The nautilus (from the Latin form of the original Ancient Greek: ναυτίλος,

'sailor') is a pelagic marine mollusc of the cephalopod family Nautilidae, the sole
extant family of the superfamily Nautilaceae and of its smaller but near equal
suborder, Nautilina.

It comprises six living species in two genera, the type of which is the genus
Nautilus. Though it more specifically refers to species Nautilus pompilius, the
name chambered nautilus is also used for any of the Nautilidae. All are protected
under CITES Appendix II.

Nautilidae, both extant and extinct, are characterized by involute or more or less
convolute shells that are generally smooth, with compressed or depressed whorl
sections, straight to sinuous sutures, and a tubular, generally central siphuncle.
[3] Having survived relatively unchanged for hundreds of millions of years,
nautiluses represent the only living members of the subclass nautiloidea, and are
often considered "living fossils".

The first and oldest fossil of Chambered Nautilus displayed at Philippine National
Museum.
The word nautilus is derived from the Greek ναυτίλος nautílos and originally
referred to the paper nautiluses of the genus Argonauta, which are actually
octopuses. The word nautílos literally means "sailor", as paper nautiluses were
thought to use two of their arms as sails.[4]

Contents
1 Anatomy
1.1 Cirri
1.2 Digestive system
1.3 Circulatory system
1.4 Nervous system
1.5 Shell
1.6 Size
2 Physiology
2.1 Buoyancy and movement
2.2 Senses
2.3 Brain and intelligence
2.4 Reproduction and lifespan
3 Ecology
3.1 Range and habitat
3.2 Diet
4 Evolution
4.1 Fossil genera
5 Taxonomy
5.1 Dubious or uncertain taxa
6 Conservation status and human use
7 See also
8 References
8.1 Notes
8.2 Bibliography
9 External links
Anatomy

Diagram of the anatomical structure of a female N. pompilius including most of its

internal organs.
Cirri
The "tentacles" of the nautiluses are actually cirri (singular: cirrus), composed
of long, soft, flexible appendages which are retractable into corresponding
hardened sheaths. Unlike the 8–10 head appendages of coleoid cephalopods,
nautiluses have many cirri. In the early embryonic stages of nautilus development a
single molluscan foot differentiates into a total of 60–90 cirri, varying even
within a species.[5] Nautilus cirri also differ from the tentacles of some coleoids
in that they are non-elastic and lack pads or suckers. Instead, nautilus cirri
adhere to prey by means of their ridged surface. Nautiluses have a powerful grip,
and attempts to take an object already grasped by a nautilus may tear away the
animal's cirri, which will remain firmly attached to the surface of the object. The
main cirri emerge from sheaths which cohere into a single firm fleshy mass. Also,
the pair of cirri before the eye (pre-ocular) and the pair of cirri behind the eye
(post-ocular) are separate from the others. These are more evidently grooved, with
more pronounced ridges. They are extensively ciliated and are believed to serve an
olfactory purpose.[6][7][8]

Digestive system
The radula is wide and distinctively has nine teeth.

The mouth consists of a parrot-like beak made up of two interlocking jaws capable
of ripping the animal's food— mostly crustaceans— from the rocks to which they are
attached.[9]:p. 105 Males can be superficially differentiated from females by
examining the arrangement of tentacles around the buccal cone: males have a spadix
organ (shaped like a spike or shovel) located on the left side of the cone making
the cone look irregular, whereas the buccal cone of the female is bilaterally
symmetrical.[9]:pp. 115–130

The crop is the largest portion of the digestive tract, and is highly extensible.
From the crop, food passes to the small muscular stomach for crushing, and then
goes past a digestive caecum before entering the relatively brief intestine.

Circulatory system
Like all cephalopods, the blood of the nautilus contains hemocyanin, which is blue
in its oxygenated state. There are two pairs of gills which are the only remnants
of the ancestral metamerism to be visible in extant cephalopods.[10]:56 Oxygenated
blood arrives at the heart through four ventricles and flows out to the animal's
organs through distinct aortas but returns through veins which are too small and
varied to be specifically described. The one exception to this is the vena cava, a
single large vein running along the underside of the crop into which nearly all
other vessels containing deoxygenated blood empty. All blood passes through one of
the four sets of filtering organs (composed of one pericardial appendage and two
renal appendages) upon leaving the vena cava and before arriving at the gills for
re-oxygenation. Blood waste is emptied through a series of corresponding pores into
the pallial cavity.

Nervous system
The central component of the nautilus nervous system is the oesophageal nerve ring
which is a collection of ganglia, commissures, and connectives that together form a
ring around the animal's oesophagus. From this ring extend all of the nerves
forward to the mouth, tentacles, and funnel; laterally to the eyes and rhinophores;
and posteriorly to the remaining organs.

The nerve ring does not constitute what is typically considered a cephalopod
"brain": the upper portion of the nerve ring lacks differentiated lobes, and most
of the nervous tissue appears to focus on finding and consuming food (i.e., it
lacks a "higher learning" center). Nautiluses also tend to have rather short memory
spans, and the nerve ring is not protected by any form of brain case.[11]

Shell
Nautilus half-shell showing the camerae in a logarithmic spiral

Section cut of a nautilus shell

A nautilus shell viewed from above (left), and from underneath (right)
Nautiluses are the sole living cephalopods whose bony body structure is
externalized as a planispiral shell. The animal can withdraw completely into its
shell and close the opening with a leathery hood formed from two specially folded
tentacles. The shell is coiled, aragonitic,[12] nacreous and pressure-resistant,
imploding at a depth of about 800 m (2,600 ft). The nautilus shell is composed of
two layers: a matte white outer layer, and a striking white iridescent inner layer.
The innermost portion of the shell is a pearlescent blue-gray. The osmeña pearl,
contrarily to its name, is not a pearl, but a jewellery product derived from this
part of the shell.

Internally, the shell divides into camerae (chambers), the chambered section being
called the phragmocone. The divisions are defined by septa, each of which is
pierced in the middle by a duct, the siphuncle. As the nautilus matures, it creates
new, larger camerae and moves its growing body into the larger space, sealing the
vacated chamber with a new septum. The camerae increase in number from around 4 at
the moment of hatching to 30 or more in adults.

The shell coloration also keeps the animal cryptic in the water. When seen from
above, the shell is darker in color and marked with irregular stripes, which helps
it blend into the dark water below. The underside is almost completely white,
making the animal indistinguishable from brighter waters near the surface. This
mode of camouflage is called countershading.

The nautilus shell presents one of the finest natural examples of a logarithmic
spiral, although it is not a golden spiral. The use of nautilus shells in art and
literature is covered at nautilus shell.

Size
N. pompilius is the largest species in the genus. One form from Indonesia and
northern Australia, once called N. repertus, may reach 25.4 cm (10.0 in) in
diameter.[13] However, most nautilus species never exceed 20 cm (8 in). Nautilus
macromphalus is the smallest species, usually measuring only 16 cm (6 1⁄2 in). A
dwarf population from the Sulu Sea (Nautilus pompilius suluensis) is even smaller,
with a mean shell diameter of 11.56 cm (4.55 in).[14]

Physiology
Buoyancy and movement
File:Nautilus.ogv
Nautilus locomotion
File format: Ogg
File size: 1.29 MB
Duration: 5 seconds

Nautilus with extended tentacles and hyponome visible

To swim, the nautilus draws water into and out of the living chamber with its
hyponome, which uses jet propulsion. This mode of propulsion is generally
considered inefficient compared to propulsion with fins or undulatory locomotion,
however, the nautilus has been found to be particularly efficient compared to other
jet-propelled marine animals like squid and jellyfish, or even salmon at low
speeds.[15] It is thought that this is related to the use of asymmetrical
contractile cycles and may be an adaptation to mitigate metabolic demands and
protect against hypoxia when foraging at depth.[16] While water is inside the
chamber, the siphuncle extracts salt from it and diffuses it into the blood. The
animal adjusts its buoyancy only in long term density changes by osmosis, either
removing liquid from its chambers or allowing water from the blood in the siphuncle
to slowly refill the chambers. This is done in response to sudden changes in
buoyancy that can occur with predatory attacks of fish, which can break off parts
of the shell. This limits nautiluses in that they cannot operate under the extreme
hydrostatic pressures found at depths greater than approximately 800 metres (2,600
ft), and in fact implode at about that depth, causing instant death.[14] The gas
also contained in the chambers is slightly below atmospheric pressure at sea level.
[17] The maximum depth at which they can regulate buoyancy by osmotic removal of
chamber liquid is not known.[18]

The nautilus has the extremely rare ability to withstand being brought to the
surface from its deep natural habitat without suffering any apparent damage from
the experience. Whereas fish or crustaceans brought up from such depths inevitably
arrive dead, a nautilus will be unfazed despite the pressure change of as much as
80 standard atmospheres (1,200 psi). The exact reasons for this ability, which is
thought to be coincidental rather than specifically functional, are not known,
though the perforated structure of the animal's vena cava is thought to play an
important role.[9]:p. 188

Senses

Head of N. pompilius showing the rudimentary eye, which functions similarly to a

pinhole camera
Unlike many other cephalopods, nautiluses do not have what many consider to be good
vision; their eye structure is highly developed but lacks a solid lens. Whereas a
sealed lens allows for the formation of highly focused and clear, detailed
surrounding imagery, nautiluses have a simple pinhole eye open to the environment
which only allows for the creation of correspondingly simple imagery.

Instead of vision, the animal is thought to use olfaction (smell) as the primary
sense for foraging and for locating and identifying potential mates.[19]

The "ear" of the nautilus consists of structures called otocysts located

immediately behind the pedal ganglia near the nerve ring. They are oval structures
densely packed with elliptical calcium carbonate crystals.

Brain and intelligence

Main article: Cephalopod intelligence
Nautiluses are much closer to the first cephalopods that appeared about 500 million
years ago than the early modern cephalopods that appeared maybe 100 million years
later (ammonoids and coleoids). They have a seemingly simple brain, not the large
complex brains of octopus, cuttlefish and squid, and had long been assumed to lack
intelligence. But the cephalopod nervous system is quite different from that of
other animals, and recent experiments have shown not only memory, but a changing
response to the same event over time.[20][21][22]

In a study in 2008, a group of nautiluses (N. pompilius) were given food as a

bright blue light flashed until they began to associate the light with food,
extending their tentacles every time the blue light was flashed. The blue light was
again flashed without the food 3 minutes, 30 minutes, 1 hour, 6 hours, 12 hours,
and 24 hours later. The nautiluses continued to respond excitedly to the blue light
for up to 30 minutes after the experiment. An hour later they showed no reaction to
the blue light. However, between 6 and 12 hours after the training, they again
responded to the blue light, but more tentatively. The researchers concluded that
nautiluses had memory capabilities similar to the "short-term" and "long-term
memories" of the more advanced cephalopods, despite having different brain
structures.[20][21][22] However the long-term memory capability of nautiluses was
much shorter than that of other cephalopods. The nautiluses completely forgot the
earlier training 24 hours later, in contrast to octopuses, for example, which can
remember conditioning for weeks afterwards. However, this may be simply the result
of the conditioning procedure being suboptimal for sustaining long-term memories in
nautiluses. Nevertheless, the study showed that scientists had previously
underestimated the memory capabilities of nautiluses.[22]

Reproduction and lifespan

Nautiluses reproduce by laying eggs. Gravid females attach the fertilized eggs,
either singly or in small batches, to rocks in warmer waters (21-25 Celsius),
whereupon the eggs take eight to twelve months to develop until the 30-millimetre
(1.2 in) juveniles hatch.[23] Females spawn once per year and regenerate their
gonads, making nautiluses the only cephalopods to present iteroparity or polycyclic
spawning.[24]

Nautiluses are sexually dimorphic, in that males have four tentacles modified into
an organ, called the "spadix", which transfers sperm into the female's mantle
during mating. At sexual maturity, the male shell becomes slightly larger than the
female's.[25] Males have been found to greatly outnumber females in practically all
published studies, accounting for 60 to 94% of all recorded individuals at
different sites.[14]

The lifespan of nautiluses may exceed 20 years, which is exceptionally lengthy for
a cephalopod, many of whom live less than three even in captivity and under ideal
living conditions.[26] However, nautiluses typically do not reach sexual maturity
until they are about 15 years old, limiting their reproductive lifespan to often
less than five years.[14]

Left: Frequency distribution of N. pompilius shell diameter at Osprey Reef, part of

the Coral Sea Islands, based on 2067 captured individuals. Shells ranged in size
from 76 to 145 mm, with a mean of 128.6±28.01 mm.[14]
Right: Shell diameter of mature male and female N. pompilius caught at Osprey Reef.
Males (n = 870) had a mean shell diameter of 131.9±2.6 mm, compared to 118.9±7.5 mm
in females (n = 86). The Osprey Reef N. pompilius population is the second smallest
known in terms of mean shell diameter, after the dwarf form from the Sulu Sea
(130.7 mm and 115.6 mm, respectively).[14]
Ecology
Range and habitat

Number of captured N. pompilius at various depths around the Osprey Reef Seamount,
Coral Sea. The data was collated from 271 trapping events spread across all months
of the year. Nautiluses were most common at 300–350 m (1,000–1,100 ft). No
specimens were recovered from a depth of less than 150 m (500 ft) during 18
trapping efforts.[14]
Nautiluses are found in only the Indo-Pacific, from 30° N to 30° S latitude and 90°
E to 175° E longitude. They inhabit the deep slopes of coral reefs.

Nautiluses usually inhabit depths of several hundred metres. It has long been
believed that nautiluses rise at night to feed, mate and lay eggs, but it appears
that, in at least some populations, the vertical movement patterns of these animals
are far more complex.[27] The greatest depth at which a nautilus has been sighted
is 703 m (2,306 ft) (N. pompilius).[27] Implosion depth for nautilus shells is
thought to be around 800 m (2,600 ft).[14][27] Only in New Caledonia, the Loyalty
Islands, and Vanuatu can nautiluses be observed in very shallow water, at depths of
as little as 5 m (15 ft).[18][27] This is due to the cooler surface waters found in
these southern hemisphere habitats as compared to the many equatorial habitats of
other nautilus populations – these usually being restricted to depths greater than
100 m (300 ft).[18][27] Nautiluses generally avoid water temperatures above 25 °C
(75 °F).[27]

A pair of N. pompilius feeding on two-spot red snapper (Lutjanus bohar) bait during
daytime at 703 m (2,306 ft) depth. This observation constitutes the deepest record
of any nautilus species.
Diet
Nautiluses are scavengers and opportunistic predators.[28][29] They eat molts of
lobsters, hermit crabs, and carrion of any kind.[18]

Evolution

Shell characters of the genera Nautilus and Allonautilus

Section cut of a nautilus shell

Fossil records indicate that nautiloids have not evolved much during the last 500
million years. Many were initially straight-shelled, as in the extinct genus
Lituites. They developed in the Late Cambrian period and became a significant group
of sea predators during the Ordovician period. Certain species reached over 2.5 m
(8 ft) in size. The other cephalopod subclass, Coleoidea, diverged from the
nautiloids long ago and the nautilus has remained relatively unchanged since.
Nautiloids were much more extensive and varied 200 million years ago. Extinct
relatives of the nautilus include ammonites, such as the baculites and goniatites.

The family Nautilidae has its origin in the Trigonocerataceae (Centroceratina),

specifically in the Syringonautilidae of the Late Triassic[3] and continues to this
day with Nautilus, the type genus, and its close relative, Allonautilus.

Fossil genera

Eutrephoceras dorbignyanum
The fossil record of Nautilidae begins with Cenoceras in the Late Triassic, a
highly varied genus that makes up the Jurassic Cenoceras complex. Cenoceras is
evolute to involute, and globular to lentincular; with a suture that generally has
a shallow ventral and lateral lobe and a siphuncle that is variable in position but
never extremely ventral or dorsal. Cenoceras is not found above the Middle Jurassic
and is followed by the Upper Jurassic-Miocene Eutrephoceras.

Eutrephoceras is generally subglobular, broadly rounded laterally and ventrally,

with a small to occluded umbilicus, broadly rounded hyponomic sinus, only slightly
sinuous sutures, and a small siphuncle that is variable in position.

Next to appear is the Lower Cretaceous Strionautilus from India and the European
ex-USSR, named by Shimankiy in 1951. Strionautilus is compressed, involute, with
fine longitudinal striations. Whorl sections are subrectangular, sutures sinuous,
the siphuncle subcentral.

Also from the Cretaceous is Pseudocenoceras, named by Spath in 1927.

Pseudocenoceras is compressed, smooth, with subrectangular whorl sections,
flattened venter, and a deep umbilicus. The suture crosses the venter essentially
straight and has a broad, shallow, lateral lobe. The siphuncle is small and
subcentral. Pseudocenoceras is found in the Crimea and in Libya.

Carinonautilus is a genus from the Upper Cretaceous of India, named by Spengler in

1919. Carinonautilus is a very involute form with high whorl section and flanks
that converge on a narrow venter that bears a prominent rounded keel. The umbilicus
is small and shallow, the suture only slightly sinuous. The siphuncle is unknown.
Obinautilus has also been placed in Nautilidae by some authorities, though it may
instead be an argonautid octopus.[30][31]

Taxonomy
Photo of profiles of three progressively larger nautilus shells
Nautilus shells: N. macromphalus (left), A. scrobiculatus (centre), N. pompilius
(right)
The family Nautilidae contains up to six extant species and several extinct
species:

Genus Allonautilus
A. perforatus
A. scrobiculatus
Genus Nautilus
N. belauensis
†N. cookanum
N. macromphalus
N. pompilius (type)
N. p. pompilius
N. p. suluensis
†N. praepompilius
N. stenomphalus
Recent genetic data has pointed to there being only three extant species: A.
scrobiculatus, N. macromphalus, and N. pompilius, with N. belauensis and N.
stenomphalus both subsumed under N. pompilius, possibly as subspecies.[14]

Dubious or uncertain taxa

The following taxa associated with the family Nautilidae are of uncertain taxonomic
status:[32]

Binomial name and author citation Current systematic status Type locality
Type repository
N. alumnus Iredale, 1944 Species dubium [fide Saunders (1987:49)] Queensland,
Australia Not designated [fide Saunders (1987:49)]
N. ambiguus Sowerby, 1848 Species dubium [fide Saunders (1987:48)] Not
designated Unresolved
N. beccarii Linné, 1758 Non-cephalopod; Foraminifera [fide Frizzell and Keen
(1949:106)]
N. calcar Linné, 1758 ?Non-cephalopod; Foraminifera Lenticulina Adriatic Sea
Unresolved; Linnean Society of London?
N. crispus Linné, 1758 Undetermined Mediterranean Sea Unresolved; Linnean
Society of London?
N. crista Linné, 1758 Non-cephalopod; Turbo [fide Dodge (1953:14)]
N. fascia Linné, 1758 Undetermined Adriatic Sea Unresolved; Linnean
Society of London?
N. granum Linné, 1758 Undetermined Mediterranean Sea Unresolved; Linnean
Society of London?
N. lacustris Lightfoot, 1786 Non-cephalopod; Helix [fide Dillwyn (1817:339)]
N. legumen Linné, 1758 Undetermined Adriatic Sea Unresolved; Linnean
Society of London?
N. micrombilicatus Joubin, 1888 Nomen nudum
N. obliquus Linné, 1758 Undetermined Adriatic Sea Unresolved; Linnean
Society of London?
N. pompilius marginalis Willey, 1896 Species dubium [fide Saunders (1987:50)]
New Guinea Unresolved
N. pompilius moretoni Willey, 1896 Species dubium [fide Saunders (1987:49)] New
Guinea Unresolved
N. pompilius perforatus Willey, 1896 Species dubium [fide Saunders (1987:49)]
New Guinea Unresolved
N. radicula Linné, 1758 ?Non-cephalopod; F. Nodosaria Adriatic Sea Unresolved;
Linnean Society of London?
N. raphanistrum Linné, 1758 Undetermined Mediterranean Sea Unresolved;
Linnean Society of London?
N. raphanus Linné, 1758 Undetermined Adriatic Sea Unresolved; Linnean
Society of London?
N. semi-lituus Linné, 1758 Undetermined Liburni, Adriatic Sea Unresolved;
Linnean Society of London?
N. sipunculus Linné, 1758 Undetermined "freto Siculo" Unresolved;
Linnean Society of London?
N. texturatus Gould, 1857 Nomen nudum
Octopodia nautilus Schneider, 1784 Rejected specific name [fide Opinion 233, ICZN
(1954:278)]