U.S. Department 

of Commerce 

April 2001 

NOAA Technical Report NMFS 152


A Scientific Paper of the FISHERY BULLETIN


Distribution, Relative Abundance 

and Developmental Morphology 

of Paralarval Cephalopods in the 

Western North Atlantic Ocean 

Michael Vecchione 

Clyde F. E. Roper 

Michael J. Sweeney 

C. C. Lu 
NOAA Technical Report NMFS 152 

A Technical Report of the Fishery Bulletin 

Distribution, Relative Abundance and 

Developmental Morphology of 

Paralarval Cephalopods in the 

Western North Atlantic Ocean 

Michael Vecchione 

Clyde F. E. Roper 

Michael J. Sweeney 

C. C. Lu 

April 2001 

U.S. Department of Commerce 

Seattle, Washington 
CON

TENTS


Introduction 

Materials and methods 

Species accounts 

Family Sepiolidae 

Semirossia tenera 

Stoloteuthis leucoptera 

Family Loliginidae 

Loligo pealeii 

Lolliguncula brevis 

Family Lycoteuthidae 

Selenoteuthis scintillans 

Family Enoploteuthidae 

Abralia cf. veranyi 

Abraliopsis cf. pfefferi 

Family Ancistrocheiridae 

Ancistrocheirus lesueurii 

Family Pyroteuthidae 

Pyroteuthis margaritifera 

Pterygioteuthis sp. 

Family Octopoteuthidae 

Octopoteuthis sp. 

Family Onychoteuthidae 

Onychoteuthis cf. banksii 

Onykia carriboea 

Family Gonatidae 

Gonatus fabricii 

Family Histioteuthidae 

Histioteuthis spp. 

Family Bathyteuthidae 

Bathyteuthis abyssicola 

Family Chtenopterygidae 

Chtenopteryx sicula 

Family Brachioteuthidae 

Brachioteuthis sp. 

Family Ommastrephidae 

Illex sp. 

Rhynchoteuthion Type A′ 

Rhynchoteuthion Type B′ (revised) 

Family Chiroteuthidae 

Chiroteuthis sp. 

Family Mastigoteuthidae 

Mastigoteuthis hjorti 

Family Cranchiidae 

Leachia lemur Teuthowenia megalops 

Heliococranchia cf. papillata 

Other Cranchiidae 

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iii

CONTENTS (continued) 

Family Octopodidae 39


Octopodid spp. 39


Family Tremoctopodidae 40


Tremoctopus violaceus 40


Family Argonautidae 40


Argonauta argo/Argonauta hians 40


Distributional patterns 43


Acknowledgments 50


Literature cited 51


iv 
1 

Abstract–Paralarval and juvenile cephalopods collected in plankton samples on 

21 western North Atlantic cruises were 

identified and enumerated. The 3731 

specimens were assigned to 44 generic 

and specific taxa. This paper describes 

their spatial and temporal distributions 

and their developmental morphology. 

The smallest paralarvae recognized for a 

number of species are identified and illustrated. The two most abundant and most 

frequently collected taxa were identifiable 

to species based on known systematic characters of young, as well as on distribution 

of the adults. These were the neritic 

squids Loligo pealeii and Illex illecebrosus collected north of Cape Hatteras, both valuable fishery resources. Other abundant 

taxa included two morphotypes of ommastrephids, at least five species of enoploteuthids, two species of onychoteuthids, 

and unidentified octopods. Most taxa 

were distributed widely both in time and 

in space, although some seasonal and 

mesoscale-spatial patterns were indicated. 

The taxa that appeared to have distinct 

seasonal distribution included most of 

the neritic species and, surprisingly, the 

young of the bathypelagic cranchiids. In 

eight seasonal cruises over the continental shelf of the middle U.S. Atlantic states, 

neritic taxa demonstrated approximately 

the same seasonal patterns during two 

consecutive years. Interannual differences 

in the oceanic taxa collected on the shelf 

were extreme. The highest abundance 

and diversity of planktonic cephalopods 

in the oceanic samples were consistently 

found in the vicinity of the Gulf Stream. 

Only eight of the oceanic taxa appeared to 

have limited areal distributions, compared 

with twelve taxa that were found throughout the western North Atlantic regions 

sampled in this study. Many taxa, however, were not collected frequently enough 

to describe seasonal or spatial patterns. 

Comparisons with published accounts of 

other cephalopod surveys indicate both 

strengths and weaknesses in various sampling techniques for capturing the young 

of oceanic cephalopods. Enoploteuthids 

were abundant both in our study and in 

other studies using midwater trawls in several areas of the North Atlantic. Thus, this 

family probably is adequately sampled over 

its developmental range. In contrast, octopoteuthids and chtenopterygiids are rare 

in collections made by small to mediumsized midwater trawls but are comparatively common in plankton samples. For 

families that are relatively common in 

plankton samples, paralarval abundance, 

derived similarly to the familiar ichthyoplankton surveys of fisheries science, may 

be the most reliable method of gathering 

data on distribution and abundance. 

Distribution, Relative Abundance and 

Developmental Morphology of Paralarval 

Cephalopods in the Western North Atlantic Ocean 

Michael Vecchione 

Systematics Laboratory


National Marine Fisheries Service, NOAA


National Museum of Natural History


Washington, DC 20560


e-mail: vecchione.michael@nmnh.si.edu 

Clyde F. E. Roper 

Michael J. Sweeney 

Department of Invertebrate Zoology – Mollusks


National Museum of Natural History


Smithsonian Institution


Washington, DC 20560


C.C. Lu 

Department of Zoology


National Chung Hsing University


Taichung, 40227 Taiwan


Introduction jects in the western North Atlantic 

Ocean. This study is not an analysis 

The value of life-history studies ex- of detailed distribution of individual 

ceeds the knowledge gained on the species, but rather it presents largeindividual species studied. While pri- scale distribution patterns made posmary goals may include answering sible by the broad diversity of projlimited questions about the repro- ects and cruises. Additional, specific 

duction or population dynamics of details are provided in the references 

a particular species, ultimately such cited. Our objectives in this study 

studies should be directed toward were as follows: 

distinguishing general patterns with 

which we can understand the organ- 1) To assign each morphological type 

isms in their ecosystems. Vecchione to the lowest possible taxonomic 

(1986) and Sweeney et al. (1992) category. 

reviewed studies on the early life his- 2) To determine the spatial and temtories of cephalopods. Although these poral patterns of distribution and 

authors could draw some generaliza- relative abundance of these taxa. 

tions from the diverse literature on Seasonal distributions need to be 

the subject, some major deficiencies determined so that inferences can 

were observed. Important among the be made about spawning seasondeficiencies pointed out were taxo- ality. Determination of large-scale 

nomic problems and contradictory and meso-scale geographical disstatements on seasonal and geograph- tributions enables us to discuss 

ical distributional patterns. possible causes for the observed 

We have examined cephalopods patterns. 

collected during several major zoo- 3) To address several species-specific 

plankton and ichthyoplankton pro- hypotheses that have been pro-
2 NOAA Technical Report NMFS 152 

posed in the literature, as presented in the individual 

species accounts. 

4) To describe developmental morphology and provide 

illustrations of taxonomic voucher specimens for 

poorly known or previously undescribed paralarvae. 

Materials and methods 

This study includes cephalopods and associated data 

collected during several projects with diverse goals: 

1) The National Marine Fisheries Service’s (NMFS) 

Marine Resources Monitoring, Assessment, and Prediction (MARMAP) Program, a fisheries study of the 

Northwest Atlantic Fisheries Management Zone of 

the United States; 

2) A multidisciplinary baseline study of the continental 

shelf in the middle U.S. Atlantic states, funded by the 

Bureau of Land Management (BLM) and conducted 

by the Virginia Institute of Marine Science (VIMS); 

3) A study of the continental slope and rise in the 

vicinity of Norfolk Canyon by VIMS, funded by the 

National Science Foundation (NSF); 

4) A VIMS study of the zooplankton of the lower Chesapeake Bay, funded by the Commonwealth of Virginia 

to determine long-term changes in the zooplankton 

of the Bay; and 

5) Several oceanic cruises to the tropical western North 

Atlantic, the area between the middle U.S. Atlantic 

states and Bermuda, and to the area from Delaware 

Bay to the Scotian Shelf. 

Areas covered 

The areas covered by all sampling programs are shown 

in Fig. 1. 

The Marine Resources Monitoring, Assessment, and 

Prediction (MARMAP) Program The zooplankton 

and ichthyoplankton of the Gulf of Maine, Georges 

Bank, southern New England, and the middle U.S. 

Atlantic states were surveyed by the National Marine 

Fisheries Service Northeast Fisheries Science Center 

(NEFSC) periodically from 1961–1993 (Sherman et al., 

1998). The primary sampling gear was 61-cm bongo 

frames rigged with 505-μm and 333-μm mesh nets. This 

gear was used for oblique subsurface sampling of the 

entire water column. A neuston sampler with 947-μm 

mesh netting was used at selected stations to sample the 

surface layer, as well. 

We have obtained 2087 cephalopod specimens from 

four MARMAP cruises over the continental shelf of 

the area described above: R/V Albatross cruise 7106, 

R/VAlbatross cruise 7308, R/V Delaware cruise 7219, R/V 

Wieczno cruise 7410. One of these cruises, R/V Delaware 

cruise 7219 (DEL7219), also sampled the continental 

shelf of the southern U.S. Atlantic states. We understand that some of the MARMAP material was sent to 

the Polish Oceanographic Sorting Center and a portion 

might not have been returned for all cruises. Therefore, in the absence of precise records, it is not clear for 

which of the cruises the data might be incomplete. This 

wealth of specimens has been relied upon primarily for 

the systematic descriptions of life history stages and the 

illustrations that we present here. The data on presence 

of taxa, in spite of any gaps, also have been used in the 

distributional and seasonal analyses. 

The Bureau of Land Management (BLM) study This 

two-year study began in the autumn of 1975 and was 

based on four quarterly cruises per year (Vecchione 

and Grant, 1983). During the first year of the study, six 

24-hour stations were occupied on a cross-shelf transect 

off Atlantic City, New Jersey, that extended from shallow 

inshore waters to the shelf break at about 200 m depth. 

Surface collections were made at each of these stations 

every three hours with a neuston frame rigged with 

a one meter 505-μm mesh net that sampled approximately the top 12 cm of the sea surface. Double-oblique 

subsurface tows (from 1 m below the surface to the 

bottom and back to 1 m to exclude the surface layer) 

were made at night with 60-cm opening-closing bongo 

systems rigged with 202-μm and 505-μm mesh nets. 

During the second year, two stations to the north of 

Atlantic City and a second transect of four stations off 

Wachapreague, Virginia, were added. Although sampling intensity varied among stations during the second 

year (Vecchione and Grant, 1983), nine of the twelve 

stations were occupied for 24 hours on each cruise, 

comparable to those of the first year. In addition, four 

replicate subsurface tows were collected at three stations on each cruise during the second year. Volumes of 

water filtered for each sample were calculated based on 

flowmeter readings. Surface temperature and salinity 

were measured concurrently with all surface samples. 

We have examined all 831 cephalopods collected in 

the 726 samples from the eight cruises of this program. 

Because we are confident of this data set’s completeness, these data are central to our analyses of variability 

in distribution and abundance. Presence of taxa in the 

other data sets discussed below, for which we have less 

complete data, are used to supplement the BLM data to 

extend seasonal information and geographic ranges. 

Continental slope and rise VIMS conducted cruises 

to the vicinity of Norfolk Canyon in September 1975 

and January 1976 to study demersal fish communities 

in that area (Middleton and Musick, 1986). Zooplankton was collected on these cruises at stations randomly 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 3 

Figure 1 

Location of 21 cruises (9 plotted on map, 12 in box) along western North Atlantic Ocean from which paralarval cephalopods were studied. 
4 NOAA Technical Report NMFS 152 

selected from strata based on bottom depth. Subsurface 

collections were made at each station by an oblique tow 

of a 60-cm diameter bongo frame rigged with 202-μm 

and 333-μm mesh nets, and a surface sample was collected with a 1-m neuston frame rigged with a 202-μm 

mesh net. Volumes of water filtered for subsurface samples were monitored as described above, as were surface temperature and salinity. 

Chesapeake Bay Cruises were conducted by VIMS 

during March and August 1978 as part of the Lower 

Bay Zooplankton Monitoring Program (LBZMP). These 

sampling months were selected based on previous determinations of peaks in zooplankton abundances in the 

lower Chesapeake Bay, with a winter-spring zooplankton community peak in March and a summer-fall community peak in August (Grant and Olney, 1979). Each 

cruise sampled ten daytime and ten nighttime stations 

selected at random from a grid of several hundred possible stations. Each station consisted of a surface tow 

with a 505-μm mesh neuston net and an oblique subsurface tow with a 61 cm diameter bongo sampler rigged 

with 333-μm and 202-μm mesh nets. Volumes of water 

filtered were estimated with flowmeter values for all 

samples, and surface temperature and salinity were 

recorded. Sixty-three cephalopods were collected in 80 

samples. 

Oceanic cruises R/V Albatross IV Cruise 7206 (ALB 

7206) was a survey of the tropical western North Atlantic between the Lesser Antilles and Cape Canaveral, 

Florida. Sampling was conducted by NMFS during July 

and August 1972 with a 60-cm diameter 505-μm mesh 

bongo net and a neuston net, presumably with 947-μm 

mesh. We have 458 planktonic cephalopod specimens 

from this cruise. 

R/V Albatross IV Cruise 7302 (ALB7302) employed 

bongo and neuston nets (the same as those described 

for ALB7206) to survey the southern Sargasso Sea between the Bahamas and Cape Hatteras. Because this 

cruise was conducted by NMFS during February 1973 

and overlapped the area sampled by the summer cruise 

(ALB7206) described above, seasonal comparisons are 

possible for the southern Sargasso Sea. We have 164 

cephalopods that were collected on ALB7302. 

R/V Trident Cruise 130 (TRI130) was a transect of six 

60-cm diameter bongo net (mesh size=505 μm) stations 

between the middle U.S. Atlantic states and the southern Sargasso Sea. This cruise, which was conducted in 

January 1973, collected 29 specimens. 

R/V Columbus Iselin Cruise 7802 (CI7802) was a 

deep-sea trawling cruise from Bermuda to Rhode Island to the Caryn Seamount to Norfolk, Virginia. It was 

conducted by VIMS during February and March 1978. 

Zooplankton samples collected on this cruise included 

several subsurface tows with a modified Tucker trawl 

rigged with a one-meter, 505-μm mesh net, and 19 surface collections made with a one-meter, 505-μm mesh 

neuston net. Volumes of water filtered were monitored 

by flowmeter, and surface temperature and salinity were 

measured concurrently with surface collections. Twenty-nine cephalopod specimens were collected in the 

plankton samples on this cruise. 

R/V Kaiyo-Maru Cruise 8201 (KAI8201) was conducted during January and February 1982 in waters of 

greater than 1000 m depth offshore between Delaware 

Bay and the Scotian Shelf. The objective of this cruise, 

which was a joint operation by the governments of Japan, Canada, and the United States, was to study the 

ecology of the squid Illex illecebrosus (Vecchione and 

Roper, 1986). Sixty-three oblique tows were made between 0 and 200 m depth with a 61-cm bongo frame 

rigged with 505-μm mesh nets on both sides. Flowmeters were used to estimate volume of water filtered for 

each sample. Additional oblique samples between 0 

and 1000 m depth were obtained at night with a 3-m 

rectangular midwater trawl which had dual 5.6 mm 

mesh cod ends. Physical data collected included vertical profiles of temperature, salinity, and dissolved 

oxygen. Because the samples were divided arbitrarily 

among the sponsoring nations, the 70 specimens we 

have are an unrecorded proportion of the planktonic 

cephalopods collected. 

All samples were fixed in buffered formalin solution, 

following standard plankton preservation procedures. 

The cephalopod paralarvae were preserved in 50% isopropyl alcohol following initial sorting from the original plankton samples. 

Species accounts 

The 3731 specimens collected on the 21 cruises were 

identified to 44 taxa (Table 1). In the presentation that 

follows, we describe the distribution of those taxa for 

which we feel the data are adequate (e.g., a sufficient 

number of specimens, with a range of sizes, that are 

in adequate condition to represent a developmental 

sequence and to enable inferences about distribution). 

We also present descriptions of taxa for which the early 

stages are rare in plankton collections or have not been 

adequately described previously. We describe as many 

sizes as possible within a given species or taxon, with 

particular emphasis on the smallest specimens available. 

Descriptions are presented in ascending-size order. In 

general, descriptions are most detailed for smaller sizes, 

with descriptions for larger specimens emphasizing 

newly developed characteristics, advances in development of earlier-formed characters, or changes in character states. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 5 

Table 1 

Taxa represented in this study; 44 taxa, 3731 specimens. Classification follows that of Sweeney and Roper (1998). 

Family Histioteuthidae Verrill, 1881 

Histioteuthis spp. 

Family Bathyteuthidae Pfeffer, 1900 

Bathyteuthis abyssicola Hoyle, 1885 

Family Chtenopterygidae Grimpe, 1922 

Chtenopteryx sicula (Verany, 1851) 

Family Brachioteuthidae Pfeffer, 1908 

Brachioteuthis sp. 

Family Ommastrephidae Steenstrup, 1857 

Illex sp. 

Illex illecebrosus (Lesueur, 1821) 

rhynchoteuthion Type A′ 

rhynchoteuthion Type B′ 

unidentified ommastrephids 

Family Chiroteuthidae Orbigny, 1841 

Chiroteuthis sp. 

Family Mastigoteuthidae Verrill, 1881 

Mastigoteuthis hjorti Chun, 1913 

Family Cranchiidae Prosch, 1847 

Leachia lemur (Berry, 1920) 

Teuthowenia megalops (Prosch, 1847) 

Helicocranchia cf. papillata (Voss, 1960) 

?Bathothauma lyromma Chun, 1906 

?Taonius/Galiteuthis sp. 

unidentified cranchiids 

Order Octopodida Leach, 1818 

Suborder Incirrina Grimpe, 1916 

Family Octopodidae Orbigny, 1840 

Octopus spp.-type 

“Macrotritopus”-type 

unidentified octopodids 

Family Tremoctopodidae Tryon, 1879 

Tremoctopus violaceus Chiaie, 1830 

Family Argonautidae Tryon, 1879 

Argonauta argo Linnaeus, 1758/ 

Argonauta hians Lightfoot, 1786 

Class Cephalopoda Cuvier, 1797 

Subclass Coleoidea Bather, 1888 

Order Spirulida Stolley, 1919 

Family Spirulidae Owen, 1836 

?Spirula spirula (Linnaeus, 1758) 

Order Sepiolida Fioroni, 1981 

Family Sepiolidae Leach, 1817 

Semirossia tenera (Verrill, 1880) 

Stoloteuthis leucoptera (Verrill, 1878) 

Order Teuthida Naef, 1916 

Suborder Myopsina Orbigny, 1841 

Family Loliginidae Lesueur, 1821 

Loligo sp. 

Loligo pealeii Lesueur, 1821 

Lolliguncula brevis (Blainville, 1823) 

unidentified loliginids 

Suborder Oegopsina Orbigny, 1845 

unidentified oegopsids 

Family Lycoteuthidae Pfeffer, 1908 

Selenoteuthis scintillans Voss, 1959 

unidentified lycoteuthids 

Family Enoploteuthidae Pfeffer, 1900 

Abralia cf. veranyi (Ruppel, 1844) 

Abraliopsis cf. pfefferi Joubin, 1896 

Ancistrocheirus lesueurii (Orbigny, 1842) 

Pyroteuthis margaritifera (Ruppell, 1844) 

Pterygioteuthis sp. Fischer, 1896 

unidentified enoploteuthids 

Family Octopoteuthidae Berry, 1912 

Octopoteuthis sp. 

Family Onychoteuthidae Gray, 1849 

Onychoteuthis cf. banksii (Leach, 1817) 

Onykia carriboea Lesueur, 1821 

Family Cycloteuthidae Naef, 1923 

?Discoteuthis sp. 

Family Gonatidae Hoyle, 1886 

Gonatus fabricii (Lichtenstein, 1818) 

The data are presented by families. The format includes the species name, author, and date (or higher 

taxon name if species identification was not possible) 

and the figure number(s) associated with that taxon. 

Each species account begins with a list of the voucher 

specimens upon which the observations on developmental morphology are based in increasing dorsal mantle length (ML, in mm), the ship acronym, cruise and 

station number, and the USNM catalog number of 

the voucher specimen(s). This is followed by a summary of the distributional data, the morphological descriptions of the voucher specimens, the specific figure 

reference(s), and remarks, principally on growth and 

development of taxonomic characters. 

The abbreviations of anatomical features, primarily 

measurements and counts, are as defined in Roper and 

Voss (1983) and Sweeney et al. (1992). Terminology 

and definitions are as presented in Roper et al. (1984). 

Family Sepiolidae 

Semirossia tenera (Verrill, 1880) 

Voucher specimens 2.3 mm mantle length (ML), 

WIE7410 sta. 12, USNM 730512; 6.9 mm ML, ALB7208 

sta. 13, USNM 816714; 7.5 mm ML, ALB7208 sta. 14, 

USNM 730513. 

Species distribution Semirossia tenera was collected 

throughout the year. Although early juveniles of this 

species are represented in our samples only from north 

of Cape Hatteras, breeding probably is not limited to 

northern latitudes, since the species is common in the 

Straits of Florida (Cairns, 1976). The 21 specimens of 
6 NOAA Technical Report NMFS 152 

Figure 2 

Semirossia tenera (Verrill, 1880). A, dorsal view, 2.25 mm ML, USNM 730512, R/V Wieczno sta. 7410-12; B, oral view, same 

specimen; C, transversely elongate bilobed light organ on anterior surface of ink sac, male 7.5 mm ML, USNM 730513, R/V 

Albatross IV sta. 7208-14. 

S. tenera we examined were captured in neritic waters 

from coastal stations to the outer shelf stations. The 

range of bottom depths for stations at which specimens 

were collected was 23–138 m. At shelf-break stations, S. 

tenera appeared to be replaced by Stoloteuthis leucoptera, 

a species found only where depths exceeded 138 m. 

Behavioral studies reviewed by Boletzky (1977a) indicate that benthic sepioids, such as Semirossia, produce 

benthic hatchlings. Some of the young specimens of S. 

tenera considered here, however, were collected a considerable distance above the bottom. Time-Depth-Recorder 

(TDR) records from the subsurface tows of the BLM 

program indicate that two specimens were captured at 

least 40–50 m above the bottom (bottom depth minus 

maximum depth of tow), and eight specimens were collected at least 10–15 m above the bottom. No sepiolids 

were collected in surface samples. 

Morphology of voucher specimens 2.3 mm ML 

(Fig. 2A,B): Mantle muscular, very plump, rounded posteriorly, MW nearly equals ML. Chromatophores thickly 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 7 

and evenly distributed over entire mantle, head, and 

arms. Fins lateral, widely separated, round; FL approximately 50% ML. Funnel long, muscular, tubular, extends to anterior of mid-eye. Head large, HW = MW. 

Eyes large, round (not ovoid), prominent, covered with 

very thin transparent membrane. Eyelid not developed. 

Arms long, robust, II=III>I=IV. Tentacles missing, due 

to damage. Light organ on anterior ventral surface of 

ink sac developing as transverse, elongate, translucent, 

bilobed organ backed with reflective tissue; appears as 

two nearly separate organs. 

6.9 mm ML: Mantle very plump, rounded; MW = ML. 

Fins lateral, separated, rounded, FL ca. 82% ML. Funnel large, tubular, extends nearly to bases of Arms IV. 

Eyes covered with thin membrane; eyelid present. Arms 

elongate, left Arm I hectocotylized with minute suckers. 

Light organ bilobed, well developed on anterior of ink 

sac, pores obvious. 

7.5 mm ML (Fig. 2C): Mantle short, muscular, broad; 

MW = 86% ML. Fins separate, very large, FL = 82% ML. 

Funnel very muscular, tubular, long, extends to bases of 

Arms IV. Left Arm I hectocotylized with minute suckers, 

curled. Transversely elongate bilobed light organ on anterior surface of ink sac; reflective tissue golden. Papillose tubes with pores overlie light organ lobes. Ink sac 

covered with bronze-colored reflective tissue. 

Remarks Sepiolid taxonomy currently is in need of 

extensive revision. As is the general case with sepiolids 

(Boletzky, 1974; 1977a), all specimens were very similar to adults of the Rossia/Semirossia complex in external morphology. Nearly all of the rossiin sepiolids examined in this project had either ink-sac light organs 

or patches of reflective tissue where such light organs 

would develop. Because of this primary character, the 

species is identified as S. tenera, which is the only light 

organ-bearing Rossinae in these waters. 

Stoloteuthis leucoptera (Verrill, 1878) 

Voucher specimen 9.8 mm ML, Z77.044, USNM 

816715. 

Species distribution Stoloteuthis leucoptera was found 

only where bottom depths exceeded 138 m in our samples. It occurs in the western North Atlantic Ocean 

from the straits of Florida to the Gulf of St. Lawrence, 

in the eastern North Atlantic in the Bay of Biscay, and 

in the Mediterranean Sea. 

Morphology of voucher specimen 9.8 mm ML 

(Figs. 3–4). Mantle fused dorsally to head; muscular, 

plump, broad, globular; MW = 110% ML; ventral mantle 

margin (ventral shield) protrudes anteriorly to level of 

eye opening; ventral mantle length = 123% dorsal ML. 

Dark, broad stripe of densely packed chromatophores 

along dorsal midline, more scattered laterally. Ventral 

shield covers entire ventral surface of mantle, very darkly pigmented with closely set chromatophores; lateral 

and posterior surfaces covered with golden reflective 

tissue. Fins widely separated, broad, elongate-rounded, 

long, FL = 96% ML. Fin base attached to anterior part of 

mantle; posterior edges do not extend posterior to mantle. Funnel tip narrow, barely extends anterior to ventral mantle opening; base broad, muscular. Head short, 

about 50–60% ML, broad, HW = MW; eyes large; eyelids 

distinct. Arms short, robust, bluntly tipped; deep web 

connects Arms I–IV, but not between Arms IV; order 

III=II>IV>I; suckers biserial, globular, reduced in diameter toward tip, closely packed. Tentacles robust, especially on proximal half, clubs short, tip bluntly pointed, 

suckers very small, closely packed, in 12 to 16 rows. Narrow median adductor muscle connects ventral mantle 

wall to viscera. Large bilobed photophore covers ventral 

surface of ink sac. 

Remarks The three specimens of Stoloteuthis leucoptera were identified based on their distinctive color pattern on the ventral shield, which could be discerned 

easily even after years of preservation in alcohol. 

Family Loliginidae 

Loligo pealeii Lesueur, 1821 

Species distribution The sympatry of several species of Loligo south of Cape Hatteras (Cohen, 1976) 

prevents absolute identification of specimens from the 

southern part of our study area. However, the most likely species other than L. pealeii that may be present to 

the south is L. plei. Other Loligo species, L. ocula and 

L. roperi, are restricted to Caribbean islands. Lolliguncula brevis and Sepioteuthis sepioidea are identifiable at 

small stages (Hanlon et al., 1992). Loligo pealeii is by far 

the most abundant loliginid species north of Cape Hatteras, thus the many specimens of Loligo collected in the 

middle U.S. Atlantic states and the Gulf of Maine are assumed to be L. pealeii (Vecchione, 1981). 

The distribution, size, and relative abundance of 

young stages of this commercially important species 

have been described previously based on the BLM samples (Vecchione, 1981). Briefly, they were captured 

throughout the spring, summer, and fall in waters with 

a salinity range of 31.5–34.0 × 10–3 and were confined 

to coastal waters except where surface current conditions resulted in strong offshore transport. The higher 

relative abundances in surface samples and larger specimens in subsurface samples are indicative of an ontogenetic migration, whereby hatchlings leave bottom 

hatching sites (Griswald and Prezioso, 1981), rise to the 

surface layer for a short period, and then move deeper 

into the water column as they grow larger. 

Loligo pealeii was the cephalopod species most abundant and most frequently collected in the samples 

we studied. The maximum abundance recorded from 

quantitative samples was 71 per 100 m3 of water filtered. 
8 NOAA Technical Report NMFS 152 

Figure 3


Stoloteuthis leucoptera (Verrill, 1878). A, dorsal view, 9.8 mm ML, USNM 816715, sta. Z77.044; B, ventral view, same specimen.


Peak abundance, however, may be even higher; a single 

surface tow off Georgia in August 1972 collected 967 loliginid specimens that likely were this species. 

The MARMAP samples from north of Cape Hatteras 

allow us to expand these observations on the early life 

history of L. pealeii. Because of the limited area sampled 

by the BLM program, large-scale geographic patterns 

could not be addressed by Vecchione (1981). However, 

in the MARMAP samples six specimens were captured 

on Georges Bank in October 1973, and 51 specimens 

came from the area of Georges Bank in September– 

October 1974. Four specimens were collected in the 

Gulf of Maine in October 1974. These captures indicate 

spawning to the northern limits of the species’ range, as 

far north as Nova Scotia (Roper et al., 1984). No published data were available on details of spawning areas 

or on water masses present or moving through the sampling area at that time. Transport of paralarvae from 

southern spawning areas is an alternative hypothesis to 

explain the presence of these specimens in the Gulf of 

Maine and Georges Bank, but such transport seems unlikely, as it is contrary to the normal drift of water masses on the continental shelf from northeast to southwest 

(Bishop and Overland, 1977). 

Vecchione (1981) proposed that diel variability in 

abundance observed in surface samples probably was 

a result of changes in the ability of young squids to 

avoid the sampler in different light conditions. Two 

sets of MARMAP diel subsurface tows do not contradict 

this hypothesis (Fig. 5). If daytime decreases in surface 

abundance were a result of diel vertical migration instead of net avoidance, subsurface abundance should 

increase during the day and decrease at night. Similarly, if the clear temporal patterns seen in the surface 

collections resulted only from diel periodicity in hatching, abundance in subsurface samples should increase 

at night and decrease during the day, concurrent with 

the pattern of the surface samples. Although variability 

among samples is evident in the time-series of MARMAP subsurface samples (Fig. 5), no consistent diel pattern is apparent. Sampling variability, however, is quite 

high for young squids. In three sets of closely spaced 

samples that approximate replication, the median coefficient of variability (standard deviation divided by 

mean) in abundance for L. pealeii was 1.83, indicating a 

patchy distribution on our sampling scale. 

Although L. pealeii was not collected in winter north of 

Cape Hatteras, young Loligo were collected throughout 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 9 

Figure 4 

Stoloteuthis leucoptera (Verrill, 1878). Right half of oral 

view, 9.8 mm ML, USNM 816715, sta. Z77.044. Figure 5 

Time series of oblique subsurface tows in two October 

cruises for captures of Loligo pealei. Note absence of consistent diel patterns. 

the year in the southern part of our study area. Again, 

in spite of minor differences in chromatophore patterns 

among hatchling loliginids (McConnathy et al., 1980), 

taxonomic problems (Vecchione, 1988) preclude determination of whether the southern specimens represent year-round spawning by L. pealeii in warm waters or 

spawning by more than one species. 

Remarks The early stages of this species have been 

described elsewhere (Verrill, 1882; McConnathy et al., 

1980; Vecchione, 1981). They have been compared with 

Lolliguncula brevis (McConnathy et al., 1980; Vecchione, 

1982; Hanlon et al., 1992) and with Sepioteuthis sepioidea 

(Hanlon et al., 1992), which are the other genera of Loliginidae in our study area. Early stages of loliginids can 

be identified based on morphometry and the presence 

of a protective membrane over the eyes (Naef, 1923). 

Young Loligo are distinguishable in the western North 

Atlantic Ocean based on chromatophore patterns, especially the presence of a trapezoidal arrangement of 

four chromatophores posterior to each eye on the ventral surface of the head. Morphometric changes during growth, though nonlinear (Vecchione, 1981), are 

gradual and not adequate for specific or generic identification. Vecchione (1988) presented evidence that 

chromatophore patterns cannot be used confidently 

to distinguish paralarval L. pealeii from its occasionally 

sympatric congenor L. plei. 

Lolliguncula brevis (Blainville, 1823) 

Species distribution Planktonic specimens of L. brevis were collected only during August–October in the 

middle U.S. Atlantic states, the period of maximum water temperatures. Because 40°N is the northern limit 

for this euryhaline species (Haefner, 1964), low temperatures probably limit this seasonal distribution. Near 

the center of its range, in the northern Gulf of Mexico, 

planktonic specimens are collected from early spring 

through late fall (Vecchione, 1982; 1991). 

In spite of extensive surface sampling throughout the 

lower Chesapeake Bay, all specimens collected inside 

the bay were in subsurface samples from the eastern 

side of the bay. Such a distribution is characteristic of 

coastal plankton intruded into the bay (Grant, 1977). 

Conversely, specimens from over the continental shelf, 

collected only at the station closest to shore off Virginia, 
10 NOAA Technical Report NMFS 152 

were taken only in surface samples. Based on what is 

known of the two-layer circulation in the bay and nearshore areas (Pritchard, 1952; Beardsley and Hart, 1978; 

Boicourt, 1982), these distributions probably indicate 

coastal spawning near the mouth of the bay. 

Remarks Chromatophore patterns of hatchling L. 

brevis have been described by McConnathy et al. (1980), 

and development of the species has been described in 

detail by Vecchione (1982), based in part on the specimens considered here. The most useful character for 

identification is the triangular arrangement of chromatophores posterior to the eyes on the ventral surface 

of the head. 

The enoploteuthid group of families 

The families Enoploteuthidae, Ancistrocheiridae, and 

Pyroteuthidae were long considered to be subfamilies of 

Enoploteuthidae. We follow Young and Harman (1998) 

in treating them as separate families which, together 

with Lycoteuthidae, comprise the enoploteuthid group 

of families. This group of families often comprises the 

most abundant cephalopod paralarvae in oceanic zooplankton samples (e.g., Piatkowski et al., 1993). 

Family Lycoteuthidae 

Selenoteuthis scintillans Voss, 1958 

Voucher specimens 3.8 mm ML, ALB7206 sta. 56, 

(no longer extant); 6.3 mm ML, ALB7206 sta. 71, USNM 730510; 9.4 mm ML, ALB7206 sta. 71, USNM 

730510. 

Species distribution Because only four specimens 

were collected in this program, all on a single cruise, 

little can be inferred about the early life history of this 

species. 

Morphology of voucher specimens 3.8 mm ML 

(Fig. 6E): Tentacles and Arms I–III long and attenuate, 

Arms IV shorter. Eye light organ 3 (counting from anterior to posterior) large and brownish-golden with clear 

center. Eye light organs 2 and 4 about one half or less 

diameter of 3, very pale color (Fig. 6E). No trace of precursor to eye light organs 1 and 5. No trace of light organs on viscera. 

6.3 mm ML: Tentacles and all arms long. Eye light 

organ 3 large and purplish-brown with central bright 

spot, 2 and 4 about one half diameter of 3 and slightly 

pigmented, 5 visible only as a greyish bump of tissue, 

1 barely visible. Medial visceral light organs present as 

minute anlagen only. Posterior visceral and gill light organs completely undeveloped. 

9.4 mm ML (Fig. 6A–D): Arms and tentacles long. 

Five well-developed eye light organs: number 3 largest, 

very dark with blue-green bright spot in center, 1, 2, 

4, and 5 round to oval and bronze-colored. Pair of relatively large, oval, pearly white light organs medial to 

bases of gills just anterior to branchial hearts. Small 

brownish light organs lateral to base of each gill, connected to medial light organs with dark brown strands. 

One conical median light organ in posterior mantle 

cavity very near tip and ventral to gonad anlagen. Pair 

of triangular light organs anteriorly, one on each side of 

anus. 

Remarks The early development of a dark photophore on the ventral surface of the eye at position 

3, followed by 2 and 4, is useful to separate lycoteuthids 

from enoploteuthids. The planktonic forms that we 

have here seem conspecific in view of Toll’s (1983) synonymy of Oregoniateuthis with Lycoteuthis. Villanueva and 

Sanchez (1993) synonymized Lycoteuthis diadema (Chun, 

1900) with Lycoteuthis lorigera (Steenstrup, 1875). Furthermore, females of Selenoteuthis scintillans seem indistinguishable from Lycoteuthis springeri (Voss, 1956). 

Family Enoploteuthidae 

Abralia cf. veranyi (Ruppell, 1844) 

Voucher specimens 3.1 mm ML, ALB7302 sta. 79, 

USNM 730511; 4.5 mm ML, ALB7302 sta. 79, USNM 

730511. 

Species distribution In our samples, Abralia cf. veranyi was collected year-round and throughout the study 

area. It was found in the continental shelf waters to 

bottom depths as shallow as 85 m, but it was not identified from any surface samples. Abralia veranyi is distributed in the tropical and subtropical eastern and western North Atlantic Ocean and the Mediterranean Sea; 

it is associated with continental shelf and slope waters 

and does not occur in open ocean. Two other species, 

A. redfieldi and A. grimpei, occur in the western Atlantic 

but their systematics and distribution are not well understood. Some of the specimens we refer to as Abralia 

cf. veranyi might include the other Abralia species whose 

paralarvae currently are not distinguishable. 

Morphology of voucher specimens 3.1 mm ML 

(Fig. 7A): Arms I–III long and attenuate with about 14 

suckers but no hooks; four chromatophores on aboral 

surface of each arm. Arms IV much less developed with 

six to eight suckers, no hooks. Arm order II>I=III>IV. 

Tentacular stalks very long and robust with row of large 

aboral chromatophores. Club region undifferentiated, 

minute suckers along distal one fourth of tentacle. A 

few small, integumentary photophores, evenly distributed over ventral and ventro-lateral surfaces of head, 

mantle, and funnel, most in association with small chromatophores. Three round, moderately well differentiated light organs (Fig. 7A) on ventral surface of eyes, anterior (#1) largest, posterior (#5) intermediate, central 

(#3) smallest; no trace of light organs in positions 2 or 4. 

Fins are very small, terminal flaps; meet posteriorly. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 11 

Figure 6 

Selenoteuthis scintillans Voss, 1958. A, dorsal view, 9.4 mm ML, USNM 730510, R/V Albatross IV sta. 

7206-71; B, tentacular club, same specimen; C, internal light organs, same specimen; D, light organs 

on ventral surface of eye (numbered 1–5 from top to bottom), same specimen; E, eye light organs, 

3.8 mm ML, R/V Albatross IV sta. 7206-56. 

4.5 mm ML (Figs. 7B, 8A–C): Arms II>I=III>IV, long, phores on Arms I–III and tentacular stalks. Tentacular 

attenuate. Relatively few suckers on arms, about 20 on stalks long and robust. Carpus with four to five suckers, 

I–III, 16 on IV; no hook development. Large chromato- four rows of suckers on manus with six to eight median 
12 NOAA Technical Report NMFS 152 

Figure 7 

Abralia cf. veranyi (Ruppell, 1844). A, light organs on 

ventral surface of eye, 3.1 mm ML, USNM 730511, R/V 

Albatross IV sta. 7302-79; B, light organs on ventral surface of eye, 4.5 mm ML, USNM 730511, R/V Albatross IV 

sta. 7302-79. 

suckers enlarged; no hook development. Head narrower than mantle opening. Seven large dark chromatophores on dorsal and lateral surface of head. Small photophores in longitudinal rows on ventral and lateral 

surface of head; one row extends onto Arms IV. Three 

well-defined light organs on ventral surface of eyes (Fig. 

7B); those in positions 1 and 5 largest and oblong, 3 

about one third smaller than 1 and 5 and round, no 

trace of organs in positions 2 and 4. Funnel strongly developed, extends to level of posterior edge of eye. Six 

small photophores on ventral surface of funnel. Mantle 

elongate and muscular, with broad opening that tapers 

to blunt posterior end. Many small chromatophores 

evenly distributed over ventral and ventro-lateral surface of mantle. Large chromatophores in bands around 

mantle, corresponding to photophores on ventral and 

lateral surfaces. Very large chromatophore on each 

postero-lateral end of mantle ventral to posterior part 

of fins. Fins terminal, muscular, short, triangular with 

rounded angles; meet at posterior end of mantle. 

Remarks Unlike some species of enoploteuthids, 

Abralia is comparatively rare in midwater-trawl collections in the Atlantic (Roper and Young, 1975). However, 

the abundance of this genus among the identified enoploteuthids from these plankton samples is comparable 

to that of the other members of the group of families 

(except lycoteuthids). There were 16 Abralia cf. veranyi 

vs. 17 Pterygioteuthis sp., 21 Abraliopsis pfefferi, 26 Ancistrocheirus lesueurii, and 30 Pyroteuthis margaritifera. If these 

proportions hold fairly constant among the 128 unidentified enoploteuthids from this study (another genus, 

Enoploteuthis, probably occurs among these specimens 

as well), then Abralia probably is more abundant in the 

western North Atlantic than is indicated by trawl studies, even though its distribution is restricted in comparison to other species. Abralia andamanica commonly 

is collected in plankton samples from the Indian and 

Pacific oceans (Silas, 1968; Yamamoto and Okutani, 

1975). Abralia spp. also are common in the plankton off 

Hawaii, where Young and Harman (1985) described the 

paralarval development of A. trigonura and A. astrosticta. 

Abraliopsis cf. pfefferi Joubin, 1896 

Voucher specimens 3.0 mm ML, Z76-247 sta. E3, 

USNM 816946; 5.0 mm ML, ALB7308 sta. 135, USNM 

816691; 13.0 mm ML, ALB7302 sta. 41, USNM 730509. 

Species distribution Specimens were distributed 

widely in both time and space. Abraliopsis pfefferi also was 

one of the few oceanic taxa collected during both years 

of the BLM program. However, it was not collected in 

any of the surface samples. Thus, the absence of this 

taxon from cruises for which all or most samples were 

collected with a neuston net (e.g. CI7802) may be a 

sampling artifact. In addition to oceanic areas, Abraliopsis pfefferi was found on the continental shelf over bottom depths as shallow as 66 m. 

Morphology of voucher specimens 3.0 mm ML: 

Mantle muscular, tapers posteriorly to blunt tip with 

minute terminal papilla. Fins very small, round. Funnel 

large, extends to posterior level of eyes. Arms extremely 

long, well developed. Arms IV with slight swelling near 

tip, the precursor of the light organ. Suckers only (9–16), 

no hooks, on Arms I–IV. Arm order III>II>I>IV. Integumentary photophores originally present, but deteriorated during preservation. Tentacles very long, stalks robust. Club with 10 biserial suckers proximally, then three 

hooks developing in next three pairs of suckers (two 

ventral, one dorsal), followed by about 20 suckers diminishing in diameter distally, some in three rows, with sucker anlagen at the tip. Eye light organs #1 (anterior), 3, 5 

developing as dark bronze reflective swellings. 

5.0 mm ML: Arms very long and well-developed. Arms 

IV with two small terminal light organs and median row 

of photophores on proximal half. Hooks absent on Arms 

IV. Four hooks present on Arms I, seven hooks on Arms 

II, nine hooks on Arms III. Photophores absent on Arms 

I–III. Three rows of photophores on head, row on ventral midline most conspicuous. Eye photophores 1 (anterior), 3, and 5 developed, 2 and 4 anlagen present. 

13.0 mm ML (Fig. 9): Arms IV with three terminal 

light organs, two rows of photophores that extend one 

half arm length, and 16 hooks. Arms I with 13 hooks, II 

with 17 hooks; no photophores. Arm order IV>III>II>I. 

Tentacular club with four large hooks in ventral row 

and five small hooks in dorsal row, carpus with eight 

suckers, dactylus with three to four rows of small suckers. Aboral keel of club developing. Two kinds of pho-
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 13 

Figure 8 

Abralia cf. veranyi (Ruppell, 1844). A, ventral view, 4.5 mm ML, USNM 730511, 

R/V Albatross IV sta. 7302-79; B, tentacular club, same specimen; C, oral view, same 

specimen. 
14 NOAA Technical Report NMFS 152 

Figure 9 

Abraliopsis cf. pfefferi Joubin, 1896. A, ventral view, 13.0 mm ML, USNM 730509, 

R/V Albatross IV sta. 7302-41; B, tentacular club, same specimen. 

tophores (large, spherical, dark; and small, spherical, 

translucent) in about nine indistinct rows on ventral 

surface of head. Ventral periphery of eyes with five photophores in single row; 1 and 5 largest, 2 and 4 small, 

3 intermediate. Thirteen small integumentary photophores around eyelids. Photophores on mantle and 

funnel similar to those on head, two kinds in indistinct 

rows; dense ventrally, sparse dorsally on mantle. Ventral 

midline devoid of photophores. 

Remarks In this study, all Abraliopsis paralarvae have 

been placed in A. pfefferi by default. Nesis (1987) considered pfefferi to be a subspecies of A. hoylei Pfeffer, 1884. 

Abraliopsis pfefferi is the most common species of this genus in the western North Atlantic (Roper, 1977; Lu and 

Roper, 1979) and our specimens probably belong to this 

species. In all, 21 paralarvae were identified as A. pfefferi. 

Nesis (1987) described A. atlantica from the tropical western Atlantic, Caribbean Sea, and Gulf of Mexico. Some 

of our southernmost material might belong to this species, but its paralarvae and juveniles are unknown. This 

genus also is common in plankton samples from the California Current (Okutani and McGowan, 1969) and the 

Indian Ocean (Silas, 1968). The paralarval development 

of two species of Abraliopsis from Hawaiian waters has 

been described by Young and Harman (1985). 

Family Ancistrocheiridae 

Ancistrocheirus lesueurii (d’Orbigny, 1842 in Ferussac & d’Orbigny 1834–1848) 

Voucher specimens 2.8 mm ML, BLM08W sta. L6, 

USNM 884242; 3.8 mm ML, GI7508 sta. 7, USNM 

884243; 5.6 mm ML, ALB7106 sta. 26, USNM 730528; 

8.2 mm ML, ALB7308 sta. 126, USNM 730529. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 15 

Species distribution Whereas this species is not very 

abundant in midwater-trawl samples (Roper and Young, 

1975), the 26 specimens reported here rank eleventh 

in abundance of planktonic cephalopods from the western North Atlantic Ocean, and eighth if only oceanic 

taxa are considered. This is one of the few species we examined that had a paralarval distribution that was limited in time and space. Although A. lesueurii was collected during May–November, no specimens were found in 

any winter samples. Similarly, this species was not identified from any sample taken south of Cape Hatteras but 

was found commonly in Gulf Stream waters from Virginia north. Yamamoto and Okutani (1975) reported 

young Thelidioteuthis alessandrinii (=A. lesueurii) absent 

from surface plankton samples, while present deeper, 

but we collected several specimens at the surface. Data 

ancillary to these surface samples indicate a temperature range of 17.6–20.9°C and a salinity range of 

33.6–35.8 × 10–3 for the young of this species. 

Morphology of voucher specimens 2.8 mm ML: 

Mantle rounded, cup-shaped, devoid of photophores. 

Fins terminal, small, weakly muscled. Tentacles long, robust, without photophores. Eleven suckers, no hooks 

on club; proximal sucker small, next five enlarged almost to diameter of tentacle stalk, distal five small with 

several anlagen distally. No hooks or photophores on 

arms. Arm suckers few and large; no suckers on proximal portions of arms. Five suckers on Arms I, six suckers 

on Arms II (none on proximal one third), two suckers 

on Arms III. Arms IV very short with only a few sucker 

anlagen. Photophores absent on head. Eyes and buccal 

assemblage stalked with gelatinous material filling spaces between stalks. 

3.8 mm ML: Mantle without obvious photophores. 

Fins small, rounded. Tentacles long, robust, without 

photophores. Fifteen suckers on club plus several distal anlagen, no hooks; all suckers large except proximal 1 and distal 2. Arm order II>I=III>>IV. Arm suckers large, few in number, none on proximal sections 

of arms. Eight suckers on Arms I, 12 on Arms II, 9 on 

Arms III, none on Arms IV. Head lacks detectable photophores. Eyes and buccal assemblage stalked with gelatinous material between stalks. 

5.6 mm ML (Fig. 10A–C): Mantle short, broad, bluntly rounded posteriorly, muscular; MW = 83% ML; 12 

photophores on ventral surface: four along anterior 

margin, four pairs that form two zig-zag rows posteriorly to the tip. Fins small, elongate, semilunar, posterior. Funnel tubular, base broad. Head broad, HW = MW. 

Two rows of five photophores on ventral surface of head 

in an arc from posterolateral corner to base of Arms 

IV. Arms long, robust, attenuate; arm order III>II>I>IV. 

Number of suckers on Arms I-15, II-16, III-22, IV-8; no 

hooks. Suckers relatively large, on long stalks. Tentacles 

long, robust to the attenuate tip. Suckers on club begin with one small proximal sucker, set apart from the 

rest. Manal suckers relatively large, biserial proximally, 

enlarge gradually to maximum diameter in third-sixth 

pairs, then diminish to tip; lateral suckers larger than 

medial suckers; about 27 suckers in total, no hooks. Six 

small spherical photophores embedded along the tentacular stalk. 

8.2 mm ML (Fig. 10D): Mantle muscular, broadest 

anteriorly, tapers evenly to bluntly rounded posterior 

tip; MW = 64% ML. Eighteen small, spheroidal photophores on ventral surface of mantle in distinct pattern 

of transverse rows, anterior to posterior: 4,2,2,4,2,2,2. 

The posterior-most photophores form at the very tip as 

elevated knobs. Fins rounded, triangular, terminal; FL = 

43% ML. Funnel large, base broad, tube extends to posterior level of eyes. Head large, HW > MW; eyes prominent, no ocular photophores. At least five photophores 

in arc on each side of ventral surface of head. Specimen 

damaged, so exact number of photophores on head and 

Arms IV cannot be determined. Arms very long, robust, 

attenuate; Arms I–III subequal, longer than IV. Armature on Arms I: one proximal sucker, five hooks, 13+ distal suckers; Arms II: 8–9 hooks, 14+ suckers; Arms III: 11 

hooks, 16+ suckers; Arms IV: 0 hooks, about 20 suckers. 

Tentacles long, robust to tip. Club with four pairs biserial carpal suckers; four transverse rows with two medial 

hooks and two large marginal suckers each (i.e., eight 

hooks) on manus; 8 to 10 diminishing biserial suckers 

on dactylus. Low aboral keel extends from level of first 

manal row to tip of dactylus. Tentacular stalk with nine 

small embedded spheroidal photophores. 

Remarks The early stages of A. lesueurii are easily 

confused with those of Octopoteuthis spp. (Figs. 16–17), 

because both are characterized by few large suckers on 

the tentacular clubs and by eyes and buccal assemblages 

on stalks embedded in gelatinous tissue. Octopoteuthis 

species, however, have a smaller number of suckers on 

the tentacular clubs. Bello (1992) presented a nomenclatural history of the names applied to this species. 

Family Pyroteuthidae 

Pyroteuthis margaritifera (Ruppell, 1844) 

Voucher specimens 1.6 mm ML, ALB7206 sta. 15, 

USNM 816692; 1.9 mm ML, ALB7302 sta. 64, USNM 

730520; 2.8 mm ML, station data missing; 3.8 mm ML, 

ALB7206 sta. 75, USNM 816693; 4.9 mm ML, ALB7302 

sta. 42, USNM 816694; 7.3 mm ML, ALB7302 sta. 42, 

USNM 816694; 13.8 mm ML, WIE7410 sta. 94, USNM 

816696. 

Species distribution Although P. margaritifera appeared to be widely distributed based on presence or 

absence in samples, 27 of the 30 specimens considered 

here were collected in the southern Sargasso Sea. Paralarval specimens collected off New England and Nova 
16 NOAA Technical Report NMFS 152 

Figure 10 

Ancistrocheirus lesueurii (d’Orbigny, 1842 in Ferussac & d’Orbigny 1834–1848). A, ventral view, 5.6 mm 

ML, USNM 730528, R/V Albatross IV sta. 7106-26; B, tentacular club, same specimen; C, oral view, same 

specimen; D, ventral mantle, 8.2 mm ML, USNM 730529, R/V Albatross IV sta. 7308-126. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 17 

Scotia probably were beyond their normal range, a result of northeastward transport by Gulf Stream meanders. This distribution contrasts with that of older specimens; P. margaritifera was very abundant in midwater net 

samples collected by Roper (1977) near Bermuda, off 

the middle U.S. Atlantic states (Lu and Roper, 1979), 

and off Nova Scotia (Vecchione, personal observ.). 

Young P. margaritifera were taken during all seasons. 

Similar to the vertical distribution found in older individuals (Roper and Young, 1975), none of these planktonic specimens were collected at the surface. 

Morphology of voucher specimens 1.6 mm ML: 

Anal light organs very small, separated, pigmented but 

non-reflective. Gill, intestinal, and posterior light organs undifferentiated. Eye light organs uncertain due 

to damage. Arms very short; Arms II and III with eight 

suckers, Arms I with six suckers, Arms IV with four buds. 

Tentacles missing (broken off). Eyes antero-lateral and 

tubular. Brachial/buccal crown tubular. Funnel very 

large. 

1.9 mm ML (Fig. 11A–E): Anal light organs small, 

spherical, pigmented, reflective, and separated. Light 

organs absent on gill, intestinal, and posterior areas. 

Seven light organs or anlagen (probably numbers 2, 3, 

4, 5, 6, 7, and 9; see Chun, 1910) detectable on each 

eye. Eye photophore 4 pigmented, 2,3,5 reflective (2–5 

shown in Fig. 11C); others just anlagen tissue knobs. 

Arms I–III with about eight suckers, Arms IV with about 

six buds. Tentacles long with 8 to 10 suckers on clubs; 

tentacular stalk with a bend, or slightly swollen elbow, 

at proximal one third, marking site of photophore anlagen. Eyes, brachial crown, and funnel as above. 

2.8 mm ML: Anal light organs posterior to anus, large, 

spherical, and separated. Gill light organs minuscule 

anlagen, non-pigmented, non-reflective. No indication 

of intestinal or posterior light organs. Nine eye light organs; 1, 6, and 8 non-reflective and non-pigmented, 9 

primordial. Eyes directed antero-laterally. Arms relatively long, without hooks, 14 to 16 suckers on each. Arms 

IV much shorter than I–III. Tentacles missing (broken 

off). Mantle/gladius tip blunt, almost rounded. Funnel 

large, long. Fins small, round, broadly separated. 

3.8 mm ML (Fig. 12): Anal light organs well developed, large, spherical, wide-set, slightly posterior to anus. 

Gill light organ anlagen visible only as spheres of differentiated tissue. Intestinal and posterior light organs not 

visible. Nine eye light organs, 1 and 8 non-pigmented 

and minute. All arms relatively long, no hooks, each 

arm has 12 to 18 suckers. Arm order III>II>I=IV. Tentacles long, clubs very short with 20 to 25 suckers, no 

carpal suckers, no hooks. Head broad. Funnel long. Fins 

round, paddle shaped. Mantle and gladius tip blunt. 

4.9 mm ML: Anal light organs fully developed. Gill 

light organs pigmented and small, about one fourth diameter of anal light organs. Precursor of intestinal light 

organ a faint strand of stained tissue in inverted-V between gill light organs. Nine eye light organs plus two 

anlagen (10 and 11). Arms with 18 to 22 suckers, fourth 

ventral sucker on Arms IV transformed into hook. Tentacle clubs with about 45 suckers, including one small 

carpal sucker. 

7.3 mm ML: Anal light organs large, well-developed. 

Gill light organs small. Intestinal light organs a single 

minute ring of differentiated tissue. Posterior visceral 

light organ a single very small anlage at terminal location. Nine eye light organs, plus two anlagen. Arms with 

20 to 26 suckers, two hooks at seventh pair on Arms II, 

three hooks at fifth pair on Arms III. No hooks on Arms 

I and IV. Tentacular club without hooks but with 55 to 

60 suckers and two enlarged carpal suckers two to three 

times the diameter of suckers on manus. Mantle and 

gladius tip elongate, pointed. 

13.8 mm ML (Fig. 13A–D): Juvenile female. Anal 

light organs large, fully developed, posterior to anus. 

Gill light organs at base of gill small, fully developed, 

sub-spherical. Three intestinal light organs developed, 

not fused; central photophore large, oval, pearly colored; laterals small, spherical, bronze-colored. Three 

posterior visceral light organs: terminal photophore 

in tip of mantle small, well-developed, spherical, darkpigmented; central and anterior photophores minute, 

round, unpigmented anlagen. Twelve eye light organs 

(nine large, three small), all reflective. Arm hooks in 

dorsal/ventral rows: Arms I-8/9; Arms II-8/8; Arms 

III-8/8; Arms IV-9/11. Tentacles longer than arms, 

slender, stalks with four embedded photophores. Four 

hooks on right tentacle, three on left in ventral median 

rows. Three enlarged carpal suckers larger than manal 

suckers. Mantle long, conical; gladius tip pointed and 

long. Head broad. Ventral and dorsal surfaces of head, 

arms, and bases of tentacles covered with very silvery reflective tissue. 

Remarks Early stages of P. margaritifera are easily 

confused with those of Pterygioteuthis spp. They can be 

distinguished by the relative sizes of the anal and gill 

light organs (ALO>>GLO in Pyroteuthis, GLO = or> 

ALO in Pterygioteuthis) and the shape of the gill light 

organs (round in Pyroteuthis, with a protuberance in 

Pterygioteuthis). 

Pterygioteuthis sp. 

Voucher specimens 2.3 mm ML, ALB7206 sta. 13, 

USNM 816697; 3.5 mm ML, ALB7302 sta. 52, USNM 

730524; 5.0 mm ML, ALB7302 sta. 79, USNM 730523; 

6.0 mm ML, ALB7308 sta. 144, USNM 730525; 9.0 mm 

ML, ALB7308 sta. 144, USNM 730525. 

Distribution Whereas Lu and Roper (1979) concluded that P. gemmata probably spawns during the 

spring in the middle U.S. Atlantic states, planktonic 

Pterygioteuthis were found throughout the year in our 
18 NOAA Technical Report NMFS 152 

Figure 11 

Pyroteuthis margaritifera (Ruppell, 1844). A, dorsal view, 1.9 mm ML, USNM 730520, 

R/V Albatross IV sta. 7302-64; B, visceral light organs, same specimen; C, vertical row 

of light organs on ventral surface of eye, same specimen; D, tentacle, same specimen; E, oral view, same specimen. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 19 

samples. This taxon also was collected throughout the 

study area. Yamamoto and Okutani (1975) reported 

that P. giardi off Japan was absent from surface plankton samples but present in the upper 200 m. Of our 17 

specimens of Pterygioteuthis sp., only a single specimen 

was collected in a surface net. Species of this genus are 

mesopelagic, but may ascend to the lower epipelagic 

zone (Young, 1978; Nesis, 1987). 

Morphology of voucher specimens 2.3 mm ML: 

Anal light organs already well-developed, silvery, pigmented. Gill light organs swollen, with minuscule 

black pigmented protuberance just developing. Intestinal and posterior light organs absent. Eyes directed 

antero-laterally; six to seven photophores. Arms I–III 

with eight suckers each, Arms IV very short and smooth, 

without suckers. Tentacle stalks long and robust, club 

tiny, terminal, with 15–20 suckers, none on carpus. 

3.5 mm ML (Fig. 14A–B): Anal light organs small, 

close-set, pigmented, lens developed. Gill light organs 

well-developed elongate spheroids; minute pigmented 

protuberance on postero-dorsal surface. Intestinal light 

organ developing, non- pigmented. Posterior light organs not developed. Eye light organs damaged, probably nine large plus two small anlagen. Arms short; Arm 

order III>II>I>IV. Arms II and III with about 20 suckFigure 12 

Pyroteuthis margaritifera (Ruppell, 1844). Ventral view, 

3.8 mm ML, USNM 816693, R/V Albatross IV sta. 

7206-75. 

ers on each, Arms I with 16, Arms IV with 10. Tentacles 

missing (broken off). Funnel short. Tip of mantle and 

gladius bluntly pointed. Head narrow. 

5.0 mm ML (Fig. 15A–B): Anal light organs large, 

well-developed. Gill light organs large spheres with 

very small black postero-dorsal protuberances. Intestinal light organ present, faintly pigmented. Posterior intestinal light organs not developed. Eyes directed laterally and adult-like. Nine large eye light organs, plus two 

small. Arm suckers: Arms I-16, Arms II-18, Arms III-18, 

Arms IV-10. Tentacle stalks very long, clubs very short 

with more than 40 suckers in four rows plus two to three 

carpal suckers. Tip of mantle plus gladius short, bluntly 

pointed. 

6.0 mm ML: Anal light organs with reflective silver 

anterior, black posterior. Gill light organs oblong, light 

brown with black postero-dorsal protuberance. Intestinal light organ spherical and pigmented; posterior intestinal light organs not developed. Eyes lateral. Eye 

light organs damaged, nine large, plus two small, plus 

two anlagen. Arm suckers: Arms I-17, Arms II-16, Arms 

III-17, Arms IV-approx.15. Protective membranes very 

well-developed; trabeculae robust. Tentacles missing distally but enlarged chromatophore present at base. Tip 

of mantle and gladius pointed, beginning to elongate. 

9.0 mm ML (Fig. 15C): Anal light organs large, half 

silvery, half black. Gill light organs large. Intestinal light 

organ well-developed, slightly ovoid, very black. Terminal intestinal posterior light organ present, small, 

black. Other two posterior light organs not present. 

Nine large eye light organs, plus four small. Hooks absent on arms; suckers on Arms I-24, Arms II-26, Arms 

III-23, Arms IV-17. Tentacles missing (broken off). Posterior mantle/gladius very pointed, adult-like. 

Remarks Two species, Pterygtioteuthis gemmata and P. 

giardi occur in the western North Atlantic. Our material 

may include either or both species. 

Family Octopoteuthidae 

Octopoteuthis sp. 

Voucher specimens 3.0 mm ML, KAI8201 sta. 47 

USNM 884239; 6.0 mm ML, KAI8201 sta. 45, USNM 

816716; 8.8 mm ML, BLM06W sta. A2, USNM 884240. 

Distribution The 30 young Octopoteuthis sp. reported here are of particular interest because adults and advanced juveniles are seldom captured in nets. Whereas 

Octopoteuthis spp. were rare in midwater-net samples examined by Cairns (1976), Clarke (1977), Roper (1977), 

and Lu and Roper (1979), stomach contents from 

sperm whales indicate that this genus is comparatively 

common in the North Atlantic (Clarke, 1977). The relative frequency of Octopoteuthis in the plankton samples 

we examined, as well as those reported by Okutani and 

McGowan (1969) from the California Current, indicates 
20 NOAA Technical Report NMFS 152 

Figure 13 

Pyroteuthis margaritifera (Ruppell, 1844). A, ventral view, 13.8 mm ML, USNM 

816696, R/V Wieczno sta. 7410-94; B, tentacular club, same specimen; C, pattern of light organs on ventral surface of eye, same specimen; D, left half of oral 

view, same specimen. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 21 

Figure 14


Pterygioteuthis sp. A, dorsal view, 3.5 mm ML, USNM 730524, R/V Albatross IV sta. 7302-52; B, oral view, same specimen.


that sampling for the young may be an adequate approach to a rough estimation of comparative abundance of this taxon, as has been done with other cephalopods (e.g. Kubodera and Okutani, 1977). Octopoteuthis 

sp. was widespread in our samples but never abundant; 

it was found in 27 samples but never more than two 

specimens in any one sample. It was collected throughout the year and throughout the western North Atlantic. However, 24 of 30 specimens were taken in the 

southern Sargasso Sea. Specific identities cannot be determined for our specimens, because four nominal species of Octopoteuthis are known to occur in the temperate and subtropical western North Atlantic (Stephen 

and Jefferts, 1992). 

Morphology of voucher specimens 3.0 mm ML 

(Fig. 16): Mantle short, broad, round posteriorly; MW 

about 75% of ML. Fins terminal, very short, broad, laterally ovoid; FL about 33% of ML; FW of both fins 

about 45% of ML. Tentacles long, robust proximally, 

thin distally with eight suckers. From proximal to distal, 

first two suckers minute, next two strikingly large (>tentacle diameter), followed by two intermediate in size, 

one slightly smaller, then one minute terminal sucker. 

Arms long, attenuate, with swollen areas at tips (anlagen to light organs). Biserial suckers on arms developing into hooks, some well-developed. Eyes and buccal 

assemblage stalked, with stalks embedded in gelatinous 

tissue of anterior head. 

6.0 mm ML (Fig. 17): Mantle thin walled, muscular, 

conical, widest at anterior opening, tapers to blunt tip; 

MW 60% of ML. Fins very muscular, fused along dorsal midline of mantle; blunt mantle tip extends very 

slightly beyond posterior border of fins. Fins longest at 

point of fusion with mantle; FL 60% of ML. Fins very 

broad, oval, FW 112% of ML. Funnel long, tapers to 

narrow opening that reaches mid-level of head, poste-
22 NOAA Technical Report NMFS 152 

rior to eyes; HW 60% of ML. Eyes on antero-laterally 

directed stalks; buccal stalk elongate. Eye and buccal 

stalks embedded in gelatinous matrix. No photophores 

Figure 15 

Pterygioteuthis sp. A, tentacular club, 5.0 mm ML, USNM 

730523, R/V Albatross IV sta. 7302-79; B, light organs on 

ventral surface of eye, same specimen; C, visceral light 

organs, 9.0 mm ML, USNM 730525, R/V Albatross IV sta. 

7308-144. 

nor photophore anlagen present on mantle or head. 

Arms long, attenuate; arm order II>III>I>IV, (most tips 

broken). Arms II and III, especially, are very robust 

in proximal half. Arms with biserial globular suckers 

that develop into hooks; hooks and suckers present on 

Arms II and III, suckers only on I and IV. Arm tips 

with swellings that are anlagen to photophores. Tentacles with robust stalks, but not nearly as muscular as 

arms; TL approximately equal to length of Arm I, longer than IV, considerably shorter than II and III. Club 

region unexpanded; two to three minute biserial suckers on proximal club, followed by two enormous suckers, each with a diameter greater than that of the tentacle where they are attached. Proximally are two more 

enlarged suckers, then one intermediate and one small 

sucker at the bluntly rounded tip of the club. No sucker anlagen occur on the tip. All sucker rings have relatively narrow openings and very broad walls. Anlagen 

of photophores on ink sac beginning to differentiate 

as a different appearance of the ventral-most reflective 

tissue. 

8.8 mm ML: Fin length (6.3 mm) 72% of ML. Fin 

width (about 12 mm) 136% of ML. Tentacles missing, 

but long shreds of tissue remain. Biserial hooks on 

all arms. Arm tips damaged, apparently from abrasion, 

causing loss of photophores. Eyes and buccal assemblage stalked, embedded in gelatinous tissue. 

Remarks Octopoteuthid development has been described in detail by Stephen (1985). Generic identifications can be determined easily for even the smallest 

paralarvae because of the huge suckers on the tentacular clubs (but see Ancistrocheirus); large post-paralarvae 

collected in plankton samples can be identified based 

on adult characteristics: absence of tentacles, presence 

of arm hooks and arm-tip photophores, large fin size, 

and body photophores. Very small specimens differ 

greatly from the adults because they lack the arm-tip 

photophores and the greatly enlarged fins, and they 

possess functional tentacles. The general appearance of 

Octopoteuthidae is quite similar to the early stages of 

Ancistrocheiridae, but they are distinguished by differences in armature of the tentacles. 

Family Onychoteuthidae 

Onychoteuthis cf. banksii (Leach, 1817) 

Voucher specimens 2.0 mm ML, Z7802, sta. 20, 

USNM 884235; 3.6 mm ML, KAI8201 sta. 75; 5.4 mm 

ML, KAI8201 sta. 78, USNM 884236. 

Species distribution The 94 specimens of Onychoteuthis cf. banksii in this study constitute the most abundant species collected except for the loliginids and the 

ommastrephids. This species was found in oceanic waters throughout the study area and was collected from 

February through August. Most of the specimens came 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 23 

Figure 16 

Octopoteuthis sp. A–C, aboral, oral, and lateral view of club, 3.0 mm ML, 

USNM 884239, R/V Kaiyo-Maru sta. 8201-47. 

Figure 17


Octopoteuthis sp. A, dorsal view, 6.0 mm ML, USNM 816716, R/V Kaiyo-Maru sta. 8201-45; B, ventral view, same specimen.

24 NOAA Technical Report NMFS 152 

Figure 18 

Onychoteuthis cf. banksii (Leach, 1817). A, dorsal view, 

2.0 mm ML, R/V Kaiyo-Maru sta. 8201-75; B, ventral 

view, same specimen. 

Figure 19 

Onychoteuthis cf. banksii (Leach, 1817). Ventral view, 5.4 

mm ML, KAI8201 sta. 78, USNM 884236. 

from the Sargasso Sea, although a few were collected 

in Gulf Stream waters. Specimens from surface samples were collected only at high salinities (S > 35.5 × 

10–3) and relatively high temperatures (18.2–28.8°C). 

Although O. cf. banksii was collected in 56 samples, all 

but three samples contained only one to three specimens (maximum=7). Thus the presumed “shoaling” by 

young juveniles of this species (Clarke, 1966) was not 

found within the planktonic size range (2–7 mm ML) 

available to us. 

Morphology of voucher specimens 2.0 mm ML 

(Fig. 18): Mantle very elongate, narrow, muscular; MW = 

42% ML. Posterior third of gladius and mantle bent ventrally. Posterior “spike” of gladius extends beyond very 

small, paddle-shaped, laterally directed fins. Funnel very 

large, well-developed, covers much of postero-ventral 

surface of head, extends anteriorly to level of lenses of 

eyes. Head narrow; HW = 28% ML, dominated by large, 

sessile, laterally oblong eyes. Arms III and IV not well 

developed, Arms II>I. Five suckers on Arms I, six on 

Arms II. Tentacles slightly longer than arms, with eight 

suckers. No hooks on clubs nor light organs on viscera 

or eyes. 

3.6 mm ML: Head completely retracted inside mantle, which is tightly sealed around head; only arms and 

tentacles protrude. Posterior third of gladius bent ventrally, and “spike” bent back dorsally. Fins small, round, 

well-developed. Arms III and IV mere protuberances. 

Arms I and II and tentacles short, stubby, all approximately of equal length. Suckers on arms and tentacles 

small, 12 each on Arms I and II and tentacles. Tentacular suckers appear to be biserial. 

5.4 mm ML (Fig. 19): Head narrow, partially withdrawn into mantle; HW 24% ML. Eyes lateral, relatively 

large. Mantle opening constricted to about two-thirds 

maximum mantle width; MW at opening 39% ML, MW 

at widest part in posterior half of mantle = 48% ML. 

Fins well-developed, broadly heart-shaped, FL = 14% 

ML, FW = 33% ML. Gladius tip still protrudes as “spike” 

beyond posterior end of mantle and fins; spike length = 

9% ML. Arm formula: I=II>>III>>IV. Arm suckers small, 

numerous on Arms I and II, few on Arms III, none on 

Arms IV. Tentacles slightly longer than longest arms. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 25 

Small biserial suckers along entire oral surface of tentacles, a few in central region beginning to elongate into 

hooks. Nuchal folds not yet developed. Intestinal light 

organs undeveloped. 

Remarks Young (1972) presented evidence that “O. 

banksii” in the western North Atlantic might represent 

a species complex of at least two species, but this has 

not yet been confirmed. If this is so, then our specimens 

may include more than one species. Furthermore, Ancistroteuthis lichtensteinii (Ferussac, 1835) now is known 

to be more abundant in the western Atlantic Ocean 

than previously recognized (Vecchione, personal observ.). Paralarval morphology of Onychoteuthis from Hawaiian waters indicates that a complex of very similar 

species is present there as well (Young and Harman, 

1987). Clarke (1992) presented additional observations 

on paralarval onychoteuthids. 

Onykia carriboea Lesueur, 1821 

Voucher specimens 1.5 mm ML, ALB7302 sta. 25, 

USNM 817004; 4.0 mm ML, TRI130, sta. 6, USNM 

816717; 7.5 mm ML, KAI8201, sta. 64, USNM 884241. 

Species distribution The young of this neustonic 

squid were collected in oceanic waters throughout the 

western North Atlantic and during all seasons. In the 

surface samples for which we have ancillary data, this 

species was taken at temperatures above 27.5°C and salinities greater than 35 × 10–3. Of the 51 specimens we 

examined, 16 were collected in a single 10-min neuston 

tow at 35°30′N, 74°00′W in the Gulf Stream off Cape 

Hatteras. 

Morphology of voucher specimens 1.5 mm ML: 

Mantle very stout, MW = 83% ML; rounded posteriorly, no point or rostral spike. Fins tiny, terminal, 

truncate-rounded. A line of chromatophores extends 

along dorsal midline of mantle. Funnel muscular, tube 

reaches posterior level of eyes. HW = 60% ML; eyes 

prominent, antero-laterally directed. Dorsal surface of 

head covered with several large chromatophores; one 

medium-sized chromatophore at base of dorsal arms. 

Arms I and II short, stubby, bluntly attenuate; suckers 

biserial, minute, few in number. Arms III minute, pointed and Arms IV merely blunt buds; no suckers. Tentacles longer than Arms II, slightly more slender, armed 

with a few biserial minute suckers along the stalk, club 

undifferentiated, sucker anlagen minute, numerous. 

4.0 mm ML (Fig. 20A–B): Mantle broad, bluntly 

pointed posteriorly; tip does not extend beyond level 

of fins; MW = 50% ML. Dorsal surface of mantle, 

head, and arms thickly covered with large and small 

chromatophores in two layers, the smaller overlying 

the larger. Small, retracted chromatophores scattered 

over ventral surface of mantle, head, and Arms IV. Fins 

well-developed, short, terminal, round, FW = 47% ML; 

FL = 19% ML. Eyes very large, lateral, completely covered with reflective tissue. Arm suckers biserial, decrease in size distally. Arm formula: II>III>I>IV. Tentacle length greater than Arms II. Tentacle suckers along 

proximal oral surface of stalk are biserial, become triserial, decrease in size and become quadraserial distally. 

7.5 mm ML: Mantle, funnel, arms, and tentacles all 

covered dorsally and ventrally with numerous small 

chromatophores. A few large chromatophores on dorsal surface of head. No nuchal folds. Mantle muscular. 

Fins subterminal, round, and large. Conus of gladius 

extends as a very short “spike” beyond posterior junction of fins and mantle. No hooks on tentacles. Arms 

short but robust, II>III>I>IV. Arm suckers biserial, darkly pigmented, well-developed along entire length of 

arms. Sucker counts: Arms I-12, Arms II-22, Arms III-18, 

Arms IV-10. Left tentacle missing. Length of right tentacle nearly equals Arms III. Tentacle suckers also darkly 

pigmented, all approximately equal in size but slightly 

smaller than arm suckers, quadraserial, numerous, covering all except base of oral surface of tentacle. Numerous sucker anlagen at distal tip of tentacle. 

Remarks Tsuchiya and Okutani (1992) hypothesize 

that some Onykia might be the young stages of Moroteuthis. 

Family Gonatidae 

Gonatus fabricii (Lichtenstein, 1818) 

Voucher specimen 6.9 mm ML, ALB7404 sta. 15, 

USNM 730519. 

Species distribution Gonatus was common only on 

Kaiyo-Maru cruise 8201 in February–March 1982, with 

six specimens collected. Two additional specimens were 

collected on separate cruises over the continental shelf, 

one in March and one in August. All specimens came 

from boreal slope water north of the Gulf Stream at the 

extreme northern part of our study area. 

Morphology of voucher specimen 6.9 mm ML 

(Fig. 21A–C): Mantle moderately long, bluntly rounded posteriorly: MW = 47% ML. Mantle chromatophores 

elongate: row of 13 to 14 encircle mantle just posterior 

to opening, three pairs along posterior half of dorsal 

midline, two pairs lateral to midline pairs, one very 

large chromatophore at base of each fin, a few others 

scattered on lateral and ventral surfaces. Fins subterminal, very small, wide, rounded; FW = 35% ML, FL = 10% 

ML. Funnel broad, robust, extends to mid-point of eyes. 

Eyes large, with ventral half covered with golden reflective tissue; diagonally directed anteriorly. Head short, 

broad; HW = 42% ML. Arms I–III well developed, especially II; Arms IV very short, much less developed than 

other arms. Arm formula: II>I=III>>IV. Basal two pairs 

of suckers on Arms I–III biserial; suckers increase to 

three to four rows at midpoint, then decrease to two 

rows of precursors at tips. Suckers on Arms IV biserial, 

minute. Arm sucker counts: I-32, II-36, III-36, IV-12. 
26 NOAA Technical Report NMFS 152 

Figure 20


Onykia carriboea Lesueur, 1821. A, ventral view, 4.0 mm ML, USNM 816717, R/V Trident sta. 130-6; B, oral view, same specimen.


Tentacles very long, robust, prominent; oral surface 

covered with more than 75 very small suckers that extend from near proximal base to distal tip. Proximal 

suckers widely spaced in two rows, then become closely 

packed in three then four rows. No evidence of enlarged suckers or hook development. 

Remarks Gonatus fabricii is common in ArcticSubarctic waters of the western North Atlantic. A borealzone eastern Atlantic species, G. steenstrupi, has been reported as far west as Newfoundland (Kristensen, 1981). 

Based on the known distributions of the adults, the Gonatus reported here probably are G. fabricii. Falcon et 

al. (2000) found that the most useful characters to distinguish between the paralarvae of the two species are 

the shape of the dorsal pad of the funnel organ (lateral 

sides straight in G. fabricii; curved in G. steenstrupi) and 

chromatophores on the ventral surface of the head (a 

pair slightly anterior to the ocular axis present in G. fabricii; absent in G. steenstrupi). 

Family Histioteuthidae 

Histioteuthis spp. 

Voucher specimens 1.6 mm ML, ALB7302 sta. 57, 

USNM 730527; 3.2 mm ML, ALB7302 sta. 87, USNM 

730526. 

Distribution Several species of this genus are found 

in the western North Atlantic (Voss, 1969; Voss et al., 

1992a). We collected Histioteuthis throughout the year 

in plankton samples, but these specimens are not identifiable to species. The 11 specimens of Histioteuthis in 

this study probably include several species because the 

development of characters is quite different for about 

the same mantle length, e.g., fin shape and dimensions. 

Seven specimens came from the southern Sargasso Sea 

and the other four were associated with the Gulf Stream 

in the central and northern regions of the study area. 

Based on midwater trawling in the middle U.S. Atlantic 

states, Lu and Roper (1979) concluded that H. reversa 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 27 

Figure 21 

Gonatus fabricii (Lichtenstein, 1818). A, dorsal view, 6.9 mm ML, USNM 730519, R/V Albatross IV 

sta. 7404-15; B, tentacular club, same specimen; C, oral view, same specimen. 

probably spawns in the spring. Clarke (1966) suggested that, because the young of Calliteuthis (=Histioteuthis) 

reversa usually are collected close to land, this species 

spawns on the continental slope. 

Morphology of voucher specimens 1.6 mm ML 

(Fig. 22A): Mantle muscular, very short, broad. Fins 

very small in proportion to mantle, close-set, wide, 

rounded only at tips. Funnel very large, muscular, 

broad; extends anteriorly nearly to base of arms. Head 

narrow, long. Eyes “sub-tubular,” dorso-ventrally elongate, antero-laterally directed. Arms I and II robust, attenuate, each with 12 suckers, central six somewhat enlarged and globular. Arms III small with eight suckers, 

only two to four slightly enlarged. Arms IV very short 

with two minute suckers. Tentacles missing distally (broken off) but proximal stalks robust. 

3.2 mm ML (Fig. 22B–E): Mantle muscular, very 

broad, rounded, much longer dorsally than ventrally; 

MW = 70% ML. Fins attached to each other, round, 

muscular, large; extend posteriorly well beyond man-
28 NOAA Technical Report NMFS 152 

Figure 22 

Histioteuthis spp. A, dorsal view of mantle, 1.6 mm ML, USNM 730527, R/V Albatross IV sta. 7302-57; B, dorsal view, 3.2 mm 

ML, USNM 730526, R/V Albatross IV sta. 7302-87; C, ventral view of mantle, same specimen; D, tentacular club, same specimen; E, oral view, same specimen. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 29 

tle; FW = 56% ML; FL = 31% ML. Reddish chromatophores thickly distributed on mantle, head, and funnel, especially on dorsal surfaces. Funnel very large, 

broad, but tapers to small opening. Head narrower 

than mantle, eyes “sub-tubular,” antero-laterally directed. Arms long, slender, tapered, bluntly attenuate. Arm 

formula: II>I>III>IV. Arms III only about two-thirds the 

length of II. Arm suckers large and few, except for 

Arms II which have much smaller suckers. Arms I with 

18 to 20 suckers, proximal six pairs enlarged; Arms II 

with 24 suckers, seven pairs enlarged; Arms III with 

16 to 18 suckers, five pairs slightly enlarged; Arms IV 

very short with four to six small suckers plus a few anlagen. Enlarged suckers very globular, spherical, with 

narrow openings. Tentacles robust, long, about onefourth longer than Arms II, suckers very small, proportionately few. Tentacular stalk with two single suckers 

near base; pairs of suckers begin sparsely at mid-stalk, 

then increase in numbers, first as pairs, then in more 

than two rows, up to six to eight rows across distal manus where suckers become smaller; minute suckers and 

anlagen buds only at dactylus. 

Remarks One useful feature to recognize Histioteuthis is the much smaller size of the tentacular suckers 

compared with the arm suckers. Species common in the 

western North Atlantic Ocean include: reversa, corona, 

arcturi and bonnellii (Vecchione, 2001). 

Family Bathyteuthidae 

Bathyteuthis abyssicola Hoyle, 1885 

Voucher specimen 4.3 mm ML, ALB7302 sta. 59, 

USNM 730514. 

Species distribution Our three specimens add little 

to what is known of the paralarval distribution of this 

species presented by Roper (1969). 

Morphology of voucher specimen 2.8 mm ML 

(Fig. 23A–C): Mantle muscular, relatively narrow, very 

blunt posteriorly, MW = 46% ML. Fins widely separated, minute, paddle-shaped, attached laterally to mantle 

anterior to tip. Funnel very large, long; tube extends at 

least to anterior level of eyes. Head small, short. Eyes 

antero-laterally directed, semi-stalked, with very large, 

bulbous red lenses. Numerous small red-brown chromatophores closely packed over all surfaces of mantle 

and head. Elongate ink sac completely enveloped in 

golden bronze reflective tissue. Arms very short, stubby, connected by web in proximal half; approximate 

order IV>III>II>I. Arm suckers on distal two-thirds of 

arms biserial but with the appearance of one zigzag 

row, six suckers on each arm. Tentacles very long, robust, blunt at tips. Club very small, confined to tips of 

tentacles; 18 minute suckers in two rows. 

Remarks Our three specimens, especially the described voucher, constitute among the smallest speciFigure 23 

Bathyteuthis abyssicola Hoyle, 1885. A, dorsal view, 4.3 

mm ML, USNM 730514, R/V Albatross IV sta. 7302-59; 

B, tentacular club, same specimen; C, oral view, same 

specimen. 

mens of this species to be described. The development 

and distribution of this species have been described by 

Roper (1969). 
30 NOAA Technical Report NMFS 152 

Family Chtenopterygidae 

Chtenopteryx sicula (Verany, 1851) 

Voucher specimens 2.2 mm ML, ALB7302 sta. 45A, 

USNM 816718; 3.2 mm ML, ALB7302 sta. 5, USNM 

816719; 3.5 mm ML, ALB7302 sta. 34, USNM 816720; 

3.6 mm ML, ALB7302 sta. 69, USNM 730522. 

Species distribution Little is known of the life history or ecology of this unusual species (Clarke, 1966; 

Roper and Young, 1975). The 37 specimens we collected 

provide some insight into the ecology of the paralarvae. 

These specimens were widely distributed both in time 

and in space; they were collected throughout the western 

North Atlantic Ocean and during all seasons. Although 

never particularly abundant (the maximum number of 

specimens taken in a single tow was six) the presence of 

this species in 23 samples from six cruises indicates the 

ubiquitous distribution of C. sicula. These results contrast 

with the paucity of adult C. sicula in trawl surveys. 

Morphology of voucher specimens 2.2 mm ML: 

Mantle large, muscular, broad, tapers to a point posteriorly; mantle much longer than head and arms combined. Fins minute, terminal flaps. Funnel large, robust, muscular, extends to level of mid-eye. Head short, 

dorso-ventrally compressed. Eyes prominent, tubular, 

sub-spherical, slightly dorso-ventrally elongate. Tentacles short, robust. Clubs terminal, broad, round, nearly 

equal in diameter to eyes, with about 25 suckers in 

round cluster of five to six suckers across. Arms IV longest. Arms II and III subequal, very short, stubby at 

bases; sudden attenuate papilla-like tips; two to three 

suckers. Arms I are minute papillae, just developing. 

Digestive gland sub-spherical. Ink sac well-developed, 

spherical, with concentration of bronze reflective tissue 

on ventral surface (precursor to photophore). 

3.2 mm ML: Fins very small, with short bases, but extend well posterior to mantle tip; about 10 short muscular supports (ribs) extend from muscular bases, connected by thin, easily torn membrane. Funnel very large, 

robust; extends anterior to base of arms. Head very short. 

Eyes tubular, directed antero-laterally, slightly elongate 

dorso-ventrally. Tentacle stalks robust, longer than arms. 

Clubs terminal, expanded, round, with about 20 suckers 

and a papilla-like dactylus. Arms short with attenuate tips 

and few very small suckers. Digestive gland globular, slightly elongate dorso-ventrally. Photophore anlage a round, 

reflective, bronze patch at ventral tip, ringed with black. 

3.5 mm ML: Fins terminal and lateral, extend far posteriorly beyond mantle tip; joined at posterior tip of mantle to form a deep notch. Series of about 12 muscular 

supports (“ribs”) connected by thin membranous tissue 

that is easily torn, giving comb-like appearance to fins. 

3.6 mm ML (Fig. 24): Tentacular clubs with attenuate 

dactylus that contains a few sucker anlagen. Otherwise 

similar to smaller specimens described above. 

Remarks Naef (1923) and Clarke (1966) presented 

a series of life history stages of C. sicula. 

Family Brachioteuthidae 

Brachioteuthis sp. 

Voucher specimens 3.0 mm ML, KAI8201 sta. 77, 

USNM 884237; 6.4 mm ML, BLM07W sta. J1, USNM 

884238. 

Distribution The 10 specimens of Brachioteuthis captured in this study were scattered among seven cruises. 

Thus, although rarely taken, paralarvae occur during 

each season throughout the western North Atlantic 

Ocean. 

Morphology of voucher specimens 3.0 mm ML: 

Mantle long, relatively broad, sac-like, muscular; MW = 

57% ML. Mantle covered with chromatophores. Fins 

very small, separate, postero-lateral, elongate flaps; FW = 

17% ML. Neck long, very narrow; neck length equals 

33% ML, neck width equals 10% ML. Head bulbous, 

long, broad; dorsal hump posterior to eyes; HL = 33% 

ML, HW = 30% ML. Eyes strongly oval in antero-ventral 

axis. Tentacles prominent, stalks robust, clubs slightly 

expanded, tips somewhat attenuate; 13 to 15 minute 

manal suckers in two to three rows; dactylus with anlagen only. TL = 73% ML; tentacle width = 13% ML. Arms 

I and II very short, II longer. Arms III and IV merely 

anlagen, papillae-like. Arms I with 0–1 suckers, Arms II 

with one to two very small suckers; several anlagen. 

6.4 mm ML: Neck length about 25% ML. Tentacles 

long, robust; clubs expanded with drawn out dactyli. 

Numerous (about 60) small suckers in up to six series 

across clubs at widest part, numerous anlagen distally. 

Arms very short; arm formula: II>>I=III>IV. Few (0–4) 

suckers on arms. Eyes sessile, antero-ventrally elongate. 

Fins small, round. 

Remarks Two species of Brachioteuthis, B. riisei, and 

B. beani are reported in the region covered by our study, 

but specimens are so rare in collections that the systematics of the family are not well known, even at the adult 

level. Consequently, we are not able to assign specific 

identifications to our material. See Roper and Sweeney 

(1992) for illustrations of paralarval Brachioteuthis. Shea 

(1995) presented information on taxonomy and distribution of paralarval Brachioteuthis spp. in the central 

North Atlantic. 

Family Ommastrephidae 

Illex sp. 

Taxonomy The development of Illex was described 

in detail by Roper and Lu (1979) and Vecchione 

(1979b), based on the MARMAP and BLM specimens, 

respectively. Roper and Lu (1979) used the MARMAP 

specimens from the middle U.S. Atlantic states to com-
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 31 

Figure 24 

Chtenopteryx sicula (Verany, 1851). A, dorsal view, 3.6 mm ML, USNM 730522, R/V 

Albatross IV sta. 7302-69; B, visceral light organ, same specimen; C, oral view, same 

specimen; D, tentacular club, same specimen; E, fin, same specimen. 

pare the morphology of paralarval Illex with the other for identification of Illex are: 1) all eight suckers at the 

genera of ommastrephids likely to be found in the west- tip of the fused tentacles (proboscis) are small and of 

ern North Atlantic. The characters that are most useful equal size, that is, the two lateral suckers are not en-
32 NOAA Technical Report NMFS 152 

larged as in some other genera of ommastrephids; and 

2) the fused tentacles are relatively short and robust 

(tentacle length index = 25.0–71.0). Furthermore, no 

photophore development occurs on either the eyes or 

the digestive gland. The stages in the development of Illex sp. are presented in Vecchione (1979b). 

Distribution Roper and Lu (1979) and Vecchione 

(1979b) also presented preliminary reports on the 

planktonic distribution of I. illecebrosus. These authors 

independently reached similar conclusions that the 

general distribution of planktonic I. illecebrosus was centered in the slope water between the shelf edge and 

the Gulf Stream. However, they reached different conclusions on the seasonality of young I. illecebrosus. Lu 

and Roper (1979) originally felt that the smallest specimens occurred only in winter and spring based on collections from a restricted area in the middle U.S. Atlantic states. However, after finding additional small 

rhynchoteuthion paralarvae from a broader area, Roper and Lu (1979) concluded that spawning takes place 

over an extended period, perhaps year-round. Based 

on a relatively large sample size, Vecchione (1979b) 

concluded that the winter-spring specimens are I. illecebrosus, but the summer specimens are too distinct 

ecologically to be assigned to I. illecebrosus without additional seasonal and regional captures. Two other Illex 

species, I. oxygonius and I. coindetii, are known from the 

western North Atlantic Ocean and could be included. 

Furthermore, Vecchione (1979a) believed that the predominant abundance of planktonic I. illecebrosus along 

the shelf break and continental slope and the absence 

of eggs in the plankton samples supported earlier proposals that this species spawns on the slope, perhaps in 

contact with the bottom. Illex illecebrosus has since been 

shown to be a pelagic spawner (O’Dor et al., 1982) 

with a large gelatinous egg mass. The egg mass could 

easily be pushed aside by the pressure wave of a bongo 

sampler and thus not be collected. Spawning on the 

bottom at the continental slope, therefore, is not a likely 

explanation for the observed distribution of planktonic 

Illex. 

More recent data, including our specimens from 

Kaiyo-Maru cruise 8201, indicate a close relationship 

between Illex paralarval distribution and the interface 

between the slope water and the Gulf Stream (Dawe 

and Beck, 1985; Rowell and Trites, 1985; Vecchione 

and Roper, 1986; Dawe and Stephen, 1988; Perez and 

O’Dor, 2000). Our specimens from the Kaiyo-Maru 

cruise were collected at stations with a wide range 

of surface temperatures, about 7–20°C. However, at 

all of these stations, temperature values converged to 

13–18°C at 150–200 m depth (see Vecchione and Roper, 

1986). Because the samples were collected by oblique 

tows between 0–200 m, we do not know the precise 

depths at which the specimens were collected. 

The distribution of I. illecebrosus in the BLM samples 

was very similar to that of pteropod species known to migrate vertically to depths of 200 m or more (Vecchione 

and Grant, 1983). Furthermore, almost all of the BLM 

specimens came from subsurface samples; relatively few 

were collected at the surface (Vecchione, 1979b). O’Dor 

et al. (1982) suggested that I. illecebrosus spawns at the 

interface between slope water and the Gulf Stream. This 

interface would be encountered at the deep end of the 

depth range sampled in our study. The temperatures 

and salinities at depths of 150–200 m at the KAI8201 

stations from which we have planktonic I. illecebrosus indicate that this is a region of isopycnal mixing at σt of 

approximately 26.7 (Vecchione and Roper, 1986), between what Wright and Parker (1976) refer to as the 

“slope-water thermostad” and Gulf Stream water. 

Further support for this relationship between Illex 

paralarval distribution and the slope water/Gulf Stream 

interface has been presented by Hatanaka et al. (1985), 

based on logged entries of field identifications of all 

specimens from Kaiyo-Maru cruise 8201. Rowell and 

Trites (1985), described paralarval distribution in the 

area of overlap of I. illecebrosus and I. oxygonious distribution between Cape Canaveral and Cape Hatteras. In addition, the few specimens of paralarval Illex that were 

taken in the surface samples on Columbus Iselin cruise 

7802 also came from the interface between slope water 

and the Gulf Stream. Our own long-term research in 

the waters off Ft. Pierce, Florida, confirms that Illex 

paralarvae occur only in the narrow band of “transitional” water that lies just shoreward of the Florida Current 

(Gulf Stream) against the outer boundary of the Florida Coastal Water mass (slope water) (Adams, 1997). 

The absence of Illex specimens in both surface and 

subsurface samples from unmixed slope water during 

the VIMS winter cruise to the Norfolk Canyon area (see 

Ruzecki, 1979 for water-mass analysis), reinforces the 

hypothesis that it is the interactions between the slope 

water and the Gulf Stream that are of critical importance to Illex spawning, paralarval distribution, and 

recruitment. Because the physical characteristics of 

the “slope-water thermostad” and the Gulf Stream remain relatively unchanged year-round, prolonged or 

non-seasonal spawning by I. illecebrosus is quite possible, 

as suggested by Lu and Roper (1979). 

Remarks Based on the distribution of the specimens discussed in Roper and Lu (1979) and Vecchione 

(1979b), both reports probably described the northern 

species, Illex illecebrosus. Two other species have been 

reported from the warmer waters of western North Atlantic, I. oxygonius and I. coindetii (Roper et al., 1969). 

Therefore, until comparisons can be made among specimens of known parentage, the morphological characters agreed upon by Roper and Lu (1979) and by Vecchione (1979b) must be considered to be of generic 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 33 

importance. Two recent studies have elucidated the systematics of I. coindetii (Roper and Mangold, 1998) and 

of the entire genus of Illex (Roper et al., 1998). 

Rhynchoteuthion Type A′ 

Distribution Twenty-seven specimens identified as 

rhynchoteuthion Type A′ under-represent the relative 

abundance of this taxon; 124 additional rhynchoteuthion paralarvae could be identified only as Ommastrephidae, but undoubtedly these included many specimens of 

rhynchoteuthion Type A′. Rhynchoteuthion Type A′ was 

collected during all seasons, but taxonomic difficulties 

preclude definite statements about temporal or spatial 

distribution. As noted below, rhynchoteuthion Type A′ 

probably is Ommastrephes bartramii. This typically oceanic 

taxon was found over bottom depths as shallow as 85 

m, a result of cross-shelf transport involving Gulf Stream 

eddies (Vecchione and Grant, 1983). Such transport, together with mixing of shelf and oceanic water masses, is 

reflected in the very broad salinity range (S = 32.6–36.6 

× 10–3) for specimens collected in surface samples. The 

temperature range for this taxon at the surface was limited to 18.2–25.3°C. On the transect from Bermuda to 

Rhode Island (cruise CI7802), rhynchoteuthion Type A′ 

was collected only near the offshore edge of the Gulf 

Stream. Similarly, off Nova Scotia (cruise KM8201) this 

species was taken predominantly at the southernmost 

stations, again indicating a tendency to be concentrated 

at the offshore edge of the Gulf Stream. 

Taxonomy Roper and Lu (1979) described larvae as 

“rhynchoteuthion Type A′” from the MARMAP collections, which were similar to Yamamoto and Okutani’s 

(1975) “Type A.” The two lateral suckers at the tip of 

the fused tentacles are enlarged, two to three times 

the diameter of the remaining six suckers. The length 

of the fused tentacles is short to moderate (tentacle 

length index = 30.0–70.0). A reflective patch is located 

on the ventral surface of each eye, and a small, round, 

well-defined light organ is located near the anterior 

end of the intestine, between the intestine and the ventral surface of the digestive gland. This combination of 

characters leads to the suggestion that this is the paralarval form of Ommastrephes. Young and Hirota (1990) described development of Pacific Ommastrephes bartramii, 

but no visceral photophore was present. Detailed descriptions and illustrations of rhynchoteuthion Type A′ 

paralarvae in the mantle length range of 1.00–7.25 mm 

are given in Roper and Lu (1979). Wormuth et al. 

(1992) presented some additional information. 

Remarks Goldman and McGowan (1992) reported 

the summer distribution of rhynchoteuthion paralarvae 

off the Florida Keys, based on discrete-depth samples. 

They found that the peak of rhynchoteuthion abundance occurred at the Gulf Stream front. The most 

abundant type was Type A′, which tended to be caught 

at shallower depths than Type B′. Similarly, Saito and 

Kubodera (1993) found vertical separation of rhynchoteuthion types in the vicinity of the Kuroshio Current off Japan. 

Rhynchoteuthion Type B′ (revised) 

Distribution This morphotype was found throughout the year and probably is distributed throughout 

the study area. As noted for the other ommastrephids 

in this study, rhynchoteuthion Type B′ occasionally was 

transported into the continental shelf waters to depths 

as shallow as 46 m. 

Taxonomy Roper and Lu (1979) described as “Rhynchoteuthion Type B′” larvae that they thought belong to 

Ornithoteuthis antillarum. We now believe that some rhynchoteuthion Type B′ include paralarvae of Sthenoteuthis 

and possibly of Hyaloteuthis. Refer to Roper and Lu (1979) 

for the detailed description and illustration of rhynchoteuthion Type B′ from which the following is summarized. Rhynchoteuthion characterized by having the two 

lateral suckers on the tip of the fused tentacles (proboscis) equal in size or slightly larger than the remaining six 

suckers, but not enlarged to twice the diameter of the 

small suckers. The fused tentacles are very thin, narrow, 

and moderately long to very long (tentacle length index 

= 54.0–137.0). A single relatively large, round, raised photophore occurs on the ventral surface of each eye. A distinct round photophore is present between the intestine 

and the digestive gland. The variability in relative sizes of 

suckers and tentacles indicates possible inclusion of multiple species in rhynchoteuthion Type B′. 

Harman and Young (1985) described paralarvae of 

Sthenoteuthis oualaniensis from Hawaiian waters that are 

very similar to Roper and Lu’s (1979) Type B′. Two 

species belong to this genus: S. pteropus which occurs 

broadly in the tropical/subtropical Atlantic Ocean from 

approximately 40°N to 30°S and S. oualaniensis, a widespread species throughout the tropical Pacific and Indian oceans. Thus it seems safe to assume that the species 

represented here includes S. pteropus. Ommastrephids 

in our study area, besides Illex spp., O. bartramii, and S. 

pteropus, include Ornithoteuthis antillarum and Hyaloteuthis pelagica. The latter two species also may be included 

in rhynchoteuthion type B′. 

Remarks Among the ommastrephids that we have 

identified, almost as many specimens were rhynchoteuthion Type B′ (21), as were rhynchoteuthion Type 

A′ (27). Thus, a substantial number of the 124 unidentified rhynchoteuthion specimens in this study are 

likely to include Sthenoteuthis, Ornithoteuthis, and Hyaloteuthis. Contrary to inferences from sampling for adults 

(Clarke, 1966), O. antillarum probably is not particularly rare in the western North Atlantic based on observations from submersibles (Vecchione and Roper, 1992), 

from which O. antillarum is seen frequently. 
34 NOAA Technical Report NMFS 152 

Family Chiroteuthidae 

Chiroteuthis sp. 

Voucher specimen 5.6 mm ML, ALB7302 sta. 71, 

USNM 730518. 

Distribution Only three specimens were collected, 

so nothing can be ascertained about the distribution except where they were caught. 

Morphology of voucher specimen 5.6 mm ML 

(Fig. 25): Mantle long, narrow, thin-walled, with scattered 

chromatophores; tapers abruptly to anterior insertion of 

fins, extends posteriorly into long spike-like tail (broken). 

Fins short, broad, elliptical. Tip of gladius broken (tail 

missing). Funnel short, narrow, weakly developed. Neck 

long, cylindrical, semi-gelatinous, with a few scattered 

chromatophores; about 3.4 mm long from dorsal mantle 

margin to posterior edge of eyes. Head small, narrow. 

Eyes occupy entire anterior portion of head, bulge ventrally; antero-ventral surface of eyes covered by large chromatophores. Eye light organs not distinguishable. Brachial crown displaced dorsally. Arms very short, stubby. Arm 

formula I=II>IV>>III (Arms III present only as minute anlagen). Arms I and II each with two relatively large suckers that occupy entire oral surface. One sucker each on 

Arms IV. Tentacles very long, robust; oral surface covered 

with minute suckers, beginning with one “large” proximal 

sucker, two to three times diameter of other tentacular 

suckers, at base of stalk, then a single small sucker, followed distally by six pairs of equal-sized suckers. Manus 

suckers in two rows proximally, increasing rapidly to three 

then four rows; suckers end abruptly at dactylus where 

only minute, very closely packed precursors occur; manus 

suckers total about 70. Digestive gland bulbous, elongate, 

vertically oriented; precursor of light organ present as 

2-tone bronze reflective tissue on ventral half; ventral-most 

part set off from upper part by dark ring. 

Remarks Paralarval chiroteuthids are referred to as 

a “doratopsis” stage. Doratopsis paralavae also occur in 

Planktoteuthis, Asperoteuthis, and Grimalditeuthis. The most 

common species in our study area is C. veranyi, but C. 

capensis and C. joubini have been reported from the area 

as well (Passarella and Hopkins, 1992). Vecchione et al. 

(1992) described tail morphology of a Chiroteuthis paralarva (not C. veranyi) collected in the western Atlantic by submersible. Young (1992) provided detailed descriptions 

and comparisons of Hawaiian doratopsis paralarvae. 

Family Mastigoteuthidae 

Mastigoteuthis hjorti Chun, 1913 

Voucher specimen 6.0 mm ML, ALB7302 sta. 71, 

USNM 730521. 

Species distribution Nothing can be inferred about 

the distribution of this species based on the two specimens collected at a single station. Adults, however, apFigure 25 

Chiroteuthis sp. A, dorsal view, 5.6 mm ML, USNM 

730518, R/V Albatross IV sta. 7302-71; B, tentacular club, 

same specimen; C, oral view, same specimen. 

pear to be cosmopolitan in tropical and subtropical waters of all major oceans, although the species is not 

found in Hawaiian waters. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 35 

Figure 26 

Mastigoteuthis hjorti Chun, 1913. A, dorsal view, 6.0 mm ML, USNM 730521, 

R/V Albatross IV sta. 7302-71; B, ocular light organ, same specimen; C, tentacular club, same specimen; D, oral view, same specimen. 

Morphology of voucher specimen 6.0 mm ML light organ on ventral surface of each eye. Arm formu­

(Fig. 26): Mantle long, narrow; tapers gradually to an- la: II>I>IV>>III (Arms III present only as minute anlaterior insertion of fins. Fins evenly rounded; fin length gen buds). Tentacles long, robust, about four times the 

about 25% of ML (exclusive of tail); tail very long, length of Arms II. Clubs with about 54 small suckers in 

spike-like, nearly three times as long as fins. Skin mostly two rows proximally, graded to six rows along “manus.” 

rubbed off but fragments around fins have scattered tu- Suckers end abruptly, followed by a number of sucker 

bercles. Head small, eyes protrude antero-laterally; one anlagen on dactylus. 
36 NOAA Technical Report NMFS 152 

Figure 27 

Leachia lemur (Berry, 1920). A, ventral view, 13.8 mm ML, USNM 816942, 

R/V Columbus Islen sta. 7802-20; B, tentacular club, same specimen; C, oral 

view, same specimen. 

Remarks The presence of a photophore on each 

eye suggests identification of these Mastigoteuthis as M. 

hjorti, which has two ocular photophores as subadults. 

Family Cranchiidae 

The comparative morphology and systematics of larval 

cranchiids have been examined in detail by Voss (1980) 

and Voss et al. (1992b). Our material includes specimens of Leachia lemur (Berry, 1920) (Fig. 27), Teuthowenia megalops (Prosch, 1849) (Fig. 28), Helicocranchia cf. 

papillata (Voss, 1960) (Fig. 29), and ?Taonius/Galiteuthis 

sp. 

Leachia lemur (Berry, 1920) 

Voucher specimen 13.8 mm ML, CI7802 sta. 20, 

USNM 816942. 

Species distribution Generic distribution circumglobal in tropical, subtropical waters; L. lemur occurs 

in the North Atlantic Subtropical Region, west of 

about 50°W (north and south Sargasso Sea) (Voss et 

al., 1992b). Our 11 specimens were collected only in 

winter. 

Morphology of voucher specimen 13.8 mm ML 

(Fig. 27): Mantle elongate, sharply tapered posteriorly; 

gladius broad posteriorly then tapers to short, stout, 

sharp spike; muscular tissue of mantle ends near anterior borders of fins; MW = 28% ML. Fins small, transversely elliptical, length equals width on each fin. Single, narrow, tubercular, cartilaginous strip on ventral 

mantle extends posteriorly from each funnel-mantle 

fusion; occupies about 35% of ML; tubercles simple, 

a single point, two points, and three points aligned 

transversely with axis of strip. Funnel very large. 

Head per se minute, but dominated by very large 

antero-lateral, transparent, tubular eye stalks and very 

long, narrow muscular arm-crown stalk. Eyes relatively 

small, dorso-ventrally elliptical; small patch of golden 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 37 

Figure 28 

Teuthowenia megalops (Prosch, 1847). A, ventral view, 6.2 mm 

ML, USNM 816721, R/V Albatross IV sta. 7203-59; B, oral view, 

same specimen; C, eye with ocular light organ anlage, same 

specimen. 

reflective tissue developed on ventral surface, anlagen 

of two photophores barely detectable. Arms minute, 

order III>>IV>II>I; seven suckers on Arm III, globular 

grading to tiny, three on IV and one to four or “variable” each on I and II. Tentacles extremely long, robust: TL = 52% ML. Clubs not expanded; suckers develop at extreme proximal area of tentacular stalk with 

a zig-zag line of nine minute suckers followed distally 

by the tiny manal suckers in four rows, and minute anlagen at the tip. 

Remarks Paralarval Leachia have been referred to 

as a “Pyrgopsis” stage. The simplicity of the tubercles 

on the ventral mantle cartilaginous strips, as well as 

the geographical localities of captures, lead us to identify this species as L. lemur. The smallest specimen of 

Leachia identified in our samples was 12.5 mm ML. 

Teuthowenia megalops (Prosch, 1847) 

Voucher specimen 6.2 mm ML, ALB7203 sta. 59, 

USNM 816721. 

Species distribution Teuthowenia megalops is found 

in the North Atlantic Ocean in subarctic and highly 

productive areas of the North Temperate region (Voss, 

1985). Paralarvae in the present material were collected 

only in the winter. 

Morphology of voucher specimen 6.2 mm ML 

(Fig. 28): Mantle stout sac-like, blunt posteriorly. Fins 

(broken) very small, separate. Funnel very large, broad. 

No tubercles on funnel-mantle fusion at this stage. 

Head small, but eye stalks stout, relatively short; eyes 

small, dorso-ventrally elliptical, reflective tissue developing, very slight ventral rostrum. Arm-crown stalk short, 

stout. Arms very small, order I=II>>III=IV. Arms I and 
38 NOAA Technical Report NMFS 152 

Figure 29 

Helicocranchia cf. papillata (Voss, 1960). A, ventral view, 15.6 mm ML, 

USNM 816722, R/V Albatross IV sta. 7206-74; B, tentacular club, same 

specimen; C, oral view, same specimen. 

II stout, conical with 5–6 suckers, globular grading to near tip. Large chromatophores are typical for this spesmall; Arms III and IV very small, conical; two to three cies, beginning at early paralarval stages. 

minute suckers and two to four sucker anlagen on III, Remarks This species is the only Teuthowenia that 

one to two sucker anlagen on IV. Tentacles moderately occurs in the western North Atlantic region sampled by 

large, stalks covered with suckers from proximal base in the programs from which we have material. Voss (1985) 

two rows that rapidly become four rows along stalk to described in great detail all growth stages and the distri-
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 39 

bution of this species. Teuthowenia megalops commonly 

is seen from submersibles off New England (Vecchione 

and Roper, 1992). 

Helicocranchia cf. papillata (Voss, 1960) 

Voucher specimens 5.5 mm ML, ALB7206 sta. 44, 

USNM 816723; 15.6 mm ML, ALB7206 sta. 74, USNM 

816722. 

Species distribution Helicocranchia papillata is a 

warm-water form distributed widely in the western 

North Atlantic Subtropical Region, Caribbean Sea, and 

Gulf of Mexico (Voss et al., 1992b). Our specimens were 

caught only in the summer in the eastern and southernmost area of our samples. 

Morphology of voucher specimens 5.5 mm ML: 

Mantle elongate, thin walled, muscular, bluntly rounded 

at tip; MW = 33% ML. Fins subterminal to mantle, wide, 

paddle shaped, inserted on short, stout shield-shaped 

posterior rostrum of gladius; FL = 13% ML; FW = 

35% ML. Funnel very large; extends to level of buccal 

mass. Head small. Eyes covered with reflective tissue, 

dorso-ventrally elongate, with developing ventral rostrum; eyes on short, stout stalks. Arm crown short. Arms 

short, order II>I>III>>IV; Arms IV merely conical buds. 

Sucker number on arms: 6 on Arm I, 11 to 12 on Arm 

II, 6 on Arm III. No enlarged suckers on Arms III at 

this stage. Tentacles long, robust; TL = at least 65% ML; 

minute suckers extend entire length of tentacular stalk, 

none enlarged at this stage. 

15.6 mm ML (Fig. 29): Posterior tip of rostrum of 

gladius broad, rounded; dorsal rostrum shield-shaped 

or lanceolate; posterior borders of fins united by thin 

membrane. Funnel huge, extends beyond level of buccal mass. Dorsal element of funnel organ with swollen 

round limbs and apex; one short, sharp papilla on apex 

and on each limb; ventral pads oval. Eyes large, on short 

stalks; with prominent ventral rostrum. Arms small, order III>II>I>>IV; Arms III with four pairs of enlarged 

suckers in distal third, beginning with eighth pair. Tentacles very long, robust proximally, tapered distally; a few 

suckers enlarged along ventral row of manus; TL > ML. 

Remarks This species is identified as Helicocranchia 

based on the arrangement of the fin attachment to the 

broad, short rostrum of the gladius, the funnel organ 

shape and papilla size, and the enlarged suckers on 

Arms III and the club (in the larger specimen). 

Other Cranchiidae 

Distribution These are general comments on our 

unidentified cranchiids. The scarcity of cranchiids in 

these samples, especially the “Pyrgopsis” stage of Leachia 

spp., is somewhat surprising. Based on other studies 

(Okutani, 1974; Clarke and Lu, 1975; Roper and Young, 

1975; Young, 1975; Lu and Roper, 1979), these taxa 

should be relatively common in the near-surface plankton. However, we collected only 11 specimens of Leachia 

lemur, all in the “Pyrgopsis” stage of development. Other 

cranchiids were quite rare, with five or fewer specimens 

of each taxon. 

The cranchiids were among the few taxa which appeared to be seasonal in distribution. Leachia lemur, Teuthowenia megalops, and ?Taonius were collected only in 

the winter, whereas Heliocranchia cf. papillata was collected only in summer in the southernmost region. Although Leachia lemur was found throughout the western 

North Atlantic, no cranchiids were collected in waters 

over the continental shelf. In the samples from Columbus Iselin cruise 7802 and Kaiyo-Maru cruise 8201, young 

Leachia lemur appear to be concentrated near the offshore edge of the Gulf Stream. 

Family Octopodidae 

Octopodid spp. 

General comments on identification Recent work 

on paralarval octopod systematics, especially Pacific species, has shown that chromatophore patterns and sucker arrangements are promising characters for identification of species (Young and Harman, 1989; Hochberg 

et al., 1992). The systematics of all octopodids however, 

still is in serious need of revision. 

Sixty octopodids were collected in our study. In spite 

of morphological diversity among these specimens, we 

are unable to identify them to species due to taxonomic 

problems such as bleaching of chromatophores that are 

important species-distinctive characters. However, two 

basic morphotypes were evident. 

Voucher specimens Octopus sp.-type (all arms subequal in length): 1.9 mm ML, ALB7302 sta. 74, USNM 

816724 (Fig. 30); 7.4 mm ML, ALB7404 sta. 18, USNM 

816866 (Fig. 31); 7.9 mm ML, WIE7410 sta. 11, USNM 

816865. “Macrotritopus”-type (Arms III greatly elongated 

and enlarged): 2.4 mm ML, DE7219 sta. 129, USNM 

816867 (Fig. 32); 6.6 mm ML, WIE7410 sta. 94, USNM 

730507 (Fig. 33). 

Species distribution The few “Macrotritopus”-type 

octopods were widely scattered both in time and in 

space. Young Octopus-type octopods are very common in 

plankton samples, although usually not abundant (e.g. 

Rees, 1950; Silas, 1968; Lu and Roper, 1979). In the 

present study, these octopodids were collected on 11 of 

21 cruises. Furthermore, they were found in all sampled areas where bottom depths were greater than 100 

m and during all seasons. Octopodids comprised the 

fifth most abundant “taxon” (60 specimens) in our collections and were found in both surface and subsurface samples. At the surface, they were collected at 

19.2–22.5°C and at S = 33.2–36.4 × 10–3. 

Remarks Uncertainty existed about the identity 

of the “Macrotritopus”-type octopods for many years. 
40 NOAA Technical Report NMFS 152 

Rees (1954) suggested they probably were the young 

of Scaeurgus unicirrhus (d’Orbigny, 1840). Boletzky 

(1977b), however has reared S. unicirrhus and found 

that it does not have a “Macrotritopus” paralarva. Hanlon et al. (1979) captured and raised a “Macrotritopus” 

which proved to be Octopus defilippi Verany, 1851. It is 

possible however that this morphotype is characteristic 

of more than one octopodid species. 

Family Tremoctopodidae 

Tremoctopus violaceus Chiaie, 1830 

Voucher specimens 2.3 mm ML, WIE7410, sta. 94, 

USNM 730508; 7.5 mm ML, ALB7302, sta. 71, USNM 

816868. 

Species distribution Thirty-six specimens were taken in the southern Sargasso Sea in February and July; a 

single additional specimen came from the continental 

shelf off New England in the summer. 

Morphology of voucher specimens 2.3 mm ML 

(Fig. 34): Mantle short, broad, covered with small, evenly spaced chromatophores. Large internal chromatophores on viscera visible through dorsal mantle. Funnel 

extends to base of Arms IV. Head short, broad, with 12 

large chromatophores on dorsal surface. Arms I disproportionately enlarged, length greater than ML; all but 

distal two suckers are large, globular; third from base 

greatly enlarged, about two times diameter of next largFigure 30 

Octopus sp.-type (all arms subequal in length). A, ventral 

view, 1.9 mm ML, USNM 816724, R/V Albatross IV sta. 

7302-74; B, oral view, same specimen. 

est. Proximal three to four suckers on Arms I uniserial, 

remainder biserial. Arms II–IV much shorter; thinner, 

with small suckers, uniserial proximally, but biserial toward tips. Arm formula I>>>II>IV>>III. Sucker counts: 

Arms I-15, Arms II-6, Arms III-2, Arms IV-5. Narrow web 

connects all arms, deepest between Arms I, proportionally shallower with each pair ventrally. Single row of 

large chromatophores on aboral surface of each arm; 

one large chromatophore on base of each sucker. 

7.5 mm ML: Two pairs of pores on head: one pair between bases of Arms I and II, one pair at bases of Arms 

IV on each side of funnel opening. Mantle plump, triangular; mantle opening very wide, extends dorsally to 

level with eye lens. Head wider than mantle opening. 

Eyes large; occupy entire lateral surface of head. Funnel large, tapers to anterior of eyes, tip free. Arms I and 

II extremely long. Arm formula: I>II>>IV>III. Suckers 

on all arms small, numerous, biserial, widely separated. 

Deep web connects dorsal arms and Arms I and II. 

Remarks Thomas (1977) reviewed the genus Tremoctopus and described the development of the two 

recognized species, T. violaceus and T. gelatus, both of 

which are found in the western North Atlantic Ocean. 

Of these species, T. violaceus is by far the most common, 

and our 10 specimens have the characteristics of T. violaceus. Hochberg et al. (1992) provide illustrations of all 

growth stages and references to detailed descriptions. 

Family Argonautidae 

Argonauta argo Linnaeus, 1758/Argonauta hians 

Lightfoot, 1786 

Voucher specimens 1.9 mm ML (juvenile female), 

WIE7410, sta. 89.1, USNM 730515; 1.5 mm ML (?mature male), ALB7106, sta. 33, USNM 730516; 13 mm ML 

(immature female), CI7802, sta. BRI2, USNM 816869. 

Species distribution Argonauts are surface dwellers 

that are cosmopolitan in tropical and subtropical waters (Roper and Young, 1975). The specimens collected 

in these plankton surveys included adult females with 

“shells” (= egg cases), belonging to two species, A. argo 

and A. hians. Most of the 13 specimens, though, were 

either males or immature females without shells and 

thus could not be identified to species. Argonauts 

were found during all seasons and in all areas, except 

from the northernmost cruise off New England and 

Nova Scotia. Specimens from surface samples were 

limited to a relatively narrow range of temperatures 

(18.5–22.3°C) but were found across a broad range of 

salinities (S=33.1–36.7 × 10–3). The low-salinity captures 

occurred on the continental shelf (minimum bottom 

depth 65 m) during September 1976. 

Morphology of voucher specimens 1.9 mm ML 

(juvenile female) (Fig. 35A–B): General appearance 

very similar to paralarval Octopus-type octopodid. Mantle 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 41 

Figure 31 

Octopus sp.-type (all arms subequal in length). A, ventral view, 7.4 mm ML, USNM 816866, 

R/V Albatross IV sta. 7404-18; B, oral view, same specimen. 

Figure 32 

“Macrotritopus”-type (arms III greatly elongated and enlarged). A, ventral view, 2.4 mm ML, 

USNM 816867, R/V Delaware II sta. 7219-129; B, oral view, same specimen. 

short, very broad, widest at anterior end. Head short, of suckers, probably a condition as precursor to the 

wider than mantle. Eyes large, protrude laterally, cov- shell-secreting modifications of Arms I of female (“shell 

ered with silvery reflective tissue. Funnel moderately web”). Arms all connected by shallow web. 

large, extends to anterior level of eyes, tip not yet free. 1.5 mm ML (male, possibly mature) (Fig. 35C): Much 

Arms very short, with eight to nine suckers each, bi- like immature specimen described above but without 

serial. Arms I longest, distal elongations are devoid naked elongations to Arms I and with left Arm III hecto-
42 NOAA Technical Report NMFS 152 

Figure 33 

“Macrotritopus”-type (arms III greatly elongated and enlarged). A, dorsal view, 6.6 mm ML, 

USNM 730507, R/V Wieczno sta. 7410-94. 

cotylized and contained in a large sac (right Arm III was 

hectocotylized on one specimen). All arms and hectocotylus sac connected by web. 

13 mm ML (immature female) (Fig. 36): Mantle very 

muscular, conical, very broad at anterior margin; attaches to head at postero-dorsal border of eyes. Funnel 

very large, muscular, extends to base of Arms IV; funnel tip free, but membrane from dorsal edge of Arms 

IV connects to funnel just posterior to the tip. Eyes 

very large, bulge laterally. All outer surfaces of mantle, 

funnel, head, and arms are covered with numerous 

dense, small chromatophores. Arms II–IV long, slender, subequal in length; each has a single small basal 

sucker followed by biserial suckers, very numerous, 

closely-packed, the proximal three to four pairs particularly large; suckers toward the distal tip minute and 

very thickly set. Arms I very long, thick, much longer 

than others, but much contorted and twisted in preservation due to strong contraction of “shell web.” Basal 

one to two suckers small, next four pairs noticeably enlarged, following pairs with reduced diameters. Suckers 

extremely minute distally with longitudinal series widely separated; distal tip devoid of suckers, a ridge-like 

supporting structure for “shell web.” “Shell web” begins 

as narrow membrane at base of Arms I, broadens noticeably at level where suckers become small (about the 

fifth to sixth pair), then becomes very expanded in distal one third of arm (full extent and dimension cannot 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 43 

Figure 34 

Tremoctopus violaceus Chiaie, 1830. A, dorsal view, 2.3 mm ML, USNM 730508, R/V Wieczno sta. 7410-94; B, oral view, same specimen. 

be described because “shell webs” are very tightly contracted and covered with dense chromatophores). Ovaries developing, ova extremely minute. Viscero-pericardial membrane covered with dark chromatophores. 

Remarks Argonauts are sexually dimorphic and are 

the only octopod species of which mature adults are 

commonly collected in plankton samples. One of the 

most distinctive features of Argonauta paralarvae is the 

presence of a “pit-like” locking apparatus on the funnel. 

Additional descriptions and illustrations are available 

in Hochberg et al. (1992), Naef (1928), and Grimpe 

(1928). 

Distributional patterns 

A conceptual framework for consideration of the distribution of micronekton, such as young cephalopods, 

was developed by Haury et al. (1976). They proposed 

scales of temporal and spatial variability appropriate 

for various size categories of pelagic organisms. They 

suggested that the range of spatial (geographical or 

linear) scales important for micronekton was roughly 

hundreds of meters to hundreds of kilometers, whereas 

the important temporal scales ranged from hours to 

tens of thousands of years. Our data allow us to examine 

the distribution of young cephalopods within this range 

of spatial scales and on a temporal scale of a few hours 

to several years. 

Replication of zooplankton collections by repeated, 

consecutive tows actually represents an interaction of 

short-term temporal and small-scale spatial variability. 

Although several attempts at such replication were 

made during the BLM and MARMAP programs, only 

three of these “replicate” data sets collected cephalopods. Thus, we have these data only for the most 
44 NOAA Technical Report NMFS 152 

Figure 35 

Argonauta argo Linnaeus, 1758/Argonauta hians Lightfoot, 1786. A, dorsal view, 1.9 mm ML (juvenile female), 

USNM 730515, R/V Wieczno sta. 7410-89.1; B, oral view, 

same specimen; C, oral view, 1.5 mm ML (?mature 

male), USNM 730516, R/V Albatross IV sta. 7106-33. 

abundant taxon, the loliginids. The high variance in 

abundance of loliginids (relative to mean abundance) 

among these “replicates” highlights a serious unresolved 

problem in field studies of the early life histories of 

cephalopods. Okutani and McGowan (1969) and Bower et al. (1999) found similarly high variability among 

consecutive tows. If, as seems likely, such variability represents spatial patchiness, sampling should be designed 

around either more numerous tows of shorter duration 

to sample smaller volumes of water, in order to resolve 

small-scale variability, or long-duration tows to filter extremely large volumes of water, in order to eliminate 

small-scale variability. However, even the most abundant species seem under represented in samples of 

any size when compared with ecologically similar fishes 

and shrimps. An investigator consequently must be prepared to examine many small-volume samples that do 

not contain cephalopods in order to resolve small-scale 

variability. In any case, this variability means that, regardless of the scale being considered, a large number 

of samples must be included in any discussion of the details of the distribution of paralarval cephalopods. Such 

details were not a goal of this paper. 

In contrast to the distinct pattern of diel variability 

which Vecchione (1981) found for Loligo pealeii in surface samples, time series of oblique subsurface tows 

showed no consistent diel patterns, either for L. pealeii 

(Fig. 5) or for enoploteuthids and ommastrephids (Fig. 

37). The patterns in Fig. 37 were tested for periodicity 

using the nonparametric Runs Test (Siegel, 1956) and 

the null hypothesis of randomness could not be refuted. Random variability in the time series of oblique 

samples does not fit the assumptions of diel vertical migration to and from the surface. This suggests that diel 

changes in the inferred abundance of juvenile cephalopods in surface samples (e.g., Okutani, 1968) may actually result from changes in visual avoidance of the 

sampler, rather than from vertical migration by these 

very young stages. The determination of when a species begins diel vertical migration awaits analyses based 

on high-resolution sets of discrete-depth samples and 

most likely will vary from species to species (e.g., Shea, 

1995). 

In spite of these problems with small-scale and shortterm variability, the large size of the data set that we 

have assembled allows us to look for seasonal and interannual temporal patterns as well as mesoscale to 

large-scale spatial patterns. Both winter and summer 

samples are available from the southern Sargasso Sea. 

Two years of data from quarterly cruises over the continental shelf and slope of the middle U.S. Atlantic states 

are available, as well as from six other shelf and slope 

cruises that cover a period of six years. The shelf data 

extend from Nova Scotia to Cape Canaveral. Furthermore, the winter oceanic samples extend from the Bahamas nearly to Newfoundland. Thus, despite some serious sampling problems, such as a lack of completely 

standardized sampling methods, we feel that the coverage is adequate to elucidate some patterns. 

Many of the taxa for which we have the most data 

were collected throughout the year (Table 2). The taxa 

that appeared to have distinct seasonal distributions 

probably reproduce seasonally and included most of 

the neritic species (e.g., the loliginids collected north 

of Cape Hatteras) and, surprisingly, the young of two 

meso-bathypelagic cranchiids. Taxa with uncertain seasonality included those that were collected on only one 

cruise (e.g., Mastigoteuthis hjorti) and those with the 

most difficult taxonomic problems (such as the loliginids collected south of Cape Hatteras). 

The BLM cruises collected 25 taxa from a total of 44 

taxa for the entire study. Neritic taxa, such as Loligo pealeii, Illex illecebrosus, and Semirossia tenera, were taken with 

approximately the same seasonal patterns during both 

years. However, interannual differences in the oceanic 

taxa collected were extreme (Table 3). Only Abraliopsis 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 45 

Figure 36 

Argonauta argo Linnaeus, 1758/Argonauta hians Lightfoot, 1786. A, ventral view, 13 mm ML (immature female), USNM 

816869, R/V Columbus Islen sta. 7802-BRI2; B, oral view, same specimen. 

pfefferi and Pterygioteuthis sp. were caught 

during both years. Therefore, although 

ample evidence existed that cross-shelf 

intrusions occurred throughout both 

years (Vecchione and Grant, 1983), either the chances of cross-shelf transport 

varied for most cephalopod species or 

the sampling program failed to collect 

many species at one time or another. 

Many more taxa were collected during 

the expanded sampling of the second 

year, but a few taxa were found only during the first year. 

Cross-shelf transport is determined 

largely by the dynamics of the Gulf 

Stream which vary from year to year. Further, meanders, eddies, and rings of the 

Gulf Stream are associated with the formation of patchy environments that can 

be important to distribution and growth 

of species in the western North Atlantic, 

young Illex for example. Because mesoFigure 37 

Time series of oblique subsurface tows in October cruise for captures of 

Enoploteuthidae and Ommastrephidae. Note absence of consistent diel 

patterns. 

scale distribution is so patchy, macro-scale distribution 

sometimes might be masked in cruises with a grid sampling design, since meso-scale spatial conditions might 

change during the course of the sampling. We interpret 

the interannual variability as further evidence that extensive sampling is necessary to estimate distribution of 

young cephalopods, especially of rarely caught species. 

The dominant features that control faunal distributions in the western North Atlantic Ocean are the continental shelf and the Gulf Stream. Mesoscale spatial vari-
46 NOAA Technical Report NMFS 152 

Table 3 

Seasonal and interannual differences in BLM collections. 

Fall Winter Spring Summer 

Taxon ’75 ’76 ’76 ’77 ’ 76 May ’77 ’76 ’77 

Unidentified octopod X X X 

“Macrotritopus” X 

Argonauta argo/Argonauta hians X 

Semirossia tenera X X X X 

Stoloteuthis leucoptera X 

Lolliguncula brevis X 

Loligo pealeii X X X X X X 

Illex sp. X 

Illex illecebrosus X X X X 

rhynchoteuthion Type A′ X 

rhynchoteuthion Type B′ X 

unidentified rhynchoteuthion X 

oegopsid sp. A X X 

Abraliopsis cf. pfefferi X X 

Abralia cf. veranyi X 

Pterygioteuthis sp. X X X 

Ancistrocheirus lesueurii X X X 

Histioteuthis spp. X 

Brachioteuthis sp. X 

Octopoteuthis sp. X 

?Discoteuthis sp. X 

Gonatus fabricii X 

unidentified oegopsid X 

Total taxa per cruise 3 4 2 5 3 6 5 14 

Total taxa per season 6 7 6 16 

Table 2 

Patterns of seasonal distribution. 

Taxa collected throughout the year: 

Semirossia tenera 

Abralia cf. veranyi 

Abraliopsis cf. pfefferi 

Pyroteuthis margaritifera 

Ptyerygioteuthis sp. 

Octopoteuthis sp. 

Onychoteuthis cf. banksii 

Onykia carriboea 

Histioteuthis spp. 

Chtenopteryx sicula 

Brachioteuthis sp. 

Illex sp. 

rhynchoteuthion Type B′ 

Chiroteuthis sp. 

unidentified octopods 

Tremoctopus violaceus 

Argonauta spp. 

Taxa collected only in winter: 

Stoloteuthis leucoptera 

Leachia lemur 

Teuthowenia megalops 

Taxa not collected in winter: 

Lolliguncula brevis 

Loligo pealeii 

Ancistrocheirus lesueurii 

Discoteuthis? sp. 

Helicocranchia cf. papillata 

“Macrotritopus” 

Taxa of uncertain seasonality: 

?Spirula spirula 

unidentified Loliginidae 

Selenoteuthis scintillans 

Gonatus fabricii 

Bathyteuthis abyssicola 

Mastigoteuthis hjorti 

?Taonius/Galiteuthis sp. 

?Bathythauma lyromma 

Nov Nov Feb Mar Jun Sep Aug 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 47 

Table 4 

Patterns of geographical distribution. 

Taxa found throughout the western North Atlantic: 

Abralia cf. veranyi 

Abraliopsis cf. pfefferi 

Pyroteuthis margaritifera (see comments in text) 

Pterygioteuthis sp. 

Octopoteuthis sp. 

Onychoteuthis cf. banksii 

Onykia carriboea 

Histioteuthis spp. 

Chtenopteryx sicula 

Brachioteuthis sp. 

Leachia lemur 

unidentified octopods 

Taxa limited to southern portion of study area: 

Pyroteuthis margaritifera (see comments in text) 

Chiroteuthis sp. 

Helicocranchia cf. papillata 

Tremoctopus violaceus 

Taxa limited to southern and middle portions of study area: 

Argonauta spp. 

Taxa limited to northern and middle portions of study area: 

Illex illecebrosus 

Teuthowenia megalops 

Gonatus sp. 

Note–no taxa were found only in northern portion of study 

area. 

Taxa of uncertain distribution: 

Spirula spirula 

Semirossia tenera 

Stoloteuthis leucoptera 

Loliginidae (including Loligo spp.) 

Selenoteuthis scintillans 

Ancistrocheirus lesueurii 

?Discoteuthis sp. 

Bathyteuthis abyssicola 

Ommastrephidae 

Mastigoteuthis hjorti 

Bathothauma lyromma 

“Macrotritopus” 

ability in the distribution of planktonic cephalopods 

was greatest perpendicular to these features rather than 

latitudinally (Table 4). This contrasts with the situation 

in the eastern North Atlantic (Lu and Clarke, 1975) 

where latitudinal variability in cephalopod distribution 

was great. Latitudinal variability would not be expected 

to be so great in the western Atlantic, because the entire Gulf Stream system acts as a strong dispersal agent 

for subtropical and temperate offshore neritic and oceanic species. Furthermore, our samples from boreal water were quite limited, as were samples from the southernmost area sampled. Perhaps if this study had been 

extended northward (and even southward into the Caribbean Sea), distributional variability associated with 

latitude might become a more important factor. 

Across-shelf transport of paralarvae of tropical, oceanic species probably is limited by the maximum depth 

at which these species occur in the water column. Zooplankton that migrate vertically to depths greater than 

the bottom depth on the shelf typically are excluded 

when Gulf Stream eddies transport the oceanic plankton community onto the shelf (Vecchione and Grant, 

1983). Species that are strictly neustonic also are usually excluded, because the direction of the surface flow 

of shelf water is predominantly offshore (Vecchione, 

1979b). Therefore, in our study area, the planktonic 

cephalopods can be separated into two distinct groups 

based on the depth of the water column (Fig. 38). One 

group is confined to water with depths greater than 

Figure 38 

Cumulative number of planktonic cephalopod taxa 

collected relative to increasing bottom depth. Two distinct groups were found, separated at the shelf/slope 

interface. 
48 NOAA Technical Report NMFS 152 

Figure 39 

Kaiyo-Maru cruise 8201. Station locations, surface and 200 m isotherms, and 

the occurrence of the five most abundant cephalopod taxa. See Fig. 1 for 

specific location. 

350 m (i.e., offshore of the shelf break, which in the 

western North Atlantic occurs at about 200 m depth); 

this group includes neustonic species such as Onykia 

carriboea and species that probably are deep in the water 

column, at least during part of the day (e.g., Histioteuthis spp., Chiroteuthis sp., and Bathyteuthis abyssicola). The 

second group is found over shallower bottom depths 

and includes the species that spawn on the continental 

shelf (e.g., the loliginids and Semirossia tenera) and those 

species that probably spend most of their time at intermediate depths of 50–200 m (e.g., ommastrephids and 

the enoploteuthid group of families) and consequently 

are subject to transport onto the shelf. 

The other pattern of spatial segregation indicated by 

the data was the distribution of species across the Gulf 

Stream. Many species seemed to be associated in one 

way or another with this feature (Figs. 39 and 40). Species such as Illex spp. are found predominantly along 

the shoreward edge of the Gulf Stream in the strong 

thermal gradient between slope water and the Gulf 

Stream, whereas rhynchoteuthion Type A′ and Leachia 

lemur often are caught along the offshore edge of the 

Gulf Stream in the more poorly defined interface with 

Sargasso Sea water (Vecchione and Roper, 1986). 

Strong vertical gradients exist in physical parameters 

such as temperature, salinity, ambient light, and dissolved oxygen, both on the continental shelf and offshore. Therefore, discrete-depth sampling is required to 

delimit the dimensions of a species’ niche along these 

gradients. Our only discrete-depth data are from the 

surface tows, for which temperature and salinity were 

measured concurrently. These data indicate broad overlapping ranges of temperature and salinity for many 

species captured in surface waters (Fig. 41). Surface 

tows may have sampled the extreme end of the range. 

Or, vertical migration may have caused some individuals to cross water-mass boundaries into the surface layer 

where conditions were not within the preferred ranges 

for the species. For example, most Illex were collected 

in subsurface samples, but the few that were collected 

at the surface indicated a tolerance for very broad ranges of temperature and salinity. Probably temperature 

and salinity were much less variable at the depths at 

which most Illex were caught. Similar explanations may 

be true for the other taxa presented in Fig. 41. 

A comparison of the results of this study with other 

studies of cephalopod species distributions is difficult 

because of the taxonomic problems discussed above 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 49 

and inherent to varying degrees in virtually all studies. 

However, comparisons of relative abundance at the family level can be made. We used Spearman’s nonparametric correlation coefficient (Siegel, 1956) to compare 

the rank-order of family abundance of our data with 

those of other studies of planktonic cephalopods and 

of trawling studies from the North Atlantic Ocean. The 

results were as follows: 

Study Type Location Spearman’s r 

Sanchez and Moli, 

1985 plankton Mediterranean .754 

Guerra et al., 1985 plankton E Atlantic .733 

Cairns, 1976 trawling NW Atlantic .732 

Okutani and 

McGowan, 1969 plankton E Pacific .640 

Dawe and Stephen, 

1988 plankton N Atlantic .601 

Dawe and Stephen, 

1988 trawling N Atlantic .528 

Okutani, 1968 plankton W Pacific .523 

Yamamoto and 

Okutani, 1975 plankton W Pacific .511 

Lu and Roper, 1979 trawling NW Atlantic .503 

Roper, 1977 trawling NW Atlantic .430 

Clarke, 1977 trawling NE Atlantic .351 

Kubodera and 

Jefferts, 1984 plankton N Pacific .327 

Thus, the rank-order of family abundance that we found 

was most similar to that of the study of planktonic cephalopods off the Mediterranean coast of Spain by Sanchez and Moli (1985). Our results were least similar to 

those of Kubodera and Jefferts (1984) from the northern North Pacific Ocean, where the planktonic cephalopods are very strongly dominated by gonatids. The 

studies by Guerra et al. (1985) and Cairns (1976) in 

the eastern and western North Atlantic Ocean, respectively, show a close similarity to our results. Interestingly, though, our results were more similar to those 

of the plankton studies from the Pacific Ocean (Okutani, 1968; Okutani and McGowan, 1969; Yamamoto 

and Okutani, 1975) than to several other trawling studies from the North Atlantic Ocean. One conclusion is 

that it seems there are some families of cephalopods 

for which plankton net methods sample quite differently from trawling methods. In fact, different midwater trawling techniques provide different rank orders 

of captures within the same geographical region, as 

pointed out in comparative studies by Roper (1977) 

and Wormuth and Roper (1983). Clearly, additional 

carefully structured comparative studies are necessary 

in order to determine the most effective techniques to 

sample different taxonomic groups and different life 

history stages. 

Octopoteuthids and chtenopterygids, for instance, are 

rare in trawl collections but comparatively common in 

Figure 40 

Columbus Islen cruise 7802. Station locations, surface 

isotherms, and the occurrence of the seven most abundant cephalopod taxa. See Fig. 1 for specific location. 

plankton samples. The frequency of octopoteuthid remains taken in sperm whale stomachs has been compared with their scarcity in trawl samples to emphasize 

the difficulty in sampling for oceanic cephalopods 

(Clarke, 1977). For species in families that are relatively 

common in plankton samples, early-life-history studies, 

similar to the familiar ichthyoplankton surveys of fisheries science, may be the most reliable method of gathering data on distribution and abundance. An example of 

the utility of such methods is Kubodera and Okutani’s 

(1981) study of Pacific gonatids. 

The enoploteuthid group of families and onychoteuthids rank among the 10 most abundant families in 

all of these studies, using both trawling and plankton 

methods. Furthermore, Young et al. (1985) showed 

that it is possible to collect and to identify enoploteuthid eggs. Consequently, it may be possible to sample 

enoploteuthid species adequately throughout all their 

developmental stages using different selective sampling 
50 NOAA Technical Report NMFS 152 

Figure 41 

Median and range of surface temperature and surface salinity for captures of cephalopod taxa in neuston 

samples. 

techniques. These species should be considered as likely candidates for studies of the population dynamics of 

oceanic squids (e.g., Bigelow, 1991). 

Acknowledgments 

We wish to acknowledge and thank those who provided 

the specimens and associated data upon which this 

study was based. Kenneth Sherman, National Marine 

Fisheries Service, provided the MARMAP material and 

George Grant, Virginia Institute of Marine Science, 

supplied the specimens from projects funded by the 

Bureau of Land Management, National Science Foundation, and the Commonwealth of Virginia. Funding 

and laboratory space were provided by the Smithsonian 

Marine Station, Ft. Pierce, FL, Mary Rice, Director, for 

part of the work, and this paper comprises SMS Contribution number 329. 

The illustrations were made by Molly Ryan and 

Marilyn Schotte, Department of Invertebrate Zoology, 

NMNH, and by Ann E. Lacy, under contract. We appreciate their work. 

The manuscript received much-appreciated pre-submission reviews from R. E. Young, University of Hawaii, 

and U. Piatkowski, Institut fur Meereskunde, Kiel, Germany. We also thank three anonymous reviewers for 

their thorough reviews and comments. 
Vecchione et al.: Distribution, Relative Abundance and Developmental Morphology of Paralarval Cephalopods 51 

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