Cephalopod dermal structures

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LepidoJoubin2 (rotated).jpg
Lepidoteuthis grimaldii scales.jpg
Mantle of Lepidoteuthis grimaldii with the end of the dermal scales indicated by an arrow. A closeup of the scales is shown on the right. This was the first cephalopod species whose dermal structures were studied in detail. [1] [2] [3]

Cephalopods exhibit various dermal structures on their mantles and other parts. These may take the form of conspicuous warts, cushions, papillae or scales, though in many species they are microscopic tubercles. [4] The most elaborate forms are found among the oceanic squid of the order Teuthida. [1] [5]

Contents

Morphology and composition

Most cephalopod dermal structures take the form of tubercles, and these are the only cartilaginous dermal structures (the various "dermal cushions" being composed of other forms of connective tissue). All three main types of cartilage found in vertebrates are represented among the different squid species: hyaline, elastic and fibrocartilage. Tubercles of hyaline cartilage are primarily associated with cranchiid or glass squid. [1]

The vast majority of cephalopod dermal structures have a thin, overlying epidermal layer, though this is often damaged or missing in captured specimens. [1]

Dermal structures of selected squid species [1]
SpeciesFamilyStructureShapeSize in mm ( ML in mm)TissueEpidermis over structure
Asperoteuthis acanthoderma Chiroteuthidae TuberclesConical1.0 × 0.4 (144)Hyaline-like cartilageYes
Cranchia scabra Cranchiidae TuberclesRound, triangular, or rectangular bases with 2–5 projections0.4–0.8 × 0.2–0.4 (94)Hyaline cartilageYes
Histioteuthis meleagroteuthis Histioteuthidae TuberclesKnob-likefrom mantle ridge: 0.4–0.8 × 1.3–1.9; from arm ridge: 0.6–2.2 × 0.7–2.3 (38)Elastic cartilageYes
Leachia cyclura Cranchiidae TuberclesSmall conical to large complex with numerous conical projections0.3–0.4 × 0.1–0.2 (55)Hyaline cartilageYes
Lepidoteuthis grimaldii Lepidoteuthidae Dermal cushions ("scales")Rhomboid to hexagonal2.0 × 0.5 (180)Connective tissue in honeycomb arrangementYes
Liocranchia reinhardti Cranchiidae TuberclesConical0.2–0.3 × 0.15–0.2 (29)Hyaline cartilageYes
Mastigoteuthis cordiformis Mastigoteuthidae TuberclesConical0.2 × 0.3 (87)Elastic or fibrocartilageYes
Mastigoteuthis hjorti Mastigoteuthidae TuberclesConical0.1–0.2 × 0.05–0.1 (93)Elastic or fibrocartilageYes
Pholidoteuthis adami Pholidoteuthidae Dermal cushionsRound to pentagonal0.5 × 0.3 (300)Connective tissue in honeycomb arrangementYes
Pholidoteuthis massyae Pholidoteuthidae Tubercles and papillaeRoundish0.3 × 0.15 (100)Dense connective tissue with chondrocytes; elastic cartilageNo; acellular cuticle

Other cephalopods with prominent dermal structures include: Brachioteuthis spp.; Galiteuthis glacialis (which has round tubercles); Histioteuthis meleagroteuthis and Histioteuthis miranda (tuberculate ridges); Mastigoteuthis danae (large tubercles in advanced paralarvae); [6] Onykia aequatorialis , Onykia ingens , Onykia lonnbergii , and Onykia robsoni (irregular warts); [7] Slosarczykovia circumantarctica (fibrous integumental net); [8] and all members of the subfamily Cranchiinae (strips of cartilaginous tubercles in various arrangements). [1] Among octopuses, Ocythoe tuberculata is noted for the tubercles and ridges on its mantle. [1]

Function

The mantle of Cranchia scabra is covered in numerous multi-pointed cartilaginous tubercles Cranchia scabra.jpg
The mantle of Cranchia scabra is covered in numerous multi-pointed cartilaginous tubercles
Ventral view of an adult Galiteuthis glacialis from the Ross Sea, with a mantle length of 32.1 cm Galiteuthis glacialis (Ross Sea, Antarctica).jpg
Ventral view of an adult Galiteuthis glacialis from the Ross Sea, with a mantle length of 32.1 cm

Different cephalopod dermal structures are hypothesised to play roles in buoyancy, locomotion, and even pseudoskeletal support. [1]

Buoyancy aid

Two fundamentally different buoyancy mechanisms associated with dermal structures have been proposed. [1]

Buoyancy vest

The mantle of Cranchia scabra is covered in multi-pointed cartilaginous tubercles. [9] [10] An anti-predator function has been proposed in the past, but this is thought unlikely given the small size and transparent nature of the tubercles. The tubercles of this species are covered by a thin, epidermal sheath that is often lost during capture. It has been speculated that in the live animal the interstitial space is filled with a buoyant fluid (likely an ammonium chloride solution) and acts as a "buoyancy vest". The hard tubercles may serve to maintain the shape of this structure. It has been estimated that in a C. scabra measuring 10 cm in mantle length (ML), the buoyancy vest could contribute an additional 4% to the animal's total buoyant fluid (most of the remainder being located in the coelom), probably sufficient to achieve neutral buoyancy. [1]

A similar mechanism may be utilised by the much larger Galiteuthis glacialis , which has a very similar combination of tubercles and overlying epidermal sheath. [1]

Lepidoteuthis grimaldii female measuring 61.7 cm in mantle length and weighing more than 4 kg Lepidoteuthis grimaldii 617 mm ML.jpg
Lepidoteuthis grimaldii female measuring 61.7 cm in mantle length and weighing more than 4 kg
Closeup of the overlapping dermal scales of the same specimen Lepidoteuthis grimaldii dermal scales.jpg
Closeup of the overlapping dermal scales of the same specimen

Fluid-filled dermal cushions

The overlapping "scales" of Lepidoteuthis grimaldii are actually dermal cushions with a vacuolate internal structure that are continuous with a similarly vacuolate underlying layer of mantle tissue. [1] [11] Ammonium ions (NH4+) are present in the mantle of this species at a measured concentration of 172  mM. [12] Structurally very similar (though non-overlapping) dermal cushions are found in Pholidoteuthis adami . [1] [13] [14] It has been proposed that these two species achieve buoyancy by means of the fluid stored in their vacuolate dermal cushions and upper mantle layer. [1] Given their spongy form, these cushions may also play a secondary protective role. [1]

Drag reduction

Pholidoteuthis massyae (41.1 cm ML) Pholidoteuthis massyae 411 mm ML.jpg
Pholidoteuthis massyae (41.1 cm ML)

The complex [15] dermal structures of Pholidoteuthis massyae may play a role in reducing hydrodynamic drag. More specifically, they may be involved in maintaining laminar flow by preventing or delaying boundary layer separation along the mantle. [1]

It is possible that a similar locomotory mechanism is present in Mastigoteuthis cordiformis and Mastigoteuthis hjorti , though the small size of the tubercles in these species may preclude such a function. [1]

Pseudoskeletal support

Liocranchia reinhardti Liocranchia reinhardti.jpg
Liocranchia reinhardti

In the cranchiids Leachia cyclura and Liocranchia reinhardti , the dermal tubercles are not distributed throughout the mantle but arranged in discrete cartilaginous bands. A role in buoyancy control is therefore unlikely. One possibility is that these rigid bands play a pseudoskeletal role, maintaining the shape of parts of the mantle during swimming contractions or providing attachment points for mantle muscles or visceral tissue (such as the septum covering the coelom). [1]

The very dense tuberculate ridges found on the arms and dorsal mantle of Histioteuthis meleagroteuthis may similarly provide insertion points for muscles, and are probably most important in juvenile animals, which lack well-developed musculature. [1] In the two other Histioteuthis species with tuberculate ridges— H. meleagroteuthis and H. miranda —these structures likely have the same function. [1]

Unknown function

The mantle of Asperoteuthis acanthoderma is covered in minute, widely spaced tubercles of hyaline-like cartilage. In a 1990 study of dermal structures in squid, Clyde F. E. Roper and C. C. Lu wrote that they were "unable to suggest a function" for the tubercles of this species, but that due to their small size and spacing they were unlikely to be involved in buoyancy or locomotion. [1]

Related Research Articles

<span class="mw-page-title-main">Squid</span> Superorder of cephalopod molluscs

A squid is a mollusc with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting these criteria. Like all other cephalopods, squid have a distinct head, bilateral symmetry, and a mantle. They are mainly soft-bodied, like octopuses, but have a small internal skeleton in the form of a rod-like gladius or pen, made of chitin.

<span class="mw-page-title-main">Cephalopod</span> Class of mollusks

A cephalopod is any member of the molluscan class Cephalopoda such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral body symmetry, a prominent head, and a set of arms or tentacles modified from the primitive molluscan foot. Fishers sometimes call cephalopods "inkfish", referring to their common ability to squirt ink. The study of cephalopods is a branch of malacology known as teuthology.

<i>Spirula</i> Species of cephalopod known as the rams horn squid

Spirula spirula is a species of deep-water squid-like cephalopod mollusk. It is the only extant member of the genus Spirula, the family Spirulidae, and the order Spirulida. Because of the shape of its internal shell, it is commonly known as the ram's horn squid or the little post horn squid. Because the live animal has a light-emitting organ, it is also sometimes known as the tail-light squid.

<span class="mw-page-title-main">Whip-lash squid</span> Family of cephalopods known as whip-lash squid

The Mastigoteuthidae, also known as whip-lash squid, are a family of small deep-sea squid. Approximately 20 known species in six genera are represented, with members found in both the mesopelagic and bathypelagic zone of most oceans. Originally described by Verill in 1881, it was later lowered by Chun (1920) to a subfamily (Mastigoteuthinae) of the Chiroteuthidae. However, Roper et al. (1969) raised it back to the family level, and this has not been changed since. The taxonomy of this family is extremely unstable, and there have been at times one genus, two genera and four subgenera(Salcedo-Vargas & Okutani, 1994), two genera and several 'groups', five genera and one species with an uncertain placement, or six genera.

<span class="mw-page-title-main">Hooked squid</span> Family of squids

The hooked squid, family Onychoteuthidae, currently comprise about 20–25 species, in six or seven genera. They range in mature mantle length from 7 cm to a suggested length of 2 m for the largest member, Onykia robusta. The family is characterised by the presence of hooks only on the tentacular clubs, a simple, straight, funnel–mantle locking apparatus, and a 'step' inside the jaw angle of the lower beak. With the exception of the Arctic Ocean, the family is found worldwide.

<i>Lepidoteuthis grimaldii</i> Species of squid

Lepidoteuthis grimaldii, also known as the Grimaldi scaled squid, is a large squid growing to 1 m in mantle length. It is named after the Grimaldi family, reigning house of Monaco. Prince Albert I of Monaco was an amateur teuthologist who pioneered the study of deep sea squids by collecting the 'precious regurgitations' of sperm whales. The Grimaldi scaled squid was first collected from the stomach contents of a sperm whale. It is a widely distributed species in tropical and subtropical areas of the North and South Atlantic, the southern Indian Ocean and the Pacific Ocean, where it has been recorded off Japan and in the west Pacific.

<span class="mw-page-title-main">Myopsida</span> Order of squids

Myopsida is one of the four orders of squid. It consists of two families: the monotypic Australiteuthis and the diverse and commercially important Loliginidae. Some taxonomists classify this taxon as a suborder of the order Teuthida, in which case it is known as Myopsina. This reclassification is due to Myopsina and Oegopsina not being demonstrated to form a clade.

<span class="mw-page-title-main">Histioteuthidae</span> Family of cephalopods

Histioteuthidae is a family of Oegopsid squid. The family was previously considered to be monotypic but the World Register of Marine Species assigns two genera to this family.

<i>Onykia ingens</i> Species of squid

Onykia ingens, the greater hooked squid, is a species of squid in the family Onychoteuthidae. It occurs worldwide in subantarctic oceans.

<i>Onykia robusta</i> Species of cephalopod known as the robust clubhook squid

Onykia robusta, also known as the robust clubhook squid and often cited by the older name Moroteuthis robusta, is a species of squid in the family Onychoteuthidae. Reaching a mantle length of 2 m (6.6 ft), it is the largest member of its family and one of the largest of all cephalopods. The tentacular clubs are slender, containing 15–18 club hooks. Arms of the species contain 50–60 suckers, and grow to 90–100% of the mantle length. It is found primarily in the boreal to Temperate Northern Pacific.

<i>Australiteuthis</i> Genus of squids

A. aldrichi is a small species of squid found in northern Australian waters. The species was described by Chung Cheng Lu in 2005 based on specimens collected in the inshore waters of Northern Australia. The largest known individual of this species is a mature female measuring 27.6 mm (1.09 in) in mantle length (ML). The holotype is a mature male of 21.3 mm (0.84 in) ML. A live specimen of A. aldrichi has yet to be recorded. A. aldrichi is a member of the class Cephalopoda and part of the subclass Coleodia. Within this class there are two orders, the Myopsida and Oegopsida, which both fall under the superorder Decapodiformes. A. aldrichi falls under the order of Myospida, and is the only member of its genus, Australiteuthis, and family, Australiteuthidae.

<i>Magnapinna talismani</i> Species of squid

Magnapinna talismani is a species of bigfin squid known only from a single damaged specimen. It is characterised by small white nodules present on the ventral surface of its fins.

<i>Asperoteuthis acanthoderma</i> Species of squid

The thorny whiplash squid, known as Asperoteuthis acanthoderma is a large species of squid belonging to the family Chiroteuthidae. It is characterised by the tiny, pointed tubercules present on its skin and a Y-shaped groove in the funnel locking apparatus.

<i>Histioteuthis bonnellii</i> Species of cephalopod

Histioteuthis bonnellii, the umbrella squid, is a species of cock-eyed squid belonging to the family Histioteuthidae.

<i>Sepiolina nipponensis</i> Species of mollusc

Sepiolina nipponensis, also known as the Japanese bobtail squid, is a bobtail squid and one of two species in the genus Sepiolina. It is found in the Western Pacific in apparently widely separated populations, the most southerly of which is in the Great Australian Bight in South Australia and Western Australia, and there are populations from the Philippines northwards to Taiwan, Fujian and southern Honshū.

<span class="mw-page-title-main">Funnel–mantle locking apparatus</span> Structure found in many cephalopods

The funnel–mantle locking apparatus is a structure found in many cephalopods that connects the mantle and hyponome (funnel) and restricts their movement relative to each other. It consists of two interlocking components: one located on the mantle and the other on the funnel. The apparatus may permit some anterior–posterior displacement or prevent movement altogether.

<i>Teuthowenia megalops</i> Species of squid

Teuthowenia megalops, sometimes known as the Atlantic cranch squid, is a species of glass squid from the subarctic and temperate waters of the northern Atlantic Ocean. They are moderately sized squid with a maximum mantle length of 40 cm (16 in). Their very large eyes are the source for the specific name megalops. Like other members of the genus Teuthowenia, they are easily recognizable by the presence of three bioluminescent organs (photophores) on their eyeballs.

<span class="mw-page-title-main">Cephalopod fin</span>

Cephalopod fins, sometimes known as wings, are paired flap-like locomotory appendages. They are found in ten-limbed cephalopods as well as in the eight-limbed cirrate octopuses and vampire squid. Many extinct cephalopod groups also possessed fins. Nautiluses and the more familiar incirrate octopuses lack swimming fins. An extreme development of the cephalopod fin is seen in the bigfin squid of the family Magnapinnidae.

<i>Slosarczykovia</i> Genus of squids

Slosarczykovia is a monotypic genus of squid, its sole representative being Slosarczykovia circumantarctica. Slosarczykovia is placed in the family Brachioteuthidae.

Histioteuthis meleagroteuthis is a species of small to medium squids that have a dark, wine-red skin pigment. Females at maturity average at 114 mm (4.5 in) in length, while males at maturity average at 65 to 102 mm in length. This species is characterized by tubercles, photophores, and asymmetric features. This species can be found in circumglobal, mesopelagic waters.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Roper, C.F.E. & C.C. Lu (1990). "Comparative morphology and function of dermal structures in oceanic squids (Cephalopoda)" (PDF).Smithsonian Contributions to Zoology, Number 493: 1–40.
  2. Clarke, M.R. (1960). Lepidoteuthis grimaldii—a squid with scales. Nature188: 955–956 doi:10.1038/188955a0
  3. Clarke, M.R. & G.E. Maul (1962). A description of the "scaled" squid Lepidoteuthis grimaldi Joubin 1895. Proceedings of the Zoological Society of London139(1): 97–118. doi:10.1111/j.1469-7998.1962.tb01824.x
  4. Young, R.E., M. Vecchione & K.M. Mangold (2001). Cephalopod Mantle Armature. Tree of Life Web Project.
  5. Roper, C.F.E. & C.C. Lu (1989). "Systematic status of Lepidoteuthis, Pholidoteuthis and Tetronychoteuthis (Cephalopoda: Oegopsida)" (PDF). Archived from the original (PDF) on 2013-04-02. Retrieved 2012-12-31.Proceedings of the Biological Society of Washington102(3): 805–807.
  6. Vecchione, M. & R.E. Young (2007). Mastigoteuthis danae (Joubin, 1933). Tree of Life Web Project.
  7. Bolstad, K.S.R., M. Vecchione & R.E. Young (2011). Onykia Lesueur, 1821. Tree of Life Web Project.
  8. Lipinski, M. & R.E. Young (2011). Slosarczykovia. Tree of Life Web Project.
  9. Person, P. (1969). Cartilaginous dermal scales in cephalopods. Science164(3886): 1404–1405. doi : 10.1126/science.164.3886.1404
  10. Dilly, P.N. & M. Nixon (1976). The dermal tubercles of Cranchia scabra (Mollusca, Cephalopoda); surface structure and development. Journal of Zoology179(3): 291–295. doi : 10.1111/j.1469-7998.1976.tb02297.x
  11. Young, R.E. & M. Vecchione (1999). Lepidoteuthis grimaldii: Dermal Cushions. Tree of Life Web Project.
  12. Lipiński, M. & K. Turoboyski (1983). The ammonium content in the tissues of selected species of squid (Cephalopoda: Teuthoidea). Journal of Experimental Marine Biology and Ecology69(2): 145–150. doi : 10.1016/0022-0981(83)90064-3
  13. Goldman, D.A. (1995). A juvenile of the scaled squid, Pholidoteuthis adami Voss, 1956 (Cephalopoda: Oegopsida), from the Florida Keys. Proceedings of the Biological Society of Washington108(1): 136–146.
  14. Young, R.E. & M. Vecchione (1999). Pholidoteuthis adami Dermal Cushions. Tree of Life Web Project.
  15. Young, R.E. & M. Vecchione (1999). Pholidoteuthis massyae Mantle Dermal Cushions. Tree of Life Web Project.