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to shallow tide pools, research
has recently revealed some of the
mysterious behaviors of two
famed cephalopods, the Giant
Squid and the deadly Blue-ringed
octopus (click image below for an
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Lophotrochozoa : Mollusca
Squids, octopuses, nautilus, and ammonites
Cephalopods are the most intelligent, most mobile, and the
largest of all molluscs. Squid, octopuses, cuttlefish, the
chambered nautilus, and their relatives display remarkable
diversity in size and lifestyle with adaptations for predation,
locomotion, disguise, and communication. These "brainy"
invertebrates have evolved suckered tentacles, camera-like
eyes, color-changing skin, and complex learning behavior.
Their lengthy evolutionary history spans an impressive 500
million years and the abundant fossils they've left behind
(mostly shelled nautiloids and ammonoids) record repeated
speciation and extinction events. From myths about their
enigmatic fossilized remains to fantastic accounts of
tentacled sea monsters, cephalopods also figure prominently
in the literature and folklore of human societies around the
world. Today, biologists and paleontologists continue to
captivate the human mind and imagination with details of
these molluscs' behavior, natural history, and evolution.
There are about 17,000 named species
of fossil cephalopods, compared to the
800 identified living species of
cephalopods. Clearly the lineages of
extinct taxa were prolific and diverse. So
diverse in fact, that paleontologists have
identified three distinct fossil clades that
are entirely extinct: Endoceratoidea,
Actinoceratoidea, and Bactritoidea
(cladogram A, at right). All members of
these clades were squid-like, but had
straight external shells called
orthocones. They flourished in Paleozoic
oceans between the Ordovician (488
mya) and Triassic periods (200 mya)
with shells that, in some species,
reached nearly 10 meters in length.
More familiar to us in the fossil record are the nautiloids, ammonoids, and belemnites.
Nautiloids and ammonoids
Nautiloids are the earliest cephalopods found in the fossil record, appearing by the Late Cambrian. The
earliest forms were orthoconic (having straight shells), but during the Ordovician the nautiloids
experienced a rapid diversification and evolved a planispiral (coiled in a single plane) shell shape. All
have shells with nacre and interconnected internal chambers, similar to what we see in the modern
nautilus. This morphology is very similar to many of the ammonoids, which first appear in coiled form
FUN FACT: Ammonoids, like
belemnites, have also played a
notable role in folklore. During
the Middle Ages, their coiled
shells were interpreted by the
English, who encountered them
in Jurassic-aged rocks exposed
throughout Great Britain, as
lithified snakes (called "snake
stones"). Similarly, ammonite
fossils encountered by the early
Romans were mistaken for horns,
and termed "ammonites" for the
coiled horns of the Egyptian ram-
in the Devonian Period. Though nautiloids and ammonoids may appear the same, they are easily
distinguished by the location of their siphuncle and the shapes of their sutures.
The photo on the left shows the position of the siphuncle in ammonoids. The other two are of nautiloids exhibiting
the simple sutures typical of the group.
The siphuncle is an internal tube that runs through and connects the chambers of the shell. In
nautiloids, it runs through the center of the shell chambers, while in almost all planispiral ammonoids,
it is found along the shell's outer edge (above left). Sutures are contact lines between shell chamber
walls (called septa) and the inner shell wall of nautiloid and ammonoid shells. In nautiloids these lines
are straight and are called simple sutures (above center and right). In contrast, ammonoid sutures dip
and fold in undulations called lobes and saddles (below left). The most undulated, complex sutures are
found in the prolific ammonoids of the Cretaceous, the ammonites (below right). One pattern of
change in the evolution of ammonoids as a clade is that their suture morphology became more
complex with time. That is, ammonoids from the Cretaceous have sutures with more intricate lobes
and saddles than those of their relatives from the Permian 200 million years earlier.
Hildoceras bifrons (left) illustrates the typical lobes and saddles of ammonoid sutures. The sutures on the
ammonite (right) are even more complex.
Suture patterns and lifestyle: The diversity of this suture
geometry has inspired paleontologists to investigate its
function. In one study, paleontologists (Daniel et al. 1997)
used modeling to determine if there was a relationship
between the pattern of lobes and saddles and buoyancy and
the ability to withstand pressure. They found that simple
sutures, like those in nautiloids and early ammonoids, can
withstand great pressure but have poor buoyancy control.
They interpret that these animals lived at depth and were not
fast moving. In contrast, complex sutures like those in
ammonites of the Cretaceous did not withstand pressure
well, but allowed for very effective buoyancy control. They
infer that this reflects a lifestyle at shallower depths. Based
on this evidence, it appears that many ammonoid lineages
evolved over millions of years, beginning in deep water
habitats and evolving to inhabit relatively shallow ones.
Shell morphology: Another characteristic unique to
ammonoid cephalopods is that although most were
planispiral, the shells of some species were wide, open-
coiled, kinked, twisted, or hooked. The implications of these shapes on the animals' behavior and
The rostrum of belemnites is the part most frequently preserved (left). At right, a
long belemnite rostrum in a block containing other marine fossils.
lifestyle could be profound, but how could it be tested? The answer is through inference. One way
paleontologists infer function from an organism's shape or form, is through experimentation and
comparison with living animals. In this case, paleontologists measured buoyancy and hydrodynamic
characteristics of Nautilus, and compared their results to actual tests of ammonoid shells and models
in a water flume. They found that planispiral shell shapes (like a discus) were able to move through
the water quickly, while wider and more open shell shapes moved more slowly. These morphologies
could mean that planispiral cephalopods lived an active, pelagic lifestyle and wider shelled ammonoids
lived near the ocean bottom and were slow movers. Dramatic curves, twists, as well as highly ornate
ammonoid shells also suggest that those animals were relatively slow movers inhabiting a quiet
The fossil record of most
cephalopods in the clade Coloidea
(squid, cuttlefish, octopuses, and
their relatives) is poor, especially
when compared to their shelly
relatives. Their hard parts, if any,
are internal, can be greatly
reduced in size, and often lack
calcification. The extinct
belemnites, however, are the
exception. These squid-like
animals (below) swam with
ammonoids and nautiloids in
oceans of the Triassic, Jurassic, and Cretaceous Periods and are considered by paleontologists to be
the ancestors of the Coleoidea. Like orthocones, belemnites had a straight shell, but it was internal,
not external. It was made of three parts, a proostracum and phragmocone followed by a rostrum.
Being highly resistant, the posterior bullet-shaped rostrum is most often preserved and can be found
in great quantity and concentration in Mesozoic marine sediments (see photos). Before these bullet-
shaped fossils were understood as fossils, early Europeans explained them as the products of lightning
hitting the ground and named them "thunderbolts" or "thunderstones."
Despite the great diversity in size, shell
morphology, behavior, and lifestyle,
cephalopods are all united by a suite of
shared molluscan characters. They are
marine, predatory, and have at least
eight arms derived from the molluscan
foot. All have a modified radula, and a
horny, parrot-like beak for subduing
prey. Their mantle is modified into a
siphon for movement via jet-propulsion,
and their highly developed nervous and
A female Octopus digueti.
FUN FACT: In 2006, a UC
Berkeley graduate student
reported that the Indonesian
Coconut Octopus, Octopus
marginatus, can move along the
sensory systems include complex eyes
and a centralized brain. Their name,
Cephalopod or "head-foot" in Greek,
reflects the unique relationship between
the cephalopod head and foot: the arms
encircle the animals' head.
Life history & ecology
Three main clades are typically identified
within the Cephalopoda (cladogram B,
left): the Ammonoidea (an extinct and
shelled clade), Nautiloidea (with only
one living shelled genus, Nautilus) and
Coleoidea (squids, cuttlefish, octopuses,
the Ram's Horn Squid, the "paper
nautilus," and an extinct clade, the
belemnites). The ammonoid lineage
survived for 300 million years in the
oceans of the Paleozoic and Mesozoic. Most had planispiral (coiled in a single plane) external shells,
and throughout their evolutionary history these plentiful predators shared the seas with the nautiloids,
a clade of less diverse shelled cephalopods. By the end of the Cretaceous Period however, extinction
had wiped out the ammonoids entirely and left only one surviving nautiloid clade, the genus Nautilus.
Today, the only living representatives of shelled cephalopods are a few species of Nautilus. These
molluscs are slow-moving, restricted to deep water, and have coiled shells that are similar to those of
their fossil ancestors.
From left, the exterior shell of the extant Nautilus; a cross-section of the Nautilus shell; the internal shells of the Ram's
Horn Squid (Spirula spirula); and the secreted external papery shell-like brood pouch of the female Paper Nautilus
Members of the Coleoidea are probably the best known of the
Cephalopoda, as this group contains the squids (Teuthoidea)
and octopuses (Octopoda) (right). Also included in this clade
are the lesser-known cuttlefish (Sepioidea), the Ram's Horn
Squid, which has an internal coiled shell and floats head down
in the water, and an enigmatic deep water genus called the
"vampire squid" (Vampyromorpha). The majority of coleoids
are squid species, and most of these animals are torpedo-
shaped, fast moving, and have a thin, flexible internal shell
called a pen. Cuttlefish (sometimes called "cuttles") look like
squid, but have stouter bodies with a broad internal shell called
a cuttlebone or sepion. They move mostly by undulating their
body fins and can live in the water column or at the sediment
surface. Octopuses are adapted to a benthic lifestyle, have no
shell, can mimic their surroundings, and even "walk" on two of their eight arms. Belemnites are
sometimes considered the sister group of extant (living) coleoids, but have also been interpreted as
Based on molecular and morphological data, the extant
Coleoidea is a monophyletic group (cladogram C, below). It is
the sister group of the living species of Nautilus, which is also
a monophyletic clade, albeit of only a few species (D, below).
The phylogenetic relationships within the coleoids is less
clear. Some cladograms depict the Sepioidea (cuttlefish),
Teuthoidea (squid), Vampyromorpha (vampire "squid"), and
ocean floor using only the tips of
its arms, almost like "walking."
See Crissy Huffard's "Walktopus."
Octopoda (octopuses) as coupled into two sets of sister
groups, the Decabrachia and Octopodiformes (E, below).
While the cladograms from another study (Lindgren et al.
2004) illustrate that although the coleoids are monophyletic,
the cuttlefish (Sepiodea) are nested within the squids
(Teuthoidea), making the Teuthoidea clade paraphyletic (F,
More on morphology
The external shell. One of the most obvious differences in body
type, which for cephalopods does reflect shared ancestry, is the
presence or absence of an external shell. Clades without an
external shell are called endocochleate and include the coleoids;
squids, cuttlefish, and octopuses. The internal shell of these taxa,
called a gladius, can be cartilaginous, calcareous, chitinous or
absent entirely. Presence of an external shell, such as that of the
Nautiloids, Ammonoids, and orthoconic cephalopods, is considered
a plesiomorphic, or ancestral state.
The eye. No presentation of cephalopods would be complete
without a discussion of the cephalopod eye. This structure is
probably the most sophisticated eye of all invertebrates and is as complex as the vertebrate eye,
though the two are not homologous. For their body size, cephalopod eyes are relatively large. They
contain an iris, pupil, and lens, but not necessarily a cornea. Octopuses are the only cephalopods with
a completely protected "closed" cornea. That means that the eyes of squids and sepioids (cuttlefish,
etc.) are in direct contact with sea water! The pupil in cephalopods is unique in that its morphology is
different in octopuses, cuttlefish, and squid. Octopuses have a slit-shaped rectangular pupil. In
cuttlefish it is W-shaped, and in squid it is round (see below). In Nautilus the eye is much simpler. It is
mounted on a stalk, has no lens, and has a very small pupil (1-2 mm). It can narrow and widen in
different brightnesses but resolves images poorly, so probably is useful only to detect light.
Differing eye morphologies in cephalopods. From left, a squid (Loligo), octopus, cuttlefish, and Nautilus. Note the
hyponome below the octopus eye — this is a muscular tube, that when contracted, expels water in a jet, propelling the
octopus backwards. The hyponome can be aimed in various directions, giving the octopus finer control over its escape
Arms and tentacles. Arms and tentacles are another distinguishing cephalopod characteristic. All
cephalopods have arms, but not all cephalopods have tentacles. Octopuses, cuttlefish, and squid have
eight non-retractable arms, but only cuttlefish and squid (Sepioidea and Teuthoidea) have tentacles
(two each). Arms usually have cirri (fleshy papillae/palps), often suckers, and sometimes hooks
(modified suckers) along their undersides. These can be attached to the arm directly or by a flexible
stalk and are used to adhere to substrates and catch prey. Tentacles are longer than arms, are
retractable, and usually have a blade-shaped or flattened tip, called a club, which is covered in
suckers. Squid and cuttlefish have one pair of tentacles and they use these to strike quickly at prey.
FUN FACT: The largest
hamiltoni, (Fig. 17) called the
colossal squid, is longer than a
city bus, while the smallest
cephalopod, Idiosepius notoides,
the pygmy squid, could fit on
FUN FACT: A hectocotylus is a
cephalopodic arm of a male,
modified to deliver a
containing sac) to a female. In
some taxa, part of this arm can
detach from the male and remain
inside the female. In 1829, the
famous naturalist George Cuvier
first identified and described a
hectocotylus in the paper nautilus
(an octopus of the genus
Argonauta) but, at the time he
thought that this detached arm
was a parasitic worm! He named
it Hectocotylus octopodis. When it
was discovered that the "worm"
was really a cephalopod arm, the
name "hectocotylus" was kept
but its meaning changed to refer
to the spermatophore-
transferring arm of any male
Top: The chromatophores on this three-
millimeter-long, 12-hour-old baby blue-ring
octopus are clearly visible. Bottom: Click on this
photo to see an enlargement — you'll need to in
order to see the chromatophores (the dark
stippling) on the skin of this California Two-spot
Octopus (Octopus bimaculoides).
Sex and reproduction. Sex and reproduction in
cephalopods is in many ways quite different than in other
molluscs. First, sexes are separate and mating usually
includes a courtship that often involves elaborate color
changes. This is followed by the transfer of a spermatophore
(sperm packet) by a male to a female through her mantle
opening. The spermatophore is transferred by the male using
either a penis or a modified arm called a hectocotylus. Most
females then lay large yolky eggs in clusters on the ocean
floor or on any other hard substrate. Eggs develop by
dividing unequally instead of in the spiral pattern of other
molluscs. It is thought this is a derived mode of
development. After a period of development within the egg,
juveniles hatch out directly without the swimming larval stage common to many other molluscs. Most
males and females die shortly after spawning.
Cephalopods have an
amazing ability to
change color very
accomplish this feat
found in the skin,
and expand and
contract to reveal or
conceal small dots of
color (left). They can
be so densely
200 may be found in
a patch of skin the
size of a pencil
an iridescent dermal
tissue can also be
manipulated by some
cephalopods to aid in
courtship rituals, or
accompany color changes.
Some pelagic squids possess an additional color-changing structure; the light organ. A pair of these
light organs is located within the mantle cavity on the underside of the squid. Each contains a crypt
and a lens. A crypt is a small sac that houses luminous bacteria and a lens is a complex stack of tiny
reflecting plates that controls the luminescence of this bacteria. Light from the bacteria projects
downward and the squid can manipulate its intensity to match any light coming from above. This
masks the squid's own silhouette, protecting it from potential predators.
The brain. Finally, one of the most intriguing aspects of cephalopods is their intelligence. With a
centralized brain, the largest of all invertebrates, and highly developed eyes and other sense organs,
they are able to remember and learn by example or through trial and error.
References and resources
Daniel, T.L., B.S. Helmuth, W.B. Saunders, and P.D. Ward. 1997. Septal complexity in
ammonoid cephalopods increased mechanical risk and limited depth. Paleobiology 23(4):470-
Lee, P.N., P. Callaerts, H.G. de Couet, and M.Q Martindale. 2003. Cephalopod Hox genes and
the origin of morphological novelties. Nature 424:1061-1065.
Lindgren, A.R., G. Giribet, and M.K. Nishiguchi. 2004. A combined approach to the phylogeny of
Cephalopoda (Mollusca). Cladistics 20(5):454-486.
Mangold (1922-2003), K.M., and R.E. Young. 1996. Idiosepiidae Appellof, 1898. Pygmy squids.
http://tolweb.org/Idiosepiidae/19985/ in The Tree of Life Web Project, http://tolweb.org.
Ruppert, E.E., R.S. Fox, and R.D. Barnes. 2004. Invertebrate Zoology : A Functional
Evolutionary Approach. Brooks Cole, California. 963 pp.
Sual, L.R., and C.J. Stadum. 2005. Fossil Argonauts (Mollusca: Cephalopoda: Octopodida) from
late Miocene siltstones of the Los Angeles basin, California. Journal of Paleontology 79(3):520-
Wei, S.L., and R.E. Young. 1989. Development of symbiotic bacterial bioluminescence in a
nearshore cephalopod, Euprymna scolopes. Marine Biology 103:541-546.
Crissy Huffard's "Walktopus".
National Geographic News page with photos of the first live giant squid.
About the hectocotylus.
University of Michigan's Museum of Zoology Animal Diversity Web information on
Vampyroteuthis infernalis, the vampire squid.
National Geographic News story on a light-producing squid.
Sex identification and mating in the blue-ringed octopus, Hapalochlaena lunulata.
Original text by Jann Vendetti, UCMP, 2006. Blue-ringed octopus (adult and baby) and Octopus digueti photos by Roy Caldwell,
Integrative Biology and UCMP, UC Berkeley; ammonoid, nautiloid, belemnite, Nautilus shell, Ram's Horn Squid, and paper nautilus
photos by Jann Vendetti, UCMP; cephalopod eye morphology and Octopus bimaculoides photos © Larry Jon Friesen.
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