Nautilus

File:Nautilus diagram-en.svg

The arm crown of modern nautilids (genera Nautilus and Allonautilus) is very distinct in comparison to coleoids. Unlike the ten-armed Decabrachia or the eight-armed Octopodiformes, nautilus may possess any number of tentacles (cirri) from 50 to over 90 tentacles depending on the sex and individual.{{cite journal |last1=Kröger |first1=Björn |last2=Vinther |first2=Jakob |last3=Fuchs |first3=Dirk |title=Cephalopod origin and evolution: A congruent picture emerging from fossils, development and molecules: Extant cephalopods are younger than previously realised and were under major selection to become agile, shell-less predators |journal=BioEssays |date=August 2011 |volume=33 |issue=8 |pages=602–613 |doi=10.1002/bies.201100001|pmid=21681989 |s2cid=2767810 }}{{cite book |last1=Sasaki |first1=T |last2=Shigeno |first2=S|last3=Tanabe |first3=K |date=2010 |title=Cephalopods - Present and Past |publisher=Tokai University Press |chapter=Anatomy of living Nautilus: reevaluation of primitiveness and comparison with Coeloidea |name-list-style=amp}} These tentacles are classified into three distinct categories: ocular, digital, and labial (buccal).{{cite book |last1=Owen |first1=Richard |title=Memoir on the pearly nautilus (Nautilus pompilius, Linn). With illustrations of its external form and internal structure |date=1832 |publisher=Richard Taylor |location=London |url=https://www.biodiversitylibrary.org/bibliography/10441}} There are two sets of ocular tentacles: one set in front of the eye (pre-ocular) and one set behind the eye (post-ocular). The digital and labial tentacles are arrayed circularly around the mouth, with the digital tentacles forming the outermost ring and the labial tentacles in between the digital tentacles and the mouth. There are 19 pairs of digital tentacles that, together with the ocular tentacles, make up the 42 appendages that are visible when observing the animal (not counting the modified tentacles that form the hood). The labial tentacles are generally not visible, being smaller than the digital tentacles, and more variable both in number and in shape.{{Cite journal |last=Griffin |first=Lawrence E. |date=1899 |title=XXVI.— Notes on the tentacles of Nautilus pompilius |journal=Annals and Magazine of Natural History |language=en |volume=3 |issue=14 |pages=170–176 |doi=10.1080/00222939908678098 |issn=0374-5481|url=https://www.biodiversitylibrary.org/part/58804 }} Males modify three of their labial tentacles into the spadix, which delivers spermatophores into the female during copulation.

The tentacle is composed of two distinct structures: the first structure, a fleshy sheath that contains the second structure: an extendable cirrus (plural: cirri).{{Citation |last=Kier |first=William M. |title=The Functional Morphology of the Tentacle Musculature of Nautilus pompilius |date=2010 |url=https://doi.org/10.1007/978-90-481-3299-7_18 |work=Nautilus: The Biology and Paleobiology of a Living Fossil, Reprint with additions |pages=257–269 |editor-last=Saunders |editor-first=W. Bruce |access-date=2023-11-28 |series=Topics in Geobiology |volume=6 |place=Dordrecht |publisher=Springer Netherlands |language=en |doi=10.1007/978-90-481-3299-7_18 |isbn=978-90-481-3299-7 |editor2-last=Landman |editor2-first=Neil H.|url-access=subscription }} The sheaths of the digital tentacles are fused at their base into a single mass referred to as the cephalic sheath. The digital cirri can be fully withdrawn into the sheath and are highly flexible, capable of extending just over double their fully retracted length and show a high degree of allowable bendability and torsion. Despite not having suckers, the digital tentacles show strong adhesive capabilities. Adhesion is achieved through the secretion of a neutral (rather than acidic) mucopolysaccharide from secretory cells in the ridges of the digital cirri.{{Citation |last=Fukuda |first=Yoshio |title=Histology of the Long Digital Tentacles |date=2010 |url=https://doi.org/10.1007/978-90-481-3299-7_17 |work=Nautilus: The Biology and Paleobiology of a Living Fossil, Reprint with additions |pages=249–256 |editor-last=Saunders |editor-first=W. Bruce |access-date=2023-11-28 |series=Topics in Geobiology |volume=6 |place=Dordrecht |publisher=Springer Netherlands |language=en |doi=10.1007/978-90-481-3299-7_17 |isbn=978-90-481-3299-7 |editor2-last=Landman |editor2-first=Neil H.|url-access=subscription }}{{Cite journal |last1=Muntz |first1=W. R. A. |last2=Wentworth |first2=S. L. |date=1995 |title=Structure of the Adhesive Surface of the Digital Tentacles of Nautilus Pompilius |journal=Journal of the Marine Biological Association of the United Kingdom |language=en |volume=75 |issue=3 |pages=747–750 |doi=10.1017/S0025315400039163 |bibcode=1995JMBUK..75..747M |s2cid=83497566 |issn=1469-7769}}{{Cite journal |last1=von Byern |first1=Janek |last2=Wani |first2=Ryoji |last3=Schwaha |first3=Thomas |last4=Grunwald |first4=Ingo |last5=Cyran |first5=Norbert |date=2012-02-01 |title=Old and sticky—adhesive mechanisms in the living fossil Nautilus pompilius (Mollusca, Cephalopoda) |journal=Zoology |volume=115 |issue=1 |pages=1–11 |doi=10.1016/j.zool.2011.08.002 |issn=0944-2006 |pmc=3311398 |pmid=22221553|bibcode=2012Zool..115....1V }} Release is triggered through contraction of the tentacle musculature rather than the secretion of a chemical solvent, similar to the adhesion/release system in Euprymna, though it is unclear whether these adhesives are homologous.{{Cite journal |last1=von Byern |first1=Janek |last2=Cyran |first2=Norbert |last3=Klepal |first3=Waltraud |last4=Nödl |first4=Marie Therese |last5=Klinger |first5=Lisa |date=February 2017 |title=Characterization of the adhesive dermal secretion of Euprymna scolopes Berry, 1913 (Cephalopoda) |url=https://linkinghub.elsevier.com/retrieve/pii/S0944200616300836 |journal=Zoology |language=en |volume=120 |pages=73–82 |doi=10.1016/j.zool.2016.08.002|pmid=27646066 |bibcode=2017Zool..120...73V |url-access=subscription }} The ocular tentacles show no adhesive capability but operate as sensory organs. Both the ocular tentacles and the eight lateral digital tentacles show chemoreceptive abilities; the preocular tentacles detect distant odor and the lateral digital tentacles detect nearby odor.{{Cite journal |last1=Basil |first1=Jennifer |last2=Bahctinova |first2=Irina |last3=Kuroiwa |first3=Kristine |last4=Lee |first4=Nandi |last5=Mims |first5=Desiree |last6=Preis |first6=Michael |last7=Soucier |first7=Christian |date=2005 |title=The function of the rhinophore and the tentacles of Nautilus pompilius L. (Cephalopoda, Nautiloidea) in orientation to odor |journal=Marine and Freshwater Behaviour and Physiology |language=en |volume=38 |issue=3 |pages=209–221 |doi=10.1080/10236240500310096 |bibcode=2005MFBP...38..209B |s2cid=33835096 |issn=1023-6244}}{{Cite journal |last1=Ruth |first1=Peter |last2=Schmidtberg |first2=Henrike |last3=Westermann |first3=Bettina |last4=Schipp |first4=Rudolf |date=March 2002 |title=The sensory epithelium of the tentacles and the rhinophore of Nautilus pompilius L. (cephalopoda, nautiloidea) |journal=Journal of Morphology |volume=251 |issue=3 |pages=239–255 |doi=10.1002/jmor.1086 |issn=0362-2525 |pmid=11835362|s2cid=38377665 }}

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.{{Cite book

|title=The anatomy of Nautilus pompilius

|last=Griffin

|first=Lawrence E.

|volume=8

|year=1900

|publisher=Government Printing Office

|place=Washington, D.C.

|url={{google books |plainurl=y |id=IHJBAQAAIAAJ}}

|oclc=18760979

|doi=10.5962/bhl.title.10466

}}{{rp|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.{{rp|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.

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.{{cite journal

|url=http://www.zmuc.dk/inverweb/Galathea/Galathea_p5.html

|format=Link to free full text + plates

|last=Wingstrand

|first=KG

|year=1985

|title=On the anatomy and relationships of Recent Monoplacophora

|journal=Galathea Rep.

|volume=16

|pages=7–94

|access-date=2009-04-21

|archive-url=https://web.archive.org/web/20160303182836/http://www.zmuc.dk/inverweb/Galathea/Galathea_p5.html

|archive-date=2016-03-03

|url-status=dead

}}{{rp|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.

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). Nautili also tend to have rather short memory spans, and the nerve ring is not protected by any form of brain case.{{cite journal

| last =Young

| first =J. Z.

| title =The Central Nervous System of Nautilus

| journal =Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences

| volume =249

| issue =754

| pages =1–25

| date =27 May 1965

| jstor =2416631

| doi =10.1098/rstb.1965.0006

| bibcode =1965RSPTB.249....1Y

| s2cid =83951270

| doi-access =

}}

Image:NautilusCutawayLogarithmicSpiral.jpg|alt=]]File:Nautilus Section cut.jpg

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| footer = A nautilus shell viewed from above (left), and from underneath (right)

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Nautili 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,{{Cite journal| last1 = Buchardt| first1 = B.| title = Diagenesis of aragonite from Upper Cretaceous ammonites: a geochemical case-study| journal = Sedimentology| volume = 28| issue = 3| pages = 423–438| year = 1981| doi = 10.1111/j.1365-3091.1981.tb01691.x| last2 = Weiner| first2 = S.|bibcode = 1981Sedim..28..423B }} nacreous and pressure-resistant, imploding at a depth of about {{cvt|800|m|ft|-2}}. The nautilus shell is composed of two layers: a matte white outer layer with dark orange stripes,{{cite web |url= https://marinesanctuary.org/blog/sea-wonder-chambered-nautilus/ |title= Sea Wonder: Chambered Nautilus |date= 21 August 2021 |website= National Marine Sanctuary Foundation |access-date= 26 November 2022}} 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.{{Citation needed|date=August 2024}}

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.

N. pompilius is the largest species in the genus. One form from Indonesia and northern Australia, once called N. repertus, may reach {{cvt|25.4|cm}} in diameter.Pisor, D.L. (2008). Registry of World Record Size Shells. Fifth edition. ConchBooks, Hackenheim. 207 pp. {{ISBN|0615194753}}. However, most nautilus species never exceed {{cvt|20|cm|in|0}}. Nautilus macromphalus is the smallest species, usually measuring only {{cvt|16|cm|in|frac=2}}. A dwarf population from the Sulu Sea (Nautilus pompilius suluensis) is even smaller, with a mean shell diameter of {{cvt|11.56|cm}}.

File:Nautilus.ogv

Image:Nautilus oceanworld thailand.png

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.{{Cite journal|title=Swimming mechanics and propulsive efficiency in the chambered nautilus|journal=Royal Society Open Science|volume=5|issue=2|page=170467|doi=10.1098/rsos.170467|pmc=5830708|pmid=29515819|year = 2018|last1 = Neil|first1 = T. R.|last2=Askew|first2=G. N.|bibcode=2018RSOS....570467N}} 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.{{Cite book|last=Askew|first=Graham|date=2017|title=Dataset associated with study on swimming mechanics and propulsive efficiency in the chambered nautilus|chapter-url=http://archive.researchdata.leeds.ac.uk/305/|language=en|publisher=University of Leeds|doi=10.5518/192|type=Data Set|chapter=B120}} 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 {{convert|800|m}}, and in fact implode at about that depth, causing instant death. The gas also contained in the chambers is slightly below atmospheric pressure at sea level.{{cite book |last1= Saunders |first1= W. Bruce |last2= Landman |first2=Neil H. |orig-date= 1987 |year=2009 |title= Nautilus: The Biology and Paleobiology of a Living Fossil | edition = 2nd |publisher= Springer Netherlands |page= 552 |isbn=978-90-481-3298-0 }} The maximum depth at which they can regulate buoyancy by osmotic removal of chamber liquid is not known.

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 {{convert|80|atm|psi|lk=in}}. 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.{{rp|p. 188}}

File:Nautilus pompilius (head).jpg]]

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.{{Cite journal| last1 = Grasso | first1 = F.| last2 = Basil | first2 = J.| title = The evolution of flexible behavioral repertoires in cephalopod molluscs| journal = Brain, Behavior and Evolution| volume = 74| issue = 3| pages = 231–245| year = 2009| pmid = 20029186| doi = 10.1159/000258669| s2cid = 13310728}}

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.

{{Main|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.{{cite magazine|url=https://www.newscientist.com/article/dn14033-simpleminded-nautilus-reveals-flash-of-memory.html|title=Simple-Minded Nautilus Shows Flash of Memory|author=Ewen Callaway|date=2 June 2008|magazine=New Scientist|access-date=7 March 2012}}{{cite journal|url=http://jeb.biologists.org/content/211/12/iii.full.pdf#page=1&view=FitH |archive-url=https://web.archive.org/web/20121029181222/http://jeb.biologists.org/content/211/12/iii.full.pdf |archive-date=2012-10-29 |url-status=live|title=Living Fossil Memories|author=Kathryn Phillips|date=15 June 2008|page=iii|volume=211|doi=10.1242/jeb.020370 |journal= Journal of Experimental Biology|issue=12|s2cid=84279320|doi-access=free|bibcode=2008JExpB.211Y...3P }}

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. 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.{{cite journal|author1=Robyn Crook |author2=Jennifer Basil |name-list-style=amp |year=2008|title=A biphasic memory curve in the chambered nautilus, Nautilus pompilius L. (Cephalopoda: Nautiloidea) |journal=The Journal of Experimental Biology|volume=211|pages=1992–1998|doi=10.1242/jeb.018531 |url=http://jeb.biologists.org/content/211/12/1992.full.pdf#page=1&view=FitH |archive-url=https://web.archive.org/web/20110817221242/http://jeb.biologists.org/content/211/12/1992.full.pdf |archive-date=2011-08-17 |url-status=live|issue=12 |pmid=18515730|s2cid=6305526 |doi-access=free|bibcode=2008JExpB.211.1992C }}

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 {{convert|30|mm|adj=on}} juveniles hatch.{{cite book |last1=Hanlon |first1=Roger T. |last2=Messenger |first2=John B. |title=Cephalopod Behaviour |date=1998 |publisher=Cambridge University Press |location=Cambridge, UK |isbn=9780521645836 |page=178}} Females spawn once per year and regenerate their gonads, making nautiluses the only cephalopods to present iteroparity or polycyclic spawning.{{Cite journal | last1 = Rocha | first1 = F. | last2 = Guerra | first2 = Á. | last3 = González | first3 = Á. F. | doi = 10.1017/S1464793101005681 | title = A review of reproductive strategies in cephalopods | journal = Biological Reviews of the Cambridge Philosophical Society | volume = 76 | issue = 3 | pages = 291–304 | year = 2001 | pmid = 11569786| s2cid = 5777682 }}

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.{{Cite journal | last1 = Bruce Saunders | first1 = W. | last2 = Spinosa | first2 = C. | title = Sexual Dimorphism in Nautilus from Palau | journal = Paleobiology | volume = 4 | issue = 3 | pages = 349–358 | year = 1978 |jstor=2400210| doi = 10.1017/S0094837300006047 | bibcode = 1978Pbio....4..349S | s2cid = 85899974 }} 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.

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.{{cite journal |author=Saunders WB |title=Nautilus Growth and Longevity: Evidence from Marked and Recaptured Animals |journal=Science |volume=224 |issue=4652 |pages=990–992 |date=June 1984 |pmid=17731999 |doi=10.1126/science.224.4652.990 |bibcode = 1984Sci...224..990S |s2cid=40891271 }} 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.{{cite journal |last1=Dunstan |first1=A.J. |last2=Ward |first2=P.D. |last3=Marshall |first3=N.J. |date=February 2011 |editor1-last=Solan |editor1-first=Martin |title=Nautilus pompilius life history and demographics at the Osprey Reef Seamount, Coral Sea, Australia |journal=PLOS ONE |volume=6 |issue=2 |pages=e16312 |pmid=21347356 |pmc=3037366 |bibcode = 2011PLoSO...616312D |doi=10.1371/journal.pone.0016312 |doi-access=free }}

Nautilus male has a reproductive organ named Van der Hoeven's organ. Nautilus female has two reproductive organs whose functions are unknown, the Organ of Valenciennes and Owen's laminated organ.{{cite book |author=Arthur Willey |url=https://books.google.com/books?id=Jwo4AQAAMAAJ&pg=PA778 |title=Zoological Results Based on Material from New Britain, New Guinea, Loyalty Islands and Elsewhere: The anatomy and development of Peripatus novae-britanniae |publisher=University Press |year=1902 |pages=778–9}}

{{multiple image

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| footer = 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.

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).

| image1 = Size frequency distribution for Nautilus pompilius at Osprey Reef.png

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File:Comparison of Nautilus capture rates with trapping depth.png. The data was collated from 271 trapping events spread across all months of the year. Nautiluses were most common at {{cvt|300|-|350|m|ft|-2}}. No specimens were recovered from a depth of less than {{cvt|150|m|ft|-2}} during 18 trapping efforts.]]

Nautiluses are found only in 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.{{Cite journal | last1 = Dunstan | first1 = A. J. | last2 = Ward | first2 = P. D. | last3 = Marshall | first3 = N. J. | editor1-last = Solan | editor1-first = Martin | title = Vertical distribution and migration patterns of Nautilus pompilius| doi = 10.1371/journal.pone.0016311 | journal = PLOS ONE | volume = 6 | issue = 2 | pages = e16311 | year = 2011 | pmid = 21364981| pmc =3043052 | bibcode = 2011PLoSO...616311D | doi-access = free }} The greatest depth at which a nautilus has been sighted is {{cvt|703|m|ft}} (N. pompilius). Implosion depth for nautilus shells is thought to be around {{cvt|800|m|ft|-2}}. Only in New Caledonia, the Loyalty Islands, and Vanuatu can nautiluses be observed in very shallow water, at depths of as little as {{cvt|5|m|ft|round=5}}. 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 {{cvt|100|m|ft|-2}}. Nautiluses generally avoid water temperatures above {{cvt|25|°C|round=5}}.

While nautiloids were once common worldwide, their numbers declined and their distribution became restricted to their current habitats during the Pleistocene, largely due to the diversification of pinnipeds.{{Cite journal |last=Kiel |first=Steffen |last2=Goedert |first2=James L. |last3=Tsai |first3=Cheng-Hsiu |date=2022 |title=Seals, whales and the Cenozoic decline of nautiloid cephalopods |url=https://onlinelibrary.wiley.com/doi/10.1111/jbi.14488 |journal=Journal of Biogeography |language=en |volume=49 |issue=11 |pages=1903–1910 |doi=10.1111/jbi.14488 |issn=1365-2699}}

File:Deepest record of Nautilus – 703 meters.png (Lutjanus bohar) bait during daytime at {{cvt|703|m|ft}} depth. This observation constitutes the deepest record of any nautilus species.]]

Nautiluses are scavengers and opportunistic predators.{{Cite journal | last1 = Saunders | first1 = W.B. | title = The role and status of Nautilus in its natural habitat: Evidence from deep-water remote camera photosequences | journal = Paleobiology | volume = 10 | issue = 4 | pages = 469–486 | doi = 10.1017/S0094837300008472| year = 1984 | jstor = 2400618| bibcode = 1984Pbio...10..469S | s2cid = 87096394 }}{{Cite journal | last1 = Wells | first1 = M. J. | last2 = Wells | first2 = J. | last3 = O'Dor | first3 = R. K. | doi = 10.1017/S0025315400037723 | title = Life at low oxygen tensions: The behaviour and physiology of Nautilus pompilius and the biology of extinct forms | journal = Journal of the Marine Biological Association of the United Kingdom | volume = 72 | issue = 2 | pages = 313–328 | year = 2009 | s2cid = 85601385 }} They eat lobster molts, hermit crabs, and carrion of any kind.{{cite book |last1= Ward |first1= P.D. |year= 1987 |title= The Natural History of Nautilus |location= London, United Kingdom |publisher= Allen and Unwin |isbn=9780045000364 }}

File:Allonautilus vs Nautilus.png

File:Section cut Nautilus.jpg

Fossil records indicate that nautiloids have experienced minimal morphological changes over the past 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 {{cvt|2.5|m|ft|0}} 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. The ancestors of all Coleoidea (shell-less Cephalopods) once possessed shells, and many early cephalopod species are only known from shell remains. Following the K-Pg extinction event most nautiloid species went extinct, while members of Coleoidea managed to survive. Following the mass extinction, the nautilus became the only extant species of nautiloids.{{Cite journal|url=https://www.academia.edu/93538950|title=Comparative morphology among representatives of main taxa of Scaphopoda and basal protobranch Bivalvia (Mollusca)|first=Luiz Ricardo L.|last=Simone|date=January 25, 2009|journal=Papéis Avulsos de Zoologia|volume=49|issue=32|pages=405–457|doi=10.1590/S0031-10492009003200001 |via=www.academia.edu|doi-access=free}}{{Cite journal|url=https://sjpp.springeropen.com/articles/10.1007/s13358-016-0112-7|title=Nautilus: biology, systematics, and paleobiology as viewed from 2015|first1=Peter|last1=Ward|first2=Frederick|last2=Dooley|first3=Gregory Jeff|last3=Barord|date=March 25, 2016|journal=Swiss Journal of Palaeontology|volume=135|issue=1|pages=169–185|via=sjpp.springeropen.com|doi=10.1007/s13358-016-0112-7|bibcode=2016SwJP..135..169W |s2cid=87025055 |url-access=subscription}}

The family Nautilidae has its origin in the Trigonocerataceae (Centroceratina), specifically in the Syringonautilidae of the Late Triassic and continues to this day with Nautilus, the type genus, and its close relative, Allonautilus.

File:Eutrephoceras dorbignyanum (Forbes in Darwin, 1846) - Santiago specimen.jpg]]

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.Teichert, C. & T. Matsumoto (2010). The Ancestry of the Genus Nautilus. In: W.B. Saunders & N.H. Landman (eds.) Nautilus: The Biology and Paleobiology of a Living Fossil. Springer. pp. 25–32. {{doi|10.1007/978-90-481-3299-7_2}}{{cite journal | last1 = Saul | first1 = L.R. | last2 = Stadum | first2 = C.J. | year = 2005 | title = Fossil argonauts (Mollusca: Cephalopoda: Octopodida) from Late Miocene siltstones of the Los Angeles Basin, California | journal = Journal of Paleontology | volume = 79 | issue = 3| pages = 520–531 | doi = 10.1666/0022-3360(2005)079<0520:FAMCOF>2.0.CO;2 | bibcode = 2005JPal...79..520S | s2cid = 131373540 }}

Image:Nautilus species shells.png

The family Nautilidae contains up to nine extant species and several extinct species:{{Cite journal|title = Miocene Nautilus (Mollusca, Cephalopoda) from Taiwan, and a review of the Indo-Pacific fossil record of Nautilus| date=2022 | doi=10.1111/iar.12442 | last1=Goedert | first1=James L. | last2=Kiel | first2=Steffen | last3=Tsai | first3=Cheng-Hsiu | journal=Island Arc | volume=31 | issue=1 | s2cid=247532223 | doi-access=free | bibcode=2022IsArc..31E2442G }}

• Genus Allonautilus
• A. perforatus
• A. scrobiculatus
• Genus Nautilus
• †N. altifrons
• †N. balcombensis
• N. belauensis
• †N. butonensis
• †N. campbelli
• †N. cookanus
• †N. geelongensis
• †N. javanus
• N. macromphalus
• N. pompilius (type)
• N. p. pompilius
• N. p. suluensis
• †N. praepompilius
• Nautilus samoaensis Barord et al., 2023Gregory J. Barord, David J. Combosch, Gonzalo Giribet, Neil Landman, Sarah Lemer, Job Veloso et Peter D. Ward, « [https://zookeys.pensoft.net/article/84427/ Three new species of Nautilus Linnaeus, 1758 (Mollusca, Cephalopoda) from the Coral Sea and South Pacific] », ZooKeys, vol. 1143, 2023, p. 51-69. – Samoa
• N. stenomphalus
• †Nautilus taiwanus Huang, 2002– Taiwan
• Nautilus vanuatuensis Barord et al., 2023 – Vanuatu
• Nautilus vitiensis Barord et al., 2023 – Fiji

Genetic data collected in 2011 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, though this was prior to the description of three additional species (samoaensis, vanuatuensis and vitiensis).

The following taxa associated with the family Nautilidae are of uncertain taxonomic status:Sweeney, M.J. 2002. [http://tolweb.org/accessory/Nautilidae_Taxa?acc_id=2324 Taxa Associated with the Family Nautilidae Blainville, 1825.] Tree of Life web project.

Nautilus are collected or fished for sale as live animals or to carve the shells for souvenirs and collectibles, not for just the shape of their shells, but also the nacreous inner shell layer, which is used as a pearl substitute.{{Cite journal |last1=Nijman |first1=Vincent |last2=Lee |first2=Paige Biqi |date=December 2016 |title=Trade in nautilus and other large marine molluscs as ornaments and decorations in Bali, Indonesia |url=https://www.researchgate.net/publication/311466511 |journal= Raffles Bulletin of Zoology |volume=64 |pages=368–373 |via= ResearchGate}}{{Cite journal|last=De Angelis |first=Patricia |year=2012 |title=Assessing the impact of international trade on chambered nautilus|journal=Geobios |volume=45 |issue=1 |pages=5–11 |via=Elsevier Science Direct |doi=10.1016/j.geobios.2011.11.005 |bibcode=2012Geobi..45....5D }}{{Cite journal |last1=Freitas |first1=B. |last2=Krishnasamy |first2=K. |title=An Investigation into the Trade of Nautilus |url=https://www.traffic.org/publications/reports/an-investigation-into-the-trade-of-nautilus/ |year=2016 |location=Washington DC, USA |journal=TRAFFIC Report |publisher=TRAFFIC Network / World Wildlife Fund }} In Samoa, nautilus shells decorate the forehead band of a traditional headdress called tuiga.{{Cite web |title=Tuiga: a Samoan ceremonial headdress |publisher=Museum of New Zealand |url=https://collections.tepapa.govt.nz/topic/10595 |access-date=Aug 23, 2023 }} Nautilus shells were popular items in the Renaissance and Baroque cabinet of curiosities and were often mounted by goldsmiths on a thin stem to make extravagant nautilus shell cups.

The low fecundity, late maturity, long gestation period and long life-span of nautiluses suggest that these species are vulnerable to overexploitation and demand for the ornamental shell is causing population declines.{{Cite journal|last1=Dunstan|first1=Andrew|last2=Bradshaw|first2=Corey J. A.|last3=Marshall|first3=Justin|date=2011-02-10|title=Nautilus at risk – Estimating population size and demography of Nautilus pompilius|journal=PLOS ONE|volume=6|issue=2|doi=10.1371/journal.pone.0016716|issn=1932-6203|pmc=3037370|pmid=21347360|page=e16716|bibcode=2011PLoSO...616716D|doi-access=free}} The threats from trade in these shells has led to countries such as Indonesia legally protecting the chambered nautilus with fines of up to US$8,500 and/or 5 years in prison for trading in this species. Despite their legal protection, these shells were reported to be openly sold at tourist areas in Bali as of 2014. The continued trade of these animals has led to a call for increased protection{{cite web|url=https://blogs.scientificamerican.com/extinction-countdown/nautilus-protection/|title=Nautilus Finally Moves toward Endangered Species Protection|last=Platt|first=John|website=Scientific American}} and in 2016 all species in Family Nautilidae{{cite web|url=https://cites.org/sites/default/files/eng/cop/17/prop/060216/E-CoP17-Prop-48%20Rev.pdf |archive-url=https://web.archive.org/web/20170118111839/https://cites.org/sites/default/files/eng/cop/17/prop/060216/E-CoP17-Prop-48%20Rev.pdf |archive-date=2017-01-18 |url-status=live|title=Inclusion of the Family Nautilidae|author1=Fiji|author2=India|author3=Palau|author4=the United States of America|year=2016|publisher=CITES|quote=because all species are being proposed for listing, this proposal seeks to list the Family Nautilidae in Appendix II.}} were added to CITES Appendix II, regulating international trade.{{cite web|url=https://www.fws.gov/le/pdf/12-21-2016-COP-17-Changes.pdf |archive-url=https://web.archive.org/web/20161228203239/https://www.fws.gov/le/pdf/12-21-2016-COP-17-Changes.pdf |archive-date=2016-12-28 |url-status=live|title=Notice to the Wildlife Import/Export Community}}{{cite magazine|url=https://blogs.scientificamerican.com/extinction-countdown/cites-cop17-wrapup/|title=Great News for Rhinos, Pangolins, Parrots, Sharks and Chambered Nautilus|last=Platt|first=John R.|date=2016-10-05|magazine=Scientific American|access-date=2017-01-15|quote=The chambered nautilus—victim of intense overharvesting for their beautiful shells—has been added to CITES Appendix II, which means all trade will now take place under a permit system, allowing the industry to be monitored for sustainability.}}

File:Berlín KGM nautilus.JPG|Baroque nautilus cup of Aleksander Kęsowski, abbot of Oliwa, 1643–1667

File:Nautilus shell carved and painted C19th Poldi Pezzoli Museum.jpg|Nautilus shell carved and painted with fanciful scenes of human figures and animals (spider, dragonfly, dog, butterfly, sawfly, fly), bronze pendant mount, nineteenth century. Poldi Pezzoli Museum, Milan

File:Nautilus cup in the form of a chicken.jpg|A nautilus shell in the form of a chicken, circa 1550

Palauans see nautili ({{langx|pau|kedarm}}) as a symbol of vulnerable or fragile character from a belief that they easily die even from slight bumps on ocean rocks; hence someone who gets quickly angry after being pranked is compared to one ({{lang|pau|ng ko er a kedarm, el di metirem e metord}}).{{cite journal |title=Proverbs of Palau |first1=Robert K. |last1=McKnight |journal=The Journal of American Folklore |date=1968 |volume=81 |issue=319 |pages=22 |doi=10.2307/537435|jstor=537435 }}

• Cephalopod size, for maximum shell diameters
• Historia animalium by Conrad Gessner, first book with fossil illustrations
• The Nautilus, a malacological journal
• The Chambered Nautilus, a poem of Oliver Wendell Holmes
• Nautilus Pompilius, a Russian rock band
• Nautilus (fictional submarine) from Jules Verne’s classic 1870 science-fiction novel Twenty Thousand Leagues Under the Seas, describing the voyage of Captain Nemo’s Nautilus submarine.

{{notelist}}

{{Reflist|colwidth=30em|refs=

{{cite web|author=Marcin Latka |title= Abbot Kęsowski's cup |url=https://artinpoland.weebly.com/en/abbot-kesowskis-cup |work=artinpl |access-date=26 July 2019}}}}

• Ward, P.D. 1988. In Search of Nautilus. Simon and Schuster.
• {{Cite book | title = Nautilus: the biology and palaeontology of a living fossil |author1=W. Bruce Saunders |author2=Neil H. Landman |name-list-style=amp |series=Topics in Geobiology |volume=6 |year=2010 | isbn = 978-90-481-3299-7 |location=Dordrecht |publisher=Springer Science+Business Media B.V | doi = 10.1007/978-90-481-3299-7 |s2cid=237330597 }}

{{CephBase Family|Nautilidae}}

• [https://www.tonmo.com/community/forums/nautilidae.58/ Nautilidae discussion forum], tonmo.com
• [https://web.archive.org/web/20060924223738/http://www.waquarium.org/MLP/root/pdf/MarineLife/Invertebrates/Molluscs/Nautilus.pdf Waikïkï Aquarium: Marine Life Profile: Chambered Nautilus], waguarium.org
• [https://web.archive.org/web/20050426132101/http://www.cephbase.utmb.edu/refdb/pdf/8036.pdf A molecular and karyological approach to the taxonomy of Nautilus], utmb.edu

{{Edible molluscs}}

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Category:Extant Triassic first appearances