303 mm M.L.) is immature and the hectocotylus is probably only partially formed. All specimens available show some signs of hectocotylus formation which begins with the loss of suckers at the extreme tip of the arm. The largest specimen has 27 suckers on the proximal part of the arm which is followed by a suckerless area which would normally carry 7 suckers. Distal to this is a single row of 6 suckers on the dorsal margin, and the remaining portion of the arm lacks suckers. Distal to the proximal series of suckers, the protective membranes do not appear to have trabeculae. It is likely that this is not a completely mature hectocotylus.

The tentacles bear an aboral keel which runs along the tentacular stalk and is continuous with the dorsoaboral keel of the club. Trabeculate protective membranes occur along both margins of the oral face of the tentacular stalk for most of its length; however, the membrane of the dorsal margin is much more extensive. These membranes extend to the tip of the club along the lateral margins. The clubs are expanded and elongate. The suckers of the manus are arranged in 4 longitudinal rows except at the proximal end where they are irregularly aligned and extend onto the tentacular stalk for a short distance. The suckers on the dactylus are arranged in 4 rows which are continuous with those of the manus. The dactylus suckers grade in size from the largest on the ventral margin to the smallest on the dorsal margin. At the tip of the dactylus, there is a pad of 17 smooth-ringed suckers. At the proximal end of the manus, along the ventral margin, is a row of 2–4 smooth-ringed carpal suckers alternating with 2–4 pads. Proximal to this series lie 5–6 toothed suckers.

The suckers of the medial rows on the manus are nearly 3 times the diameter of the marginal suckers. Numerous fingerlike papillae arise from the fleshy outer edges of these medial suckers. These papillae vary in size and thickness and are often joined at their bases in groups of 4–5, each set looking like a human hand. The papillae are also present on the marginal suckers but are located in only 2 tufts, one on each dorsolateral corner with the biggest tuft nearest the margin of the club. Papillae are not present on the dactylus suckers. On 2 of the largest suckers of the manus 123 and 128 slats were counted in the outer ring. These slats have small knobs at their bases.

The inner rings of the largest suckers of the manus have 4 enlarged teeth, one in each quadrant. Occasionally an additional enlarged tooth may be observed. There are generally 7–10 smaller teeth, which may vary considerably in size, in each quadrant between the large teeth. The marginal suckers have about 25–30 teeth with those of the distal border being much larger than those of the proximal border. Generally, minute teeth may be observed alternating with the distal teeth. The suckers of the dactylus have the same dentition as the marginal suckers on the manus.

The buccal membrane has 7 lappets. The buccal connectives attach to the dorsal borders of arms I, II, and IV, and to the ventral borders of arms III.

The animal is speckled with numerous, small, reddish brown chromatophores. The ventral surface of the mantle bears an elongate, sharply delineated, silvery strip that extends from the anterior edge of the mantle posteriorly to the level of the anterior edge of the fins. The ventral surface of the head also has a silvery background that extends down arms IV. The dorsal surface of the mantle and the dorsal midportion of the fins have a deep-purple coloration that terminates very abruptly laterally. The dorsal surface of the head is similarly colored, as are the dorsalmost portions of arms I–III and tentacles. The lateral sides of the head and mantle have a silvery, iridescent layer benath the chromatophores.

Numerous photophores are imbedded in the musculature near the ventral surface of the mantle. These photophores are not visible in preserved specimens unless the skin is removed, and often it is necessary to chemically macerate the muscle tissue. The photophores are small, closely packed, and extremely irregular in shape. Along the ventral midline of the mantle, there is a strip extending from the conus of the pen to nearly the free margin of the mantle in which the number of photophores is reduced. In the posterior quarter of this strip photophores are lacking.

The concentration of photophores is greatest near the free margin of the mantle. This group of photophores extends along the lateral sides of the mantle margin becoming progressively less numerous. There are no photophores dorsally.

An irregular patch of photogenic tissue is situated on either side of the head at the point of attachment of the muscular bands which pass from the ventral surface of the head to the funnel near the locking-cartilages. At least several photophores are located in the integument and connective tissue immediately anterior to the foveola. Unfortunately, in the specimen examined, the integument was stripped away from most of this area; therefore, there may be more photophores in an intact specimen. Another extremely irregular patch of photogenic tissue is situated on the head near the base of each ventral arm. There is also a long band of photogenic tissue that extends from the base of each ventral arm nearly to the arm tip within the base of the lateral keel.

Within the funnel there is a patch of photophores, clustered approximately into a V-shape, that is located above (dorsal to) the dorsal pad of the funnel organ.

With growth, the patches of photophores on the ventral surface of the head increase in size and change somewhat in shape. The specimen described and illustrated here (Plate 6C) has a M.L. of only 210 mm; therefore, the size of the patches on the head, at least, can be expected to be more extensive in mature specimens.

DISCUSSION.—Three species are presently recognied in the genus Ommastrephes: O. bartramii (Lesueur 1821), O. pteropus (Steenstrup 1855), and O. caroli Furtado, 1887. The latter two are known only from the Atlantic Ocean, whereas O. bartramii has been reported from the Atlantic, Pacific, and Indian oceans. The broad distribution of O. bartramii indicated that it might possibly represent a species complex. I have carefully compared the present specimens against specimens of O. bartramii captured off Florida and find that the eastern Pacific specimens unquestionably differ from those of the Atlantic. The tentacular clubs of the Pacific specimens are distinctly broader than those from the Atlantic. The suckers of the Pacific specimens are about 20% larger than those from the Atlantic specimens of the same mantle length. A specimen of 303 mm M.L. from the Pacific was immature; the hectocotylus was only partially formed and there were no spermatophores present in the genital tract. A specimen of 302 mm M.L. from the Atlantic had a completely formed hectocotylus and spermatophores. There were several minor differences in the structure of the beaks, but it is not known whether these fall within the normal variation for either form.

It is possible that a more complete comparison of all stages of development will reveal additional differences. At present, however, the differences do not seem sufficient to indicate anything more than geo graphical variation.

There is considerable confusion concerning the relationship between O. bartramii and O. caroli. It is even possible that they represent different growth stages of the same species. If this is the case, it is curious that the distributions of O. caroli and O. bartramii do not more closely coincide. O. bartramii can be distinguished from O. pteropus by the presence of 4–6 suckers on the tentacular stalk proximal to the first knob of the locking apparatus compared to 0–2 suckers in O. pteropus. The silvery strip on the ventral surface of the mantle is distinctive of O. bartramii, as is the photophore pattern. The photophore pattern in O. pteropus was described by Roper (1963) and Clarke (1965). The most obvious distinguishing features of the photophore pattern in this latter species are the presence of a large oval patch composed of densely packed photogenic spherules situated on the dorsal surface of the mantle beneath the integument near the nuchal region, and the presence of small, spherical photophores, which are broadly spaced, on the ventral surface of the mantle.

TABLE 6.—Measurements in mm of Ommastrephes bartramii (Lesueur, 1821).

DISTRIBUTION.—There are a number of records of O. bartramii from cold water (Pfeffer 1912, Grimpe 1925, Berry 1912b); however, as there has been considerable confusion regarding the identification of this species these records retain an element of doubt. O. bartramii is common in tropical and subtropical waters, and I consider that this is the usual habitat for the species, although, being powerful swimmers, it would not be unexpected to find stragglers occasionally in high latitudes. In the present study, 5 specimens were captured at one night-light station off Guadalupe Island.

Genus Symplectoteuthis Pfeffer, 1900

DIAGNOSIS.—Funnel locking-cartilage with typical ommastrephid inverted T-shape, but fused in its midportion to the mantle locking-cartilage.

TYPE SPECIES.—Loligo oualaniensis Lesson, 1830.

Symplectoteuthis luminosa Sasaki, 1915

PLATES 5A; 6A, B; 7A–I

Symplectoteuthis oualaniensis.—Watase, 1906, p. 195. Symplectoteuthis luminosa Sasaki, 1915, p.144, text fig. 4, pl.4, figs. 7–13; 1916, p.106; 1929, p.293, text fig. 140, pl.24, figs. 4–5.

Eucleoteuthis luminosa.—Berry, 1916, p.60.—Okada, 1927, p.4.

DESCRIPTION.—The mantle is long and slender (width 15–20% of length); anterior to the fins it is nearly cylindrical, but posteriorly it tapers to a point. The free margin is only slightly produced near the ventral locking-cartilages and not at all in the nuchal region. The mantle wall is extremely thick and muscular.

The fins are about 45–50% of the mantle in length and about 55–60% in total width. They are muscular and attenuate posteriorly. The fins extend to the posterior tip of the mantle. The anterior lobes are free.

The funnel is short and muscular. It is fused to the ventroposterior side of the head by a narrow muscular band extending forward anterior to the funnel locking-cartilage. The funnel locking-cartilage has an inverted T-shaped groove which is fused with the mantle locking-cartilage in the posterior portion of the longitudinal groove. The dorsal pad of the funnel organ has an inverted V-shape with only a slight trace of an anterior papilla. The lateral arms are very long and extend onto the funnel retractor muscles. The ventral pads are rather slender and extremely elongate. The anterior and posterior extent of the ventral and dorsal pads coincides. A large funnel valve is present.

The head is approximately the same diameter as the mantle. The eyes are large and occupy almost the entire lateral sides of the head. A large anterior sinus is present on each eyelid, and there is a small “window” in the skin above and below each eye. A nuchal crest bears 4 folds which are connected posteriorly. An oblique ridge, the “olfactory” organ, lies on the posterior extension of the second fold from the funnel. A distinct funnel groove is present. The foveola has 8 low ridges on its anterior wall. Only a single, small, side pocket can be found on each side of the foveola. The nuchal cartilage broadens at the anterior end and has a median ridge with a central groove.

The arms are short (30–40% of the M.L.) and in the order 2=3=4>1. Arms I–III all have well-developed aboral keels. An additional keel is present along the lateral side of each arm I. Large lateral keels are present on arms IV.

Trabeculate protective membranes are well developed on all arms. On arms I–III, the membranes on the ventral margins are broader than on the dorsal margins. These are most fully developed on arms III where they are wider than the greatest width of the aboral keel of the same arm. The long, slender, supporting trabeculae extend to the margin of the membrane. The trabeculate membranes of arms IV are equally developed on both margins.

The suckers of the arms are biserially arranged and total 43–48 on each of arms I–III and 56–58 on arms IV. The basal few suckers on each arm generally have 3–5 broad, truncate teeth on the upper margin of the chitinous ring and a smooth lower margin. Each broad tooth is the homologue of 2 pointed teeth on the more distal suckers. At rows 5 and 6 on the arms, the teeth have approximately 9–12 sharply pointed teeth on the distal margin that merge with 1–3 broad, generally truncate, teeth on each lateral margin; the proximal margin is smooth. The teeth are more or less of a uniform size, except for an occasional small tooth situated near a larger tooth, although more frequently these are fused, forming a single, somewhat lopsided tooth. The number of teeth is somewhat irregular; some of the large suckers of arms II may have small broadly spaced teeth on the proximal half of the sucker ring. This appears to be a variable feature. Suckers of the seventeenth and eighteenth rows each have 5–6 long pointed teeth on the distal half of their inner rings. The largest suckers of arms IV are roughly half the diameter of the largest suckers of arms II. The largest suckers of arms I and III are only slightly smaller than those of arms II.

The tentacles are short and muscular.. An aboral keel extends the full length of the tentacular stalk and onto the club where it is continuous with the dorsoaboral keel of the dactylus. The clubs are only slightly expanded and bear roughly 115–123 suckers. The suckers of the manus are arranged in 4 longitudinal rows except at the proximal end where the number of suckers rapidly diminishes to a few irregularly arranged suckers that extend onto the tentacular stalk. There are no carpal suckers although a single carpal knob was found near the base of the manus close to the dorsal border of both clubs in one specimen, but not in the other. The suckers in the 2 median rows on the manus are greatly enlarged, the largest being 2½ to 3 times the diameter of the marginal suckers. These suckers each bear a single enlarged tooth on the distal margin of the sucker ring. The remaining margin has low jagged edges which form teeth lacking a regular pattern or shape. The dentition becomes more regular on the smaller suckers. The marginal suckers bear about 30–35 teeth around the entire circumference of the chitinous ring. The teeth of the distal margin are slender, pointed, and roughly alternate between long and short sizes. The teeth on the proximal margin are all short and uniform. The dactylus has 4 longitudinal rows of suckers which are continuous with those of the manus. The suckers grade in size in a transverse series from the largest in the ventral row to the smallest in the dorsal row. The sucker dentition is the same as in the marginal suckers of the manus, although the number of teeth decreases as the size of the suckers diminishes near the tip of the dactylus. There is a small pad of about 7–8 smooth ringed suckers at the tip of the dactylus.

The buccal membrane has 7 lappets. The buccal connectives attach to the dorsal borders of arms I, II, IV, and to the ventral borders of arms III.

This species has a very distinctive photophore pattern. Two broad luminous strips extend almost the full length of the mantle on either side of the ventral midline. Each strip consists of 3 separate parts: a small anterior segment is situated at the free margin of the mantle; a short gap separates this from the second part which extends to approximately the level of the lateral angles of the fins; and this is barely separated from the third portion which continues to approximately the level of the conus of the pen. Near the anterior edge of the mantle are two small patches, one situated on either side of the ventral midline, but medial to the anterior portion of the long strips. Also near the anterior margin, but lateral to each of the long strips, is a medium-size photophore. There is a large photophore on the ventral surface of the head at the base of each arm IV, and another is located on midventral surface of each arm IV.

Many small photophores, generally invisible without dissection, are scattered over the mantle, funnel, head, and arms. These form almost a complete line along the anterodorsal margin of the mantle. They are scattered irregularly over the ventral surface of the mantle, but are absent from its dorsolateral surface. They are scattered along the dorsal midline from the anterior mantle margin to the fins. The inner surface of the funnel has a single photophore on each dorsolateral side at approximately the level of the emergence of the cephalic vein. There is a photophore at the funnel attachment of each of the 2 bridles and 2 small photophores are imbedded in the funnel valve. On the head, a few photophores are scattered lateral to the base of the funnel. I have found only a single photophore in the integument immediately in front of the foveola; however, since most of the tissue there has been torn away, it is quite likely that at least several others are present in intact specimens. A small distinct photophore is present at the medial base of each ventral arm very near the point where the arms unite. A line of irregularly shaped photophores extends along the lateral edge of each ventral arm at the base of the keel in the proximal half of the arm. There is also a series of small photophores along the surface of the axial nerve of each tentacle.

TABLE 7.—Measurements in mm of Sympletocteuthis luminosa Sasaki, 1915.

There can be no assurance that all photophores have been located, since they are extremely difficult to see. For example, Okada (1927) in a histological examination found photophores in the fins, but I have not been able to find them by gross dissection.

The larger specimen examined is a gravid female with greatly enlarged nidamental glands. The mature eggs measure just under 1 mm.

The animal is covered with numerous, small, reddish brown chromatophores. The dorsal surface is darker than the ventral surface, and the lateral sides of the animal have a distinct silvery appearance.

TYPE LOCALITY.—Off Misaki, Japan.

LOCATION OF TYPE.—In Tokyo Imperial University.

DISCUSSION.—Within the genus Symplectoteuthis there are, presently, only 2 recognized species: S. luminosa Sasaki, 1915, and S. oualaniensis (Lesson 1830). (A third species very similar to S. oualaniensis has recently been reported (Clarke 1965), but not described.) The long, luminous strips on the ventral surface of the mantle are present on the former, but not the latter, and easily allow their separation. The present specimens agree with Sasaki’s description except for some differences concerning dentition of the arm and club suckers, the number of side pockets in the funnel groove, and the number of carpal knobs. These features, however, are probably all variable. The small photophores buried within and beneath the integument were first reported by Okada (1927).

DISTRIBUTION.—S. luminosa is known only from the waters off Japan (Sasaki 1929), as possible record from the Kermadec Islands (Berry 1916), and from the California Current (Okutani 1967). It is here recorded off Guadalupe Island at 29°37′ N, 188°49′ W.

Family NEOTEUTHIDAE Naef, 1921

DIAGNOSIS.—Arms with biserial suckers; tentacular club with 4 rows of suckers on the distal part of the manus and dactylus and numerous small smuckers at the proximal end of the manus. Buccal connectives attach to the dorsal borders of arms IV. Funnel locking-cartilage with a straight, simple groove. Anterior fin lobes lacking; posterior fin lobes free. Photophores absent.

Genus Neoteuthis Naef, 1921

DIAGNOSIS.—Fins more than 60% of the mantle length. Club length more than 60% of the mantle length. Most of manus bearing many hundreds of minute suckers.

TYPE SPECIES.—Neoteuthis thielei Naef, 1921.

Neoteuthis species

PLATE 8A–C

DESCRIPTION.—The mantle is cylindrical anteriorly and gradually tapers to a point posteriorly. The mantle wall is thick and moderately muscular; anteriorly it is produced slightly at the nuchal and funnel locks. The pen is visible in the dorsal midline.

The fins are peculiar: they are very elongate (66% of M.L.), but very narrow (29% of M.L.); and they lack anterior lobes, but possess small posterior lobes. The posterior attachment of the fins is slightly in advance of the pointed tip of the mantle. The fins are attached to the lateral sides of the mantle and are, thus, broadly separated from one another in the dorsal midline.

The funnel is small: it reaches just past the level of the posterior margin of the eyes. The funnel locking-cartilage has a simple, straight groove which opposes a straight ridge on the mantle. The dorsal pad of the funnel organ has roughly an inverted V-shape. A small papilla is present near the rounded apex. On each lateral arm of the dorsal pad a ridge is present that slopes from the lateral margin toward the midline, then gradually back to the lateral margin. The lateral arms are very long and extend onto the funnel retractor muscles. The ventral pads are also very large and have a teardrop shape with the more pointed end posterior. The funnel is free from the head laterally.

The head is rounded and bears large eyes which occupy almost the entire lateral sides of the head. A sinus is present on the anterior border of each eyelid, but is extremely small. There is no indication of nuchal folds or a nuchal crest. An “olfactory” papilla is present on either side of the head immediately lateral to the funnel. The nuchal cartilage is broader anteriorly and bears a median ridge with a central groove.

The long arms (60–80% of M.L.) are rather brittle in preservation, and in the order of 4>3>?2>1. Arms I–III bear low, broad, gelatinous protective membranes with gelatinous trabeculae that project well beyond the membrane. Protective membranes are barely detectable on arms IV. The arms have been rubbed, and no evidence of aboral keels remains. Arms IV have low, gelatinous lateral keels.

All arms bear biserial suckers. The suckers are large except for those on the ventral arms, the largest of which is less than half the diameter of the largest suckers of arms III. On arms I–III the suckers are largest in the middle of the arms, but on arms IV the suckers are largest near the base of the arms. Although the sucker rings have almost been destroyed by the fixative, it is still possible to detect about 13–15 apparently semilunar or truncate teeth on the distal two-thirds of each ring; the proximal portion seems to be smooth.

The tentacles are long (64% of the M.L.) and bear very large clubs. The clubs are expanded and the manus, which tapers gradually onto the stalk, bears many hundreds of minute suckers arranged in numerous irregular rows. Approximately 35 suckers are present in a transverse series at about two-fifths of the club length from the proximal end. The most distal of the small manus suckers have very broad outer rings. Each inner ring seems to bear triangular teeth around its entire margin, with those of the distal margin enlarged. The suckers in the middle of the manus have teeth which are all nearly equal in size. The suckers decrease in size at the proximal and lateral margins of the manus, but at the distal end the suckers increase abruptly in size, the largest being about 4 times the diameter of the smaller suckers. Except for the area where the manus and dactylus blend together, the dactylus has suckers arranged in 4 longitudinal rows with the suckers of the ventral margin 2 to 3 times as large as the suckers of the dorsal margin. The medial suckers grade in size between the marginal rows. Except for an irregular circle of smooth-ringed suckers at the tip of the dactylus, all suckers of the dactylus have roughly 26–28 sharply pointed teeth around the entire margin of each inner ring. There is a large dorsoaboral keel on the dactylus that extends onto the distal end of the manus.

The buccal membrane connectives attach to the dorsal borders of arms I, II, and IV, and to the ventral borders of arms III.

DISCUSSION.—The genus Neoteuthis and the family Neoteuthidae were erected by Naef (1921:48). The only diagnosis, given in a footnote, was based on 3 young specimens which were not given a specific name. Later in the same year, Naef named Neoteuthis thielei form a description that Thiele (1921) had given of a small squid from Antarctic waters identified as Octopodoteuthis sp.; therefore, Neoteuthis thielei becomes the type species of the genus by subsequent monotypy. The diagnosis given by Naef and the description given by Thiele greatly lack in detail. Although Naef’s diagnosis states the club has 4 rows of suckers, it seems likely that he merely overlooked the manus suckers since in young specimens the tentacles are disproportionately small and the manus suckers are extremely difficult to see. No new information has appeared on this group since Naef’s work. Fortunately, at Miami there are specimens captured in Antarctic waters by the USNS ELTANIN that look very similar to Thiele’s sketch. Except for this material, it probably would have been impossible to correctly identify the California form.

Although Neoteuthis has a very distinctive appearance, it is closely related to the genus Alluroteuthis, the sole genus of the family Alluroteuthidae. The similarities are: in both, the buccal membrane connectives attach to the dorsal borders of arms IV. Both genera have very distinctive fins which arise from the lateral sides of the mantle and are broadly separated from one another in the dorsal midline. The fins are also slightly subterminal, lack anterior lobes, and have free posterior lobes. The fins in the 2 genera are identical, except that they are much longer and more slender in Neoteuthis. Likewise, the clubs differ only in degree. Alluroteuthis has a cluster of minute suckers at the proximal portion of the manus which corresponds to the much more elaborate manus of Neoteuthis. The appearance of a large broad head is similar in both groups as in the armature of the arms, including the occurrence of reduced suckers on the ventral arms.

These features are sufficient to indicate the close relationship between Neoteuthis and Alluroteuthis. Their families must therefore be considered synonymous, and the more recent family Alluroteuthidae Odhner, 1923, must be submerged into the older Neoteuthidae.

The present species is very close to N. thielei. The specimens of N. thielei that are available for comparison, however, are fragmentary and the original description offers little of specific value. Thus, I am unable to determine whether the present form is conspecific with N. thielei and, therefore, must refrain from giving it a specific name.

Data from the single specimen of Neoteuthis sp. here described are: Sex, ; M.L., 83; M.W., 18; H.W., 23; F.L., 55; F.W., 24; Arms I, 49; II, 56+; III, 64; IV, 68; Tent. L., 118; Club L., 53.

DISTRIBUTION.—The genus Neoteuthis has been reported from Antarctic waters (Thiele 1921), and from the Mediterranean Sea (Naef 1921). The specimen described here came from 30°23′ N,118°29′ W (VELERO Station 10840). Several specimens are present in the collections at Miami which were taken at the following localities: 3°30′ N, 4°41′ E, Gulf of Guinea (PILLSBURY Station 306); 0°31′ S, 11°02′ W and 13°31′ S, 18°03′ W, South Atlantic (DANA collections); and 46°30′ S, 170°9′ W, South Pacific (ELTANIN collections).

This broad distribution suggests mat more than one species may be present in the genus.

Family HISTIOTEUTHIDAE Verrill, 1881

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with suckers generally arranged in more than 4 rows. Large, anteriorly directed photophores over the surface of the mantle, head and arms. Left eye considerably the larger. Buccal connectives attach to the dorsal borders of arms IV. Funnel locking-cartilage with an approximately straight, somewhat broad groove.

Genus Histioteuthis Orbigny, 1839

DIAGNOSIS.—The characters coincide with those of the family.

TYPE SPECIES.—Cranchia bonnellii Férussac, 1835.

Histioteuthis heteropsis (Berry 1913)

PLATES 9A; 10D–G

Melagrotcuthis hoylei —Berry, 1912b, p.305, text figs. 13–16; pl.50, figs. 1–3; pl.51, figs. 1–5; pl.52, figs. 6–7.

Calliteuthis (Meleagroteuthis) heteropsis Berry, 1913, p.75.

Histioteuthis heteropsis.—Voss, 1969, p.815.

DESCRIPTION.—The mantle is short and broad; it tapers gradually posteriorly to a blunt tip. The mantle margin projects slightly in the region of the dorsal and ventral locking-cartilages. The mantle wall is very thick, but moderately gelatinous.

The fins are short, about 35–10% of the mantle in length and about 50–55% of the mantle in total width. Each fin is roughly semicircular and has free anterior and posterior lobes. Medially, the fins touch one another only at the posterior end where they extend slightly past the tip of the mantle. Anteriorly, the fin insertions are broadly separated from one another.

The funnel is short. The funnel locking-cartilage has a broad, simple, generally straight groove that is deepest in the midportion. This structure opposes a straight or slightly curved ridge on the mantle, the anterior end of which is flanked on either side by additional short broad ridges. The dorsal pad of the funnel organ has an inverted V-shape with a small papilla at the anterior apex. The ventral pads are oval. A funnel valve is present.

The head is broad and asymmetrical due primarily to the unequal development of the eyes; the left eye has almost 1 2/3 the diameter of the right. An anterior sinus is present on each eyelid. A nuchal crest and nuchal folds are not clearly developed. The nuchal cartilage is short and much broader anteriorly than posteriorly.

The arms are broad and short, although slightly greater than the mantle in length. Aboral keels are well developed only on the midportion of arms III; they are represented by faint ridges on arms I and II. The lateral keels are well developed on arms IV. Low, thick, gelatinous protective membranes, lacking distinct trabeculae, are present on all arms. The arms have biserial suckers. The suckers are globular and have relatively narrow outer rings, and inner rings with about 8–10 rounded to truncate teeth on the distal margin. All arms have low inner and outer webs.

The tentacles are short and have small (30–40% of M.L.) expanded clubs. The manus has 7 irregular longitudinal rows of suckers. The third and fourth rows from the dorsal margin bear the largest suckers; the remaining suckers become progressively smaller toward the club margins. Sucker rows near the ventral margin of the club extend slightly farther proximally than the others. Proximally, the dactylus has 7–8 irregular longitudinal rows of suckers; however, these are not all continuous with the rows on the manus. At the base of the dactylus, a new row begins along the dorsal margin giving a total of 8 rows for a short distance; however, rows that have continued on from the manus begin terminating, and approximately at the midpoint of the dactylus the number has dropped to 6 rows, then shortly to 5, and finally to 4 rows at the tip of the dactylus. The extreme tip is occupied by a circular pad of small suckers. The largest suckers of the manus have large, pointed or slightly truncated teeth around the entire margin of the inner chitinous ring. The teeth of the distal margin are only slightly larger than those of the proximal margin in large specimens. In smaller specimens, the size difference is much greater. A trabeculate protective membrane extends along the entire ventral margin of their club, but dorsally it disappears about the middle of the dactylus. A dorsoaboral keel is present on the dactylus.

The carpal locking-apparatus begins in an oblique line of 3 suckers with alternating pads along the dorsal-basal part of the manus. Three to 4 suckers follow with alternating pads that extend proximally in a single line along the dorsal margin of the club and onto the tentacular stalk. Medial to the end of this series is a single small suckers and pad aligned longitudinally and situated slightly to the ventral side of the midline. The major series of suckers and pads crosses from the dorsal margin of the stalk to the ventral margin. Elements transversing the stalk may consist either of 2 suckers, or 2 pads, or 2 suckers and 2 pads. Along the ventral margin of the tentacular stalk, the locking-apparatus consists of a single line of pairs of suckers alternating with pairs of pads. There are 2–3 sets of suckers in this series. The last element in the series is a single sucker set some distance proximal to the alternating suckers and pads.

Histioteuthis heteropsis is covered with numerous small, anteriorly directed photophores. On the anterior portion of the ventral surface of the mantle, photophores are aligned roughly in closely packed transverse rows. There are approximately 40–45 photophores along the free edge of the mantle. On the posterior portion, the photophores are irregularly distributed. The number of photophores decreases laterally; on the dorsal surface of the mantle they are sparse and smaller in size. Photophores are absent from the ventral surface of the fins, but occur in small numbers on the dorsal surfaces. The funnel lacks photophores. The ventral surface of the head is covered with many irregularly distributed photophores. About 19–21 photophores are situated at the edge of the circular eyelid of the right eye and only a few, small, scattered light organs are present near the eye opening of the right eye and none of these lies on the edge of the eyelid. Large photophores are scattered over the right side of the head, but the left side has smaller and less numerous photophores. The dorsal surface of the head has small, scattered photo phores. At the base of the ventral arms there are 8–9 longitudinal rows of light organs. These decrease to 7 about halfway along the arm length. Arms III have 4 rows near the arm base, but only the ventral row extends to the arm tip. Arms I and II each has a distinct row along its ventral border which extends to the arm tip. Other portions of these arms bear small, scattered photophores.

TABLE 8.—Measurements in mm of Histioteuthis heteropsis (Berry 1913).

The buccal membrane has 7 lappets, and the primary buccal connectives attach to the dorsal borders of arms I, II, and IV and to the ventral border of arms III. An accessory connective attaches to the ventral border of each arm I.

This animal has a uniform purplish red coloration.

TYPE LOCALITY.—Monterey Bay, California, eastern North Pacific Ocean.

LOCATION OF TYPES.—United States National Museum and personal collection of S. S. Berry, numbers 108, 110.

DISCUSSION.—H. heteropsis can be separated from all other members of he family by the numerous small photophores on the body, head, and arms (i.e., 8–9 rows across the base of each ventral arm), combined with a lack of cartilaginous tubercules on the arms (Voss 1969).

DISTRIBUTION.—H. heteropsis is a common species off southern California. The northernmost record is from Monterey Bay (Berry 1912b) while the southernmost record in this area is from off Baja California at 27°34′ N. (Voss 1969). Within the area of the VELERO survey, the highest frequency of capture, 0.52 specimens/hour of trawling, occurred in zone 2 (Santa Catalina Basin). The other major zones also had a relatively high frequency (.22 to .37 specimens/hour) with the exception of zone 7. In this latter zone only 3 specimens were taken, giving the low frequency of 0.03 specimens/hour. Although not entirely convincing, this data suggests that H. heteropsis is primarily a cold-water form, and that its southern limit occurs somewhere off Baja California.

Voss (1969) also recorded a specimen, captured by the USNS ELTANIN, belonging to this species from off the west coast of South America at latitude 33° S. Although the data are sparse, it is possible that H. heteropsis is an antitropical species with disjunct populations, one in the temperate northeastern Pacific and the other in the temperate southeastern Pacific.

Histioteuthis dofleini (Pfeffer, 1912)

PLATES 9B; 10A–C

Calliteuthis ocellata.—Chun, 1910, p.147; pl.I, figs.1, 2; text fig.23a, b.

Stigmatoteuthis Dofleini Pfeffer, 1912, p.288.

Stigmatoteuthis dofleini.—Sasaki, 1929, p.258; text fig.126—127; pl.22, figs.1–3.

Histioteuthis dofleini.—Voss, 1967, p.74; 1969, p.784.

DESCRIPTION.—The mantle is thick walled and moderately muscular. It is conical, but tapers to a blunt point. The free margin projects at the locking-cartilages. Beneath the thin outer integument, the mantle is covered with short, broad, wartlike papillae.

The fins are short (about one-third of the mantle in length and one-half of the mantle in width). Each fin is nearly semicircular in outline and has small, free anterior and posterior lobes. The papillate tissue of the mantle extends onto the medial portion of the dorsal surface of the fins.

The funnel is short, and free from the head laterally. The dorsal pad of the funnel organ has an inverted V-shape with an inverted V-shaped central ridge terminating in a small anterior papilla. The ventral pads are oval. The funnel locking-cartilage has a straight but deep and broad central groove which opposes a large, slightly curved ridge on the mantle.

The head is large and broad. The left eye is nearly twice the size of the right. There appears to be 3 nuchal folds rising from the nuchal crest on either side of the head. The second fold bears the “olfactory” organ. The nuchal cartilage is long and slender; it bears a median ridge with a central groove. The head and aboral surfaces of the arms have weltlike papillae beneath the outer epidermis as on the mantle.

The arms are very long (over 175% of the M.L.); they are broad at the bases and attenuate at the tips. The arm formula is I = IV>II = III. Rather broad aboral keels are present on the distal half of arms III, but arms I and II have keels that are only slender ridges. Arms IV have thick, fleshy lateral keels. Arms I–III have large trabeculate protective membranes on both the dorsal and ventral margins; these are much less developed on arms IV. Arms I of the single large female examined have suckers arranged in 3 rows at the arm tips (this is not typical of the species according to N. Voss); otherwise, all arms have biserially arranged suckers. The suckers of arms I–III are much larger than those of arms IV. The inner chitinous rings have been destroyed by the fixative in the largest specimen; however, in the 14 mm M.L. specimen, about 7–10 triangular teeth are present on the distal half of the inner ring; laterally and proximally these diminish to small knobs. An outer web connecting all arms at their bases is relatively well developed, being approximately 20–25% of the mantle in length.

The tentacles are long and muscular. The tentacular clubs are slender but expanded, and nearly 40% of the M.L. The suckers of the manus are somewhat irregularly arranged, but can be generally divided into 5 longitudinal series. There is a row of very large suckers in the median portion of the club. Dorsal to this row is a very irregular row of medium-size suckers that alternate somewhat in size. Ventral to the central series is another, more regular row of medium-size suckers. Along both the dorsal and ventral margin is a row of very small suckers. Along the ventral margin, this row may be incomplete. The suckers of the dactylus also appear somewhat irregularly arranged. In a transverse line through the dactylus there are 3–4 suckers. At the tip of the club there is a small flap that carriers 7–9 suckers. The inner rings of the largest suckers of the manus have as many as 60 sharp teeth around the entire margin. A large dorsoaboral keel is present on the dactylus. The carpal locking-mechanism consists of a row of 2 suckers with alternating pads at the base of the manus along the dorsal border. Proximal to the manus there is a median row of 3–4 alternating suckers and knobs. A small sucker is located near the proximal member of this series. Proximal to the median row is a series of 2–3 pairs of suckers alternating with 2–3 pairs of knobs. The first set in this series is situated obliquely on the stalk, but the remaining lie along the ventral margin. This series terminates with a single sucker or knob.

The buccal connectives attach to the dorsal borders of arms I, II, and IV and to the ventral borders of arms III.

The ventral surface of the mantle is covered with large, well-separated photophores. A somewhat irregular row of smaller photophores extends along the free margin of the mantle. Dorsally the photophores in this series lie somewhat farther from the mantle margin and are very small. There are 22–23 photophores in this series. On the dorsal surface of the mantle, the photophores are small and scattered. I counted a total of 103 photophores on the entire mantle; however, since some are very difficult to locate in the present material, this number may not be exact. The photophores on the ventral surface of the mantle are arranged approximately in diagonal rows.

Photophores are arranged in diagonal rows on the ventral surafce of the head, except for a transverse series that extends along the posterior margin. Seventeen photophores, embedded in the margin of the eyelid, encircle the right eye. Photophores are not so regularly arranged around the enlarged left eye; unfortunately, their exact arrangement cannot be determined in the material available. A few scattered photophores are present on the dorsal surface of the head. In the proximal half of the ventral arms, photophores are arranged in 3 longitudinal rows. On the third quarter, or slightly more, of the arm, photophores are arranged in 2 rows, and on the distal segment only a single row is present.

Arms I–III all have a single row of large photophores, extending the full length of the arms along the ventral margin of the aboral surface. A second row, near the dorsal margin, of very small photophores is also present on all arms. It is possible that other photophores are present, but not detectable in the present material.

This animal has a uniform purplish red coloration.

The data for the largest specimen available are: Sex, ; M.L., 150 mm; M.W., 65 mm; H.W., 75 mm; F.L., 54 mm; F.W., 77 mm; arm lengths for arms I–IV respectively, 265 mm, 318 mm, 313 mm, 268 mm; Tent. L., 345 mm; Club L., 61 mm.

TYPE LOCALITY.—Sagami Bay, Japan.

LOCATION OF TYPE.—Not traced.

DISCUSSION.—The present material was identified by N. Voss. She informs me that this species can be separated from all others in the family by the presence of paired genitalia in males, and by the papillate skin in some specimens (i.e., this is a variable character). The short, conical mantle and the very long arms and relatively few photophores separates this species, at a glance, from H. heteropsis. A third species Histioteuthis corona, subspecies berryi has been reported once (Voss 1969) near this area (i.e., about 29° N, 126° W). It is superficially simliar to H. dofleini except for the somewhat shorter arms. This species can be easily separated from H. dofleini by the presence of four longitudinal rows of light organs along the proximal half of each ventral arm, compared to three rows in H. dofieini.

DISTRIBUTION.—H. dofieini occurs primarily in tropical and subtropical waters in both the Atlantic and Pacific oceans, although there are a few records from colder waters (Voss 1969). In the present study, only three specimens (M.L., 150 mm, 14 mm, 9 mm) were taken; these were captured in zones 1 and 7 .

Family OCTOPOTEUTHIDAE Berry, 1912

Veranyidae Chun 1910, p. 139. [Family name inadmissable because originally based on a generic name not currently valid. See Roper, Young, and Voss, 1969.]

Octopoteuthidae Berry, 1912a, p. 645.

Octopodoteuthinae Pfeffer, 1912, p.212.

DIAGNOSIS.—Arms with biserial hooks that may be replaced by suckers near the arm tips. Tentacles present only in larvae (rarely in juveniles). Buccal connectives attach to the ventral borders of arms IV. Fins more than 70% of the mantle length. Funnel locking-cartilage with a simple, approximately straight but broad groove.

Genus Octopoteuthis Ruppell, 1844

DIAGNOSIS.—Photophores present at the tips of all arms. At M.L., 15–25 mm, all trace of tentacles has been lost.

TYPE SPECIES.—Octopoteuthis sicula Rüppell, 1844.

Octopoteuthis deletron, new species

PLATES 10H–M; 11; 12A–D

Octopodoteuthis sp.—McGowan, 1967, p.209.

Octopoteuthis sicula.—Pearcy, 1965, p.261.

DESCRIPTION.—The conical mantle gradually tapers to a blunt point posteriorly. The tip of the mantle is almost always contracted and pulled ventrally and anteriorly, giving the posterior end of the body a very blunt appearance. The mantle extends slightly posterior to the fins in the larger specimens, forming a short, broad tail. The tail is not detectable in young specimens, but increases allometrically with growth in specimens larger than about 40 mm M. L. up to maturity when the tail is about 18% of the M.L.

The fins are very large and muscular. At their posterior ends, they are reduced to narrow strips that extend along each lateral edge of the tail and join at the apex of the tail. Although their anterior and posterior lateral edges are slightly flattened, the fins are nearly elliptical in outline, except for the posterior attenuation. Their width is 95% of the M.L., and their length (exclusive of the tail) is 86% of the M.L. in the holotype. Small free, anterior lobes are present. The fins fuse at the dorsal midline.

The funnel extends approximately to the level of the midpoint of the eyes. It is free from the head laterally. The funnel locking-cartilage is basically of the simple, straight type, although posteriorly the groove curves slightly and flares. The dorsal pad of the funnel organ has roughly an inverted V-shape with a prominent ridge and papilla at the apex and short ridges on the arms; the organ, however, is somewhat variable in appearance. The ventral pads are very large and oval; a funnel valve is present.

The head is large and bears large eyes which extend the entire length of the head. A small optic sinus is present at the anterior end of each circular eyelid. Along the posterior margin of each eyelid is a large, broad, crescentshape patch of muscular tissue. The “olfactory” papillae lie at the posterior edge of the head, lateral to the base of the funnel; each consists of a white, flattened terminal bulb with a short, swollen stalk. Both the mantle and head are encased in a thick layer of gelatinous tissue.

The arms are. long (over 70% of the M.L. in the holotype). The arms are brittle and almost always broken at the tips; in addition, those that can be measured show considerable variation. The arm formula, therefore, is difficult to determine with certainty; in mature specimens they appear approximately subequal. All arms have gelatinous aboral keels. Low, thick, gelatinous protective membranes are present on all arms, but are only apparent in large specimens. These gelatinous ridges are joined across the midline of the arm by membranes which alternate with the hooks; the ridges and membranes form the walls of cup-shape depressions from which the arm hooks arise. The arms bear 2 rows of small hooks which generally are partially or entirely hidden in thick-walled sheaths. Two small, but distinct, accessory cusps occur on each hook. These cusps are relatively more developed in large specimens. The hooks occupy most of the arm lengths, but near the arm tips they abruptly give way to approximately 3–11 pairs of small, globular suckers which have about 6–9 somewhat irregular, pointed teeth, including a large central tooth in each of the more proximal suckers. The number of suckers seems to depend somewhat on size and stage of development, although in the largest specimens they vary from 6–11 pairs. The extreme distal ends of the arms are bare, swell slightly, and have a small bulb at their tips. The swelling and bulb are generally more pigmented than the rest of the arms and are luminescent.

The tentacles are completely lacking in adult specimens; however, a specimen with a mantle length of 17 mm has large blunt stubs of the tentacles. Larval forms have small tentacles with a few enlarged suckers at the tips, but no details can be given due to the damaged state of the larvae available.

Octopoteuthis deletron has a considerable number of photophores; however, most are inconspicuous. A single large photophore is buried deep in the mantle tissue in the ventral midline near the posterior end of the body. This abdominal photophore is difficult to locate in specimens less than 30 mm M.L., but very careful examination will reveal its presence in specimens somewhat less than 20 mm M.L.

A large photophore is embedded deeply in the gelatinous tissue of the head just posterior to each eye. As described earlier, the tips of all the arms bear elongate photophores. An elongate, obliquely oriented photophore lies inconspicuously near the anterior medial margin of each eye on the ventral portion of the head. These latter photophores do not seem to be developed in specimens less than about 30 mm M.L. A series of photophores lies near the central core of each of the ventral 4 arms interior to the longitudinal muscles, and are often adjacent to the axial nerves. The basal photophore on each arm is larger than the following ones and is somewhat more isolated from the others. The remaining arm photophores, numbering about 25 on each arm IV, are situated at regular intervals up to the level of the distal sucker-bearing zone of the arms where the series ends. Arms II have only the basal photophore and the arms I lack any trace of this series. In preserved specimens, it is necessary to pick away the muscular tissue of the arms to see these light organs. Another pair of photophores lies near the ventral surface of the ink sac. These photophores are overlain by a pair of muscular bands.

In mature males the penis is large and appears capable of extending well beyond the opening of the mantle cavity. Gravid females have greatly enlarged nidamental and oviducal glands. The mature eggs measure about 2 mm in diameter.

Specimens are covered with reddish brown chromatophores. The external layer of chromatophores that lie in the aboral gelatinous tissue on arms III and IV are arranged in a peculiar manner, in that they lie in parallel transverse planes relative to the axes of the arms and are evenly spaced, giving the appearance of a series of lamellae. The mature female has a uniform dark-purple color.

Octopoteuthis deletron undergoes rather drastic developmental changes at a M.L. of about 30–40 mm. This is the change from the juvenile to the adult form, although maturity is not reached until a much larger size. The most apparent changes include the great increase in eye size, the thickening of the arms and body, and the acquisition of greater pigmentation. These changes are illustrated in Plate 12A–D.

The radula has a tricuspid rachidian and a bicuspid first lateral, unicuspid second and third laterals, and marginal plaques.

The primary buccal connectives attach to the ventral margins of arms IV.

TYPE LOCALITY.—33°15′ N, 118°37′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod Type No. 3.

DISCUSSION.—In the genus Octopoteuthis there are 7 nominal species: O. sicula Rüppell, 1844, near Messina, Mediterranean Sea; O. megaptera (Verrill 1885), off east coast of United States, 44 mm M.L.; O. persica Naef, 1923, off south tip of Africa, 34°31′ S, 26°0′ E, 4.7 mm M.L.; O. indica Naef, 1923, Gulf of Aden, Indian Ocean, 13°2′ N, 46°41′ E, 3.8 mm M.L.; O. danae Joubin, 1931, near Bermuda, 33°15′ N, 68°20′ W, 31 mm M.L.; O. nielseni Robson, 1948, near Cocos Island, Pacific Ocean; O. longiptera Akimushkin, 1963, from sperm whale stomach, 20° S, 25° W, Atlantic Ocean.

O. persica is based on an illustration in Chun (1910). The specimen shows distinct swellings near the tips of arms II and extremely broad fins. Both features are strongly suggestive of the genus Taningia, and I think it safe to transfer this species from Octopoteuthis to Taningia.

All members of the genus seem to have the same basic pattern of large photophores, with the exception of the abdominal series which has either 1 median or 2 lateral photophores, depending on the species. Joubin (1931) first noticed the presence of the larger photophores, and this resulted in the establishment of a new species, O. danae. O. danae seems to agree in almost all respects with O. sicula; unfortunately, the arrangement of abdominal photophores in specimens of O. sicula from the Mediterranean is unknown. Still, there would be little evidence to prevent synonymizing these 2 forms if an additional, similar species did not occur in the eastern North Atlantic. In contrast to O. danae which has 2 abdominal photophores this species has only a single abdominal photophore, and I shall henceforth refer to it as Octopoteuthis A.

The other common Atlantic species, O. megaptera, from the western north Atlantic and Gulf of Mexico can easily be distinguished by the greater size of its “tail,” by the presence of a 2 abdominal photophores, and by large accessory cusps on the arm hooks even in young stages.

O. longiptera was very inadequately described. The only character of possible specific significance is the relative dimensions of the fins; their length is 135% of the width. (This measurement apparently includes the tail.) This feature is probably only a result of the large size of the specimen (590 mm total length).The specimens from California at 30–40 mm M.L. have fins whose lengths (including the tail) are approximately 55%–65% of their widths; at 109 mm M.L. the fin length is 90% of its width. In the 167 mm male (=300 mm total length) the percentage is 105, while in the 148 mm female the percentage is 120. G. Voss informs me that the type of O. longiptera is not extant. Since O. longiptera cannot be clearly separated from any of the existing species, it must be considered a species dubia.

Unfortunately, O. indica from the Indian Ocean must be ignored for the present, since its small size prevents adequate comparison to related forms.

Therefore, we are left with at least 3 distinct species in the Atlantic Ocean: Octopoteuthis A, O. danae, and O. megaptera. One of the former 2 species (most likely O. danae) must certainly be a synonym of the Mediterranean species O. sicula. This problem, however, cannot be resolved with certainty until the photophore pattern of O. sicula is determined. In making comparisons with the Califomian species, I shall refer only to the 3 Atlantic forms and assume that O. sicula is included in one of them.

In the eastern Pacific, at least 2 additional species are present: O. deletron from California and possibly Peru (see discussion of distribution), which is characterized by a short tail and a single abdominal photophore, and O. nielseni from the region of Panama. O. nielseni was only briefly described by Robson, and the critical abdominal photophore pattern was not given. G. Voss has recently (1962) examined the type in the British Museum and informs me that the specimen is so damaged that it is not possible to determine the photophore pattern. In the DANA collections, there is a specimen which was captured at 6°48′ N, 80°33′ W off Panama. It is probably safe to assume that this species is O. nielseni. It is a short-tailed form, as Robson (1948) illustrated, and possesses 2 posteroabdominal photophores. This is a large specimen (M.L., 112 mm) which, surprisingly, has only a slight trace of accessory cusps on the arm hooks. This latter feature would seem clearly to separate O. nielseni from O. danae, which possess distinct accessory cusps at least in the adult stage (Adam 1952:73, as O. sicula), but the value of this character is somewhat uncertain; I examined a small specimen captured off the west coast of Mexico which also appears to be referable to O. nielseni, but has distinct cusps. Also specimens of O. danae from the Atlantic can be divided into young forms with accessory cusps and young forms without them. Since O. deletron is quite consistant in the presence of this feature, it is possible but perhaps unlikely that these different types represent distinct species. The reason for this confusion and difficulty in resolving the specific status of the different forms results from the presence of only a few specific systematic characters within the genus. This condition is partially the result of the almost invariable loss of the arm tips during capture.

TABLE 9.—Measurements in mm of Octopoteuthis deletron, new species.

If the difficulties concerning the accessory cusps on the hooks can be explained by individual variation, then it becomes very difficult to separate O. danae and O. nielseni. I believe, however, that O. nielseni will prove to be a valid species and, therefore, believe that its name should be retained at least until detailed data are available on all growth stages of both species.

O. deletron is thus separable from all other species except Octopoteuthis A by the presence of only a single abdominal photophore. It differs from Octopoteuthis A by having a much shorter tail at equivalent mantle lengths and by having well-developed accessory cusps on the arm hooks which are lacking in the latter species.

DISTRIBUTION.—In the present study, O. deletron had a fairly uniform frequency of capture in zones 1, 2, 3, 5, and 6 (i.e., fluctuates between 0.18 and 0.36 specimens/hour of trawling). No specimens were captured in zone 7. McGowan (1967) has recorded larvae identified as Octopodoteuthis sp, which undoubtedly belong to the present species, from 34° N to 30° N in the southern California area. Pearcy (1965) reported O. sicula from off the coast of Oregon; his specimens possess a single abdominal photophore (G. Voss, personal communication) and can, almost certainly, be referred to O. deletron. This information suggests that O. deletron is a cold-water species with its southern limit somewhere off Baja California; however, the ELTANIN has taken a large specimen of Octopoteuthis from 7°45′ S, 81°37′ W off northern Peru that appears to be identical with O. deletron. If this is the same species, present evidence would favor the occurrence of two disjunct populations rather than a continuous distribution from Oregon to Peru.

Family GONATIDAE Hoyle, 1886

DIAGNOSIS.—Arms with tetraserial armature; arm tips may be modified. Median 2 rows on arms I—III consist of hooks except at the extreme base and tip of each arm, with the exception of Berryteuthis anonychus which lacks hooks except at the bases of arms I—III in females. Funnel locking-cartilage has a simple straight groove which may flare slightly at the posterior end in some species. Buccal connectives attach to the ventral borders of arms IV. Nuchal folds present.

Gonatus Gray, 1849

DIAGNOSIS.—Five teeth present in a transverse series on the radula. Tentacular club with a large central hook and generally additional hooks in the median line. Each tentacular club bears a distinctive locking apparatus on the dorsal border of the manus which consists of a series of large, thick, transverse pads with medial alternating oval pads and suckers.

TYPE SPECIES.—Onychoteuthis? amoena Møller, 1842.

Gonatus onyx, new species

PLATES 13A; 14A, C–I; 17J

Gonatus fabricii.—Berry, 1912b, [part], p.308, pl.52, fig.4.—? Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is long and slender; posteriorly it tapers gradually to a pointed tip. The muscular portion of the mantle which is moderately thick terminates at the edge of the conus. A muscular cone with a gelatinous core forms a tail that extends posteriorly from the apex of the conus. This structure is not apparent in young specimens, but increases allometrically and becomes prominent in large specimens. The free margin of the mantle projects slightly at the mantle locking-cartilages.

The fins have free anterior lobes. The posterior margins are drawn out along the tail in large specimens. The lateral edges of the fins are rounded.

The funnel is free from the head laterally and reaches to approximately the level of the lenses of the eyes. Each funnel locking-cartilage bears a straight, simple groove. The dorsal pad of the funnel organ has the shape of an inverted V; the arms are broadest in the posterior portions; the anterior portions each have a median longitudinal ridge that extend to the apex. Near the anterior apex, which is blunt and slightly indented, a slender papilla arises. The ventral pads are oval. A large funnel valve is present.

The head is oval and bears large eyes that occupy the entire lateral sides of the head. An anterior sinus is present on each eyelid. Three pairs of nuchal folds are present; the “olfactory” organ arises from the second pair. The nuchal cartilage is elongate and rounded at the anterior end; it has a median ridge with a central groove and marginal ridges.

Arms II and III are very muscular, nearly equal in size, and 50–60% as long as the pen. The ventral arms are the same length or only slightly less than the lateral arms, but are more slender. Arms I are the smallest; they reach about 40–50% of the P.L. Arms I–III all have gelatinous aboral keels which are best developed on arms III. Arms IV have large lateral keels along their entire lengths. Well-developed protective membranes with long, slender trabeculae are present on arms I–III; they are less developed on the arms IV. The armature is in 4 longitudinal series. On arms I–III the median rows consist primarily of large hooks and the marginal rows of small suckers. The suckers arise near the distal ends of the trabeculae. At the extreme tips of the arms, the hooks are replaced by suckers. A total of 25–30 suckers and hooks occur on the proximal half of each arm I–III. Arms IV lack hooks and have 4 series of small suckers. Specimens larger than 40 mm P.L. have 30–37 suckers on the proximal half of arms IV. The arm suckers all have similar dentition. Long, pointed teeth are present on the distal margin, but decrease in size laterally. The proximal border is smooth. The total number of teeth is generally about 12–13. The suckers of the median rows on arms IV are the largest arm suckers, and the marginal suckers of arms I–III are the smallest. The marginal suckers of arms IV are intermediate in size.

The tentacles are robust and have small (approximately 20–25% of the P.L.), complex clubs. The manus possesses the 4 sucker-bearing zones typical of the genus: a ventral-marginal zone that extends the length of the manus and terminates behind the large central hook; a locking zone along the proximal half of the dorsal margin; a dorsal-marginal zone which is continuous with the dactylus distally and proximally passes medial to the locking zone; and, finally, a median zone consisting of a single row of suckers, hooks, or both. The ventral-marginal zone in its central part has 4, rarely 5, suckers that arise from each trabecula and are arranged generally in 4 longitudinal rows. The suckers in the medial row are smaller, but not less than half the diameter of the large suckers in the marginal row. Proximally, the rows converge to a single one which is continuous with the sucker row that extends along the ventral margin of the tentacular stalk. The largest suckers of the ventral marginal zone have 10–12 long, pointed teeth distally, which diminish in size laterally. The teeth are more broadly spaced than are those of the arm suckers.

The locking zone consists of 4–6 thick ridges, each with a large smooth-ringed sucker placed at its proximal end. There is usually a sucker at the distal end of the zone without the adjoining ridge. Alternating with the suckers are large rounded knobs. Proximally, this zone ends abruptly and this break is here used to indicate the proximal limit of the club.

The dorsal-marginal zone contains an irregular group of suckers that do not appear to be arranged in definite rows; distally, however, the suckers merge into 3, or occasionally 2, rows which pass lateral to the large central hook and merge with the suckers of the dactylus. Proximally, the number of suckers decreases rapidly and disappears medial to the locking zone.

The median zone bears a large central hook and a proximal row of suckers. These suckers are never transformed into hooks. Usually, there is no member of this zone distal to the large hook; however, occasionally an enlarged sucker lies behind the hook. Sometimes this sucker may bear an enlarged tooth on the distal margin of the inner ring; and, rarely, this is developed into a small, but completely formed hook. If such a hook is present, it is usually on only one club of the specimen. At the proximal end of the dactylus, suckers are irregularly arranged in 5–6 rows, but quickly decrease in number distally to form 4 regular longitudinal rows. Except for the basal few transverse rows, there is very little difference in size between members of the dorsal and ventral rows. Distally, the dactylus ends in a circle of smooth-ringed suckers at the tip of the club. The suckers from the middle of the dactylus have dentition similar to those of the manus. The total number of suckers on the dactylus (excluding the circle at the tip), the ventral marginal zone, and the dorsal marginal zone varies from 160–200. A well-developed keel is present on the dorsoaboral surface of the dactylus. A narrow, trabeculate protective membrane extends lateral to the suckers along the entire length of the ventral border of the club. On the dorsal margin, a similar membrane terminates in the proximal half of the dactylus.

The tentacular stalk bears a single series of suckers along either margin. Distally, the row along the dorsal margin consists of alternating suckers and pads with adjacent ridges that show faint resemblance to the locking apparatus of the manus; traces of the fleshy ridges accompany only the first few suckers, then disappear. The knobs are displaced medially, relative to the suckers.

The row on the ventral margin is a continuation of the ventral-margin zone on the manus. Most sucker rings in this series. have nearly smooth inner rings. The suckers that are situated one-fifth of the tentacle length from the base of the clubs are nearly smooth, but have the distal margin lightly scalloped. Both marginal rows extend almost the entire length of the tentacle. The number of suckers in the ventral row is 70–115% of the number in the dorsal row. A protective membrane, apparently lacking trabeculae, extends along the entire dorsal margin of the tentacular stalk. A trabeculate protective membrane is present distally on the ventral broder. Only a few, generally less than 10 (maximum range is 0–27), suckers are found between the rows on the oral face of the stalk.

The buccal lappets are indistinct. The buccal membrane connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

The animal is covered with numerous reddish brown chromatophores. This species seems to have a iridescent silver sheen over the lateral surfaces of the head and arms.

Hooks begin to develop on the arms between 26–28 mm P.L. and on the tentacular club between probably 17–24 mm. A spindle-shaped liver set obliquetly to the body axis is present in the larvae.

TYPE LOCALITY.—33°19′ N, 118°45′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C. Hancock collections, AHF Cephalopod Type No. 4.

DISCUSSION.—Berry (1912b) described a number of specimens under the name Gonatus fabricii. One of his illustrations (plate LII, figure 4) showing part of a tentacular club probably refers to G. onyx.

This species is unique in generally possessing only a single hook on the tentacular clubs. Occasionally, an enlarged sucker is present distal to the large median hook and sometimes this is transformed into a small hook; however, this rarely happens on both clubs of a single specimen. Suckers proximal to the large hook are never transformed into hooks. Other characters also separate it from some of its close relatives. The low number of suckers (usually less than 10) on the tentacular stalk between the marginal rows separates it from all species except G. berryi. The size at which the hooks develop on the arms and tentacles separates it from all species except G. californiensis. The number of suckers on the tentacular clubs and the number of hooks and suckers on the proximal half of all arms clearly separates it from G. californiensis. (See also, discussion of G. californiensis.) The unique and distinguishing characters listed for each of the other species of Gonatus are found in the following text.

DISTRIBUTION.—Gonatus onyx is the most abundant species in the collections. In the area under survey, its distribution is limited almost exclusively to the San Pedro, Santa Catalina, and San Nicolas basins; out of a total of 1,236 specimens only 7 have been captured outside of these 3 basins. It is apparent that the species seldom penetrates to the area of Guadalupe Island since there is not a single record from this area despite an estimated 76 hours of trawling in appropriate depths. The almost complete absence from all of the most oceanic zones (i.e., 5, 6, 7), however, indicates that this is an “inshore” species which may extend farther south in the more inshore waters which were not sampled south of latitude 32° N.

TABLE 10.—Measurements in mm of Gonatus onyx, new species.

The northern limits of distribution of this species are unknown. There is a probable record by Berry (1912b) from Monterey Bay. I examined several specimens of Gonatus collected by Pearcy from off the Oregon coast which were deposited in the Miami collections. These specimens clearly belong to G. onyx, and it is probable that the G. fabricii listed by Pearcy (1965) belongs to this species, at least in part. Therefore, the known range of G. onyx extends from about latitude 30° N to at least 43° N.

Gonatus berryi Naef, 1923

PLATES 15A; 16A, C–I; 17A–C, L

Gonatus fabricii.—Berry, 1912b, [parts], p.308; pl.52, fig.3; pl.55, fig.2.

Gonatus berryi Naef, 1923, p.245.

DESCRIPTION.—The mantle is muscular, but thin walled. The mantle margin is produced at the ventral locking-cartilages, but only slightly, if at all, dorsally. The mantle terminates in a point posteriorly; the muscular portion ends at the conus of the pen. A muscular cone with a gelatinous core forms a tail that extends posteriorly from the apex of the conus. The tail is disproportionately small in young specimens.

The fins are rounded laterally and have free anterior lobes. Posteriorly the fin becomes attenuate as it extends along the sides of the tail.

The muscular funnel, completely free from the head laterally, reaches to the level of the midpoint of the eyes. The funnel locking-cartilage bears a straight groove which flares slightly posteriorly and opposes a straight ridge, which broadens posteriorly, on the mantle. The dorsal pad of the funnel organ has an inverted V-shape with a blunt or slightly indented apex. The arms are strongly angled laterally and bear median ridges that extend anteriorly to the apex. A small papilla is present near the apex. The ventral pads are egg-shape. A large funnel valve is present.

The head is oval and bears large eyes that occupy almost the entire lateral sides of the head. Each eyelid has a well-developed anterior sinus. There are 3 pairs of nuchal folds; the second fold from the funnel on either side bears the “olfactory” organ. The nuchal cartilage is elongate, rounded at its anterior end, and bears a median ridge containing a central groove, and lateral ridges.

Arms II and III are moderately long and muscular. In specimens greater than 35 mm P.L. they vary between 60–70% of the P.L. Arms I are slightly shorter and vary between 50–60% of the P.L. Arms IV are nearly equal to arms II and III in length, but are much thinner.

The presence of aboral keels on arms I–III is uncertain due to damage to the specimens. Well-developed trabeculate protective membranes are present on arms I–III; they are less developed on arms IV. Armature is in 4 series: on arms I–III the 2 median rows have large hooks and the marginal 2 rows have small suckers; at the arm tips the hooks are replaced by suckers. On arms IV, all 4 rows have only suckers. The number of suckers on the proximal half of each arm IV is approximately 40 at a P.L. of 50 mm and 45 at a P.L. of 119 mm. The average number of hooks and suckers on the proximal half of each of arms I–III is about 25 at a P.L. of 50 mm and 33 at a P.L. of 119 mm. The suckers of the arms all have similar dentition. Long, pointed teeth, which diminish in size laterally, are present on the distal margin of the inner sucker ring. The proximal margin is smooth. The total number of teeth varies approximately from 10–14. The largest arm suckers occur in the median rows of arms IV; these are nearly 1½ times larger than the largest marginal suckers of arms III. The marginal suckers of arms IV are intermediate in size.

The tentacles are robust; the clubs are moderately large, generally between 30–37% of the mantle length. The ventral-marginal zone of the manus typically consists of 3 longitudinal rows in its broadest part. Occasionally, 1–2 small suckers are present which are the only indication of a fourth row. The suckers of the median row (occasionally rows) are minute and not more than one-fourth the diameter of the large suckers in the marginal 2 rows. The distal-most suckers of this zone are also minute. The largest suckers of the ventral-marginal zone have about 8–9 long, sharp teeth on the distal half of the inner ring which are rather broadly spaced. These diminsh in size laterally and are replaced on the proximal margin by small knobs. These suckers are distinctly larger than those of the median rows on arms IV. The locking zone consists of 5–6 broad, fleshy ridges with a large smooth-ringed sucker at the medial end of each. An additional sucker is commonly present at the distal end of the zone.

The dorsal-marginal zone has relatively few suckers. Distally, it continues past the central hook in 1–2 irregular rows and merges with the suckers of the dactylus. Proximally, the row terminates medial to the locking apparatus. The median zone contains a large central hook followed distally by a small hook and preceded proximally by a single series of suckers and hooks. One or two suckers always lie nearest to the large hook and are followed by 2–4 hooks; the largest hook is never the closest one to the large central hook. At the base of the dactylus along the dorsal border, there are 2 rows of large suckers. Between these rows and the ventral margin are scattered a few minute suckers. The number of rows increases distally to 4 regular series that extend to the tip of the dactylus. The suckers of the ventral row are much smaller than the others. This is most noticeable in the proximal portion of the dactylus and in large specimens. The largest suckers in the middle of the dactylus have similar dentition to the suckers of the manus except that the teeth are disporportionately smaller. At the tip of the dactylus, there is a small circle of smooth-ringed suckers. The total number of suckers on the dactylus (excluding the distal circle), the median zone, and the ventral-marginal zone varies from 159–181. A dorsoaboral keel is present on the dactylus. A small, trabeculate protective membrane is present along the entire ventral margin of the club. A similar membrane is present on the dorsal border of the manus, but ends along the base of the dactylus.

A locking apparatus, consisting of a series of suckers and knobs, extends almost the entire length of the tentacular stalk along its dorsal margin. There is only a slight trace of the fleshy ridges characteristic of the locking apparatus of the manus adjacent to the most distal suckers in the series. The knobs are displaced medially from the suckers. Another series of suckers extends, in a single line, along the ventral margin of the tentacular stalk in its distal half; however, the length is somewhat variable since occasionally scattered suckers extend farther down the stalk. Almost all of these suckers bear pointed teeth along the distal margins of their chitinous rings. Suckers in this series located one-fifth of the tentacle length from the base of the club have 4–6 distinct, pointed teeth on the distal margin of the inner rings. The number of suckers along the ventral margin varies from 38–58% of the number along the dorsal margin. The oral surface of the tentacular stalk between the marginal rows is completely free of suckers except for an occasional one or two. A low, protective membrane, apparently without trabeculae, is present along almost the entire dorsal margin of the tentacular stalk. A trabecular protective membrane extends along the distal half of the tentacular stalk on the ventral margin.

The buccal mambrane bears 7 indistinct buccal lappets. The buccal membrane connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

The larva first develops hooks on the arms between 8–10 mm P.L. The large central hook on the club develops at 12–17 mm P.L. The more distal hook develops between P.L. 19–28 mm, while the proximal hooks begin to develop between 25–32 mm. P.L. A spindle-shape, larval liver is never present in this species.

Photophores are absent. Members of this species are covered with numerous, reddish brown chromatophores. There is some evidence, at least in younger specimens, of the presence of iridescent silvery tissue on the sides of the head and arms.

TYPE LOCALITY.—Monterey Bay, California, eastern North Pacific Ocean.

LOCATION OF TYPE.—?In personal collection of S. S. Berry, number 98.

DISCUSSION.—Berry (1912b) described a number of specimens of squid which he identified as Gonatus fabricii. Naef (1923), noticing that one of the young specimens illustrated by Berry had fins that were unusually large for G. fabricii, established for it the new species G. berryi. No further information has been published on this species until now. The large size of the club in the type illustration as well as the illustrated details of the club structure make it very easy to connect this name with the second most abundant species of the genus taken in the present study.

G. berryi is very distinct, and there should be no difficulties in its recognition. Among its unique features are the following:

1. In the median zone of the club there are 1–2 suckers which lie near the large central hook. These are followed by several hooks that generally increase in size as the series proceeds away from the central hook. The usual case is the opposite arrangement.

2. The ventral-marginal zone of the club has 2 marginal rows of large suckers and 1–2 rows of minute suckers medially.

3. Suckers in the ventral row on the dactylus are considerably smaller than those of the dorsal row.

4. The tentacular clubs are 30–37% of the pen length.

5. The ventral-marginal row on the tentacular stalk has only about 30–60% of the number of suckers present in the dorsal-marginal row.

6. The larvae develop hooks on the arms when they have a P.L. of only 8–10 mm.

7. A spindle-shaped larval liver is not present at any stage of the life cycle.

TABLE 11.—Measurements in mm for Gonatus berryi Naef, 1923.

In addition, many features distinguish this species from one or more of its relatives. Among these are the almost complete lack of suckers on the oral face of the tentacular stalk between the marginal rows; toothed suckers along the ventral margin of the tentacular stalk; the relative length of the arms; the arrangement of suckers on the dactylus; the number of suckers on the tentacular club; the number of suckers and hooks on the proximal halves of the arms; and the P.L. at which the hooks on the club develop.

DISTRIBUTION.—This species occurs in all zones except 7. Its frequency of 0.5 specimens/hour of trawling in zone 6 is surprisingly high and compares favorably with its frequency of 0.57 in the Santa Catalina Basin. Its absence from the vicinity of Guadalupe Island, however, is significant and suggests that this area is near the southern limit of the species. The type specimen, which is the only other record of the species, was taken by the ALBATROSS in Monterey Bay. Thus, the known range of this species extends from latitude 30° to 37° N off the coasts of southern California and Baja California.

Gonatus pyros, new species

PLATES 13B; 14B, J–Q; 17G–I, K

DESCRIPTION.—The mantle is slender; in almost all specimens examined, it is contracted and tapers gradually from a flaring margin to a posterior point. The mantle margin projects ventrally in the areas of the mantle locking-cartilages, but not dorsally. The mantle wall is moderately thick and muscular.

The fins are broad. A posterior “tail” is only slightly developed in the largest specimens. Free anterior lobes are present, and the lateral fin margins are rounded. The fins are very muscular, but are almost always damaged.

The funnel is free laterally and extends forward to about the level of the posterior edge of the lens. The funnel locking-cartilage has a long, straight sulcus which opposes a straight ridge on the mantle. The dorsal pad of the funnel organ has roughly the shape of an inverted V; the lateral arms are long and angled laterally. Small, median ridges are present near anterior ends of each of the lateral arms, and a small papilla lies at the blunt anterior apex. The ventral pads are roughly oval in shape.

The head is short and rounded. Its width generally exceeds that of the mantle. The eyes are large, and occupy the entire lateral sides of the head; an anterior sinus is present on each eyelid. Each “olfactory” organ is flaplike and located at the posterodorsal end of the second nuchal fold from the funnel. The 3 nuchal folds are very feeble and can be recognized only in the most perfectly preserved specimens. The nuchal cartilage is elongate and rounded at either end. It bears 3 longitudinal ridges; 2 are near the lateral edges of the cartilage, and one is in the median line. The median ridge has a narrow, central sulcus along its midline.

Arms II and III are moderately long and muscular; their length in specimens larger than 35 mm P.L. varies between 60–70% of the P.L. Arms I are shorter and vary between 50–60% of the P.L. Arms I–III appear to have had membranous or gelatinous swimming keels. Arms IV are nearly the same length at the lateral arms, but are more slender. Thin, broad, lateral keels are present along the entire lengths of arms IV. Arms I–III all have well-developed trabeculate, protective membranes; trabeculae and membranes are less developed on arms IV. The armature of the arms is in 4 series. On arms I–III, the marginal rows consist of small suckers that arise near the ends of the trabeculae; the medial 2 rows are composed of large, alternating hooks which are replaced by suckers at the arm tips. Arms IV have suckers instead of hooks in the medial rows. All of the arm suckers have similar dentition. On the distal part of the inner ring are long, pointed teeth that grade into smaller teeth laterally. The proximal margin is smooth. The total number of teeth in each ring varies roughly from 6–9. The marginal suckers of arms I–III are larger than the medial suckers of arms IV, while the marginal suckers of arms IV are smaller than the medial suckers. The usual number of suckers and hooks on the proximal half of arms I–III varies from 29–33, while on arms IV, in specimens larger than P.L. 34 mm, the half-arm count varies from 36–41.

The tentacles are long and robust. The tentacular clubs are short, expanded, and very complex. The club length varies from 20–25% of the P.L. The locking zone has 4–6 thick transverse lamellae; adjacent to each at the base of the medial end lies a large, smooth-ringed sucker, and alternating with these are rounded knobs. Proximally, this structure is marked off from the tentacular stalk by a large gap. Distally, the thick lamellae abruptly decrease in size; generally, the last two are rudimentary, but have typical suckers. Suckers of the dorsal-marginal zone converge into approximately 3 rows which pass around the central hook to merge with the suckers of the dactylus. Suckers in this group usually are absent at the proximal edge of the club, although occasionally they extend onto the stalk. In the median zone, a single, large hook lies near the center of the club and is followed distally by a hook of intermediate size and proximally by 3–4 small hooks. The distal hook is rarely joined by an enlarged sucker with a large median tooth. The small hooks decrease in size proximally and are occasionally preceded by a proximal series of suckers. The ventral-marginal zone consists of 3 or sometimes 4 longitudinal rows of suckers in its midportion, all of which are nearly the same size. This group of suckers continues onto the tentacular stalk not only as the marginal row, but also as a more medial, sligthly irregular series. Distally, this zone terminates opposite the large central hooks.

The largest suckers from the ventral-marginal zone have large, rather broadly separated, pointed teeth on the distal margin of the inner rings which grade into smaller teeth laterally. The proximal border is smooth. The total number of teeth in each ring is roughly 9–10. These suckers are approximately the same diameter as the suckers from the medial rows on arms IV. The suckers of the middle of the dactylus have similar dentition, but with sligthly fewer teeth. The suckers at the proximal end of the dactylus are numerous and not arranged in distinct rows. Slightly distally, they become aligned in 4 longitudinal rows. The suckers in a transverse series are nearly the same size proximally, but distally the suckers of the dorsal row are slightly smaller than those of the ventral row. The tip of the dactylus has a circlet of small, smooth-ringed suckers. The total number of suckers on the dactylus (excluding the circlet), the dorsal-marginal zone, and the ventral-marginal zone varies from 151–184.

The locking apparatus on the manus continues along the dorsal margin of the tentacular stalk proximal to the gap, but much reduced. The large lamellae on the manus are represented by relatively low rounded lobes which soon disappear; but they are much more apparent than in any of the other 3 local species of Gonatus. The marginal suckers and alternating knobs decrease in size proximally, and extend along, roughly, three-fourths of the stalk. The suckers along the ventral margins continue for about half the length of the stalk. The suckers in the latter series situated one-fifth of the tentacle length from the base of the club have a few broad, slightly pointed teeth on the distal margin of the inner ring. Almost all specimens have an additional series of suckers on the tentacular stalks which are continuous with the suckers of the ventral-marginal zone of the manus. This series closely parallels the suckers of the ventral margin on the stalk. This series is variable, but usually extends about three-fourths of the length of the marginal row, although it is most clearly present in the distal part. These suckers generally are arranged in a single row, an irregular single row, or occasionally an irregular double row. Sometimes, this series lies very close to the marginal row; more frequently, it is separated from the latter by a distinct gap and rarely contains the only suckers on the tentacular stalks that lie medial to the marginal rows. More commonly, additional suckers are scattered about the oral surface of the stalk. The number of suckers in the ventral row varies between 80–110% of those in the dorsal row. The number of suckers on the stalk between the marginal rows varies betwen approximately 50–125.

The thin buccal membrane attaches far up on the bases of the arms. It has 7 lappets and the connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

A single, large, white photophore lies on the ventral surface of each eye. Each photophore is somewhat oval and covers the greater portion of the ventral surface of the eye. It is immediately distinguishable from the silver tissue surrounding the eye by its well-defined boundaries and its peculiar surface which seems to have many small pores.

Although the integument is damaged on all specimens, some indication of its color remains. The sides of the head and the aboral sides of arms II and III and, to a lesser extent, arms I, are covered by a brilliant silvery layer. This layer extends over the ventral surface of the head, but it is not as well developed there. It is not possible to tell whether or not this layer was present on the lateral surfaces of the mantle. All of the head, arms, fins, and mantle (including the silvery layers) appear to have been covered by brownish red chromatophores. The buccal membrane lacks pigmentation except for a few chromatophores on its outer surface.

Hooks on the arms are present at a P.L. of 17–22 mm; the distal hook on the club begins to develop at a P.L. of 18–23 mm; the median hook at a P.L. of 15–18 mm; and the proximal hooks at a P.L. of 21–26 mm. A spindle-shape liver lying oblique to the body axis is present in the larva.

TABLE 12.—Measurements in mm for Gonatus pyros, new species.

TYPE LOCALITY.—33°37′ N, 118°26′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod Type No. 5.

DISCUSSION.—G. pyros is separated from all other members of the genus by a large photophore on the ventral surface of each eye. Another unique feature is the double series of suckers along the ventral margin of each tentacular stalk. Also among the features which separate G. pyros from other species are: size at which the hooks develop; number of suckers and hooks on the proximal half of the arms; number of suckers on the tentacular clubs; and the length of the arms.

DISTRIBUTION.—No specimens of G. pyros were taken south of latitude 32° N, and it is assumed that this was the southern limit of the species during the period covered by this survey. In the Santa Catalina zone G. pyros is moderately abundant; its frequency is 0.47 specimens/hour of trawling. This species is known only from the present trawling program. The northern extension of the species, therefore, coincides with that of the program.

Gonatus californiensis, new species

PLATES 15B; 16B, J–P; 17D–F

DESCRIPTION.—The mantle is long, slender and moderately thick and muscular; it tapers to a point posteriorly. The free edge projects at the ventral, mantle locking-cartilage, but only slightly dorsally. The muscular portion terminates at the conus. Posterior to the conus, a fleshy column continues as a tail which is best developed in large specimens.

The fins have free anterior lobes and are attenuate posteriorly along the sides of the tail.

The funnel is short and reaches to the level of the lenses of the eyes. The funnel locking-cartilage bears a straight groove that opposes a straight ridge on the mantle. The dorsal pad of the funnel organ has an inverted V-shape. The apex is slightly indented. Each arm is angled on its lateral edge and bears a median ridge on the anterior portion. A slender papilla is present near the apex. The funnel is entirely free from the head laterally.

The head is oval. Two large eyes occupy almost the entire lateral surface of the head. A distinct sinus is present on the anterior edge of each eyelid. Three weak nuchal folds are present lateral to the funnel on either side, the second of which bears the “olfactory” organ. The nuchal cartilage is elongate, rounded anteriorly and posteriorly, and has a median ridge with a central groove and marginal ridges.

Arms II and III are muscular and moderately long; generally they are 40–50% of the P.L. Arms IV are approximately the same length as II and III, but are a weaker construction. Arms I are the shortest, being roughly 35–45% of the P.L. Arms I–III bear gelatinous aboral keels. A lateral keel is present on each arm IV and extends the full length of the arm. Well-developed trabeculate protective membranes are present on all arms, but are less developed on arms IV. The armature is in 4 series: arms I–III have 2 medial rows of hooks and 2 marginal rows of small suckers. The suckers arise from the tips of the slender trabeculae. At the arm tips, the hooks are replaced by suckers. Arms IV have 4 rows of suckers only. All arm suckers have similar dentition. Long slender teeth are present on the distal part of the inner ring which grade into smaller teeth on the sides; the proximal margin is smooth. The total number of teeth varies roughly from 9–13. The suckers of the median rows on arms IV are the largest, followed by the marginal suckers of the same arms and finally by the marginal suckers of arms III, but the size differential is not great.

The tentacles are moderately long and slender. The tentacular clubs are small (17–24% of the P.L.). The ventral-marginal zone of the club has 4–5 rows of suckers in its central part. Proxirnally, they blend into a single row which continues onto the tentacular stalk. Suckers of the 3 lateral rows are nearly the same size, while the medial row (or rows) are about half the diameter of the larger suckers. This discrepancy in size only becomes apparent in large specimens and it is somewhat variable in these. The largest suckers from the ventral marginal zone on the manus have roughly 8–10 teeth on the distal two-thirds of the inner ring. The teeth are moderately long and pointed on the distal part of the ring and grade into smaller teeth on the lateral sides. These suckers are clearly smaller than the larger suckers on the arms. The proximal margin appears to be smooth. The suckers of the dactylus have similar dentition, but apparently have slightly fewer teeth. The locking zone has 4–6 large, fleshy ridges with smooth-ringed suckers at their medial ends. Generally a single sucker is present at the distal end of the zone. The suckers alternate with rounded knobs.

The dorsal marginal zone passes lateral to the large median hook in 2–3 irregular rows that blend distally into the suckers of the dactylus. Proxirnally, the suckers are irregularly arranged and pass medial to the locking zone where they decrease in number until all suckers disappear near the base of the manus.

The median zone contains a very large central hook with a moderately large hook immediately distal to this. Proximal to the central hook are usually 3–5 small hooks followed generally by 1–2 suckers. At the base of the dactylus, suckers are arranged in approximately 7–8 irregular rows, but decrease to 4 rows about halfway out on the dactylus. The suckers of the dactylus are nearly equal in size in a transverse series. The dactylus terminates in a small circle of suckers. A swimming keel is present on the dorsoaboral border of the dactylus. A narrow, trabeculate, protective membrane lies lateral to the suckers along the entire ventral margin of the club. A similar dorsal membrane ends partway out on the dactylus. Both membranes extend down the entire length of the stalks, but are barely visible.

A series of alternating suckers and pads extends along most of the length of the tentacular stalk on its dorsal margin. At the distal end, a few suckers are accompanied by small, fleshy ridges similar to those on the locking apparatus of the manus. A single row of suckers is present along the ventral margin of the stalk for most of its length. The number of suckers in the ventral row varies roughly between 90–120% of the number in the dorsal row. Suckers in this series situated one-fifth of the tentacle length from the base of the club have smooth inner rings. Between these 2 rows there are about 40–80 scattered suckers.

Seven indistinct lappets are present on the buccal membrane. The buccal membrane connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

The species has numerous reddish brown chromatophores. There is some indication that in life the lateral sides of the head and arms have an underlying silvery iridescent layer.

TYPE LOCALITY.—33°32′ N, 118°24′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod Type No. 6.

DISCUSSION.—Gonatus berryi and Gonatus pyros have unique features which easily separate them from other members of the genus. G. onyx and G. californiensis are not as easily separated from one another. The arrangement of hooks on the tentacular clubs of the 2 species differs considerably; in order to demonstrate clearly that these differences are not merely types of variation within a single species, they have been correlated with sucker counts on the tentacles, on the proximal halves of the arms, and in the middle of the tentacular stalks (Figures 8–10). The figures also point out several other interesting features of these species. At about a P.L. of 40–45 mm in G. onyx and about 60–65 mm in G. californiensis, the number of suckers on arms IV no longer increase. Therefore, although the arms will more than double in size as the animal continues to grow (the maximum sizes of these species are not known), the number of suckers will remain constant.

FIGURE 8.—Club-sucker counts, excluding median series and circlet at tip of dactylus, for Gonatus onyx (dots) and Gonatus californiensis (triangles).

FIGURE 9.—One-half arm counts of arms IV for Gonatus onyx (dots) and Gonatus Californiensis (triangles).

FIGURE 10.—Sucker counts of the medial suckers, excluding suckers of the dorsal and ventral marginal series, on the tentacular stalks for Gonatus onyx (dots) and Gonatus californiensis (triangles).

A similar situation occurs with the number of suckers on the tentacular clubs. After the sucker buds have all appeared on the clubs between roughly 10–20 mm P.L., no additional suckers are added. Thus, at almost the first appearance of the definitive clubs the number of suckers is fixed. These features of the arms and tentacles indicate that sucker counts, although little used in the past, can provide excellent taxonomic characters, at least in Gonatus, and should be included in all future descriptions of species within this group.

DISTRIBUTION.—Gonatus californiensis is the least abundant member of the genus in the collections, yet it is the only species that has been captured in zone 7 around Guadalupe Island. Eleven specimens have been captured there, giving a catch frequency of 0.15 compared to 0.24 specimens/hour of trawling in zone 2. The indication is that this species, although primarily a northern form, extends farther to the south than any of the other species of Gonatus of this area. Since all known records of Gonatus are primarily from temperate or colder waters, it would be expected that G. calforniensis does not extend too much farther south than recorded here, although, McGowan (1967) recorded larvae identified as Gonatus fabricii as far south as 25° S off Baja California. Recently, the PILLSBURY captured a larval Gonatus (P.L., 17 mm) near the Bay of Panama (7°19′ N, 79°42′ W). This record is extremely surprising and if the specimen belongs to one of the species found off California, as would seem likely, then it can be identified as G. californiensis. The small size of the specimen would suggest that it had been spawned in the vicinity of Panama. I cannot adequately account for this record, but believe it unlikely that the normal range of this species extends so far south. This problem can only be resolved by further trawling.

TABLE 13.—Measurements in mm for Gonatus californiensis, new species.

At present G. californiensis is definitely known to extend from latitutde 34° N to 28° N off the coasts of California and Baja California.

DISCUSSION OF THE GENUS GONATUS.—Within the genus Gonatus, sensu strictu, there was previously considered to be only a single valid species, G. fabricii (Lichtenstein 1818), with a worldwide distribution. The discovery of 4 sympatric species of Gonatus living off southern California suggests that some of the other forms thought to be synonyms of G. fabricii may represent valid species. This is most likely true with G. antarcticus (Lönnberg 1899) from Antarctic waters and G. kamtschatica (Middendorff 1849) from the northwestern Pacific. It is not known whether or not there is more than a single species in the North Atlantic. Until a study of the latter area is undertaken, the true identity of G. fabricii and its accurate characterization must remain in question. Similarly, there is confusion concerning the correct identification of G. kamtschatica and G. antarcticus. Both have been inadequately described. The collections at the Institute of Marine Sciences indicate that at least 2 species are living in Antarctic waters and at least 2 additional species are living in the Bering Sea. Also, Sasaki (1929) described an adult and a juvenile specimen of “G. fabricii” from the northwestern Pacific. From his description and illustrations, it is apparent that Sasaki was dealing with 2 species and that the juvenile, at least, is different from any of the above-mentioned species. The latter form recalls in general appearance a larval gonatid described by Okutani (1966, plate 3: figure 2) called Gonatidae γ. Okutani also considered that these belonged to the same species. Therefore, at least 7 species of Gonatus are present in the north Pacific, 2 in the Antarctic and 1 in the Atlantic, with only those from off southern California adequately characterized.

With this unsatisfactory state of the genus, it is nearly impossible to properly compare the California species with those from other areas; however, G. berryi, G. onyx, and G. pyros each has a number of unique features that allow its separation from other species. Gonatus californiensis is very similar to specimens that I have seen from the North Atlantic and Antarctic oceans. It is with some hesitation that I give this species a new name, since it is not possible to separate it clearly from forms in these latter localities due to lack of sufficient material. A few differences, however, that substantiate its specific ranking have been noted. The clubs of Gonatus californiensis are larger than those of the specimens of G. “fabricii” examined from the North Atlantic, but are considerably smaller than those from specimens captured in the Antarctic. There are also differences in the sizes at which the large hooks develop on the tentacular clubs between specimens of G. “fabricii” and Gonatus californiensis.

FIGURE 11.—Size at which arm hooks develop for species of Gonatus. Empty squares indicate only suckers present. Squares with Xs show intermediate condition (first hint of hook development). Solid squares indicate hooks present including small incompletely formed hooks. Crosshatched rectangles represent estimated size range over which hooks begin to develop.

Genus Gonatopsis Sasaki, 1920

DIAGNOSIS.—Seven teeth present in a transverse series on the radula. Tentacles present only in larvae.

TYPE SPECIES.—Gonatopsis octopedatus Sasaki, 1920.

Gonatopsis borealis Sasaki, 1923

PLATES 18B; 19I–P; 36H

Gonatopsis borealis Sasaki, 1923, p.203, text-fig. p.205; 1929, p.274, pl.30, figs. 3–6.—Okutani, 1966, p.65, pl.II, figs. 3–4.—Pearcy, 1965, p.261.

FIGURE 12.—Size at which the central club hook in species of Gonatus develops.

DESCRIPTION.*—Anteriorly the mantle is cylindrical, but it begins tapering approximately at the level of the anterior margin of the fins and terminates in a blunt point just short of the posterior margin of the fins. The mantle wall is extremely thick and muscular. The free anterior margin of the mantle projects slightly at the funnel and nuchal locking cartilages.

The fins are broad (approximately 65–70% of the mantle length), moderately long (over 40% of mantle length), and muscular. The fins are sharply angled at their lateral extremities and taper posteriorly to form a broadly rounded apex where their posterior margins fuse. The anterior lobes are free.

The funnel is well developed and powerful. The funnel locking-cartilage has a long straight groove which broadens posteriorly and opposes a long ridge that widens posteriorly on the mantle. The dorsal member of the funnel organ is an inverted V-shaped pad with a small anterior papilla and a low, short ridge near the anterior end of each lateral arm. The lateral arms are long and extend onto the end of the funnel retractor muscles. The ventral pads are large and oval. A funnel valve is present.

The head is cylindrical and has a well-defined, inverted U-shape funnel groove. A nuchal crest is present from which arise 4 nuchal folds on either side of the head. The 2 dorsal folds on each side join posteriorly. The second fold from the funnel curves dorsally and bears a small “olfactory” papilla. The ventral-most fold is continuous with the margin of the funnel groove. The eyes are very large and extend nearly the entire length of the head. A large eye sinus is present on the anterior-ventral margin of each eyelid. The nuchal cartilage is large, slightly constricted in the middle of its length and bears a long, straight median ridge with a central groove.

FIGURE 13.—Size at which the distal club hook in species of Gonatus develops.

FIGURE 14.—Size at which the first proximal club hook in species of Gonatus develops.

FIGURE 15.—Pattern of hook development for the four Californian species of Gonatus: C. hook, central club hook; D. hook, distal hook; and P. hooks, proximal hooks.

The arms are long, muscular, and in the order II= III>I>IV. Arms II and III are roughly 40–50% of the mantle in length. Aboral keels are well developed on arms III, but are less apparent on arms II and form only a small ridge on arms I. Arms IV have well-developed lateral keels. All arms have low, trabeculate protective membranes. Each arm I–III has 39–48 hooks arranged in 2 medial rows and 2 marginal rows of small suckers that arise from the trabeculae of the protective membranes. Near the tips of the arms, the hooks are replaced by large suckers. The 4 rows of suckers thus formed extend to the tips of the arms and usually consist of 56–63 suckers. At the base of the arms, proximal to the hooks are 5–10 suckers. Each arm IV has 4 rows of suckers with 2 median rows somewhat enlarged. All suckers have approximately 12–15 sharply pointed teeth on the distal half of the inner chitinous ring. The proximal margins are generally slightly irregular except on the suckers from the tips of arms III, at least, where the proximal border has distinct small knobs. Peculiarly, in the smaller suckers it is the narrow edge of each tooth which faces the exterior while the broad face lies parallel to the adjacent teeth.

Tentacles are absent except in the larvae.

The largest male, although not fully mature, has a long (95 mm), slender penis that reaches nearly to the mantle opening. There is no evidence of hectocotylization. The female has paired oviducts and typical nidamental glands.

There are 7 lappets on the buccal membrane. The buccal membrane connectives attach to the dorsal sides of arms I and II and to the ventral sides of arms III and IV

The color in preservation is provided by numerous very small purple-brown chromatophores.

The juvenile at 34 mm M.L. is very similar in appearance to the adult; the major exceptions are that only suckers are present on the arms and that the fins are proportionally smaller.

In larvae the arms are proportionally shorter, much broader, and bear 4 distinct rows of suckers. Tentacles are present. In the youngest forms they are short and broad, but with pointed tips. The oral face of the tentacle is flattened and bears 3–5 irregular longitudinal rows of suckers. The rows break up proximally and near the base of the tentacle lie 1–3 enlarged suckers. Although differing in the manner in which they arise from the head, the tentacles are very similar to the arms in size and shape.

In the more advanced larva the tentacles are slenderer and much longer than the short arms IV, but only slightly longer than arms III. The enlarged suckers at the base of the club are very prominent. At a later stage, the tentacle is much more slender than any of the arms and is equal in length or slightly shorter than the arms which are roughly subequal at this stage. The enlarged proximal suckers are still prominent. There is considerable variation in the size of the larvae at which these stages of tentacle development occur. At roughly 8–11 mm M.L. the tentacles are lost abruptly.

TYPE LOCALITY.—Kushiro, Hokkaido.

LOCATION OF TYPE.—In Hokkaido Imperial University.

DISCUSSION.—The genus Gonatopsis was established by Sasaki (1920) to include a new species Gonatopsis octopedatus from the western North Pacific. This gonatid was particularly distinctive in its lack of tentacles and the possession of 8–12 rows of minute suckers on the distal one-third of each arm. In 1923 Sasaki described another member of the genus, G. borealis, which also lacked tentacles, but had only 4 rows of suckers at the tips of the arms. Okutani and Nemoto (1964) have described a subspecies of the latter, Gonatopsis borealis makko, from specimens taken from the stomachs of sperm whales in the North Pacific. This form differs from G. b. borealis primarily in having a slenderer mantle, longer arms (two-thirds of the M.L. versus two-fifths of the M.L.), a slightly different arm formula, and smaller fins. These differences are sufficient to indicate that G. b. borealis and G. b. makko should be considered as distinct species and not as subspecies. The distinctness of G. makko is further enhanced by the recent description (Okutani 1966: 66, plate III: figure 1) of a larval squid, thought to belong to this species, which differs greatly in shape from the larvae of G. borealis.

TABLE 14.—Measurements in mm of Gonatopsis borealis Sasaki, 1923.

The present material of G. borealis differs from Sasaki’s description in the slightly smaller size of arms IV and in the slightly greater number of suckers and hooks on the arms. At best, these differences can only be considered to represent geographical variation.

DISTRIBUTION.—Previously, G. borealis has been recorded off California (Rice 1963) from a whale stomach, off Oregon (Pearcy 1965), the Bering Sea, off Kamchatka (55°40′ N, 180°00′ and 49°50′ N, 165°49′ E) (Okutani 1966), and off Japan (Sasaki 1929).

Sasaki’s comments (1929) on G. borealis are worth repeating:

Specimens examined have mostly come from Kushiro, but also include those from Nemuro and Tokachi. These localities cover the eastern coast of Hokkaido strongly influenced by a cold current ‘Oyashio.’ Most of the specimens of Yamaguchi and Shirai referred to were caught on 28th July, 15–30 miles off Kushiro, where the water temperature showed 11.6°C. at the surface and 2.5°C. at the depth of 20 fathoms below the surface. According to the reports of Mr. Yamaguchi and Mr. Shirai who made researches on the habits of the species by angling from the experimental boat “Sangyo-maru” 1922, whenever Ommastrephes sloani pacificus [=Todarodes pacificus] was abundant, Gonatopsis borealis was very rare, the reverse being also the case, or when both the species happened to be met with at the same region of the sea, the latter species was always angled from a much greater depth than the former. All these facts point to the conclusion that Gonatopsis borealis is physiologically adapted to much colder water than Ommastrephes sloani pacificus, which generally inhabits water above 10°C in temperature. [Spelling has been corrected.]

In the present study the frequency of capture of G. borealis in the Santa Catalina Basin was 0.40 specimens/hour of trawling. The adjacent basins (zones 1 and 3) show surprisingly low frequencies (0.06 and 0.05 specimens/hour, respectively). The reason for the discrepancy is uncertain; however, the relatively high frequency of capture in the Santa Catalina Basin, the most intensively studied area, indicates that this species is fairly abundant in the northern sector of the area under study. Only 4 specimens have been taken in zone 6 and only a single one in zone 7 (frequency 0.08 and 0.01 specimens/hour). Although the data is somewhat confusing due to the low frequency of capture in zones 1 and 3, I think that it is safe to assume that zone 7 roughly coincides with the southern limits of this species in the eastern North Pacific.

Family ONYCHOTEUTHIDAE Gray, 1849

DIAGNOSIS.—Arms with biserial suckers. Tentacular clubs with tetraserial armature of which the 2 marginal rows may disappear during growth. Clubs, where known, bear hooks in the medial rows. Buccal connectives attach to the ventral sides of arms IV. Funnel locking-cartilage with a simple, straight groove. Nuchal folds present.

Genus Onychoteuthis Lichtenstein, 1818

DIAGNOSIS.—Tentacular clubs with 2 rows of hooks; marginal suckers absent except in early juveniles. Arm suckers with smooth chitinous rings (teeth lacking). Photophores present on viscera and a single photophore present on the ventral bulbus of each eye.

TYPE SPECIES.—Onychoteuthis bergii Lichtenstein, 1818.

Onychoteuthis borealijaponicus Okada, 1927

PLATES 18A; 19A–G

Onychoteuthis banksii.—Berry, 1912c, p.83, text figs.44–48; 1913, p.77.—?Pfeffer, 1912, p.79, p.85.—Sasaki, 1916, p.90; 1929, p.228, pl.20, fig.12, pl.30, figs. 1–2.

Onychoteuthis borealijaponicus Okada, 1927, p.4.

Onychoteuthis banksi?—Clarke, 1962, p. 424.—Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is long and slender (width 22–27% of length); the anterior half is cylindrical, and the posterior portion tapers to a terminal point. The mantle wall is thick and muscular. The free margin projects in the nuchal region and at the mantle cartilages. The muscular portion of the mantle terminates at the conus. The pen is clearly visible the entire length of the mantle in middorsal line. The rostrum of the pen extends posterior to the conus and is easily visible in ventral view. The rostrum is surrounded by a cone of gelatinous connective tissue

The fins are 54–60% of the M.L. in length and 70–79% of the M.L. in width. The anterior lobes arefree. Posteriorly, the fins terminate at the tip of the mantle.

The funnel is small; it extends to the level of the middle of the eyes. The funnel cartilage is pointed anteriorly, flares slightly posteriorly, and bears a straight median groove. The dorsal pad of the funnel organ has an inverted V-shape. The posterior portion of the lateral limbs broaden abruptly; the anterior portion bears a weak lateral ridge. A low, conical papilla is present at the anterior apex. The ventral pads are very large and oval. The funnel is free from the head laterally and contains a large valve.

The head is relatively small, being equal to or less than the mantle width. The large eyes occupy the entire lateral sides of the head. Each eyelid has a large anterior sinus. Distinct dorsal and ventral “windows” are present on either side of the head. A funnel groove with well-defined margins is present; in the median anterior line the margin turns forward to form an acute angle. There are 9–10 folds on the nuchal crest on either side of the head; the second bears the “olfactory” papilla. The second to fifth or sixth folds are joined posteriorly. The first fold connects with the midportion of the second. The nuchal cartilage is elongate, constricted slightly laterally, and bears a prominent median ridge with a central groove.

The arms are short (longest arms of the largest specimens are 40% of the M.L.) and in the order 2=3=4>1. Aboral keels are well developed on arms II and III, but are present only on the distal halves of arms I. Lateral keels are present on arms IV. Low trabeculate protective membranes are present on all arms. The armature consists of biserial suckers throughout the length of each arm. The inner chitinous rings of the suckers are smooth; the outer rings are reduced. The suckers of arms IV are distinctly smaller than those of the other arms.

The tentacles are short and robust. The tentacular clubs measured to the proximal end of the carpus are 24–27% of the M.L. The club is not clearly divided into manus and dactylus. It bears biserial hooks and no suckers except for a small cluster of 15–16 smooth-ringed suckers at the tip of the club. Marginal rows of small suckers, however, are present on the club in juveniles up to a M.L. of at least 33 mm. The chitinous rings of these suckers are smooth. The first 5 hooks of the ventral row increase gradually in size distally and have symmetrical bases. The following 2–3 hooks increase greatly in size and have asymmetrical bases. The succeeding hooks gradually decrease in size. Each hook distal to the largest hook frequently bears a small knob near the ventral border of its base, although this is somewhat variable. In the dorsal row, the second hook is usually slightly larger than the first. The following hooks gradually decrease in size to the fifth or sixth hook. The succeeding hooks increase slightly in size, then gradually diminish in size. With the possible exception of the first few proximal hooks, the hooks of the dorsal row are much smaller than those of the ventral row. The carpus occupies an oval patch clearly marked off from the surrounding tissue. It contains 8–10 suckers and pads. The tentacular stalk is bare. A protective membrane is present along the full length of the ventral margin of the club. The proximal portion is much more expanded. A low membrane is present along the dorsal border lateral to the 5 proximal hooks. A large dorsoaboral keel is present. An additional separate keel extends along the aboral surface of the tentacular stalk.

The buccal membrane bears 7 lappets and the connectives attach to the dorsal border of arms I and II and to the ventral borders of arms III and IV.

A large photophore is present over the ventral surface of each eye. An oval photophore is present on the ventral surface of the ink sac and an ill-defined photophore is present on the ventral surface of the ink duct just before the duct enters the intestine. Part of the ink sac and the ink duct between the photophores is covered by a band of tissue with an iridescent gold sheen which Okada thought was also photogenic tissue. O. banksii, a closely related form, has a very large and well-defined anal and intestinal photophores with no iridescent tissue between them. The animal is speckled with reddish brown chromatophores.

TYPE LOCALITY.—Tokachi, Hokkaido, Japan.

LOCATION OF TYPE.—In the collection of Hokkaido Imperial University.

DISCUSSION.—In recent years Onychoteuthis banksii has been considered the only valid member of its genus. Throughout its worldwide distribution it can easily be recognized by its robust appearance, large hooks on the tentacular clubs, a large photophore under each eye, and 2 photophores next to the intestine. In 1927, Okada described a new species, O. borealijaponicus, from Japan based primarily on the slightly different form and structure of the light organs. Sasaki (1929), in his monograph of the Japanese cephalopods, placed this species in the synonymy of O. banksii indicating that Okada’s distinguishing characters might be a result of developmental or seasonal differences. I have examined specimens from 12–100 mm M.L. and find that the appearance of the photophores can easily separate the 2 species throughout this size range. O. banksii also is much more robust than is O. borealijaponicus.

I have attempted to find additional characters to separate the 2 species, but have run into difficulties. The “O. banksii” from Florida waters clearly fall into 2 separate groups and, therefore, the former worldwide species “O. banksii” now seems to consist of at least three species: O. borealijaponicus from the North Pacific and 2 species from Florida waters. It seems likely that “O. banksii’ will eventually be found to consist of a number of additional species, and until this complex can be worked out it will be difficult to completely characterize O. borealijaponicus. O. borealijaponicus, however, can be separated from the Florida specimens, by the light organs, and by the tentacular clubs. I have counted the number of hooks on 37 specimens of “O. banksii” from off the coast of Florida and the results are as follows:

Number of club hooks

19

20

21

22

23

Number of clubs examined

4

28

28

13

1

Voss (personal communication) has examined the type of O. banksii and finds that each club carries 21 hooks. The number of hooks on the clubs of the relatively few large specimens of O. borealijaponicus available to me clearly fall outside of this range:

Number of club hooks

25

26

27

Number of clubs examined

2

8

4

In addition, the relative sizes of the fins, arms, and tentacular clubs separate O. borealijaponicus from each of the Florida species, but in different ways.

DISTRIBUTION.—O. borealijaponicus was taken throughout the area under study, although most records (72 of 95 specimens) came from zones 5 and 6. This species is a powerful swimmer and presumably epipelagic, and is thus ill-suited for capture by the Isaacs-Kidd mid-water trawl. The collection consists primarily of larval forms which probably are taken incidentally when the trawl is being lowered and retrieved. Considering this method of collection, the results are rather difficult to interpret, but they seem to indicate that the young, at least, are more abundant in the more oceanic zones of the study.

TABLE 15.—Measurements in mm of Onychoteuthis borealijaponicus Okada, 1927.

Pearcy (1965) recorded O. banksii from off Oregon. I have examined one of his specimens now at the Institute of Marine and Atmospheric Sciences, Miami, and find that it can be referred to O. borealijaponicus. O. borealijaponicus is also known from off the western coast of Canada (Roper, personal communication) and the eastern coast of Hokkaido, Japan. Other records of O. banksii from the high latitudes in the North Pacific (Clarke 1966) probably belong to this species. I examined a specimen of a more typical “O. banksii” from 16° N, 123° W, and Berry (1914) reported the more typical form from the Hawaiian Islands. It appears that O. borealijaponicus replaces “O. banksii” in the colder waters of the North Pacific.

Family MASTIGOTEUTHIDAE Verrill, 1881

DIAGNOSIS.—Arms with biserial suckers. Tentacles whiplike, with long unexpanded clubs bearing many thousands of minute suckers. Funnel locking-cartilage with an oval depression, usually bearing 1 or 2 knobs (tragus, antitragus). Buccal connectives attach to the ventral borders of arms IV. Small light organs generally present on the surface of the mantle, head, and arms.

Genus Mastigoteuthis Verrill, 1881

DIAGNOSIS.—Cartilaginous tubercules not present on surface of mantle, head, funnel, and arms.

TYPE SPECIES.—Mastigoteuthis agassizii. Verrill, 1881b.

Mastigoteuthis pyrodes, new species

PLATES 25; 26I–Q

DESCRIPTION.—The mantle is cylindrical for most of its length, but posteriorly it tapers rapidly, and the muscular portion terminates in a blunt point just posterior to the anterior end of the fins. The mantle continues past this point as a narrow cylindrical cord of tissue which merges into the tail. The tail extends for a short distance posterior to the fins, but its exact extent is uncertain since it is invariably damaged. The pen appears to extend to the tip of the tail. Anteriorly, the mantle projects strongly in the nuchal region, but less so in the areas of the mantle locking-cartilages.

The fins are large, about 60% (excluding the tail) of the M.L., and nearly circular in outline. The anterior lobes are attached for the most part, but project anteriorly very slightly.

The funnel reaches approximately to the level of the posterior edges of the lenses. It contains a large valve and a well-developed funnel organ with an inverted V-shaped dorsal pad bearing a small anterior papilla, and 2 oval ventral pads. The funnel locking-cartilage has an oval depresion with a strong tragus on the medial margin. Occasionally, a low antitragus can be detected; however, it is usually not apparent.

The head is slightly broader than long. Its width is nearly equal to the M.W., and its length is approximately 20% of the M.L. The eyes are very large, occupying almost the full length and depth of the head. An eye sinus is not apparent on the eyelid. A crescentshape patch of tissue borders the posterior margin of the eyelid, and a small “olfactory” papilla is located posterior to each eye.

The arms are large, the ventral pair extremely so, being comparable to the mantle in length. The arm formula is IV>II>III>I. Although the arms are generally damaged, they appear to contain rather feeble protective membranes supported by flabby, triangular trabeculae. Arms IV have large lateral keels. It is not possible to determine the presence or absence of swimming keels on the other arms due to damage. The suckers are arranged biserially and have slender stalks arising from large conelike bases which are fused with the lateral trabeculae. The oral face of each arm IV is very narrow and the suckers are forced nearly into a single row, particularly in the distal half of the arm. The number of suckers in the proximal half of each arm in the largest female (165 mm M.L.) is 36, 36, 39, and 38 for arms I through IV, respectively. Each sucker ring bears sharply pointed teeth on the distal margin and truncate teeth on the lateral margins. The total number of teeth varies roughly between 10 and 15. The proximal edge of the chitinous ring is smooth. There is no evidence of hectocotylization in males.

The tentacles are very long, over twice the mantle length, and very slender. The club is unexpanded and approximately 50% of the tentacle length. The club possesses a microscopic protective membrane with tiny trabeculae which defines the borders of the sucker-bearing portion of the club. The proximal end of the club is damaged in all specimens, so its exact structure cannot be determined with certainty; however, it appears that the sucker-bearing area tapers rapidly to an abrupt end. Distally, the suckers progressively cover increasingly more of the club surface until they completely surround the tentacle a short distance from its tip. The smallest suckers occur at the tip of the club. The largest suckers occur in a broad area about one-third of the club length from its distal end and are arranged in approximately 18–22 rows. The suckers have long, slender stalks which result in a disorderly arrangement in preservation and makes an exact determination of the number of rows difficult. The club suckers are elongate in the proximal-distal plane. The outer chitinous ring is broad and each lateral margin of its inner edge has 2–3 greatly enlarged, blunt teeth that project into the aperature of the sucker. The inner ring has 3 distinct blunt teeth on the distal margin; the proximal and lateral margins are smooth. The largest suckers on the club measure 0.28–0.32 mm in the long diameter of the holotype (110 mm M.L.). The club in its middle portion has a diameter about 7 times the width of the largest suckers of this portion.

The numerous photophores present are almost exclusively in the outer integument and, therefore, are easily lost with only a small amount of abrasion. As a result of such injury to the present specimens, it is impossible to determine the exact distribution of light organs. The ventral surface of the mantle is covered with small photophores which appear to diminish in number laterally. On the ventral surface of each fin, adjacent to the mantle, photophores are lacking, while lateral to this area scattered light organs appear, but gradually diminish toward the lateral margins of the fins. Thus, there appears to be a somewhat narrow patch of photophores down the ventral surface of each fin. Photophores are abundant on the dorsal surface of the fins, particularly on the lateral halves. Both the dorsal and ventral surfaces of the funnel are sprinkled with photophores as are the ventral and lateral surfaces of the head. A large light organ is present deep in the tissue near the anteroventral margin of each eyelid. It is oval in shape and may be completely hidden by an aggregation of dark chromatophores. Numerous light organs are present on the ventral surface of arms IV.

The buccal membrane attaches to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

The color in preservation consists of a reddish purple background pigmentation upon which dark reddish purple chromatophores occur in the outer integument.

TYPE LOCALITY.—33°32′ N, 118°23′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California, U.S.C Hancock collections, AHF Cephalopod Type No. 7.

DISCUSSION.—It is often impossible to identify with certainty many species of Mastigoteuthis because of several factors. These animals are soft and apparently inhabit a rather deep zone in the ocean. As a result, specimens are usually brought to the surface badly damaged; their long tentacles are often lost along with the delicate epidermis that frequently (if not always) is covered with photophores. Although these squid apparently are common inhabitants of the deep sea, only rarely is more than 1 or 2 specimens taken at a time. These difficulties, compounded by the large number of species within the genus, have led to the present systematic jumble. Many species are known only from a single mutilated specimen, and most species are inadequately described.

It is with some hesitation that I add a new name to a group that is in such a state of instability; however, there appears to be no other course available.

In the following brief review of the 15 nominal species, pertinent characters are listed only when they are derived from the type material.

Mastigoteuthis agassizii Verrill, 1881b. Club suckers are armed with 2–3 sharp teeth on one side. Arm suckers have smooth inner rings. Fins are 60–65% of the M.L. in length. The arm formula is IV>III=II>I. At the proximal end of the club, the suckers are in 2 rows. Light organs are distributed over the entire body. Tragus and antitragus are both weakly developed. Type locality: Off east coast of United States (about 34° N; 76° W). I examined the smaller of Verrill’s syntypes which unfortunately lacks tentacles, and find that there is an enlarged photophore imbedded in the anteroventral margin of each eyelid. This species can be easily separated from M. pyrodes by the lack of dentition on its arm suckers.

Mastigoteuthis grimaldii (Joubin 1895). Tentacles are not known. Arm suckers have 13 pointed teeth on the distal margin of the inner ring. The fins are 54% of the M.L. in length. The arm formula is IV>III=II>I. Light organs are distributed over the entire body. No antitragus is present. Type locality: Near Azores (39°43′ N; 33°22′ W). This species was described from a juvenile specimen that had a M.L. of approximately 26 mm. Due to its lack of described characters, it cannot be separated with any certainty from 5 of the other species of the genus and, therefore, must be considered a species dubia.

Mastigoteuthis levimana Lönnberg, 1896. Tentacles are not known. Arm suckers have broad, truncate teeth around the entire circumference of the inner ring. The fins are 38% of the M.L. The arm formula is IV>III=II>I. Arms IV are more slender than others and almost whiplike. Suckers are present only on the proximal portions of arms IV and are arranged in intervals of 2 to 4 mm. Type locality: North Atlantic (43° N, 24° W). The fragile structure of the ventral arms with their peculiar arrangement of suckers, the small fins, and the sucker dentition, indicate that this species does not belong in Mastigoteuthis and is here placed in the genus Valbyteuthis (Chiroteuthidae).

Mastigoteuthis dentata Hoyle, 1904. Tentacles are not known. The arm suckers have 13 teeth on distal half of the inner ring. Fins are 43–67% of the M.L. in length. The arm formula is IV>III=II>I. Type locality: Off Cape Mala, Panama (7°21′ N, 79°35′ W), and east of the Galápagos Islands (0°36′ S, 86°46′ W). As can be seen from the preceding diagnosis, Hoyle gave very few characters of specific value in the original description. Also, judging by the great discrepancy in fin length between his 2 specimens, it appears that he probably had 2 species. In order to clarify this situation, I here designate his syntype from off Cape Mala (number 7957, male, 72 mm M.L.) as the lectotype. I examined this specimen which is deposited in the United States National Museum (according to C. F. E. Roper, Associate Curator of Mollusks, United States National Museum, the other syntype has been lost). Although the animal has undergone complete desiccation, it is still possible to determine from bits of integument still clinging to the animal that the entire body is covered with photophores and that an enlarged photophore is imbedded in the anteroventral margin of each eyelid.

Recently the PILLSBURY captured 2 specimens from off Cape Mala which agree in all discernible features with M. dentata. From these specimens, which unfortunately were badly damaged, I can add the following characters to the description of this species: the clubs, although badly scraped in places, appear to occupy about half of the tentacular length. The largest suckers occur about one-third of the club length from the distal end. This area of the club bears suckers in roughly 35–40 rows. Suckers near the base of the club are smaller and less thickly spaced, but their precise arrangement cannot be determined. Suckers also decrease in size near the lateral margins of the club. The inner chitinous rings of the club suckers are smooth (Plate 26T). The inner edge of the outer ring has 12–14 small knobs around its entire margin that project slightly into the aperature of the inner ring (Plate 26S). The club suckers are very small. The long diameter across the outer ring of the largest club suckers from a specimen with P.L. of 92 mm was from 0.13–0.16 mm. A minute trabeculate protective membrane is present. The funnel locking-cartilage has a tragus and only a weakly developed antitragus, although the posterior end of the cartilage is well undercut. The 2 specimens had fin lengths of 66% and 67% of the M.L.

I had originally thought that the specimens of Mastigoteuthis from southern California belonged to this species; however, after examining the specimens from Panama, there can be no doubt that the 2 forms belong to separate species. The much smaller size of the tentacular suckers as well as the differences in the dentition of the inner and outer chitinous rings clearly verifies their separate status.Mastigoteuthis talismani (Fischer and Joubin 1906). Arm suckers are smooth (?). The fin is 89% of the M.L. in length. Photophores are present on the fins and probably over the entire body. There is no distinct antitragus. Type locality: South of Azores (34°46′ N, 36°11′ W). The original and only specimen of this species was very badly multilated. The club length is given as about one-tenth of the tentacle length; however, this is almost certainly a misinterpretation resulting from the poor condition of the animal and should not be considered a specific character. The larger size of the fins easily separates this form from M. pyrodes.

Mastigoteuthis glaukopis Chun, 1908. The club is seven-tenths of the tentacle length. Club suckers have 10–12 small teeth around the margin of the inner ring. Club suckers measure 0.1 mm in a specimen with a M.L. 37 mm. Arm suckers have 5–7 large teeth on the distal portion of the inner ring and somewhat smaller teeth on the proximal side. The arm formula is IV>II>III>I. The fins are 50% of the M.L. in length. A single photophore is imbedded in the anteroventral border of each eyelid. Other photophores are apparently absent. No distinct antitragus is present. Type locality: Region of the east African coast. Due to the small size of Chun’s specimen, it is not certain whether the dentition of the club suckers refers to the inner or outer ring. The dentition of the arm and club suckers, fin size, and possibly some of the other characters separates this species from M. pyrodes.

Mastigoteuthis flammea Chun, 1908. Tentacles are not known. Arm suckers have 3–5 teeth on distal margin of inner ring. The arm formula is: IV>II>III≥I. Tragus and antitragus are present. Photophores are distributed over the entire body. Eyes are reduced in size. Fins are 50% of the M.L. Type locality: Gulf of Guinea (0°25′ N, 7°00′ W; 1°14′ N, 2° 10′ W. This species can easily be distinguished from all others now known by the small size of the eyes.

Mastigoteuthis cordiformis Chun, 1908. The club is two-thirds of the tentacle length. Club suckers have numerous pointed teeth around the entire margin of the inner ring with those of the distal border being much larger. Diameter of the suckers is 0.5 mm in a specimen of M.L. 83 mm. Arm suckers have about 30, very small, rounded teeth on the distal two-thirds of the inner ring. The fins are 78% of the M.L. in length. A tragus is present and an antitragus is present, but small. The arm formula is: IV>II>III = I. Proximal suckers of the club are arranged in 2 rows. The middle portion of the club has approximately 25 rows of suckers. The whole body is covered with small light organs. Type locality: South of Sumatra (0°15′ N, 98°8′ E). This species is easily separated from M. pyrodes by its large fins and by the dentition of the club suckers. Descriptions by Chun (1910), Sasaki (1929), Adam (1954), and Voss (1963) indicates that this species exhibits considerable variation.

Mastigoteuthis famelica (Berry 1909). Tentacles are not known. Arm suckers are “minutely toothed.” The arm formula is: IV>II>III≥I. Fins are longer than broad. Type locality: Vicinity of Kauai Island, Hawaii. This species, based on a single specimen of 39 mm M.L., is inadequately known. It differs from California specimens of the same mantle length by having much shorter arms (each arm IV in M. famelica is 20 mm; in M. pyrodes of the same size each arm IV is 35 mm). A redescription of M. famelica is badly needed; unfortunately, the type, which was deposited in the United States National Museum, has been destroyed (Roper, personal communication). Therefore, the further elucidation of this species depends on the capture of new material from the Hawaiian Islands.

Mastigoteuthis magna Joubin, 1913. The club is nine-tenths of the tentacle length. Club suckers are so small that they are invisible to the naked eye. The inner rings of the club suckers are smooth. Arm suckers have smooth inner rings. The fin is 65% of the M.L. The arm formula is IV>II>III=I. The funnel locking–cartilage bears a straight groove anteriorly that enters a bowl–shape depression posteriorly. Photophores are apparently absent. Type locality: Middle of North Atlantic Ocean (31°45′ N, 42°39′ W). This species is very distinctive and can easily be separated from all other species by, among other things, its peculiar funnel locking–cartilage.

Mastigoteuthis hjorti Chun, 1913. Tentacles have suckers of 0.25 mm diameter at the proximal end of the club in a specimen with a mantle length of 95 mm. Tentacular suckers are arranged in 18–20 rows. Arm suckers have 9–10 teeth on the distal half of the inner rings. Fins are equal to the M.L. The arm formula is IV>III>II>I. Two large photophores attached to the bulbus of each eye. Type locality: North Atlantic Ocean (31°24′ N, 34°47′ W). This species is easily separated from all others by its enormous fins and by the presence of photophores on the eyes.

Mastigoteuthis latipinna (Sasaki 1916). The club is three–fifths of the tentacle length. The proximal part of the club is slightly flattened and has broad protective membranes with trabeculae. The proximal suckers of the club are equal to large arm suckers in size and bear 10 blunt, minute teeth on the distal margin of the inner rings. Distally, the suckers become smaller and more extensively toothed. The arm suckers have smooth inner rings. The arm formula is IV>II>III>I. The tragus and antitragus are very faint. The fins are 84% of the M.L. The left eye is much the larger. Type locality: Sagami Sea. This species is very distinctive and can be distinguished from M. pyrodes by almost any of the characters mentioned above.

Mastigoteuthis schmidti Degner, 1925. The club is three-fifths of the tentacular length. The club suckers have smooth inner rings and lack inward projecting knobs on the outer rings. Arm suckers have 13–14 slender teeth on the distal border and 11–12 blunt teeth on the proximal border. The arm formula is IV>III=II>I. The fin is 65% of the M.L. Tragus and antitragus are present. Photophores are distributed over the entire body. From 20–23 suckers are present in transverse series in the distal third of the club. Type locality: Eastern North Atlantic (46°30′ N, 7°00′ W). This species can be easily separated from M. pyrodes by the dentition of the inner and outer rings of the tentacular suckers.

Mastigoteuthis atlantica Joubin, 1933. Tentacular suckers have 2 or 3 elongate knobs that project inward from the inner edge of the chitinous ring. The inner chitinous ring is apparently smooth. Sucker diameter is 0.36 mm in a specimen with a mantle length of 112 mm. Arm suckers have about 11 sharply pointed teeth on the distal border of the inner ring; proximal borders are smooth except for a possible cleft in the median proximal line. The arm formula is IV>II>III>I. A photophore is embedded in the anteroventral margin of each eyelid. No antitragus is present. Fin is 71% of the mantle length. Type locality: Near Bay of Biscay (46°28′ N, 8°01′ W). Joubin′s measurements state that the club is 31% of the tentacle length; however, his illustration indicates that it is much longer. Joubin originally placed this form as a subspecies of M. glaukopis, owing to the presence of a large photophore embedded in each eyelid in both forms. It is now apparent that this feature is widespread throughout the genus and that M. glaukopis atlantica cannot remain at the subspecific level; it is here elevated to specific status. The differences in sucker dentition and possibly fin size are sufficient to verify that M. atlantica and M. glaukopis require separate specific status. This species is very close to M. pyrodes, but differs primarily in the dentition of the inner rings of the tentacular suckers and in its locality.

Mastigoteuthis iselini MacDonald and Clench, 1934. Tentacles are not known. Arm suckers have 13 pointed teeth on the distal margin of the inner ring which are replaced by 6 blunt teeth on each side of the ring. The median proximal portion is slightly crenulated. There is no antitragus. Fins are 67% of the mantle length. The arm formula is IV>III≥II>I.

TABLE 16.—Measurements in mm for Mastigoteuthis pyrodes, new species.

Type locality: 39°04′ N, 71°29′ W. This form is inadequately described and cannot be distinguished with certainty from M. atlantica, M. dentata or M. schmidti and must be considered a species dubia.

DISTRIBUTION.—M. pyrodes is known only from the present trawling program. One specimen each was taken from zones 1 and 2, none from zones 3 and 4, 4 from zone 5, 5 from zone 6, and 16 from zone 7. Apparently this species is more abundant in the offshore zones and, in particular, the more southern zones.

Family CHIROTEUTHIDAE Gray, 1849

DIAGNOSIS.—Arms with biserial suckers except occasionally at the arm tips. Tentacular clubs with tetraserial suckers. Funnel locking–cartilage oval and with 1 or 2 distinct knobs (tragus, antitragus) projecting toward the center of the cavity. Buccal connectives attach to the ventral borders of arms IV.

Genus Chiroteuthis Orbigny, 1845

DIAGNOSIS.—Arms IV greatly enlarged. Photophores present at least on eyes and arms IV. Funnel locking-cartilage generally with both tragus and antitragus. Tentacles greatly elongated and with secondary clubs (larval clubs) lost.

TYPE SPECIES.—Loligopsis veranyi Férussac, 1835.

Chiroteuthis calyx, new species

PLATES 20; 21A, B; 22A–K, N–Q

Chiroteuthis veranyi.—Berry, 1963, p. 128, text figs. 1–6.— Pearcy, 1965, p.262.

DESCRIPTION.—The mantle is short and conical; its musculature is weak and terminates in a rounded point just past the anterior end of the fins. The pen continues posterior to the fins, but has been broken off in all specimens so that its full length is unknown. The mantle margin protrudes in the middorsal line and very slightly near the mantle locking–cartilages.

The fins are nearly circular in outline, 55–65% of the M.L. in length, and have free anterior lobes. On the posterior margin of the fins, where the pen emerges, there is a slight indentation of the musculature covered by a transparent membrane.

The funnel is moderate in size, and is free from the head laterally. The funnel locking–cartilage is oval and bears both a tragus and an antitragus. The dorsal pad of the funnel organ has an inverted V-shape with short, wide diverging arms and a prominant anterior papilla. The ventral pads are oval. A small, delicate valve is present.

The head is elongate and cylindrical, although swollen slightly in the region of the eyes. Nuchal folds are lacking. The “olfactory” papillae which arise from the ventral surface of the head lateral to the exhalant opening of the funnel have long stalks and a bumpy, terminal swelling. Each eye is large and the eyelid bears only a slight indication of an anterior sinus. The posterior margin of the eyelid consists of a large muscular patch of tissue.

The arms are very large and in the order of IV>III>II>I. Arms IV are extremely long, over 1½ times the mantle length, and broad. Arms I–III have low protective membranes supported by fleshy trabeculae which fuse with the broad bases of the sucker stalks. Protective membranes and trabeculae are absent from arms IV. Arms I–III possess thick, low, keels. Arms IV have usually broad, lateral keels which exceed the arm diameter in width. The suckers on each arms I–III are biserial except at the arm tip. The arm tip (i.e., approximately one-tenth of the M.L. on each arm) bears small, closely packed suckers in 3 longitudinal rows. The suckers on each arm IV are broadly separated and arranged in 2 alternating series which tend to converge distally into nearly a single line at the arm tip. Very large globular suckers occur on the middle third of arms I and II, but are somewhat more distally situated on arms III. The suckers on arms IV are largest at the base of the arms, then gradually diminish in size distally.

The enlarged suckers of arms I–III have nearly smooth rings, but may show regular indentations in the margin indicating that they have been formed by the fusion of broad, truncate teeth. On the arm proximal to the enlarged suckers, the inner rings have roughly 18–20 broad, truncate teeth which are separate from one another on the distal two-thirds of the sucker margin; the proximal one-third is smooth. On the arm distal to the enlarged suckers, individual teeth on the suckers also can be recognized. Near the tip of the arms, the suckers have relatively longer and slenderer truncate teeth on the distal margins and nearly smooth but slightly irregular proximal margins. Distinct truncate teeth also occur on all suckers of the ventral arms. The largest suckers of arms III are more than twice the diameter of the basal suckers of arms IV and are more than 3 times the diameter of the adjacent suckers on arms IV.

The tentacles are extremely long, as much as 5–6 times the mantle length; however, their length depends greatly on their state of contraction at the time of fixation. The right and left tentacles in one specimen at hand differ in length by 10 cm, while those of the holotype differ by 6 cm. Thick oval pads are loosely adherent along the length of the tentacle stalks, except near the club where they become embedded in the stalk and decrease in size. It is impossible to get an accurate count of their numbers since many are invariably lost during capture and preservation. It is estimated, however, that there are over 50 of these organs on each tentacle in specimens of about 60 mm M.L.

The club is long (55–65% of the M.L.) and bordered by protective membranes with large broad trabeculae. The membrane is divided into a proximal and a distal portion of nearly equal lengths by a slight decrease in width and by a change in the structure of the trabeculae. Each proximal trabecula is split into 2 or 3 parts in its outer half, except for the small trabeculae in the initial 10% of the club. Each proximal trabecula is separated by a small gap from its neighbors. The trabeculae of the distal portion of the club are undivided and broadly separated from one another at their bases. The club is not differentiated into a dactylus, manus, and carpus. The suckers of the club are arranged in 4 longitudinal rows. Each sucker is set on a long stalk composed of 2 portions. A rather broad cylindrical basal portion terminates in a dark purple, pleated “skirt” from which arises the slender distal portion of the stalk that bears the sucker. The basal portions of the marginal stalks are nearly twice the size of the medial ones and each has a narrow, transparent keel along its lateral edge which runs onto the protective membrane between the trabeculae. The number of suckers varies from 134–142 on the adult clubs in specimens examined. The inner chitinous ring of each sucker bears a large recurved median tooth on its distal margin along with approximately 10–12 smaller lateral teeth. The proximal margin is smooth.

Two large photophores lie embedded in the ink sac, one to either side of the intestine. The ventral surface of each eye is provided with 2 longitudinal strips of luminous tissue which parallel each other on the inner and outer eye margins. Lying between these strips at either end is an oval luminous patch. An additional small luminous organ is situated between the 4 other organs and slightly in front of the posterior patch (Plate 22N). The large pads on the tentacular stalks may be photogenic. The tip of the club is occupied by a very large photophore which faces aborally. Large spherical photophores, each covered by a layer of dark-brown chromatophores, line the lateral-oral edges of arms IV along the base of the lateral membrane.

The buccal membrane attaches to the dorsal borders of arms I and II, and to the ventral borders of arms III and IV.

In preservation C. calyx is speckled with numerous red-brown chromatophores.

Apparently all members of the genus possess a larval stage that attains an unusually large size. C. calyx is no exception; its larval form (it is a “doratopsis” larva) has a maximum size of about 60 mm M.L. This larva has recently been described by Berry (1963), and it is necessary to mention here only the following points. The late doratopsis stage has enlarged suckers on the arms, the largest of which have many closely set, slender, truncate teeth. The full compliment of adult club suckers is present proximal to the larval club. The larval club has approximately 63–65 suckers, tetraserially arranged and set on short stalks. The suckers of the ventral row are considerably larger than those of the other rows. At the tip of the club is a small lobe which carries 5 suckers.

TYPE LOCALITY.—Santa Catalina Basin?, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod Type No. 8.

DISCUSSION.—Presently, there are 8 recognized species belonging to the genus Chiroteuthis in addition to C. calyx. Three species, C. imperator Chun, 1910; C. picteti Joubin, 1894; and C. macrosoma Goodrich, 1896, are easily distinguished from C. calyx by the presence of 3 rows of oval photophores on the ventral surface of each eye (many other characters separate them equally well). C. capensis Voss, 1967, seems more closely related to the above 3 species although the eye photophore pattern, which is imperfectly known, seems to be more similar to the pattern in C. calyx. At any rate, it can easily be separated from the California species by the dentition of the suckers (8–10 long, slender, sharp teeth on the upper edge of the ring and about 20 low, blunt teeth on the lower edge of the sucker ring), by the lack of photophores on the viscera, by the lack of a striped “skirt” on the stalks of the club suckers, and by the different proportional lengths of the arms, mantle, and clubs.

C. atlanticus (MacDonald and Clench 1934) is known only from a single mutilated specimen; however, its elongate mantle clearly indicates its relationship to the above-mentioned species. C. joubini Voss, 1967, can easily be distinguished by the unique photophore pattern on the eye (10 oval photophores in 2 series) and by the presnce of a single, median visceral photophore.

The remaining 2 species, C. lacertosa Verrill, 1881, and C. veranyi (Férussac 1835), are much more closely related to C. calyx than are the other 6 species. C. lacertosa was considered a subspecies of C. veranyi by Pfeffer (1912). These 2 forms are unquestionably very close and even the validity of C. lacertosa still has not been definitely established. The 2 forms differ in the shape of the fins (they are round in C. veranyi, but elongated—60 mm long by 40 mm wide—in C. lacertosa); otherwise, the animals closely agree in most characters. Unfortunately, the holotype of C. lacertosa lacks tentacular clubs; however, if the clubs found by Verrill on other occasions belong to this species, as Verrill believed, then it would appear that the species is more similar to the California species, particularly regarding the arrangement of the lateral trabeculae on the club, and that C. lacertosa is certainly a valid species. It is, however, necessary to examine more material from the type locality of C. lacertosa (i.e., Brown’s Bank off Nova Scotia) before its relationships can be unquestionably resolved. At present, C. calyx can be separated from C. lacertosa by the differences in sucker dentition (sharply pointed teeth in C. lacertosa and truncate or fused teeth in C. calyx, different shape of the fins (circular in C. calyx, elongate in C. lacertosa), and by the greater size of the enlarged suckers on arms I–III in C. calyx.

Of the clearly established species, C. calyx is most closely related to C. veranyi. Relative to the latter species, the suckers in the middle section of arms I–III in C. calyx are considerably enlarged, while the suckers of arms IV are relatively reduced in size. The ratio of the diameter of suckers at the midsection of arms III to adjacent suckers on arms IV is greater than 3:1 in C. calyx, while it is less than 2:1 in C. veranyi (Plate 22B, I, L, M). This difference between the 2 species is large enough that it can be detected at a glance. In C. calyx, the sucker rings have distinctly truncated teeth on the distal margin, which on the largest arm suckers may all fuse giving a completely smooth ring. In C. veranyi, the suckers have many sharply pointed teeth on the distal half of the ring.

TABLE 17.—Measurements in mm for Chiroteuthis calyx, new species.

The final major differences between the 2 species concern the protective membranes of the tentacular club. In C. veranyi, the proximal portion of the membrane is much broader than in C. calyx. Also, in C. veranyi, the individual supporting trabeculae in this membrane can be distinguished only at their bases since they are subdivided for most of their length; these subdivisions are all closely aligned making it impossible to separate the subdivisions of one trabecula from those of its neighbors (Plate 21C). In C. calyx, the trabeculae are subdivided only in their distal half and the trabeculae remain separated from their neighbors by a distinct gap (Plate 21B).

It is worth noting that the arrangement of photophores is identical in C. veranyi, C. lacertosa, and C. calyx.

DISTRIBUTION.—I examined a specimen of C. calyx taken by Pearcy off Oregon at 44°40′ N and 125° 16′ W, and his record of C. veranyi (1965) from Oregon certainly can be referred to this species. The only other record of C. calyx is from off southern California (Berry 1963, as C. veranyi).

In the present study, the frequency of capture was fairly even in zones 1, 2, 3, and 5 (i.e., 0.16–0.26 specimens/hour). The highest frequency of 0.52 specimens/hour occurred in zone 6; however, this high figure is largely the result of a single, unusual trawl capture that yielded 15 larvae. If this trawl was ignored, the catch frquency for zone 6 would be 0.24 specimens/hour. No specimens were captured in zone 7.

Although the data are somewhat contradictory, the complete lack of specimens from zone 7 coupled with the records from Oregon suggests that this is a cold-water species with a southern limit of distribution off Baja California.

Genus Valbyteuthis Joubin, 1931

DIAGNOSIS.—Funnel locking–cartilage oval. Antitragus present, tragus absent. Tentacular club short, compact and with suckers in 4 rows. Club suckers on short stalks. Fins subterminal. Photophores absent. Funnel valve absent.

TYPE SPECIES.—Valbyteuthis dana Joubin, 1931.

Valbyteuthis oligobessa, new species

PLATES 23A, B; 24H–N, Q, R

DESCRIPTION.—The thin, delicate mantle is saclike, except posteriorly where it tapers to a point. The anterior mantle margin is broad and only loosely surrounds the head and funnel.

The fins are small, about 23–33% of the M.L. in length. Each is roughly semicircular and has only slightly projecting anterior and posterior lobes. The broad, tubular conus of the pen projects posteriorly past the fins. It has been broken in all specimens so that its full extent is unknown. The fins join in the dorsal midline.

The funnel is very large; it reaches anteriorly to the level of the anterior margins of the eyes in mature specimens, but is smaller in immature forms. The funnel locking-cartilage is distinctive of the genus; the cartilage has an oval depression that deepens posteriorly where it is overhung by a small antitragus. The opposing mantle cartilage has the appearance of a human nose with a single nostril; the “nostril” locks with the antitragus. The dorsal member of the funnel organ is very small relative to the large size of the funnel. The organ is approximately shield-shape with a rough or papillate surface and an anterior papilla. This papilla, at its base, is slender, but distally it broadens in the transverse plane and is slightly cupped. The ventral pads are somewhat variable, but generally are elongate and oval and slant obliquely toward the midline. There is no funnel valve.

The head is elongate and cylindrical. The eyes are small; in all specimens, they have been damaged. The paired bridles and cephalic vein are prominent along the ventral surface of the head beneath the funnel. “Olfactory” papillae are present lateral to the funnel and considerably posterior to the eyes. Each consists of a very elongate stalk with a terminal swelling. The nuchal cartilage is straight, slender, and elongate. A central ridge with a median groove is barely detectable on its anterior half.

The arms are short, extremely delicate, and in the order IV>III>II>I. In young specimens, arms IV are greatly elongate (up to 130% of the M.L.), but in the mature animal they are less than the length of the mantle and only slightly longer than arms III. There is only a slight indication of protective membranes, and keels appear to be lacking from arms I–III, although lateral keels are present on arms IV. The suckers on the arms are biserially arranged along the entire length of each arm I–III. On arms IV, the suckers are broadly separated and arranged in apparent single file. These are preset only on the proximal end of the arms and number only 2-A on each arm.

The outer chitinous ring of each arm sucker consists of an outer row of parallel scales followed medially by 3–4 rows of scales with generally square bases. The latter bear large knobs which have enlarged, flattened ends in the most medial series of the distal half of the ring. On the proximal side of the ring, the knobs are replaced by hollow cylinders that have a portion of one side missing (Plate 24R). The inner chitinous ring has about 25–35 small, blunt teeth around the distal three-fourths of the margin. The proximal portion is nearly smooth.

The tentacles are relatively robust and long. Each tentacular club is short, only slightly expanded, and is not differentiated into a manus and a dactylus. Low, protective membranes are present on both borders. The suckers are arranged in 4 regular longitudinal rows over most of the club, but become slightly irregular at the tip and at the base of the club. The suckers of the ventral row are the largest. The suckers become progressively smaller in a transverse series, although the suckers of the dorsal row are still only slightly smaller than those of the ventral rows. Five or six suckers, arranged in 2 alternating rows, are present in the carpus. The larger of these is sligthly larger than the suckers of the club.

The buccal membrane connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

No photophores could be found. This animal is covered with numerous reddish brown chromatophores.

The holotype is a gravid female. The mantle cavity is occupied almost entirely by a large mass of mature eggs and large nidamental glands. The ripe eggs are about 1.5 mm in diameter. Several large sperm reservoirs can be found in the enlarged ovary and, surprisingly, one can be seen protruding through each nephridiopore. Paired oviducts are present with enlarged oviducal glands. This is the first record of a mature specimen in the genus.

TABLE 18.—Measurements in mm for Valbyteuthis oligobessa, new species.

TYPE LOCALITY.—32°17′ N, 120°12′ W, eastern North Pacific Ocean.

LOCATION OF TYPE.—University of Southern California. U.S.C. Hancock collections, AHF Cephalopod Type No. 9.

DISCUSSION.—In addition to Valbyteuthis oligobessa there are, presently, only 2 other described species in the genus: V. danae Joubin 1931, from the Pacific Ocean off Panama and V. levimana Lönnberg, 18%, from the North Atlantic; however, at least 2 undescribed species are also present in the Atlantic. It seems that this genus will eventually prove to be rather speciose. V. oligobessa differs from V. danae in a numbr of respects. The most obvious distinguishing feature is the much smaller size of the fins in V. oligobessa (23–33% of the M.L. versus 50–55% of the M.L. in V. danae). Other characters include the number of suckers on the ventral arms (2–4 in V. oligobessa and at least 12–15 in V. danae), the nnumber and shape of teeth on the inner chitous rings of each arm sucker (25–35 narrow teeth in V. oligobessa and 7–10 broad teeth in V. danae), the relative size of the tentacular club suckers compared to the size of the largest suckers of each arm (Plate 24B–F, H–L) and the size of the sucker orifice (Plate 24B–F, H–L). This latter character is particularly noticeable on the distal half of each arm in V. danae where the size of the orifice relative to the size of the sucker rapidly decreases, whereas the sucker orifice in V. oligobessa show very little relative reduction. In addition, the suckers of the ventral row on the tentacular clubs of V. oligobessa are slightly larger than those of the dorsal row. In V. danae both rows have equal-size suckers.

It is more difficult to separate the Californian species from V. levimana, since the latter is known only from the type description which was based on two mutilated specimens. The only definite characters of specific value are the greater number of suckers (6–8) on each arm IV in V. levimana and the dentition of the arm suckers, which is described in V. levimana as consisting of broad, truncate teeth around the entire aperture.

DISTRIBUTION.—V. oligobessa is known only from the present trawling program. Four specimens were taken in zone 2, 1 in zone 4, 5 in zone 5, 3 in zone 6, and 5 in zone 7. If the number of trawling hours in each zone is considered, it is apparent that this species is most abundant in the offshore zones. This distribution probably results from the circumstance that Valbyteuthis is one of the “deepest living” species taken in the program.

Valbyteuthis danae Joubin, 1931

PLATES 23C; 24A-G

Valbyteuthis danae Joubin, 1931, p. 188.—Roper and Young, 1967, p.l.

DESCRIPTION.—The mantle is cylindrical anteriorly, but tapers rapidly posteriorly; the muscular portion terminates at roughly the midpoint of the fins. The pen is broken at the posterior end of the fins, so its full length is unknown. The free margin projects slightly at the nuchal and ventral mantle locks.

The fins are large (52% of the M.L. in length) and muscular. Each is nearly semicircular and has free anterior and posterior lobes.

The funnel reaches to approximately the level of the middle of the eyes. The funnel locking-cartilage has an oval depression and a distinct antitragus. This opposes a cartilage on the mantle that is shaped like a human nose with a single nostril; the latter locks with the antitragus. The dorsal pad of the funnel organ is nearly triangular, but has a slight posterior indentation. The ventral pads are oval and very large. There is no funnel valve.

The head is elongate and broadest posteriorly. The bridles and cephalic vein are prominent on the ventral surface of the head up to the level of the middle of the eyes. The head has been damaged; no other details can be ascertained.

The arms are slender and delicate. The arm formula appears to be IV>III>II>I; however, all arms have been damaged and it is impossible to get accurate measurments. All of the arms have low trabeculate protective membranes and biserial suckers. On arms I—III, the suckers near each arm base have relatively larger aperatures than the more distal suckers. Near the arm tips the suckers become globular and have reduced aperatures and reduced outer chitinous rings. Suckers are present only on the proximal portions of arms IV. These suckers are distinctly biserial and number 12 on the left arm and 13 on the right arm. The inner chitinous rings of the arm suckers have about 7–9 broad, truncate teeth on the distal margin; the proximal margin is smooth.

The tentacles are long and relatively robust; the clubs are short and compact. Suckers are arranged tetraserially and possess broad outer rings and inner rings with narrow aperatures and smooth margins. The club is not divided into a manus and dactylus. The carpus has 6 suckers arranged in 2 alternating rows. A thick protective membrane lines either side of the club.

The buccal connectives attach to the dorsal borders of arms I and II and to the ventral borders of arms III and IV.

TYPE LOCALITY.—Off Panama, 7°30′ N, 79°19′ W.

LOCATION OF TYPE.—Copenhagen Museum.

DISCUSSION.—The above description is based on a single, damaged specimen (, 44 mm M.L.). It agrees in all respects with specimens of V. danae captured at the type locality off Panama. The illustrations used in this report are taken from Roper and Young (1967) and are based on specimens taken from the eastern tropical Pacific Ocean. For comparisons with the other species presently known, see discussion of Valbyteuthis oligobessa.

DISTRIBUTION.—V. danae was previously known only from the Pacific Ocean off Panama at about latitude 7° N and off Peru at about latitude 8° S. Within the Panamanian region, however, this species appears to be fairly common. The single specimen described here was taken in the Santa Catalina Basin (VELERO Station 8120) at 32°20′ N, 118°43′ W. There seems to be little doubt that this specimen is a straggler from farther south.

Family GRIMALDITEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Arms with biserial suckers. Tentacles absent. Buccal connectives attach to the ventral borders of arm IV. Funnel and mantle locking–cartilages fused. Nuchal articulation free.

Genus Grimalditeuthis Joubin, 1898

DIAGNOSIS.—The characters coincide with those of the family.

TYPE SPECIES.—Grimalditeuthis richardi Joubin, 1898.

Grimalditeuthis bomplandii (Vérany, 1837)

PLATES 27B; 24O, P

Loligopsis Bomplandii Vérany, 1837, p.99, pl. 1a.

Chiroteuthis Bonplandii.—Férussac and d’Orbigny, 1839, p.326.—Gray 1849, p.44.

Grimalditeuthis Richardi Joubin, 1898a, p. 101, figs. 1,2; 1899, p.71; 1900, p.79, pl.4, figs. 1–2, pl.5, figs. 1–13, pl. 10, figs. 1–2.

Grimalditeuthis Bonplandii.—Pfeffer, 1900, p.188.—Chun, 1910, p.217; 1913, p.8.

Grimalditeuthis bonplandii.—Voss, 1956, p.151, text fig.13b.

DESCRIPTION.—The soft, gelatinous mantle is slender (width is 16% of M.L.) and tapers gradually toward the posterior end. Near the level of the anterior edges of the fin, the muscular layers of the mantle diminish. The gelatinous portion of the mantle continues as a thick cylindrical mass of tissue which is divided internally into a number of compartments giving the appearance of a honeycomb.

The funnel and mantle locking-cartilages are fused to one another and thus form a permanent union between the mantle and funnel. The mantle is free in the nuchal region.

The fin is large (46% of the M.L. in length and 61% of the M.L. in width) and relatively muscular. It is subcircular in outline. The anterior lobes are almost completely attached. Posterior to the fins, the mantle and pen have been broken off. At this point, the pen is very broad and undoubtedly continues for some distance in the intact animal. Joubin (1900) had an intact specimen in which the pen and mantle continued posteriorly to support a second pair of fins.

The funnel is very small. No trace remains of a funnel organ or funnel valve due to damage.

The head is cylindrical, elongate, and bears a pair of small, lateral eyes. A pair of “olfactory” papillae is present at the posterior edge of the head. Each papilla lies immediately lateral to the funnel and consists of a terminal swollen bulb set on a slender stalk.

The arms are long and slender; arms IV have exceptionally attenuate tips. All arms lack protective membranes, trabeculae, and swimming membranes. Each sucker stalk arises from a large base from which project 3 conical papillae. The largest suckers on arms I–III occur at roughly two-thirds the distance along the arm from its base. On arms IV, the largest suckers are very near the base of the arm and are smaller than the largest suckers of the other arms. Near the arm bases, the suckers are globular and have very small apertures. Distally, the apertures become relatively much larger. The suckers have about 12–13 slender, pointed teeth on the distal two-thirds of the inner chitinous rings. The proximal margin is smooth. The tips of all the arms are bare; however, the length of these tips are longer in the ventral arms than any of the others. The tips show no evidence of being photogenic. A large sucker from the right arm III has 13 sharply pointed teeth.

Tentacles are apparently lacking in this species. They are represented by minute stubs located between arms III and IV.

The buccal membrane has 7 lappets which attach to the dorsal borders of arms I and II, and to the ventral borders of arms III and IV.

The preserved specimen lacks pigmentation except for a few small, scattered, brown chromatophores.

The data for the single specimen from off California are: Sex, ; M.L., 89 mm; M.W., 14 mm; H.L., 23 mm; H.W., 16 mm; F.L., 41 mm; FW., 54 mm; Arm L. for arms I-IV respectively, 54 mm, 59 mm, 56 mm, 54 mm; ½ A.C. for arms I–IV respectively, 40, 40, 42, 39.

TYPE LOCALITY.—29° N, 39° W, North Atlantic Ocean.

LOCATION OF TYPE.—Not traced.

DISCUSSION.—Pfeffer (1912) synonymized the 2 nominal species in the family. Grimalditeuthis bomplandii (Vérany, 1837) and Grimalditeuthis richardi Joubin, 1898a. There has been some doubt concerning the validity of this action, since only G. richardi is known to have photophores at the tips of the arms. Judging from the present specimen, it seems likely that this difference represents merely an ontogenetic change. The VELERO specimen lacks photophores at the tips of the arms, but all of the arm tips are free of suckers. This condition is peculiar, but could be easily explained if one assumed that the arm tips were developing into photophores. I compared the VELERO specimens with Atlantic material, but have not been able to detect any differences.

This species is believed to be without tentacles; however, delicate stubs of tentacles are present on all specimens available to me from both oceans which comprise a rather large size range. This indicates that either extremely weak tentacles are present, but are always lost during capture or, that the stubs represent rudimentary tentacles which persist, probably due to their retention of some function which is perhaps sensory. In this regard, it should be emphasized that in other species that lack tentacles (i.e., Octopoteuthis, Gonatopsis) the stubs are present in young forms, but are quickly lost with growth.

DISTRIBUTION.—This species is know only from a very small number of specimens. It has been reported from the north Atlantic at 29° N, 39° W (Vérany 1837), 34°51′ N, 33°1′ W (Chun 1913), 38°55′ N, 23°39′ W (Joubin 1898a), from the Gulf of Mexico (Voss 1956), and from the South Atlantic at 28° S, 28° W (Pfeffer 1912). The VELERO specimen was taken from the Santa Catalina Basin at 33° 13′ N, 118°28′ W.

Family BATHYTEUTHIDAE Pfeffer, 1900

DIAGNOSIS.—Suckers on arms arranged in 2 rows proximally, increasing to 4 irregular rows distally. Tentacular clubs with minute suckers in approximately 8–10 rows. Minute suckers present on buccal lappets. Fins small and separate with convex posterior margins. Buccal connectives attach to dorsal borders of arms IV.

Genus Bathyteuthis Hoyle, 1885

DIAGNOSIS.—The characters coincide with those of the family.

TYPE SPECIES.—Bathyteuthis abyssicola Hoyle, 1885.

Records of Additional Pelagic Species from off Southern California

A number of other species of pelagic cephalopods occur in the southern California area, but were not included in this study since material was either lacking or inadequate for proper treatment. These species consist of the following: Dosidicus gigas, Moroteuthis robusta, Loligo opalescens, Thelidioteuthis sp., Pterygioteuthis giardi, Argonauta pacifica, Branchioteuthis sp., Taonius sp., Liocranchia sp., and Eledonella sp.

Dosidicus gigas commonly occurs off the western coasts of Central and South America and Mexico. Occasionally, it is found in the waters off California and has been reported as far north as Monterey Bay. In the years 1934–1937, it was particularly abundant in local waters. Croker (1937) in recording its occurrence stated: “The plague of jumbo squid has been worse in 1937. Set line, net and troll commercial fishermen are still bothered by them, and in addition sport fishermen have been harassed all spring. No sooner does a pleasure boat start to fish than a horde of squid appears on the scene to crowd the game fish away and seize all the baits. When one is hooked, it proceeds to shower boat and fishermen with ink and water, and then delights in biting its captor with its parrot-like beak. Several fishermen have been seriously bitten this year. Although squid fishing is hilarious sport for a few minutes, it becomes too much of a good thing day after day.” Dosidicus gigas attains considerable size: 4½ feet (Classic 1949) and a weight of more than 30 pounds (Croker 1937) have been recorded for specimens off California, but much larger specimens have been reported off South America. Specimens of D. gigas can easily be distinguished from other local ommastrephids by their attenuate arm tips bearing many small suckers which increases the total sucker count to well over 200 suckers per arm. This is more than three times the sucker count of other local ommastrephids.

Moroteuthis robusta has been captured off the west coast of north America, along the Aleutian Islands, and off northern Japan. Although this species has been recorded a number of times from California, only one record (Croker 1934) occurs as far south as the present study area. This species attains a mantle length of at least 7½ feet (Berry 1912b: 315). Although the I.K.M.T. (Isaacs-Kidd Midwater Trawl) is perhaps not well suited for capturing specimens of M. robusta, it is somewhat surprising that no larvae were taken. It seems probable that southern California is at the extreme limit of its distribution. M. robusta can easily be distinguished from Onychoteuthis borealijaponicus, the other local onychoteuthid, by the relatively greater length of the arms, by the larger number of hooks on the tentacular clubs (about 36 in M. robusta and 25–27 on O. borealijaponicus), and by the lack of photophores.

Loligo opalescens is probably the most abundant cephalopod species occurring off the California coast. This species is epipelagic and fast moving, and therefore difficult to capture with an I.K.M.T. Over 80 specimens, however, were taken during the present study, but most were larval forms under 10 mm M.L. Fields (1965) did a detailed study of the biology of this species, and nothing could be added from the present collection. L. opalescens is fished commercially off California, primarily at Monterey. Over the past 17 years, the average catch has been approximately 6,500 tons per year; the maximum catch of 19,000 tons occurred in 1946 (Fields 1965).

L. opalescens is the only member of the suborder Myopsida occurring off California. It can be easily separated from all other squid of this area by the presence of a cornea covering the lens of each eye.

The several specimens of Taonius sp. taken during the present program were too badly damaged to treat systematically. Taonius has also been reported from off Oregon (Pearcy 1965). This cranchiid appears very similar to Galiteuthis, but instead of the central club suckers being modified into hooks, each looks like a set of claws (i.e., a number of elongate teeth on each sucker).

Berry (1912b) recorded a small, finned octopod captured off San Diego, California as “? Stauroteuthis sp.” The identity of this form is still uncertain, since no additional material has been taken. Judging, however, from the small size of the fins and the large size of the eyes, it is possible that this is a juvenile specimen of Opisthoteuthis californiana Berry, 1949.

Argonauta pacifica has been reported from off California on a number of occasions (see Berry 1912b). It is not yet certain whether this form is specificially distinct from the Atlantic Argonauta argo. Several larval specimens of the “paper nautilus” are present in the VELERO collections from off Baja California, but can only be referred to Argonauta sp. Specimens of Argonauta can easily be recognized. The tips of the dorsal arms in females are each provided with an extensive flaglike web which is responsible for secreting the paper-thin shell that serves both as a broad chamber and a flotation device. Males are dwarfed and lack the modifications of the dorsal arms; however, the third left arm is hectocotylized, and enrolled in a sac; this feature distinguishes Argonauta from all other cephalopods.

Recntly McGowan (1967) has indicated the presence of Pterygioteuthis giardi in the southern California area. This species is very similar to P. gemmata, but can easily be separated by the presence of biserial hooks in the middle of the arms. Apparently, this warm-water species occurs off southern California only during certain years.

Other species taken in the present study include larval specimens of Liocranchia sp., Thelidioteuthis sp. from zone 7, Brachioteuthis sp. from zone 6, and one damaged specimen of Eledonella sp. from zone 3.

Checklist of Cephalopods Definitely Known to Occur in Southern Californian or Neighboring Waters

Order Sepioidea

Family Sepiolidae

Rossia pacifica Berry, 1911 †

Order Teuthoidea

Suborder Myopsida

Family Loliginidae

Loligo opalescens Berry, 1911

Suborder Oegopsida

Family Enoploteuthidae

Abraliopsis falco, new species*

Abraliopsis felis McGowan and Okutani,1968*

Pyroteuthis addolux, new species *

Pterygioteuthis gemmata Chun, 1908 *

Pterygioteuthis giardi Fischer, 1895

Thelidioteuthis sp.

Family Ommastrephidae

Dosidicus gigas d’Orbigny, 1835

Ommastrephes bartramii (Lesueur, 1821)*

Symplectoteuthis luminosa Sasaki, 1915 *

Family Neoteuthidae

Neoteuthis sp.*

Family Histioteuthidae

Histioteuthis heteropsis (Berry, 1913)*

Histioteuthis dofleini (Pfeffer, 1912)*

Histioteuthis corona berryi Voss, 1969

Family Octopoteuthidae

Octopoteuthis deletron, new species *

Family Gonatidae

Gonatus onyx, new species *

Gonatus berryi Naef, 1923 *

Gonatus pyros, new species *

Gonatus californiensis, new species *

Gonatopsis borealis Sasaki, 1923 *

Family Onychoteuthidae

Onychoteuthis borealijaponicus Okada, 1927 *

Moroteuthis robusta Verrill, 1876

Family Brachioteuthidae

Brachioteuthis sp.

Family Mastigoteuthidae

Mastigoteuthis pyrodes, new species *

Family Chiroteuthidae

Chiroteuthis calyx, new species *

Valbyteuthis oligobessa, new species *

Valbyteuthis danae Joubin, 1931 *

Family Grimalditeuthidae

Grimalditeuthis bomplandii (Vérany, 1837)*

Family Bathyteuthidae

Bathyteuthis berryi Roper, 1968 *

Family Cranchiidae

Cranchia scabra Leach, 1817 *

Leachia dislocata, new species *

Galiteuthis phyllura Berry, 1911 *

Galiteuthis pacifica Robson, 1948 *

Helicocranchia pfefferi Massy, 1907 *

Taonius sp.

Liocranchia sp.

Order Octopoda

Suborder Cirrata

Family Opisthoteuthidae

Opisthoteuthis californiana Berry, 1949

Suborder Incirrata

Family Bolitaendiae

Japetella heathi (Berry, 1911)*

Japetella sp.*

Eledonella sp.

Family Ocythoidae

Ocythoe tuberculata Rafinesque, 1814*

Family Alloposidae

Alloposus mollis Verrill, 1880 *

Family Argonautidae

Argonauta pacifica

Family Octopodiae

Octopus bimaculatus (Verrill, 1883)

Octopus bimaculoides Pickford and McConnaughey, 1949

Octopus leioderma (Berry, 1911)

Octopus californicus (Berry, 1911)

Octopus rubescens Berry, 1953

Octopus micropyrsus Berry, 1953

Octopus dofleini (Wülker, 1910)

Octopus penicillifer Berry, 1954

Octopus pricei Berry, 1913

Order Vampyromorpha

Family Vampyroteuthidae

Vampyroteuthis infernalis Chun, 1903 *

For a complete review of the early work done along the Pacific coast of North America, the reader is referred to the excellent paper of Berry (1912b).

Summary

SYSTEMATICS

1. Thirty-three species of pelagic cephalopods and the pelagic larval stage of a benthic octopus are described, and a key to their identification is presented.

2. The following 10 species are described as new to science: Abraliopsis falco, Pyroteuthis addolux, Octopoteuthis deletron, Gonatus onyx, Gonatus pyros, Gonatus californiensis, Chiroteuthis calyx, Valbyteuthis oligobessa, Mastigoteuthis pyrodes, and Leachia dislocata.

3. It is demonstrated that Abraliopsis felis and Watasenia scintillans do not belong in the same subgenus.

4. Pyroteuthis margaritifera oceanica, Pyroteuthis margaritifera aurantiaca are considered to be invalid subspecies.

5. Pterygroteuthis gemmata is shown to undergo considerable geographical variation in the number of arm hooks.

6. The photophore patterns of Ommastrephes bartramii and Symplectoteuthis luminosa are described.

7. The genus Neoteuthis is delineated and the family Alluroteuthidae is synonymized with the family Neoteuthidae.

8. Octopoteuthis longiptera is declared a doubtful species. It is suggested that all identifications of Octopoteuthis sicula from the Atlantic Ocean must be held in question until the photophore pattern of specimens from the type locality in the Mediterranean is determined. Octopodoteuthopsis is submerged for the second time into Octopoteuthis.

9. The formerly monotypic genus Gonatus is shown to consist of at least 10 species, 4 of which live sympatrically off California. The number of arm and club suckers in members of the genus are shown to become fixed at a very early stage and constitute an important systematic character. Each species off California is shown to have a distinctive pattern at the time when various arm and club hooks appear.

10. Larval Gonatopsis borealis are shown to possess tentacles which are lost abruptly at 8–11 mm M.L. Gonatopsis borealis makko is elevated to specific status.

11. Onychoteuthis borealijaponicus is shown to be a valid species. It is further suggested that “O. banksii” represents a complex of species.

12. The formerly monotypic genus Valbyteuthis is shown to consist of at least 5 species, 2 of which are undescribed.

13. The enlarged photophore embedded in the eyelid of certain Mastigoteuthis is shown to be a widespread feature in the genus. M. grimaldii and M. iselini are declared doubtful species. M. levimana is transferred to the genus Valbyteuthis. Mastigoteuthis glaukopis atlantica is elevated to specific status. M. dentata is briefly redescribed from material taken at the type locality off Panama.

14. Pyrgopsis is shown to represent a larval stage of Leachia.

15. Galiteuthis phyllura is considered to be a valid species, although evidence for its separation from G. armata is not conclusive. Taonidium pacificum is shown to be a larval Galiteuthis and a valid species.

16 Helicocranchia is demonstrated to be a valid and distinctive genus. The enigmatical genus Ascocranchia is shown to be a close relative of Helicocranchia. The identification of Helicocranchia pfefferi off California needs confirmation through the comparison of adults from the Atlantic and Pacific oceans.

17. Two sympatric species of Japetella are shown to occur off California, but their taxonomic separation has been incomplete.

18. The abundant octopus larvae appear to belong predominantly to one species, but it has not been identified.

19. It is suggested that Berry’s Strauroteuthis sp. may be the young of Opisthoteutnis californiana.

20. Species known to occur off California, but not encountered in the present trawling program, are briefly distinguished from those described in this paper.

DISTRIBUTION

1. Northern zones within the area studied have predominantly Subarctic and Transitional water species.

2. Southern zones have predominantly tropical species. The two most numerous of which appear to be associated with the north Pacific central water.

3. Forty-two species of pelagic cephalopods are now known from the waters off southern California, of which 36 have been identified to the specific level. Only 10 of these species appear to be cosmopolitan; however, in only 4 of the 10 species were detailed comparisons with Atlantic specimens possible. All 4 showed marked geographical variation.

4. All 10 apparently cosmopolitan species are associated primarily with tropical and subtropical waters.

5. It is suggested that Histioteuthis heteropsis and Octopoteuthis deletron are antitropical species.

6. Gonatus onyx, G. califoriensis, and G. pyros appear to occur primarily in the inshore waters over the Continental borderland while Onychoteuthis borealijaponicus, larval Abraliopsis felis, and Pyroteuthis addolux showed preference for more oceanic waters.

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Appendix 1

After the present manuscript was completed, a paper by Okutani and McGowan (1969) on the systematics and distribution of larval squid from the California current was published. Fourteen species of larval squid are described and information is presented on spawning and distribution of some of these. The authors attempted to identify the larvae by tracing successive developmental stages to the identifiable adult (page 7). This is a difficult task under the best circumstances; however, it becomes doubly difficult when the systematics of the adults is in an unstable state as it was when these authors undertook their project. Laboring under these handicaps, it is inevitable that errors will be made; however, as long as the following emendations are considered, little information will be lost from their paper because of systematic difficulties.

1. Specimens identified as Octopodoteuthopsis sp. almost certainly belong to Octopoteuthis deletron.

2. Onychoteuthis banksii should read Onychoteuthis borealijaponicus. Their illustration (figure 9c) shows a tentacular club with 26 hooks, a feature distinctive of this latter species. Their records of this species are shown to extend far south to the tip of Baja California. The most southernly records possible belong to a member of the “O. banksii” complex. This is somewhat suggested by the shape of the individual in figure 9h (no locality data are given for this figure); however, even if this proves to be the case, it will not significantly effect the conclusions reached by Okutani and McGowan.

3. The larvae identified as Gonatus fabricii probably belong to the most dominant species in the genus off California, Gonatus onyx, and this is attested to by there figure of a club belonging to this species (page 28, figure 10a). Larvae of G. pyros and G. californiensis are extremely close to those of G. onyx and since these species are quite common, I suspect some of their specimens belong to these species also. The authors mention that 10 larvae had more slender and opaque mantles. I suspect that these are larvae of Gonatopsis borealis and not Berryteuthis magister as is suggested. Okutani and McGowan state that this latter animal is known from the California coast. I have not been able to find this record, but its occasional occurrence here would not be surprising since it has been reported from Oregon. Even if the species gets as far south as southern California, I doubt that it breeds here as no larvae or juveniles have been taken in the University of Southern California’s trawling program.

4. Juveniles identified as Meleagroteuthis heteropsis consist of two species: Histioteuthis heteropsis (figure 12c) and H. dofleini (figure 12d).

5. The specific identification of their Ctenopteryx must be held in question. The genus is in a systematic jumble at present.

6. Larvae identified as Chiroteuthis veranyi almost certainly belong to C. calyx.

7. Species identified as Pyrogopsis pacificus belong to Leachia dislocata as evidenced by figure 16b which shows the displaced tubercle characteristic of the species.

8. Species identified as Teuthowenia megalops almost certainly belong to Galiteuthis phyllura.

These authors add two species to those now known from southern Californian and adjacent waters. These are Liguriella sp. from 28°17′ N, 118°58′ W and 29°12′ N, 115°38′ W, and Ctenopteryx sp. from 27°33′N and 115°53′W.

Appendix 2

PLATES 1–38

PLATE 1.—A, Abraliopsis falco, ventral view, male, M.L. 37 mm; B, same specimen, left side of head; C, Abraliopsis felis, ventral view, male, M.L. 41 mm; D, same specimen, left side of head.

PLATE 2.—A–G, Abraliopsis falco: A, tentacular club, M.L. approximately 40 mm; B. arms IV, mature male; C, Largest hook from arm III; D, distal sucker, arm III; E, largest club hook; F, dactylus sucker (C, D, F maintain relative sizes, E has half enlargement of others, all M.L. approximately 40 mm); G, ventral view of eye. H–N, Abraliopsis felis: H, tentacular club, M.L. 40 mm; I, arms IV, M.L. 41 mm; J, largest hook from arms III; K, distal sucker, arm III; L, largest club hook; M, dactylus sucker (J, K, M maintain relative sizes, L has half enlargement of others, all M.L. 40 mm); N, ventral view of eye.

PLATE 3.—A, Pterygioteuthis gemmata, ventral view, M.L. 23 mm; B, Pyroteuthis addolux, ventral view, holotype, M.L. 30 mm.

PLATE 4.—A–H, Pterygioteuthis gemmata: A, tentacular club, M.L. 28 mm; B, medial view, hectocotylus, M.L. 27 mm; C, largest hook, arm III; D, ventral view of eye; E, chitinous plate from hectocotylus, M.L. 24 mm; F. chitinous plate from hectocotylus, M.L. 27 mm; G, inner ring from largest sucker, arm III; H, inner ring, largest sucker, manus (C, G, H maintain relative sizes, M.L. 23 mm). I–Q, Pyroteuthis addolux: I, tentacular club, holotype; J, hectocotylus, holotype; K, spermatophore, holotype; L, enlargement of tip of ejaculatory apparatus, holotype; M, largest club hook, tentacular club; N, inner ring of largest club sucker; O, largest hook, arm III; P, inner ring, second proximal sucker, arm IV, (M-P maintains relative size, M.L. 45 mm); Q, ventral view of eye. R. Pyroteuthis margaritifera, second hook proximal to pad on ventral row of hectocotylus, Gulf of Mexico specimen, M.L. 28 mm; S. Pyroteuthis addolux, second hook proximal to pad on ventral row of hectocotylus, holotype, M.L. 30 mm.

PLATE 5.—A, Symplectoteuthis luminosa, ventral view, M.L. 136 mm; B, Ommastrephes bartramii, ventral view, M.L. 211 mm.

PLATE 6.—A, Symplectoteuthis luminosa, photophore pattern, ventral view, M.L. 163 mm; B, Same, dorsal view; C,Ommastrephes bartramii photophore pattern, ventral view, M.L. 215 mm.

PLATE 7.—A–I, Symplectoteuthis luminosa, M.L. 136 mm; A, tentacular club; B, inner ring of large sucker from 6th row of arm III; C, inner ring of basal sucker of arm III; D, inner ring of sucker from 18th row of arm III; E, inner ring of sucker from middle of dactylus, dorsal row; F, enlargement of E; G, inner ring of largest sucker, manus, marginal row; H, inner ring of largest sucker, manus, medial row; B–E, G–H maintain relative sizes); I, right funnel locking-cartilage showing area of fusion. J–R, Ommastrephes bartramii from off Guadalupe Island, Pacific Ocean, M.L. 303 mm: J, right funnel locking-cartilage; K, inner ring of sucker from middle of dactylus, ventral row; L, large medial sucker of manus showing fleshy, marginal papillae; M, tentacular club; N, inner ring of basal sucker, arm III; O, inner ring of 5th row sucker, arm III; P. inner ring of 14th row sucker, arm III; Q, inner ring of largest sucker, marginal row of club (K, N–Q maintain relative sizes); R, inner ring of largest sucker, medial row of club, enlargement half that of other suckers; S, Ommastrephes bartramii from off Florida, M.L. 302 mm, inner ring of largest sucker, marginal row of club; T, same specimen, inner ring of largest sucker, medial row of club (enlargement of S and T is the same as Q, and R, respectively).

PLATE 8.—Neoteuthis sp., M.L. 83 mm: A, ventral view; B, tentacular club; C, dorsal view.

PLATE 9.—A, Histioteuthis heteropsis, ventral view, photophore pattern only approximate, M.L. 35 mm; B, Histioteuthis dofleini, ventral view, M.L. 150 mm.

PLATE 10.—A, Histioteuthis dofleini, tentacular club, M.L. 150 mm; B, same, large club sucker, M.L. 150 mm; C, same, inner ring of sucker from arm III, 6th row, M.L. 14 mm; D, Histioteuthis heteropsis, tentacular club, M.L. 35 mm; E, same, inner ring of large club sucker, M.L. 65 mm; F, same, inner sucker ring, arm III, 11th row; G, same, inner sucker ring, arm IV, eleventh row (E–G maintain relative sizes). H–M, Octopoteuthis deletron; H, tip of arm III, M.L. 167 mm; I, ventral view of mantle and fins, M.L. 167 mm; J, inner ring from tip of arm III, fifth row from hooks, M.L. 167 mm; K, large hooks, arm III, M.L. 20 mm?; L, large hook, arm III, M.L. 109 mm; M, large hook, arm III, M.L. 167 mm.

PLATE 11.—A, Octopoteuthis deletron, holotype, M.L. 109 mm; B, same showing photophore pattern.

PLATE 12.—A–D, Octopoteuthis deletron showing metamorphosis, relative sizes maintained, M.L. 31, 32, 34, 39 mm, respectively; E, Octopoteuthis A, large arm hook. M.L. 49 mm; F, Octopoteuthis A, ventral view of mantle and fins. M.L. 49 mm; G, Octopoteuthis neilseni, ventral view of mantle and fins, M.L. 36 mm; H, Octopoteuthis danae, ventral view of mantle and fine, M.L. 53 mm; I, Octopoteuthis megaptera, ventral view of mantle and fins, M.L. 38 mm.

PLATE 13.—A, Gonatus onyx, ventral view, P.L. 69 mm; B, Gonatus pyros, ventral view, P.L. 39 mm.

PLATE 14.—A, Gonatus onyx, tentacle, P.L. 69 mm; B,Gonatus pyros, tentacle, P.L. 35 mm?; Gonatus onyx, P.L. 69 mm relative sizes maintained except D: C, inner ring of large marginal sucker, arm III; D, large hook, arm III, enlargement half that of suckers; E, inner ring of large medial row sucker, arm IV; F, inner ring of large marginal sucker, arm IV; G, inner ring of largest sucker of ventral marginal zone of club; H, inner ring of sucker from middle of dactylus, ventral row; I, inner ring of sucker taken one-fifth of tentacle length from base of club J–Q, Gonatus pyros, P.L. 39 mm, relative sizes maintained except for L: J, inner ring of arm III sucker tilted to show frequent bicuspid nature of teeth; K, inner ring of large marginal sucker, arm III; L, large hook, arm III, enlargement half that of suckers; M, inner ring of large medial row sucker, arm IV; N, inner ring of large marginal sucker arm IV; O, inner ring of largest sucker from ventral marginal zone on club; P, inner ring of sucker from middle of dactylus, ventral row; Q, inner ring of sucker taken one-fifth of tentacle length from base of club.

PLATE 15.—A, Gonatus berryi, ventral view, P.L. 119 mm; B, Gonatus californiensis, ventral view, holotype, P.L. 112 mm.

PLATE 16.—A, Gonatus berryi, tentacle, P.L. 119 mm; B, Gonatus californiensis, tentacle, holotype, P.L. 112 mm. C–I. Gonatus berryi, relative sizes maintained except for D: C, inner ring of large marginal sucker, arm III; D, large hook, arm III, enlargement half that of suckers; E, inner ring of large medial row sucker, ventral arm; F, inner ring of large marginal sucker, arm IV; G, inner ring of largest sucker of ventral marginal zone club; H, inner ring of sucker from middle of dactylus, dorsal row; I, inner ring of sucker taken one-fifth of tentacle length from base of club. J–P, Gonatus californiensis, holotype, relative sizes maintained except for K: J, inner ring of large marginal sucker, arm I: K, large hook, arm I, enlargement half that of suckers; L, inner ring of large medial row sucker, arm IV; M, inner ring of large marginal sucker, arm IV; N, inner ring of largest sucker of ventral marginal zone of club; O, inner ring of sucker from middle of dactylus, ventral row; P, inner ring of sucker taken one-fifth of tentacle length from base of club.

PLATE 17.—A–C, club hooks Gonatus berryi, P.L. 119 mm (relative sizes maintained): A, central hook, enlargement half that of other hooks; B, distal hook; C, first proximal hook. D–F, Gonatus californiensis, P.L. 112 mm, club hook (relative sizes maintained): D, central hook; E, distal hook; F, first proximal hook. G–I , Gonatus pyros, P.L. 39 mm, club hooks (relative sizes maintained): G, central hook; H, distal hook; I, proximal hook. J, Gonatus onyx, P.L. 69 mm?, central hook of club; K, Gonatus Pyros, P.L. 37 mm, ventral view of eye showing photophore; L, Gonatus berryi, larva, PL. 10 mm.

PLATE 18.—A, Onychoteuthis borealijaponicus, ventral view, M.L. 50 mm; B, Gonatopsis borealis, ventral view, M.L. 240 mm.

PLATE 19.—A–G, Onychoteuthis borealijaponicus, M.L. 81 mm; A, tentacular club; B, ninth hook from proximal end of club in ventral row from club in A; C–F, largest suckers from arms I–IV respectively (B–F retain relative sizes); G, visceral photophores. H, Onychoteuthis “banksii” horn off Florida, visceral photophores, M.L. 80 mm. I—P, Gonatopsis borealis: I, larva, M.L. 9 mm; J, Tentacle from larva, M.L. 7 mm; K–P, Gonatopsis borealis, M.L. 200 mm, relative sizes retained except in M: K, inner ring of largest sucker, arm IV, median row;L, inner ring of largest sucker, arm IV, marginal row; M, largest hook, arm III, enlargement half that of suckers. N, inner ring of sucker, arm III tip, medial row; O, inner ring of largest sucker, arm III, marginal row;P, inner ring of sucker, arm III tip, marginal row.

PLATE 20.—Chiroteuthis calyx: A, advanced doratopsis stage, ventral view, M.L. 49 mm?; B, ventral view, subadult, M.L. 54 mm.

PLATE 21.—A, Chiroteuthis calyx, oral view of tentacular club, M.L. 60 mm; B, same specimen, aboral view of tentacular club; C, Chiroteuthis veranyi from Mediterranean Sea (DANA 4030 XVIII), M.L. 61 mm, aboral view of tentacular club.

PLATE 22.—Chiroteuthis calyx except L and M which are C. veranyi. Inner sucker rings in A–I maintain relative sizes except in E, M.L. 60 mm?. A, arm III, second row; B, arm III, thirteenth row; C, arm III, eighteenth row; D, arm III, fourteenth row; E, D enlarged four times; F, arm I, sixth row; G, arm II, tenth row; H, arm IV, first row; I, arm IV, twelfth sucker. J, inner ring of large club sucker; K, largest arm III sucker from advanced doratopsis larva, M.L. 56 mm; L, C. veranyi from Mediterranean Sea (DANA 4139 IV), M.L. 68 mm, inner sucker ring, twelfth sucker of arm IV; M, same, inner sucker rings, thirteenth row of arm III; N, ventral view of eye showing photophores, holotype; O, left funnel locking-cartilage, M.L. 60 mm; Q, medial and marginal tentacular suckers and stalks, holotype.

PLATE 23.—A, Valbyteuthis oligobessa, mature female, holotype; B, Valbyteuthis danae, ventral view, redrawn from Roper and Young (1967), M.L. 34 mm; C, Valbyteuthis oligobessa, juvenile, M.L. 23 mm.

PLATE 24.—A-G. Valbyteuthis danae redrawn from Roper and Young (1967); A, tentacular club, M.L. 47 mm; B-E, largest suckers from arms I-IV, respectively, M.L. 47 mm; F, largest club sucker M.L., 47 mm, (B-F maintain relative sizes); G, mantle and funnel locking-cartilages, M.L. 47 mm. H-N, Valbyteuthis oligobessa, V-10976, M.L. 27 mm; H—K, largest suckers from arms I-IV, respectively; L, largest club sucker (H—L maintain relative sizes); M, mantle and funnel locking-cartilages; N, tentacular club; O, Grimalditeuthis bomplandii, large sucker and base from arm III, M.L. 89 mm; P, same specimen, inner ring of large sucker from arm III; Q, Valbyteuthis oligobessa, section of distal portion of outer ring of arm sucker, M.L. 34 mm; R, same specimen, scale from proximal portion of outer ring of arm sucker (slightly diagrammatic).

PLATE 25.—Mastigoteuthis pyrodes, holotype, M.L. 110 mm.

PLATE 26.—A–H, Bathyteuthis berryi: A, right arm I, holotype; B, tentacular club, M.L. 19 mm; C—F, suckers from base of arms I-IV, respectively, relative sizes maintained, holotype; G, inner ring of buccal sucker from dorsal lappet, holotype; H, club sucker, M.L. 19 mm. I—Q., Mastigoteuthis pyrodes: I—L, largest suckers from arms I-IV respectively, relative sizes maintained, holotype; M, inner ring of club sucker, holotype; N, club sucker enlarged three times relative to arm suckers, holotype; O, left funnel locking-cartilage, holotype; P, base of tentacular club of holotype; Q, aboral view of tentacular club of holotype; R, Mastigoteuthis dentata from off Cape Mala, Panama (PILLSBURY 510), large sucker from arm III; S, same specimen, club sucker; T, same specimen, inner ring of club sucker.

PLATE 27.—A, Bathyteuthis berryi, ventral view, holotype; B, Grimalditeuthis bomplandii, ventral view, M.L. 89 mm.

PLATE 28.—A, Heliocranchia pfefferi, ventral view, M.L. 41 mm; B, Cranchia scabra, ventral view, pattern and shape of tubercules diagrammatic, M.L. 107 mm.

PLATE 29.—A–I, Heliochanchia pfefferi, male, P.L. 39 mm: A, tentacular club; B, left arm I; C, left arm II; D, largest club sucker; E, inner ring of same sucker; F—I, largest suckers from arms I-IV, respectively, relative sizes maintained. J—Q, Cranchia scabra, male, M.L. 107 mm: J, tentacular club; K, hectocotylized right arm IV; L, largest sucker from tentacular club; M, enlarged sucker from tip of hectoctylus; N—Q, largest suckers from arms I-IV, respectively. L—Q maintain relative sizes.

PLATE 30.—A, Galiteuthis pacifica, ventral view, P.L. 42 mm; B, Lechia dislocata, VELERO 8028, ventral view, P.L. 110 mm.

PLATE 31.—Lechia dislocata: A, Larva, “pyrgosis stage,” P.L. 27 mm; B, mature female, P.L. 145 mm, tips of arms III added from another specimen.

PLATE 32.—Lechia dislocata: A, Immature hectocotylus of holotype; B, cartilagenous band and tubercules, left side, P.L. 136 mm: C, right tentacular club of holotype; D, largest sucker from arm III, female, P.L. 136 mm; E—H, largest suckers from arms I-IV, respectively, relative sizes maintained, holotype; I, enlarged sucker from tip of hectoctylus, P.L. 137 mm; J, marginal sucker from manus, holotype, enlargement double that of arm suckers; K, medial sucker from manus, holotype, same enlargement as arm suckers. L–S, development of eye photophores showing lateral and ventral views of eye at each stage: L, P.L. 16 mm; M, P.L. 22 mm; N, P.L. 25 mm; O, P.L. 35 mm; P, P.L. 44 mm; Q, P.L. 64 mm; R, P.L. 125 mm, holotype; S, P.L. 143 mm.

PLATE 33.—Galiteuthis phyllura, ventral views: A, M.L. 79 mm; B, M.L. 33 mm; C, M.L. 170 mm.

PLATE 34.—A, Galiteuthis phyllura, tentacular club, P.L. 79 mm; B, same, tentacular club, P.L. 359 mm; C, Galiteuthis paciftca, tentacular club, P.L. 36 mm; D, G. phyllura, larva, P.L. 13 mm; E—H, G. phyllura, P.L. 211 mm, largest suckers from arms I-IV, respectively, relative sizes maintained; I, same specimen; inner ring of largest dactyl sucker, enlarged four times relative to arm suckers; J, G. paciftca, largest dactyl sucker, P.L. 36 mm; K, G. paciftca, largest sucker of arms III, P.L. 65 mm; L, G. phyllura, cartilage at funnel-mantle fusion, P.L. 173 mm; M, G. paciftca, cartilage at funnel-mantle fusion, P.L. 65 mm.

PLATE 35.—A, Vampyrouthis infemalis, dorsal view, P.L. 50 mm; B, Japetella heathi, lateral view.

PLATE 36.—A, Japetella sp., V-9661, lateral view, M.L. 72 mm; B, same specimen, radula; C, same specimen, viscera; D, Japetella heathi, viscera, M.L. 75 mm; E, Vampyroteuthis infernalis, upper beak from Atlantic Ocean specimen, P.L. 78 mm; F, same, Californian specimen, P.L. 74 mm; G, same, Atlantic Ocean specimen, P.L. 64 mm; H, Gonatopsis borealis, juvenile, M.L. 35 mm.

PLATE 37.—Ocythoe tuberculata: A, radula, M.L. 80 mm; B, immature male, M.L. 14 mm; C, immature female, M.L. 80 mm.

PLATE 38.—A, Octopus sp. pelagic juvenile. M.L. 19 mm; B, Alloposus mollis, female, M.L. 115 mm.

____________

*K. N. Nesis (1971) has proposed that G. borealis be placed in a new subgenus, Boreoteuthis. The work of M. Okiyama (1969) and W. Fields and V. Gauley (in press) further indicates that present generic definitions in the Gonatidae are no longer adequate.

†Berry has divided R. pacifica, a benthic sepiolid, into two subspecies: R. p. pacifica and R. p. diegensis.

*Species described in this report.

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