Mollusc A

Mollusca
Snails, clams, mussels, squids, octopi, chitons, and tusk

shells
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ancestor of the other groups in the tree. This ancestor diversified over time into several
descendent subgroups, which are represented as internal nodes and terminal taxa to the right.
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Containing group: Bilateria
Gastropoda
Caenogastropoda
Pterotracheoidea Rafinesque, 1814
Heteropoda Lamarck, 1812, heteropods, sea elephants
Roger R. Seapy
• Atlantidae Rang, 1829

• Carinariidae Blainville 1818
• Pterotracheidae Rafinesque, 1814
Atlantidae Rang, 1829

Roger R. Seapy
• Atlanta Lesueur, 1817

• Protatlanta souleyeti (Smith 1888)
• Oxygyrus keraudreni Benson 1835
• Carinariidae Blainville 1818

• Roger R. Seapy
•
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•
Pterotracheidae Rafinesque, 1814

Roger R. Seapy
• Firoloida Lesueur 1817

• Pterotrachea Niebuhr (in Forskål) 1775
• Heterobranchia
•
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Cephalopoda Cuvier 1797

Octopods, squids, nautiluses, etc.
Richard E. Young, Michael Vecchione, and Katharina M. Mangold (1922-2003)

pages.
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Containing group: Mollusca
Introduction
The Cephalopoda is an ancient and very successful group of the Mollusca. Cephalopods have
been among the dominant large predators in the ocean at various times in geological history.
Two groups of cephalopods exist today: The Nautiloidea with a few species of the pearly
nautilus, and the Coleoidea, containing the squids, cuttlefishes, octopods and vampire squids,
which is represented by about 700 species. Cephalopods are the most active of the molluscs
and some squids rival fishes in their swimming speed. Although there are relatively few
species of living cephalopods, they occupy a great variety of habitats in all of the world's
oceans. Individual species are often very abundant and provide major targets for marine
fisheries.
Cephalopods first appeared about 500 million years ago in the Upper Cambrian Period.
Although considerable uncertainity still exists, the two extant lineages may have separated
470 mya with the possible origin of the Bactritida or earlier. The long separation of the two
lineages has, today, resulted in lineages with cephalopods that are very different in structure.
Characteristics
1. A funnel derived from the molluscan foot (ref.?) or the molluscan neck
region (Shigeno et al., 2007).
2. Circumoral arms that are derived from the molluscan head (J.Z. Young,
1965) or the molluscan foot (Shigeno et al., 2007).
3. Chitinous beaks.
Figure. Oral view of the buccal mass of the giant squid Architeuthis sp. The dark,
chitinous, bird-like beaks are visible on the surface of the white muscle mass.
Photograph by Mark Norman.
4. Shell.
1. Shell with a phragmocone and siphuncle.

Figure. A longitudinal cut through the shell of the pearly nautilus (Nautilus
sp.)showing the flotation chambers (phragmocone) and the siphuncle (partially
lost from the larger chambers) that passes through the septa separating
individual chambers. The shell has become internal and highly modified in
most living cephalopods. Photograph by R. Young.
5. Image-forming eyes.
Classification
Much of the higher classification of Recent cephalopods is unstable. Various authors have
suggested highly varying arrangements. We adopt a conservative arrangement that does not
differ much from that of Naef (1921-23). Except for the position of the Octopodiformes and
its two orders, we have a questionable phylogenetic basis for accepting this or any other
scheme. We suggest, however, for the sake of stability, that the following classification be
used until this or an alternative arrangement can be derived from cladistic analyses. The
analyses, whether molecular or morphological, however, must be robust and must survive
considerable scrutiny before changes in classification should be adopted.
• Class: Cephalopoda Cuvier, 1797

o Subclass: Nautiloidea Agassiz, 1847





 Fam: Nautilidae Blainville,
1825
o Subclass: Coleoidea Bather, 1888
 Division: Neocoleoidea Haas, 1997
 Superorder: Octopodiformes Berthold and Engeser,
1987
 Order: Vampyromorpha Robson, 1929


Fam: Vampyroteuthidae

Thiele, in Chun, 1915
 Order: Octopoda Leach, 1818
 Suborder: Cirrata Grimpe, 1916

 Fam: Cirroteuthidae
Keferstein, 1866
 Fam: Stauroteuthidae
Grimpe, 1916
 Fam: Opisthoteuthidae
Verrill, 1896
 Suborder: Incirrata Grimpe, 1916

 Fam: Amphitretidae Hoyle,
1886
 Fam: Bolitaenidae Chun,
1911
 Fam: Octopodidae Orbigny,
1839 In: Ferussac and
Orbigny, 1834-1848
 Fam: Vitreledonellidae
Robson, 1932
o




 Superfamily: Argonautoidea Naef,
1912
 Fam: Alloposidae Verrill,
1881
 Fam: Argonautidae
Cantraine, 1841
 Fam: Ocythoidae Gray, 1849
 Fam: Tremoctopodidae
Tryon, 1879
•
o

 Superorder: Decapodiformes Leach, 1817
 Interactive KEY to families of the Decapodiformes
 Order: Oegopsida Orbigny, 1845


 Fam: Architeuthidae Pfeffer,
1900
 Fam: Brachioteuthidae
Pfeffer, 1908
 Chiroteuthid families
 Fam: Batoteuthidae Young
and Roper, 1968
 Fam: Chiroteuthidae Gray,
1849
 Fam: Joubiniteuthidae Naef,
1922
 Fam: Magnapinnidae
Vecchione and Young, 1998
 Fam: Mastigoteuthidae
Verrill, 1881
 Fam: Promachoteuthidae
Naef, 1912
 Fam: Cranchiidae Prosch,

1847
 Fam: Cycloteuthidae Naef,
1923
 Enoploteuthidae families
 Fam: Ancistrocheiridae
Pfeffer, 1912
 Fam: Enoploteuthidae
Pfeffer, 1900
 Fam: Lycoteuthidae Pfeffer,
1908
 Fam: Pyroteuthidae Pfeffer,
1912
 Fam: Gonatidae Hoyle 1886
 Histioteuthid families
 Fam: Histioteuthidae Verrill,
1881
 Fam: Psychroteuthidae
Thiele, 1920
 Lepidoteuthid families
 Fam: Lepidoteuthidae Naef,
1912
 Fam: Octopoteuthidae Berry,
1912
 Fam: Pholidoteuthidae Voss,
1956
 Fam: Neoteuthidae Naef,

1921
 Fam: Ommastrephidae
Steenstrup, 1857
 Fam: Onychoteuthidae Gray,
1847
 Fam: Thysanoteuthidae
Keferstein, 1866
 Order: Myopsida Naef, 1916



 Fam: Australiteuthidae Lu,
2005
 Fam: Loliginidae Lesueur,
1821
 Order: Sepioidea Naef, 1916


 Suborder: Sepiida Keferstein, 1866
 Fam: Sepiidae Keferstein,
1866
 Suborder: Sepiolida Naef, 1916
 Fam: Sepiadariidae Fischer,
1882
 Fam: Sepiolidae Leach, 1817
 Order: Spirulida Haeckel, 1896



 Fam: Spirulidae Owen, 1836
 Order uncertain


 Superfamily: Bathyteuthoidea nov.
 Fam: Bathyteuthidae Pfeffer,
1900
 Fam: Chtenopterygidae
Grimpe, 1922
 Fam: Idiosepiidae Fischer,

1882
Nautiloidea
Nautilidae Blainville 1825

Pearly nautiluses
Richard E. Young
Only these few living species represent this once abundant subclass.
• Nautilus belauensis
• Nautilus macromphalus
• Nautilus pompilius
• Nautilus repertus ?
• Nautilus stenomphalus
• Allonautilus scrobiculatus
Coleoidea Bather, 1888
Octopods, squids, cuttlefishes and their relatives
The Coleoidea contains all living cephalopods except for species of pearly nautilus.
• Belemnoidea
• Neocoleoidea
o Decapodiformes Leach, 1817 (squids, cuttlefishes and their
relatives)
o Octopodiformes Berthold and Engeser, 1987 (vampire squid and
octopods)
Information on FOSSIL neocoleoids can be found under Octopodiformes

Containing group: Cephalopoda
Introduction
The Coleoidea contains two subdivisions. One, the Belemnoidea, had members that were
common inhabitants of neritic waters during the Cretaceous but became extinct and the end
of this Period.

Figure. Left - Side view of a fossil belemnoid "Acanthoteuthis" from Solenhofen, Germany
and in the collection of the U. S. National Museum of Natural History. The equal arms are
lined with hooks as is characteristic of this group. Right - Fossil hooks of Belemnoteuthis (?)
from Lyme Regis, Dorsetshire, England and in the collection of the U. S. National Museum
of Natural History. The shape of the hooks is seen along with the way they may have lined up
on the arm. Photographs by R. Young.
The other subdivision, the Neocoleoidea, contains two extant groups: the Octopodiformes
(octopods and vampire squid) and the Decapodiformes (squids and cuttlefishes). In addition,
a variety of fossil groups exist that may fall into either of these two groups of neocoleoids.
The extant coleoids occupy all major habitats in the ocean from intertidal to great depths
(deepest record is 7279m, Aldred et al., 1983; Voss, 1988,p. 266) and from south to north
polar regions. Although some loliginid squids and some sepiolids can tolerate reduced
salinity, no cephalopod occurs in fresh water. In many regions of the oceans, coleoids are
dominant members of the ecosystem and are often targets of major fisheries.
Coleoids generally use mantle-generated jet propulsion to obtain high swimming speeds. This
is an effective but energetically expensive means of locomotion (Webber and O'Dor, 1985)
and, as a result, some groups utilize the fins as the primary locomotor organ. This is
especially common in deep-sea species (e.g., cirrate octopods, octopoteuthid squid,
magnapinnid squid) where, in dark waters, the value of speed is reduced.
When cephalopods first evolved, the primary feature that separated them from other molluscs
was a shell with buoyancy chambers. This allowed cephalopods to invade the pelagic realm
where predators and competitors were few. When the shell became internal in coleoids the
bouyancy chambers were retained in belemnoids. In most present day neocoleoids, the
buoyant shell has been abandoned (it is retained in Spirula and the Sepiidae) and many
squids and octopods are negatively buoyant. While this is not a problem for those octopods
that are benthic, negatively buoyant squids, however, must swim constantly to stay afloat.
Some squids and pelagic octopods, however, have gained near-neutral buoyancy via the
retention of light ions (especially ammonium in place of sodium) and the reduction of heavy
tissues (Clarke, et al., 1979). This type of neutral buoyancy is found in many deep-sea
species since reduction of muscular and cartilagenous tissues can be tolerated in this habitat.
In a few cases oil provides buoyancy (Bathyteuthidae) and in one case (Ocythoidae) a true
swimbladder has evolved (Packard and Wurtz, 1994).
Coleoid arms are unique grasping structures that possess a variety of different types of
suckers and hooks. The arms and suckers are also provided with a variety of sensory organs
that can be a primary means of investigating the immediate environment. This is especially
true of the octopods where the arms can hunt for unseen prey under rocks. Prey are seized
and passed to the mouth then cut into pieces by sharp beaks and swallowed. Because of the
circumesophageal organization of the molluscan central nervous system and its elaboration
into a complex brain in cephalopods, the cephalopod esophagus passes through the center of
the brain; this arrangement limits the maximum size of particles that can be swallowed.
Survival in the sea depends not only on obtaining food but also on avoiding predators.
Although preyed upon by many vertebrates (especially marine mammals) and other animals,
coleoids excel at camouflage and deception (see Hanlon and Messenger, 1996). They easily
exceed all other animals in their ability to change color quickly through the use of
chromatophoric organs (chromatophores) and various types of reflectors. The
chromatophores contain muscles under nervous control; quick muscle contraction produces
rapid color changes that immediately changes the appearance of the animal. This camouflage,
combined with their ability to secrete clouds or globs of ink, and the ability of many to alter
their skin texture and arm postures, provides them exceptional concealment abilities.
Another, perhaps surprising feature of coleoids, since many species reach a large size, is their
short life span. Most species grow rapidly and live for one year or less. Reasons for this are
not well understood. However, this is part of a life-history strategy that seems designed for
rapid increase in population size. One result of this is that cephalopod fisheries typically
fluctuate wildly in annual catch rates.
Characteristics
1. Shell located internally.

2. Muscular mantle for propulsion and respiration.
3. Brachial crown consisting of ten arms.

Figure. The diagramatic illustrations show the arm-crown modifications found in the
major groups of coleoid cephalopods. The Belemnoidea represent the primitive
condition with 10 equal arms. In the Vampyromorpha arms II are modified and in the
only living representative, Vampyroteuthis, these are a pair of slender, sensory
filaments that arise from the aboral surface of the web. In the Octopoda arm pair II is
lost leaving 8 arms, and in the Decapodiformes, arms IV are modified into tentacles
with suckers generally confined to distal clubs.
4. Fins present*
5. Chromatophores present*
6. Ink sac present.
7. Eyes with lenses.
* presence not established in Belemnoidea.
Comments
Many of these characters have been secondarily lost in various taxa.
Arm pairs are counted from dorsal to ventral as indicated in the illustration above. However,
the numbering of arms in taxonomic descriptions leads to some ambiguity. In the
Decapodiformes, the fourth of five pairs of arms has been modified as tentacles, leaving four
pairs of functional arms. As a result, the ventral arms are the true fifth pair but functionally
are the fourth pair. Similarly, the second pair of arms is modified in vampyroteuthids and lost
in octopods causing a difference between true (i.e., homologous) and functional arm-pair
numbering. True arm-pair numbering is used in the above illustration while functional
numbering is used in descriptions of taxa.
Nomenclature
General consensus on the names of higher taxonomic catagories in the Coleoidea has not
been achieved (eg, see Berthold and Engeser, 1987; Clarke, 1988; Doyle, et al., 1994; Haas,
2002).
Discussion of Phylogenetic Relationships
Extant coleoids form a monophyletic group, the Neocoleoidea, as evidenced by the uniform
presence of suckers. Their sister group, the Belemnoidea, is characterized by the presence of
hooks that are derived, perhaps, from muscular papillae, the homologues of trabeculae or cirri
that lie lateral to the suckers in living coleoids (e.g. Engeser and Bandel, 1988, see also Haas,
1989).
The Decapodiformes is supported by only a single morphological character that has been
demonstrated to be apomorphic (i.e. derived): the modification of the fourth pair of arms as
tentacles (Young and Vecchione, 1996). The monophyly, however, is supported by molecular
cladistic studies (Bonnaud et al. 1997; Carlini and Graves, 1999). The problem with
morphology, however, is not due to a lack of shared character states among decapods but
instead results from our inability to polarize these states. The outgroup on which most
polarity decisions are based is the Nautiloidea. Unfortunately, this group is morphologically
very different from coleoids and many characters are not applicable to both groups.
Young and Vecchione (1996, 2002) found five unambiguous character states that supported
the monophyly of the Octopodiformes:
1. Presence of an outer statocyst capsule.

2. Superior buccal lobes adjacent (fused at edges) or fused to posterior
buccal lobes of the brain.
3. Arms II modified as filaments. (This assumes an "ordered" evolutionary
relationship of the character states: arms II unmodified - arms II modified -
arms II absent.)
4. Primary gill lamellae possess a partial or complete median septum.
5. Secondary gill lamellae possess a partial or complete median septum.
An evolutionary study by Hass (2002) also placed the Vampyromorpha and Octopoda
together as a monophyletic group on the basis of a combination of characters four and five
above (he independently recognized these characters but defined them slightly differently).
Molecular studies support the monophyly of this group (Carlini and Graves, 1999) or give it
equivocal support (Bonnaud et al., 1997).
Other Names for Coleoidea Bather, 1888
Decapodiformes Leach, 1817

Squids, cuttlefishes and their relatives

The Decapodiformes contains about 95 genera and 450 species in 31 families. Many of these
families seem to form natural groups and these are included below without formal names as
the validity of the groups has not yet been demonstrated by cladistic methods.
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Containing group: Coleoidea
Introduction
The most distictive feature of decapods is the modification of the fourth pair of arms into
tentacles (see title photographs). In some species these are long slender structures that are
several times the length of the body and probably function much like a fishing line. In some
others, they are short, muscular structures that can be quickly extended (literally shot
forward) to grasp prey then completely retracted into pockets in the head.
The Decapodiformes includes species with a wide range of body forms and habits. Muscular
sepioids bury in the sand during the day; gelatinous squids swim via slow jet propulsion in
the deep-sea while in the same habitat some squids with weak mantle muscles use large
muscular fins for swimming; surface-dwelling squid with powerful jet propulsion may
undertake migrations of a thousand miles and, when disturbed, dart from the water and glide
over the ocean surface like a flying fish. In size they range from Idiosepius with a mantle
length of 8 mm at maturity to the giant squid Architeuthis which reaches nearly 18 m in total
length. Cephalopods are primarily visual animals even though they possess lateral line
analogs, very low frequency hearing and olfaction. The dominance of the visual system is
particularly true of decapods. For example, the eye diameter of the small Iridoteuthis is over
40% of the mantle length and large squids have eyes that surpass the eye-size of all other
animals big or small. Among decapods, the oegopsid squids characteristically produce small
pelagic eggs and the paralarvae develop in the near-surface plankton whereas myopsids and
sepioids have benthic eggs that often produce large demersal hatchlings.
The suckers of all decapods have horny rings. These rings often carry sharp claw-like teeth
and in some species these have been modified into hooks. The advantages of hooks are
uncertain. They could be designed for sinking into soft-bodied prey, such as other squid, or
they could act as grappling hooks for hard-body spiny prey where suckers would be
ineffective (Young, et al., 1999).

Figure. Oral-lateral view of a portion of the tentacular club of a preserved Ancistrocheirus
lesueurii showing hooks with sheaths. Photograph by R. Young.
Characteristics
1. Arms/tentacles
1. Arms IV modified as tentacles (see title photographs).
2. Suckers bilaterally symmetrical.
3. Suckers with horny rings.
4. Suckers stalks with constricted necks.
Figure. Arm suckers. Left - Oral view of arm suckers of Sthenoteuthis

oualaniensis, preserved. The reddish, toothed portion is the inner horny ring;
the yellow surrounding slats form the outer horny ring. Right - Lateral view of
an arm sucker and stalk of Histioteuthis hoylei, preserved. The skin has been
removed from the sucker stalk to show its shape: a broad base tapering to a
constricted neck where the stalk attaches to the sucker. Photographs by R.
Young.
5. Arms IV without connecting web.
2. Buccal crown
1. Buccal crown present.
Figure. Oral view of the brachial (=arm) crown surrounding the buccal crown
of Histioteuthis sp. Arrow points to one of seven buccal supports of the buccal
crown.
3. Head
1. Statocyst with single capsule.
2. Superior and posterior buccal lobes widely separated.
3. Inferior frontal lobe system of brain absent.
4. Photosensitive vesicles
1. Photosensitive vesicles lie within cephalic cartilage.
5. Fins
1. Fins present but without cartilagenous core.
2. Fin cartilage present at base of fins often as a plate.
6. Shell
1. Shell a phragmocone, gladius or absent.
7. Viscera
1. Nidamental glands present.
2. Crop absent.
3. Oviducal glands bilaterally symmetrical.
4. Digestive gland duct appendages surrounded by nephridial coelom.
5. Nephridial coeloms joined to form a single coelom.
6. Visceropericardial coelom extensive.
Comments
Comparisons of the major taxa within the Decapodiformes
Buccal Sucker
Corn Branchia Tentacle Interstellate
support circular
ea l canal pocket connective
suckers muscle
Oegopsida No Yes No No No Yes

Bathyteutho
No Yes Yes No Yes Yes
idea
Myopsida Yes Yes Yes/No Yes Yes Yes
Idiosepiidae Yes No No No Yes ?
Spirulidae No No No No Yes No
Sepioidea Yes No Yes/No Yes Yes No
A key to the families of the Decapodiformes can be found here.
At present we recognize four major lineages within the Decapodiformes but the relationships
between them are unresolved. Two additional taxa, Bathyteuthoidea and Idiosepiidae, which
have subordinal rankings are included here as their ordinal relationships are uncertain. The
Bathyteuthoidea shows relationships with both the Oegopsida and the Myopsida while the
Idiosepiidae shows relationships with the Myopsida and the Sepioidea. Among the orders, the
Myopsida shows relationships with both the Oegopsida and the Sepioidea and the Sepioidea
and the Spirulida show relationships.
Among the latter two orders, the closeness of the relationship is uncertain. The Sepiidae of
the Sepioidea and the Spirulidae of the Spirulida have substantial fossil records and these
records argue for a close relationship between these two orders. In both cases, the presence of
an internal calcareous shell with a phragmocone, however, represents the retention of an
ancient and plesiomorphic character state, and does not indicate that the animal is an early
derivative of the decapod clade or close relationships. The details of the shell structure do
support a close relationship.
Spirula spirula, the only living member of the Spirulida, is a peculiar cephalopod that has
stimulated much discussion regarding its phylogenic position. While the relationships are still
unresolved, Young, et al. (1998) support its close relationship to the Sepiidae and other
sepioid families based on arguments mostly put forward by Naef (1922). A cladistic study
based on morphology has not been made.

Figure. Drawings from Young et al. (1998) modified from Naef (1922) show the similarities
between Spirulirostra, Spirulirostrina (fossil relatives of Spirula) and Belosaepia (a fossil
relative of Sepia) and Sepia. Note the gray-colored "guard-like sheath" on all species
illustrated (it forms the rostrum, shield,etc. on the Sepia cuttlebone). The cuttlebone of
Belosaepia lacks the cross-struts of the Sepia cuttlebone and has distinct septa on the ventral
side of the siphuncle. The siphuncle is also more ventrally curved than in Sepia. The
presumed attachment of the mantle muscle (red) to the ventral process of the sheath is shown
in Spirulirostra and Spirulirostrina. The mantle attaches similarly to the recurved ventral
process of the cuttlebones.
Both the spirulid and sepiid shells have a distinctive ventral curvature of the apical end and
the asymmetrical elaboration of the sheath that includes such similarities as a rostrum and a
ventral process (Young, et al., 1998).
Bathyteuthoidea Vecchione, Young and

Sweeney, 2004
Richard E. Young and Michael Vecchione

• Bathyteuthidae Pfeffer 1900
• Chtenopterygidae Grimpe 1922
Containing group: Decapodiformes
Introduction
These are small mesopelagic to bathypelagic squids.
Brief diagnosis:
A decapodiform ...
• with suckers on the buccal membrane.

• without carpal locking-apparatus on tentacles.
• with branchial canals in gills.
Characteristics
1. Arms
1. Suckers in four or more series at some point on arms I-III.
2. Suckers without circularis muscles.
2. Tentacles
1. Club not divided into manus and dactylus.
2. Carpal locking-apparatus absent.
3. Club suckers in more than 7 irregular series.
3. Buccal crown
1. Buccal supports bear small suckers.
Figure. Side view of buccal crown and oral views of arms II and III of
Bathyteuthis berryi. Arrows point to suckers on the buccal supports.
Photograph by R. Young.
4. Head
1. Head with tentacle pockets.
5. Mantle
1. Mantle locking-apparatus reaches anterior mantle edge.
6. Fins
1. Fins with posterior lobes.
7. Shell
1. Shell a gladius with conus present in Chtenopteryx but absent in
Bathyteuthis.
8. Viscera
1. Gills with branchial canals.
2. Oviducts paired.
9. Eggs
1. Egg masses unknown.
Comments
Buccal connectives attach to the ventral borders of Arms IV in the Chtenopterygidae and to
the dorsal borders in the Bathyteuthididae. Suckers on the buccal supports are found in
members of only four families (Bathyteuthidae, Chtenopterygidae, Loliginidae, Sepiidae).
The suckers look much like the arm suckers but are much smaller.
The two genera that represent these families (Bathyteuthis, Cthenopteryx) were, at one time,
placed within the same family (e.g., Naef, 1921) because they share a number of similar
features. They especially show strong similarities in the structure of the tentacular clubs, the
sucker arrangement on the arms and the morphology of their gladii. Roper (1969) considered
them to belong to separate families and emphasized the differences in the attachment of the
buccal crown to arms IV. Others (e.g., Clarke, 1988) have considered them to be distantly
related. Their close relationship to one another, however, has been confirmed by a molecular
study (Carlini, 1998).
The relationship of the Bathyteuthoidea to other groups is uncertain as they share some
characters with both the Oegopsida and the Myopsida.
Other Names for Bathyteuthoidea Vecchione, Young and Sweeney, 2004
• Bathyteuthoida
Idiosepiidae Appellof, 1898

Pygmy squids
Katharina M. Mangold (1922-2003) and Richard E. Young

This family contains the single genus Idiosepius with about eight species:
• Idiosepius biserialis
• Idiosepius macrocheir
• Idiosepius mimus (?)
• Idiosepius notoides
• Idiosepius picteti
• Idiosepius paradoxus
• Idiosepius pygmaeus
• Idiosepius thailandicus
Introduction
This Idiosepiidae contains the smallest species within the Decapoda. Males in some species
mature at lengths of 6 mm ML and females at 8 mm ML (Nesis, 1982/7). These cephalopods
are elongate but have a rounded posterior mantle and separate fins. A unique attachment
organ is present on the dorsal surface of the mantle which is used for attaching the animal to
seaweed. They are found in shallow water in the Indo-West Pacific.
Brief diagnosis:
A decapodiform ...
• with an adhesive organ on the dorsal surface of the mantle.
Characteristics
1. Arms
1. Both arms IV hectocotylized.
Figure. Oral view of hectocotylized arms IV of Idiosepius pygmaeus, 18 mm

ML, off Philippines. Drawing from Voss (1963).
2. Arms with suckers in two series.

3. Arm and tentacle suckers without circularis muscles.
2. Tentacles
1. Tentacles without carpal (= proximal) locking-apparatus.
2. Tentacular clubs with suckers in 2 - 4 series.
3. Head
2. Eyes with corneas.
4. Funnel
1. Funnel without lateral funnel adductors.
5. Mantle
1. Adhesive organ on dorsal surface (see title drawing).

Figure. Top - Dorsal view of Idiosepius sp., preserved, with an enlargement
of a portion of the dorsal adhesive organ showing the glandular (white
reflection) epithelium. 12 mm ML, mature female, 05°27'S, 134°27'E.
Photograph by R. Young. Bottom -Side view of Idiosepius paradoxus in an
aquarium attached by its adhesive organ to a blade of seagrass. Photograph by
Takashi Kasugai, Port of Nagoya Public Aquarium.
2. Mantle not fused to head in nuchal region, but nuchal cartilage

absent.
3. Mantle locking-apparatus does not reach anterior mantle margin.
6. Fins
1. Fins completely separate; with posterior lobes.
7. Shell
1. Shell a delicate gladius not reaching anterior or posterior ends of
mantle.
8. Viscera
1. Right oviduct present but not functional.
2. Accessory nidamental glands present.
3. Gills without branchial canals.
9. Eggs
1. Eggs attach to substrate in flat masses.
Nomenclature
A list of all nominal genera and species in the Idiosepiidae can be found here. The list
includes the current status and type species of all genera, the current status, type repository
and type locality of all species and all pertinent references.
Life History
I. paradoxus spawns repeatedly in captivity. A single individual may lay as many as 42

batches over a 70 day period (Kasugai, 2006).
Figure. Side view of a spawning female I. paradoxus in an aquarium. The squid is attached
to a blade of seagrass by its adhesive organ and is attaching its eggs to the underside of the
same blade. Another I. paradoxus is swimming nearby, presumably a male. Photograph by
Takashi Kasugai.
Other Names for Idiosepiidae Appellof, 1898
• Pygmy squids
References
Kasugai, T. 2006. Spawning mode and reproductive output of the Japanese pygmy squid
Idiosepius paradoxus (Cephalopoda: Idiosepiidae). Program and Abstract Book, CIAC '06,
Hobart Tasmania.
Nesis, K. N. 1982. Abridged key to the cephalopod mollusks of the world's ocean. 385,ii pp.
Light and Food Industry Publishing House, Moscow. (In Russian.). Translated into English
by B. S. Levitov, ed. by L. A. Burgess (1987), Cephalopods of the world. T. F. H.
Publications, Neptune City, NJ, 351pp.
Voss, G. L. 1963. Cephalopoda of the Philippine Islands. Bull. U. S. Nat. Mus., 234: 1-180.
Title Illustrations
Scientific
Idiosepius notoides
Name
Location off Australia
Acknowledge provided by Mark

ments Norman
Copyright © 1996 David Paul
Scientific
Idiosepius pygmaeus
Name
Location Off Philippine Islands
Voss, G. L. 1963. Cephalopoda of the Philippine Islands. Bull. U. S.

Reference
Nat. Mus., 234: 1-180.
Sex Male
View Dorsal and ventral

Size 18 mm ML
Myopsida Naef, 1916

Michael Vecchione and Richard E. Young
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Introduction
Myopsid squids are neritic, often in very shallow water, or upper slope demersal species.
Many species are strong swimmers, occur in large schools and are fished commercially for
food. The Loliginidae contains many species some of which reach a rather large size (at least
90 cm ML in Loligo forbesii) but those in Pickfordiateuthis, are dwarf species where males
may mature at less than 14 mm ML (Brachoniecki, 1996). The Australiteuthidae contains a
single species that is also a dwarf with males that mature as small as 17 mm ML (Lu, 2005).
Brief diagnosis:
A decapodiform ...
• with corneal membranes covering eye lenses.

• without secondary eyelids.
• with well-developed gladius.
Characteristics
1. Arms
1. Suckers of arms (and tentacles) with circularis muscles (unknown in
Australiteuthidae).
2. Tentacles
1. Club without proximal (= carpal) locking-apparatus.
Figure. Oral view of the tentacular club of Loligo plei, 105 mm ML,
preserved. Photograph by R. Young.
3. Head
1. Head with tentacle pocket.
2. Eyes with corneal membranes covering lenses.
3. Eyes without secondary (= ventral) eyelid.
Figure. Left - Dorsal view of the head of Sepioteuthis sp. (Loliginidae)

showing cornea covering lens. Photograph by Mark Norman. Right -
Ventrolateral view of the head of Lollicuncula diomedeae showing the
absence of a ventral eyelid. Photograph by R. Young.
4. Funnel
1. Funnel without lateral adductor muscles.
5. Mantle
1. Mantle locking-apparatus extends to mantle edge (except
Australiteuthidae).
6. Shell
1. Shell a gladius, extending the full length of the mantle.
7. Viscera
1. Gills with branchial canal (except Pickfordiateuthis, unknown in
Australiteuthis).
2. Right oviduct absent.
3. Females with accessory nidamental glands.
Figure. Diagramatic cross-section through gills. Drawing modified from Naef

(1921-23).
8. Eggs
1. Eggs, where known, attached to substrate.
2. Embryo with large external yolk sac.
Australiteuthidae Lu, 2005
Australiteuthis aldrichi Lu, 2005

Chung Cheng Lu and Richard E. Young
Containing group: Myopsida
Introduction
Australiteuthis aldrichi was described from numerous specimens collected from the inshore
waters of Northern Australia. This is a dwarf species, the largest squid described was a
mature female of 27.6 mm ML. The holotype is a mature male of 21.3 mm ML. The small
squid has never been observed alive.
Brief diagnosis:
A myopsid squid ...
• with peculiar funnel locking-apparatus containing a boomerang-shaped

groove.
• with fins separated from one-another and with posterior lobes.
• with dumbell-shaped photophore on ink sac.
Characteristics
1. Funnel
1. Funnel locking apparatus with subcircular shape and "bisected by a
boomerang-shaped groove running anteroposteriorly. Anterior wing
of the boomerang-shaped groove deep, with wide anterior end,
groove deepens and becomes slightly narrower posteriorly to form
deep, broad pit. Pit becomes a narrow and shallow groove
posteriorly and turns medially forming posterior limb of the
boomerang, posterior end of groove slightly widened." (Lu, 2005).
2. Mantle component does not reach anterior mantle margin.
Figure. Frontal views of the funnel/mantle locking-apparatus of A. aldrichi,

17.3 mm ML, mature male, paratype. Top - Mantle cut open showing the
funnel and mantle components of the locking-apparatus on the left side.
Photograph from Lu (2005). The photograph also shows that there is no lateral
adductor muscle connecting the funnel to the head and that there is no ventral
secondary eyelid on the eye, a character state similar to that of loliginids but
not sepiids and sepiolids. Bottom left - Funnel component. Bottom right -
Mantle component. Bottom drawings were of the right locking apparatus but
the illustration here has been flipped to coincide with the photograph.
Drawings from Lu (2005).
2. Fins
1. Fins large, round, separate (length 47-63% of ML, width 80-104% of
ML); separation at posterior end inconspicuous.
2. Fins with free anterior and posterior lobes.
Figure. Dorsal and ventral views of the fins, holotype. Enlargement from the
title photograph.
3. Photophores
1. Large dumbell-shaped photophore present. The photophore appears
to contain a large population of bacteria (Lu, 2005).
Figure. Photophore of A. aldirchi, 22.3 mm ML, mature male. A - Ventral

view of the mantle cavity with some viscera removed showing the two lobes
of the photophore (arrows) and rectum between the lobes. B -
Figure. Histological section through one arm of the photophore. Note the
inksac surrounding the photophore, the laminated structure (at the left) that
presumably houses luminescent bacteria and the lens-like tissue (center top).
At the far right is the rectum. Photograph from Lu (2005).Shell
Comments
More details of the description can be found here.
Distribution
Type locality: Joseph Bonaparte Gulf, Western Australia at 14°07.30S, 128°02.00'E. In

addition to Joseph Bonaparte Gulf, squids have been collected from inshore waters near
Darwin and Paupa New Giunea, which suggests a rather broad distribution in shallow waters
of this general region.
Other Names for Australiteuthis aldrichi Lu, 2005
• Australiteuthidae Lu, 2005
Loliginidae Lesueur, 1821


At present the Loliginidae is divided into ten genera with 42 nominal species. The
generic affinities of three species currently are not resolved.
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Containing group: Myopsida
Introduction
The family Loliginidae includes many species that are important in trophic systems, fisheries,
and biomedical studies. Brakoniecki (1996) merged the Pickfordiateuthidae with the
Loliginidae.
Loliginids are mostly neritic squids (i.e., they occupy waters over the continental shelves).
Most are very muscular, "squid-like" in appearance and range in size from about 3 to 100 cm
ML. All members of the family have a cornea that covers the lens of each eye. Among
decapods, this feature is shared with all members of the Sepioidea but is absent in all
oegopsid squids. Loliginids differ from sepioids in having a gladius that extends the full
length of the mantle and a gill that (except in Pickfordiateuthis) has a branchial canal.
Brief diagnosis:
A myposid ...
• with straight funnel locking-apparatus.

• with mantle locking-apparatus reaching mantle margin.
Characteristics
1. Buccal crown
1. buccal supports usually bear suckers.
2. Funnel
1. Funnel locking-apparatus with a straight groove.
3. Mantle
1. Mantle locking-apparatus reaches anterior mantle margin.
4. Fins
1. Fins usually joined posteriorly and, except Pickfordiateuthis,
without posterior lobes.
5. Viscera
1. Bacterial photophores on viscera present or absent. If present, form
single, apparently unconnected, pair.
6. Egg masses
1. Egg masses, where known, with eggs organized in finger-like
masses and attached to substrate.
Generic characters
Characters that Vecchione et al. (1998) agreed were of generic importance within the
Loliginidae include:
1. Fins.
1. lateral.
2. terminal.
2. Tail-like extension of the posterior mantle.

1. extends beyond fins.
2. fins along sides to posterior tip.
3. absent.
3. Eggs.
1. large (longest axis >5mm).
2. small (longest axis <4mm).
4. Arm-sucker rings, proximal margin.

1. semicrescent plate.
2. square teeth.
3. pointed teeth.
5. Arm-sucker rings, distal margin.

1. square teeth.
2. pointed teeth.
6. Hectocotylus, ventral row of suckers.

1. reduction of suckers and elongation of sucker stalks along modified
portion of arm to form papillae.
2. ventral crest formed by fusion of the protective membrane with
ventral row of papillae such that original form of conical papillae is
completely obscured.
3. unmodified.
7. Hectocotylus, proximal region.

1. suckers and/or stalks modified.
2. unmodified.
8. Photophores on ventral ink sac.

1. one pair present.
2. absent.
9. Spermatophore, cement body.
1. short.
2. long.
Comments
The morphology of the gladius, as used formerly to distinguish the genus Doryteuthis, is no
longer considered to be of generic importance, although it is important at the specific level
(see also Alexeyev, 1989). The gladius is quite variable, both within and among species, and
could be highly adaptive in response to differences in swimming behavior. Other traditional
characters that Vecchione et al. (1998) consider too variable to be of use in the generic
systematics of the Loliginidae include spermatophore deposition site, presence of suckers on
the buccal lappets, adult chromatophore patterns (e.g. lateral "flame stripes"), and presence of
a longitudinal mid-ventral ridge on the mantle.
Although the preliminary generic classification of Vecchione et al. (1998) represented an

important step toward consensus on the genera of this family, subsequent analyses of DNA
sequences (Anderson 2000) have indicated that a holophyletic classification requires
recognician of generic-level species groups defined primarily on distributional characteristics.
Nomenclature
Lesueur (1821) erected a family that he called Loligoidea (afterwards emended to

Loliginidae, see Vecchione et al., 1998) to fit the "Loligos" (Leachia, Loligo and Onykia)
into Cuvier's "natural order". He stated (Lesueur, 1821:88)--
"It is of little consequence what characters we select for the distribution of these
animals into families and genera, if our arrangement is the most convenient, and
exhibits, as near as possible, a graded transition from one to the other."
A consequence of the age of this family, and its early inclusion of all squids, is that many
taxa, both correctly and incorrectly described, have been considered loliginids at one time or
another. A summary of this history is found here. Therefore, taxonomic nomenclature in the
squid literature can be very confusing.
A thorough nomenclatural review of the family can be found in Sweeney and Vecchione
(1998).
Our present classification of the Loliginidae comes from a consensus report by the world's
experts on he systematics of this family that was published in the Phuket Marine Biological
Center Research Bulletin, No. 66, 2005. A copy of this paper can be found here.
A list of all nominal genera and species in the Loliginidae can be found here. The list
includes the current status and type species of all genera, and the current status, type
repository and type locality of all species and all pertinent references.
The systematics of the myopsid squid family Loliginidae have long been in disorder. In
addition to many problems with differentiation of species, loliginid systematics have been
hampered throughout the last several decades of the 20th Century by the presence of two
systems of generic-level classification. The differences between these two systems primarily
involved a question of the importance of gladius structure at the generic level. Both systems
have been used widely in the scientific literature, although many authors consistently
qualified their use by stating that the family was badly in need of revision based upon a
worldwide review.
Three separate morphological revisions have been completed (Natsukari, 1984b;

Brakoniecki, 1986; Alexeyev, 1991); all concluded that a correct generic-level classification
of the family is radically different from either of the previous classifications. Unfortunately,
these new classifications also differed substantially from each other, were presented in
unpublished dissertations and not widely disseminated, and none has gained full acceptance.
Obviously, the existence of five contrasting systems of classification can cause hopeless
confusion to researchers studing these squids, in addition to obfuscating real relationships
among the species. Species groupings into subgenera and genera have been based only on
similarity. In a workshop publication, Vecchione et al. (1998), recognized six genera. Four of
the genera are divided into subgenera: Lolliguncula (Lolliguncula and Loliolopsis);
Uroteuthis (Uroteuthis and Photololigo); Loliolus (Loliolus and Nipponololigo); Loligo
(Loligo and Alloteuthis). The generic affinities of several species were unresolved.
A follow-up workshop consensus on generic and sub-generic taxonomy of the family was
published by Vecchione et al. (2005), based primarily on the phylogenetic research of
Anderson (1996; 2000a; 2000b) and Alexeyev (1989; 1991). Both of those authors built
upon the previous workshop proceedings. The most noteworthy differences in Vecchione et
al. (2005) from the classification of Vecchione et al. (1998) include the following: (1)
removal of mercatoris from Lolliguncula based primarily on gladius characters, DNA
sequence data and biogeography, and recognition of Afrololigo Brakoniecki as a valid genus
for this species; (2) elevation of Alloteuthis Wülker from sub-generic to full generic status
based on gladius structure and DNA sequence data; (3) removal of American species from
the genus Loligo because DNA sequence analyses indicate that Loligo sensu Vecchione et al.
(1998) is probably paraphyletic. The generic name with priority for the American species is
Doryteuthis Naef. This genus further comprises two natural subgroups based on differences
in gladius and hectocotylus structure; these subgroups are considered here to be the
subgenera Doryteuthis Naef and Amerigo Brakoniecki. Doryteuthis sanpaulensis does not
belong in either of these subgenera and is therefore considered to be the sole recognized
species in an undescribed subgenus; (4) removal of bleekeri from Loligo and recognition of
Heterololigo Natsukari as a valid genus based on DNA sequence analysis; (5) removal of
noctiluca from subgenus Uroteuthis of genus Uroteuthis and recognition of Aestuariolus
Alexeyev as a valid monotypic subgenus of Uroteuthis primarily because of differences in
photophore structure from the rest of the genus.
Life History
The egg masses typically are groups of finger-like gelatinous masses containing a few to
many eggs each often woven together and always attached to the ocean floor.

Figure. Left - Side view of an egg mass of a loliginid that has been pulled out of its anchor in
the sand, found at low tide off San Felipe, Baja California. Note the root like anchors at the
bottom of the picture. Photograph by Mike Lang. Right - Ventral views of a late embryo
within the swollen egg chorion, and a recently hatched paralarva of Loligo pealeii.
Photographed by Clyde Roper.
Loligo Lamarck, 1798

Inshore squid
The following three species are present in the genus.

pages.
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Type species.-- Loligo vulgaris Lamarck, 1798, by subsequent designation of

Hoyle (1910).
Containing group: Loliginidae
Introduction
Loligo species are the common inshore squids of the eastern Atlantic Ocean. Several species
are targets of substantial fisheries. Although the biology of some species of Loligo (e.g., L.
vulgaris, L. reynaudii ) is among the best known of any cephalopod, considerable
controversy remains about their taxonomy and systematic relationships.
The genus is defined by geography and the absence of specific modifications that define other
genera.
Brief diagnosis:
A loliginid ...
• with rhomboidal fins in adults, longer than broad (length <70% of ML);
mantle elongate; without photophores.
• with an Eastern Atlantic distribution.
Characteristics
1. Arms
1. Hectocotylus
1. Ventral crest absent.
2. Proximal suckers unmodified.
3. Modified suckers of reduced size and sucker stalks elongated
to form papillae in either dorsal or both dorsal and ventral
series.
2. Tentacles
1. Tentacular clubs expanded; suckers in four series.
3. Mantle
1. Elongate, posterior tip blunt.
4. Fins
1. Fins in adults rhomboidal and longer than broad, tapering
posteriorly.
2. Fins extend to posterior tip of mantle.
Figure. Ventral view of a mature male of L. vulgaris. Drawing from Naef

(1921-3).
5. Photophores
1. Photophores absent.
6. Viscera
1. Eggs less than 4 mm.
2. Spermatophore cement body short.
Comments:
Alloteuthis is morphologically very similar to Loligo. However, molecular analyses by

Anderson (2000) indicate that Alloteuthis should be considered a separate genus. This
supports earlier inferences by Naef (1921-23) and Alexeyev (1989) that the presence of a
gladial conus in Alloteuthis should be considered a generic character. The conus in
Alloteuthis would clearly separate the genera but it can be very difficult to find and in some
Alloteuthis specimens may be reduced until it is essentially absent. The posterior tip of the
mantle in Alloteuthis is elongate into a tail-like structure that tends to be more pointed than in
Loligo. This is particularly true for males but identification of this character state can be
subjective in females.
Nomenclature
According to Lesueur (1821), use of the name "loligo" for squids can be traced to the
classical Roman philosopher, Pliny (lib. IV, cap. XIX; see also Naef, 1923:197). Linnaeus
established Sepia loligo in 1758 to encompass all cephalopods with a cylindrical body and
fins (Hoyle, 1910:410). Subsequently, squids in general were often referred to as "loligos"
(e.g. Lesueur, 1821). Many authors have interpreted Schneider's (1784) use of the name
"Loligo" (capitalized and not preceded by a generic name) as the establishment of the genus
Loligo. However, Hemming (1954) showed that Loligo Schneider, 1784 resulted from a
misinterpretation of Schneider's work. Schneider's intent was to remove Nautilus Linnaeus,
1758 and Argonauta Linnaeus, 1758 from the univalve molluscs and to include them with all
other cephalopods in a new genus, Octopodia. Thus, Schneider actually proposed the new
combination Octopodia loligo but did not follow the convention of always using the generic
name and not capitalizing the species. Loligo was then formally described by Lamarck in
1798 to distinguish four squid species, L. vulgaris, L. subulata, L. sagittata and L. sepiola
from the octopods (for which he erected the genus Octopus Lamarck, 1798) and from the
cuttlefishes of the genus Sepia.
The only other squid genera that predate 1821 are Onychoteuthis Lichtenstein, 1818, which
was established to distinguish cephalopods in which hooks replace some of the suckers on the
arms or tentacles, and Cranchia Leach, 1817, a very unusual squid.
Except for geographic distribution, Loligo species are unified in their lack of the characters
distinctive of some other genera (photophores, spermatophore with long cement body, large
eggs, hectocotylus with ventral crest, etc.). The most distinct of the three species is L.
forbesi. The southern African species, L. reynaudii, is very similar to L. vulgaris and , based
on electrophoresis, Augustyn and Grant (1988) concluded that L. reynaudii should be
considered a subspecies of L. vulgaris. However, L. reynaudii and L. vulgaris are allopatric
and distinct morphologically; we consider them to be distinct species.
Distribution
Loligo includes species of the eastern Atlantic Ocean and forms a nearly continuous
distribution from southern Africa to 60°N.
Other Names for Loligo Lamarck, 1798
• Vernacular Names: Inshore squid
Loligo forbesii Steenstrup 1856

Veined squid
Containing group: Loligo
Introduction
Maximum mantle length 90 cm in males and 41 cm in females. Off the Azores,

length at first maturity is 57 cm in males and 34 cm in females. In the English
Channel, the size of summer spawning females is 20 cm and 29 cm in winter
spawning females.
Characteristics
1. Mantle
1. Mantle long, moderately slender, cylindrical.
2. Fins
1. Fins rhomboidal, their length three quarters that of mantle, their
posterior borders slightly concave.
3. Arms:
1. Largest arm sucker rings with 7 or 8 teeth.
2. Left ventral arm hectocotylized in its distal third by modification of
suckers into long papillae which gradually decrease in size distally.
4. Tentacles
1. Suckers on manus of tentacular club subequal in size; sucker rings
with 13 to 18 sharp, conical teeth.
Habitat and biology
A species of subtropical and temperate waters (it avoids temperatures below

8.5°C) occurring over the shelf in the temperate part of its distributional range,
but found in deeper waters in subtropical areas; the entire depth range
extending from about 100 to 400 m.
The population in the northeastern Atlantic is known to carry out seasonal
migrations, spending the summer in the North Sea and the eastern part of the
English Channel and overwintering in the western part of the Channel. In daytime
squids aggregate near the bottom, dispersing at night throughout the water
column.
Spawning occurs almost throughout the year in the English Channel, showing a
peak in winter (December and January, at temperature of 9 to 11 °C) and
another one in summer. The eggs are attached to hard objects on sandy or
muddy bottoms; hatching occurs after 30 to 40 days. Juveniles hatched in
January and February attain sizes of approximately 11.5 cm in June; by August,
the females measure about 14 cm, the males about 15 cm mantle length, and in
November about 25 to 30 cm respectively. Both sexes are then mature (the
males beginning in October).
Loligo forbesii feeds on small and juvenile fishes, and to a minor extent on
other cephalopods, crustaceans, and polychaetes; cannibalism is common. Off
the Azores, the most important fish species in their diet is blue jack mackerel
(Trachurus picturatus). (Lepidopus caudatus). Other fishes preyed upon include
bogue (Boops boops), and silver scabbardfish
Geographical distribution
Eastern Atlantic: 20 °N to 60 °N (excluding the Baltic Sea), Azores Islands, and
along west African coast South to the Canary Islands; Mediterranean Sea;
southern boundary is unknown. .
Remarks
The population in the Azores is genetically distinct from those of mainland

Europe.
Other Names for Loligo forbesii Steenstrup 1856
• Veined squid
• Vernacular Names: En: Veined squid, Fr: Encornet veiné, Sp: Calamar
veteado, Local species names: Portugal and Azores: Lula.
Title Illustrations

Scientific
Loligo forbesii
Name
C. F. E. Roper, Michael J. Sweeney and Cornelia E. Nauen. 1984.

Acknowledge Cephalopods of the World. An Annotated and Illustrated
ments Catalogue of Species of Interest to Fisheries. FAO Fisheries
Synopsis No 125, Vol. 3. 277 pp. Rome: FAO 1984.
Loligo reynaudii Orbigny 1845

Cape Hope squid
Introduction
A loliginid of southern African waters. Maximum mantle length 40 cm, weight
more than 1 kg.
Characteristics
1. Mantle
1. Mantle narrow, elongate.
2. Fins
1. Fins long, 65% or more of mantle length.
3. Arms:
1. Arms short (in comparison with L. vulgaris).
4. Tentacles
1. Tentacles long; clubs expanded.
2. Club suckers on the manus (medial series) greatly enlarged.
3. Club sucker rings smooth or with 16-20 teeth.
Habitat and biology
This species has been a subject of numerous studies because of the importance of the jig
fishery. It spawns primarily during September to December on sand beds in shallow bays
along the south coast of South Africa. Size at maturity is highly variable, depending on
location and time of year. Males may be mature at 9 cm or immature at 25 cm ML, females
mature at 10-18 cm ML. Mature squid arrive on spawning grounds in sexually segregated
schools. Individual squid may spawn over a period of weeks or months. No post-spawning
mass mortality has been observed.
Inshore/eastward-offshore/westward migrations are correlated with water temperature and

wind direction. Offshore distribution reaches depths of 200-300 m.
Eastern central Atlantic: From South Africa to southern Namibia.
Remarks
Loligo reynaudi has previously been proposed to be a subspecies of L.

vulgaris.
Other Names for Loligo reynaudii Orbigny 1845
• Cape Hope squid

• Local species names: Chokker squid
• Vernacular Names: En: Cape Hope squid, Fr: Calmar du Cap, Sp:
Calamar del Cabo
Loligo vulgaris Lamarck 1798

European squid

Introduction
The European squid is the type species of the genus Loligo. Maximum mantle
length is about 42 cm in males, 32 cm in females; maximum weight 1.5 kg.
Characteristics
1. Mantle
1. Mantle long, moderately slender, cylindrical.
2. Fins
1. Fins rhomboid, their length two thirds of mantle length, posterior
border slightly concave.
3. Arms:
1. arm sucker rings with 20 teeth, distal teeth large and pointed,
proximal teeth minute or absent.
2. Left ventral arm hectocotylized along its distal 1/3 - 2/4 by
modification of suckers into papillae that decrease in size distally
4. Tentacles
1. Manus of tentacular clubs with 4 longitudinal series of suckers, 2
median series with 6 enlarged suckers each.
2. sucker rings of median series on manus with approximately 30
irregularly-sized teeth; club with about 36 transverse rows of
minute suckers.
Habitat and biology
Ranging in depth from the surface to approximately 500 m, and most abundant
between 20 and 250 m; deepest in winter. European squid is known to migrate
vertically and horizontally in response to changes in environmental conditions.
The stock in the northeastern Atlantic overwinters in deeper waters off Portugal,
approaches the French coast in spring, and migrates from May through June
farther north into the North Sea where spawning takes place in depths from 20
to 80 m. A southward migration takes place in fall. The stock on the west
Saharan fishing grounds likewise overwinters in deeper offshore waters and
migrates onshore for spawning in spring and fall. Accordingly, juveniles appear to
recruit into the fishery in February and March and between July and September.
In the western Mediterranean, European squid migrate into deeper water in late
fall; the largest individuals begin their onshore migration as early as in January
and February, followed in summer by the smaller ones.
Spawning extends for most of the year with peaks in early summer and early fall.
Females produce up to 20000 small eggs (diameter about 2 mm) that are
deposited in gelatinous tubes containing tens of eggs and attached to debris and
other hard objects on sandy to muddy bottoms. The eggs hatch after an
incubation period ranging between 25 days (at 22° C) and 45 days (at 12 to
14°C). The number (up to 800) and size of spermatophores is directly related to
the size of the male. Males reproducing for the second time usually carry more
than those reproducing for the first time.
Growth is faster in summer than in winter in both sexes; the rate is greater in
males as compared with females. In the Atlantic, June hatchlings attain
approximately 12 cm mantle length in December and 13 or 14 cm in the
following April. By August, males reach about 17.5 cm and in April of their
second year 21 cm, compared to 17 cm in females. In the western
Mediterranean, juvenile females and males migrating onshore in July measure
about 7 and 8 cm respectively (ranges 6.6 to 8 cm, and 7 to 8.3 cm) and grow to
about 15 and 16 cm in December (ranges 14 to 16 cm and 15 to 17 cm). On the
other hand, juveniles approaching the coast in October are approximately 5.7 cm
long (range 5 to 6.3 cm) growing to 8.3 cm (range 7.8 to 8.9 cm) towards the
end of December when they leave the shallow waters. By March they have
attained a length of 13 cm. After May they mix with the other group in a new
onshore migration to where spawning occurs.
Longevity is 2 years in females and about 3 years in males.
European squid feed on fishes and crustaceans. Cannibalism is common.
Eastern Atlantic: From approximately 20° S to 55°N; around the British Isles,
North Sea; Mediterranean Sea.
Remarks
Loligo reynaudi has previously been proposed to be a subspecies of L.

vulgaris.
Other Names for Loligo vulgaris Lamarck 1798
Afrololigo mercatoris (Adam, 1941)

Guinean thumbstall squid
A single species is presently recognized in Afrololigo.
Type species. -- Lolliguncula mercatoris Adam, 1941 by original designation.
Introduction
A. mercatoris is a small species with females reaching 5 cm ML and males 3.5 cm ML. In
general appearance it resembles species of Lolliguncula and Loliolus. Afrololigo mercatoris
is known only from the west African coast where it has been collected at depths of less than
50 m on mud and sandy mud bottoms.
Brief diagnosis:
A loliginid ...
• with short mantle, rounded posteriorly; fins wider than long but without
posterior lobes.
• without a ventral crest on the hectocotylus.
• with distribution restricted to waters off West Africa.
Characteristics
1. Arms
1. Arms I extremely short compared to other arms.
2. Arm sucker rings with square, plate-like teeth around entire margin.
3. Suckers on mid-sections of lateral arms of males greatly enlarged.

Figure. Dorsolateral view of the brachial crown of Afrololigo mercatoris,
male, 04°06'N, 05°58'E.
4. Hectocotylus (left arm IV):

2. Proximal half with 6-12 pairs of normal suckers.
3. Distal half with elongate papillae replacing suckers, papillae
of dorsal series more strongly developed.
2. Tentacles
1. Tentacular clubs narrow, small; suckers in four series.
2. 4 or 5 pairs of medial suckers on manus much larger than lateral
suckers.
3. Club sucker rings with 15-25 teeth.
3. Head
1. Head short.
2. Buccal supports without suckers.
4. Mantle
1. Mantle bluntly rounded posteriorly; without posterior tail-like
elongation.
5. Fins
1. Fins rounded, short (length 40 - 45% ML), broad (55 - 65% ML) with
convex posterior margins.
6. Photophores
7. Viscera
1. Eggs small.
2. Spermatophores with long cement body.
Comments
Many of the morphological characters of Afrololigo mercatoris are shared with

species of Lolliguncula, the genus in which it was originally described. DNA
sequence analysis by Anderson (2000) supports Brakoniecki's (1986) conclusion
that Afrololigo is a distinct genus.
Distribution
Eastern central Atlantic: limited to the west coast of Africa from Rio de Oro (Mauritania) to
Lüderitz Bay (Namibia).
Remarks
“Loligo” abulati Adam 1955 in the Red Sea appears to be closely related to A.
mercatoris, but is too poorly known to include in the species accounts.
Other Names for Afrololigo mercatoris (Adam, 1941)
• Vernacular Names: Guinean thumbstall squid
Alloteuthis Wulker, 1920

Image: Photograph by Peter Wirtz
Alloteuthis contains three species.

pages.
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Type species. -- Loligo media Linnè, 1758 by original designation of Naef in an

unpublished manuscript cited by Wulker [see Vecchione et al. (1998)].
Introduction
Small loliginids of the eastern Atlantic.
Brief diagnosis:
A loliginid ...
• with posterior mantle elongated as a tail-like structure; fins longer than

broad.
• with a small conus sometimes apparent on gladius.
• with an Eastern Atlantic distribution
Characteristics
1. Arms
1. Hectocotylus:
3. Proximal region with 10 - 12 (usually 11) normal suckers in
ventral series followed distally by papillae.
Figure. Oral view of the hectocotylus of Alloteuthis media, Gulf of Tunis,

preserved. Photograph by R. Young.
2. Tentacles
1. a Tentacular clubs expanded, suckers in four series; two medial
manal series with larger suckers.
3. Head
4. Mantle
1. Mantle long, relatively narrow; posterior end drawn out into narrow,
pointed tail, up to 6 cm in adults.
Figure. Ventral view of A. subulata showing dimorphism in the relative sizes and
shapes of mature males. Top - Small male which we presume to be a sneaker male
(sneakers are known in L. reynauldii - Hanlon, et al., 2002). Bottom - Large male
with elongate tail. Females are approximately the same size and shape as the small
male. Drawings from Naef (1921-3).
5. Fins
1. Fins reach posterior tip of mantle.
2. Fins heart-shaped, lateral angles rounded, posterior borders
concave, extending posteriorly along tail.
6. Photophores
7. Viscera
1. Eggs small.
2. Spermatophore with small cement gland.
Comments:
Alloteuthis is morphologically very similar to Loligo and was considered by Vecchione et al.
(1998) to be a subgenus. However, molecular analyses by Anderson (2000) indicate that
Alloteuthis should be a separate genus. This supports earlier inferences by Naef (1921-23)
and Alexeyev (1989) that the presence of a gladial conus in Alloteuthis should be considered
a generic character. The conus in Alloteuthis would clearly separate the genera but it can be
very difficult to find and in some Alloteuthis specimens may be reduced until it is essentially
absent. The posterior tip of the mantle in Alloteuthis is elongate into a tail-like structure that
tends to be more pointed than in Loligo. This is particularly true for males but identification
of this character state can be subjective in females.
Distribution
Eastern Atlantic Ocean from about 55°-60°N to 20°S.
Alloteuthis africanus Adam 1950

Containing group: Alloteuthis
Introduction
A tropical loliginid of the eastern Atlantic. Maximum mantle length 19 cm in
males, 9 cm in females.
Characteristics
1. Mantle
1. Mantle long and narrow, mantle width 20 - 25% ML in juveniles,
15% in adult females, 5% in adult males
2. Anterior ventral mantle margin squarish in outline.
3. Tail (fins and posterior mantle projection) very long and pointed in
females (37% of dorsal mantle length in juveniles and 58% in
adults) and extremely long and spike-like in males (35% in
juveniles, 73% in adults).
2. Fins
1. Fins oval in outline.
2. Fin width index (width of both fins as a percentage of dorsal mantle
length) 23% in adult females and 10% in adult males.
3. posterior border of fins concave.
3. Buccal crown
1. Buccal lappets without suckers.
4. Arms:
1. Arms very short.
2. Arm suckers with 6 to 10 square teeth on dista1 half, smooth on
proximal half.
3. Left ventral arm hectocotylized by modification of dista1 2/5 of
length; 8 to 11 pairs of normal suckers proximally followed by 2
longitudinal series of elongate papilla that gradually decrease in
size distally.
5. Tentacles
1. Diameters of club suckers of median 2 series 3 times greater than
lateral ones.
2. Sucker rings with 20 to 30 blunt teeth.
Habitat and Biology
Found at depths of 20-100 m, temperatures of 16-26 °C. Reproduction is

apparently year-round. Feeds on small fishes.
Geographical Distribution
Atlantic: from 20 °S to 25 °N.
Other Names for Alloteuthis africanus Adam 1950
• Vernacular Names: En: African squid, Fr: Casseron africain, Sp:

Calamarin africano
Alloteuthis media (Linnaeus 1758)

Midsize squid

Introduction
This small eastern Atlantic species is the type species of the genus Alloteuthis.
Maximum mantle length 12 cm; females grow bigger than males. Length at first
maturity is about 9.5 cm in females, and 5.5 cm in males.
Characteristics
1. Mantle
1. Mantle long, relatively narrow, its posterior end drawn out into a
narrow, pointed tail, up to 6 cm long in adults.
2. Fins
1. Fins heart-shaped, their lateral angles rounded, posterior borders
concave, extending posteriorly along tail.
3. Buccal crown
1. Buccal suckers absent.
4. Arms:
1. Left ventral arm hectocotylized: 10 to 12 (usually 11) normal
suckers in ventral row followed distally by papillae.
5. Tentacles
1. Tentacles long, robust; clubs large, expanded.
2. Large suckers in the 2 median manal series.
Habitat and biology
Found on sandy and muddy grounds. Depth range from the surface to 350 m, usually
between 20 and 200 m. Like other neritic squids, it performs seasonal migrations between
offshore and inshore areas. In the Mediterranean, spawning extends throughout the year, but
two groups of spawners can usually be distinguished by size and time of peak spawning, even
though mixing occurs. Large individuals are encountered in February at depths between 150
and 200 m; from March to April, they start migrating into shallower waters, where spawning
takes place on sandy grounds and to a minor extent, in Posidonia grass beds. The second
group, composed of smaller individuals, migrates onshore in June and July and spawn later in
the year. By late fall, depending on environmental temperatures, they start returning to deeper
waters.
Large males may carry up to 170 spermatophores, large females 8 to 30 eggs. The eggs are
shed in several batches and are encapsuled in rather short gelatinous capsules (each
containing up to 1400 eggs) that are attached to hard objects on the substrate (shells, corals,
stones) in batches of up to 12 capsules.
This squid feeds on crustaceans, molluscs and small fishes.

Longevity is about 1 year in males, 1 1/2 years in females.
Eastern Atlantic: Irish Sea, English Channel (very rare in North Sea) South into
Mediterranean Sea.
Remarks
Recent genetic analyses indicate that A. media and A. subulata may represent
extremes of a morphological gradient in a single species.
Other Names for Alloteuthis media (Linnaeus 1758)
• Synonymy: Loligo marmorae, Loligo parva, Loligo urceolatus

• Vernacular Names: En: Midsize squid, Fr: Casseron bambou, Sp:
Calamarín menor, Local species names: France: Petit encornet; Italy:
Totariello; Monaco: Totanitu; Spain: Luria; UK: Little squid.
Title Illustrations
Scientific
Alloteuthis media
Name
C. F. E. Roper, Michael J. Sweeney and Cornelia E. Nauen. 1984.

Acknowledge Cephalopods of the World. An Annotated and Illustrated
Alloteuthis subulata (Lamarck 1798)

Introduction
Maximum mantle length 20 cm in males, 12 cm in females.
Characteristics
1. Mantle
1. Mantle long and narrow.
2. Anterior ventral mantle margin shallowly curved.
3. Tail long and pointed in adult females (length of posterior extension
plus fins 66% of dorsal mantle length) and very long and spike-like
in adult males (72% of dorsal mantle length).
2. Fins
1. Fin width index (width of both fins as a percentage of dorsal mantle
length) 23% in adult females and 10% in adult males.
2. Posterior borders concave and extending along tail.
3. Buccal crown
1. Buccal lappets without suckers.
4. Arms:
1. Arm length medium to short.
2. Arm suckers with 6 to 10 square teeth on dista1 half, smooth on
proximal half.
3. Left ventral arm hectocotylyzed, with 6 to 8 pairs of normal suckers
proximally, followed distally by 2 longitudinal series of fine papillae.
5. Tentacles
1. Tentacles short, delicate.
2. Clubs small, narrow.
3. Median manal suckers small.
Habitat and Biology
Usually associated with sandy and hard bottoms, occurring at depths down to
200 m. In the North Sea, males and females arrive together in inshore waters in
early summer. The sole spawning season is restricted to June and July. Juveniles
of 2 mm ML hatch after a few weeks; they first occur in plankton samples
toward the end of July; after about 15 to 30 days they shift to the demersal life
habits of the adults; by November, at an age of 3 months (size about 3 cm ML),
they leave the North Sea until the following spring (size about 5 cm ML).
Longevity is between 1 and 2 years. The species feeds on small and juvenile
fishes.
Geographic Distribution
Eastern Atlantic: North Sea and western Baltic Sea south to the Sahara Banks,
extending into the Mediterranean Sea.
Other Names for Alloteuthis subulata (Lamarck 1798)
• Vernacular Names: En: European common squid, Fr: Casseron commun,

Sp: Calamarín picudo, Local species names: Italy: Calamaretto
Title Illustrations
Scientific Alloteuthis
Name subulata
Off Cascais,
Location
Portugal
Specimen
Live Specimen
Condition
Identified By Peter Wirtz
View Dorsal-oblique
© 2007 Peter
Copyright
Wirtz
Scientific
Alloteuthis subulata
Name
Acknowledge C. F. E. Roper, Michael J. Sweeney and Cornelia E. Nauen. 1984.

Cephalopods of the World. An Annotated and Illustrated
Doryteuthis Naef, 1912

Doryteuthis contains eight species placed into three subgenera one of which is
unnamed.
pages.
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Type species. -- Loligo plei Blainville, 1823; species first mentioned and
subsequent designation of Naef (1923:193)].
Introduction
Doryteuthis species are the common inshore squids of American waters. Several species
(e.g., D. pealeii, D. opalescens) are targets of substantial fisheries. The genus is defined by
geography and the absence of specific modifications that define other genera.
Brief diagnosis:
A loliginid ...
• with fins in adults rhomboidal, longer than broad, (length <70% of ML);
mantle elongate, pointed posteriorly; cement body of spermatophore
short.
• with a distribution in waters off the Americas.
Characteristics
1. Arms
1. Hectocotylus:
3. Distal suckers of reduced size, sucker stalks elongated to
form papillae in either dorsal or both dorsal and ventral
series.
2. Tentacles
1. Tentacular clubs expanded, suckers in four series.
3. Head
1. Beaks: Descriptions can be found here: Lower beak; upper beak.
4. Fins
Figure. Dorsal view of D. (Doryteuthis) plei in an aquarium. Photograph by

Roger Hanlon.
5. Photophores
6. Viscera
Comments
The genus is subdivided into three subgenera, two of which are named, D. (Doryteuthis) and
D. (Amerigo). Species of D. (Doryteuthis) have the modified portion of the hectocotylized
arm extending to the arm tip, and the edges of gladius vane are thickened, while in D.
(Amerigo) neither of these conditions hold. Doryteuthis sanpaulensis does not belong in
either of these subgenera and was considered by Vecchione et al. (2005) to be the sole
recognized species in an undescribed subgenus.
Figure. Oral view of the distal region of the hectocotylus of D. (Doryteuthis) plei, 105 mm
ML, off S.E. USA, preserved. Photograph by R. Young.
Although we tend to think of squids as constantly swimming, D. (A.) pealeii is often
observed sitting on the ocean floor or the bottom of an aquarium.
Figure. Side view of D. (Amerigo) pealeii sitting on the substrate in an aquarium.

Photograph by Clyde Roper.
Distribution
American waters of the western Atlantic and eastern Pacific Oceans.
References
Vecchione, M., E. Shea, S. Bussarawit, F. Anderson, D. Alexeyev, C.-C. Lu, T. Okutani, M.

Roeleveld, C. Chotiyaputta, C. Roper, E. Jorgensen and N. Sukramongkol. 2005. Systematics
of Indo-West Pacific loliginids. Phuket Mar. Biol. Cent. Res. Bull. 66: 23-26.
Heterololigo Natsukari, 1984
Heterololigo bleekeri (Keferstein, 1866)


A single species is present in Heterololigo.
Type species. -- Loligo bleekeri Keferstein, 1866 by original designation.
Introduction
H. bleekeri is one of the more common loliginids in Japan.
Brief diagnosis:
A loliginid ...
• of moderately large size (to 40 cm ML) with fins in adults rhomboidal,

longer than broad, not round (length <70% of ML); mantle elongate,
pointed posteriorly; without photophores.
• with unusual hectocotylus.
• with a Northwest Pacific distribution.
Characteristics
1. Arms
1. Arms very short.
2. Arm suckers with blunt teeth around distal ring.
3. Hectocotylus:
3. Distal suckers of reduced size and sucker stalks elongated to
form papillae in dorsal series.
4. Dorsal series of papillae and trabeculae at distal tip form
bicuspid lamelliform flaps separated from ventral series of
suckers by serrated membrane.

Figure. Oral view of the hectocotylus of H. bleekeri, ca. 300 mm ML, Sagami Bay.
Drawing from Sasaki (1929).
Figure. Oral view of the tip of the hectocotylus of H. bleekeri. Enlargement from
drawing above.
2. Tentacles
1. Tentacular clubs narrow
2. Club suckers small, in four series; suckers in transverse row nearly
uniform in size.
Figure. Oral view of tentacular club of H. bleekeri, ca 300 mm ML, mature

male, Sagami Bay. Drawing from Sasaki (1929).
3. Fins
4. Photophores
5. Viscera
2. Spermatophore cement body short
Distribution
Off Japan, excluding northern half of Hokkaido; off southern Korea.
Other Names for Heterololigo bleekeri (Keferstein, 1866)
• Heterololigo Natsukari, 1984
References
Vecchione, M., E. Shea, S. Bussarawit, F. Anderson, D. Alexeyev, C.-C. Lu, T. Okutani, M.

Roeleveld, C. Chotiyaputta, C. Roper, E. Jorgensen and N. Sukramongkol. 2005. Systematics
of Indo-West Pacific loliginids. Phuket Mar. Biol. Cent. Res. Bull. 66: 23-26.
Title Illustrations
Scientific
Heterololigo bleekeri
Name
Location Sagami Bay, Japan
Sasaki, M. 1929. A Monograph of the Dibranchiate Cephalopods of the

Referenc
Japanese and Adjacent Waters. Journal of the College of Agriculture,
e
Hokkaido Imperial University, 20(supplement):357 pages.
Sex Male
Life
Cycle Mature
Stage
View Ventral
Size ca. 30 cm
Loliolus Steenstrup, 1856
Michael Vecchione
Loliolus has six species placed into two subgenera.
pages.
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Type species.-- Loliolus hardwickei (Gray) [Note: according to Lu et al. (1985),

Steenstrup's originally designated type species, L. typus Steenstrup 1856 =
Loligo hardwickei Gray 1849].
Introduction
These are generally small squids with heart-shaped fins.
Brief diagnosis:
A loliginid ...
posterior lobes.
• with ventral crest on hectocotylus formed by fusion of the protective
membrane with the ventral row of papillae.
• with an Indo-Pacific distribution.
Characteristics
1. Arms
1. Arm sucker rings with square plate-like teeth on margin.
Figure. Oral view of large arm III suckers of L. hardwickei, 33 mm ML,

male. The left sucker is from the dorsal series and the right sucker from its
pair in the ventral series. Drawings from Lu, et al. (1985).
2. Hectocotylus:
1. Ventral crest present, formed by fusion of protective
membrane with ventral row of papillae such that original
form of conical papillae completely obscured (see below).

Figure. Top - Aboral-medial view of the hectocotylus of Loliolus (Loliolus)
hardwickei showing the crest that reaches the full length of the arm, male, off
Madras, India, preserved. Photograph by R. Young. Bottom - Oral-lateral
view of the hectocotylus of L. (L.) hardwickei showing the crest, male, 33 mm
ML, Indian Ocean south of Calcutta. Drawing from Lu et al. (1985).
2. Tentacles
3. Mantle
1. Mantle rounded posteriorly; without tail-like elongation.
4. Fins

Figure. Dorsal view of L. (L.) hardwickei, 33 mm ML, male. Drawing from
Lu, et al. (1985).
5. Photophores
6. Viscera
1. Eggs small.
comments
Hectocotylization includes a crest along a portion of the ventral surface of the arm and a
modification of part of the dorsal row of suckers into papillae. Modification of the proximal
portion of the hectocotylus varies between subgenera. In Loliolus (Loliolus) the modified
portion of the hectocotylus occupies the entire length of the arm whereas in Loliolus
(Nipponololigo) some proximal suckers are unmodified.
Figure. Oral view of the hectocotylus of Loliolus (Nipponololigo) japonica showing the
crest restricted to the distal region of the arm, male, Hakkodate fish market. Photograph by R.
Young.

Figure. Oral (top) and medial (bottom) views of a portion of the hectocotylus crest of L.
(N.) japonica, same hectocotylus as pictured above. Photographs by R. Young.
Nomenclature
Steenstrup (1856) erected the genus Loliolus for his new species typus. Lu et al. (1985)
showed that L. typus is a junior synonym for Loligo hardwickei Gray, 1849, making the
latter species the type for the genus Loliolus.
Because the percentage of the arm hectocotylized is considered to be a cline within this group
of species with otherwise similar hectocotyli and arm-sucker dentition, Loliolus and
Nipponololigo were combined by Vecchione et al. (1998) into a single genus that, by
priority, is named Loliolus.
Loliolus was originally erected to include species of small loliginids with heart-shaped fins in
which the entire left ventral arm of the males is modified. However, as discussed by
Brakoniecki (1986), clines exist in the percentage of the arm hectocotylized within species
groups with similar hectocotyli. Thus, this character state is not sufficient for generic status.
The presence, though, of a distinctive character, a ventral crest on the hectocotylus formed by
fusion of the protective membrane with the ventral row of papillae such that the original form
of the conical papillae is completely obscured, indicates a close relationship among these
species (Natsukari, 1983).
Distribution
The species in this genus are all found in the Indo-West Pacific.
Lolliguncula Steenstrup, 1881
Brief squids
Loliguncula contains four species placed in two subgenera.
pages.
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Type species.--Loligo brevis Blainville, 1823, by original designation.
Introduction
Species of Lolliguncula are rather small squids that generally occupy warm, shallow waters
and some invade regions of low salinity.
Brief diagnosis:
A loliginid ...
posterior lobes.
• with a long cement body in the spermatophores.
• without ventral crest on hectocotylus.
• with distribution only in American waters.
Characteristics
1. Arms
1. Sucker rings with square teeth on distal or entire margin.
Figure. Oral view of arm III sucker of L. argus, male, 30 mm ML. Drawing
from Brakoniecki and Roper (1985).
2. Suckers on mid-sections of lateral arms of males not greatly
enlarged.
3. Hectocotylus:
2. Unmodified proximal region of hectocotylus present (in
subgenus Lolliguncula) or absent (in subgenus Loliolopsis).
3. Suckers of modified region reduced, sucker stalks elongated
to form papillae on either dorsal or both dorsal and ventral
rows.
2. Tentacles
3. Mantle
1. Mantle posteriorly rounded; posterior tail-like elongation absent.
4. Fins
1. Fins posterior, broadly rounded posteriorly.
2. Fins wider than long in adults.
5. Photophores
6. Viscera
1. Eggs less than 3mm.
2. Spermatophore cement body long.
Figure. Spermatophore of L. brevis, 90 mm ML, showing the size of the cement body
(amber color). Drawing modified from Hess (1987).
Comments
The cement body of Lolliguncula species, where known, varies roughly between 30-50% of
the spermatophore length. In other loliginid genera with small cement bodies, the cement
body typically is between about 3-15% of the spermatophore lenght.
Members of Lolliguncula (Lolliguncula) have the modified portion of the hectocotylus less
than the entire arm (ie, the proximal portion is not modified) while Lolliguncula
(Loliolopsis) has the hectocotylus modified along the entire arm.

Figure. Ventral view of Lolliguncula (Loliolopsis) diomedeae (holotype of Loliolopsis
chiroctes), preserved. Photograph by R. Young.
Nomenclature
Berry (1929) described a new species that he named chiroctes and erected the genus
Loliolopsis because of the distinctiveness of the new species. Voss (1971) recognized that L.
chiroctes is a junior synonym of Hoyle's Loligo diomedeae but that Berry was correct in
establishing a new genus. Therefore, the type species of Loliolopsis (now a subgenus of
Lolliguncula) is Loliolopsis diomedeae (Hoyle, 1904).
The genus Lolliguncula was established to distinguish L. brevis from the species of Loligo
then known, based on body and fin shape and spermatophore deposition site. These
characters have since proven to exhibit considerable variability, both within and among
species. However, all of the species in this genus are united and distinguished from all other
loliginids by the long cement body of their spermatophores. Variability exists in hectocotylus
morphology, and this led Berry (1929) to erect the genus Loliolopsis and Brakoniecki (1986)
to erect Afrololigo. This variability is extreme in L. diomedeae, but Brakoniecki (1986) has
pointed out the similarities in hectocotylization between this species and the others of
Lolliguncula.
Many of the morphological characters of Afrololigo mercatoris are shared with species of
Lolliguncula, the genus in which it was originally described. However, DNA sequence
analysis by Anderson (2000) supports Brakoniecki's (1986) conclusion based on hectocotylus
morphology that Afrololigo is a distinct genus.
Distribution
These are all American species, with their low-latitude distribution centered at Panama. One
species, L. brevis, is found on the Atlantic side of the Americas whereas three, L.
panamensis, L. diomedeae and L. argus are found on the Pacific side. All of these species
are found in warm, shallow, inshore waters. The genus is noteworthy among cephalopods for
its euryhalinity.
Other Names for Lolliguncula Steenstrup, 1881
• Brief squids
Pickfordiateuthis Voss, 1953
Grass squid
Michael Vecchione
The following four species have been described (one has not been named) in
Pickfordiateuthis.
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Type species.-- Pickfordiateuths pulchella Voss, 1953, by monotypy.

Introduction
These are small (22 mm ML max.), muscular squids that occupy neritic waters often in
association with small patch reefs and seagrass (Voss, 1953). In spite of the accessible
habitat, the squid is rarely captured and little is known of its biology.
Brief diagnosis:
A loliginid ...
• of small size and with posterior fin lobes; fins not joined posteriorly.
• with club suckers on manus in two series.
Characteristics
1. Arms
1. Suckers in two series.
2. Tentacles
1. Club suckers on manus in two series; suckers on dactylus in four
series or in two series.
Figure. Oral view of the tentacular club of P. pulchella, 22 mm ML, female,

holotype, off Florida Keys. Drawing from Voss (1953).
3. Head
4. Fins
1. Fins subterminal.
2. Fins oval with free anterior and posterior lobes.

Figure. Left - Ventral view of the posterior mantle and fins of P. pulchella, 22 mm
ML, holotype, preserved. Photograph by R. Young. Right - Dorsal view of the same
specimen. Drawing modified from Voss (1953).
5. Photophores
1. Absent.
Comments
The general shape varies greatly between species. P. pulchella is broad with relatively large
arms and fins; P. bayeri and P. vossi are slender with small arms and fins. The latter two look
look much like a young Doryteuthis.
Brakoniecki (1996) submerged the family Pickfordiateuthidae into the Loliginidae.
Figure. Dorsal view of P. bayeri in a shipboard aquarium.
Other Names for Pickfordiateuthis Voss, 1953
• Grass squid
References
Brakoniecki, T. F. 1996. A revision of the genus Pickfordiateuthis Voss, 1953

(Cephalopoda; Myopsida). Bull. Mar. Sci. 58: 9-28.
Roper, C.F.E. and M. Vecchione. 2001. Pickfordiateuthis bayeri n. sp., a new species of squid
(Cephalopoda: Loliginidae) from the western North Atlantic Ocean discovered by
submersible. Bull. Biol. Soc. Wash. 10:301-310.
Voss, G. L. 1953. A new family, genus and species of myopsid squid from the Florida keys.
Bull. Mar. Sci. Gulf Carib. 2: 602-609.
Title Illustrations
Scientific Pickfordiateuthis
Name pulchella
Location off Florida
Specimen
Live Specimen
Condition
© 1996 Clyde F. E.
Copyright
Roper
Sepioteuthis Blainville, 1824

Reef squids

Three nominal species are present in Sepioteuthis.
pages.
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Type species. -- Loligo sepioidea Blainville, 1823 by original designation.

Introduction
Sepioteuthis is distinctive; unlike in many loliginid genera, controversy does not exists over
which species should be included (Vecchione et al., 1998). These loliginids have a rather
broad, posteriorly rounded mantle and fins that extend nearly the full length of the mantle.
This gives them a Sepia-like appearance. They also lay eggs that are much larger than those
of other loliginids. They usually are found in shallow tropical or subtropical regions.
Brief diagnosis:
A loliginid ...
• with fins in adults that occupy >85% ML; combined fin shape broadly
elliptical.
Characteristics
1. Arms
1. Arm sucker rings with pointed teeth around entire margin.
2. Hectocotylus:
3. Distal suckers reduced, sucker stalks elongated along
modified portion of arm to form papillae on both dorsal and
ventral rows.
2. Tentacles
1. Tentacular clubs expanded, with suckers in four series.
3. Head
1. Beaks: Descriptions can be found here: Lower beak; upper beak.
4. Mantle
1. Posterior mantle not elongated into tail-like structure.
5. Fins
1. Extend nearly full length of mantle (>85% of ML) except in very
young squid.

Figure. Dorsal view of S. lessoniana, adult showing the large fins. Drawing
from Roper, et al. (1984)
6. Photophores
7. Viscera
1. Eggs larger than 5 mm.
2. Eggs lie in single, straight series within egg strand.

Relationships among species are uncertain. Segawa et al. (1993) suggest that S.
lessioniana represents a species complex.
Distribution
Because of their association with coral reefs, the common name for these species is "reef
squids". The distribution of the genus generally follows that of coral reefs, warm, clear,
tropical and subtropical waters, but coral reefs are not a habitat requirement for all of these
squids. For example, S. lessoniana is common around the southern part of Honshu and
Kyushu in Japan, but is not associated with coral reefs in those areas and occupies temperate
waters of southern Australia. There is one species, S. australisS. sepioidea, in the western
Atlantic; the remaining species are found in Indo-West Pacific waters.
Other Names for Sepioteuthis Blainville, 1824
• Reef squids
References
Nesis, K. N. 1982. Abridged key to the cephalopod mollusks of the world's ocean. 385+ii pp.
Light and Food Industry Publishing House, Moscow. (In Russian.). Translated into English
by B. S. Levitov, ed. by L. A. Burgess (1987), Cephalopods of the world. T. F. H.
Publications, Neptune City, NJ, 351pp.
Segawa, S., S. Hirayama and T. Okutani. 1993. Is Sepioteuthis lessoniana in Okinawa a
single species? Pp. 513-521. In: T. Okutani, R. K. O'Dor and T. Kubodera (eds). Recent
Advances in Cephalopod Fisheries Biology. Tokai University Press, Tokyo.
Vecchione, M., T. F. Brakoniecki, Y. Natsukari and R. T. Hanlon. 1998. A provisional

generic classification of the family Loliginidae. Smithson. Contr. Zool., 586
Title Illustrations
Scientific Sepioteuthis
Name sepioidea
Specimen
Live Specimen
Condition
View Anterodorsal
Copyright © Roger Hanlon
•
o Containing Groups
o Loliginidae
o Myopsida
o Decapodiformes
o Coleoidea
o Cephalopoda
o Mollusca
o Bilateria
o Animals
o Eukaryotes
o Life on Earth
•
•
o Other Loliginidae
o Loligo
o Afrololigo mercatoris
o Alloteuthis
o Doryteuthis
o Heterololigo bleekeri
o Loliolus
o Lolliguncula
o Pickfordiateuthis
o Sepioteuthis
o Uroteuthis
•
•
Uroteuthis Rehder, 1945

Michael Vecchione
The following fourteen nominal species, divided into three subgenera, are presently included
in Uroteuthis; however this generic and subgeneric arrangement is controversial.
pages.
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Type species.--Uroteuthis bartschi Rehder, 1945, by original designation.
Introduction
These common inshore squids of the Indo-West Pacific are targets of important large-scale
and artesanal fisheries. Uroteuthis spp. are the only squids in the Loliginidae that possess
photophores.
Brief diagnosis:
A loliginid ...
• with a pair of photophores on the ink sac.
Characteristics
1. Arms
1. Proximal margin of sucker rings with semicrescent plate; distal
margin with square teeth.
2. Hectocotylus:
1. Two rows of papillae present; ventral crest absent.
2. Tentacles
3. Mantle
1. Mantle generally elongate and pointed posteriorly.
4. Fins
1. Fins in adults generally rhomboidal and longer than broad, tapering
posteriorly.
2. Fins may or may not reach posterior tip of mantle.
5. Photophores
1. Pair of organs on ventral surface of ink sac; intestine separates
members of the pair.
Figure. Ventral view of the visceral photophores (arrow) of (probably) Uroteuthis

edulis, Japanese fish market. Photograph by Sabrina Pankey.
6. Viscera
1. Eggs small.
Comments
Uroteuthis (Uroteuthis) is distinguished from the other two subgenera by a greatly elongate
"tail" with "lateral" fins which to not extend to the tip of the tail. In Uroteuthis (Photololigo)
and Uroteuthis (Aestuariolus) the elongation of the posterior mantle is variable but not tail-
like and when the mantle is elongate, the fins extend along the sides to the posterior tip.

Figure. Dorsolateral view of U. (U.) bartchi, holotype, 199 mm ML, male, preserved,
Philippine waters at 06°05'N, 121°00'E, showing elongate tail and lateral fins. The bend in
the tail is an artifiact of preservation. Photograph by R. Young.
Uroteuthis (Photololigo) differs from Uroteuthis (Aestuariolus) in that the modified portion
of hectocotylus is less than entire arm; that is, the proximal portion is not modified. In
Uroteuthis (Aestuariolus) the entire arm is modified. The structure of the photophores also
differs somewhat between the two subgenera.
Figure. Top - Oral view of the hectocotylus of Uroteuthis (Photololigo) edulis, 73 mm ML,
male, preserved, East China Sea at 21°42'N, 114°50'E, preserved. Note the normal suckers in
the proximal region of the arm. Bottom - Oral view of the hectocotylus of Uroteuthis
(Aestuariolus) nocticula , 53 mm ML, male, preserved, Bass Strait, Australia at 37°51'S,
144°56'E. Note that only modified suckers are in the proximal region of the arms.
Photographs by R. Young.

Figure. Dorsal view of Uroteuthis (Aestuariolus) noctiluca, paratype, preserved. Photograph
by R. Young.
Nomenclature
From Vecchione et al., 1998:
The pair of photophores on the ventral surface of the ink sac in all of these species is perhaps
the strongest indication of a close relationship among species to be found in this family. This
relationship is further supported by similarities in the hectocotylus and arm-sucker dentition.
These characters led Natsukari (1984) to establish the genus Photololigo for the Indo-West
Pacific species with photophores, formerly assigned to either Loligo or Doryteuthis.
However, Uroteuthis bartschi also has similar photophores, a similar hectocotylus and arm
suckers, and an Indo-West Pacific distribution. The elongation of the posterior mantle, which
was given generic status in the description of Uroteuthis, is the extreme example in a cline of
mantle elongation, similar to that found in the eastern Atlantic where Alloteuthis is the
extreme example. This elongation is so extreme in Uroteuthis as to cause a discontinuity in
the cline. It thus warrants separate taxonomic status, but the sole recognized species in this
taxon possesses all of the generic characters of Photololigo. Thus, it should be included in the
genus that Natsukari described. Unfortunately, priority therefore goes to Uroteuthis as the
name of the genus, rather than the more descriptive Photololigo. Aside from the photophores,
Uroteuthis s.s. and Alloteuthis can be separated based upon position of the fins, lateral versus
terminal respectively, and tentacular ring dentition. Therefore, elongation of the mantle end
in these two taxa appears to be analogous, not homologous.
In a revision of Loliolus, Lu et al. (1985) described a new species: L. noctiluca. They

included this species in Loliolus because its hectocotylization was a modification of the
entire length of the arm, the original diagnostic character of the genus Loliolus. However, the
hectocotylus of L. noctiluca is otherwise very different from those of the other species of
Loliolus in that it lacks a ventral crest and both the dorsal and ventral rows of suckers are
modified into papillae. The hectocotylus of L. noctiluca is thus similar to those of the
Uroteuthis and Photololigo species except that more of the arm is modified, making it the
extreme in a cline of the percentage of arm length that is modified by hectocotylization.
Furthermore, L. noctiluca was so named because of structures on the ink sac believed to be
photophores. These characters, together with similarity in arm-sucker dentition, indicate that
this species should be included in the subgenus Photololigo of the genus Uroteuthis.
Yeatman (1993) has presented evidence that some species of Uroteuthis (Photololigo) are
actually species complexes.
Distribution
The Indo-West Pacific distribution of all of these species supports the contention
that they comprise a natural, closely related species group.
References
Alexeyev, D.O. 1991. Systematics, Phylogeny, Distribution, Biology, and Prospects for
Fishery of the Squid Suborder Myopsida. Candidate in Biological Sciences Thesis, Institute
of Oceanography of the Russian Academy of Sciences, Moscow. 129 pages. (In Russian).
Alexeyev, D.O. 1992. The Systematic Position of Bioluminescent Squids of the Family
Loliginidae (Cephalopoda, Myopsida). Zoologeskiy Zhurnal, 71(11):12-23. (In Russian).
Lu, C.C., C.F.E. Roper, and R.W. Tait. 1985. A Revision of Loliolus (Cephalopoda;
Loliginidae), Including L. noctiluca, a New Species of Squid from Australian Waters.
Proceedings of the Royal Society of Victoria, 97(2):59-85.
Natsukari, Y. 1984. Taxonomical and Morphological Studies on the Loliginid Squids-- IV.
Two New Genera of the Family Loliginidae. Venus, The Japanese Journal of Malacology,
43(3):229-239.
Vecchione, M., T. F. Brakoniecki, Y. Natsukari and R. T. Hanlon. 1998. A Provisional

Generic Classification of the Family Loliginidae. Smithson. Contr. Zool., 586
Yeatman, J. 1993. Genetic and Morphological Aspects of Australian Photololigo spp.

(Loliginidae: Cephalopoda). Ph.D. Thesis, James Cook University, North Queensland,
Australia.
Title Illustrations
Scientific
Uroteuthis chinensis
Name
Location off Australia
© 1996 Mark D.
Copyright
Norman
Oegopsida Orbigny, 1845

Richard E. Young and Michael Vecchione

pages.
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Introduction
Among cephalopods, oegopsid squids dominate the pelagic, oceanic environment. Diversity
at the family and generic levels is very high with 24 families and 69 genera. Although
pelagic, some species are consistently associated with and some occasionally sit on the ocean
floor.
Diagnosis
Decapodiforms ...
• without cornea covering eyes.

• with a gladius.
Characteristics
1. Arms
2. Tentacles
1. Tentacular club usually with carpal (= proximal) locking-apparatus.
Figure. Oral view of the club base of Cycloteuthis serventyi. The red line
encircles the suckers and knobs of the carpal locking-apparatus. Drawing
modified from Young and Roper (1969).
2. Suckers without circularis muscles (see Sepioidea page).

3. Buccal crown
1. Buccal supports lack suckers.
4. Head
1. Head without tentacle pockets.
2. Eyes without corneal membranes covering lenses.
Figure. Dorsal view of the head of Pyroteuthis addolux with the protruding
eye lenses directly exposed to sea water. Photograph by R. Young.
5. Funnel
6. Mantle
1. Mantle locking-apparatus, with rare exceptions, reaches anterior
mantle margin.
7. Fins
1. Fins usually joined posteriorly and usually without posterior lobes.

Figure. Ventral view of mantle and fins of Ancistroteuthis lichtensteini, 175
mm ML, Mar-Eco cruise, R/V G. O. SARS, North Atlantic. Photograph by R.
Young.
8. Shell
1. Shell a gladius.
9. Viscera
1. Gills with branchial canal.

(1921-23).
2. Oviducts paired.
3. Females without accessory nidamental glands.
10.Eggs
1. Eggs in pelagic egg masses.
Comments
Comparisons of the major groups of the Oegopsida
Buccal Carpal
Funnel locking- Gladius Anterior fin
connective locking-
cartilage conus attachment
to arms IV apparatus
1°
Architeuthidae Dorsal Straight Yes Mantle
conus
Brachioteuthida 2°
Ventral Straight Yes/No Gladius
e conus
Variable,
Chiroteuthid usually oval, 2°
Ventral No Gladius
families often with conus
knobs
2°
Cranchiidae Ventral Fused Yes conus / Gladius/Mantle
No
2°
Cycloteuthidae Ventral Triangular Yes conus / Gladius
No
Enoploteuthid 1°
Dorsal Straight Yes Gladius
families conus
1°
Gonatidae Ventral Straight Yes Gladius
conus
Histioteuthidae
Dorsal Straight Yes No Gladius
families
Lepidoteuthidae 2°
Ventral Straight No Gladius
families conus
1°
Neoteuthidae Dorsal Straight Yes Mantle
conus
Ommastrephida 1°
Dorsal Inverted T Yes/No Gladius
e conus
Onychoteuthida 1°
Ventral Straight Yes Gladius
e conus
Thysanoteuthid
Ventral Lazy T Yes No Mantle
ae
A key to the families of the Decapodiformes can be found here.

Title Illustrations
Pyroteuthis margaritifera, Promachoteuthis sp., Magnapinna ?,

Scientific
Helicocranchia pfefferi, Joubiniteuthis portieri, Onychoteuthis sp.,
Name
Todarodes sagittatus
Acknowledge J. portieri photograph by Frank Parish; Magnapinna (?)

ments photograph from N.U.R.C., others photographed by R. Young.
This media file is licensed under the Creative Commons

Image Use
Attribution-NonCommercial License - Version 3.0.
Copyright © 2004 Richard E. Young
Sepioidea Naef, 1916

pages.
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Introduction
We place three families in the sepioid clade: Sepiidae, Sepiadariidae and Sepiolidae. The
latter two families are related and placed in the suborder Sepiolida. Two additional families
(Idiosepiidae, Spirulidae) have often been included in the Order Sepioidea. However
considerable uncertainty exists concerning the relationships of the Idiospiidae in general and
the closeness of the relationship of the Spirulidae to the sepioid families. Members of the
sepioidea are mostly neritic and upper slope benthic species although one group
(Heteroteuthinae) is pelagic.
Brief diagnosis:
A decapodiform ...
• with corneal membranes covering eye lenses.

• without branchial canals in gills.
• with circularis muscles in suckers.
Characteristics
1. Arms
1. Suckers with circularis muscles.
2. Tentacles
1. Tentacular club without proximal (= carpal) locking-apparatus.
2. Suckers in more than four series (except in some Sepiolidae).
3. Suckers with circularis muscles.
Figure. Sections through the club suckers. Left - Sepia officinalis, arrow
points to circularis muscle which encircles the sucker. Middle -
Ommastrephes bartramii. Note absence of a circularis muscle. Right -
Sepia officinalis, section at right angles to left figure, circularis muscle in
blue; the asymmetry in the muscle is revealed by the angle of section.
Histological sections made by Barbara Littman; photographed by R. Young.
Drawing modified from Naef (1921-23).
3. Buccal crown
1. Buccal supports with or without suckers.
4. Head
1. Head with tentacle pocket.
2. Eyes with corneal membranes covering lenses.
3. Eyes with secondary (= ventral) eyelid (except in some Sepiolidae).

Figure. Lateral view of head of Rossia sp., 19 mm ML, preserved, showing
secondary eyelid covering the ventral region of the transparent cornea.
5. Funnel
1. Funnel with lateral adductor muscles (except in some Sepiolidae).
Figure. Ventral view of mantle cavity of Semirossia tenera (Sepiolidae).

Photograph by M. Vecchione.
6. Mantle
1. Mantle locking-apparatus does not reach anterior mantle margin
(see short arrow in above photograph which marks the anterior end
of the mantle locking-apparatus) except in some Sepiolidae.
2. Interstellate connective absent (ie, no direct nerve connective
passes directly from one stellate ganglion to the other).
7. Fins
1. Fins completely separate from one another; usually with posterior
lobes.
8. Shell
1. Shell a flattened phragmocone (=cuttlebone), a gladius or absent.
9. Viscera
1. Gills without branchial canal.

(1921-23).

10.Eggs
1. Eggs, where known, attached to substrate singly or in unorganized
groups.
Synapomorphies
Many of the above characteristics of the Sepioidea are shared with the Myopsida. The
characters that unite the Sepiidae and the Sepiolida and are considered to be apomorphic
(newly derived) characters are (characters 3 and 5 seem to be independently derived in a few
Myopsida):
1. Eyes with secondary eyelids.

2. Funnel with lateral adductor muscles.
3. Mantle locking-apparatus that does not reach the mantle margin.
4. Absence of an interstellate connective.
5. Gills without a branchial canal
Title Illustrations

Scientific Sepioloidea lineolata, Sepia apama,
Name Iridoteuthis iris
Copyright © 2004 Mark Norman, Thomas burch
Spirulida Haeckel, 1896, Spirulidae Owen, 1836
Spirula spirula Linnaeus, 1758

Richard E. Young
The order Spirulida contains a single living species.

Introduction
Spirula spirula is a small, muscular species (45 mm ML) found in mesopelagic waters of the
tropical open ocean. In the title photographs, the skin of the mantle is missing. The intact
mantle is covered with regularly aligned collagen fibers that produce a silvery sheen, as seen
on the head in the title photographs (Herring et al., 1981). Spirula carries an unusual internal
shell that is calcareous and has the shape of a horn coiled in a single plane without the coils
touching one another (open planispiral).
Figure. Left - Side view of the shell of Spirula. Photograph by R. Young of a shell found on
a Florida beach. Right - Cut-away view of the shell provided by ray tomography showing the
phragmocone with septa and siphuncle. Photograph taken by the atomic and nuclear physics
group from the University of Fribourg in Switzerland, and provided by Hans Ueli Johner.
The shell, which retains the phragmocone and siphuncle of its distant ancestors, is used as a
buoyancy device. The posterior position of the shell within the body causes the animal to
generally orient vertically with the head downward. The unusual general appearance of
Spirula with a narrow arm crown, bulging eyes, the peculiar structure of the mantle, the
transverse orientation of the fins and the presence of the coiled shell makes this species very
different in appearance from all other cephalopods.
The large posterior guard-like sheath of fossil relatives of Spirula seems to be designed to
function as a counterweight to maintain the animal in a horizontal position. Such an
orientation is particularly important for a bottom-associated animal that swims just above the
ocean floor (Naef 1921-23). Presumably the ancestors of Spirula were bottom associated and
some remnants of this behavior apparently remains in their life history and distribution
(Young, et al., 1998). A small remnant of the sheath exists on the Spirula shell and a remnant
of the ancestral habitat remains in Spirula's apparent benthic spawning (Young, et al., 1999).
Brief diagnosis:
A decapodiform ...
• with a calcareous shell that is a phragmocone and has the shape of a
coiled horn.
Characteristics
1. Arms
1. Arms with suckers in four series.
2. Both arms IV hectocotylized.
2. Tentacles
1. Tentacular clubs without proximal locking apparatus.
2. Tentacular clubs with suckers in 16 series; not divided into manus
and dactylus.
3. Club suckers without circularis muscles.
Figure. Oral view of tentacular club of S. spirula. Drawing from Chun (1910).
3. Head
2. Eyes without corneas.
4. Funnel
5. Mantle
1. Mantle locking-apparatus does not reach anterior mantle margin.
6. Fins
1. Fins separate, terminal and lie in a plane nearly transverse to body
axis.
2. Fins with posterior lobes.
7. Photophores
1. Large photophore at posterior end of body.
8. Shell
1. Calcareous shell a phragmocone, curved ventrally in open
planispiral; round in cross section and possessing a ventral
siphuncle and transverse septa.
9. Viscera
1. Gills without brachial canal.
4. Radula absent.
5. Ink sac reduced.
6. Anal flaps absent.
7. Digestive gland paired; esophagus passes between members of the
pair.
Nomenclature
A list of all nominal genera and species in the Spirulidae can be found here. The list includes
the current status and type species of all genera, and the current status, type repository and
type locality of all species and all pertinent references.
Behavior
Spirula is able to withdraw its head and arms completely within the mantle; the mantle
opening can then be closed by folding over the large dorsal and ventrolateral extensions (=
flaps) of the mantle margin (Bruun, 1943). The photophore at the posterior end of the body is
known to glow for hours at a time (Schmidt, 1922). When swimming slowly downward, head
first, the terminal fins are pointed upward (i.e. posteriorly) and move with a rapid "waving or
fluttering motion" that propels the animal downward (Bruun, 1943).
Virtually all cephalopods have mobile irises but for an oceanic, pelagic decapodiform the
extreme contraction seen below in Spirula is unusual.
Figure. Views of the iris of Spirula spirula, 23 mm ML, western North Atlantic. Left -
Dorsal view. Right - Dorsal-oblique view. Note the wide-open eyelid and the strongly
contracted iris leaving a tiny pupil. Photogaphs by M. Vecchione.
Life History
Mature ovarian eggs are 1.7 mm in the longest diameter (Bruun, 1943). Bruun (1943)
suggests that the eggs are laid on the ocean floor and the capture of very small animals in
deep water supports this idea (Bruun, 1943; Clarke, 1970). Clarke (1970) described a small
squid with a mantle length of just over 2 mm and Bruun (1943) illustrated a similar squid that
had a shell with just two chambers. These squid are, or are very close to being, hatchlings.
The eyes in these individual are small and the buccal mass is very large with the beaks
extending beyond the short arms. Note that only three arm pairs are present.
Figure. Views of paralarvae of Spirula spirula. Left - Ventral view. Drawing from Clarke
(1970). Right - Drawing from Bruun (1943).
The shift in size frequency distribution over time in the region of the Canary Islands suggests
to Clarke (1970) that this cephalopod has a life-span of 18-20 months. The ventral arms of
the male are hectocotylized: They are much longer and thicker than the other arms (see title
photograph) and possess a variety of unusual flaps and papillae but lack suckers.
Distribution
Vertical distribution
Clarke (1969) found that Spirula occurs at depths between 550 and 1000 m during the day
(peak distribution is at 600-700 m) and at night migrates into the upper 100-300 m (peak at
200-300 m).

Figure. Graph of 70 Spirula spirula captured with opening/closing nets near the Canary
Islands, eastern North Atlantic (Clarke, 1969). Yellow circles- Day captures. Blue triangles-
Night captures. Oversized symbols represent two captures.
Most captures come from the tropical Atlantic and tropical IndoWest Pacific Oceans. Bruun
(1943) found that Spirula is distributed mostly over the slopes of continents or islands where
the bottom depth is between 1000 and 2000 m. The most numerous captures came from the
vicinity of oceanic islands. Indeed, this animal was the most abundant midwater cephalopod
collected in the region of the Canary Islands by Clark (1969).
Other Names for Spirula spirula Linnaeus, 1758
• Spirulida Haeckel, 1896

• Spirulidae Owen, 1836
References
Bruun, A. F. 1943. The biology of Spirula spirula (L.). Dana-Report No. 24. 49pp.
Clarke, M. R. 1969. Cephalopoda collected on the SOND cruise. J.Mar. Biol. Ass. U.K., 49:
961-976.
Clarke, M. R. 1970. Growth and development of Spirula spirula. Jour. Mar. Biol. Ass. U. K.
50: 53-64.
Herring, P. J., M. R. Clarke, S. von Boletzky and K. P. Ryan. 1981. The light organs of
Sepiola atlantica and Spirula spirula (Mollusca: Cephalpooda): bacterial and intrinsic systems
in the order Sepioidea. J. Mar. Biol. Ass. U.K., 61: 901-916.
Naef, A. 1922 -- Die Fossilen Tintenfische. Gustav Fischer, Jena. 322 pp.
Schmidt, J. 1922. Live specimens of Spirula. Nature, 110, No. 2271.
Young, R. E., M. Vecchione and D. Donovan. 1998. The evolution of coleoid cephalopods
and their present biodiversity and ecology. South African Jour. Mar. Sci., 20: 393-420.
Title Illustrations
Scientific
Spirula spirula
Name
Sex Male
View lateral
© 1998 Martina A. Compagno Roeleveld (ca.

Copyright
1946-2006)
Refere from Chun,

nce 1910
Sex Female
View dorsal
Locatio
western North Atlantic
n
View Side
Size 23 mm ML
Image This media file is licensed under the Creative Commons Attribution-
Use NonCommercial License - Version 3.0.
Copyrig
© 2004 Michael Vecchione
ht
Octopodiformes Berthold and Engeser,

1987
Vampire Squid and Octopods
Richard E. Young, Michael Vecchione, and Dirk Fuchs
pages.
close box
Containing group: Coleoidea
Introduction
The Octopodiformes consists of a large group of living cephalopods, the Octopoda which
contains over 200 species, the Vampyroteuthidae which contains a single species (a
phylogenetic relict) and a large variety of extinct cephalopods in the Prototeuthidina and
Loligosepiina. Information on the living and extinct forms are treated separately below.
Characteristics
See separate sections on living and extinct Octopodiformes below.
Characteristics of the living Octopodiformes

Authors: R. Young and M. Vecchione
1. Arms
1. True arms II modified as retractile filaments or absent.
2. True arms IV unmodified.
3. Suckers radially symmetrical.
4. Horny rings in suckers absent.
5. Functional arms IV united by web (absent in some argonautoids).
2. Buccal crown
1. Buccal crown absent.

Figure. Oral view of the region surrounding the buccal mass of
Vampyroteuthis infernalis showing the absence of a buccal crown.
3. Head
1. Head fused to mantle dorsally.
2. Outer statocyst capsule present.
3. Superior buccal lobes adjacent (fused at edges) or completely fused
to posterior buccal lobes.
4. Inferior frontal lobe system of brain present or insepient.
4. Photosensitive vesicles
1. Lie outside of cephalic cartilage: dorsal to funnel (Vampyromorpha)
or on stellate ganglia in mantle cavity (Octopoda).
5. Fins
1. When present, with cartilagenous axial support (cartilage lost in
subadult Vampyroteuthis).
6. Shell
1. Shell a gladius, cartilage-like fin support, pair of stylets, or absent.
7. Viscera
1. Nidamental glands absent.
2. Crop usually present.
3. Oviducal glands radially symmetrical.
4. Digestive gland duct appendages lie "outside" nephridial coelom.
5. Nephridial coeloms separate.
Origin of the Octopoda

Drawing modified from Young, Vecchione and Donovan, 1999.
Naef (1923) felt that the origin of the Octopoda would "forever be obscure." But Naef was
unaware of the significance of Vampyroteuthis which was believed to be a cirrate octopod at
the time. We now know that the Vampyromorpha is the sister group of the Octopoda and that
Vampyroteuthis provides telling clues to the origin of the octopods. Young, et al. (1999)
suggest two possible methods for this origin as illustrated here. In one (benthic route) the
pelagic ancestor becomes benthic in a horizontal attitude (similar to the sepiolids today) and
subsequently the arms and mouth rotate under the head. In the other alternative (pelagic
route) the pelagic ancestor develops an "oral" orientation (i.e. laterally spread arms) here
pictured with the oral end downward (as in octopods today) and subsequently becomes
benthic with the oral end down.
Evidence for the latter route is found in the structure of the brain. Octopods have a series of
lobes in the brain, the inferior frontal lobe system, that is derived from the posterior buccal
lobe. These lobes process complex chemotactile information from the arms. Vampyroteuthis
has an "insipient" inferior frontal lobe system (J. Young, 1977). This bathypelagic animal
apparently has a relatively advanced system for processing chemotactile information from the
arms, one that surpasses that of shallow-living benthic decapods. Apparently Vampyroteuthis
uses its arms in an unusual manner. Young, et al. (1999) suggest that like its immediate
ancestor (a "pre-octopod"), associates with pelagic jellyfish or tunicates and uses its arms and
suckers to adhere to and/or explore the surfaces of these gelatinous animals. That is, it has an
oral orientation for contact with surfaces. The pre-octopod, then was pre-adapted for settling
on the ocean floor in an oral-end down configuration. This oral orientation enabled octopods
to become the effective crawling animals that most are today. However the manner in which
Vampyroteuthis actually uses its arms and its possible association with gelatinous animals
has yet to be demonstrated.
Young, et al. (1999) argue, therefore, that the benthic habitat was the primitive one within the
Octopoda and that pelagic species, which comprise nine of the eleven octopod families, are
secondarily pelagic. In the Cirrates, these authors point out the compaction of viscera, loss of
jet propulsion, presence of a fully-formed inferior frontal lobe system, single oviduct, fusion
of the head and mantle and reduction of the shell as evidence of a quasi-benthic ancestry. In
the Incirrates, they point to the presence of corneas (or their remnants), absence of shell, fins
and cirri, fully formed inferior frontal lobe system, probably stalked chorions and brooding as
evidence of a fully-benthic ancestry.
J. Z. Young (1989), on the basis of anatomical differences, removed the Cirrata from its
traditional place in the Octopoda and put it in its own order, Cirroctopodida. Young and
Vecchione (1996), however, using cladistic analyses demonstrated that the inclusion of the
Cirrata and Incirrata in the Octopoda is justified. That is, the order Octopoda, including the
suborders Cirrata and Incirrata is a monophyletic group. This conclusion was more recently
strengthened by the inclusion of additional characters in the analyses (Young & Vecchione,
2002).
Nomenclature
A variety of names have been proposed for this group (e.g., Octobrachia, Fiorini, 1981;
Octopodiformes, Berthold and Engeser, 1987; Vampyromorphoidea, Engeser and Bandel,
1988; Vampyropoda, Boletzky, 1992). Young, et al.(1999) concluded that Octopodiformes
was the most appropriate.
Characteristics of the extinct Octopodiformes

Author: Dirk Fuchs
Rachiteuthis donovani. © Dirk Fuchs

Gladius morphotypes of fossil Octopodiformes. © Dirk Fuchs
Vampyroteuthis infernalis, as the single living species of the Vampyromorpha, clearly

indicates with its well-developed gladius that both the unpaired shell vestige of the Cirrata
and the paired stylets of some Incirrata morphogenetically might have evolved from a gladius
of an ancient group of Octopodiformes (Haas 2002, Bizikov 2004, Fuchs et al. 2009).
Cephalopod gladii are well-known from the Mesozoic fossil record. In general, fossil gladii
consist of an anteriorly diverging median field, hyperbolar zones and lateral fields (= “wings”
or “vanes” in other terminologies). As can be observed from growth increments, fossil gladii
grow isometrically. Compared to modern gladii, the median field of fossil gladii is thicker
and wider (except in the families Muensterellidae and Palaeololiginidae that exhibit a rachis-
like median field). Rostrum-like or septal-like structures as present in many modern gladii
(see Toll 1998 or Bizikov 2008) are unknown in fossil gladii.
Among fossil gladii, three different morphotypes can be recognized: a prototeuthidid, a

loligosepiid and a teudopseid gladius. Whereas the relationships between the Prototeuthidina,
the Loligosepiina, and the Teudospseina are still uncertain, the general gladius morphology
within these three subgroups is remarkably uniform (in contrast to the extremely high
morphological plasticity in modern gladii; see Toll 1998), i.e. only three main evolutionary
pathways are distinct. Accordingly, newly discovered gladii can be easily ascribed to one of
the three subgroups.
The gladius is hence the most important systematic character complex, although preservation
of soft tissues is very common in so-called Lagerstaetten (La Voulte-sur-Rhone, Solnhofen,
Nusplingen, Hakel, Hadjoula). Soft-part characters such as the shape of the fins or the length
of the arms revealed a high morphological variability even on the species-level.
The diversity, taxonomy, morphology and phylogeny of Mesozoic gladius-bearing coleoids

has been recently revised by Fuchs et al. (2003), Fuchs (2006 a, b, c, 2007, 2009, in press),
Fuchs et al. (2007a, b, in press); Fuchs & Schultze (2008) and Fuchs & Weis (2008; 2009, in
press).
Relationships of extinct Octopodiformes to other Extinct Coleoid Squids

Phragmoteuthis bisinuata. © Dirk Fuchs
Figure: Head and arm crown of Leptotheuthis gigas. © Dirk Fuchs
The Phragmoteuthida is a group of Triassic-Jurassic belemnoids with a breviconic

phragmocone, relatively few chambers, a weakly developed guard, and a broad three-lobed
proostracum (Jeletzky 1966, Donovan 2006). It is widely accepted that the ten-armed
phragmoteuthidids are closely related to the Mesozoic gladius-bearing coleoids (Jeletzky
1966, Donovan 1977, Donovan & Toll 1988, Doyle et al. 1994, Fuchs 2006a). The loss of
secreting calcareous shell material in a phragmoteuthidid subgroup would produce a purely
organic structure that is almost identical with the gladius of early Jurassic loligosepiids.
Since some gladii from different subgroups appear similar to those of living teuthoids, they
have been long time grouped as “fossil teuthids”. However, this systematic position has been
first doubted by Bandel & Leich (1986), because preserved arm crowns in association with
gladii from the Late Jurassic Limestones of Solnhofen (Germany) never show more than
eight arms. Earlier illustrations i.e. by Naef (1922), showing ten arms including a pair of
ventrolateral tentacles have been proved to be wrong. Apart from this, the Loligosepiina
exhibit a type of gladius that can be simply compared with Vampyroteuthis. Indeed, despite
dozens of well preserved arm crowns of different species from different subgroups, different
localities and different stratigraphic levels, an unambiguous pair of ventrolateral tentacle-like
arms is still unknown from the fossil record (the presumed tentacle reported in Vecchione et
al. 1999 is regarded as ambiguous; see Haas 2002 and Fuchs 2006a).
Although the “Octopodiformes-Theory” has been meanwhile well-established (Engeser 1988,

Haas 2002, Bizikov 2004, Fuchs 2006a), the systematic position of those gladius-bearing
Mesozoic groups is nevertheless still under heavy debate. Supporters of the “Teuthoid-
Theory” argue that the tentacles might have been lost before the dead squid touched the sea
floor or that “fossil teuthids” would have lost their tentacles during life time as e.g. living
Octopoteuthididae do. For supporters of the “Octopodiformes-Theory”, these arguments are
not convincing, because among dozens of well-preserved arm crowns from various families
there should be at least one specimen that preserved nine or ten arms (or at least a modified
arm in ventrolateral position).
Considering Mesozoic gladius-bearing coleoids as Octopodiformes means that unambiguous

teuthoids are virtually unknown from the fossil record (with the exception of some teuthid-
like statoliths from Cenozoic deposits).
Other Names for Octopodiformes Berthold and Engeser, 1987
• Vampire Squid and Octopods
References

Mollusc A

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Mollusca

Snails, clams, mussels, squids, octopi, chitons, and tusk

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• Atlantidae Rang, 1829

Atlantidae Rang, 1829

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• Atlanta Lesueur, 1817

• Carinariidae Blainville 1818

Pterotracheidae Rafinesque, 1814

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• Firoloida Lesueur 1817

Cephalopoda Cuvier 1797

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Containing group: Mollusca

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• Class: Cephalopoda Cuvier, 1797

 Fam: Cranchiidae Prosch,

 Fam: Gonatidae Hoyle 1886

 Fam: Neoteuthidae Naef,

 Order: Myopsida Naef, 1916

 Order: Sepioidea Naef, 1916

 Order: Spirulida Haeckel, 1896

 Fam: Idiosepiidae Fischer,

Nautilidae Blainville 1825

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Information on FOSSIL neocoleoids can be found under Octopodiformes

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1. Shell located internally.

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* presence not established in Belemnoidea.

Many of these characters have been secondarily lost in various taxa.

1. Presence of an outer statocyst capsule.

Other Names for Coleoidea Bather, 1888

Decapodiformes Leach, 1817

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Containing group: Coleoidea

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Figure. Arm suckers. Left - Oral view of arm suckers of Sthenoteuthis

5. Arms IV without connecting web.

Oegopsida No Yes No No No Yes

Myopsida Yes Yes Yes/No Yes Yes Yes

Idiosepiidae Yes No No No Yes ?

Sepioidea Yes No Yes/No Yes Yes No

A key to the families of the Decapodiformes can be found here.

Discussion of Phylogenetic Relationships

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Bathyteuthoidea Vecchione, Young and

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Containing group: Decapodiformes

These are small mesopelagic to bathypelagic squids.

• with suckers on the buccal membrane.

Discussion of Phylogenetic Relationships

Other Names for Bathyteuthoidea Vecchione, Young and Sweeney, 2004

Idiosepiidae Appellof, 1898

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Containing group: Decapodiformes

• with an adhesive organ on the dorsal surface of the mantle.

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Figure. Oral view of hectocotylized arms IV of Idiosepius pygmaeus, 18 mm