NOAA Technical Report NMFS 73

Marine Flora and Fauna

of the Eastern United States

Mollusca: Cephalopoda

February 1989

Michael Vecchione, Clyde F.E. Roper, and Michael J. Sweeney

U.S. Department of Commerce
NOAA TECHNICAL REPORT NMFS

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5285 Port Royal Road, Springfield, VA 22161. Although the contents have not been copyrighted and may be reprinted entirely, reference to source is appreciated.

34. Additions to a revision of the shark genus Carcharhinus: Synonymy of Aprionodon and Hypoprion, and description of a new species ofCarcharhinus (Carcharhinidae),

by J. A. F. Garrick. November 1985, 26 p.

35. Synoptic review ofthe literature on the Southern oyster drill Thais haemastoma

j/oridana, by Philip A. Butler. November 1985, 9 p.

36. An egg production method for estimating spawning biomass of pelagic fish:

Application to the northern anchovy, Engraulis mordax, by Reuben Lasker (editor).

December 1985, 99 p.

37. A histopathologic evaluation of gross lesions excised from commercially important North Atlantic marine fishes, by Robert A. Murchelano, Linda DespresPatanjo, and John Ziskowski. March 1986, 14 p.

38. Fishery atlas of the northwestern Hawaiian Islands, by Richard N. Uchida and

James H. Uchiyama (editors). September 1986, 142 p.

39. Survey of fish protective facilities at water withdrawal sites on the Snake and

Columbia Rivers, by George A. Swan, Tommy G. Withrow, and Donn L. Park. April

1986, 34 p.

40. Potential impact of ocean thermal energy conversion (OTEC) on fisheries, by

Edward P. Myers, Donald E. Hoss, Walter M. Matsumoto, David S. Peters, Michael

P. Seki, Richard N. Uchida, John D. Ditmars, and Robert A. Paddock. June 1986,

33 p.

41. A stationary visual census technique for quantitatively assessing community structure of coral reef fishes, by James A. Bohnsack and Scott P. Bannerot. July 1986, 15 p.

42. Effects oftemperature on the biology of the northern shrimp, Pandalus borealis,

in the Gulf of Maine, by Spencer Apollonio, David K. Stevenson, and Earl E. Dunton,

Jr. September 1986, 22 p.

43. Environment and resources of seamounts in the North Pacific, by Richard N.

Uchida, Sigeiti Hayasi, and George W. Boehlert (editors). September 1986, 105 p.

44. Synopsis of biological data on the porgies, Calamus arctifrons and C. proridens

(Pisces: Sparidae), by George H. Darcy. September 1986, 19 p.

45. Meristic variation in Sebastes (Scorpaenidae), with an analysis of character

association and bilateral pattern and their significance in species separation, by Lochai Chen. September 1986, 17 p.

46. Distribution and relative abundance of pelagic nonsalmonid nekton off Oregon

and Washington 1979-84, by Richard D. Brodeur and William G. Pearcy. December

1986, 85 p.

47. Reproduction, maturation, and seed production of cultured species: Proceedings

of the twelfth U.S.-Japan meeting on aquaculture, Baton Rouge, Louisiana, October

25-29, 1983, by Carl J. Sindermann (editor). February 1987,73 p.

48. Widow rockfish: Proceedings of a workshop, Tiburon, California, December

11-12, 1980, by William H. Lenarz and Donald R. Gunderson (editors). January 1987,

57 p.

49. Reproduction, movements, and population dynamics of the southern kingfish,

Mentici"hus americanus, in the northwestern Gulf of Mexico, by Stephen M. Harding

and Mark E. Chittenden, Jr. March 1987,21 p.

50. Preparation of acetate peels of valves from the ocean quahog, Arctica islandica,

for age determinations, by John W. Ropes. March 1987, 5 p.

51. Status, biology, and ecology of fur seals. Proceedings of an international

workshop, Cambridge, England, 23-27 April 1984, by John P. Croxall and Roger

L. Gentry (editors). June 1987,212 p.

52. Limited access alternatives for the Pacific groundfish fishery, by Daniel D.

Huppert (editor). May 1987, 45 p.

53. Ecology of east Florida sea turtles: Proceedings of the Cape Canaveral, Florida,

sea turtle workshop, Miami, Florida, February 26-27, 1985, by Wayne N. Witzell

(convener and editor). May 1987, 80 p.

54. Proximate and fatty acid composition of 40 southeastern U.S. finfish species,

by Janet A. Gooch, Malcolm B. Hale, Thomas Brown, Jr., James C. Bonnet, Cheryl

G. Brand, and Lloyd W. Reiger. June 1987,23 p.

55. Proximate composition, energy, fatty acid, sodium, and cholesterol content of

finfish, shellfish, and their products, by Judith Krzynowek and Jenny Murphy. July

1987, 53 p.

56. Some aspects of the ecology of the leatherback turtle Dermochelys coriacea at

Laguna Jolova, Costa Rica, by Harold F. Hirth and Larry H. Ogren. July 1987, 14 p.

57. Food habits and dietary variability of pelagic nekton off Oregon and Washington,

1979-1984, by Richard D. Brodeur, Harriet V. Lorz, and William G. Pearcy. July

1987, 32 p.

58. Stock assessment of the Gulf of Menhaden, Brevoortia patronus, fishery, by

Douglas S. Vaughan. September 1987, 18 p.

59. Atlantic Menhaden, Brevoortia tyrannus, purse seine fishery, 1972-84, with a

brief discussion of age and size composition of the landings, by Joseph W. Smith,

William R. Nicholson, Douglas S. Vaughan, Donnie L. Dudley, and Ethel A. Hall.

September 1987, 23 p.

60. Gulf Menhaden, Brevoortia potronus, purse seine fishery, 1974-85, with a brief

discussion of age and size composition of the landings, by Joseph W. Smith, Eldon

J. Levi, Douglas S. Vaughan, and Ethen A. Hall. December 1987, 8 p.

61. Manual for starch gel electrophoresis: A method for the detection of genetic variation, by Paul B. Aebersold, Gary A. Winans, David J. Teel, George B. Milner, and

Fred M. Utter. December 1987, 19 p.

62. Fishery publication index, 1980-85; Technical memoradum index, 1972-85, by

Cynthia S. Martin, Shelley E. Arenas, Jacki A. Guffey, and Joni M. Packard.

December 1987, 149 p.

63. Stock assessment of the Atlantic Menhaden, Brevoortia tyranllUS, fishery, by

Douglas S. Vaughan and Joseph W. Smith. January 1988, 18 p.

64. lliusrrated key to penaeoid shrimps of commerce in the Americas, by Isabel Perez

Farfante. April 1988, 32 p.

65. History of whaling in and near North Carolina, by Randall R. Reeves and Edward

Mitchell. March 1988, 28 p.

66. Atlas and zoogeography of common fishes in the Bering Sea and Northeastern

Pacific, by M. James Allen and Gary B. Smith. April 1988, 151 p.
NOAA Technical Report NMFS 73

Marine Flora and Fauna

of the Eastern United States

Mollusca: Cephalopoda

Michael Vecchione

Clyde F.E. Roper

Michael J. Sweeney

February 1989

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

William E. Evans, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

James Brennan, Assistant Administrator for Fisheries
Foreword _

This NOAA Technical Report NMFS is part of the subseries "Marine Flora and Fauna ofthe Eastern

United States" (formerly "Marine Flora and Fauna of the Northeastern United States"), which

consists of original, illustrated, modem manuals on the identification, classification, and general

biology of the estuarine and coastal marine plants and animals of the eastern United States. The

manuals are published at irregular intervals on as many taxa of the region as there are specialists

available to collaborate in their preparation. These manuals are intended for use by students, biologists,

biological oceanographers, informed laymen, and others wishing to identify coastal organisms for

this region. They can often serve as guides to additional information about species or groups.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates of the Woods

Hole Region," edited by R.I. Smith, and produced in 1964 under the auspices of the Systematics

Ecology Program, Marine Biological Laboratory, Woods Hole, Massachusetts. Geographic coverage

of the "Marine Flora and Fauna of the Eastern United States" is planned to include organisms from

the headwaters of estuaries seaward to approximately the 200-m depth on the continental shelf from

Maine to Florida, but can vary somewhat with each major taxon and the interests of collaborators.

Whenever possible, representative specimens dealt with in the manuals are deposited in the reference

collections of major museums.

The "Marine Flora and Fauna of the Eastern United States" is being prepared in collaboration

with systematic specialists in the United States and abroad. Each manual is based primarily on recent and ongoing revisionary systematic research and a fresh examination ofthe plants and animals.

Each manual, treating a separate major taxon, includes an introduction, illustrated glossary, uniform

originally illustrated keys, annotated checklist (with information, when available, on distribution,

habitat, life history, and related biology), references to the major literature of the group, and a

systematic index.

Manuals are available from the National Technical Information Service, United States Department of Commerce, 5285 Port Royal Road, Springfield, VA 22161, either as a paper copy or

microfiche, for a charge. Manuals are not copyrighted, and so can be photocopied from the NOAA

Technical Report NMFS Circulars and Reports available in most major libraries and listed on

p. 23 of this manual.

The National Marine Fisheries Service (NMFS) does not approve, recommend

or endorse any proprietary product or proprietary material mentioned in this

publication. No reference shall be made to NMFS, or to this publication furnished

by NMFS, in any advertising or sales promotion which would indicate or imply

that NMFS approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose an intent to cause

directly or indirectly the advertised product to be used or purchased because of

this NMFS publication.

11
Contents

Introduction 1

Glossary 2

Key to neritic cephalopods of the northeastern United States 7

Annotated systematic list 16

Some recently published comprehensive works on cephalopods 20

Citations 20

Systematic index 21

Acknowledgments 21

Coordinating Editor's comments 22

List of published manuals in the Marine Flora and Fauna series 23

iii

Marine Flora and Fauna

of the Eastern United States

Mollusca: Cephalopoda

MICHAEL VECCHIONE

National Marine Fisheries Service. NOAA

Systematics Laboratory

National Museum of Natural History

Washington. DC 20560

CLYDE F.E. ROPER

MICHAELJ.SWEENEY

Department of Invertebrate Zoology-Mollusks

National Museum of Natural History

Smithsonian Institution

Washington, DC 20560

Abstract

The cephalopods found in neritic waters of the northeastern

United States include myopsid and oegopsid squids, sepiolid

squids, and octopods. A key with diagnostic illustrations is provided to aid in identification of the eleven species common in

the neritic waters between Cape Hatteras and Nova Scotia; included also is information on two oceanic species that occur over

the continental shelf in this area and that can be confused with

similar-looking neritic species. Other sections comprise a glossary of taxonomic characters nsed for identification of these

species, an annotated systematic checklist, and checklists of the

89 other oceanic species and 18 Carolinian and subtropical

neritic species that might occur occasionally off the northeastern

United States.

'A pile offresh squid lay on the deck, and the larger cod thinks very well of a little

shiny piece ofsquid tentacle at the tip of a clam-baited hook. Next day they caught

many fish. and met the "Carrie Pittman, " to whom they shouted their luck, and she

wanted to trade-seven cod for one fair-sized squid; but Disko would not agree at

the price. and the "Carrie" dropped sullenly to leeward and anchored half a mile

away. in hope of striking on to some for herself'

Rudyard Kipling

Captains Courageous

Introduction _

Cephalopods are very important, both to the marine ecosystem and

to humans. They can be abundant predators that are large, active,

and voracious; conversely, they are important as prey for many

marine mammals, birds, and fishes, including economically important species. Substantial squid fisheries have existed in North

America almost exclusively to provide bait for other fisheries.

Cephalopods also are important as food for people in many parts

of the world. Furthermore, they are important in biomedical

research; much of what is known about neurophysiology results

from experiments with the giant axon ofsquids, a contributing factor

in the founding of the Marine Biological Laboratory at Woods Hole,

Massachussetts.

The classification of recent taxa of the molluscan Class Cephalopoda (see Voss 1977) includes the squids (Order Teuthoidea), the

octopods (Order Octopoda), and the cuttlefishes and sepiolid or bobtailed squids (Order Sepioidea), as well as the bizarre vampire squid

in the monotypic Order Vampyromorpha and the chambered nautilus

in the separate subclass Nautiloidea. Teuthoid squids are further

distinguished based on whether their eyes are covered by a transparent cornea (Suborder Myopsida) or are exposed (Suborder

Oegopsida). The fauna of the northeastern United States lacks

nautiloids and true cuttlefishes but does include myopsid and oegopsid squids, octopods, and sepiolid squids; the vampyromorph Vampyroteuthis infemalis occurs in this geographic area but is limited

to the deep waters of the bathypelagic realm.

Although the cephalopods of the northeastern United States have

not been reviewed since the works of Verrill (1880, 1881), considerable collecting has been done in this region. Many species of

cephalopods have been collected off the northeastern United States

and can be divided into two ecological groups:

1) Species of the neritic zone with life histories restricted to

obligate relationships with the comparatively shallow waters of the

continental shelf (some of these species can also be found on the

continental slope); and

2) oceanic species found primarily in open oceanic waters away

from the shelf and slope (some of these species can also be found

in shelf and slope waters).

We discuss here the cephalopod species commonly found in neritic

waters of the northeastern United States. This work is based largely upon extensive collections of cephalopods deposited throughout

the last century in the National Museum of Natural History

(NMNH). These collections are a result of efforts by fishermen,

individual researchers, institutional research programs, and government agencies. The old U.S. Commission of Fish and Fisheries

and its descendents, the Bureau of Commercial Fisheries and, fmalIy, the National Marine Fisheries Service (NMFS) have conducted

comprehensive trawling along the northeastern United States for

groundfish surveys, and cephalopods from these surveys have been

deposited at the National Museum. The area of coverage for this

paper extends from Cape Hatteras to Nova Scotia.
The scope of this presentation does not include the many oceanic

species of squids and octopods normally found in deep waters

beyond the continental slope, although any of these species could

move inshore to the slope or shelfunder conditions such as impingement of a Gulf-Stream eddy onto the shelf. Examination of the

NMNH collections demonstrates, howeyer, that a few oceanic

species are sufficiently common in neritic waters and similar enough

to neritic species to cause confusion. Therefore, two ofthese oceanic

species are included in the key, but this is not intended to be a comprehensive guide to all oceanic species that can be transported into

neritic waters by Gulf-Stream eddies or other mechanisms.

Cephalopod morphology changes substantially during growth.

Some species hatch as planktonic "paralarvae" (Young and Harman

1988), and the greatest morphological changes occur during the

transition between the paralarval and juvenile phases of the life

cycle. Paralarval morphology and ecology have not been determined

for many species (Okutani 1987, Vecchione 1987). Thus, the key

is designed for identification of adults and advanced juveniles but

does not apply to paralarvae and early juveniles.

Glossary _

This glossary is derived from a more complete illustrated listing

oftaxonomic characters presented by Roper et aI. (1984). Interested

readers also should consult Roper and Voss (1983).

Anal flaps Pair of fleshy palps that arise at the sides of the anus

in teuthoid and sepioid squids (Fig. 1).

Anterior Toward the head end or toward the arm tips.

Carpal knobs Small rounded protuberances on the carpus of the

tentacular club to which carpal suckers from the opposite club

adhere during the locking-together of the clubs.

Carpus Proximal zone ofthe tentacular club which often has small

suckers and/or knobs (Fig. 2).

Cartilaginous tubercles Semi-rigid to rigid structures in the skin

of certain squids; can be overlapping and scale-like, or simple

to multifaceted knobs or papillae.

Cirri Fleshy erectile protuberances of the skin, usually elongate

and finger-like (Fig. 3).

Corneal membrane Thin, transparent skin that covers the eyes

of myopsid and sepioid squids and octopods (Fig. 2).

Dactylus Distal, terminal zone of the tentacular club, often

characterized by comparatively small suckers (Fig. 2).

Dorsal Uppermost surface, opposite the ventral surface where the

funnel is located.

Fin angle Angle between the longitudinal axis of the mantle and

the posterior border of one fin (Fig. 4).

Fins Pair of muscular flaps that arise along the dorsolateral sur·

face of the mantle, usually in its posterior half (Fig. 2).

Foveola Transverse membranous fold ofskin that forms a pocket

in the funnel groove of some oegopsid squids (Fig. 5).

Funnel Ventral tube through which water is expelled from the

mantle cavity (Fig. 2).

Funnel organ Glandular structures on the internal dorsal and

ventral surfaces of the funnel (Fig. 6).

Gladius Feather- or rod-shaped chitinous supporting structure in

the dorsal mantle of squids (Fig. 7).

Hectocotylus One or more arms in males of some cephalopod

species, modified for transferring spermatophores to the female;

modifications can involve the suckers, sucker stalks, protective

membranes, and/or trabeculae (Fig. 8).

Hooks Chitinous claw-like structures derived from the suckers

on the arms and/or tentacles of some oegopsid squids (Fig. 2).

2

Light organ Simple or complex structure that produces bioluminescence (also termed photophore).

Ligula Spatulate, or spoon-shaped, terminal structure of the hectocotylus of octopods (Fig. 9).

Mantle Tubular or sac-like muscular body of cephalopods (Fig. 2).

Mantle length Standard measure of size, is abbreviated ML (Fig.

10).

Posterior Toward the closed end of the mantle, away from the

head and arms.

Rachis Thickened central axis that usually extends the entire length

of the gladius. The free rachis is the portion that does not support vanes (Fig. 7).

Side pockets Structures formed by small membranous folds of

integument lateral to the foveola in the funnel groove of some

oegopsids (Fig. 5).

Suckers Muscular suction-cups on the arms and tentacles and occasionally on the the membrane around the mouth. Suckers of

squids are mounted on thin muscular stalks and include chitinous

rings with teeth (Fig. 11).

Tentacles Elongate pair of appendages located between the ventral and ventrolateral arms (Arms ill and IV) ofsquids and cuttlefishes (Fig. 2); with a proximal stalk usually devoid of suckers

or hooks, and with a distal club armed with suckers and occasionally hooks.

Tentacular club Terminal, usually expanded, portion of tentacle,

armed with suckers or, in some groups, with suckers and/or hooks

(Fig. 11).

Trabeculae Muscular rods that support the protective membranes

on the arms and tentacular clubs; occasionally the membranes

are reduced and trabeculae are elongated so they project beyond

the edge of the membrane as papillae (Fig. 8).

Vane Thin lateral expansion of the gladius that arises from the

rachis (Fig. 7).

Ventral Lower surface of the animal; side on which the funnel

is located.

Water pores Small orifices in the integument at the bases ofsome

arms on some pelagic octopods (Fig. 12).

Web Membrane that extends between the arms of many octopods

and sepiolid squids (Fig. 13).
anal flaps

~tr--ink sac

Figure 1

Ventral aspect of a squid visceral mass

(schematic).

suckers

keelbuccal membrane

funnel groove

Oegopsid eye

(orbital pore)

funnel--+-..,--/._

funnel-mantle fusion

funnel locking cartilage

mantle locking cartilage _-I-...-........-r,

photophores

fin (posteriorly concave)

sheathed

hooks

~,-- tentacle

fin lobe

tail

Figure 3

Lateral aspect of an octopod.

fin angle

Figure 4

Measurement of fin angle.

Figure 2

Composite diagram illustrating basic squid features, ventral aspect.

3
\)\\\\\1\ favea I.

~favea\

a. Todarodinae \ b. Ommastrephinae

Figure 5

Funnel grooves of some Ommastrephidae.

funnel valve

Figure 6

Interior of funnel, opened to show funnel organ.

greatest

width of

rachis

only enlarged

trabeculae,

no suckers

modified portion

starts here

·'U\>lofIH-t-+- suckers

trabeculae (transverse

fleshy supports)

Figure 8

Hectocotylized arm of a squid.

ligula calamus sucker

greatest

I--#--~ width of

vane

v

Figure 7

Gladius, showing measurement of rachis and vane.

4

Figure 9

Hectocotylized arm of an octopod.

@
mantle

head length

arm

length

tail

fin

length

arm

length

mantle

length

total

length

Figure 10

Some standard measurements on cephalopods. Squid (left); octopod (right).

sucker of squid

8 suckers

in (oblique)

transverse

rows

sucker of octopus club

Figure 11

Suckers and configuration ofsuckers (schematic) on tentacular club.

5
web

water

pores

Figure 12

The pelagic octopod Tremoctopus.

dorsal

eye oceJJus web __a~ms,_

--1--------11 -~ '-'

'-") suckers

-~II

'®~ ---- /~-~.o..J':--hec:cotYIUS \:-__ -- - "'Z'--.:~~ III

, -:_~~_~-~~~~Ia

~IV

-l~__ /_\_

outer gill

lamellae

(internal)

Figure 13

Schematic lateral aspect of octopod features.

6
Key to neritic cephalopods of the northeastern United States _

1 Suckers with narrow stalks and with chitinous rings that might be modified into hooks; usually ten circumoral appendages

comprised of eight arms and two tentacles (tentacles, however, can be lost during development)-squids and sepioids .....4

1 Suckers without stalks and without chitinous rings or hooks; eight circumoral appendages (arms only)-octopods 2

2(1) Any of the following characters present: fins; body gelatinous; water pores located on ventral surface of head at base of

arms; broad membranous flaps on dorsal arms; left third arm coiled in sac below eye oceanic octopods

2(1) None of the characters listed above 3

3(2) Ink sac absent in adults; arms short; suckers small; a large cirrus located dorsal to each eye; right arm III of males hectocotylized with huge, broad, spade-shaped ligula with prominent transverse ridges Bathypolypus arcticus (Fig. 14)

Figure 14

Bathypolypus arcticus (lateral aspect).

3(2) Ink sac present; arms moderately long; suckers large; cirri above eyes small or absent; ligula of hectocotylized right arm

III of males very small Octopus vulgaris (Fig. 15)

Figure 15

Octopus vulgaris (lateral aspect).

7
4(1) Any of the following characters present: suckers modified into hooks; numerous conspicuous light organs on surface of

mantle, head, arms, or tentacles; tissues gelatinous; mantle surface with cartilaginous tubercles oceanic squids

4(1) None of the characters listed above 5

5(4) Fins extend to posterior tip of mantle, joined or nearly joined posteriorly 9

5(4) Fins do not extend to posterior tip of mantle, broadly separate posteriorly 6

6(5) Mantle fused with head dorsally; all arms except ventral pair united by broad web; ventral mantle broadly flattened into

dark shield-like structure Stoloteuthis leucoptera (Fig. 16)

Figure 16

Stoloteuthis leucoptera. Ventral aspect (left); dorsal aspect (right).

6(5) Mantle free from head dorsally; only arms III and IV united by a web; mantle rounded ventrally 7

8
7(6) Light organs present on ink sac Semirossia tenera (Fig. 17)

Figure 17

Semirossia tenera. Dorsal aspect (left); oral aspect of male (right).

7(6) No light organs on ink sac 8

8(7) Dorsal pad of funnel organ short, without posterior extensions; anal flaps large, with expanded blades (Fig.18 left) ..

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rossia palpebrosa (Fig. 19)

Figure 18

Anal palps, funnel organs, and funnel valves. Rossia palpebrosa (left); Rossia megaptera (right).

9
Figure 19

Rossia palpebrosa (dorsal aspect).

8(7) Dorsal pad of funnel organ long, with angles and posterior extensions on both sides; anal flaps small, without expanded

blades (Fig. 18 right) Rossia megaptera (Fig. 20)

Figure 20

Rossia megaptera (dorsal aspect).

10
9(5) Eye covered by a cornea 10

9(5) Eye without a cornea 12

10(9) Fins in adults wider than long Lolliguncula brevis (Fig. 21)

Figure 21

Lolliguncula brevis. Ventral aspect (left); dorsal aspect (right).

10(9) Fins in adults as long as or longer than wide 11

II
11(10) Ratio of greatest width of gladius vane to greatest width of free rachis is 2.4-3.7 (Fig. 22); lateral margin of gladius vane

usually thin although sometimes slightly darkened in preserved specimens; modified suckers in hectocotylized left ventral

arm of males do not extend to arm tip __ Loligo pealei (Fig. 23)

Figure 22

Gladii. Loligo pealei (left);

Loligo plei (right).

Figure 23

Loligo pealei. Ventral aspect (left);

dorsal aspect (right).

12
11(10) Ratio of greatest width of gladius vane to greatest width of gladius free rachis is 1.5-2.4; lateral margin of gladius vane

usually thickened; modified suckers in hectocotylized left ventral arm of males extend to arm tip Loligo ptei (Fig. 24)

Figure 24

Loligo ptei. Ventral aspect (left); dorsal aspect (right).

Figure 25

Comparison of ventral aspects of mantle and fin outline.

Illex illecebrosus (left); 1. oxygonius (right).

12(9) Dactylus of tentacular club with eight longitudinal rows of small suckers; funnel groove smooth, without foveola or side

pockets 13

12(9) Dactylus of tentacular club with four longitudinal rows of small suckers; funnel groove with foveola and side pockets .... 14

13
13(12) Fin angle 40-50° (Fig. 25 left); modified portion of hectocotylized ventral arm in mature males occupies about 15-25 %

of hectocotylized-arm length; head not noticably wider than mantle opening; suckers on lateral arms of males not enlarged

.................................................................................. lilex illecebrosus (Fig. 26)

13(12) Fin angle 25-40° (Fig. 25 right); modified portion of hectocotylized ventral arm in mature males occupies about 30-35 %

of hectocotylized-arm length; head wider than mantle opening; suckers on lateral arms of males enlarged .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . lilex oxygonius (Fig. 27)

Figure 26

lilex illecebrosus. Ventral aspect (left); dorsal aspect (right).

Figure 27

lilex oxygonius (dorsal aspect).

14(12) Large, diffuse, compound light organ beneath integument on anterodorsal surface of mantle; not more than two suckers

occur proximal to knobs on carpus of tentacular club; no reflective stripe on midline of ventral mantle .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ommastrephes pteropus (Fig. 28)

14(12) No large dorsal light organ; longitudinal reflective stripe present along ventral midline of mantle surface; four to seven

suckers occur proximal to first knob on carpal part of tentacular club Ommastrephes bartrami (Fig. 29)

14
Figure 28

Ommastrephes pteropus. Ventral aspect (top); carpal

region of tentacular club (bottom).

15

Figure 29

Ommastrephes bartrami. Ventral aspect (top); carpal

region of tentacular club (bottom).
Annotated systematic list

Class Cephalopoda _

The generic status of at least some species in each family of

cephalopods discussed here is currently unsettled. Species-level

systematics also might change in the Octopodidae, Sepiolidae, and

Ommastrephidae from this area when thorough revisions have been

completed. A review of the published literature and an examination of the collections of the National Museum of Natural History

reveal that, in addition to the eleven species of neritic cephalopods

we list from the northeastern United States, 91 oceanic species might

occasionally be transported into these waters, and 18 other neritic

species that generally are confined to Carolinian or subtropical

waters might stray north of Cape Hatteras under unusual oceanographic circumstances.

1 Neritic species _

Order Sepioidea

FAMILY SEPIOLIDAE

Bobtailed squids are small benthic animals more closely related to

the cuttlefishes than to the true teuthoid squids. Species-level

systematics are confused, and the entire family, with many species

around the world, is in need of revision.

Rossia megaptera VerriU 1881, Fig, 20. This little-known species

occurs from New York to Greenland on the outer shelf and slope

at depths of 125-1500 m, with a maximum size of about 40 mm

ML.

Rossia palpebrosa Owen 1834, Fig. 19. An Arctic-Boreal species

known from Greenland and the Canadian Arctic to South Carolina

in the western Atlantic and from Spitzbergen to the North Sea

in the eastern Atlantic. In the NMFS Groundfish-Survey collections, it is scattered along the outer shelf at depths of 88-175 m.

Grows to about 45 mm ML. Rossia glauposis Loven 1846 and

R. sublaevis Verrill 1878 probably are junior synomyms (Nesis

1987).

Semirossia tenera (Verrill 1880), Fig. 17. Common on sand and

mud bottoms on the central shelf at depths of 85-135 m, S. tenera

is known to occur from the Gulf of Maine to the Gulf of Mexico

and Caribbean Sea. Also reported from Surinam and French

Guiana (Okutani 1983). Attains a size of about 50 mm ML. This

is the only species of Semirossia from the northeastern United

States, but its relationships with several other nominal species

in the southern part of its range are confusing.

Stoloteuthis leucoptera (Verrill 1878), Fig. 16. This beautiful little

cephalopod has seldom been reported since Yerrill's description.

Apparently, however, it is not rare, as several hundred specimens

are included in the NMFS Groundfish-Survey collections.

Stoloteuthis leucoptera seems to be relatively common on the outer

shelf and slope of the Gulf of Maine at depths of 175-340 m,

and is found as far south as the Straits of Florida. This species,

which reaches 25 mm ML, can be distinguished easily from all

other neritic sepiolids in the Atlantic by the ventral shield-like

structure of the mantle.

16

Order Teuthoidea

FAMILY LOLIGINIDAE

The systematics of the Loliginidae currently are in turmoil. In a

little more than a decade, at least four different generic-level

classifications have been proposed for species that include those

we refer to as Loligo (Cohen 1976; Natsukari 1984; G.L. Voss,

Univ. Miami, pers. commun., 1984; Brakoniecki 1986). We retain

this generic designation for both species as a matter of continuity

until such problems are resolved.

Although L. pealei and L. plei are distinct species, they are very

similar and often are difficult to separate. Cohen (1976) examined

50 taxonomic characters and concluded that L. pealei and L. plei

overlap in all but two: the structure of the hectocotylus and the ratio

of vane width to rachis width in the gladius. Five other characters

were found that, although overlapping, differed significantly

between the two species. Furthermore, the two species are most

similar in geographical areas where they are sympatric.

Loligo pealei Lesueur 1821, Fig. 23. A very abundant species

in the waters off the northeastern United States, L. pealei is the

target of important commercial fisheries. Also the primary source

of giant axons for neurological research in this region.

Loligo pealei occurs over the continental shelf and upper slope

from the surface to about 400 m depth. It is found throughout

the western North Atlantic from Newfoundland to Brazil, but apparently not around the islands of the Caribbean Sea (Cohen

1976). Vecchione (1981) described the early life history of L.

pealei, and Summers (1983) reviewed the many published studies

of its biology and ecology. Attains a maximum size of 500 mm

ML.

Loligo plei BlainviUe 1823, Fig. 24. Loligo plei tends to be limited

to warmer waters than L. pealei. Where the two species are sympatrie, L. pealei is usually found in deeper water than L. plei,

although considerable overlap occurs. The maximum depth

reported for L. plei is 370 m. Loligo plei is rare north of Cape

Hatteras but has been collected as far north as the New York

Bight. Its range extends south at least to southern Brazil and

includes the islands of Bermuda, the Bahamas, and the Caribbean Sea, where it is sympatric with two other species currently

assigned to the genus Loligo. Reported to reach a size of at least

350 mm ML.

Lolliguncula brevis (Blainville 1823), Fig. 21. A species that is

noteworthy for its euryhalinity, L. brevis is common in coastal

waters from Delaware Bay south to Brazil and is the only species

of cephalopod frequently caught in estuaries. Throughout its

range, it is found in waters less than 20 m deep. Haefner (1964)

compared the morphometries ofL. brevis, which attains a relatively small size of 120 mm ML, with the larger-growing L. pealei

from the Middle Atlantic Bight; Vecchione (1982) described the

development of L. brevis.
Order Teuthoidea (continued)

FAMILY OMMASTREPHIDAE

The family Ommastrephidae is currently under revision by several

authors; numerous questions need to be resolved on a worldwide

basis at both the generic and the species levels. Most members of

the family are oceanic, but a few genera, including Illex, are strongly

associated with the continental shelf and slope (although occasionally

they are found far offshore). This is the only family included in

the key whose species spawn pelagically.

lllex illecebrosus (Lesueur 1821), Fig. 26. This very abundant

species is important in commercial fisheries. Its depth range extends from the surface to probably about 1000 m. A seasonal

migration transports I. illecebrosus from the Gulf Stream right

up to the shoreline. Geographical range includes the western

Atlantic from the Straits of Florida to Greenland and Iceland and

extends eastward to the British Isles; however, the eastern Atlantic

populations may represent a separate species, I. coindetii, or

subspecies.lllex illecebrosus grows to 350 mm ML. O'Dor (1983)

reviewed the life history of I. illecebrosus, and its larval development and ecology have been discussed by Roper and Lu (1979),

Vecchione (1979), and Vecchione and Roper (1986).

lllex oxygonius Roper, Lu, and Mangold 1969, Fig. 27. Except

for mature adult males, I. oxygonius is very difficult to distinguish

from I. illecebrosus, especially where the two species are sympatric. IIlex oxygonius, however, has been found only as far north

as Delaware and appears to be rare north of Cape Hatteras. It

has been reported only as far south as Key West, Florida, although

this could simply be an artifact of the difficulty in distinguishing

I. oxygonius from I. illecebrosus and I. coindetii. IIlex oxygonius

has been reported from waters with bottom depths of 50-550 m

and attains a size of 230 mm ML. Its distribution was summarized by Roper et al. (1984).

Order Octopoda

FAMILY OCTOPODIDAE

Octopod morphology is quite variable, making the taxonomy difficult. Morphometric indices, such as relative lengths ofthe arms,

are often used in describing species but can be very misleading in

any particular specimen because of the octopod's ability to extend

or contract arms independently. Furthermore, artifacts of preservation can exacerbate this problem. Similarly, skin texture and color

patterns can be of use in distinguishing among species, but, because

these characters are related to different behavioral patterns and also

can change dramatically during fixation and preservation, they also

can be misleading. The generic-level systematics of the family are

currently under revision.

We recommend that a series ofspecimens be obtained, including

(if possible) mature adults of both sexes. For example, adult males

of Octopus vulgaris and Bathypolypus arcticus are easily distinguished based on the hectocotylus. Once the adult males of these

species have been compared, it will be easier to identify the less

easily distinguished juveniles without having to dissect them to determine if the internal ink sac is present. Differences in ovarian egg

size are consistent and useful for identification of adult females.

The eggs of O. vulgaris are 3 mm or less in length whereas those

of B. arcticus are about 13 mm long.

17

Octopus vulgaris Cuvier 1797, Fig. 15. Although this species occurs primarily south ofCape Hatteras, it is distributed as far north

as Connecticut. The NMFS Groundfish-Survey collections from

the Middle Atlantic Bight include several large specimens. As

currently defined, this species has been reported from shallow

(intertidal to 200-m depth) temperate and tropical waters of both

the eastern and western North and South Atlantic Oceans, the

Mediterranean Sea, Red Sea, Indian Ocean, and western Pacific

Ocean. Possibly, though, this broad range might actually represent a complex ofseveral similar, though currently unrecognized,

species (Roper 1985). Octopus vulgaris has been reported to reach

a size of 400 mm ML. The life cycle was reviewed by Mangold

(1983).

Bathypolypus arcticus (Prosch 1849), Fig. 14. The most abundant neritic octopod in the waters of the northeastern United

States. It is a cold-water species that also is found in the deep

water ofthe continental slope as far south as the Straits of Florida.

A species with considerable morphological variation, it has been

reported from eastern Canada, Greenland, Iceland, the North Sea,

and Spitzbergen. It is usually caught in trawls on mud bottoms

at depths of 14 to 1000 m, and attains a maximum size of about

100 mm ML. O'Dor and Macalaster (1983) reviewed the life

history of B. arcticus.

2 Oceanic species _

Both of these species are common in epipelagic and mesopelagic

waters of the Gulf Stream. They are transported onto the continental shelf of the northeastern United States by Gulf-Stream eddies.

Order Teuthoidea

FAMILY OMMASTREPHIDAE

Ommastrephes barlrami (Lesueur 1821), Fig. 29. Found in midlatitude waters ofthe North and South Atlantic, Pacific, and Indian

Oceans. It is absent from equatorial waters as well as from waters

with temperatures of less than WOC. In oceanic waters, O. bartrami occurs at the surface at night to about 1500 m depth.

Ommastrephes pteropus (Steenstrup 1855), Fig. 28. A PanAtlantic species of tropical and warm-temperate waters, O.

pteropus is found where surface waters are warmer than 22°C.

It is known to migrate vertically between the surface and deep

waters and has been reported at depths down to 1500 m. Nesis

(1987) has proposed that this species be assigned to the genus

Sthenoteuthis.
3 Other oceanic species _

Species that might be found in the waters off northeastern United

States and that have been recorded in oceanic waters north of26°N

and west of 6O oW.

Order Sepioidea

FAMILY SPIRULIDAE

Spirula spirula (Linnaeus 1758)

FAMILY SEPIOLIDAE

Heteroteuthis dispar (Ruppell 1844)

Nectoteuthis pourtalesi Verrill 1883

Order Teuthoidea

FAMILY LYCOTEUTIDDAE

Lycoteuthis diadema (Chun 1900)

Selenoteuthis scintillans Voss 1958

Lampadioteuthis megaleia Berry 1916

FAMILY ENOPLOTEUTIDDAE

Abralia veranyi (Ruppell 1844)

Abralia redfieldi Voss 1955

Abralia grimpei Voss 1958

Abraliopsis hoylei (Pfeffer 1884)

Abraliopsis pfefferi Joubin 1896

Pyroteuthis margaritifera (Ruppell 1844)

Pterygioteuthis giardi Fischer 1895

Pterygioteuthis gemmata Chun 1908

Enoploteuthis anapsis Roper 1964

Enoploteuthis leptura (Leach 1817)

Ancistrocheirus lesueuri (d'Orbigny 1839)

Thelidioteuthis alessandrini (Verany 1851)

(probably = A. lesueun)

FAMILY OCTOPOTEUTHIDAE

Octopoteuthis megaptera (Verrill 1885)

Octopoteuthis danae Joubin 1931

Taningia danae Joubin 1931

FAMILY ONYCHOTEUTHIDAE

Onychoteuthis banksii (Leach 1817)

Onykia carribaea Lesueur 1821

Ancistroteuthis lichtensteini (d'Orbigny 1839)

FAMILY CYCLOTEUTIDDAE

Cycioteuthis sirventi Joubin 1919

Discoteuthis discus Young and Roper 1969

Discoteuthis laciniosa Young and Roper 1969

18

FAMILY GONATIDAE

Gonatus fabricii (Lichtenstein 1818)

Gonatus steenstrupi Kristensen 1981

FAMILY LEPIDOTEUTmDAE

Lepidoteuthis grimaldi Joubin 1895

Pholidoteuthis adami Voss 1956

Tetronychoteuthis dussumieri (d'Orbigny 1839)

Tetronychoteuthis massyae Pfeffer 1912

FAMILY HISTIOTEUTIDDAE

Histioteuthis bonnelli (Ferussac 1835)

Histioteuthis celetaria (Voss 1960)

Histioteuthis corona (Voss and Voss 1962)

Histioteuthis dofleini (Pfeffer 1912)

Histioteuthis elongata (Voss and Voss 1962)

Histioteuthis meleagroteuthis (Chun 1910)

Histioteuthis reversa (Verrill 1880)

FAMILY NEOTEUTHIDAE

Neoteuthis thielei Naef 1921

FAMILY BATHYTEUTHIDAE

Bathyteuthis abyssicola Hoyle 1885

FAMILY CTENOPTERYGIDAE

Ctenopteryx sicula (Verany 1851)

FAMILY BRACIDOTEUTHIDAE

Brachioteuthis beanii Verrill 1881

Brachioteuthis riisei (Steenstrup 1882)

FAMILY ARCIDTEUTHIDAE

Architeuthis princeps Verrill 1875

(as well as other poorly defined species of Architeuthis)

FAMILY OMMASTREPIDDAE

lllex coindetii (Verany 1837)

Todarodes sagittatus (Lamarck 1799)

Ommastrephes caroli Furtado 1887 (may = O. bartrami)

Omithoteuthis antillarum Adam 1957

Hyaloteuthis pelagica (Bose 1802)
Order Teuthoidea (continued)

FAMILY THYSANOTEUTHIDAE

Thysanoteuthis rhombus Troschel 1857

FAMILY CHIROTEUTHIDAE

Chiroteuthis veranyi (Ferussac 1835)

FAMILY MASTIGOTEUTHIDAE

Mastigoteuthis grimaldii (Joubin 1895)

Mastigoteuthis hjorti Chun 1913

Mastigoteuthis magna Joubin 1913

FAMILY GRIMALDITEUTHIDAE

Grimalditeuthis bomplandii (Veranyi 1837)

FAMILY JOUBINITEUTHIDAE

Joubiniteuthis portieri (Joubin 1916)

FAMILY CRANCHIIDAE

Cranchio scabra Leach 1817

Liocranchio reinhardti (Steenstrup 1856)

Leachio cyclura Lesueur 1821

Leachio atlantica (Degner 1925)

Egea inermis Joubin 1933

Helicocranchio pfefferi Massy 1907

Galiteuthis armata Joubin 1898

Bathothauma lyromma Chun 1906

Sandalops melancholicus Chun 1906

Taonius pavo (Lesueur 1821)

Megalocranchia abyssicola (Goodrich 1896)

Megalocranchio oceanica (Voss 1960)

Megalocranchio speculator Chun 1906

Teuthowenio megalops (Prosch 1849)

Order Vampyromorpha

FAMILY VAMPYROTEUTHIDAE

Vampyroteuthis infemalis Chun 1903

Order Octopoda

FAMILY CIRROTEUTHIDAE

Cirroteuthis muelleri Eschricht 1838

Cirrothauma murrayi Chun 1911

19

FAMILY STAUROTEUTHIDAE

Stauroteuthis syrtensis Verrill 1879

Grimpoteuthis grimaldii (Joubin 1903)

Grimpoteuthis megaptera (Verrill 1885)

Grimpoteuthis umbellata (Fischer 1883)

FAMILY OPISTHOTEUTHIDAE

Opisthoteuthis agassitii Verrill 1883

FAMILY BOLITAENIDAE

Eledonella pygmaea Verrill 1884

Japetella diophana Hoyle 1885

FAMILY VITRELEDONELLIDAE

Vitreledonella richardi Joubin 1918

FAMILY OCTOPODIDAE

Benthoctopus piscatorum (Verrill 1879)

FAMILY TREMOCTOPODIDAE

Tremoctopus violaceus Delle Chiaje 1830

FAMILY OCYTHOIDAE

Ocythoe tuberculata Rafinesque 1814

FAMILY ARGONAUTIDAE

Argonauta argo Linnaeus 1758

Argonauta hians Lightfoot 1786

FAMILY ALLOPOSIDAE

Alloposis mollis Verrill 1880
4 Carolinian and subtropical neritic species _

These might occasionally occur north of Cape Hatteras.

Order Sepioidea

FAMILY SEPIOLIDAE

Semirossio equalis (Voss 1950)

Rossio bullisi Voss 1956

Rossio antillensis Voss 1955

Rossio tortugaensis Voss 1956

Order Teuthoidea

FAMILY PICKFORDIATEUTHIDAE

Pickfordioteuthis pulchel!a Voss 1953

FAMILY LOLIGINIDAE

Loligo ocula Cohen 1976

Loligo roperi Cohen 1976

Sepioteuthis sepioidea (B1ainville 1823)

Order Octopoda

FAMILY OCTOPODIDAE

Octopus hummelincki Adam 1936

Octopus joubini Robson 1929

Octopus briareus Robson 1929

Octopus burryi Voss 1950

Octopus macropus Risso 1826

Octopus defilippi Verany 1851

Danoctopus schmidti Joubin 1933

Scaeurgus unicirrhus (d'Orbigny 1840)

Pteroctopus tetracirrhus (Delle Chiaje 1830)

Tetracheledone spinicirrhus Voss 1955

20

Some recently published comprehensive works on cephalopods _

Boyle, P.R. (editor)

1983 Cephalopod life cycles. Vol. I, Species accounts. Acad. Press, London,

475 p.

1987 Cephalopod life cycles. Vol. II, Comparative reviews. Acad. Press, London, 441 p.

Hanlon, R.T. (editor)

1988 Life history, systematics and zoogeography of cephalopods in honor of

S. Stillman Berry. Malacologia 29(1), 307 p.

Nesis, K.N.

1987 Cephalopods of the world; squids, cuttlefishes, octopuses and allies. [Engl.

transl.] T.F.H. Publ., Neptune City, NJ, 351 p.

Nixon, M., and j.B. Messenger (editors)

1977 The biology of cephalopods. Symp. Zool. Soc. Lond. 38, 615 p.

Roper, C.F.E., C.C. Lu, and F.G. Hochberg (editors)

1983 Biology and resource potential of cephalopods. Mem. Natl. Mus. Vic. 44.

311 p.

Roper, C.F.E., M.j. Sweeney, and C.E. Nauen

1984 FAO species catalogue. Vol. 3, Cephalopods of the world. FAO Fish.

Synop. (125)3, 277 p.

Citations _

Brakoniecki, T.F.

1986 A generic revision of the family Loliginidae (Cephalopoda: Myopsida)

based primarily on the comparative morphology of the hectocotylus. Diss.,

Univ. Miami, Coral Gables, FL 33124, 163 p.

Cohen, A.C.

1976 The systematics and distribution of Loligo (Cephalopoda, Myopsida) in

the western North Atlantic, with descriptions of two new species. Malacologia

15(2):299-367.

Haefner, P.

1964 Morphometry of the common Atlantic squid, Loligo pealei, and the brief

squid, Lolliguncula brevis, in Delaware Bay. Chesapeake Sci. 5: 138-144.

Mangold, K.

1983 Octopus vulgaris. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol. I,

Species accounts, p. 335-364. Acad. Press, London.

Natsukari, Y.

1984 Taxonomical and morphological studies on the loliginid squids - IV. Two

new genera of the family Loliginidae. Venus 43(3):229-239.

Nesis, K.N.

1987 Cephalopods of the world; squids, cuttlefishes, octopuses and allies. [Engl.

transl.] T.F.H. Publ., Neptune City, NJ, 351 p.

O'Dor, R.K.

1983 Illex illecebrosus. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol. I,

Species accounts, p. 175-199. Acad. Press, London.

O'Oor, R.K., and E.G. MacAlaster

1983 Bathypolypus arcticus. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol.

I, Species accounts, p. 401-410. Acad. Press, London.

Okutani, T.

1983 Mollusks. In Takeda, M., and T. Okutani, Crustaceans and mollusks

trawled off Suriname and French Guiana. Jpn. Mar. Fish. Resour. Res. Cent.,

Tokyo, 354 p.

1987 Juvenile morphology. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol.

II, Comparative reviews, p. 33-44. Acad. Press, London.

Roper, C.F.E.

1985 Is there a squid in your future: perspectives for new research. Am. Malacol.

Bull., spec. ed. 1:93-100.

Roper, C.F.E., and C.C. Lu

1979 RhynchoteutlUon larvae of ommastrephid squids ofthe western North Atlantic, with the first description of larvae and juveniles of Illex illecebrosus. Prac.

BioI. Soc. Wash. 91(4):1039-1059.

Roper, C.F.E., M.j. Sweeney, and C.E. Nauen

1984 FAO species catalogue. Vol. 3. Cephalopods of the world. FAO Fish.

Synop. (125)3, 277 p.

Roper, C.F.E., and G.L. Voss

1983 Guidelines for taxonomic descriptions of cephalopod species. Mem. Natl.

Mus. Vic. 44:49-63.
Summers, W.C.

1983 Loligo pealei. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol. I, Species

accounts. p. 115-142. Acad. Press, London.

Toll, R.B.

1982 The comparative morphology of the gladius in the Order Teuthoidea

(Mollusca: Cephalopoda) in relation to systematics and phylogeny. Diss.• Univ.

Miami, Coral Gables, FL 33124, 390 p.

Vecchione, M.

1979 Larval development of IIIex Steenstrup, 1880, in the northwestern Atlantic, with comments on IIIex larval distribution. Proc. BioI. Soc. Wash. 91(4.):

1060-1075.

1981 Aspects of the early life history of Loligo pealei (Cephalopoda; Myopsida). J. Shellfish. Res. 1(2):171-180.

1982 Morphology and development of planktonic Lolliguncula brevis

(Cephalopoda: Myopsida). Proc. BioI. Soc. Wash. 95(3):601-608.

1987 Juvenile ecology. In Boyle, P.R. (ed.), Cephalopod life cycles. Vol. II,

Comparative reviews, p. 61-84. Acad. Press, London.

Vecchione, M., and C.F.E. Roper

1986 Occurrence of larval IIIex iIIecebrosus and other young cephalopods in

the Slope Water/Gulf Stream interface. Proc. BioI. Soc. Wash. 99(4):703-708.

Verrill, A.E,

1880 The cephalopods of the north-east coast of America. Part I. The gigantic

squids (Archileulhis) and their allies; with observations on similar large species

from foreign localities. Trans. Conn. Acad. Sci. 5(5):177-257.

1881 The cephalopods of the north-east coast of America. Part II. The smaller

cephalopods, including the "squids" and the octopi, with allied forms. Trans.

Conn. Acad. Sci. 5(6):259-446.

Voss, G.L.

1977 Classification of Recent cephalopods. In Nixon, M., and J.B. Messenger

(eds.), The biology of cephalopods, p. 575-579. Symp. Zool. Soc. Lond. 38,

615 p.

Young, R.E., and R.F. Harman

1988 "Larvae", "paralarva" and "subadult" in cephalopod terminology.

Malacologia 29(1):201-207.

Systematic index _

Note This is an index to the species included in the key and

discussed in the text, but does not include the extralimital species

listed on p. 18-20.

Bathypolypus arcticus 7

Illex

illecebrosus 13, 14

oxygonius 13, 14

Loliginidae 16

Loligo

pealei 12

plei 13

Lolliguncula brevis II

Octopodidae 17

Octopus vulgaris 7

Ommastrephes

bartrami 14, 15

pteropus 14, 15

Ommastrephidae 17

Rossia

megaptera 9, 10

palpebrosa 9, 10

Semirossia tenera 9

Sepiolidae 16

Stoloteuthis leucoptera 8

21

Acknowledgments _

Preparation of this manual was supported in part by a grant from

the National Science Foundation to the Editorial Board of the

"Marine Flora and Fauna of the Eastern United States."

The present manual has benefited from the artistic talents of Alan

D. Hart (Figs. 16, 18-20, 25, 28, and 29), Cynthia Mason (Fig.

14), Keiko H. Moore (Fig. 22, which was redrawn from Toll 1982),

and Ann Davis (Figs. 21, 23, and 26). Thanks are expressed to

Walter Fischer for allowing the use of Figures 1-13 and 27, which

were previously published by Roper et al. (1984); and to Wolfgang

Sterrer for allowing the use of Figure 15. Figure 17 was originally

published in Verrill (1881), and Figure 24 came from an unpublished

thesis by Edward T. LaRoe. Donald D. Flescher has kindly sent

many specimens from the NMFS Groundfish Survey over the past

decade. Reviews ofthe manuscript of this manual by Bruce B. Collette, John B. Pearce, Warren F. Rathjen, Ronald B. Toll, and

Richard E. Young are appreciated.

Preparation of manuals in the "Marine Flora and Fauna of the

Eastern United States" subseries is coordinated by the following

Board:

Coordinating Editor

Melbourne R. Carriker, College of Marine Studies, University of

Delaware, Lewes, DE 19958.

Editorial Advisers

A. Ralph Cavaliere, Department of Biology, Gettysburg College,

Gettysburg, PA 17325.

Arthur G. Humes, Boston University Marine Program, Marine

Biological Laboratory, Woods Hole, MA 02543.

David L. Pawson, Department of Invertebrate Zoology, National

Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Marine Science and Maritime Studies Center,

Northeastern University, East Point, Nahant, MA 01908.

Ruth D. Turner, Museum of Comparative Zoology, Harvard University, Cambridge, MA 02138.

Robert T. Wilce, Department of Botany, University of Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine Flora and

Fauna of the Eastern United States," the Board invites systematists

to collaborate in the preparation of manuals, reviews manuscripts,

and advises the Scientific Editor of the National Marine Fisheries

Service.

We regret to report that one of the original Editorial Advisers

is no longer with us: Marie B. Abbott passed away in 1987. We

would like to express our deep thanks to Marie for her enthusiasm

and support during the formative stages of the "Marine Flora and

Fauna of the Northeastern United States" and for her many subsequent editorial contributions.

We are pleased to report that a new systematist, Dr. Kenneth

P. Sebens, has joined the Board of Editorial Advisers, and we look

forward with much pleasure to his participation in the work of the

Board.
Coordinating Editor's comments _

Publication of the "Marine Flora and Fauna of the Eastern United

States" is most timely in view of the growing universal emphasis

on work in the marine environment and the critical need for precise

and complete identification of organisms related to this work. It

is essential, if at all possible, that organisms be identified accurately

to species. Accurate scientific names of plants and animals unlock

the great quantities of biological information stored in libraries,

obviate duplication of research already done, and often make possible prediction of attributes of organisms that have been inadequately

studied.

Michael Vecchione went to sea as a cabin boy on a three-masted

schooner in Maine at the age of 16. He completed undergraduate

studies in biology at the University of Miami in 1972, and then

spent four and a half years in the United States Army. He has been

working on cephalopods since his graduate studies on planktonic

molluscs during 1976-79 at the Virginia Institute of Marine Science,

School of Marine Science, College of William and Mary. After

receiving the Ph.D. degree, he worked briefly for the U.S. Fish

and Wildlife Service before accepting a faculty position at McNeese

State University. There he studied cephalopods, zooplankton, and

ichthyoplankton in addition to teaching during 1981-86. In 1986

22

he moved to his present position as Cephalopod Biologist at the

National Marine Fisheries Service Systematics Laboratory at the

National Museum of Natural History.

Clyde F.E. Roper's interest in marine biology developed while

he was a commercial lobster fisherman in New Hampshire during

summers through high school and undergraduate school at Transylvania College, Lexington, Kentucky. Lured to work as a research

assistant in the summer following his first year of graduate school

at the University of Miami's Institute of Marine Sciences (now

Rosenstiel School of Marine and Atmospheric Science), Roper was

captivated by cephalopods. He completed the M.S. and Ph.D.

degrees at Miami, working primarily on oceanic and deep-sea

squids. Roper has continued research on the systematics, biology,

and behavior of cephalopods of the world at the Smithsonian Institution's National Museum of Natural History where he has been

a Curator of Molluscs since 1966.

Michael 1. Sweeney has worked at the National Museum of

Natural History since 1968. His research interests are oceanic cephalopods and collections management. Restoration of antique automobiles occupies his spare time.
Published manuals

Marine Flora and Fauna of the Northeastern United States

Annelida: Oligochaeta

Cook, David G., and Ralph O. Brinkhurst

Protozoa: Ciliophora

Borror, Arthur C.

Higher Plants of the Marine Fringe

Moul, Edwin T.

Pycnogonida

McCloskey, Lawrence R.

Crustacea: Stomatopoda

Manning, Raymond B.

Crustacea: Decapoda

Williams, Austin B.

Tardigrada

Pollock. Leland W.

Cnidaria: Scyphozoa

Larson, Ronald J.

Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes

Cavaliere, A.R.

Copepoda: Harpacticoida

Coull, Bruce C.

Sipuncula

Cutler, Edward B.

Echinodermata: Holothuroidea

Pawson, David L.

Copepoda: Lernaeopodidae and Sphyriidae

Bo, Ju-Shey

Copepoda: Cyclopoids Parasitic on Fishes

Bo, Ju-Shey

Crustacea: Branchiura

Cressey, Roger F.

Protozoa: Sarcodina: Amoebae

Bovee, Eugene c.. and Thomas K. Sawyer

Crustacea: Cumacea

Watling, Les

Arthropoda: Cirripedia

Zullo, Victor A.

Cnidaria: Scleractinia

Cairns, Stephen D.

Protozoa: Sarcodina: Benthic Foraminifera

Todd, Ruth, and Doris Low

Turbellaria: Acoela and Nemertodermatida

Bush, Louise F.

Lichens (Ascomycetes) of the Intertidal Region

Taylor, Ronald M.

Echinodermata: Echinoidea

Serafy, D. Keith, and F. Julian Fell

Marine Flora and Fauna of the Eastern United States

Cephalopoda

Vecchione, Michael, Clyde F.E. Roper, and Michael J. Sweeney

23

NOAA Tech Rep.

NMFS Circular no. NTIS no.

374 COM 73 50670

378 7350888

384 7450019

386 7450014

387 7450487

389 7451194

394 PB 257 987

397 261 839

398 268036

399 268714

403 273062

405 274999

406 280040

409 281 969

413 222923

419 285538

423 296460

425 297676

438 124520

439 225053

440 219 387

446 124 735

NOAA Tech Rep.

NMFS no.

33 PC A03/MF AOI

73