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Bigfin Reef Squid

Sepioteuthis lessoniana d'Orbigny 1826

Behavior

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Bigfin reef squid are typically solitary, limiting the amount of communication that occurs between individuals. However, when in close proximity to others, as in feeding groups or breeding shoals, this species does display distinct communication behaviors. It uses rapid, vivid changes in its body color pattern (via chromatophores) to convey information about potential threats, food sources, or dominance. Physical intimidation, such as chasing or biting of other individuals, as well as additional visual displays of dominance, such as genital presentation and limb-spreading, have also been observed in shoals and in mating groups. They do not possess any specialized organs for sound detection or production (although vibrations are perceived), instead relying upon their highly developed sense of sight for communication and perception of their environment. In addition, bigfin reef squid possess sensitive chemoreceptors near their mouth and on the suckers (more developed in octopi and cuttlefish), which helps to distinguish between prey and non-prey items.

Communication Channels: visual ; tactile ; chemical

Other Communication Modes: photic/bioluminescent ; vibrations

Perception Channels: visual ; polarized light ; tactile ; vibrations ; chemical

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
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Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Associations

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Some common predators of bigfin reef squid are large fish, such as blue-fin (Thunnus orientalis) and skipjack (Katsuwonus pelamis) tuna, Atlantic blue marlin (Makaira mazara), and sharks, as well as birds, whales, and humans. They are most vulnerable to predation during larval stages. Squid hatchlings are often cannibalistic, which is one of the main sources of predation on young squid. However, the rate of cannibalism is lower in this species than in other squid species.

Newly hatched young will swim in shoals and can interact with one another to increase chances of survival, by warning each other of predators and helping one another find food sources. During all life stages, bigfin reef squid rely on camouflage, changing their coloration based on their surroundings using their chromatophores. They may also use differently polarized skin patterns as a means of communication between individuals, which would allow them to communicate with each other while remaining camouflaged from predators that are unable to perceive polarized light. The ink produced by threatened squid can cause predators to lose their sight and sense of smell, due to the color and alkalinity of the ink.

Known Predators:

  • blue-fin tuna (Thunnus orientalis)
  • skipjack tuna (Katsuwonus pelamis)
  • albacore tuna (Thunnus alalunga)
  • Atlantic blue marlin (Makaira mazara)
  • oval squid (Sepioteuthis lessoniana)

Anti-predator Adaptations: cryptic

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
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Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Morphology

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Bigfin reef squid possess a characteristic cephalopod body plan that is fusiform in shape, with a large mantle, attached head, and multiple arms. The genus name Sepioteuthis describes its cuttlefish (Sepia officinalis)-like appearance. Its main body mass is contained in the mantle (sometimes referred to as the "hood"), where the organs for reproduction, respiration, digestion, circulation, excretion, and propulsion are found. The mantle is a highly muscular organ which surrounds the entire body of the squid. The muscles of the mantle are made of two different types (circular and radial) of fibers that have different orientations. The majority of the mantle is made up of the circular muscle fibers. The remnant of an ancestral shell is also found within the mantle, and is known as the internal gladis (commonly called the "pen"). The distinguishing "big fins" referred to in the common name are located on the superior portion of the mantle. The fins extend along over 90% of the length of the mantle and give the squid a characteristic oval appearance. The maximum reported mantle length in males is 422 mm and 382 mm in females. The mantle length is regularly 200-300 mm. When fully grown, this small squid can weigh anywhere from 1 pound to 5 pounds.

The head contains the brain, eyes, and buccal mass (the beak/mouth and salivary glands) and is located below the mantle. Squid species have complex eyes that are capable of focusing and forming clear images. The pupils are adjustable, based on the amount of light entering the eye. Eight arms (not generally of identical length) and two tentacles are attached to the bottom portion of the head. The tentacles are primarily used for the capture of prey. Each of the arms is lined with toothed suction cups for manipulating prey, and the two tentacles possess toothed suction cups at their distal end.

Between the head and the mantle is the hyponome, which is a funnel that water is forced through as a means of propulsion. The giant axon controls part of the water jet propulsion system in squid. Action potentials in the giant axon cause contraction of mantle muscle fibers, forcing water though the hyponome. The squid can adjust the position of the hyponome to change the direction of propulsion.

Oxygen and carbon dioxide are exchanged at the gills, which are located within the mantle. Bigfin reef squid (and other squid species) have a closed circulatory system with three hearts, two of which are brachial hearts found at the base of their gills. These are used to pump oxygen deficient blood through the gills, where carbon dioxide in the blood is exchanged for oxygen and pumped out of the body through the siphon. The third heart pumps oxygenated blood throughout the rest of the body. Hemocyanin, rather than hemoglobin, is the oxygen carrying protein in this species' blood. Hemocyanin is copper based, making the blood appear blue.

Like all cephalopods, bigfin reef squid have a complex digestive system, consisting of a stomach, a caecum, and a liver. Following digestion and absorption of nutrients by these structures, solid waste is passed out of the rectum. Both the mantle and the digestive gland tissues are used in energy storage.

Energy expenditure shifts towards the reproductive system gains priority as squid mature. Female bigfin reef squid have a large ovary (which can be seen through the transparent mantle tissue) and a nidamental gland for the production of egg cases and nutrients. Males have a large testis and a spermatophore gland and sac. Males also have modified arms, with the presence of hectocotyli to store spermatophores. This is one of the key characteristics used in determining the sex of individuals.

Bigfin reef squid can control their pigmented skin cells, called chromatophores, to rapidly change their body color and pattern. This species also has an ink sac, which can release a dark cloud of ink to disorient predators.

Range mass: 100 to 1400 g.

Average mass: 400 g.

Range length: 4 to 33 cm.

Other Physical Features: endothermic ; ectothermic ; heterothermic ; bilateral symmetry

Sexual Dimorphism: male larger; sexes colored or patterned differently; ornamentation

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
author
Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Life Expectancy

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Bigfin reef squid generally complete their entire life cycle within 4 to 6 months of birth. Water temperature may affect rates of growth and sexual maturation in squid. In hot, equatorial waters, they have a shorter lifespan and reach smaller body sizes than individuals that live in “cooler” subtropical waters.

Range lifespan
Status: captivity:
315 (high) days.

Average lifespan
Status: wild:
171 days.

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
author
Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Habitat

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Bigfin reef squid live in warm (typically 16°C to 34°C) coastal waters. At night, when they are most active, they are commonly found in shallow areas (0 to 100 m) around reefs, sea grass beds, sandy bottoms, or rocky shorelines. At the shore, where there are greater fluctuations in salinity, bigfin reef squid are found in waters varying from 28 ppt to 36 ppt. They surface at night, when they are less likely to be detected by predators. During the day, they generally move to deeper waters or near any form of cover, such as floating driftwood, reefs, rocks, or grasses.

Range depth: 0 to 100 m.

Habitat Regions: tropical ; saltwater or marine

Aquatic Biomes: reef ; coastal

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
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Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Distribution

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This species, known as either the bigfin reef or oval squid, is generally found in Indo-West Pacific waters, including the tropical waters of the Indian Ocean around the Red Sea and the seas between northern Australia, New Zealand and Asia, which connect the Indian and Pacific Oceans. Oval squid also occur as far north as the Mediterranean Sea and in eastern portions of the Pacific Ocean near the Hawaiian Islands.

Biogeographic Regions: australian (Native ); indian ocean; pacific ocean (Native ); mediterranean sea (Native )

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Trophic Strategy

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Bigfin reef squid are strictly carnivorous. They primarily feed on other mollusks and fish, but also consume non-insect arthropods, zooplankton, and other marine invertebrates, depending upon life cycle stage and prey availability. This species has also been known to cannibalize smaller conspecifics. Squid use their specialized tentacles to capture live prey. The tentacles are then used to guide the prey to the specialized buccal region and the sharp, muscular beak, which is specialized for the tearing and crushing of prey.

Animal Foods: fish; mollusks; aquatic crustaceans; other marine invertebrates; zooplankton

Primary Diet: carnivore (Piscivore , Molluscivore )

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Associations

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Bigfin reef squid are predators of a number of species, primarily pisfishes and mollusks, but also eat zooplankton and other marine invertebrates. This species also serves as prey to a number of common predatory species. They can consume up to 50% of their body mass during their growth period and they have fast metabolic and growth rates, so they assist in the transfer of energy through trophic levels of the food web in marine ecosystems.

Commensal/Parasitic Species:

  • copepods (Doridicola similis)
  • Dicyema koshidai (Phylum Dicyemida)
  • Dicyema orientale (Phylum Dicyemida)
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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Benefits

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This species is among the most commercially important squid species. Since it is not a seasonal species, it can be fished throughout the year. It is used not only for food, but also as bait for many other species. Since bigfin reef squid have a rapid growth rate, short life span, low rates of disease, low rates of cannibalism, and tolerance to handling and captivity, they are commonly observed and raised in labs. This not only assists in learning about the lifecycle and reproduction of the squid, but also aids in determining the effects of temperature, salinity, and different food sources on the squid. Another positive aspect of raising bigfin reef squid in labs is that they offer a constant supply of giant squid axons, which are used in neuroscience and physiology research.

Positive Impacts: food ; research and education

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
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Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Benefits

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Squid species in general do not threaten humans in any measurable way. They do not actively prey upon humans and are rarely provoked to attack. There are no known adverse economic effects of bigfin reef squid.

Negative Impacts: injures humans (bites or stings)

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
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Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Life Cycle

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Egg capsules incubate for about 3 weeks, depending on temperature. The incubation period can last anywhere between 15 to 22 days. Upon hatching, the paralarvae are planktonic and are about 4.5 to 6.5 mm in mantle length (excluding tentacles), with fully functioning fins and ink sacs. They resemble miniature adults and are already strong swimmers.

Development - Life Cycle: indeterminate growth

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Conservation Status

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Bigfin reef squid are not found on any threatened or endangered species list. Their robust population and wide distribution makes it unlikely that it will become a threatened species in the near future.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
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Stacey Kennedy, Radford University
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Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Reproduction

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When bigfin reef squid enter their reproductive stage, they tend to exhibit the same shoaling behavior exhibited when they first hatched. The group breeds in shoals during the day and separates at night to feed. During mating, male and female bigfin reef squid form close pairs that can last up to several days. Mating can occur in two different ways: "head-to-head” or “male-parallel” mating. In “head-to-head” mating, the male swims upside down and lunges forward towards the female, having already ejected several spermatophores from his funnel onto his hectocotyli, which he attempts to deposit into the female’s mouth funnel. The female will then transfer the spermatophores to her oviduct for fertilization. “Male-parallel” mating involves the male and female swimming side by side. The male then moves below the female, clasps her neck with his arms, and inserts his hectocotylus into the mantle cavity of the female, attaching the spermatophores at the opening of the oviduct.

Elaborate breeding behaviors are known in this species, which largely involve the display of complex body patterns during courtship. One of the body patterns displayed is referred to as the “accentuated gonads” pattern, which occurs when squid reduce their body coloration and enhance the colors of their reproductive organs. This is likely an initial sign of reproductive maturity and receptiveness to other individuals. There are also sex-specific chromatophore displays used by males and females for breeding. Male oval squid use a “Stripe” pattern and a “Flicker” display, while females show a display known as the sexual “Saddle”. The “Stripe” display, similar to the “Zebra” display discussed below, indicates a willingness to mate to females. The male swims parallel to the female displaying this pattern to get her attention. Females respond to the “Stripe” pattern with the sexual “Saddle” pattern, which indicates that the female is mature and ready to breed. It is known as the “Saddle” because the female lightens her entire body except for a central band where the male grasps the female for mating. The male then indicates his intent to pass spermatophores to the female with a “Flicker” chromatophore display.

Prior to mating, male bigfin reef squid can become aggressive and they will battle other males using certain body patterns and postures to determine who will breed with the mature females. Males display color pattern variations to deter or compete against other intruding males. The “Zebra” display is characterized by the presence of irregular dark stripes on the squid’s skin and is agnostic towards other male, warding them off from breeding with paired females. Unpaired males will also sometimes mimic color displays of paired females, to get a paired male to waste spermatophores, leaving the unpaired male open to mate with the paired female. Males guard their mate while she lays her eggs, exhibiting additional agonistic display behaviors to ward off other males. These displays involve both body positioning, such as spreading their arms to look larger, and chromatophore displays. The male usually leaves once the female has spawned.

Mating System: polygynous

Bigfin reef squid can lay eggs year round and the onset of the major spawning season can vary by location. In warmer waters, such as around India, spawning can occur as early as January while in cooler waters near Japan, spawning can begin as late as September. Females can release 20 to 1180 eggs per individual, in egg capsules that contain up to 13 eggs each. These capsules are laid in single straight strands on rocks, corals, plants, submerged branches, and other surfaces along shore lines. Once the female lays her eggs, her body usually deteriorates and she usually dies before she can mate again, but a male can usually mate with several more females before he dies. Sexual maturity in the wild is reached, on average, 171 days after hatching, but this can vary from 161 to 315 days. In captive populations, males reached sexual maturity at approximately 140 days after hatching, while females began spawning when they were 156 to 196 days old. Both males and females mature earlier in captivity than in the wild. Growth to sexual maturity is distinguishable by size and close examination only, as juveniles closely resemble adults.

Breeding interval: Bigfin reef squid can begin mating as early as January to as late as September, depending on the location. Additional spawning can occur throughout the year.

Breeding season: Breeding occurs in the spring.

Range number of offspring: 20 to 1180.

Average number of offspring: 680.

Range gestation period: 15 to 22 days.

Range age at sexual or reproductive maturity (female): 156 to 196 days.

Average age at sexual or reproductive maturity (female): 171 days.

Range age at sexual or reproductive maturity (male): 100 to 140 days.

Key Reproductive Features: semelparous ; seasonal breeding ; year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal ); oviparous ; sperm-storing

Paired males generally remain with the females for protection while the eggs are being laid, but mating partnerships are only temporary. Females coat the eggs in a gelatinous substance, forming an egg capsule that offers protection and nourishment while the egg develops. The eggs are layed in areas where the hatchlings will have cover to hide, in to attempt to avoid predation, but there is still a high risk of mortality. There is no parental care, imprinting, or protection from predators or the environment post-hatching.

Parental Investment: no parental involvement; precocial ; pre-fertilization (Protecting: Male); pre-hatching/birth (Provisioning: Female)

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Filcek, K. and S. Kennedy 2012. "Sepioteuthis lessoniana" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Sepioteuthis_lessoniana.html
author
Kimberly Filcek, Radford University
author
Stacey Kennedy, Radford University
editor
Christine Small, Radford University
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Jeremy Wright, University of Michigan-Ann Arbor
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Brief Summary

provided by EOL authors

The bigfin reef squid or oval squid, Sepioteuthis lessoniana, is a common, small to medium-sized loliginid squid 3.8 - 33 cm (1.5 to 13.0inches) long that is native to temperate and tropical Pacific and Indian Ocean waters.It has recently been documented as spreading to the Mediterranean through the Suez Canal. They live near the shore, in areas with rocks or coral reefs, and eat mostly crustaceans and small fish.Taxonomic studies in the 1990s indicate that S. lessoniana is probably a cryptic species complex made up of several very similar and closely related species.Of the three species in the genus Sepioteuthis, S. lessoniana is has the largest distribution (Wikipedia 2014 and references therein).

As do all species in genus Sepioteuthis, bigfin reef squids have a large ovoid fin circling almost all the way around their mantle.This distinguishes them from other squids, and gives them a cuttlefish-like appearance, reflected in the genus name Sepioteuthis (Sepio is Greek for cuttlefish, teuthis means squid).Bigfin reef squid show social behavior such as schooling and shoaling of multiage individuals and less cannibalism than most other squid species (Boal and Gonzalez 2010; Ikeda et al. 2008). When the juveniles hatch they are miniature adults able to change body color and pattern (metachrosis) as do full adults. Bigfin reef squids have the fastest recorded growth rates of any large marine invertebrate, reaching 600g (1.3lb) in four months. They are a short-lived species, with a maximum recorded lifespan of 315 days (Wikipedia 2014; Boal and Gonzalez 2010; Izeda et al. 2008).

Sepioteuthis lessoniana are one of the most economically important squid species, fished year round in vast quantities for human food especially in Asia. They are regarded as a promising species for mariculture, because of their rapid growth rate, short life span, and tolerance to handling and captivity (Wikipedia 2014 and references therein).These squid have axons hundreds of times larger than other animals, used for very fast muscle contractions to enable their escape mechanisms.These giant axons play a valuable role in medical research in neuroscience (Lee et al. 1994). Sepioteuthis lessoniana are also noted as potential indicators of warming water temperatures, as population sizes respond very rapidly to temperature increase (Wikipedia 2014).

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Bigfin reef squid

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Sepioteuthis lessoniana, commonly known as the bigfin reef squid, glitter squid or oval squid, is a species of loliginid squid. It is one of the three currently recognized species belonging to the genus Sepioteuthis. Studies in 1993, however, have indicated that bigfin reef squids may comprise a cryptic species complex. The species is likely to include several very similar and closely related species.

Bigfin reef squids are characterised by a large oval fin that extends throughout the margins of its mantle, giving them a superficial similarity to cuttlefish. They are small to medium-sized squids, averaging 3.8 to 33 centimetres (1.5 to 13.0 in) in length. They exhibit elaborate mating displays and usually spawn in May, but it can vary by location. The paralarvae resemble miniature adults and are remarkable for already having the capability to change body colouration upon hatching. Bigfin reef squids have the fastest recorded growth rates of any large marine invertebrate, reaching 600 g (1.3 lb) in only four months. They are a short-lived species, with a maximum recorded lifespan of 315 days.

The diet of bigfin reef squids comprises mainly crustaceans and small fish. They are found in the temperate and tropical waters of the Pacific and Indian Oceans, and have recently been introduced into the Mediterranean as a Lessepsian migrant. They are commonly found near the shoreline, near rocks, and coral reefs. They are fished extensively for human consumption in Asia. Because of their rapid growth rate, short life span, and tolerance to handling and captivity, bigfin reef squids are regarded as one of the most promising species for mariculture. They are also a valuable source of giant axons for medical research.

Taxonomy and nomenclature

Bigfin reef squids are also known as northern calamary in Australia and New Zealand, to distinguish them from the southern reef squid (or southern calamary), Sepioteuthis australis.[6][7] Other common names include green-eyed squid in English; koonthal in Malayalam; oosi kanava in Tamil;[8] calmar tonnelet in French; calamar manopla in Spanish;[9] Großflossen-Riffkalmar in German;[10] mu he`e in Hawaiian; 莱氏拟乌贼 in Chinese;[11] torak in Malaysian;[12] アオリイカ (aori-ika) in Japanese;[13] kinn mon in Burmese;[14] and 무늬오징어 (munuiojing-eo), 흰꼴뚜기 (huinkkolttugi), or 미즈이카 (mizuika) in Korean.[15][16]

Sepioteuthis lessoniana is one of the three currently recognised species classified under the genus Sepioteuthis of the pencil squid family, Loliginidae. It belongs to the suborder Myopsina of the squid order Teuthida.[17] Sepioteuthis literally means 'cuttlefish squid', from Greek: σηπία (sēpía, 'cuttlefish') and τευθίς (teuthis, 'squid').[18]

It was first described by the French naturalist André Étienne d'Audebert de Férussac and named after René Primevère Lesson. The type specimen was collected by Lesson off the coast of New Guinea during the circumnavigational voyage of the French corvette La Coquille (1822–1825) under the command of Louis Isidore Duperrey.[19] Numerous other species of Sepioteuthis were described from the Pacific and Indian Oceans in the late 19th century and in the early 20th century. In 1939 the Belgian malacologist William Adam examined the specimens of Sepioteuthis recovered from the tropical western Pacific. He synonymised the twelve species then considered valid under Sepioteuthis lessoniana.[20]

A study in 1993 by Segawa et al. revealed that the population of S. lessoniana in Okinawa may actually be composed of three distinct species.[21] This was confirmed in genetic studies by Izuka et al. in 1994. Triantafillos and Adams in 2005 also showed that S. lessoniana in Shark Bay, Australia is composed of two species.[6] These findings indicate that S. lessoniana may actually comprise several very similar and closely related species. It is now believed that S. lessoniana is a cryptic species complex.[17][20][22]

Description

Like other members of the genus Sepioteuthis, bigfin reef squids are easy to distinguish from other squids in that they possess thick and muscular oval fins that extend around almost the entire mantle.[23] The fins extend about 83 to 97% of the mantle length and are 67 to 70% of the mantle length in width.[24][25] Because of these fins, bigfin reef squids are sometimes mistaken for cuttlefish, a fact reflected by their scientific names. A narrow blue or white line is visible at the point of attachment of the fins to the mantle.[9] A fleshy ridge is also present where the fins meet at the back of the squid.[25]

The mantles of bigfin reef squids are cylindrical, tapering to a blunt cone at the posterior. The mantle is usually 4 to 33 cm (1.6 to 13.0 in) long in males and 3.8 to 25.6 cm (1.5 to 10.1 in) long in females.[22] Both sexes can reach a maximum mantle length of 38 cm (15 in).[2][9] Adult males weigh 403.5 to 1,415 g (0.890 to 3.120 lb), while adult females are 165 to 1,046 g (0.364 to 2.306 lb).[22] Both sexes can attain a maximum documented weight of 1.8 kg (4.0 lb).[2][9] The forward margin of the mantle on the ventral side is concave.[25]

Their eyes are large and covered entirely by a transparent secondary cornea.[26] They are greenish at the base.[27] A pair of prominent ridges (olfactory crests) are present on the ventral surface of the head at the rear edge of the eyes.[24][25] The mouth area is supported by seven triangular flaps (buccal lappets), each with 0 to 7 suckers of less than 0.2 mm in diameter and 18 to 25 teeth. The strong, curved, and short beaks (rostra) are mostly black to dark brown. The radula has seven rows of teeth.[26]

Gladius of S. lessoniana

The spermatophores of males are about 4.5 mm (0.18 in) long and 0.15 mm wide. The ink sac is pear-shaped, with a silvery blue-green outer layer. The vane of the gladius (the rigid internal remnants of the mollusc shell) is oval-shaped and pointed at both ends (lanceolate). It has a broad midrib (rachis).[26][27]

The eight arms are thick, tapering to a narrow point. They are unequal in length, with arm pair I the shortest, followed by arm pair II and arm pair IV, and arm pair III the longest.[27] All of them possess two rows of suckers. Each sucker has a diameter less than 2 mm (0.08 in), decreasing distally, and a ring of 17 to 28 sharp acute teeth. The left arm of pair IV in males is modified into a sexual organ known as the hectocotylus. They bear long fleshy protrusions (papillae) with toothless suckers at the distal portion.[25] The tentacles are thick and long, extending the length of the mantle when retracted. They are slightly compressed laterally.[27] A prominent ridge (a keel) is present on the outer surface of each of the tentacle clubs (the wide tip of the tentacles). There are four rows of suckers on the manus (proximal part of the club) and the dactylus (distal part of the club). The larger suckers in the centre of the manus have 17 to 18 widely spaced teeth.[25]

Coloration

A bigfin reef squid from the Komodo National Park showing vivid iridescence. They are often attracted to divers' lights at night.

Large chromatophores densely cover the upper surfaces of the head, mantle and arms. They are distributed more sparsely on the ventral side.[26][27] The fins do not possess chromatophores on the underside.[24] Living specimens range in colour from translucent creamy white through pale yellow to brownish pink and brownish violet.[24][26][27]

Like some other cephalopods, bigfin reef squids are capable of metachrosis – rapidly changing body colouration and patterns through voluntary control of chromatophores.[28] They also possess iridophores (particularly in the head), a form of structural colouration that produces iridescent metallic greens and red when illuminated.[29] They are also possibly one of two squid species with leucophores. Leucophores are a reflector-type structural colouration that reflects ambient light, such that they are white in white light, green in green light, and so on.[30] Bigfin reef squids are remarkable for having the ability to produce complex body patterns from the moment they hatch. In comparison, other loliginid squid species do not produce complex body patterns at less than four months of age. The patterns produced by bigfin reef squids, however, are less diverse than those of the Caribbean reef squids.[31]

Bigfin reef squids do not possess photophores, and are thus not truly bioluminescent.[26]

Sexual dimorphism

It is often difficult to superficially distinguish between male and female bigfin reef squids. Some authors say that females are generally smaller than males,[22] but this distinction is not observed in other studies.[28] Closer examination of sexually mature specimens, however, will usually distinguish males from females by the presence of the hectocotylus on the fourth left arm in males, and the nidamental glands and the pale ovaries within the mantle in females. Males also purportedly show a more conspicuous pattern of transverse streaks on their dorsal side.[27]

Ecology

Distribution and habitat

A bigfin reef squid among corals in the Red Sea of Egypt

The bigfin reef squid is a neritic warm water-dwelling squid. They are usually found 0 to 100 m (0 to 328 ft) below the water's surface.[25] They tend to remain close to the shoreline, near rocks and reefs.[31][32] They are slightly more active during the night and will move to deeper waters or find cover in daytime. Large numbers of juveniles can often be found hiding beneath floating driftwood.[12]

The bigfin reef squid is the most widespread species in the genus Sepioteuthis. It is found in temperate and tropical regions of the Indian Ocean and the western Pacific Ocean.[2] Their original range extends east to the Hawaiian Islands, west to the Red Sea, north to Japan, and south to Australia and New Zealand (42°N to 42°S and 32°E to 154°W).[2] The range has also expanded to include parts of the Mediterranean Sea. In 2002, bigfin reef squids were first documented in the Gulf of İskenderun of the southeastern Mediterranean Sea. They may have already existed in significant populations in the area prior to their discovery in 2002, as they have acquired a common name among the fishermen of the Aegean Seaσουπιοκαλάμαρο (soupiocalamaro, literally "cuttlefish-like squid"). It is a Lessepsian migrant, reaching the Mediterranean Sea through the Suez Canal.[24]

Diet and predators

The bigfin reef squid eats a variety of different marine organisms. Its main prey are usually prawns and other crustaceans, and fish.[33] Captive specimens were observed to consume one fish every 2 to 25 hours.[28]

Bigfin reef squids are, in turn, preyed upon by tuna, marlin, swordfish, and other predator fish and groundfish.[34][35]

Parasites

Bigfin reef squids serve as hosts to the copepod ectoparasite Doridicola similis and the worm-like dicyemid endoparasites Dicyema koshidai and Dicyema orientale.[36]

Biology and behaviour

A school of bigfin reef squids from the island of Réunion, off the eastern coast of Madagascar
Schooling bigfin reef squids from the Suma Aqualife Park, Kobe, Japan

Bigfin reef squids are closely related to the Caribbean reef squid (Sepioteuthis sepioidea), a species noted for its complex social interactions. Like Caribbean reef squids, bigfin reef squids also exhibit elaborate mating displays.

Bigfin reef squids also exhibit both schooling and shoaling behaviours. Very young bigfin reef squids will also stay close together (shoaling), but do not swim together parallel to each other (schooling). Unlike most other squid species, bigfin reef squids are rarely cannibalistic. Shoals can include animals of different sizes without the threat of larger members attacking and consuming the smaller members. Whether bigfin reef squids recognise each other individually still remains unknown.[28]

Phototaxis

Bigfin reef squids exhibit strong positive phototactic behaviour (attraction to light) and will move readily within a certain distance of a light source. Studies have proposed that this might be an involuntary stimulus behaviour, as the squids immediately stop all other movements once a light source is switched on. The colour of the light does not matter, but it has been shown that they react more strongly to underwater lights between the intensities of 1.5 to 2.5 lx, with peak ranges of 2.5 to 10.0 lx.[37]

Hearing

Bigfin reef squids and the common octopus (Octopus vulgaris) were studied to resolve a century-long debate over whether cephalopods can hear. Unlike fishes, cephalopods do not possess air-filled swim bladders, which might amplify sound waves travelling in water.[38] The results were published in 2009. It showed that bigfin reef squids and octopuses utilise their statocysts for detecting vibrations, an organ primarily used for maintaining spatial orientation. The common octopus can hear sounds between 400 Hz and 1000 Hz. Bigfin reef squids have a slightly better hearing range of 400 Hz to 1500 Hz. Both hear best at a frequency of 600 Hz. Relatively, their hearing is comparable to prawns and some other invertebrates but is less sensitive than that of most fishes.[39]

The difference in the hearing ranges for octopus and bigfin reef squids may be explained by the difference in their habitats. The octopus is demersal (bottom-dwelling) with excellent camouflage capabilities. Bigfin reef squids, on the other hand, are usually in open water with limited hiding places. Hearing would thus be more important for the squids to escape predators. The ability to hear is particularly relevant for avoiding mammalian predators of the suborder Odontoceti (particularly dolphins), who use echolocation to find prey.[38][39]

Mating behaviour

Video of swimming bigfin reef squids

Bigfin reef squids exhibit two most common social body patterning and posturing behaviours related to mating.[28]

The first is dubbed "accentuated gonads", in which they will sometimes increase the visibility of their gonads while reducing the rest of their body colouration. This makes their reproductive organs appear bright white through the transparent mantle. It may indicate the reproductive condition of the signalling squid.[28]

Another common behaviour, primarily seen in males, is dubbed "spread arms", in which the squid will slightly tilt its body forward, head down and arms spread widely and raised above. The mantle is darkened. This behaviour is exhibited mostly when the squids are chasing or following another individual. It is thought to be a signal of reproductive arousal or aggression, similar to the "zebra display" behaviour of Sepioteuthis sepioidea, the "intense zebra display" behaviour of Sepia officinalis, and the "lateral display" of Loligo plei. Females will also sometimes use this display to rebuff courting males.[28]

There are three known courtship behaviours in bigfin reef squids, dubbed "male-upturned" mating, "male-parallel" mating, and "head-to-head" mating.[40] Actual insertion in each position lasts for only a few seconds.[28][40]

"Male-upturned" mating involves rapid back and forth swimming by the courting male beside a slower-swimming female. The male will then flip over so that he is swimming upside down and quickly lunge forward towards the female. He will quickly eject several spermatophores from his funnel into his hectocotylus and attempt to deposit them on the female's mouth funnel, then jet away from the female.[28] "Head-to-head" mating is regarded as a variation of this tactic.[40]

"Male-parallel" mating involves the male and female swimming side by side. The male will then raise one or two of his arm pair I upwards and swing them back and forth. He moves below the female and clasps the female's neck with his arms. In contrast to the previous behaviours, in this position the male actually inserts his hectocotylus into the mantle cavity of the female, attaching the spermatophores right at the opening of the oviduct rather than at the mouth. Possibly for this reason, it is usually more successful in fertilizing the female than other mating behaviours.[40]

In addition to the above, males will often engage in "sneaking" behaviour. In this scenario, a smaller male will attach spermatophores to the female's mouth area while she is being courted by a larger male using the "male-upturned" behaviour. Even when successful, the male using this strategy is usually chased away by the larger male afterwards.[40]

The spermatophores usually remain embedded near the mouth of the female. Mating usually occurs well before spawning, but may also happen on the spawning grounds themselves. In those cases, the male will stay near the female's side as she lays eggs.[28]

Males have been observed to exhibit mating behaviours with other males. Some males have been found with numerous spermatophores embedded in their mouth funnels.[22][28] Since bigfin reef squids distinguish sex by visual cues, this may be a form of deception. The smaller males (termed "female mimics" or "sneaker males") might have assumed body patterning typical of females in order to trick larger males. Believing they are females, they will then waste their spermatophores on them.[41] This behaviour has also been observed in other cephalopods.[22]

Reproduction and life cycle

Egg strands of bigfin reef squids

The main spawning season for bigfin reef squids usually begins in May, but they lay eggs all year round and spawning seasons can vary by location.[33][42][43] A single female can spawn more than once in her lifetime.[22] Females can release 20 to 1180 eggs per individual and will die soon afterwards.[22]

The females spawn by passing eggs from their oviducts. These eggs are then coated in gelatinous substances from the nidamental glands and oviducal glands, forming an egg 'capsule'. The egg capsules of the bigfin reef squids contain two to nine eggs each.[44] These are laid in single straight strands on rocks, corals, aquatic plants, submerged branches and other surfaces.[23][45] At this point, the eggs are 3 mm (0.12 in) in diameter and the egg capsules about 58.2 mm (2.29 in) in length and 12.6 mm (0.50 in) in width, on average.[46]

A small (possibly subadult) bigfin reef squid from East Timor

The capsules incubate for about 3 weeks, depending on temperature. In warmer Indonesia, the incubation period was recorded to be only 15 to 16 days, while in Thailand it takes around 20 to 22 days. They gradually enlarge by absorbing water, reaching around 82.4 mm (3.24 in) in length and 14.6 mm (0.57 in) in width. Unfertilised eggs remain milky white and do not develop further. Fertilised eggs undergo cell division reaching a diameter of 16 mm (0.63 in) with the developing embryo at 11 mm (0.43 in) on the day before hatching. Upon hatching, the paralarvae are 6 mm (0.24 in) in mantle length (excluding tentacles), with fully functioning fins and ink sacs.[46] They resemble miniature adults and are already strong swimmers.[33] They exhibit schooling behaviour two weeks after hatching.[31]

Hatchlings are often cannibalistic. This is regarded as the main cause of death in young squids, particularly in dense populations.[22] However, cannibalism usually happens only when eaten individuals were already weakened significantly or dead, so the actual cause of death may have been something else.[31] Subadults are usually recognisable by their size, ranging from 20 to 60 mm (0.79 to 2.36 in) in length.[33] They reach sexual maturity at less than 210 days in the wild. Males reach sexual maturity earlier than females. In captive populations, males mature 140 days after hatching at most. Females will begin spawning at around 156 to 196 days after hatching. Both males and females mature earlier in captivity than in the wild. Water temperature may play an important role in the earlier sexual maturation of captive specimens. High temperatures may induce shorter lifespans and smaller body sizes, while cooler temperatures favour longer lifespans and larger individuals.[22][47]

Bigfin reef squids have one of the fastest recorded growth rates for any large marine invertebrate. They can reach 600 g (1.3 lb) in only four months.[48] Nonetheless, size can not often be reliably correlated with age, as variations of body size within a generation is fairly common.[22] In captivity, bigfin reef squids have a lifespan of 161 to 315 days for both sexes.[22][28]

Economic importance

Stir-fried bigfin reef squid
Bigfin reef squid caught off Pekan, Malaysia by jigging.

Commercial fishing and human consumption

Bigfin reef squids are one of the most commercially important squid species,[49] and are widely consumed as human food. They are usually caught in large numbers by trawling, seine fishing, or fixed net traps.[25] In small-scale fishing, they are caught by jigging, drive-in nets, slingshot-driven spearguns, or with squid pots.[50]

Fishing operations for bigfin reef squids (particularly in jigging) are usually done at night and utilise bright lights, taking advantage of their attraction to illumination.[37][51][52] They are especially abundant during the full moon and in foggy weather. Populations of bigfin reef squids are not seasonal, and they can be fished throughout the year. They are also used as fish bait in hook and line fishing.[23]

Because of their rapid growth rate, short life span, and tolerance to handling and captivity, bigfin reef squids are regarded as one of the most promising species for mariculture. Although there have been several studies about this, there have been no reported commercial-scale cultures, as of 2011.[42][48]

Biomedical research

The bigfin reef squid is the first squid species to have been cultured for more than one generation. It is remarkable for its ability to readily adapt to being confined in tanks,[31][53] and is one of the few squid species of which the entire life span has been observed under laboratory conditions.[54]

Bigfin reef squids are also valuable sources for squid giant axons used in research in neuroscience and physiology. Unlike axons of other animals, squid axons are very large. Those of bigfin reef squids can range in diameter from 350 to 560 μm (in contrast to the typical 1 μm for humans).[31][55] In life, these giant axons are used by the squids to coordinate escape jetting behaviour, enabling the squid to contract its muscles in a split second directly from the brain.[56]

Global warming

Bigfin reef squids adapt to warmer temperatures by laying more eggs, making them a good indicator species for climate change.[47][57] In conjunction with their rapid growth rates and short lifespans, bigfin reef squid populations may rise dramatically in response to global warming. Overfishing may also play an important role. In the Gulf of Thailand, the fishing industry has been forced to adapt to the large numbers of bigfin reef squids now present in the area, believed to be the result of overfishing of the squid's natural predators. The Australian scientist George Jackson describes them as "the weeds of the sea."[34]

Warmer waters may also accelerate the squid's expansion into areas in which it was not previously native. Its recent discovery as a Lessepsian migrant in the Mediterranean Sea may be an example.[35][58]

See also

Wikimedia Commons has media related to Sepioteuthis lessoniana.
Wikispecies has information related to Sepioteuthis lessoniana.

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Bigfin reef squid: Brief Summary

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Sepioteuthis lessoniana, commonly known as the bigfin reef squid, glitter squid or oval squid, is a species of loliginid squid. It is one of the three currently recognized species belonging to the genus Sepioteuthis. Studies in 1993, however, have indicated that bigfin reef squids may comprise a cryptic species complex. The species is likely to include several very similar and closely related species.

Bigfin reef squids are characterised by a large oval fin that extends throughout the margins of its mantle, giving them a superficial similarity to cuttlefish. They are small to medium-sized squids, averaging 3.8 to 33 centimetres (1.5 to 13.0 in) in length. They exhibit elaborate mating displays and usually spawn in May, but it can vary by location. The paralarvae resemble miniature adults and are remarkable for already having the capability to change body colouration upon hatching. Bigfin reef squids have the fastest recorded growth rates of any large marine invertebrate, reaching 600 g (1.3 lb) in only four months. They are a short-lived species, with a maximum recorded lifespan of 315 days.

The diet of bigfin reef squids comprises mainly crustaceans and small fish. They are found in the temperate and tropical waters of the Pacific and Indian Oceans, and have recently been introduced into the Mediterranean as a Lessepsian migrant. They are commonly found near the shoreline, near rocks, and coral reefs. They are fished extensively for human consumption in Asia. Because of their rapid growth rate, short life span, and tolerance to handling and captivity, bigfin reef squids are regarded as one of the most promising species for mariculture. They are also a valuable source of giant axons for medical research.

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Habitat

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Reference

van der Land, J. (ed). (2008). UNESCO-IOC Register of Marine Organisms (URMO).

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