Vertical Distribution

of Pelagic Cephalopods

*

CLYDE F. E. ROPER

and

RICHARD E. YOUNG

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 209
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 20 9

Vertical Distribution

of Pelagic Cephalopds

Clyde F. E. Roper

and Richard E. Young

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Roper, Clyde F. E., and Richard E. Young. Vertical Distribution of Pelagic

Cephalopods. Smithsonian Contributions to Zoology, number 209, 51 pages, 31

figures, 1975.—The vertical distributions of pelagic cephalopods are analyzed,

based primarily on studies undertaken in the waters off California, Bermuda,

and Hawaii. Much of the information is derived from midwater trawl nets

equipped with closing apparatus. Each family that contains pelagic or quasipelagic species is discussed; selected records from the literature are included

where necessary.

A wide variety of patterns of vertical distribution exists among pelagic cephalopods; patterns may be associated with stage of maturity, diel rhythms, or seasonal

behavior. The study revealed that several basic types of vertical distributional

patterns occur: near-surface dwellers, first order diel vertical migrators, second

order diel vertical migrators, diel vertical shifters, diel vertical spreaders, nonmigrators, vertical wanderers, species associated with the ocean bottom, species

exhibiting ontogenetic descent.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 5389. SERIES COVER DESIGN:

The coral Montastrea cavemosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Roper, Clyde F. E. 1937-

Vertical distribution of pelagic Cephalopods.

(Smithsonian contributions to zoology; no. 209)

Bibliography: p.

Includes index.

1. Cephalopoda—North Pacific Ocean. 2. Cephalopoda—Bermuda Islands. I. Young, Richard

E., joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to

zoology; no. 209.

QL1.S54 no. 209 [QL430.2] 591'.08s [594'.5'092] 75-6936
Contents

Page

Introduction 1

Materials and Methods 3

California Study 3

Bermuda Study 5

Hawaii Study 6

Subclass Nautiloidea 6

Nautilidae 6

Subclass Coleoidea 6

Order Sepioidea 6

Spirulidae 6

Sepiidae 7

Sepiadariidae, Sepiolidae, Idiosepiidae 7

Order Teuthoidea 8

Suborder Myopsida 8

Loliginidae 8

Pickfordiateuthidae 9

Suborder Oegopsida 9

Lycoteuthidae 9

Enoploteuthidae 9

Ommastrephidae 13

Histioteuthidae 14

Psychroteuthidae 15

Neoteuthidae 15

Architeuthidae 16

Bathyteuthidae 16

Gonatidae 17

Octopoteuthidae 21

Onychoteuthidae 22

Thysanoteuthidae 23

Cycloteuthidae 23

Ctenopterygidae 23

Brachioteuthidae 23

Lepidoteuthidae 24

Joubiniteuthidae 24

Chiroteuthidae 24

Mastigoteuthidae 27

Promachoteuthidae 28

Batoteuthidae 28

Grimalditeuthidae 28

Cranchiidae 28
i v SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Page

Order Vampyromorpha 32

Vampyroteuthidae 32

Order Octopoda 33

Suborder Cirrata 33

Cirroteuthidae and Stauroteuthidae 33

Opisthoteuthidae 34

Suborder Incirrata 35

Bolitaenidae 35

Vitreledonellidae 35

Amphitretidae 36

Allopsidae 36

Tremoctopodidae 36

Ocythoidae 36

Argonautidae 36

Discussion 37

Literature Cited . 45

Index 49
Vertical Distribution

of Pelagic Cephalopds

Clyde F. E. Roper and Richard E. Young

Introduction

Information concerning the vertical distribution

of pelagic cephalopods is relatively sparse. Several

factors are primarily responsible: (1) Oceanic

cephalopods are often large, fast-swimming animals that are able to avoid being captured by

conventional sampling gear. (2) Until recently,

most specimens were taken in open nets that, in

addition to the desired sampling depth, sample the

entire water column during setting and retrieval;

as a result the true depth of capture is difficult to

define precisely. (3) In many groups only larvae

and juveniles are captured and adults are rare or

unknown. (4) Most sampling has been conducted

along geographical transects where a given locality is generally sampled only once, at one time of

day and at one season of the year. Compounding

these sampling problems are the complexities of

the distributional patterns of the cephalopods.

To understand the vertical distributions of

cephalopods (as indeed of many oceanic organisms) a knowledge is required of more than

merely the bathymetric range, with carefully defined upper and lower limits, over which specimens of a species are known to occur. Often the

bathymetric range will extend over a great vertical

distance, but the vast majority of a population will

occupy only a restricted depth zone. In addition,

larvae of a species may inhabit one or more depth

Clyde F. E. Roper, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, D.C. 20560. Richard E. Young, Department of

Oceanography, University of Hawaii, Honolulu, Hawaii 96822.

zones that are uninhabited by adults. Therefore,

because distributional patterns often differ

markedly between growth stages, it is necessary to

separate and distinguish between depth records of

larval and juvenile-adult populations. Seasonal or

daily differences in depth distributions also may

occur; for example, some species migrate to nearshore waters in the summer to feed and/or spawn,

then migrate into deeper, offshore waters during

winter; many species in the day inhabit depths that

differ from their nighttime depths. Because of

these factors, it is extremely difficult to draw conclusions from most of the literature.

A factor that has led to considerable confusion

in the literature is the inability of many workers

to recognize the phenomenon of diel vertical migration in which animals ascend toward the surface

zone at night and descend to deeper water during

the day. Although vertical migrations are well

documented in several groups of pelagic animals,

evidence of this phenomenon in cephalopods has

been inconclusive until recently. The first indication of vertical movements of cephalopods was

given in 1841 by d'Orbigny who noted that cephalopods occurred at the surface at night but were

absent during the day (vide G. L. Voss, 1967:511).

Similar observations have been recorded since that

time. For instance, Sasaki (1914:92) stated that

Watasenia scintillans comes to the surface at night,

where it may be captured by fishermen, and disappears to greater depths during the day. More

recently Pearcy (1965) presented trawling data to

suggest that some oceanic species show greater frequencies of occurrence in the surface layer
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(0-200 m) during night than during the day, while

in the layer of 0-500 m abundance of these species

is greater during the day than during the night.

Clarke (1969) compared day-night captures using

several different types of sampling gear, among

them a closing-net, during the SOND cruises in the

eastern tropical Atlantic. His results indicate more

clearly the existence of vertical migration.

Our studies presented here confirm the occurrence of vertical migration in cephalopods and

indicate that this phenomenon is widespread in

oceanic squid. These conclusions have relied most

heavily on sampling programs conducted in three

different oceanic regions of the world. The first

study was conducted off southern California by the

University of Southern California and involved

approximately 900 hours of sampling using open

midwater trawls. The second study, known as the

Ocean Acre Program, was conducted off Bermuda

by the Smithsonian Institution in cooperation

with the University of Rhode Island and the U.S.

Navy. This study was based upon the use of closing

midwater trawls. Projects off Hawaii using closing

trawls and open trawls yielded additional

information.

This paper reviews the vertical distribution of

cephalopods and relies heavily on information derived from the above programs. Much of our data

is based on the capture of juvenile specimens. In

most species examined, however, juveniles appear

to exhibit the same distributional patterns as the

adults. Therefore, in most cases, this data can be

considered representative of the species.

The larvae of most species of pelagic cephalopods occur in the near-surface waters both during

the day and at night and as such do not demonstrate the same distributional patterns as their

juvenile and adult forms. In order to determine

the distributions of juveniles and adults, therefore,

the larvae for each species must be eliminated from

the calculations. All specimens that were equal in

size (mantle length) to, or smaller than, the largest

"larva" captured in near-surface waters during the

daytime (indicative of nonmigratory behavior)

were eliminated from the calculations.

We have attempted to include pertinent information from the literature. Our treatment of the

literature, however, has not been exhaustive because of the problems discussed above, as well as of

the danger of interpreting data based on records

about which we have no knowledge of the circumstances related to the capture.

The vertical distribution of cephalopods is surveyed by discussing representative species in each

family. Knowledge is insufficient to allow a presentation based upon recognized distributional patterns or vertical faunal zones in the open ocean.

The systematic approach allows us to present data

wherever they exist without having to force them

into distributional concepts. Hopefully, within the

next few years it will be possible to discuss vertical

distribution of cephalopods in relation to the physical and biological parameters of the pelagic

environment.

In addition to these primary sources listed above

we have utilized data from: (1) a closing-net program in the Mediterranean (Roper, 1974); (2)

open midwater trawl sampling of the Antarctic

Research Program; (3) bottom trawl surveys in

the tropical western Atlantic (National Marine

Fisheries Service); (4) a survey with a large, open

Engel trawl (WALTHER HERWIG); and (5) our collections from various other areas.

The Ocean Acre and Hawaii data presented

here are partial results of continuing programs.

The data represent some of the information that

was accumulated through January 1972.

For representative illustrations of the various

groups of pelagic squid discussed in the following

report, the reader may consult Roper, Young, and

Voss (1969).

The vertical distributional data are summarized

in Figures 24-31. As the graph generalizes data, it

should be used in conjunction with the text. The

distributions are represented by bars, and only

reference to the text will indicate the number of

data points available and consequently the reliability of the suggested distributions. Not all species

mentioned in the text are included in the graph.

The families included in the graph are not necessarily in the same order as in the text.

The following symbols are used in Figures 24-31. Solid bar: distribution of animals not

associated with bottom, based primarily on midwater tows. Bars represent centers of distribution

where the vast majority of the population occurs.

Hachured bar: distribution of animals associated

with bottom, based primarily on bottom trawls.

Dotted bar: presumed zone of distribution based

on limited data. Horizontal dotted line: distribu-
NUMBER 209

tional boundary uncertain. Horizontal dashed

line: single open-net record. Solid vertical line (or

narrow hachured bar): total distributional range

occupied by very small portion of population.

Dashed vertical line: few data points scattered over

broad range of depths; resolution of actual distribution not possible.

Plots for most species show distributions for day

(D) and night (N). Where data indicate no daynight differences, they are combined. Since a few

species show seasonal changes, their summerwinter (S-W) distributions are plotted. Data for

ontogenetic differences in distribution are also

plotted where available. "S-»" indicates increasing

size of non-adult specimens.

ACKNOWLEDGMENTS.—We are most grateful to the

many individuals and institutions whose aid has

made the completion of this study possible

through physical and financial support or by supplying data:

University of Southern California Deep-Sea Biology Program; especially J. Garth, J. Lavenberg,

M. Murillo, J. McGinnis, and D. Robins; students

who participated in cruises of Velero IV; National

Science Foundation Grant Numbers G-10691 and

G-23467.

The Ocean Acre and Mediterranean Biological

Studies Programs of the Smithsonian Institution

(U.S. Navy Contract Numbers: N00140-72-C-0315

and N00014-67-A-0399-0007); R. Gatton, M.

Sweeney, R. Goodyear, R. Gibbs, Jr., D. Brown

and all participants in the numerous cruises.

The University of Hawaii deep sea biology

projects; T. Clarke, R. Qrigg, and students who

participated in "Teuthis" cruises, especially J.

Walters, S. Amsbury, and F. Riggs; National

Science Foundation Grant Numbers GB-20933

(Young); GB-23931 (Clarke).

C. C. Lu, Marine Sciences Research Laboratory,

Memorial University of Newfoundland; G. L.

Voss, E. McSweeny, U.S. Antarctic Research Program Grant, Rosenstiel School of Marine and

Atmospheric Sciences, University of Miami; P.

Rancurel, ORSTOM, New Caledonia; A. Cohen,

Student Research Associate, Smithsonian Institution; L. Burgess, National Marine Fisheries Service Biological Laboratory, Hawaii; H. Bullis and

associates, National Marine Fisheries Service Exploratory Laboratory, Pascagoula; G. Krefft and

associates, WALTER HERWIG, Institut fur Seefischeri,

Hamburg; A. G. Smith, R. R. Talmadge, California Academy of Sciences.

K. Lamb and M. Sweeney, Division of Mollusks,

National Museum of Natural History, Smithsonian Institution, kindly checked the manuscript

for consistency and accuracy of numbers.

The manuscript has been reviewed by M. R.

Clarke, Marine Biological Laboratory, Plymouth;

K. Mangold, Laboratoire Arago, Banyuls-sur-Mer;

T. Okutani, Tokai Regional Fisheries Research

Laboratory, Tokyo; G. L. Voss, University of

Miami. We sincerely appreciate their comments.

Materials and Methods

CALIFORNIA STUDY

Material from California was collected by the

University of Southern California over a six-year

period from 1960 to 1966. During the program,

445 midwater tows were made with an open

Isaacs-Kidd midwater trawl representing over 900

hours of "fishing" at depths from the surface to

about 1200 m (Figure 1). The program yielded

over 5000 specimens of pelagic cephalopods. The

systematic study of these cephalopods, including

the descriptions of a number of new species, was

published by Young (1972a).

The reconstruction of the vertical distribution

of the species integrates catches over the entire

sampling period. Probably, at any given time the

vertical extent of these distributions is considerably less than shown. A correction factor that compensates for unequal trawling time at each depth

interval has been added to the charts of vertical

distribution (white bars). The correction factor

represents the catch that would be expected if all

depth intervals were sampled as intensively as the

most heavily sampled depth. The actual catch

(black bars) is presented on the same chart as the

correction factor. This technique is useful when

low numbers of specimens are available. Because

of the low numbers, the actual catch is discussed

rather than the adjusted catch except in a few

cases where the adjusted catch gives a slightly different picture of the vertical distribution.

Trawling times generally varied between two

and four hours; deep tows generally spent four

hours at depth. Any tow that was in the water at

0600 hours or at 1800 hours is considered a twi-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

light tow. For example, if a deep tow began at

0115 hours and terminated at 0615 hours, it is

classified as a twilight tow and is excluded from

the day-night analyses.

The depth limit of the sampling program was

approximately 1200 m. A number of tows undoubtedly extended below 1200 m; because it is

difficult to determine actual fishing depth of the

net, all specimens caught in tows below 1200 m

are lumped into the 1100-1200 m depth zone in

the charts. The data are handicapped by occasional lack of precise depth of capture records.

In these cases, depth of capture was considered to

be one-third of the wire out. This factor was determined by plotting values of wire out against

trawl depths that were obtained from a series of

trawl hauls with a depth recording instrument

present. The localities of trawling stations off

southern California are given in Young (1972a).

The oceanographic regime off southern California is unusually complex. The continental

borderland, 80-257 km in width from the shoreline

to the continental slope, contains 12 complete basins, ranging from about 500-2100 m in depth,

several troughs, numerous banks and eight islands.

At the western edge of the borderland, the continental slope drops steeply to a depth of about

3500 m. The waters over the borderland are derived from the complex intermixing of several

water masses. Surface waters are influenced by the

southerly flowing California Current that is derived largely from Subarctic waters and modified

through mixing with Central waters. Surface

waters also are modified by wind-induced upwelling that brings nutrient-rich waters to the surface

from a maximum depth of 200-300 m. Below

1200

0 40 60 80

Hours at Depth

FIGURE 1.—Hours of trawling at various depths (in 100 m increments) off southern California.

(Black bar = hours of fishing at night; hatchured bar = hours of fishing during the day; white

bar = total hours of fishing (day, night, twilight).)
NUMBER 209

150-200 m a deep countercurrent of Pacific Equatorial Water flows northward along the coast to at

least 40°N and progressively mixes with the lower

layers of the California Current. (Above discussion based on Reid, Roden, and Wyllie, 1958;

Emery, 1960).

The vertical temperature structure of the water

shows considerable variation, often in association

with large internal waves (Emery, 1956). In general, however, the average surface temperature

varies seasonally from about 14° to 20° C. The

10° C isotherm, which marks the approximate bottom of a strong shallow thermocline, usually lies

between 100 m and 200 m. The 8° C isotherm generally fluctuates around the 300 m level, the 6° C

isotherm lies between 700 m and 800 m, and the

4° C isotherm occurs at about 1000 m.

Salinity at the surface varies around 33.6 %0 and

gradually increases with depth to about 34.5 %c at

1000 m.

Oxygen concentration at the surface is approximately 6.0 ml/L, and between 500 m and 1000 m

it reaches a minimum that is less than 0.5 ml/L.

Emery (1960) gives an oxygen value of 0.2 ml/L

below sill depth in the San Pedro Basin and 0.4

ml/L below sill depth in the Catalina Basin.

Emery and Hiilsemann (1962) found that oxygen

does not appear to be a limiting factor on the

abundance of the benthic megafauna in the Santa

Barbara Basin in concentrations above 0.3 ml/L.

Almost no megafauna was present, however, in

less than 0.1 ml/L. A rich cephalopod fauna occurs

in the oxygen minimum zone; and no correlation

is apparent between the patterns of vertical distribution of cephalopods and the oxygen minimum

layer.

BERMUDA STUDY

The cephalopod data utilized here from off

Bermuda are part of a study known as the Ocean

Acre Program. Details of the long-term program

are given in Gibbs and Roper (1970) and Gibbs,

Roper, Brown, and Goodyear (1971).

The principal sampling gear was a 3 m IsaacsKidd midwater trawl (3 m IKMT) equipped

with a cod-end closing device. The fourchambered, electronically actuated sampler is described in detail by Aron, Raxter, Noel, and

Andrews (1964). The apparatus allows the collection of three discrete samples at a precise depth

and a fourth sample during retrieval of the net

from fishing depth to the surface. The gates that

separate the chambers are closed by means of a

solenoid-actuated triggering mechanism.

The experimental design for Ocean Acre included a regime of depths and times for seasonal

discrete-depth sampling. Twelve standard depths

were established for sampling the water column

during each cruise; each depth was sampled at

least once during full daylight hours and once during the night. Trawls that occurred one and one

half hours before and after sunset and sunrise

were considered to be twilight tows.

The Ocean Acre study area is located east of Bermuda and encompasses a 1° square area centered

at 32°N, 64°W. The site has relatively simple

oceanographic characteristics, at least in comparison with other deep-water localities in the western

Atlantic. The depth of the water in Ocean Acre

ranges from about 2000 m in the northwest corner

to 4500 m in the southwest region. The physical

oceanography is summarized here from Gibbs and

Roper (1970:120).

The Surface Water Mass occupies the upper

600 m; surface temperatures range from 20° to

29° C and salinity generally is above 36.4 %c. A

seasonal shallow thermocline occurs between 50 m

and 150 m during April to November, with a peak

intensity in August. A temperature reduction of

8° C can occur within the thermocline in a vertical

distance of 50 m. Below the thermocline temperature and salinity remain nearly uniform (18°-

19° C and 36.2° %0) to about 600 m. From about

600-1100 m a permanent deep thermohalocline

occurs that is identified as North Atlantic Central

Water. Temperature changes from 18° to 6° C

and salinity from 36.2-35.0 %c over this depth

range. Intermediate water is present between

1100 m and 2000 m with temperatures of 5.5° to

4.0° C and a salinity of about 35.0 %<-. Characteristic North Atlantic Deep Water, with temperatures

below 3.5° C and a salinity around 35.0

 (/(C, occurs

below 2000 m. Oxygen levels generally are between 4.5 and 5.5 ml/L at the surface and decrease

until about 700-900 m, where an oxygen minimum

layer occurs with concentrations around 3.5 ml/L.

Below 700-900 m, oxygen concentrations again increase to greater than 6.0 ml/L at 2000-3000 m.

Characteristics of the sound scattering layer are
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

difficult to define. With a 12 kHz sound source, a

weak and variable scattering layer usually is present at 0-150 in during the day, while a deep,

strong layer occurs between 400 m and 600 m. At

night the scattering layer is very prominent between 0 m and 250 m, and the 400-600 m layer,

though diminished and narrower, is still intact.

HAWAII STUDY

In Hawaiian waters the area of study lies about

seven miles off the leeward side of Oahu where

water depth exceeds 2000 m. Sampling gear was a

modified 10' Tucker trawl with an opening-closing

system in the mouth that is actuated by acoustic

signals from the ship. An open 3 m Isaacs-Kidd

midwater trawl also was used extensively.

Most oceanographic features off Hawaii are

very stable throughout the year. The seasonal

range in surface temperature is from 23° to 26° C.

A weak, shallow thermocline at about 35 m forms

during the spring and summer, while a permanent

thermocline extends from about 100 m to about

400 m where the temperature is around 8° C.

Temperature at 1000 m is 4° C and decreases to

2° C at 2000 m. Surface salinity is between 34.7 %c

and 35.2 %0; between 400 m and 500 m a strong

salinity minimum occurs (less than 34.2 %0),

which marks the core of North Pacific Intermediate Water. The very homogeneous Pacific Deep

Water lies below about 1500 m. Dissolved oxygen

falls to a minimum of about 0.7-1.0 ml/L at about

700 m. (Above discussion based on Gordon, 1970).

Subclass NAUTILOIDEA

NAUTILIDAE

FIGURE 24: BARS 1, 2

Nautilus occurs in tropical Indo-Pacific waters.

Species of the genus apparently are not truly

pelagic, as living specimens generally are captured

near the ocean bottom. Specimens commonly are

caught in bottom traps, and in captivity they spend

most of the day at rest on the bottom of the aquarium and swim chiefly at night (Bidder, 1962;

Stenzel, 1957). Nautilus tnacromphalus is caught

in shallow water at night and occasionally may be

taken at the surface by fishermen (Rancurel, pers.

comra.), The maximum depth to which any species of Nautilus can descend is uncertain. Denton

and Gilpin-Brown (1966:755) estimated that the

shell of N. macromphalus is sufficiently strong to

withstand hydrostatic pressures to a depth of 600

m, while Collins and Minton (1967) showed experimentally that the siphuncular tube can withstand pressures equivalent to 480 m. In New Caledonia N. macromphalus occurs at least as deep

as 200 m during the day (Rancurel, pers. comm.).

Bidder (pers. comm.) captured a female specimen

of N. pompilins in a trap at 200 m in the Bay of

Rabaul, New Britain, off Vulcan Island. The trap

had been set on sandy bottom for only a few hours

during the daytime. One specimen of N. pompilius

was obtained by the Challenger off Matuku Island,

Fiji, in a dredge haul to 576 m (Moseley, 1879:

256), but Bidder (pers. comm.) feels it may have

been caught at a shallower depth as the dredge

fished up the slope.

Subclass COLEOIDEA

Order SEPIOIDEA

SPIRULIDAE

FIGURE 24: BAR 3

Bruun (1943:23) investigated the vertical distribution of Spirula spirula and concluded that it

occurs between 200 m and 1750 m, except in areas

of upwelling where it is found as shallow as 100 m.

Bruun (1955) re-examined the distribution of

Spirula and maintained that the upper limit as

previously determined was correct. He revised his

original estimate of the lower limit of occurrence

of Spirula, and, based on inconclusive data, implied that this limit probably coincides with the

10° C isotherm. In attempting to correlate the

10° C isotherm with geographical distribution

Bruun stated (1955:68): "It is obvious that all

Spirula fall within the regions where the temperature is about 10° C or higher at a depth of 400 m."

(Actually, this is not at all obvious from the chart).

His use of the term "regions" refers to geographical

regions, but this has been interpreted as vertical

region, which led to the misconception that the

lower limit of the vertical distribution of Spirula

is 400 m (G. L. Voss, 1967:516). We are left with
NUMBER 209

Bruun's suggestion that the lower limit is the 10° C

isotherm or the lower boundary of what Bruun

termed the thermosphere; this would correspond to

a maximum depth in some areas of about 700 m

(Bruun, 1957:641).

Bruun (1943:10) found that pressures of 40-65

atmospheres (about 400-650 m) imploded Spirula

shells collected from the beach; he suggested that

shells in the living animal might be stronger.

Denton and Gilpin-Brown (1971:367) found experimentally that the chambered shells of Spirula

in the intact animal are able to withstand pressures

equivalent to those occurring at around 1700 m.

Recently Clarke (1969) presented data on the

vertical distribution of Spirula based on 70 specimens captured near the southern end of the Canary Islands. Using an Isaacs-Kidd midwater trawl

equipped with a Fox ton closing device, he showed

that Spirula undergoes vertical migration. Clarke

found the daytime center of distribution to be at

500-700 m, while at night Spirula ascended to

100-300 m. No specimens were taken shallower

than 500 m during the day, but a few were captured well below the center of distribution to

depths of 950 m.

SEPIIDAE

The sepiids are quasipelagic, generally burying

themselves in the bottom during the day and

emerging at night to feed. Therefore, they are beyond the scope of this report.

SEPIADARIIDAE, SEPIOLIDAE,

IDIOSEPIIDAE

Among these three families, only certain members of the Sepiolidae are wholly pelagic. Within

the Sepiolidae, the Rossiniae and the Sepiolinae

are benthic or quasipelagic and spend much of the

time buried in the substrate (Naef, 1923).

The remaining subfamily in the Sepiolidae, the

Heteroteuthinae (Figure 24:4-7) generally possess

silvery bands along the lateral surfaces of the

mantle and head. This feature in other cephalopod groups is usually associated with a pelagic

habit in the zone of light penetration. Little information exists, however, on the actual vertical

range of members of the subfamily.

The vertical distribution of the best-known

species, Heteroteuthis dispar, was studied by Degner (1925) in the Mediterranean Sea. Captures

were made in open nets and depths recorded only

as meters of wire out. We have considered the

depth of capture to be one-half the length of wire

out; this method gives a probable maximum fishing depth (see discussion in Bruun, 1943:21).

Degner, reporting on 124 specimens, found that

5.6 percent of the specimens were captured above

35 m, 72.5 percent were captured between 35-150

m, and 21.8 percent were captured below 150 m.

Unfortunately, the sizes of individual specimens

were not reported. They ranged from at least 2-12

mm ML (mantle length), with specimens from

about 2-4 mm ML apparently predominating.

Most specimens were captured over deep water.

Some of the specimens captured below 150 m are

very likely contaminants that were caught while

the open nets passed through the populated layer.

Clarke (1969:970) has also presented evidence

based on closing nets for a shallow living habitat

in H. dispar. He found that specimens of this species with mantle lengths greater than 2 mm (24

specimens) were captured during daytime at

depths of 110-255 m and at night near the surface

(0-70 m) to 265 m.

Our closing net captures from the Mediterranean also indicate the shallow occurrence of H.

dispar, although several specimens were captured

as deep as 400-500 m during the day. Four specimens were captured between 150 and 250 m and

10 specimens were captured between 400 and 500

m during the day. At night 16 specimens were

taken between 50 and 300 m (size range of all

specimens: 3-25 mm ML).

We have scattered observations on mature or

nearly mature specimens in two of the remaining

four recognized genera in the Heteroteuthinae

(Nectoteuthis, Iridioteuthis, Stoloteuthis, Sepiolina). We have recorded 12 specimens of Stoloteuthis leucoptera taken in five otter (bottom) trawls

that fished at the following depths (in meters):

145, 156, 161, 172, 188; one specimen was taken at

night in an open midwater trawl that fished at

350-0 m. Six specimens of Stoloteuthis maoria

were captured during daytime in two otter trawls

that fished on the bottom at 420-440 m and four

specimens were taken at night in an open midwater trawl that fished from 53-0 m over water
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

500-575 m in depth. Three specimens of Nectoteuthis pourtalesi were taken in two bottom trawls

that fished at depths of 394 and 385 m.

Although it is apparent from the above records

that Heteroteuthis occurs in the upper few hundred meters during both day and night, the full

extent of its range is still uncertain. Examination

of the photosensitive vesicles of a species of Heteroteuthis (H. hawaiiensis) has revealed a small

group of vesicles similar in size to those of the

shallow living Loligo vulgaris. This further suggests to us that Heteroteuthis is a shallow living

animal. The captures of H. dispar from 400-500 m

in the Mediterranean, however, are difficult to

interpret and remain unexplained until more extensive data are obtained.

Although Heteroteuthis probably is pelagic

throughout its life cycle, the habitats of the species

in the remaining genera are much less clear. The

rather meager data indicate that at least some of

these species occur at rather shallow depths. The

preponderance of specimens taken in bottom

trawls compared to midwater captures suggests

that these species spend considerable time on or

near the bottom.

Order TEUTHOIDEA

Suborder MYOPSIDA

LOLIGINIDAE

FIGURE 24: BARS 8-14

Members of the family are primarily neritic;

i.e., generally they are not found over deep water

and apparently are restricted to waters of the

coastal margins and continental shelves (Summers,

1969). Although all of the approximately 50

known species of loliginids have been recorded

from shallow water, details of vertical distributions are known for very few species. The records

of captures are based primarily on bottom trawls.

Summers (1969) records winter captures of Loligo

pealeii from Georges Bank to Cape Hatteras in

depths of 28-366 m with the highest concentration

at 110-183 m; in summer the species migrates inshore to spawn and occurs at depths as shallow as a

few meters.

Differences in captures during the day and at

night in winter suggest that the squid disperse

vertically into the water column at night. In late

winter L. pealeii is concentrated in the canyons of

the slope at 100-200 m where water temperatures

remain at 9° to 12° C (Vovk, 1969); with wanning

waters in spring the concentrations move northward and inshore. The northern limit of distribution is about 600 km further north in summer

than in winter (Summers, 1969).

Mangold-Wirz (1963:249) records depth distributions of the following species of loliginids from

the Catalane Sea: In L. vulgaris the depth at

which peak abundance occurs (40-120 m) varies

seasonally, being shallowest in the spring when

adults come inshore to spawn. Alloteuthis media

exhibits an inshore-offshore migration similar to

that of L. vulgaris. Total ranges are approximately

20-200 m. Results of exploratory fishing off the

northwest African shelf (Porebski, 1970) give an

indication of possible diel vertical movement of

Loligo vulgaris. Bottom trawls fished from 20-280

m yielded positive captures (abundance varied

with depth and locality) during daylight hours,

but after 1900 hours no L. vulgaris were captured

in the trawls, indicating that the animals either

leave the bottom and disperse into midwater at

night, or move to depths that were not sampled.

In the English Channel waters near Plymouth

L. forbesi is taken readily in bottom trawls during

the day at 13-104 m, but is absent from tows made

at night (Holme, 1974:496).

LaRoe (1967:50) found that Loligo (= Doryteuthis) plei in the tropical western Atlantic generally occurs from the surface to 180 m; Sepioteuthis sepioidea, a very shallow living tropical

species, is associated with coral reefs and turtle

grass, and is recorded from the surface to 21 m.

Lolliguncula brevis is recorded by LaRoe (1967:

84) from 85 m (one capture) to the surface, with

the great majority in less than 18 m.

Although many species of loliginids may be

seen at the surface at night over shallow depths,

the success of capture with bottom trawls suggests

that most species live near the bottom during the

day.
NUMBER 209

PICKFORDIATEUTHIDAE

FIGURE 24: BAR 15

This monotypic family, containing only Pickfordiateuthis pulchella, has been captured in shallow tropical waters from the surface to 3-4 m

(LaRoe, pers. comm.).

Suborder OEGOPSIDA

The great majority of truly oceanic species of

cephalopods are included in the Oegopsida. The

group is very diversified and contains over half of

all families and genera of living cephalopods.

Members of the Oegopsida inhabit a wide variety

of oceanic habitats. Some live at or near the surface; some undergo diel vertical migrations; some

are restricted to the depths; some undergo shoreward seasonal migrations, and some are associated

in part with the ocean bottom.

LYCOTEUTHIDAE

FIGURE 25: BAR 1

G. L. Voss (1962:273) suggested that members

of the family occupy the zone from 300-600 m.

Oregoniateuthis springeri is recorded from a single

open net (367-0 m).

Lycoteuthis diadema is recorded from seven

captures of adults in open nets that fished at (in

meters): 366 (2 stations), 403, 589, 1500, 2195, 3000,

and two captures of larvae at 46 m and 57 m (G. L.

Voss, 1962:275-277). Lampadioteuthis megaleia is

recorded from one adult taken at night in an open

net that presumably did not descend below 200 m

(400 m of wire out) and two larvae (Young, 1964).

Selenoteuthis scintillans was previously known

from two records: 46 m at night in an IKMT and

0-3290 m (G. L. Voss, 1962:294). Some closing net

data on S. scintillans are available from off Bermuda (9 specimens, 11-32 mm ML). Five specimens were captured from 610-650 m during the

day and four specimens were taken at 90-130 m

at night.

Selenoteuthis scintillans apparently undergoes

an extensive diel vertical migration. While data on

other species are almost completely lacking, we suspect that vertical migratory behavior may be typical of the group.

ENOPLOTEUTHIDAE

FIGURES 2-5, 25: BARS 2-15

Some members of the Enoploteuthidae are

among the most commonly caught squids in midwater trawls. Three subfamilies are known, and

within each, the assemblage of species shows considerable homogeneity in size and appearance.

The subfamily Enoploteuthinae consists of four

genera: Enoploteuthis, Abralia, Abraliopsis, and

Watasenia.

We have data from California, Bermuda, and

Hawaii on three species of Abraliopsis.

Nine hundred and thirty specimens of Abraliopsis felis from open nets were examined from

California, of which 480 specimens were used in

plotting the vertical distribution; the remainder

were too small (less than 15 mm ML) or were

captured during twilight. During the daytime only

two specimens were captured above 300 m; 93 percent of the specimens were captured between 300

and 600 m with a strong peak between 400 and 500

m (Figure 2). The specimens listed below 600 m

probably are contaminants. The exact nighttime

position of the population is less clear. The specimens (27 percent) located below 500 m probably

are contaminants. Seventy-five percent of the remaining specimens are located in the upper 300 m

and 62 percent in the upper 200 m. It is uncertain

how many of the specimens taken between 200 m

and 500 m are contaminants, so it is difficult to put

a lower limit on the major portion of the population at night. The data show that Abraliopsis felis

undergoes a diel vertical migration covering an

average distance of over 300 m, ascending from a

mean daytime depth of 450 m to a mean nighttime depth of 100 m.

A great disparity exists between the number of

specimens captured during the daytime and at

night. Perhaps members of this species school during the day and break up into small groups or

completely dissociate at night in the shallower

water. Trawl data support this suggestion. Trawls

that fished at day below 300 m occasionally caught

large numbers of A. felis (e.g., 93, 63, 54, 44 specimens) while a maximum of eight specimens was

taken in a single trawl that fished above 300 m at

night.

Pearcy (1965) found that A. felis (as Abraliopsis sp.) off Oregon shows a greater frequency of
10

occurrence in the surface layer (0-200 m) during

the night than during the day; while in the layer

of 0-500 m, occurrence is greater during the day

than during the night. These data suggest a distributional pattern similar to that for the same

species off California.

Off Bermuda, closing-net captures of Abraliopsis pfefferi (15-27 mm ML) during the daytime

occur at 610-650 m (three specimens) and at night

four specimens larger than 15 mm ML were captured at 50-100 m. Therefore an extensive vertical

migration also is indicated for this species.

Ten specimens of Abraliopsis sp. (9-20 mm

ML) off Hawaii were captured in closing nets during the day between 500 m and 600 m. At night

41 specimens (19-20 mm ML) were taken in open

nets; nearly 75 percent of these were captured in

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tows that fished between 50 and 100 m. One specimen (16 mm ML) was taken in a closing net at

60 m.

Information on the other genera in the subfamily is scarce. Berry (1926) recorded Abralia

veranyi from Madeira; specimens were observed

and captured in Funchal Harbor at night during

summer, but were absent during the day. We also

have dipnetted A. veranyi at night in Bahamian

waters. A number of specimens of several species

of Abralia also have been taken at or near the surface presumably at night (Clarke, 1966:170). These

observations suggest that species of Abralia probably migrate vertically to or near the surface at

night and descend during the day.

Off Bermuda Abralia is infrequently caught.

Two closing trawl captures were made of A. redNighf

1200

120 160 200 0

No. of Specimens

FIGURE 2.—Vertical distribution of Abraliopsis felis off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)
NUMBER 209 11

fieldi at night at 50 m. Daytime captures are absent in spite of intensive sampling from the surface

to 750 m.

Abralia astrosticta from off Hawaii is known

from only a few captures. Although small specimens have been taken in midwaters, most large

specimens have been taken in bottom trawls (L.

Burgess, National Marine Fisheries Service Laboratory, Honolulu, pers. comm.). One adult (48

mm ML) was taken in a gill net on the ocean bottom at night in 180 m. Extensive pelagic trawling

around Hawaii has failed to capture any adults.

The meager data suggest that this species may

associate with the ocean bottom off oceanic islands.

Off Hawaii three specimens of A. trigonura

(30-37 mm ML) were captured in a closing net

that fished between 460-600 m during the day. In

open nets nine specimens (15-37 mm ML) were

taken between 490 and 575 m during the day and

a single specimen, probably a contaminant, was

taken from an open net that fished at 775 m. At

night, two specimens (22 mm and 37 mm ML)

were captured from 100-0 m.

Sasaki (1914:103) studied the distribution of

Watasenia scintillans in Japan and concluded that

in the spring and summer it lives deeper than 200

m during the day and in the evening it approaches

the shore or shallow sea floor to spawn. This is a

spawning migration; whether W. scintillans in

addition undergoes an oceanic daily vertical migration is as yet unknown to us.

Roper (1966) presented capture data on Enoplotenthis anapsis, which included six juvenile and

adult specimens (17-79 mm ML) that were cap

tured at night between 0-120 m. One Bermuda

specimen (15 mm ML) was taken in a closing net

at 90 m at night. Three additional specimens of

Enoploteuthis sp. were taken in closing nets at

night; two specimens at 90 m and one at 130 m.

No daytime closing-net captures have been made

to date.

The Pyroteuthinae contains two genera, Pyroteuthis and Pterygioteuthis. Pyroteuthis margaritifera is one of the most abundant squids taken in

the Ocean Acre program off Bermuda; 163 specimens (5-27 mm ML) were captured in closing

nets and their distributions are plotted on a timedepth chart (Figure 3). The boxed digits indicate

the number of specimens caught at a given time

and depth per hour of trawling throughout the

PYRUTEUTH1S MARGARITIFERA

50

100

150

200

250

300

350

'•OH

450

500

55 0

500

6

r

)0

700

750

P00

850

900

950

1000

1050

1100

1150

1200

1250

1300

1350

1*00

1450

1500

rr

NUMBER OF SPECIMENS PfcR HOUR IS DISCRETE DEPTH SAMPLES FROM ACRE 1

NUMBER OF SAMPLES 328 NUMBER OF SPECIMENS 163

BERMUDA STANDARD TIME

12:00 14:00 16:00 18:00 20:00 22:00 24:C

4 6 10 11

0 I 0 I i

0 0

[3 ED CD

cn

o

o I I DO

Q] QJ Q]

o

E2 QH CD1

o I

Ell

o o o

0 0

o I

FIGURE 3.—Vertical distribution of Pyroteuthis margaritifera off Bermuda. Time-depth plot of

positive and negative captures and sampling effort. (See page 11 for explanation of symbols.)

CD

0
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

24-hour cycle, as well as the time-depth location

of samples that caught no specimens of P. margaritifera (indicated by a zero); blank spaces represent time-depth loci that were not sampled. During

the day over 80 percent of the 43 specimens cap

tured came from 375-500 m. Sixty-two specimens

were captured at night from 75-175 m. Pyrpteuthis margaritifera, therefore, undergoes a diel

vertical migration of about 300-350 m.

During the daytime there is no size stratification

with depth. At night a possible size stratification

may occur but data are inconclusive. During the

day only a single adult was captured while at night

three were captured.

In Pterygioteuthis 130 specimens of P. gemmata

from open nets were available for analysis from

California after elimination of "larval" specimens

less than 11 mm ML and twilight captures (Figure

4). Of the 56 daytime captures, 3 percent were

taken in the upper 300 m; 56 percent were taken

from 300-600 m; 25 percent from 600-900 m; and

16 percent from 900-1200 m. A small peak is present at 700-800 m as a result of a single trawl

which likely captured the specimens during retrieval through the upper layers. The daytime capNight

Night

12°°- -- 20 40

No. of Specimens

FIGURE 4.—Vertical distribution of Pterygioteuthis gemmata

off southern California. (Black bar = number of specimens

captured; white bar = correction factor.)

600

0 2 0 2 4 6

Specimens Per Hour

FIGURE 5.—Vertical distribution of Pterygioteuthis giardi off

Bermuda. (Black bar = number of specimens captured.)

tures predominate between 300 and 600 m, and

the catches below 600 m presumably represent

contamination.

The nighttime distribution shows 88 percent of

the 74 specimens in the upper 200 m. The few

captures below this level probably are contaminants. Pterygioteuthis gemmata from California,

therefore, appears to undergo a diel migration of

about 300-400 m in vertical extent.

Another species of Pterygioteuthis, P. giardi,

has been taken during the Ocean Acre cruises off

Bermuda. One hundred four specimens of P.

giardi were captured in closing net samples (Figure 5). During daytime, six specimens (8-16 mm

ML) were captured, five from 327-475 m and one

at 260 m (9 mm ML). At night 70 specimens (5-16

mm ML) were captured between 50 and 175 m; all

but three were captured between 50 and 100 m,

and over 80 percent of the captures occurred at

50-60 m. Pterygioteuthis giardi, therefore, undergoes a diel vertical excursion of 250-300 m. There

is no apparent segregation of sizes during day or

nighttime periods.

It is difficult to explain the large disparity between number of individuals captured in day vs

night trawls. It is possible that there is an avoidance of the net during the daytime due to the

greater light intensities or that the discrepancy reflects a greater dispersion of individuals during the

day.

The Ancistrochirinae contains two species, one

of which, Thelidioteuthis alessandrinii, may be a

synonym of the other, Ancistrochirus lesueuri.
NUMBER 209 13

Four larval and juvenile specimens (15-33 mm

ML) of T. alessandrinii were captured by the

Dana Expeditions on four separate occasions in

less than 100 m at night. Two specimens (20 mm

and 21 mm ML) were taken at night off Bermuda

in closing nets in less than 125 m. One specimen

(35 mm ML) was taken in an open net at 80 m at

night off Hawaii. The nighttime distribution of

juveniles, at least, appears to be in the near-surface

layers.

The preceding data on the family Enoploteuthidae demonstrate that several species undergo

diel vertical migration of several hundred meters.

Where known, all stages beyond the larval stage

for a particular species appear to occupy the same

depth zones; even individuals as small as 5 mm

ML (e.g., Pyrotenthis margaritifera) may migrate

with adults. While daytime captures are few or

lacking for many species, nighttime captures of

individuals of many other species in near-surface

waters suggest that the migratory behavior demonstrated in a few species probably is very widespread

in the family. Indeed, no evidence exists for nonmigratory behavior of any enoploteuthid species.

OMMASTREPHIDAE

FICURE 26

We have relatively little additional information

on the vertical distribution of this large and important family since Clarke (1966) reviewed much

of the literature covering the subject.

Ommastrephes caroli, O. pteropus, and O. bartramii are frequently dipnetted at the surface at

night (Clarke, 1966; Roper, 1963; Young, 1972a).

A tentacle of O. caroli was caught by a reversing

water bottle at 1490 m, presumably during the

night, and photographs, apparently of this species,

were taken at night at 600 and 700 m (Clarke,

1966:108-109). Little is known about their daytime distribution. Baker (1960) reported that

specimens of O. pteropus (or O. caroli, see Clarke,

1966:109) were photographed and captured on

long lines at 1000 m at night at the same time

that other individuals of these species were being

captured at the surface. These are prominant species at the surface at night except during periods

of bright moonlight and rough seas (Baker, 1960).

Arata (1954) observed squid, thought to be O.

bartramii, leaping from the water and gliding for

some distance over the water surface during the

daytime.

Symplectoteuthis oualaniensis commonly is

found at the surface at night (Clarke, 1966:115).

Symplectoteuthis luminosa has also been reported

from the surface at night (Young, 1972a: 34).

Nothing is known of the daytime distribution of

these species.

Hyaloteuthis pelagica, distributed world-wide in

warm waters, is a relatively infrequently captured

ommastrephid. Off Bermuda, four specimens

(12-24 mm ML) have been taken in closing nets

at night at 100 m. Open-net captures with a large

Engel midwater trawl resulted in the capture of

85 specimens (10-90 mm ML, mostly 40-90 mm

ML), 78 of which were taken at night; the remaining 7 were twilight captures. At night 56 specimens

were taken at 150-0 m; the remaining 22 specimens

taken in trawls fishing at various depths to 1000 m,

probably are contaminants.

Dosidicus gigas commonly is seen at the surface

at night (Clarke, 1966:117). We have seen motion

pictures of this species leaping from the water

during the day off Chile and attacking fishing

lures at the surface during the day off Peru.

Ornithoteuthis antillarum has been reported at

the surface at night (G. L. Voss, 1957). Sasaki

(1929:285) reported that O. volatilis is referred to

as flying squid by Japanese fishermen who report

that it can glide over the surface of the sea.

Data are available on 43 adult specimens of O.

antillarum from the western tropical Atlantic and

Caribbean; all captures were made in bottomfishing otter trawls (21 tows) at depths of 585-

1100 m (mostly 640-825 m), and all but three were

captured during the day. The three possible nighttime captures were made in nets that fished

through twilight periods as well as night. In the

eastern south Atlantic, many specimens of O. cf.

antillarum were captured at night over deep

water in Engel trawls that fished between 100 and

600 m. The significant preponderance of daytime

vs nighttime captures in bottom trawls and the

occurrence of specimens in midwater and at the

surface at night suggest that Ornithoteuthis spends

the day on or near the bottom and spreads out

into middepths at night.

Illex illecebrosus generally is caught in shallow

water 15-30 m deep during the summer. At the
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

northern end of its range, off Newfoundland,

these squid are occasionally seen at the surface

(C. C. Lu, pers. comm.). Clarke (1966:119) stated

that Illex illecebrosus is most numerous near the

coast and probably lives on the upper slope and

shelf.

Illex illecebrosus off the Chesapeake Bight has

been captured with otter trawls at depths of

45-265 m; nearly all captures occur during the

daytime indicating that specimens leave the bottom and disperse into midwater at night (C. C. Lu,

pers. comm.). Milliman and Manheim (1968) observed dense concentrations (estimated to be 500

per 103m

3

) of squid off Cape Hatteras, North

Carolina, thought to be Illex illecebrosus, from a

submersible at depths of 220-250 m during the day

over a bottom depth of 700 m. C. R. Robins (University of Miami, pers. comm.) observed considerable numbers of squid, thought to be a species of

Illex, from a submersible at approximately 900 m

during daytime off Miami, Florida. Some of the

squid were resting on the bottom while others were

swimming immediately above the bottom.

Illex coindetti is closely associated with the bottom where it is fished at depths of about 40 to

500 m depending somewhat on the locality and

season (Mangold-Wirz, 1963:157). In the Gulf of

Guinea, /. coindetii has been captured in otter

trawls from 80-410 m during the daylight hours;

at night the specimens apparently leave the bottom and disperse into middepths (C. C. Lu, pers.

comm.). Illex oxygonius from the western Atlantic has been taken from 50-550 m with bottom

trawls (Roper, Lu, and Mangold, 1969:219); no

information is available on day-night distributions.

Todaropsis eblanae also is associated with the bottom and is fished at depths from 85-660 m

(Mangold-Wirz, 1963:168).

Todarodes sagittatus frequently is found at the

surface during the night, but during the day it can

be fished with bottom trawls from 70-800 m

(Clarke, 1966:129). It has been photographed at

depths of 1000 m presumably far off the bottom

(Clarke, 1966:129). Clarke and Merrett (1972)

record fragments of T. sagittatus from the stomach

of the bottom-living shark, Centroscymnus coelolepis, captured at 1975 m.

Todarodes pacificus commonly is caught by jig

and line at 20-30 m from the surface at night.

During the day these squid descend to around

100 m where they are occasionally fished by hand

line (Suzuki, 1963:143).

Little information is available on the vertical

distribution of species of the genus Nototodarus.

Nototodarus sloani gouldi has been taken by dipnets (Cotton, 1942). In Hawaii, N. hawaiiensis appears to be associated with the ocean bottom as

evidenced by occasional bottom photographs (400

m by R. Grigg, University of Hawaii), otter trawl

captures (400-570 m, ALBATROSS stations, vide

Berry, 1914), and trap captures. During late summer months it can be captured at the surface at

night off the island of Hawaii where it supports a

minor dipnet fishery.

The ommastrephids exhibit a wide variety of

vertical distributional patterns. Some species (e.g.,

Ommastrephes caroli) can roam from the surface

to great depths. Others appear to be restricted to

near surface waters (e.g., 7". pacificus). A number

of species are associated with the bottom, but approach the surface nightly during all or part of

the year (e.g., T. sagittatus, N. hawaiiensis). Other

species may move into shallow water seasonally

during feeding migrations (e.g., /. illecebrosus).

HISTIOTEUTHTOAE

FIGURES 6, 27: BARS 1-4

The Histioteuthidae consists of 13 species in the

single genus Histioteuthis (N. Voss, 1969). Since

most species are represented by relatively few

specimens captured in open nets from scattered

localities, information on vertical distribution is

sketchy.

Histioteuthis heteropsis from the California

study is represented by 191 specimens larger than

6 mm ML taken in open nets (Figure 6). During

the daytime 90 specimens were taken, none of

which occurred in the upper 300 m; 3 percent

were taken in 300-400 m, 13 percent in 400-500

m, 38 percent in 500-600 m, 24 percent in 600-

700 m, and 10 percent in 700-800 m. The remaining 12 percent were scattered between 800 and

1200 m and probably represent contamination.

The upper limit for the majority of the daytime

captures is 400 m and the lower limit lies around

800 m. Peak abundance occurs between 500 m and

700 m.

At nighttime 69 percent of the 101 specimens
NUMBER 209 15

Night

1200

30 0 10

No. of Specimens

FIGURE 6.—Vertical distribution of Histioteuthis heteropsis off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)

captured were taken in the upper 400 m with a

peak at 300-400 meters; 21 percent from 400-800

m, and 10 percent from 800-1200 m. Possibly all

specimens from below 400 m are contaminants.

The shallowest capture at night for specimens

larger than 20 mm ML was 200 m and most were

concentrated at 300-400 m. This species undergoes a diel vertical migration of 300 to 400 m.

From off Bermuda closing net captures of H.

dofleini were made during the day at 510 m (14

mm ML) and at night at 50 m (12 mm ML). One

H. meleagroteuthis (16 mm ML) was captured at

690 m during the day and another (32 mm ML)

was taken at 360 m at night. Two specimens of

H. corona (25 mm and 27 mm ML) were captured

during the day at 630 m. We suspect that a diel

vertical migration pattern probably is widespread

in this family.

PSYCHROTEUTHIDAE

The family consists of a single genus, Psychroteuthis, which contains two species (one undescribed) that occur in Antarctic waters and,

possibly, a third species from off the Bonin Islands

(Roper and Young, 1972:220). A large specimen

was taken through an ice hole in the Antarctic

and other specimens are known from midwater

trawl captures. Nothing definitive, however, is

known of the vertical distribution of these species.

NEOTEUTHIDAE

FIGURE 29: BAR 1

Two genera are known in the Neoteuthidae,

Neoteuthis and Allurotenthis. Allnroteiithis antarcticus is a pelagic species abundant in Antarctic

waters, but little information currently is available

on its vertical distribution.

Neoteuthis thielei is known from only a few

specimens that have been captured in open midwater trawls. Off California one specimen (83 mm

ML) was taken at 1333-0 m. In the eastern tropical Atlantic eight specimens (ca. 36-125 mm ML)

of Neoteuthis sp. were taken at night in open
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Engel trawls that fished at 1750-2000 m; one specimen (ca. 130 mm ML) was taken at 1300-0 m at

night. In the Gulf of Mexico a specimen (ca. 50

mm ML) was taken at night at 900 m in a closing

net.

ARCHITEUTHIDAE

The Architeuthidae consists of 20 nominal

species of Architeuthis, most of which are undoubtedly synonymous. Although the giant squid

has received a great deal of attention in the literature, very little is known of its vertical distribution or of any other aspect of its biology. Occasionally Architeuthis has been sighted (F. A.

Aldrich, Memorial University of Newfoundland,

pers. comm.) or captured alive at or near

the surface (Verrill, 1882a: 181), but these sightings may be unrepresentative of its habitat. The

great majority of specimens come from strandings

or whale stomachs. Perhaps the most significant

capture is that reported by Cadenat (1936) of a

specimen taken in a bottom trawl that fished at

about 200 m. Clarke and Merrett (1972) record

fragments of Architeuthis sp. from the stomach

of the deep-sea shark, Centroscymnus coelolepis,

which was captured on a bottom-set long line at

1246 m.

Two juvenile specimens of Architeuthis were

taken from stomach contents of the oceanic fish

Alepisaurus ferox (Roper and Young, 1972); this

fish is thought to hunt primarily in the upper 300

m of the open ocean (Rancurel, 1970:82), so it

seems probable that the young Architeuthis occur

within this range.

BATHYTEUTHIDAE

FIGURES 7, 27: BARS 5-7

The Bathyteuthidae consists of three species in

the genus Bathyteuthis. Details of the distribution

of these bathypelagic species are reported by

Roper (1969).

Bathyteuthis abyssicola is a very commonly

caught species in Antarctic waters and it occurs

less abundantly in other areas of the world ocean;

its maximum distribution appears to be directly

associated with areas of relatively high organic

productivity at the surface of the sea and with

waters of l°-5° C. In the Antarctic 563 specimens

were analyzed (Figure 7). All specimens were

captured in open net trawls; 87 percent of the

captures occurred between 1000 and 2500 m, and

the remainder were taken between 500 and 1000

m. A very few captures occurred below 2500 m

and are considered to be contaminants. Of the

total captures, 60 percent (340 specimens) occur

between 1500 and 2500 m.

No diel vertical movements are known to occur.

Larger animals tend to occur at greater depths

than smaller individuals. Sixty-one percent of the

specimens less than 300 mm ML live between 500-

1500 m, while 72.5 percent of those larger than

30 mm ML live between 1500-2500 m.

One specimen of B. abyssicola (30 mm ML)

2750

3000

40 60 80 100 120 140

No. of Specimens

FIGURE 7.—Vertical distribution of Bathyteuthis abyssicola

from Antarctic waters. (Black bar = number of specimens

per standardized 2-hour tow.)
NUMBER 209 17

was captured in a closing net that fished between

830 and 975 m during the day off Hawaii.

Bathyteuthis bacidifera has been taken between

680 and 1550 m from the tropical eastern Pacific

in open nets. Bathyteuthis berryi from off California has been captured between 800 and 1200 m

in open nets and probably occurs much deeper,

but data on deep water captures are lacking.

GONATIDAE

FICURES 8-12, 27: BARS 8-14

Members of the Gonatidae inhabit primarily

high latitude waters (Nesis, 1971a). Three genera

presently are known; Gonatus, Gonatopsis, and

Berryteuthis, although recent work has made the

generic boundaries uncertain (Okiyama, 1969;

Fields and Gauley, 1971; Nesis, 1971b).

Gonatus onyx was the most abundant species

taken in the California study. Seven hundred and

seventy-four specimens over 21 mm ML from open

trawls were used in determining patterns of vertical distribution (Figure 8).

Less than 2 percent of the 438 daytime captures

were shallower than 400 m; 87 percent of the

specimens were taken in a zone between 400 and

800 m. Specimens captured below 800 m are probably contaminants.

At night 3 percent of the 336 specimens were

taken in the upper 100 m; 11 percent in 100-200

m, 5 percent in 200-300 m, 18 percent in 300-400

m, and 48 percent in 400-500 m. The remaining

12 percent below 500 m probably represent contamination. Thirty-seven percent of the nighttime captures were taken from the water above the

upper limit of the daytime captures (400 m), and

the lower limit of the nighttime distribution appears to be 300 m shallower than the lower limit

of the daytime distribution.

This species apparently shifts upward at night

primarily into the 300-500 m zone, but ranges

into the upper 100 m. Trawls that caught Gonatus

onyx during the day between 400 and 800 m averaged 16.8 specimens per tow. At nighttime between 300 and 500 m the average catch was 16.0,

while between 0 and 300 m the average was only

4.0.

Gonatus californiensis was the least abundant

member of the genus in the California study. Of

the 99 specimens larger than 20 mm ML, 46 were

twilight captures unsuitable for plotting diel verNight

1200

0 40 80 120 0 40 80

No. of Specimens

FIGURE 8.—Vertical distribution of Gonatus onyx off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)

160 200 240
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tical distribution. The data are difficult to interpret and for this reason the figure showing total

captures (day, night, twilight) is included with the

day and night distribution figures (Figure 9).

During the daytime no specimens were captured

above 400 m. Seventy percent of the 30 daytime

captures were taken in the zone from 400-800 m

with the greatest proportion, 30 percent, occurring

at the 700-800 m level. The remaining 30 percent

is scattered from 800-1200 m and may represent

contamination. Even though the daylight captures

are few, the upper and lower limits of the daytime

distribution appear to be at 400 and 800 m, respectively, if twilight captures are considered.

Eighty-eight percent of the combined daytime and

twilight captures occurred between 400 and 800 m.

Only 23 specimens were suitable for the nighttime analysis. Thirty-five percent were taken from

100-400 m and 61 percent from 400-800 m; a

slight peak (48 percent of the night specimens)

occurs between 300 and 500 m. While the number

of specimens is inadequate to allow definite conclusions, this species seems to undergo an upward

shift at night.

Eighty-eight specimens of Gonatus pyros over

20 mm ML are available from the California

study (Figure 10). Only a single specimen was

captured above 400 m during the daytime; 86

percent of the 52 daytime captures were taken in

a zone between 400-700 m. The few specimens

captured below 700 m probably are contaminants.

The daytime distribution of this species appears to

be between 400 m and 700 m.

Thirty-six specimens were taken during the

nighttime; 14 percent in 100-200 m, 6 percent in

200-300 m, 42 percent in 300-400 m, and 17 perDay, Night, Twilight Day Night

1200

10 20 40 0

No. of Specimens

0 10

FIGURE 9.—Vertical distribution of Gonatus californiensis off southern California.

(Black bar — number of specimens captured; white bar = correction factor.)
NUMBER 209 19

cent in 400-500 m. The remaining 21 percent of

the specimens below 500 m possibly are contaminants. Therefore, the limited data suggest that

the nighttime distribution has an upper limit at

100 m and a lower limit at 500 m, although most

captures were at 300-400 m; this indicates that

the species shifts upward at night.

Eighty-seven specimens of Gonatus berryi from

off California were suitable for plotting vertical

distribution. Although fairly common, this species

is never caught in large numbers. The larvae of

G. berryi differ morphologically from the larvae

of the other three species of Gonatus in the area

studied, and this is reflected in their deeper living

habit. The distribution of larvae (6-15 mm ML)

shows a peak between 300 and 700 m (Figure 11).

This range is slightly shallower than that found

for the larger animals.

No subadult specimens were captured in less

than 300 m during the daytime. A single specimen

each was captured in the 300-400 m zone and the

400-500 m zone; 68 percent of the catch was in

the 500-800 m zone, while the remaining specimens below 800 m probably represent contamination. Therefore, the daytime distribution seems to

lie between 500 and 800 m. A large number of

captures in the 400-500 m zone was made in tows

that extended into the twilight period; many of

these probably entered the net during daytime, so

the upper limit may be at 400 m instead of 500 m.

At nighttime no specimens were taken in the

upper 100 m; one was taken in 100-200 m, three

in 200-300 m, and one in 300-400 m. The greatest

concentration is in the 400-500 m zone where 33

percent of the specimens were taken, while 30 percent were taken between 500-800 m. While there

might be a slight upward shift in the peak of

abundance at night, it is not as apparent as in the

other species of Gonatus. The number of captures

attributed to depths below 800 m during both

daytime and nighttime is considerably higher than

is found in the other three species of Gonatus.

Although much of this undoubtedly is contamination, a small percentage of the population probably extends to the lower limit (1200 m) of the

sampling program. Church (1971) observed a

specimen of Gonatus, apparently berryi, from a

deep-diving submersible at 915 m.

Gonatopsis borealis from off California was represented by 74 juvenile specimens between 16 and

47 mm ML (Figure 12). No specimens were cap

tured during the daytime in the upper 300 m and

only a single specimen was taken in the 300-400 m

zone. Ninety percent of the daytime captures occurred at 400-700 m, while 8 percent, probably

contaminants, were captured below 700 m. Therefore, the distribution in the daytime seems to be

between 400 and 700 m.

Only a few specimens were captured at night;

three in the upper 200 m, six in 200-400 m, one

in 400-500 m, and three, probably contaminants,

in 800-1000 m. The scanty evidence suggests that

the nighttime distribution is in the upper 400 m.

If this is correct, G. borealis undergoes vertical

migration of 200-300 m, at least during the juvenile stage.

Fourteen specimens of Berryteuthis magister are

available from the Auke Bay region of Alaska.

Some specimens were captured in otter trawls at

depths that ranged from 165-310 m during both

day and night. The specimens had the following

mantle lengths (in mm): 69, 73, 100, 152, 192,

200, 210, 250. Three specimens (57, 75, and 90 mm

Night

1200

10 20 0 10

No. of Specimens

FIGURE 10.—Vertical distribution of Gonatus pyros off southern California. (Black bar = number of specimens captured;

white bar = correction factor.)
20

ML) were taken in Isaacs-Kidd midwater trawls

at night between 3-5 m, and 3 others (62, 124, and

173 mm ML) were taken at or near the surface in

a herring seine and salmon traps. Sasaki (1929:272)

records B. magister from 180, 725, 820, and 1000 m,

in bottom trawls off Japan, Korea, and the Bering

Sea.

Berryteuthis anonychus has been recorded (as

Gonatus anonychus) only from specimens that

were captured at the surface at night off Oregon

(Pearcy and Voss, 1963).

Three of the four species of Gonatus (G. pyros,

G. onyx, G. californiensis) for which we have data,

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

exhibit very similar distributional patterns, in

which they appear to undergo a slight upward

shift in their vertical distributions at night. The

absence of strong vertical migrations in these species is somewhat surprising, because they are

among the more muscular midwater squids. It is

possible that the fourth species (G. berryi) does

not migrate vertically. Juvenile Gonatopsis berryi

seem to exhibit a more pronounced upward shift,

but nighttime data are insufficient to confirm this.

Berryteuthis magister has been taken many

times in bottom trawls and probably spends some

time associated with the ocean bottom.

LARVAE SUB-ADULTS

Day, Night, Twilight Day, Night, Twilight Day Night

20 40 0 20 40 0 10

NO. OF SPECIMENS

10 20

FIGURE 11.—Vertical distribution of Gonatus berryi off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)
NUMBER 209 21

Night

1200

0 10 20 30 0

No. of Specimens

FIGURE 12.—Vertical distribution of Gonatopsis borealis off

southern California. (Black bar = number of specimens

captured; white bar = correction factor.)

OCTOPOTEUTHIDAE

FIGURES 13, 28: BARS 5, 6

The Octopoteuthidae consists of two genera,

Octopoteuthis and Taningia.

Eighty-three specimens larger than 15 mm ML

of Octopoteuthis deletron from California were

analyzed (Figures 13, 28:bar 5). Only about half

as many specimens (26) were captured during the

daytime as at night. During the day, no specimens

were captured in the upper 200 m. The greatest

number of captures occurs between 300 and 400 m

where 13 specimens (50 percent) were taken

(when the amount of trawling at different depths

is equalized, the greatest abundance would be at

200-300 m). Three specimens each were taken in

400-500 m, 500-600 m, and 600-700 m; a single

specimen was taken in 700-800 m, and three between 1000 and 1200 m. The daytime upper limit

is clearly at 200 m, but the lower limit is more

difficult to determine.

Fifty-seven specimens were captured at night,

the majority of which were fairly evenly distributed throughout the upper 500 m; 23 percent in

the upper 100 m, 14 percent in 100-200 m, 11 percent in 200-300 m, 21 percent in 300-400 m, and

12 percent in 400-500 m. The remaining 19 percent was distributed from 600-1200 m and probably represents contamination from shallower

zones.

Certainly additional information is needed, but

the available data indicate that the species does

not undergo a "typical" diel vertical migration,

but seems to spread out over a greater range of

depths during the night than during the day.

The largest Octopoteuthis (20 mm ML) taken

off Bermuda in closing nets was captured at 230 m

at night.

Little information exists on Taningia danae, the

only species in the genus. Taningia danae grows to

a large size of at least 140 cm ML (Clarke, 1967,

table 1). A few juveniles have been captured at

night within 175 m of the surface (Clarke, 1967:

127). Remains of a large T. danae, estimated to

approach 50 kg, were taken from the stomach of

the bottom-living shark, Centroscymnus coelolepis,

Night

1200-

10 20 0 10

No. of Specimens

FIGURE 13.—Vertical distribution of Octopoteuthis deletron

off southern California. (Black bar = number of specimens

captured; white bar = correction factor.)
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

captured on a bottom-set long line at 1246 m

(Clarke and Merrett, 1972). Otherwise, nothing

definitive is known of the distribution of the

larger animals.

ONYCHOTEUTHIDAE

FIGURE 28: BARS 2-4

The family Onychoteuthidae contains five genera, the best known of which is Onychoteuthis.

Onychoteuthis banksii occasionally is found on

the decks of ships in the morning (Rees, 1949)

and individuals are frequently observed and dipnetted at the surface at night (personal observations). The animals are powerful swimmers and

adults only rarely are caught in nets.

Off Bermuda, 52 specimens of Onychoteuthis

banksii (5-57 mm ML) were taken in closing net

samples. Captures were made from the surface to

a maximum depth of 150 m; all but two specimens

were taken in the 50-100 m zone both day and

night. The largest specimen was taken at the surface at light. All daytime captures from Ocean

Acre are small individuals which may not have

adopted the juvenile or adult pattern of distribution; thus little can be determined concerning

daytime distribution. However, a specimen of 20

mm ML was taken from a depth of 800 m in midday in the Mediterranean in a closing trawl. Although it is possible that this was a contaminant,

examination of the photosensitive vesicles of

Onychoteuthis, which are rather elaborate, suggests that this animal is not restricted to epipelagic

regions.

Onykia caribaea is a relatively small species that

has been taken primarily at the surface both during the day and at night. It is commonly associated

with Sargassum, the floating seaweed (Lesueur,

1821; Bullen, 1899; personal observations). Onykia

caribaea has a bluish coloration on the dorsal surface that is characteristic of species of the neuston

(Herring, 1967:220). One specimen of O. caribaea

(34 mm ML) from St. Helena was regurgitated

from the stomach of a Fairy tern which feeds only

at the surface.

Ten closing net captures of juvenile O. caribaea

have been made during the Ocean Acre cruises

and all specimens were taken at the surface both

during the day and at night.

Chaunoteuthis mollis is known from only a few

specimens, several of which were taken dead at the

surface (Clarke, 1966:149).

The vertical distribution of Ancistroteuthis lichtensteini, a rarely caught species, cannot be determined on the basis of the scant information

available.

Several nominal species of Moroteuthis exist,

the best known of which is M. robusta from the

north Pacific. Records are available from the bottom trawl fisheries off the west coast of North

America from central California to Vancouver.

Specimens recorded in the literature have been

taken in otter trawls that fished at the following

depths (in meters): 165-275, 250-280, 295, 320,

340 (2 stations), 350, 370 (3 stations), 400 (2 stations), 460, and 550 (Pattie, 1968; van Hyning and

Magill, 1964; Smith, 1963). One specimen was

captured by skin divers in shallow water in central

California, which presumably is an abnormally

shallow occurrence for this species, since the shallow water fisheries have failed to yield additional

shallow water records.

R. R. Talmadge (pers. comm.), utilizing information and specimens provided by sole fishermen,

reports that a specimen of M. robusta was captured

at 100 m and that specimens are commonly captured from 200-275 m on down to 375 m. The

currently known distribution may well be incomplete, in that it reflects only the depths at

which fishermen concentrate their fishing efforts

for various species of sole. Fishermen report that

M. robusta may be caught both during day and

night, and that no correlation between depth of

capture and time of day seems to exist. Although

most captures occur in the fall (August to November), specimens may be caught throughout the

year. Occurrence in northern Californian waters

appears to be cyclic and the species may be absent

one year and abundant the next.

Currently, sufficient data are lacking to indicate

the deeper boundary of M. robusta, although the

upper limit appears to be around 100 m. Since all

captures have been made by bottom trawls, it is

probable that this species is associated with the

bottom, at least part of the time. Furthermore,

about half of the specimens examined by Talmadge (pers. comm.) contained in their stomachs

fragments of the sea urchin, Briaster townsendii
NUMBER 209 23

which lives on mud bottom from about 275 m to

900 m.

In summary, little can be said about the vertical

distribution of most species of onychoteuthids. It

is clear that Onychoteuthis banksii occurs in surface waters at night. Onykia caribaea apparently

is restricted to surface waters both during the day

and night. As with many ommastrephids and

loliginids, Moroteuthis robusta seems to associate

with the ocean bottom at least part of the time.

Almost no information is available on the distribution of Chaunoteuthis and Ancistroteuthis.

THYSANOTEUTHIDAE

FIGURE 28: BAR 1

Thysanoteuthis rhombus is a world-wide oceanic

species in lower latitudes. In the Japan Sea it is

taken in set nets during the fall and early winter

in near-shore waters (Nishimura, 1966:329); it is

caught in greater abundance during the night but

daytime captures are relatively common. We have

a juvenile specimen of T. rhombus (25 mm ML)

that was captured in the open ocean (bottom

depth 2750 m) in an open midwater trawl that

fished during the day at 35-55 m.

CYCLOTEUTHIDAE

FIGURE 28: BARS 7-9

This family contains three (possibly four) species in two genera, Cycloteuthis and Discoteuthis;

each species is known from only a few specimens.

Five specimens of Cycloteuthis sirventi (17-134

mm ML) have been caught at 650-0 m during the

day; 140-0 m, 150-0 m, and 100-0 m at night and

200-0 m at twilight in open nets. Five specimens

of Discoteuthis discus (24-53 mm ML), were

captured at 750-0 m and 450-0 m during the day;

400-0 and 100-0 m at night, and 600-0 m (time

unknown). These records have been taken from

scattered localities throughout the north Atlantic

(Young and Roper, 1969a). Two adults have been

taken off Bermuda in open nets at depths of 140-0

m and 950-0 m, at night.

Discoteuthis laciniosa (19-134 mm ML) has

been taken in open nets that fished at 425-0 m,

725-0 m, 950-0 m, and 1060-0 m during the day

and at 160-0 m, 350-0 m, 500-0 m, 750-0 m (2

specimens) and 1200-0 m at night. D. laciniosa

has been taken off Hawaii in open nets at 900-0 m

during the day (55 mm ML) and 215-0 m at night

(40 mm and 50 mm ML). Several specimens were

recently captured in a closing net at 650 m during

the day.

With broad scatter of depth-points for the

cycloteuthid species, little can be said about their

vertical distributions; however, all three species

have been captured in the upper 200 m at night.

CTENOPTERYGIDAE

FIGURE 28: BAR 10

The single genus, Ctenopteryx, is comprised of

several nominal species. Little is known concerning

their vertical distribution. From the Mediterranean two specimens (19 and 35 mm ML) of C.

sicula were captured in closing nets at night between 100 and 150 m. Off Bermuda a single specimen of C. sicula (22 mm ML) was taken at 100 m

at night in a closing net. Off Hawaii four specimens of Ctenopteryx sp. (23-35 mm ML) have

been taken at night from 100-0 m. These species

appear to inhabit the near-surface waters at night.

During the day only three closing net records

are available. Off Bermuda two specimens (12 and

14 mm ML) were caught at 350 m. In the Mediterranean one specimen (12 mm ML) was taken at

150 m. Off Hawaii five specimens (20—29 mm ML)

have been taken in open nets between 600-0 m

and 800-0 m during the day.

In the Atlantic Ocean in open nets three specimens (38-62 mm ML) were taken at 850-0 m and

1000-0 m during the day; at night 10 specimens

(20-58 mm ML) were taken at 100-300 m, five

specimens (20-77 mm ML) at 600-800 m, and five

specimens (42-83 mm ML) at 1300-2000 m.

BRACHIOTEUTHIDAE

FIGURE 29: BAR 2

The family contains a single genus and an indeterminate number of species. Very little is

known of the vertical distribution of any species.

Two specimens of Brachioteuthis sp. were taken

in closing nets off Hawaii, one at 830-975 m during
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the day (38 mm ML) and one at 100-150 m at

night. Open net captures of four specimens were

taken at 975-1000 m during the day (38 mm and

42 mm ML) and at 50 m and 225 m during the

night (33 mm and 47 mm ML). In the Mediterranean three specimens (34-63 mm ML) of

Brachioteuthis sp. were captured in closing nets

at 150 m during the night. The meager data suggest that Brachioteuthis spp. undergo extensive

diel vertical migration.

LEPIDOTEUTHIDAE

FIGURE 28: BARS 11-13

The family tentatively consists of the genera

Lepidoteuthis, Pholidoteuthis, and Tetronychoteuthis. Few reliable records of vertical distribution are available for these genera.

Two juvenile Lepidoteuthis grimaldii (60 and

84 mm ML) have been taken at night in open

midwater trawls that fished from 270-0 m (Clarke,

1964). A specimen (190 mm ML) was taken in an

Engel trawl at 700-0 m at night in the south Atlantic. Off Bermuda one juvenile (35 mm ML)

was captured in a closing net at 50 m at night; a

second specimen (41 mm ML) was taken in an

open net at 585-0 m during the day. All other

large specimens available have been taken from

the stomachs of large fishes and sperm whales

(Clarke, 1966:214), so we have little precise information concerning their vertical distribution.

Three specimens of Tetronychoteuthis dussumieri were captured off Bermuda; one (21 mm

ML) in a closing net at 350 m during the day, two

(44 mm and 100 mm ML) in Engel trawls at night

at 1000-0 and 100-0 m, respectively. In the eastern

Atlantic seven specimens (36-166 mm ML) were

captured in Engel trawls at night at 100-325 m,

four (31-159 mm ML) were taken at 600-800 m

and one (64 mm ML) at 1800-0 m.

Sixteen specimens of Pholidoteuthis adami (all

greater than 250 mm ML) are available from the

Gulf of Mexico, Caribbean Sea, and northeastern

South American waters. All captures were made

during the day in bottom trawls from 360-925 m

(greatest number of captures 625-750 m). No

nighttime trawl captures are known so possibly

this species leaves the near-bottom waters and disperses into middepths at night. Voss (1956:132)

records the species from 85-870 m in the Gulf of

Mexico; all captures were made with bottom

trawls. Remains of P. boschrnai were taken from

stomachs of the shark Centroscyrnnus coelolepis

captured on bottom-set long lines set at 998-1102

m, 1246 m, 1259 m, 1260 m, and 1472 m (Clarke

and Merrett, 1972).

JOUBINITEUTHIDAE

FICURE 27: BAR 15

The Joubiniteuthidae consists only of Joubiniteuthis portieri which is known from a few specimens. Captures were made in open nets that fished

at 800-900 m and 2500 m during the day; 2000 m

at night; and at 330 m, 350 m, 500 m, and 3500 m

at unknown times, although the first three are

probably night captures (Young and Roper,

1969b). A large specimen was taken at night in an

Engel trawl in the South Atlantic at 1900-0 m.

CHIROTEUTHIDAE

FIGURES 14-16, 29: BARS 3-7

The Chiroteuthidae consists of three genera;

Chiroteuthis, Chiropsis, and Valbyteuthis. Information on the vertical distribution of some species

of Chiroteuthis and Valbyteuthis is available, but

the distribution of the monotypic Chiropsis is unknown, since only a single specimen of C. mega

has been taken.

Members of the Chiroteuthidae generally have

very distinctive larval stages that may attain a relatively large size before metamorphosis.

Larvae of Chiroteuthis calyx compose the bulk

of the 136 specimens of this species captured off

California. Metamorphosis to the subadult stage

takes place when the larva attains a mantle length

of 50-60 mm. The vertical distribution of the

larvae (Figure 14) presented here is based almost

exclusively on specimens larger than 20 mm ML.

Generally the youngest specimens live in the upper

100 m, and as they grow they gradually descend to

greater depths. In the upper 100 m larvae in the

20-30 mm ML size category are most abundant.

At 300-400 m the 30-50 mm ML size range predominates, while at 500-700 m the 40-60 mm ML

size category predominates. Specimens taken below

700 m probably are contaminants.
NUMBER 209 25

The day-night distribution seems to indicate

some larval vertical migration. Specimens were

captured in the upper 100 m only at night. In successive (deeper) depth zones, the larger-sized specimens were captured predominantly at night and

smaller specimens during the day. This indicates

that at night, specimens probably shift upward

from their slightly deeper daytime levels.

The subadult population (Figure 15) appears

to occupy a zone between 500 m and 800 m during

the daytime. Within this zone 14 specimens were

captured; no subadults were taken above 500 m.

Two additional specimens, probably contaminants,

were taken between 1000 m and 1100 m. The 500-

800 m zone is approximately the level at which

the largest larvae are found.

The vertical distribution of the nighttime population of subadult C. calyx is uncertain since only

four specimens were captured. One specimen

each was captured in 0-100 m, 200-300 m, 300-

400 m, and 400-500 m. These captures occurred

above the level of the daytime population. The

Day, Night, Twilight Day, Night

1200

50 10 30

Mantle Length

FIGURE 14.—Vertical distribution of the larvae of Chiroteuthis calyx off southern California.

(Day, Night, Twilight: black bar = total captures; Day, Night: black bar = night captures,

white bar = day captures.)
26

scanty data indicate that a migration or spreading

out may occur at night. This interpretation is supported by a few captures from other areas.

Nighttime captures of two subadult Chiroteuthis sp. at 167-0 m were made off Bermuda and

one subadult Chiroteuthis sp. was captured at

150-0 m off Hawaii. One subadult C. veranyi was

taken in a closing trawl in the Mediterranean at

400 m at night. Three larval Chiroteuthis sp.

(40-45 mm ML) were captured in a closing net

during the day off Hawaii between 600 m and

660 m.

m

200-

400- 1

 1

t

 600"

«

o

800-

1000-

1200-

Day

1

•

Night

ii

1

•

•

0 10 0 10

No. of Specimens

FIGURE 15.—Vertical distribution of subadults of Chiroteuthis

calyx off southern California. (Black bar = number of

specimens captured; white bar = correction factor.)

1200

0 10 20

No. of Specimens

FIGURE 16.—Vertical distribution of Valbyteuthis oligobessa

off southern California. (Black bar = number of specimens

captured; white bar = correction factor.)

The records for other species of Chiroteuthis

are too few and too inconclusive to provide information on their vertical distributions.

Eighteen specimens of Valbyteuthis oligobessa

were taken off California in 15 different open net

tows (Figure 16). No specimens were taken in the

upper 700 m. A single specimen was taken at 700-

800 m, one at 800-900 m, three at 900-1000 m,

four at 1000-1100 m, and six at 1100-1200 m. The

shallowest capture is the smallest specimen (23 mm

ML). The lower limit of distribution undoubtedly

lies deeper than 1200 m, beyond the range of the

sampling program. Valbyteuthis is one of the
NUMBER 209 27

deepest living cephalopods taken off California.

There is no difference between daytime and nighttime distributions.

Data available on several other species of Valbyteuthis also indicate a deep-living habit (Roper

and Young, 1967). Specimens larger than 20 mm

ML were taken in open nets that fished at (in

meters): 530, 683, 1000, 1162, 1250 (2 stations),

1500 (3 stations), 1830, and 2635. Two specimens

(35 mm and 50 mm ML) were taken in closing nets

off Hawaii at 875-1220 m during the day and

780-955 m at night. Four specimens (22-45 mm

ML) were captured in open nets that fished between 800 m and 900 m during the day and 700

and 800 m during the night.

In summary, the evidence suggests that some

species of Chiroteuthis exhibit at least limited

diel vertical migration. This is rather unexpected

since they have small, weakly muscled mantles. It

appears that the larvae of C. calyx at least, descend

into deeper water as they increase in size until, by

the time of metamorphosis, they have reached the

depth of the subadult distribution.

Species of Valbyteuthis appear to be among the

deepest living of all pelagic squid and apparently

exhibit no diel vertical migration.

MASTIGOTEUTHIDAE

FIGURES 17, 29: BARS 8-11

The family Mastigoteuthidae consists of two

genera. Echinoteuthis is recorded from only a few

specimens and little is known of its distribution.

Mastigoteuthis is a speciose genus, but meaningful data on vertical distributions are available for

only a few species.

Mastigoteuthis pyrodes from California is represented by 27 specimens (Figure 17). Of the

eight specimens captured during the daytime,

none were taken above 600 m; two were captured

in 600-700 m, five in 800-900 m, and one in 1000-

1100 m. Thirteen specimens are recorded as twilight captures. Many of these captures undoubtedly occurred during daytime in trawls that

extended into the twilight period. If these data are

considered in conjunction with daytime captures,

it appears that the upper limit of the daytime distribution is around 600 m. The lower limit is

uncertain.

Of the five specimens captured at night; two

were taken at 300-400 m, one at 400-500 m, and

two at 600-700 m. Some upward movement during

the night may occur, but the data are too meager

to allow a definitive statement.

Fifteen specimens of M. hjorti have been captured off Bermuda. Closing net captures were made

at night at 450 m (27 mm and 28 mm ML) and

510 m (30 mm ML). Open net nighttime captures

were as follows; four specimens (29-37 mm ML)

at 450-0 m, three (32-155 mm ML) at 750-0 m,

and two (84 mm and 181 mm ML) at 760-0 m.

Open net daytime captures of three specimens

(27-115 mm ML) were made at 800-0 m, 950-0

m, and 1025-0 m.

Off Hawaii Mastigoteuthis sp. has been taken

in closing nets at 700-875 m during the day (two

specimens) and at night two specimens were capDay,

Night, Twilight Day Night

1200

10 0 10 0

No. of Specimens

FIGURE 17.—Vertical distribution of Mastigoteuthis pyrodes

off southern California. (Black bar = number of specimens

captured; white bar = correction factor.)
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tured from 260-325 m and two others from 690-

770 m in closing nets. Five specimens (24-55 mm

ML) have been taken in open nets that fished

from 670-850 m during the day. Two specimens

of M. jamelica were taken during the day, probably at 810 m in a closing net that fished from

700-925 m. Eleven additional specimens (25-41

mm ML) were taken during the day in open nets

between 700 and 800 m and single specimens were

taken at 600-0 and 1100-0 m. A single specimen

(37 mm ML) was captured in an open net at night

at 375-0 m.

It appears from the species examined that members of this genus are deep-living animals with the

upper limits of daytime distribution probably at

600-700 m depending on the locality. At night the

captures suggest that while some individuals remain at the daytime depths, others spread upward

into the 200-700 m zone.

PROMACHOTEUTHTOAE

FIGURE 29: BAR 12

The Promachoteuthidae has two known species

in the single genus Promachoteuthis. The species

P. megaptera is known from a single mutilated

specimen taken by the CHALLENGER in an open

bottom trawl that fished at 2440 m in the north

Pacific. Two specimens of Promachoteuthis from

Antarctic and southern Pacific waters were taken

in open midwater trawls at 1391-0 m (10.5 mm

ML) and 1830-0 m (17 mm ML), respectively.

Two specimens (greater than 25 mm ML) were

taken in open Engel trawls in the south Atlantic

at 1550-0 m and 1900-0 m at night. Based on the

anatomical features of this species, Roper and

Young (1968:213) suggested that it is a deep-sea

species; indeed, it is probably one^of the deepest

living of all squids.

BATOTEUTHIDAE

FIGURE 29: BAR 13

The Batoteuthidae is monotypic, known from

only a few specimens (Young and Roper, 1968).

Data on depths of capture in open nets are insufficient to verify vertical range; one larva (30 mm

ML) was taken at 366 m during the day and juveniles and adults (51, 90, and 95 mm ML) were

taken at 1257 m, 1866 m, and 2525 m during the

night.

GRIMALDITEUTHIDAE

FIGURE 27: BAR 16

The Grimalditeuthidae is represented only by

Grimalditeuthis bomplandii. The few previous

records provide inadequate information on the

vertical distribution of the species because depths

of capture range from the surface to 2600 m. In

the Atlantic four specimens (greater than 46 mm

ML) have been taken at night in open nets at

1275-0 m, 1300-0 m, 1690-0 m, and 2100-0 m.

Off Hawaii this species has been taken at 750-0 m

(58 mm ML) and 1435-0 m (97 mm ML) at

night, and at 1150-0 m (80 mm ML) during the

day. We suspect that G. bomplandii is a deepliving species.

CRANCHIIDAE

FICURES 18-20, 30: BARS 1-9

The Cranchiidae is a highly diversified group

that contains numerous genera and species. The

family is so diverse and so poorly known that we

shall make no attempt to survey the distributional

records of all the genera. We shall discuss only the

species for which we have accumulated direct

information.

Galitenthis phyllura off California is represented by 182 specimens (Figure 18). This species

undergoes a metamorphosis at about 60 mm ML.

The larvae exhibit ontogenetic descent whereby

increasingly larger larvae occupy successively

greater depths. In the upper 100 m larvae from

1-10 mm ML (mean 7.5 mm ML) predominate;

between 100-300 m the 10-30 mm ML size range

(mean 23.4 mm ML) predominates; at 300-500 m

the 20-40 mm ML size (mean 28.1 mm ML) predominates; and at 500-600 m the 30-50 mm ML

size (mean 33.3 mm ML) predominates. The

mean values for most of these size groups would be

larger if contamination of each depth zone from

overlying zones were eliminated.

At approximately 60 mm ML larval development is complete and the animals descend to 900
NUMBER 209 29

m or more as subadults (Figure 19). Twenty-one

percent of the larval specimens less than 50 mm

ML, however, were taken in the 1000-1200 m zone

and may descend prematurely, although contamination may be partly responsible for this peculiar

pattern. No indication of vertical migration exists

for specimens smaller than 55 mm ML.

During the daytime only a single specimen larger than 55 mm ML was taken in water

shallower than 900 m, one specimen was taken in

900-1000 m, three in 1000-1100 m, and seven in

1100-1200 m. When twilight captures are considered as well, it seems clear that the bulk of the

subadult population is located below 900 m.

Only a small number of specimens was taken at

night; three specimens were captured in the upper

200 m, seven in 500-600 m, and three in 1100-

1200 m. The captures above 900 m indicate that

some of the subadult population wander into the

upper layers at night. Five of the seven specimens

taken at 500-600 m at night are less than 100 mm

ML; since this depth corresponds approximately

to the maximum depth of larval occurrence, this

may indicate that a tendency exists for some specimens to remain there after metamorphosis.

Off Hawaii, nine specimens (20-52 mm ML) of

G. pacifica were taken in open nets that fished

below 650 m (seven day captures and two night

captures). Four additional specimens (25-258 mm

ML) were taken during the day in closing nets between 650-800 m, and another specimen (42 mm

ML) was captured in a closing net probably at

550 m (500-560 m range) during the day. In spite

of the single record at 550 m, G. pacifica appears

generally to occupy depths below 600 m during

both day and night. This species undergoes meta200

Number

of Specimens E

10

1200

100 150 200 250

Mantle Length in 10mm Increments

300

FIGURE 18.—Vertical distribution with respect to size-groups of Galiteuthis phyllura off southern

California. (Black bar = number of specimens captured.)
30

mor.phosis at about 40-45 mm ML. Data are insufficient to show the existence of ontogentic

descent.

Galiteuthis glacialis is restricted primarily to

Circumpolar Deep Water in Antarctic waters

(McSweeny, 1971:104). Based on open net data,

McSweeny (1971) concluded that the vertical

range of the species extends from 200 m to about

1400-1500 m; the bulk of the population lives between 600 and 1000 m. An indication of ontogenetic descent exists, since the mean capture

depths for successively larger 10 mm size groups

increase for animals of 20-90 mm ML McSweeny

also suggested that a diel vertical migration occurs;

maximum daytime captures occur at 800-1000 m

while maximum nighttime captures occur at 400-

600 m. McSweeny's data indicate that, rather than

a well-defined migration, the animals disperse at

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

night into a broader range of depths since nighttime captures are relatively high between 200 and

1000 m.

In the California study 102 specimens of Helicocranchia pfefferi (10-70 mm ML) were used to

plot vertical distribution (Figure 20). The specimens captured both during the day and night

with open nets are well intermixed at all depth

levels. No specimens were taken in the upper 100

m, 4 percent were captured between 100-200 m,

21 percent in 200-300 m, 38 percent in 300-400 m,

11 percent in 400-500 m, 12 percent in 500-600 m, 3

percent in 600-700 m, 8 percent in 700-800 m,

and 4 percent below 800 m. The bulk of the captures (59 percent) occurs in the zone between

200 and 400 m. Eighty-seven percent of the specimens captured in this zone are 10-40 mm ML.

Although only a small number of specimens (20)

Day, Nighf, Twilight Day Night

200h

400-

-B 600-

a>

0 10 20 30 0 10 0 10

No. of Specimens

FIGURE 19.—Vertical distribution of subadult Galiteuthis phyllura off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)
NUMBER 209 31

was captured in the 400-600 m zone, 65 percent

of these were over 40 mm ML. The data suggest

that larger animals (i.e., over 40 mm ML) live in

deeper water. A break in distribution lies at 600 m

which may be the bottom of the vertical range of

H. pfefferi. The deeper captures presumably are

contaminants. No diel vertical migration occurs.

Records for seven larval specimens of Taonius

pavo are available from Bermuda and Hawaii. Off

Bermuda three specimens (35-43 mm ML) were

captured in closing nets between 610-690 m during

the day. Off Hawaii, four daytime captures (60-

115 mm ML) were made in closing nets between

600 and 800 m.

Two specimens of Bathothauma lyromma (23

and 33 mm ML) were captured off Bermuda at

520 m at night in closing nets. Off Hawaii one

specimen (80 mm ML) was taken in a closing net

at night at 700-730 m and a mature female (100

mm ML) was captured in a closing net at 1015—

1300 m during the day.

Leachia sp. off Bermuda is represented in our

collections by 309 specimens that range from 4-

50 mm ML. During the day 159 specimens (4-28

mm ML) were taken at 50 m and 12 specimens

(25-50 mm ML) were taken at 60 m. At night

93 specimens (7-29 mm ML) were taken at 50 m;

36 specimens (5-24 mm ML) were captured at

100 m, and nine specimens (26-48 mm ML) were

taken at 135-230 m. It is worth noting that most

of the large specimens taken during the daytime

were the deepest captures. No evidence of diel

vertical movement exists.

Off Baja California, a specimen of Leachia

dislocata (100 mm ML) was dipnetted at the surface at night.

1200

10 20 30 40

No. of Specimens

50 60

FIGURE 20.—Vertical distribution of Helicocranchia pfefferi off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Five specimens of Phasmatopsis oceanica (17-

29 mm ML) were taken in closing nets off Bermuda during the day between 90 m and 200 m and

three specimens (19-27 mm ML) were captured at

night from 50-175 m. Off Hawaii, 97 specimens

(8-55 mm ML; one at 130 mm ML) of P. fisheri

were taken at night from 200-0 m, and five specimens, presumably contaminants, from 800-0 m.

Daytime captures for this species are inconclusive.

Off Hawaii, 16 specimens of Liocranchia valdiviae have been captured in closing nets. Apparently 10—20 mm ML is the size range at which the

larvae begin to assume the adult habitat. Eleven

specimens less than 20 mm ML have been taken at

scattered depths between 175 m and 750 m. Five

specimens from 21-42 mm ML have been taken in

closing nets from 650 m to about 1050 m. Sixtyeight specimens were taken in open nets of which

33 specimens larger than 21 mm ML were captured between 700 m and 1200 m. Twenty of these

came from 700-800 m, five from 800-900 m, six

from 900-1000 m, and two from 1000-1200 m.

Day and night captures are intermixed for all

specimens larger than 21 mm ML.

The species we have examined in the

Cranchiidae exhibit several different patterns of

vertical distribution. The juveniles, at least, of

some species (e.g., Leachia, Phasmatopsis) live in

near-surface waters. Some (e.g., Helicocranchia)

occupy intermediate depths. Most species, however, live in deep water below 600-700 m (e.g.,

Liocranchia, Galiteuthis, Taonius, Bathothauma).

At present no species has been shown to exhibit

strong vertical migration. One distributional pattern appears to be prominant in the family: Many

species exhibit ontogenetic descent, that is, in larval and juvenile stages, larger specimens inhabit

progressively deeper waters. Where known, adults

and subadults occupy the deepest waters.

Order VAMPYROMORPHA

VAMPYROTEUTHIDAE

FIGURES 21, 31: BAR 1

The vertical distribution of Vampyroteuthis

infernalis, the only species in the order Vampyromorpha, has been extensively investigated by Pickford (1946, 1949). She (1946:25) states that the

majority of specimens are found in water that has

a salinity between 34.7 % and 34.9 %c (maximum

range 34.5-35.5 %c), an oxygen content between

1.00 ml/L and 3.99 ml/L, a density (sigma-t) between 27.4 and 27.8 (maximum range 27.0-27.9),

a temperature between 2.0° and 6.0° C (maximum

range 1.73°-15.46° C). The species is most abundant between 1500 m and 2500 m in the Pacific

Ocean and 1000-2000 m in the Atlantic Ocean.

Vampyroteuthis infernalis was the third most

abundant species in the California study. Three

hundred and eighty-five specimens were captured

ranging in size from 8 mm to 110 mm ML (Figure

21). The daytime and nighttime captures intermingle at all depths, indicating that no vertical

migration occurs. No specimens were taken in the

upper 100 m, and only 2 percent were taken between 100 m and 400 m. This species is susceptible

to entanglement in trawl nets, and very likely many

of the records above 400 m represent contamination of this type from preceeding deeper trawls.

Six percent of the specimens were captured between 400 m and 600 m, part of which may represent contamination and part stragglers from the

main population. Forty-five percent of the captures occurred between 600 m and 800 m with those

between 700 m and 800 m accounting for 34 percent

of the captures. The 800-900 m zone accounts for

8 percent of the specimens, the 900-1000 m zone

15 percent, the 1000-1100 m zone for 16 percent,

and the 1100-1200 m zone 6 percent. When catch

per trawling effort is considered, peaks occur in

the distribution at 700-800 m and at 900-1000 m.

Sixty-five percent of the specimens with a mantle

length less than 20 mm were captured deeper than

900 m, while 76 percent of the specimens with a

mantle length of 20 mm or larger occur shallower

than 900 m. This size is the point at which the larval fins begin to be resorbed. The larval forms,

therefore, generally occupy a slightly deeper layer

than the adults.

The upper limit of most of the captures is 600

m, and the lower limit probably is 1100 m or 1200

m. Even though the water off California between

1200 m and 2200 m (around the 2° C isotherm)

would appear to be well suited to V. infernalis,

according to previous studies, the species probably
NUMBER 209 33

1200

20 40 60 80

No. of Specimens

100 120 140

FIGURE 21.—Vertical distribution of Vampyroteuthis infernalis off southern California.

(Black bar = number of specimens captured; white bar = correction factor.)

is absent or at least not abundant in this zone. The

vertical distribution falls within a temperature

range of approximately 3°-6° C, a salinity variation of 34.4

 c/cc to slightly over 34.5 %c a density

(sigma-t) range of 27.1-27.5, and oxygen values of

0.3-0.8 ml/L. These values extend the limits for

the majority of previously recorded specimens, for

all parameters except temperature. These limits,

however, are so broad that other features may be

equally or more important in controlling the distribution of this species.

Off Bermuda, 12 specimens of V. infernalis were

captured between 950 m and 1500 m in open nets.

In the south Atlantic 175 specimens (18-94 mm

ML) of V. infernalis were taken in open Engel

trawls at night; 115 specimens were taken at 1300-

2000 m, with the large majority at 1800-2000 m

(13 tows), and 10 were taken in five tows that

fished between 500 m and 1000 m. One tow captured 50 specimens at 640-0 m.

Order OCTOPODA

Suborder CIRRATA

CIRROTEUTHIDAE and STAUROTEUTHIDAE

FIGURES 22, 31: BARS 10, 11

Species in these families are rare in collections,

and the systematics are very confused; relatively

little is known of their distributions. Recently

Roper and Brundage (1972) reviewed the depth

records in the literature and added information

based on deep-sea photographs. They concluded

that Cirroteuthis spp., and probably some of the

other cirrates as well, are benthopelagic, living

just above, but not on, the ocean bottom (Figure

22). Records for species of Cirroteuthis, Grimpoteuthis, and Stauroteuthis indicate that the group

as a whole extends from about 350 m to 5300 m.

While a few species appear to live primarily shal-
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Cirrate octopods near ocean bottom in Virgin Islands Basin, 3600-3950 m.

Scale approximate. (After Roper and Brundage, 1972.)

lower than 1000 m, the large majority occur at

depths in excess of 1000 m and many of these in

excess of 2000 m (Robson, 1932:82-89). The

photographic records of Roper and Brundage

(1972) are concentrated in 3500-5200 m.

Cirrothauma murrayi, known from only a few

specimens, is pelagic (Roper and Brundage, 1972:

14) and has been taken in open midwater nets at

the following depths: 1500 m, 2900 m (night),

3600 m (day), and 4050 m (night). Specimens of

this species are very fragile and are blind. The

clear indication of deep-living habitat provided by

anatomical and trawl data is partially offset by the

capture of a specimen at the surface through the

ice in the Arctic Ocean; this may, however, be an

example of polar emergence of the widely distributed species.

OPISTHOTEUTHIDAE

The several species in this monotypic family,

represented by Opisthoteuthis, all appear to be

benthic forms; they are weak swimmers and seldom

leave the bottom (Pereyra, 1965:436). Depth

records range from 125-2251 m, but the great

majority of captures come from 400-1000 m (see

discussion in Roper and Brundage, 1972:16).

Pereyra (1965:430) reports that Opisthoteuthis

californiana is distributed from 125-825 m, the

shallower captures occurring in the northern end
NUMBER 209 35

of its range. In addition, males of O. californiana

appear to occupy shallower depths than females

and to move into shallower water during the

summer.

Suborder INCIRRATA

BOLITAENIDAE

FICURES 23, 31: BARS 2, 3

Thore (1949) monographed the family Bolitaenidae and treated the horizontal and vertical

distribution based on open net captures of one of

the most common species, Japetella diaphana. He

1200

20 40 60

No. of Specimens

FIGURE 23.—Vertical distribution of Japetella sp. and /.

heathi, off southern California. (Black bar = number of

specimens captured; white bar = correction factor.)

considered this species to be world-wide but latitudinally bounded by the 10° C isotherm at 200 m.

Thore's data on the vertical distribution of /.

diaphana indicated that young specimens less

than 25 mm ML live between 100 m and 330 m,

while larger specimens migrate to greater depths,

with a lower limit between 3000 m and 4000 m.

Two species of Japetella occur off southern

California. Nearly all specimens are /. heathi;

however, since it is impossible to separate the two

species at small sizes, their distributions are considered together. The 115 specimens (8-100 mm

ML) captured in open nets were intermixed at all

depths during the day and night (Figure 23).

No specimens were captured in the upper 200 m,

10 percent between 200 m and 600 m, 57 percent

from 600-800 m, and 33 percent from 800-1200 m.

All specimens captured shallower than 600 m

were less than 20 mm ML and were nighttime

captures. Below 600 m, 74 percent of the specimens less than 30 mm ML occur in the 600-800 m

zone. This size category comprises over 80 percent

of the total captures in the 600-800 m range. Specimens larger than 30 mm ML are evenly distributed

from 600-1200 m. Therefore, nearly all the

younger specimens seem to occupy a rather restricted vertical zone between 600-800 m, while

the larger animals occur from 600 m to the depth

limits of the sampling program and presumably

occur in greater depths.

The genus Bolitaena consists of a single poorly

known species, B. microcotyla. Off Hawaii a

brooding female (35 mm ML) was taken in a

closing net between 830 m and 975 m during the

day (Young, 1972b).

Thore (1949:62) suggests that Eledonella pygmaea has a distribution similar to that of Japetella diaphana.

VITRELEDONELLTOAE

FIGURE 31: BAR 4

According to Thore's (1949) data, specimens of

Vitreledonella richardi less than 100 mm ML occur

in the upper 300 m, while those larger than 100

mm ML occur below 1000 m with a peak concentration at 1000-1050 m. Off Hawaii one specimen

(23 mm ML) of V. richardi was captured in a

closing net during the day between 615 m and
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

690 m. Off Bermuda four larval specimens less

than 11 mm ML were captured in closing nets

during the night and twilight at depths of

200-300 m.

The pelagic and benthic captures of this gelatinous octopod seem peculiar and require further

investigation before we can gain an understanding

of its vertical distribution.

AMPHTTRETIDAE

FIGURE 31: BAR 5

This monotypic family consists of only Amphitretus pelagicus. Based on five captures, Thore

(1949:55) reported that small specimens (20 mm

ML and smaller) occur in the upper 133 m. Data

of adults are inconclusive. One specimen (35 mm

ML) was taken at night off Hawaii in a closing

net at 240-260 m.

In the South Atlantic five specimens of A.

pelagicus were captured at night in Engel trawls:

one (20 mm ML) at 300-0 m, two (30 mm and 32

mm ML) at 600-0 m and 750-0 m, and two (34

mm and 52 mm ML) at 2000-0 m.

ALLOPOSIDAE

FIGURE 31: BAR 6

The Alloposidae consists of a number of nominal species, all of which have been considered as

synonyms of Alloposus mollis in the review by

Thore (1949). The vertical distribution of A.

mollis presents a perplexing problem, since specimens have been recorded from the surface to as

deep as 3180 m.

Thore (1949:71) reported that specimens from

7-400 mm ML, taken in open midwater nets, occurred from 0-333 m; specimens less than 80 mm

ML were limited to the upper 33 m. In general,

captures of both large and small specimens are

concentrated in the near-surface waters from open

pelagic nets (0-200 m) and at around 450-1000 m

from the bottom trawls (Thore, 1949:71; Robson,

1932:84; Verrill, 1882b:366). The National Marine Fisheries Service, Hawaii, has captured as

many as 26 large specimens of A. mollis in a single

otter trawl that fished at 682 m (L. Burgess, per.

comm.).

Thore (1949:72), concluded that the species is

a cosmopolitan form that spends part of its life

on the bottom and part in open water but always

in association with landmasses.

TREMOCTOPODIDAE

FIGURE 31: BAR 8

The family is represented by Tremoctopus

violaceus and possibly by T. lucifer Akimushkin,

1963. Tremoctopus violaceus is a pelagic cosmopolitan species in all warm and temperate waters

of the world. Females attain a large size (200 mm

ML; Kramer, 1937), but males are much smaller

(about 20-30 mm ML). Most specimens recorded

in the literature are from the surface of nearsurface waters, with a few scattered to about 300 m

(Robson, 1932:85).

Off Bermuda, a juvenile T. violaceus (6 mm

ML) was captured in a closing net at 100 m during

the day, 10 juveniles (5-10 mm ML) were captured at the surface at night, and one juvenile was

taken at 250 m at night.

OCYTHOIDAE

FIGURE 31: BAR 9

Ocythoe tuberculata, the sole species in the

family, has been recorded from scattered oceanic

localities (Robson, 1932:85; Berry, 1955:219), but

nowhere does it seem to be abundant. Females

attain a large size (up to about 300 mm ML), but

males remain small (about 20 mm ML) and apparently live within pelagic salps in near-surface

waters (Hardwick, 1970). The species is pelagic

and seems to inhabit the near surface waters, but

nothing more can be stated about its vertical

distribution.

ARGONAUTIDAE

FIGURE 31: BAR 7

The several species of Argonauta, the sole genus

in the family, are all open ocean dwellers that inhabit warm waters of the world (Robson, 1932:85).

Many records in the literature indicate a surface

and near-surface habitat. We have captured spawn-
NUMBER 209 37

ing adult females (shell diameters greater than

200 mm) at the surface at night in the Florida

Current.

Four juvenile males (5-8 mm ML) of A. argo

were captured in closing nets between 50 m and

200 m off Bermuda. In the Mediterranean Sea in

closing nets, six juvenile males and females (5-13

mm ML) and one adult female (72 mm ML) of

A. argo were captured at the surface at night and

one juvenile female (6 mm ML) was captured at

155 m at night.

The species of Argonauta inhabit the surface

and near-surface waters.

Discussion

A complete analysis of the vertical distribution

of pelagic cephalopods should include information concerning physical and biological parameters

that may regulate species distributions. Of particular importance are the roles of light, temperature, productivity, competition and predation in

regulating species distributions. Also, the relationships of species distributions to vertical faunal

zones should be considered. Unfortunately, knowledge concerning vertical distribution in cephalopods is rudimentary and we are restricted to a

general discussion of approximate habitats and

day-night differences in distribution.

The information presented here reveals a variety

of patterns of vertical distribution in pelagic

cephalopods. The types of patterns presented below are not all mutually exclusive; some species

may exhibit a combination of patterns. All patterns are not necessarily sharply delimited.

NEAR SURFACE DWELLERS.—A number of species

are found at or near the surface of the ocean

throughout the diurnal cycle. Apparently only a

1&2

D.N

7

D. N

8 9 10

S W D . N D N

11 12 13

DNDND N

14

DN

15

D N

200-

400-

600-

800-

1000-

I

e±n

FIGURE 24.—Patterns of vertical migrations. I & 2 NAUTILJDAE: 1, 2, Nautilus macromphalus and

N. pompilius, various localities. 3 SPIRIULIDAE: Spirula spirula, off Canary Islands. 4-7 HETEROTEUTHINAE: 4, Heteroteuthis dispar, off Canary Islands and Mediterranean; 5, Stoloteuthis leucoptera, off eastern United States (time of bottom trawl captures assumed to be day); 6,

Stoloteuthis maoria, off New Zealand (time of bottom trawl captures assumed to be day); 7,

Nectoteuthis pourtalesi, off Florida (time of bottom trawl captures assumed to be day). 8-14

LOUCINIDAE: 8, Loligo pealeii, off eastern United States; 9, Loligo forbesi, Catalane Sea (time

of capture assumed to be day); 10, Loligo vulgaris, off northwest Africa; 11, Loligo plei, tropical

western Atlantic; 12, Alloteuthis media, Mediterranean (time of capture assumed to be day); 13,

Sepioteuthis sepioidea, tropical western Atlantic; 14, Lolliguncula brevis, off tropical western

Atlantic. 15 PICKFORDIATEUTHIDAE: PickfoTdiateuthis pulchella, tropical western Atlantic. (D =

day, N = night, S = summer, W = winter; see pp. 2-3 for explanation of symbols.)
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

single species, Onykia caribaea, presently qualifies

as a member of the neuston, living at the ocean

surface. Some other members of Onykia for which

data are lacking may also occupy this habitat.

Other cephalopods in this category occur primarily in the upper 50 m or so during the day and

night. This group includes the octopods, Argonauta spp., Tremoctopus violaceus, and Ocythoe

tuberculata, as well as some of the squid such as

Thysanoteuthis rhombus.

Other shallow-living species avoid the nearsurface waters during the day and move into these

waters during the night. These shallow diel migrators include Nautilus pompilius, N. macromphalus, Todarodes pacijicus, and probably many

of the loliginid species.

Among the Ommastrephidae, a number of open

ocean species (e.g., O. caroli) seem to occur primarily in the upper few hundred meters during

the day and night. Some individuals, however, apparently roam into great depths. Species that

exhibit roaming activities are powerful swimmers

for which data are difficult to obtain, and the

extent and magnitude of roaming are uncertain.

FIRST ORDER DIEL VERTICAL MIGRATORS.—Many

species live at moderate depths during the daytime and migrate into the upper 200 m at night.

All members of the Enoploteuthidae, where

known, are first order diel vertical migrators. During the day these species occur primarily between

300 m and 600 m, and at night they are found in

the upper 200 m. Within these zones, the distribution of a given species may be considerably more

restricted (e.g., Pyroteuthis margaritifera). At

least one member of the Lycoteuthidae, Selenoteuthis scintillans, exhibits the same pattern of

distribution, and data suggest that members of the

Cycloteuthidae also exhibit this pattern.

Strong diel vertical migration from great depths

is indicated by data for only a single genus. Species

of Brachioteuthis inhabit deep water (about 900-

1000 m) during the day and ascend into the upper

200 m at night.

Ctenopteryx sp. seems to occupy an inter2 3 4 5

D N D N D N D N I D N D,N

7 8 9 10 11 12 13 14

.NI D . N DNIDNIDNIDNID N

i

D N

200 -

— 400 -

E

600-

800-

JE2~

I

1000-

i

FICURE 25.—Patterns of vertical migrations. 1 LYCOTEUTHIDAE: Selenoteuthis scintillans, off Bermuda. 2-15 ENOPLOTEUTHIDAE: 2, Abraliopsis sp., off Hawaii; 3, Abraliopsis felis, off California;

4, Abraliopsis pfefferi, off Bermuda; 5, Abralia veranyi, off Madeira; 6, Abralia redfieldi, off

Bermuda; 7, Abralia trigonura, off Hawaii; 8, Abralia astrostica, off Hawaii; 9, Watasenia scintillans, off Japan; 10, Enoploteuthis anapsis, various localities; II, Enoploteuthis sp., off Bermuda;

12, Pyroteuthis margaritifera, off Bermuda; 13, Pterygioteuthis gemmata, off Califorina; 14,

Pterygioteuthis giardi, off Bermuda; 15, Thelidioteuthis alessandrinii, several localities. (D =

day, N = night; see pp. 2-3 for explanation of symbols.)
NUMBER 209 39

mediate position between the few migrators from

great depths and the many from mid-depths. It

migrates from 600-800 m into the upper 150 m.

SECOND ORDER DIEL VERTICAL MIGRATORS.—At

night, some species desert their daytime depths

and migrate toward the surface but generally occupy deeper waters than first order migrators. For

instance, Spirula spirula moves from 500-700 m

in the day to 200-300 m at night, and Histioteuthis heteropsis moves from 400-800 m (peak

at 500-700 m) during the day to the upper 400 m

at night, with a peak at 300-400 m. Second order

migrators also appear to occupy somewhat greater

depths during the day than most first order

migrators.

DIEL VERTICAL SHIFTERS.—Some species shift upward slightly at night with the nighttime distribution overlapping the daytime distribution.

Several species of Gonatus shift from 400-800 m

during the day to 300-500 m at night.

DIEL VERTICAL SPREADERS.—The nighttime vertical distribution in some species not only encompasses the daytime levels, but spreads well beyond

the daytime limits. For example, Octopoteuthis

deletron appears to occupy depths of 200-400 m

during day and at night spreads out over depths

from near the surface to about 500 m. Although

the data are insufficient, we suspect that species of

Mastigoteuthis are also spreaders.

NONMIGRATORS.—Many species exhibit no diel

changes in vertical distribution. Nonmigrators

include species from near-surface waters, as men200-

— 400 -

E

Q 600-

800-

D N

1000-H I

1500-

2 3 4 5 6 7 8 9

D&N D N D NID N D N D N D N D N

10

D N

3

n

D N

12

D N

13 14 15 16

D N DNW S D N

FIGURE 26.—Patterns of vertical migrations. 1-16 OMMASTREPHIDAE: 1, Ommastrephes caroli, central Atlantic; 2, Ommastrephes bartramii, several localities and Dosidicus gigas, off western

South America; 3, Symplectoteuthis oualaniensis and S. luminosa, several localities; 4, Hyaloteuthis pelagica, off Bermuda; 5, Illex illecebrosus, off Newfoundland; 6, Illex illecebrosus, off

the Chesapeake Bight, eastern United States; 7, Illex illecebrosus?, off Cape Hatteras; 8, Illex sp.,

off Miami, Florida; 9, Illex oxygonius, western Atlantic; 10, Illex coindetii, various localities in

western Atlantic and Mediterranean (time of capture assumed to be day); 11, Todaropsis

eblanae, various localities (time of capture assumed to be day); 12, Todarodes sagittatus, various

localities; 13, Todarodes pacificus, off Japan; 14, Nototodarus sloani gouldi, off Australia; 15,

Nototodarus hawaiiensis, off Hawaii; 16, Ornithoteuthis antillarum, various localities. (D =

day, N = night, W = winter, S = summer, D&N = combined day and night captures (no

marked diel differences in distribution); see pp. 2-3 for explanation of symbols.)
40

tioned above, as well as species from mid-depths

and from great depths. For example, Helicocranchia pfefferi occupies the same mid-depths

(200-600 m) during both the day and night. Nonmigrators are more prevalent in great depths than

in mid-depths. For instance, species of Bathyteuthis, Valbyteuthis, Japetella, and Vampyroteuthis infernalis are deep-living nonmigrators.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

VERTICAL WANDERERS.—Individuals of some deepliving species occasionally are found during the

night considerably above the normal range. This

pattern is displayed by Galiteuthis phyllura which

normally occurs below 900 m, but during the night

individuals may wander upward as far as the

upper 100 m.

SPECIES ASSOCIATED WITH THE OCEAN BOTTOM.—

FIGURE 27.—Patterns of vertical migrations. 1-4 HISTIOTEUTHIDAE: 1, Histioteuthis heteropsis,

off California; 2, Histioteuthis dofleini, off Bermuda; 3, Histioteuthis meleagroteuthis, off Bermuda; 4, Histioteuthis corona, off Bermuda. 5-7 BATHYTEUTHIDAE: 5, Bathyteuthis abyssicola,

Antarctic waters; 6, Bathyteuthis bacidifera, tropical eastern Pacific; 7, Bathyteuthis berryi, off

California. 8-14 GONATIDAE: 8, Conatus onyx, off California; 9, Gonatus californiensis, off California; 10, Gonatus pyros, off California; 11, Gonatus berryi, off California; 12, Gonatopsis

borealis, off California; 13, Berryteuthis magister, near Auke Bay, Alaska; 14, Berryteuthis

anonychus, off Oregon. 15 JOUBINITEUTHIDAE: Joubiniteuthis portieri, various localities (shallow

records are assumed to be nighttime captures). 16 GRIMALDITEUTHIDAE: Grimalditeuthis bomplandii, various localities. (D = day, N = night, D&N = combined day and night captures (no

marked diel differences in distribution); see pp. 2-3 for explanation of symbols.)
NUMBER 209 41

A surprisingly large number of pelagic cephalopods live at least part of the time near the ocean

bottom; these include shallow as well as moderately

deep-living forms. Some species of the Heteroteuthinae (e.g., Stoloteuthis maoria) appear to

belong to this category, as well as many species of

loliginids (e.g., Loligo pealeii and L. vulgaris)

and some ommastrephids, onychoteuthids, gonatids and mastigoteuthids.

SPECIES EXHIBITING ONTOGENETIC DESCENT.—Several species occupy progressively greater depths

during successive stages in their life history. This

is most apparent in species that have larvae which

attain an unusually large size. For example, larval

Chiroteuthis calyx with increasing size gradually

descend from the upper 100 m to 500-700 m where

they undergo metamorphosis at a mantle length

of 50-60 mm. Many cranchiids exhibit a similar

pattern (e.g., Galiteuthis phyllura, Helicocranchia

pfefferi, and Phasmatopsis fisheri).

In contrast to this gradual ontogenetic descent,

nearly all other species of pelagic cephalopods

demonstrate a different type of ontogenetic descent; larvae occur in near surface waters and at a

particular size abruptly descend to the adult

habitat. An exception to this pattern occurs in

Vampyroteuthis infernalis where the young occupy

greater depths than the adults.

Some of the above patterns seem to correlate

approximately with familial boundaries, although

many exceptions exist. Most species that live in

the upper 800 m exhibit some type of diel change

in vertical distribution. Most species that live

primarily below 800 m do not exhibit diel variations in vertical distribution. Apparently very few

species live below 1000 m.

200-

~ 400H

600-

800-

1000-

2 3 4

D NID&N D«N

5 6 1 7 8 9 I 10 I 11 12 13

D N ID N D NIDNI D N D N D N D N D N

I

FIGURE 28.—Patterns of vertical migrations. 1. THYSANOTEUTHIDAE: Thysanoteuthis rhombus,

various localities. 2-4 ONYCHOTEUTHIDAE: 2, Onychoteuthis banksii, off Bermuda and Mediterranean; 3, Onykia caribaea, various localities; 4, Moroteuthis robusta, off western North America.

5-6 OCTOPOTEUTHIDAE: 5, Octopoteuthis deletron, off California; 6, Taningia danae, various

localities. 7-9 CYCLOTEUTHIDAE: 7, Cycloteuthis sirventi, various localities; 8, Discoteuthis discus,

various localities; 9, Discoteuthis laciniosa, various localities. 10 CTENOPTERYCIDAE: Ctenopteryx

spp., various localities. 11-13 LEPIDOTEUTHIDAE: 11, Lepidoteuthis grimaldii, various localities;

12, Pholidoteuthis adami, Gulf of Mexico, Caribbean Sea, and northeastern South American

waters; 13, Tetronychoteuthis dussumieri, various localities. (D = day, N = night, D8cN =

combined day and night captures (no marked diel differences in distribution); see pp. 2-3 for

explanation of symbols.)
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1

D.N

3 4 5 6 7

jLorvae Adult

D,N S—» D,N D,N DNlD& N D N

8 9 10 11

D.N D.N D.NID. N

12

D&N

13

D.N

200-

400-

— 600-

800H

1000-

1200-

1400-

2000-

3000-

FIGURE 29.—Patterns of vertical migrations. 1 NEOTEUTHIDAE: Neoteuthis sp., eastern tropical

Atlantic and Gulf of Mexico. 2 BRACHIOTEUTHIDAE: Brachioteuthis spp., off Hawaii and Mediterranean. 3-7 CHIROTEUTHIDAE: 3, Chiroteuthis caylx, off California; 4, Chiroteuthis veranyi, Mediterranean; 5, Chiroteuthis spp., off Hawaii and Bermuda; 6, Valbyteuthis oligobessa, off

California; 7, Valbyteuthis sp., off Hawaii. 8-11 MASTICOTEUTHIDAE: 8, Mastigoteuthis pyrodes,

off Califronia; 9, Mastigoteuthis hjorti, off Bermuda; 10, Mastigoteuthis sp., off Hawaii; 11,

Mastigoteuthis famelica, off Hawaii. 12 PROMACHOTEUTHIDAE: Promachoteuthis sp., various localities. 13 BATOTEUTHIDAE: Batoteuthis scolops, Antarctic waters. (D = day, N = night, D&N =

combined day and night captures (no marked diel differences in distribution), S-> = size of

individuals increase toward the right; see pp. 2-3 for explanation of symbols.)
NUMBER 209 43

Larvae Adult

D,N|S—-» , D N

200-

400-

— 600-

800

1000

1200-

1400

3 4

D ,N|D ,N

l

5 6

D&N

D,N |S •

I

I

FIGURE 30.—Patterns of vertical migrations. 1-9 CRANCHIIDAE: 1, Galiteuthis glacialis, Antarctic

waters; 2, Galiteuthis phyllura, off California; 3, Galiteuthis pacifica, off Hawaii; 4, Taonis pavo,

off Bermuda and Hawaii; 5, Bathothauma lyromma, off Bermuda and Hawaii; 6, Helicocranchia

pfefferi, off California; 7, Phasmatopsis fisheri, off Hawaii; 8, Leachia sp., off Bermuda; 9,

Liocranchia valdiviae, off Hawaii. (D = day, N = night, S-> = size of individuals increases to

the right, D&N = combined day and night captures (no marked diel differences in distribution);

see pp. 2-3 for explanation of symbols.)
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

200-

400-

~ 600-

_c _

a

o 800-

1000-

1200-

2000-

3000-

4000-

5000

1

D&N

J A

II

2

D&N

J A

1

3

D N

4

D&N

J A

1

5

D N

6

D&N

I

i

7

D&N

8

D&N

9

D&N

10 11

D N D&N

I

! J ^

| y/y

FIGURE 31.—Patterns of vertical migrations. 1 VAMPYROTEUTHIDAE: Vampyroteuthis infernalis,

off California. 2-3 BOLITAENIDAE: 2, Japetella sp. and /. heathi, off California; 3, Bolitaena

microcotyla, off Hawaii. 4 VITRELEDONELLIDAE: Vitreledonella richardi, various localities. 5

AMPHITRETIDAE: Amphitretus pelagicus, off Hawaii (consists of juveniles and larvae only). 6

ALLOPOSIDAE: Alloposus mollis, various localities. 7 ARGONAUTIDAE: Argonauta spp., various

localities. 8 TREMOCTOPODIDAE: Tremoctopus violaceus, various localities. 9 OCYTHOIDAE: Ocythoe

tuberculata, various localities. 10 CIRROTEUTHIDAJE: Cirrothauma murrayi, various localities. 11

CIRRATA: most cirrate octopods other than Cirrothauma and opisthoteuthids, various localities.

(D = day, N = night, D&N = combined day and night captures (no marked diel differences

in distribution), J = juveniles, A = adults; see pp. 2-3 for explanation of symbols.)
NUMBER 209 45

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C. F. E., R. Gibbs, Jr., and W. Aron

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A New Species of I Ilex from the Western Atlantic

and Distributional Aspects of Other Illex Species

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A Review of the Valbyteuthidae and an Evaluation

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The Family Promachoteuthidae (Cephalopoda:

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More Giant Squids from California. California

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1970.

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1957.

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1969. Winter Population of Loligo pealei in the MidAtlantic Bight. Biological Bulletin, 137 (l):202-216.

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Thore, S.

1949. Investigations of the "Dana" Octopoda, Part I:

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van Hyning, J. M., and A. R. Magill

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48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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Index

Abralia, 10

astrosticta, 11, 38 (fig.)

redfieldi, 10, 38 (fig.)

trigonura, 11, 38 (fig.)

veranyi, 10, 38 (fig.)

Abraliopsis, 9

felis, 9, 10 (fig.), 38 (fig.)

pfefferi, 10, 38 (fig.)

sp., 10, 38 (fig.)

abyssicola, Bathyteuthis, 16 (fig.), 40 (fig.)

adami, Pholidoteuthis, 24, 41 (fig.)

Alcpisaurus ferox, 16

alcssandrinii, Thelidioteuthis, 12, 13, 38

(fig-)

Alloposidac, 36, 44 (fig.)

Alloposus mollis, 36, 44 (fig.)

Alloteuthis, 8

media, 8, 37 (fig.)

Allurotcuthis antarcticus, 15

Amphitrctidae, 36, 44 (fig.)

Amphitretus pelagicus, 36, 44 (fig.)

anapsis, Enoploteuthis, 11, 38 (fig.)

Ancistrochirus lesueuri, 12

Ancistroteuthis lichtensteini, 22, 23

anonychus, Berryteuthis, 20, 40 (fig.)

antarcticus, Alluroteuthis, 15

antillarum, Ornithoteuthis, 13, 39 (fig.)

Architeuthidae, 16

Architeuthis, 16

argo, Argonauta, 37

Argonauta, 36, 37, 38

argo, 37

spp., 44 (fig.)

Argonautidae, 36, 37, 44 (fig.)

astrosticta, Abralia, 11, 38 (fig.)

bacidifera, Bathyteuthis, 17, 40 (fig.)

banksii, Onychoteuthis, 22, 23, 41 (fig.)

bartramii, Ommastrephes, 13, 39 (fig.)

Bathothauma, 32

lyromma, 31, 43 (fig.)

Bathyteuthidae, 16, 17, 40 (fig.)

Bathyteuthis, 40

abyssicola, 16 (fig.), 40 (fig.)

bacidifera, 17, 40 (fig.)

berryi, 17, 40 (fig.)

Batoteuthidae, 28, 42 (fig.)

Batoteuthis, 28

scolops, 42 (fig.)

berryi, Bathyteuthis, 17, 40 (fig.)

Gonatopsis, 20

Gonatus, 19, 20 (fig.), 40 (fig.)

Berryteuthis, 17

anonychus, 20, 40 (fig.)

magister, 19, 20, 40 (fig.)

Bolitaena microcotyla, 35, 44 (fig.)

Bolitaenidae, 35, 44 (fig.)

bomplandii, Grimalditeuthis, 28, 40 (fig.)

borealis, Gonatopsis, 19, 21 (fig.), 40 (fig.)

boschmai, Pholidoteuthis, 24

Brachioteuthidae, 23, 24, 42 (fig.)

Brachioteuthis, 23

sp., 23

spp., 42 (fig.)

brevis, Lolliguncula, 8, 37 (fig.)

Briaster tovvnsendii, 22

californiana, Opisthoteuthis, 34, 35

californiensis, Gonatus, 17, 18 (fig.), 20,

40 (fig.)

calyx, Chiroteuthis, 24, 25 (fig.), 26 (fig.),

27, 41, 42 (fig.)

caribaea, Onykia, 22, 23, 38, 41 (fig.)

caroli, Ommastrephes, 13, 14, 38, 39 (fig.)

Centroscymnus coelolepis, 14, 16, 21, 24

Chaunoteuthis mollis, 22, 23

Chiropsis mega, 24

Chiroteuthidae, 24-27, 42 (fig.)

Chiroteuthis, 24, 27

calyx, 24, 25 (fig.), 26 (fig.), 27, 41, 42

(fig-)

sp., 26

spp., 42 (fig.)

verany, 26, 42 (fig.)

Cirroteuthidae, 33, 34, 44 (fig.)

Cirroteuthis spp., 33

Cirrothauma murrayi, 34, 44 (fig.)

coelolepis, Centroscymnus, 14, 16, 21, 24

coindetii, Illex, 14, 39 (fig.)

corona, Histioteuthis, 15, 40 (fig.)

Cranchiidae, 28-32, 43 (fig.)

Ctenopterygidae, 23, 41 (fig.)

Ctenopteryx, 23

sicula, 23

spp., 41 (fig.)

Cycloteuthidae, 23, 38. 41 (fig.)

Cycloteuthis, 23

sirventi, 23, 41 (fig.)

danae, Taningia, 21, 41 (fig.)

deletron, Octopoteuthis, 21 (fig.), 39, 41

(fig)

diadema, Lycoteuthis, 9

diaphana, Japetella, 35

Discoteuthis, 23

discus, 23, 41 (fig.)

laciniosa, 23, 41 (fig.)

discus, Discoteuthis, 23, 41 (fig.)

dislocata, Leachia, 31

dispar, Heteroteuthis, 7, 37 (fig.)

dofleini, Histioteuthis, 15, 40 (fig.)

Dosidicus gigas, 13, 39 (fig.)

dussumieri, Tetronychoteuthis, 24, 41

(fig-)

eblanae, Todaropsis, 14, 39 (fig.)

Echinoteuthis, 27

Eledonella pygmaea, 35

Enoploteuthidae, 9, 38 (fig.)

Enoploteuthis, 11

anapsis, 11, 38 (fig.)

sp., 11, 38 (fig.)

famelica, Mastigoteuthis, 28, 42 (fig.)

felis, Abraliopsis, 9, 10 (fig.), 30 (fig.)

ferox, Alepisaurus, 16

fisheri, Phasmatopsis, 32, 41, 43 (fig.)

forbesi, Loligo, 8, 37 (fig.)

Galiteuthis, 32

glacialis, 30, 43 (fig.)

pacifica, 29, 43 (fig.)

phyllura, 28, 29 (fig.), 30 (fig.), 40, 41,

43 (fig.)

gemmata, Pterygioteuthis, 12 (fig.). 38

(fig)-

giardi, Pterygioteuthis, 12 (fig.), 38 (fig.)

gigas, Dosidicus, 13, 39 (fig.)

glacialis, Galiteuthis, 30, 43 (fig.)

Gonatidae, 17, 40 (fig.), 41

Gonatopsis, 17

berryi, 20

borealis, 19, 21 (fig.), 40 (fig.)

Gonatus, 17, 19, 39

berryi, 19, 20 (fig.), 40 (fig.)

californiensis, 17, 18 (fig.) 20, 40 (fig.)

onyx, 17 (fig.), 20, 40 (fig.)

pyros, 18, 19 (fig.), 20, 40 (fig.)

grimaldii, Lepidoteuthis, 24, 41 (fig.)

Grimalditeuthidae, 28, 40 (fig.)

Grimalditeuthis bomplandii, 28, 40 (fig.)

Grimpoteuthis, 33

hawaiiensis, Heteroteuthis, 8

Nototodarus, 14, 39 (fig.)

49
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

heathi, Japetella, 35 (fig.), 44 (fig.)

Helicocranchia, 32

pfefferi, 30, 31 (fig.), 40, 41, 43 (fig.)

heteropsis, Histioteuthis, 14, 15 (fig.), 39,

40 (fig.)

Heteroteuthis, 7, 8

dispar, 7, 37 (fig.)

hawaiiensis, 8

Histioteuthidae, 14, 15, 40 (fig.)

Histioteuthis, 14

corona, 15, 40 (fig.)

dofleini, 15, 40 (fig.)

heteropsis, 14, 15 (fig.), 39, 40 (fig.)

meleagroteuthis, 15, 40 (fig.)

hjorti, Mastigoteuthis, 27, 42 (fig.)

Hyaloteuthis pelagica, 13, 39 (fig.)

Idiosepiidae, 7, 8

illecebrosus, Illex, 13, 14, 39 (fig.)

Illex, 14

coindetii, 14, 39 (fig.)

illecebrosus, 13, 14, 39 (fig.)

oxygonius, 14, 39 (fig.)

sp., 39 (fig.)

infernalis, Vampyroteuthis, 32, 33 (fig.).

40, 41, 44 (fig.)

Japetella, 40

diaphana, 35

heathi, 35 (fig.), 44 (fig.)

sp., 35 (fig.), 44 (fig.)

Joubiniteuthidae, 24, 40 (fig.)

Joubiniteuthis portieri, 24, 40 (fig.)

laciniosa, Discoteuthis, 23, 41 (fig.)

Lampadioteuthis megaleia, 9

Leachia, 32

dislocata, 31

sp., 31, 43 (fig.)

Lepidoteuthidae, 24, 41 (fig.)

Lepidoteuthis grimaldii, 24, 41 (fig.)

lesueuri, Ancistroteuthis, 12

leucoptera, Stoloteuthis, 7, 37 (fig.)

lichtensteini, Ancistroteuthis, 22, 23

Liocranchia valdiviae, 32, 43 (fig.)

Loliginidae, 8, 37 (fig.)

Loligo, 8

forbesi, 8, 37 (fig.)

pealeii, 8, 37 (fig.), 41

plei, 8, 37 (fig.)

vulgaris, 8, 37 (fig.), 41

Lolliguncula, 8

brevis, 8, 37 (fig.)

lucifer, Tremoctopus, 36

luminosa, Symplectoteuthis, 13, 39 (fig.)

Lycoteuthidae, 9, 38 (fig.)

Lycoteuthis diadema, 9

lyromma, Bathothauma, 31, 43 (fig.)

macromphalus, Nautilus, 6, 37 (fig-), 38

magister, Berryteuthis, 19, 20, 40 (fig.)

margaritifera, Pyroteuthis, 11 (fig.), 12,

13, 38 (fig.)

maoria, Stoloteuthis, 7, 37 (fig.), 41

Mastigoteuthidae, 27, 28, 41, 42 (fig.)

Mastigoteuthis, 27, 39

famelica, 28, 42 (fig.)

hjorti, 27, 42 (fig.)

pyrodes, 27 (fig.), 42 (fig.)

sp., 27, 42 (fig.)

media, Alloteuthis, 8, 37 (fig.)

mega, Chiropsis, 24

megaleia, Lampadioteuthis, 9

megaptera, Promachoteuthis, 28

meleagroteuthis, Histioteuthis, 15, 40

(fig-)

microcotyla, Bolitaena, 35, 44 (fig.)

mollis, Alloposus, 36, 44 (fig.)

Chaunoteuthis, 22, 23

Moroteuthis robusta, 22, 23, 41 (fig.)

murrayi, Cirrothauma, 34, 44 (fig.)

Nautilidae, 6, 37 (fig.)

Nautilus, 6

macromphalus, 6, 37 (fig.), 38

pompilius, 6, 37 (fig.), 38

Nectoteuthis pourtalesi, 8, 37 (fig.)

Neoteuthidae, 15, 16, 42 (fig.)

Neoteuthis, 15

thielei, 15

sp., 15, 42 (fig.)

Nototodarus, 14

hawaiiensis, 14, 39 (fig.)

sloani gouldi, 14, 39 (fig.)

oceanica, Phasmatopsis, 32

Octopodteuthidae, 21, 22, 41 (fig.)

Octopoteuthis, 21

deletron, 21 (fig.) 39, 41 (fig.)

Ocythoe tuberculata, 36, 38, 44 (fig.)

Ocythoidae, 36, 44 (fig.)

oligobessa, Valbyteuthis, 26 (fig.) 42 (fig.)

Ommastrephes, 13

bartramii, 13, 39 (fig.)

caroli, 13, 14, 38, 39 (fig.)

pteropus, 13

Ommastrephidae, 13, 14, 38, 39 (fig.), 41

Onychoteuthidae, 22, 23, 41 (fig.)

Onychoteuthis, 22

banksii, 22, 23, 41 (fig.)

Onykia, 38

caribaea, 22, 23, 38, 41 (fig.)

onyx, Gonatus, 17 (fig.), 20, 40, (fig.)

Opisthoteuthidae, 34, 35

Opisthoteuthis californiana, 34, 35

Oregoniateuthis spingeri, 9

Ornithoteuthis, 13

antillarum, 13, 39 (fig.)

volitilis, 13

oualaniensis, Symplectoteuthis, 13, 39

(fig;)

oxygonius, Illex., 14, 39 (fig.)

pacifica, Galiteuthis, 29, 43 (fig.)

pacificus, Todarodes, 14, 38, 39 (fig.)

pavo, Taonius, 31,43 (fig.)

pealeii, Loligo, 8, 37 (fig.) 41

pelagica, Hyaloteuthis, 13, 39 (fig.)

pelagicus, Amphitretus, 36, 44 (fig.)

pfefferi, Abraliopsis, 10, 38 (fig.)

Helicocranchia, 30, 31 (fig.), 40, 41, 43

(fig)-

Phasmatopsis, 32

fished, 32, 41, 43 (fig.)

oceanica, 32

Pholidoteuthis, 24

adami, 24, 41 (fig.)

boschmai, 24

phyllura, Galiteuthis, 28, 29 (fig.), 30

(fig.), 40, 41, 43 (fig.)

Pickfordiateuthidac, 9, 37 (fig.)

I'ickfordiateuthis, 9

pulchella, 9, 37 (fig.)

plei, Loligo, 8, 37 (fig.)

pompilius, Nautilus, 6, 37 (fig), 38

portieri, Joubiniteuthis, 24, 40 (fig.)

pourtalesi, Nectoteuthis, 8, 37 (fig.)

Promachoteuthidae, 28, 42 (fig.)

Promachoteuthis, 28

megaptera, 28

sp., 42 (fig.)

Psychroteuthidae, 15

Psychroteuthis, 15

pteropus, Ommastrephes, 13

Pterygioteuthis, 11, 12

gemmata, 12 (fig.), 38 (fig.)

giardi, 12 (fig.), 38 (fig.)

pulchella, Pickfordiateuthis, 9, 37 (fig.)

pygmaea, Eledonella, 35

pyrodes, Mastigoteuthis, 27 (fig.), 42 (fig.)

pyros, Gonatus, 18, 19 (fig.), 20, 40 (fig.)

Pyroteuthis, 11

margaritifera, 11 (fig.), 12, 13, 38 (fig.)

redfieldi, Abralia, 10, 38 (fig.)

rhombus, Thysanoteuthis, 23, 38, 41 (fig.)

richardi, Vitreledonella, 35, 44 (fig.)

robusta, Moroteuthis, 22, 23, 41 (fig.)

sagittatus, Todarodes, 14, 39 (fig.)

Sargassum, 22

scintillans, Selenoteuthis, 9, 38 (fig.)

Watasenia, 1, 11, 38 (fig.)

scolops, Batoteuthis, 42 (fig)

Selenoteuthis, 9
NUMBER 209 51

scintillans, 9, 38 (fig.)

Sepiadariidae, 7, 8

Sepiidae, 7

sepioidea, Sepioteuthis, 8, 37 (fig.)

Sepiolidae, 7, 8

Sepioteuthis sepioidea, 8, 37 (fig.)

sicula, Ctenopteryx, 23

sirventi, Cycloteuthis, 23, 41 (fig.)

sloani gouldi, Nototodarus, 14, 39 (fig.)

Spirula spirula, 6, 7, 37 (fig.), 39

spirula, Spirula, 6, 7, 37 (fig.), 39

Spirulidae, 6, 7, 37 (fig.)

springeri, Oregoniateuthis, 9

Stauroteuthis, 33

Stoloteuthis, 7

leucoptcra, 7, 37 (fig.)

maoria, 7, 37 (fig.), 41

Symplectoteuthis, 13

luminosa, 13, 39 (fig.)

oualaniensis, 13, 39 (fig.)

Taningia, 21

danae, 21, 41 (fig.)

Taonius, 32

pavo, 31, 43 (fig.)

Tetronychoteuthis dussumieri, 24, 41

(fig-)

Thelidioteuthis alessandrinii, 12, 13, 38

(fig)

thielei, Neoteuthis, 15

Thysanoteuthidae, 23, 41 (fig.)

Thysanoteuthis rhombus, 23, 38, 41 (fig.)

Todarodes, 14

pacificus, 14, 38, 39 (fig.)

sagittatus, 14, 39 (fig.)

Todaropsis eblanae, 14, 39 (fig.)

townsendii, Briaster, 22

Tremoctopodiae, 36, 44 (fig.)

Tremoctopus, 36

lucifer, 36

violaceus, 36, 38, 44 (fig.)

trigonura, Abralia, 11, 38 (fig.)

tuberculata, Ocythoe, 36, 38, 44 (fig.)

Valbyteuthis, 24, 27, 40

oligobessa, 26 (fig.), 42 (fig.)

sp., 42 (fig.)

valdiviae, Liocranchia, 32, 43 (fig.)

Vampyroteuthidae, 32, 33, 44 (fig.)

Vampyroteuthis infernalis, 32, 33 (fig-).

40, 41, 44 (fig.)

veranyi, Abralia, 10, 38 (fig.)

Chiroteuthis, 26, 42 (fig.)

violaceus, Tremoctopus, 36, 38, 44 (fig.)

Vitreledonella richardi, 35, 44 (fig.)

Vitreledonellidae, 35, 36, 44 (fig.)

volidlis, Ornithoteuthis, 13

vulgaris, Loligo, 8, 37 (fig.), 41

Watasenia scintillans, 1, 11, 38 (fig.)

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