Description

Phylogenetic Position:

Cephalopoda is a large class in which the majority of sub-classes and orders are extinct except the Coleoidea subclass (neocoleoids) and the Nautiloidea subclass. Nautilina is also the only sub-order of Nautilida with any living members.

The family Nautilidae contains two genera, Nautilus and Allonautilus, and six extant species in total. The six species are:

-Nautilus belauensis (Palau nautilus) (bottom left)
-Nautilus macromphalus (bellybutton nautilus) (centre)
-Nautilus pompilius pompilius/suluensis (chambered nautilus) (bottom right)
-Nautilus stenomphalus (white-patch nautilus) (left column)
-Allonautilus perforatus
-Allonautilus scrobiculatus (top right on right)


Because more is known of Nautilus sp and Allonautilus differ usually in only gills, male reproductive strategy or shell-ribbing, I will refer to Nautilus sp. more often. Unless noted otherwise, these qualities are expected to apply to Allonautilus sp.

Body form:

The shell, the same for Nautilus and Allonautilus, is formed of three layers of either organic periostracum (exterior) or calcareous aragonite crystals, is a long coil devided by septa into progressively smaller chambers, or camerae, towards the innermost chamber of the coil. This is the posterior end, and it is the oldest part of the shell, the rest of the shell was secreted with growth. In A. scrobiculatus, a thick encrusting layer of shell called periostracum is present. It is interesting to note that these shells are beautifully logarithmic with a ratio of 1: 1.618, often confused with the "Golden Ratio"


The soft body of Nautilus lives in only the most anterior region, called the body chamber. The soft body is considered everything but the shell, including all the viseral organs and the tentacles, which can be up to 90 in number! As the soft body grows and moves forward, the previous body chamber is filled with fluid, and shell is secreted to seal it except for at the siphuncle. The siphuncle is a calcareous tube enclosing a strand of tissue that is continuous throughout the shell. This allows gas to be pumped into the chambers through osmosis and gives Nautilus bouyancy as it increases in depth. Conversly, if the nautilus is climbing in depth towards to surface, it can pump fluid or gas into the chambers. Here is a diagram of all the major body parts.


Physiology, Ecology:

In order to swim, Nautilus must draw water into and out of the living chamber with the hyponome, or siphon (see left). Also known as jet propulsion! When water is inside the chamber, the siphuncle extracts salt from it and diffuses it into the blood. When water is pumped out, the siphuncle lessens the gas contained in the chamber in order to remain neutrally buoyant. The control of buoyancy in this manner limits the nautilus; they dont particularly do well in aquariums nor do they exist past depths of 800m due to the extreme hydrostatic pressure.
At the end of this entry, there is a video depicting the constantly jet-propulsed burst swimming and the tendency for nautiluses to remain near the top in aquariums. Because swimming does not involve any limbs in propulsion, they attribute very little energy to it.

Nautiluses are predators to fish, small shrimp and crustaceans and capture them using their the tentacles (see left). Because they swim so passively, they only require food once a month. The average lifespan for Nautiluses is irregularily high in comparison to other cephalopds at 20 years. In terms of reproduction they are iteroparous, they lay eggs (although never recorded in the wild), and are sexually dimorphic. A Nautilus sp. embryo removed from the egg is pictured on the left.

Unlike other cephalopods, they do not have good vision nor a highly developed brain. The structure of the eyes is highly modified but renders the visual quality alike that of a pin hole camera because it lacks a solid lens. For this reason, they are thought to rely more heavily on smell (olfaction).



Vast fossil record!!

Nautiloids are first seen in the fossil record in the Cambrian period. They left an abundant amount of fossils, suggesting the few species that remain alive today signify a great reduction in their diversity. Due to the inability of their soft bodies to fossilize, the shells are what's used to understand the evolution, classification and early nautiloid morphology.
Nautiloids are the only cephalopod with a true external shell alive today (most have an internal shell or no shell at all).The first cephalopods likely had high, curved, conical, septate shells (like Goniatites) which were lost in most lineages. The extant species of Nautilus are thought to be very similar in shell to their ancestors.
Here are some of the more interesting fossils and their representations of mostly ancestral ammonids (cephalopods) indicative of their past diversity. More information on these drawings and more of them can be found here.







Annotated Scientific Papers about the Nautilus

1. Shigeno, S., Takenori S., Takeya M. et al. 2007. Evolution of the cephalopod head complex by assembly of multiple molluscan body parts: Evidence from Nautilus embryonic development. (Nautiloidea:Nautilida:Nautilina:Nautilidae). Journal of Morphology. Published online 07.24.2007. PMID: 17654542.

  • Can certain primitive features in embryonic stages of Nautilus pompilius propose an alternative scheme for the evolution of cephalopod body plans, specifically of the morphology of the head parts? Previous evolutionary hypotheses have been proposed on the basis of combined embryological and paleontological observations as they tend to contrast. The recent onset of studies on the morphology of Nautilus embryos offered a new opportunity to test these hypotheses. as differences in Nautilus and coleiod cephalopod embryos were noted.
  • The evolution of the head complex, or the complicated incorporation of many long tentacles or arms, cupshaped eyes, statocysts, sense organs, a centralized brain and funnel in the cephalopod head, is intriguing as it seems to represent the pivotal transition from benthos to nekton. Because embryological hypotheses were previously based on coleiod cephalopods, the opportunity to study Nautilus embryology would test the universality of such hypotheses. Although there were no previous studies to suggest Nautilus embryology would be more informative of the evolution of the cephalopod head complex to date, the fact its embryology was preliminarily different than of coleiods suggested it might be.
  • Embryos of both Nautilus pompilius and of the already-studied Idiosepius paradoxus squid were compared. The results demonstrated, among many things, that 1) the distinct cephalic compartment, foot, brain cords, mantle, and shell resembled the body
    plans of monoplacophorans and basal gastropods, 2) origins of digital tentacles or arms of all other cephalopods develop not from the head but from the foot, silencing previous debates and that 3) formation of a ‘‘head complex" assembled in mid to late embryos when the primary body plans changed.
  • This was the first evidence from embryology that the morphological development of the head complex was via the assembly of multiple archetypical molluscan body parts and support that this assembly was based on transitional niche ecology.

2. Ward, P.D., Saunders, W.B. 1997. Allonautilus: A new genus of living nautiloid cephalopod and its bearing on phylogeny of the Nautilida. (Nautiloidea:Nautilida:Nautilina:Nautilidae). Journal of Paleontology. 17: 1054-1064.

  • Will soft-, as well as hard-part anatomy differ in the currently defined species of Nautilus and what light can morphological data shed on the phylogeny as opposed to genetic?Previous studies were based solely on genetic data, and had concluded that N. scrobiculatus was so distinct from the rest of the Nautilus genus that it should be separated into two unifying taxa.
  • A century and a half has passed that shell shape and colouration have been known to be different in N. scrobiculatus and the rarely discussed N. perforatus but their distinction was never formalized. Now that unambiguous soft-part observations can be made across three extant and numerous species of the genus, they may depict an intirely new phylogeny.
  • N. scrobiculatus, N. pompilius, N. belauensis were used for comparison of gross anatomy and male reproductive system while comparisons of gross shell morphology was referenced. Extinct species of N. cookanum and other outgroup nautilacean fossils were also examined. In total there were 30 character sets on which to determine philogeny twenty of which were informative. The cladistic analysis sucessfully distinguished N. scrobiculatus from N. perforatus as two separate species deserving of a seperate genus from the rest of the extant Nautilus species. Fossil records suggested that these two species are dervied from the Nautilus genus.
  • These findings lead Ward et al. to form the Allonautilus genus composed of A. scrobiculatus and A. perforatus. and place them to be more derived than the species of the Nautilus genus. This study exemplifies how genetic data in combination with morphological character cladistics can yield the best view of phylogeny.

3. Sharp, J. 2002. Spirals and the Golden Section. Nexus Network Journal, 4:59-82. DOI 10.1007/s00004-001-0005-x

  • Many types of spirals are known from mathematics, but does a logarithmic spiral always consist of a Golden Section spiral? After a detailed description of various spirals and Golden Section geometry, the Nautilus shell spiral is correctly employed to help answer this question because it is often considered a Golden Section spiral by the public.
  • The misunderstanding of Golden Section spirals often grows with facisnation, if the person has a lack of knowledge regarding types of spirals and mathematics behind them. Theusual logic of someone in such a situation ignores this misunderstanding in making great deductions. This has spread a common myth about natural logarithmic spirals, inlcuding that of Nautilus sp.
  • Sharp reviewed detailed properties of different spiral types, the abilityof spirals to vary based only on the circles from which they are approximated, and the mathematics of Golden Section spirals before attempting to justify the myth of the Nautilus shell. This preliminary review is crucial and successful in proving that the Nautilus shell is not a Golden Section spiral despite that it is logarithmic (inherent mathimatical differences are explained).
  • Sharp concludes with confidence that the Nautilus shell is not a Golden section spiral. This in turn prooves that no, logarithmic spirals are not all Golden Section spirals (spirals with a ratio of 1:1.618 ) contrary to popular misconception. This study nicely exemplifies the ability of new mathematics and geometry to dispel long-held biology myths.

Header image artwork is the intellectual property of Roger Hall as part of Inkart productions. It is in use for decorative purposes only and can be found in its original context here.