09

Cephalopod classification and

Taxonomy

V. Venkatesan and K.S. Mohamed

Molluscan Fisheries Division, Central Marine Fisheries Research Institute, Kochi-682 018

Introduction have proved to be useful based on the morphological studies

Chambered nautilus, cuttlefishes, squids and octopus are (Voss, 1988), but less significant in determining higher-level
the four major groups of cephalopods, which belong to the relationships. The following classification can be used until an
highly evolved class of phylum Mollusca. Cephalopods are the alternative arrangement can be derived from cladistic analysis.
third largest molluscan class after bivalves and gastropods Molecular or morphological based analysis must have to go
and consist of more than 800 species (Lindgren et al. 2004). considerable scrutiny before changed in classification should
The fossil record contains about 17,000 named species of be adopted.
cephalopods. Although the diversity of cephalopods is very
much reduced in the modern era, cephalopods are found to Cephalopods (Class Cephalopoda) are represented by two
occur in all the oceans of the world from the tropics to the extant subclasses, Nautiloidea (Nautilus and Allonautilus) and
polar seas and at all depths ranging from the surface to below Coleoidea and one extinct subclass, Ammonoidea. Members
5000m. Cephalopods were dominant predators millions of the subclass Coleoidea includes two subdivisions, the
of years before fish appeared. The earliest cephalopods Belemnoidea, which is the primitive form of cephalopod
were primitive shelled nautiloids which evolved in the Late possessing ink sac and ten equally sized arms, became extinct
Cambrian period. The living cephalopods range in size from during the cretaceous period and Neocoleoidea (cuttlefish,
25mm (Southern pygmy squid, Idiosepius notoides) to more squid and octopus) where the shell has been internalized
than 12m (Colossal squid, Mesonychoteuthis hamiltoni) in and reduced, completely lost. As a consequence, members
length. of Neocoleoidea rarely fossilize and very few information
pertaining to the origin and relationships of living coleoid
All cephalopods are dioecious and females are generally bigger cephalopods is available from the fossil record. The major
than males. Males have one or two modified arms known as division of Coleoidea is based upon the number of arms or
hectocotylus which are used for mating. Fertilization takes tentacles and their structure.
place in the female. Development is direct to the young ones
as miniature of adult. Many species of deep-sea cephalopods Presently, living coleoids can be segregated into two
occurring at depths of about 400 - 800m undergo vertical superorders, Decapodiformes and Octopodiformes (Berthold
migrations during the day and then rise into the uppermost and Engeser, 1987). The Decapodiformes has fourth arm
200 m or so during the night. Cephalopods are carnivores pair modified into long tentacles. The Decapodiformes
and possess a radula and powerful beaks. They eat fish, contains two orders; the order Teuthoidea, which includes
crustaceans, shellfish etc. They are major food resources for two suborders [ Myopsida(closed-eye squids) and Oeopsida
many top predators such as dolphins, whales, seals, birds and (open-eye squids)] and the order Sepioidea which includes
large fish. families like Idiosepiidae (pygmy squid), Sepiidae (cuttlefish),
Sepiolidae (bobtail squids), Spirulidae (ram’s horn squid),
Classification and Sepiadariidae (bottletail squids). The Octopodiformes
Higher-level classification of recent cephalopods is mostly includes the orders Octopoda (pelagic and benthic octopuses)
unstable. Several authors have suggested highly varying and Vampyromorpha (vamphire squid). Octopodiformes has
classification. A conservative arrangement should be modifications to second arm pair; it is drastically reduced as
accepted that do not differ much from that of Naef (1921 - a sensory filament in the Vampyromorphida, while Octopoda
23). Classification of species within subfamilies and /or genera species have totally lost that arm pair. The Octopoda

49
Cephalopod classification and Taxonomy

contains two suborders; Cirrata (deep-sea finned octopuses) Suborder Oegopsida

and Incirrata (pelagic and benthic octopuses including the
Family Onychoteuthidae
argonautiods and blanket octopuses)
Genus Onchoteuthis Onchoteuthis banksii
Squids posses elongate, cigar-shaped body with Family Ommastrephidae
posteroexternal fins and eight cirumoral arms, not joined Subfamily Ommastrephinae
at the base with a web, with two or more rows of stalked Genus Sthenoteuthis Sthenoteuthis oualaniensis
suckers bearing chitinous rings (and/or hooks) extend the
Family Thysanoteuthidae
length of the mantle. They also posses two long tentacles
with tentacular club of two or more rows of suckers (and or Genus Thysanoteuthis Thysanoteuthis rhombus
hooks) at the distal end. The cuttlefish posses a broad sac- Order SEPIODIDAE
like body with lateral fins that are either narrow and running Family Sepiidae
over the entire length (Sepiidae) or are short, round and flap-
Genus Sepia Sepia pharaonis
like (Sepiolidae). In both cases the posterior ends of the fins
are free (subterminal) and separated by the posterior end of Sepia aculeata
the mantle; ten circumoral appendages, the longest (fourth) Sepia prashadi
pair of tentacles are retractile into pockets at the ventrolateral Sepia elliptica
sides of head. The eight arms frequently bear four rows of
Sepia trygonina
stalked suckers with chitinous rings. Both eyes are covered
with a transparent membrane; shell is thick, chalky, calcareous Sepia brevimana
(cuttlebone of sepia) or thin, chitinous (Sepiolidae). Octopus Sepia arabica
posses a short, sac-like body with either no lateral fins or with Sepia kobiensis
separate paddle-like fins in some deep sea forms, and eight
Sepia prabahari
circumoral arms with no tentacles, with the bases connected
by a web and un-stalked suckers, without chitinous rings, Sepia ramani
along the length of the arms. Sepia omani
Genus Sepiella Sepiella inermis
About 210 species of cephalopods have been reported Family Sepiolidae
from India. Among these, there are about 80 species of
Genus Euprymna Euprymna stenodactyla
cephalopods of commercial and scientific interest distributed
in the Indian seas. Order OCTOPODA
Suborder Incirrata
Systematic position of potentially important cephalopods of Family Octopodidae
India
Genus Amphioctopus Amphioctopus aegina
Amphioctopus neglectus
Class CEPHALOPODA
Amphioctopus marginatus
Sub class NAUTILOIDEA
Amphioctopus rex
Family Nautilidae Nautilus pompilius
Genus Cistopus Cistopus indicus
Subclass COLEOIDEA
Cistopus taiwanicus
Order TEUTHOIDEA
Genus Haplochlaena Haplochlaena maculosa
Suborder Myopsida
Genus Callistoctopus Callistoctopus luteus
Family Loliginidae
Genus Octopus Octopus vulgaris
Genus Uroteuthis Uroteuthis (Photololigo)
duvaucelii Genus Pteroctopus Pteroctopus keralensis
U (P) sibogae Family Argonautidae
U (P) singhalensis Genus Argonauta Argonauta hians
U (P) edulis Argonauta argo
U (P) chinensis Subclass Nautiloidea
Genus Sepioteuthis Sepioteuthis lessoniana
Genus Loliolus Loliolus (Loliolus) hardwickei Shell complete external, smooth, coiled and chambered, more
Loliolus (Nipponololigo) uyii than 10 (63 - 94) circumoral appendages without suckers, a
funnel bilobed, two pairs of gills and the absence of an ink sac.
L (N) sumatrensis

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 V. Venkatesan and K.S. Mohamed

Family Nautilidae U (P) sibogae (Adam, 1954)

The ‘’chambered or pearly nautiluses ‘’ comprises single family Mantle long, narrow and slender, no ridge but chromatophore
and genus and six species. They have approximately 100 sucker- concentration ventrally along midline; fins narrow and less
less tentacles, simple eye without lenses and thick rigid hood than 60 per cent of mantle length; less than half of left ventral
used to protect the animal when retracted within the shell. arm hectocotylized distally in males; gladius narrow, sharply
accumulate posteriorly.
Subclass Coleoidea
U (P) singhalensis (Ortmann, 1891)
This subclass includes all living cephalopods - squids,
cuttlefish and octopuses, other than chambered nautiluses. Mantle is long, slender, cylindrical, and it tapers posteriorly
Key diagnostic characters are shell internal, calcareous, into as sharply-pointed tip. Mantle bout 4-7 times as long
chitinous or cartilaginous, 8-10 circumoral appendages as wide. Mantle with a ridge along midline in males; The
with suckers, only one pair of gills (dibranchiate) and funnel tentacles are short and slender. Clubs are rather short. Left
tube-like. ventral arm IV is hectocotylized distally in mature males for
40 - 45% of its length. The chitinous sucker rings are smooth
Order Teuthoidea or wavy proximally, while the distal margin bears 6-11 (most
This order contains the squids, characterized by internal commonly 9) plate-like, truncate, squared teeth.
shell (gladius or pen) chitinous feather or rod shaped,
U (P) edulis (Hoyle, 1885)
eight arms; two contractile but not retractile, pocket
absent, tentacles lost secondarily in some, fin on the Mantle more or less stout, elongate and slender. Fins large,
mantle and stalked suckers with or without chitinous rhombic with the anterior margin slightly convex, the posterior
hooks, with horny rings and constricted necks; fin lobes margin gently concave and the lateral angles rounded. Fins
fused posteriorly. Eyes either covered or open and without become slightly longer than wide in adult specimens (up to
supplementary eyelid. 70% of mantle length), gladius long, somewhat narrow, arms
somewhat long (25- 45% of mantle). More than half of left
Suborder Myopsida ventral arm hectocotylized distally in males.
Myopsid squids are characterized by eyes entirely covered by
U (P) chinensis Gray, 1849
a transparent corneal membrane. Eye cavity communicates
with the exterior through a tiny hole. Arms and tentacles have Fin length in adults greater than 60% of mantle length.
suckers only, no hooks. Mante locking apparatus is simple Hectocotylized portion of the left arm IV from 33% to 50% of
(linear) and the gladius is pen-like. total arm length. Arm sucker rings with 10-15 stout, pointed,
conical teeth distally, the proximal margin smooth; occasionally
Suborder Oegopsida with rudimentary teeth only. Although the record of this species
Oegopsid squids (Oceanic squid or Open-eyed squids) are along the Indian east coast is available in the literature, this
characterized by eyes not covered with a corneal membrane species is not recorded in the cephalopod samples of Institute.
and open to the surrounding medium, arms and tentacles
Loliolus (Loliolus) hardwickei (Gray, 1849)
bear suckers and / or hooks. Mantle locking apparatus ranges
from simple to complex to fused. Small squids. Mantle length of adults less than 60 mm;
fins heart shaped; vane of gladius conspicuously broad at
Family Loliginidae midlength
Sepioteuthis lessoniana Ferussac in Loliolus (Nipponololigo) uyii (Wakiya and
Lesson, 1831 Ishikawa, 1921)
Body elongate, cylindrical in outline; fins marginal, wide and Body short and stout; mid rib of gladius clearly visible through
muscular, very long almost running along entire length of dorsal mantle skin as a median dark line; fins 55-65 per cent
mantle; elliptical in shape of mantle length; Tentacular clubs have median manal suckers
with smooth rings; in males left ventral arm hectocotylized
Uroteuthis (Photololigo) duvaucelii
almost the entire arm; papillae on ventral margin fused with
(Orbigny, 1835)
membrane.
Body elongate, mid-rib of gladius clearly visible through
L (N) sumatrensis (D’Orbigny, 1835)
mantle skin; fin length in adults upto 60 per cent of mantle
length; tentacular clubs large median manal sucker ring with Body short, sub-cylindrical, gradually decrease in width
14 - 17 teeth; Arm sucker rings with broad, large, square posteriorly to blunt point. Head small with large eyes; fins
teeth (5 to 9) on the distal margin; in males, more than half 60-65% of mantle length; fin rhomboidal in shape; arm sucker
the length (up to75 %) of the left ventral arm hectocotylized, ring with 6-9 broad, squared teeth; in male left ventral arm
papillae not fused. hectocotylized upto 87%.

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Cephalopod classification and Taxonomy

Onychoteuthidae Sepia pharaonis Ehrenberg, 1831

Onchoteuthis banksii (Leach, 1817)
Body robust, fins broad commencing from edge of anterior
Oceanic squids with muscular body; head with nuchal folds mantle margin; tentacular clubs moderately long and well
on the dorsal side at posterior end; rachis of gladius visible expanded; 5 or 6 suckers in middle row of manus greatly
as a longitudinal ridge middorsally along the entire length of enlarged; cuttlebone broad, thick and with a midventral
mantle; tentacular clubs with two rows of hooks, marginal flattening anteriorly in striated area; striae ‘ ’ shaped; inner
suckers lacking. cone forms a conspicuous yellow flat ledge; a sharp thick
spine present; when alive, body brownish, tiger-stripe pattern
Thysanoteuthidae prominent.
Thysanoteuthis rhombus Troschel, 1857 Sepia aculeata Van Hasselt, 1835 ( in
Ferussac and d’ Orbigny, 1834 - 1848)
Funnel locking cartilage shaped consisting of a narrow
longitudinal groove and a short transverse groove branching Tentacular clubs very long, with 10-14 rows of minute sub-
from it medially. Fins broad and rhombus-shaped occupying equal suckers. Cuttlebone broad and thick with a median
nearly entire length of mantle. longitudinal edge with a faint groove running medially on
striated area; inner cone forms a ledge-like callosity.
Ommastrephidae
Sepia prashadi Winckworth, 1936
Sthenoteuthis oualaniensis (Lesson, 1830)
Body not robust, fin narrow commencing a few mm behind
Funnel and mantle cartilages of the locking apparatus fused edge of anterior mantle margin; tentacular clubs short,
together. An oval photophoric patch present middorsally near expanded; not more than 3 suckers in middle row of manus
anterior margin of mantle; muscle of mantle ventrally without greatly enlarged; cuttlebone narrow, midventral groove
embedded light organs; two intestinal photophores present. narrow and distinct, striae anteriorly broadly truncate with
lateral corners slightly produced forward; dorsal surface
Order Sepioidea pinkish in colour, a sharp thin spine present; When alive,
This order includes the cuttlefishes, characterized by an oval dusty brownish, transverse stripes less distinct.
body shape, compressed dorsoventrally and framed along
Sepia elliptica Hoyle, 1885
both sides of the body by narrow fins that do not attach at
the posterior end. The arms bear 2 to 4 rows of suckers. The Tentacular clubs moderately long, with 10 rows of small
tentacles are totally retractile into pockets. The internal shell, suckers of uniform size. Cuttlebone thin, elliptical in shape,
cuttlebone (calcareous) lies dorsally in the body below the dorsal surface smooth; two conspicuous lateral ridges more
skin. The shell is an oval in shape, thick, containing several prominent anteriorly resulting in three longitudinal furrows in
gas and /or water filled chambers for buoyancy control. striated area; spine thick, sharp, long and well curved.
Family Sepiidae Sepia trygonina (Rochebrune, 1884)
Small to medium- sized animals characterized by an oval No fleshy projections on head; fins extend upto end of mantle;
body; flattened dorsoventrally, calcareous internal shell, head tentacles with short clubs, suckers in eight rows, about five
free from dorsal mantle, fins marginal and narrow, light organ in third row enlarged. Cuttlebone lanceolite with acuminate
absent. anterior tip with edges of outer cone winged giving an arrow
head appearance; spine small.
Family Sepiolidae
Sepia brevimana Steenstrup, 1875
Small animals characterized by saccular body, wide, round
bottomed; fins circular; internal shell lacking; dorsal mantle Tentacular club short with 6-8 small subequal suckers.
and head united by a nuchal commissure; saddle-shaped light Cuttlebone flat and distinctly acuminate anteriorly, dorsal
organ present on ink sac. surface rugose, a shallow median groove in the striated
area, the striae ‘ ’ shaped with a median shallow groove
Genus Sepia broadening anteriorly; inner cone and its limbs pinkish in
Body without a glandular pore at posterior extremity; colour; spine small, sharp and slightly curved.
cuttlebone mostly with a spine (rostrum) at posterior end.
Onychoteuthidae
Sepiella inermis Van Hasselt, 1835 ( in
Ferussac and d’ Orbigny, 1834 - 1848) Onchoteuthis banksii (Leach, 1817)
Body with a district glandular pore at posterior extremely Oceanic squids with muscular body; head with nuchal folds
on ventral side; with brownish fluid oozing out; cuttlebone on the dorsal side at posterior end; rachis of gladius visible
devoid of spine. as a longitudinal ridge middorsally along the entire length of

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 V. Venkatesan and K.S. Mohamed

mantle; tentacular clubs with two rows of hooks, marginal shell; internal shell either vestigial or lacking; no great
suckers lacking. disparity between males and females in size.
Thysanoteuthidae Cistopus indicus Rapp, 1835 ( in Ferussac
and d’ Orbigny, 1834 - 1848)
Thysanoteuthis rhombus Troschel, 1857
Hectocotylized arm only slightly modified, ligula small about
Funnel locking cartilage shaped consisting of a narrow 3 per cent of arm. Small water pores leading to embedded
longitudinal groove and a short transverse groove branching pouches between bases of arms.
from it medially. Fins broad and rhombus-shaped occupying
Amphioctopus aegina (Gray, 1849)
nearly entire length of mantle.
Eyes prominent; a single large cirrus posterior to each eye.
Ommastrephidae Ligula small, 5 to 8 per cent of arm; with shallow groove;
Sthenoteuthis oualaniensis (Lesson, 1830) penis and diverticulum together form U-shaped loop;
spermatophores long and unarmed.
Funnel and mantle cartilages of the locking apparatus fused
Amphioctopus neglectus (Nateewathana
together. An oval photophoric patch present middorsally near
and Norman, 1999)
anterior margin of mantle; muscle of mantle ventrally without
embedded light organs; two intestinal photophores present. Medium-sized species characterized by elongate and ovoid
body, U-shaped iridescent transverse bar on the head between
Order Octopoda the eyes, Dark ocellus including blue ring present at base of
This order includes all octopuses, described by eight arms 2nd and 3rd arm pair, Head relatively wider in males than in
with 1 or 2 rows of suckers. Most species have web sectors female, 1 or 2 papillae present over each eye. Ligula long and
between the arms. slender.
Sub-order Cirrata Haplochlaena maculosa (Hoyle, 1883)
Finned or cirrate octopods are deep sea octopuses Body globular smaller in size; skin smooth without reticulate
characterized by round to tongue- like fins on the mantle and pattern; white fresh dusty brown in colour with prominent
single rows of suckers interspersed by cirri. Mantle aperture is bluish rings on mantle, head, web and arms.
very narrow. Only the left oviduct is developed
Suggested reading
Sub-order Incirrata Jerep, P and Roper, C.F.E. 2005. Cephalopods of the world. An annotated and
illustrated catalogue of cephalopod species known to date. Volume 1.
Incirrate octopuses are characterize by fins lacking, and have Chambered nautilus, and sepioids. FAO Species Catalogue for Fishery
Purposes. No. 4, Vol. 1. Rome, FAO. 262p.
1 or 2 rows of suckers and no cirri. Jerep, P and Roper, C.F.E. 2010. Cephalopods of the world. An annotated and
illustrated catalogue of cephalopod species known to date. Volume 2.
Family Argonautidae Myopsid and Oegopsid squids. FAO Species Catalogue for Fishery Purposes.
No. 4, Vol. 2 Rome, FAO. 605p.
This family of pelagic octopuses is known as paper nautiluses Jerep, P, Roper, C.F.E., Norman, M.D. and Julian K Finn 2014. Cephalopods of the
or Argonauts, the females of which secrete an external shell. world. An annotated and illustrated catalogue of cephalopod species known
to date. Volume 3. Octopods and Vampire squids. FAO Species Catalogue for
This calcareous external shell is brittle and white in colour Fishery Purposes. No. 4, Vol. 3. Rome, FAO. 370p.
with fine corrugations. The male is much smaller than the Berthold, T., Engeser, T., 1987. Phylogenetic analysis and systematization of the
Cephalopoda (Mollusca). Ver. Naturwissenschaftliche Vereins Hamburg 29,
female. Male lacks the external shell and possesses a large 187-220.
modified third left arm which is detached during mating. Naef, A., 1921-1923. Cephalopoda. Fauna e flora del Golfo di Napoli, Monograph.
In (translated from German by the Israel program for Scientific translations,
Family Octopodidae Jerusalem 1972), p. 917.
Lindgren, A.R., Giribet, G., Nishiguchi, M.K., 2004. A combined approach to the
This family includes tiny to very large benthic octopuses phylogeny of Cephalopoda (Mollusca). Cladistics 20, 454-486.
characterized by eight arms with 1 or 2 rows of sessile suckers Voss, G.L., 1988. Evolution and phylogenetic relationships of deep-sea octopods
(Cirrata and Incirrata). In: Clarke, M.R., Trueman, E.R., (Eds.), The Mollusca.
and modified third right arm in males, without an external Paleontology and Neontology of Cephalopods, vol. 12, pp. 253-276.

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