California sea hare

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California sea hare
Aplysia californica.jpg
Releasing ink after being disturbed
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Mollusca
Class: Gastropoda
Subclass: Heterobranchia
Clade: Euopisthobranchia
Clade: Anaspidea
Superfamily: Aplysioidea
Family: Aplysiidae
Genus: Aplysia
Species:
A. californica
Binomial name
Aplysia californica
California sea hare range.svg
Occurrence is sparse in the northern portion of this range [1]
Synonyms [2]

Aplysia nettiaeWinkler, 1959

The California sea hare (Aplysia californica) is a species of sea slug in the sea hare family, Aplysiidae. [2] It is found in the Pacific Ocean, off the coast of California in the United States and northwestern Mexico.

Contents

Distribution

A. californica is found along the coast of California, United States, and northwestern Mexico (including the Gulf of California). Aplysia species inhabit the photic zone to graze on algae, mainly the intertidal, usually not deeper than 18–20 m (59–66 ft). [3]

Description

The maximum length recorded for the California sea hare is 75 cm (30 in) when crawling, thus fully extended, although most adult specimens are half this size or smaller. Adult animals can weigh up to 7 kg (15 lb). [4] [5] A closely related species, Aplysia vaccaria , the black sea hare, can grow larger still.

A California sea hare is typically reddish-brown to greenish-brown, but the color varies based on the algae it ingests. Each sea hare houses four tentacles, with two on the head sheltering the eyes, and two on the face surrounding the mouth. The body has two folds, called parapodia, which envelop the gills for protection but enable water to get by. Below the body is a muscle that allows mobility, almost like a foot. The California sea hare also has an internal shell to protect its organs. [6]

Lifecycle

Like all sea hares, the California sea hare is hermaphroditic, simultaneously acting as male and female during mating. A. californica is known to form mating chains with up to 20 animals. The eggs are yellow-green, and change after 8–9 days into a brown color before larvae hatch. Mating is most prominent during the summer following water temperature rise to 17°C.

A. californica has a generation time of 19 weeks: Days 1-37 after hatching from the egg comprise the planktonic stage, days 34-37 are the metamorphic stage, and days 45-80 are the juvenile stage. At about day 30, the larvae move on from the planktonic stage and begin to roost on algae, typically red algae. The larvae eat enough to double their weight every 10 days for the following 3 months while they undergo metamorphosis. [7] Reproductive maturity is reached 85 days after hatching (133 days after deposition of the fertilized eggs). The development of the nervous system lasts for 140 days. [8] Often, the California sea hare dies shortly after laying eggs. [9] Cooler temperatures (14-25°C, or 57-77°F) delay spawning and have been shown to extend the lifespan.

Sexual reproduction

At the base of the right anterior tentacle is the aperture from which the penis can protrude. The genital aperture lies at the anterior end of the mantle cavity, a seminal grove arises from it and runs forward to the penis, at the base of the anterior tentacle.

Coupling lasts for hours or sometimes for days, although the actual passage of the sperm may take only a few minutes. Egg laying normally has to be triggered by copulation, but it occurs spontaneously in individuals kept in isolation for up to 3–4 months (typically these eggs are unfertilized). [8] Copulation occurs most frequently in the early morning, and rarely after 12:30 pm. [10] An individual animal weighing 2,600g was recorded to have laid about 500 million eggs at 27 separate times during less than five months.

Feeding habits

Like all Aplysia species, the California sea hare is herbivorous. Its diet consists primarily of red algae such as Laurencia pacifica, Plocamium pacificum, and Ceramium eatonianum, which give the animal its typically reddish or pinkish coloration. Some California sea hares will appear more brown, which helps them blend into their surroundings. [11] A. californica resembles the food on which it grazes, and cannot be distinguished easily from the seaweed unless the animal is moving. [8] When the sea hares eat algae, they consume toxins, which they store in themselves to repel predators. [11] They can also release these toxins as a purple ink, which will also discourage predators from attacking them. [11]

Predators

Close-up showing the rhinophores of Aplysia californica Aplysia californica rhinophore.jpg
Close-up showing the rhinophores of Aplysia californica

Because of the toxins in its body that come from consuming algae, the California sea hare has very few predators. Predators include starfish, lobsters, [12] and the ophistobranch Navanax inermis which will take juveniles. [13]

When it is considerably disturbed, the sea hare is capable of releasing two different kinds of ink from different locations within its mantle cavity, much in the way an octopus does. One ink is reddish-purple and comes from what is called the purple ink gland, while the other is milky white, comes from what is called the opaline gland, and contains the aversive chemical opaline. [14]

Protective mechanisms

Inking provides protection from spiny lobsters, a major predator of sea hares, by means of three mechanisms: [14] chemical deterrence, sensory disruption, and phagomimicry.

The typical defence response of the sea hare to a predator is the release of chemicals such as free amino acids, ink from the ink gland, and opaline from the opaline gland. Chemical deterrence involves the release of toxic chemicals that are noxious to predators and rapidly dissuades them from feeding. Ink creates a dark, diffuse cloud in the water that disrupts the sensory perception of the predator by acting as a screen or decoy. The opaline, which affects the senses dealing with feeding, causes the predator to instinctively attack the cloud of chemicals as if it were indeed food. [14] [15]

Laboratory use

A. californica has become a valuable laboratory animal, used in studies of the neurobiology of learning and memory, and is especially associated with the work of Nobel laureate Eric Kandel. [16]

Its ubiquity in synaptic plasticity studies can be attributed to its simple nervous system, consisting of just 20,000 large, easily identified neurons with cell bodies up to 1 mm in size. [17] Despite its seemingly simple nervous system, though, A. californica is capable of a variety of nonassociative and associative learning tasks, including sensitization, habituation, and classical and operant conditioning. Study typically involves a reduced preparation of the gill and siphon withdrawal reflex. [18]

Sequencing of its whole genome was approved as a priority by National Human Genome Research Institute in March 2005. [19] [20] The draft genome is available on the UCSC Genome browser. [21]

Related Research Articles

<span class="mw-page-title-main">Nudibranch</span> Order of gastropods

Nudibranchs belong to the order Nudibranchia, a group of soft-bodied marine gastropod molluscs that shed their shells after their larval stage. They are noted for their often extraordinary colours and striking forms, and they have been given colourful nicknames to match, such as "clown", "marigold", "splendid", "dancer", "dragon", and "sea rabbit". Currently, about 3,000 valid species of nudibranchs are known.

<span class="mw-page-title-main">Anaspidea</span> Clade of gastropods

The clade Anaspidea, commonly known as sea hares, are medium-sized to very large opisthobranch gastropod molluscs with a soft internal shell made of protein. These are marine gastropod molluscs in the superfamilies Aplysioidea and Akeroidea.

<span class="mw-page-title-main">Sea slug</span> Group of marine gastropods

Sea slug is a common name for some marine invertebrates with varying levels of resemblance to terrestrial slugs. Most creatures known as sea slugs are gastropods, i.e. they are sea snails that, over evolutionary time, have either entirely lost their shells or have seemingly lost their shells due to having a significantly reduced or internal shell. The name "sea slug" is often applied to nudibranchs and a paraphyletic set of other marine gastropods without apparent shells.

The Aplysia gill and siphon withdrawal reflex (GSWR) is an involuntary, defensive reflex of the sea hare Aplysia californica, a large shell-less sea snail or sea slug. This reflex causes the sea hare's delicate siphon and gill to be retracted when the animal is disturbed. Aplysia californica is used in neuroscience research for studies of the cellular basis of behavior including: habituation, dishabituation, and sensitization, because of the simplicity and relatively large size of the underlying neural circuitry.

<span class="mw-page-title-main">Opisthobranchia</span> Informal group of gastropods

Opisthobranchs is a now informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping.

<span class="mw-page-title-main">Aplysiidae</span> Family of gastropods

Aplysiidae is the only family in the superfamily Aplysioidea, within the clade Anaspidea. These animals are commonly called sea hares because, unlike most sea slugs, they are often quite large, and when they are underwater, their rounded body shape and the long rhinophores on their heads mean that their overall shape resembles that of a sitting rabbit or hare. Sea hares are however sea snails with shells reduced to a small plate hidden between the parapodia, and some species are extremely large. The Californian black sea hare, Aplysia vaccaria is arguably the largest living gastropod species, and is certainly the largest living heterobranch gastropod.

<i>Aplysia</i> Genus of sea slugs

Aplysia is a genus of medium-sized to extremely large sea slugs, specifically sea hares, which are a kind of marine gastropod mollusk.

<span class="mw-page-title-main">Rhinophore</span> Anatomy of groups of marine gastropods

A rhinophore is one of a pair of chemosensory club-shaped, rod-shaped or ear-like structures which are the most prominent part of the external head anatomy in sea slugs, marine gastropod opisthobranch mollusks such as the nudibranchs, sea hares (Aplysiomorpha), and sap-sucking sea slugs (Sacoglossa).

<i>Dolabrifera dolabrifera</i> Species of gastropod

Dolabrifera dolabrifera is a species of sea hare, a marine gastropod mollusc in the family Aplysiidae, the sea hares. Dolabrifera dolabrifera, otherwise known as a Warty Seacat. The animal goes by many names, including the common sea hare. The Hawaiian name for Dolabrifera dolabrifera, is Kualakai.

<i>Aplysia fasciata</i> Species of gastropod

Aplysia fasciata, common name the "mottled sea hare", or the "sooty sea hare", is an Atlantic species of sea hare or sea slug, a marine opisthobranch gastropod mollusk in the family Aplysiidae.

<i>Aplysia dactylomela</i> Species of gastropod

Aplysia dactylomela, the spotted sea hare, is a species of large sea slug, a marine opisthobranch gastropod in the family Aplysiidae, the sea hares.

<i>Aplysia punctata</i> Species of gastropod

The spotted sea hare is a species of sea slug in the family Aplysiidae, the sea hares. It reaches a length of up to 20 cm (7.9 in) and is found in the northeast Atlantic, ranging from Greenland and Norway to the Mediterranean Sea.

<i>Aplysia vaccaria</i> Species of gastropod

Aplysia vaccaria, also known as the black sea hare and California black sea hare, is a species of extremely large sea slug, a marine, opisthobranch, gastropod mollusk in the family Aplysiidae. It is the largest sea slug species.

<span class="mw-page-title-main">Chemical defense</span>

Chemical defense is a strategy employed by many organisms to avoid consumption by producing toxic or repellent metabolites or chemical warnings which incite defensive behavioral changes. The production of defensive chemicals occurs in plants, fungi, and bacteria, as well as invertebrate and vertebrate animals. The class of chemicals produced by organisms that are considered defensive may be considered in a strict sense to only apply to those aiding an organism in escaping herbivory or predation. However, the distinction between types of chemical interaction is subjective and defensive chemicals may also be considered to protect against reduced fitness by pests, parasites, and competitors. Repellent rather than toxic metabolites are allomones, a sub category signaling metabolites known as semiochemicals. Many chemicals used for defensive purposes are secondary metabolites derived from primary metabolites which serve a physiological purpose in the organism. Secondary metabolites produced by plants are consumed and sequestered by a variety of arthropods and, in turn, toxins found in some amphibians, snakes, and even birds can be traced back to arthropod prey. There are a variety of special cases for considering mammalian antipredatory adaptations as chemical defenses as well.

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

Phagomimicry is a defensive behaviour of sea hares, in which the animal ejects a mixture of chemicals, which mimic food, and overwhelm the senses of their predator, giving the sea hare a chance to escape. The typical defence response of the sea hare to a predator is to release two chemicals - ink from the ink gland and opaline from the opaline gland. While ink creates a dark, diffuse cloud in the water which disrupts the sensory perception of the predator by acting as a smokescreen and as a decoy, the opaline, which affects the senses dealing with feeding, causes the predator to instinctively attack the cloud of chemicals as if it were indeed food. This ink is able to mimic food by having a high concentration of amino acids and other compounds that are normally found in food, and the attack behaviour of the predator allows the sea-hares the opportunity to escape.

Aplysia morio, the Atlantic black sea hare or sooty sea hare, is a species of sea slug, a marine gastropod mollusk in the family Aplysiidae, the sea hares. It lives in warm waters in the Caribbean Sea and off the south and southeastern coast of the United States, where it feeds on seaweed.

<span class="mw-page-title-main">Opaline gland</span> Defensive gland in sea hares

Sea hares are gastropods without hard shells, using their specialized ink as their main defensive mechanism instead. Their ink has several purposes, most of which have a chemical basis. For one, the ink serves to cloud the predator's vision as well as halt their senses temporarily. In addition, the chemicals in the ink mimic food. Their skin and digestive tract are toxic to predators as well. They are also seen to change their feeding behaviours in response to averse stimuli.

<i>Phestilla minor</i> Species of gastropod

Phestilla minor is a species of sea slug in the Trichechidae family. It is a type of aeolid nudibranch under the Aeolidina suborder. Phestilla minor is a benthic sea slug that is a very small, slow-moving organism found in marine habitats all over the world.

<span class="mw-page-title-main">Aplysioviolin</span> Chemical compound

Aplysioviolin is a purple-colored molecule secreted by sea hares of the genera Aplysia and Dolabella to deter predators. Aplysioviolin is a chemodeterrent, serving to dispel predators on olfactory and gustatory levels as well as by temporarily blinding predators with the molecule's dark color. Aplysioviolin is an important component of secreted ink and is strongly implicated in the sea hares' predatory escape mechanism. While the ink mixture as a whole may produce dangerous hydrogen peroxide and is relatively acidic, the aplysioviolin component alone has not been shown to produce human toxicity.

<i>Aplysia gigantea</i> Species of mollusc in the family Aplysiidae

Aplysia gigantea is a species of sea slug, a shell-less marine gastropod mollusk in the family Aplysiidae. The species was first described in the Journal of the Malacological Society of Australia in 1869. A. gigantea is also known more commonly as the sea hare due to their posterior chemosensory tentacles resembling a hare's ear. A. gigantea is the largest known species in Australia of the opisthobranch genus. The species is known to have toxic effects on terrestrial organisms, particularly domestic dogs. Exposure to this species with dogs has been associated with the development of neurotoxicosis, with symptoms ranging from respiratory distress to tremors, muscle fasciculations, and seizures.

References

  1. Robert Hugh Morris; Eugene Clinton Haderlie (1980). Intertidal Invertebrates of California. Stanford University Press. p. 348. ISBN   978-0-8047-1045-9 . Retrieved 26 July 2013.
  2. 1 2 Rosenberg, G.; Bouchet, P. (2011). Aplysia californica J. G. Cooper, 1863. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=240765 on 2012-03-31
  3. Kandel, Eric. R. (1979). Behavioral Biology of Aplysia: Contribution to the Comparative Study of Opisthobranch Molluscs. San Francisco: W. H. Freeman and Company.
  4. MacFarland, F.M. (1966). Studies of the opisthobranchiate mollusks of the Pacific coast of North America. Mem. Calif. Acad. Sc. VI, p. 1-596
  5. Aplysia californica from the Sea Slug Forum
  6. Dice, Samantha. "Aplysia californica". Animal Diversity Web. Retrieved 2022-10-20.
  7. Pacific, Aquarium of the. "California Brown Sea Hare". www.aquariumofpacific.org. Retrieved 2022-10-20.
  8. 1 2 3 Kandel, Eric Behavioral biology of Aplysia. W.H. Freeman and Co., San Francisco.
  9. Pacific, Aquarium of the. "California Brown Sea Hare". www.aquariumofpacific.org. Retrieved 2022-10-20.
  10. Newby 1972
  11. 1 2 3 "Insided Look: California Sea Hare". Catalina Island Marine Institute. 2021-06-03. Retrieved 2022-10-20.
  12. Watkins et al. Lobster attack induces sensitization in the sea hare, Aplysia californica. J Neurosci. 2010 Aug 18;30(33):11028-31.
  13. Janet L. Leonard and Ken Lukowiak. The Behavior of Aplysia californica Cooper (Gastropoda; Opisthobranchia): I. Ethogram. Behaviour Vol. 98, No. 1/4 (Aug., 1986), pp. 320-360
  14. 1 2 3 Derby, Charles D.; Kicklighter, Cynthia E.; Johnson, P. M. & Xu Zhang (29 March 2007). "Chemical Composition of Inks of Diverse Marine Molluscs Suggests Convergent Chemical Defenses" (PDF). Journal of Chemical Ecology. 2007 (33): 1105–1113. doi:10.1007/s10886-007-9279-0. PMID   17393278. S2CID   92064. Archived from the original (PDF) on 15 November 2009. Retrieved 9 May 2015.
  15. Inman, Mason (29 March 2005). "Sea Hares Lose Their Lunch". Sciencemag.org. Retrieved 10 May 2015.
  16. Edythe McNamee and Jacque Wilson (14 May 2013). "A Nobel Prize with help from sea slugs". CNN. Retrieved 2020-10-31.
  17. Redish, Edward F. (2003). Teaching physics : with the physics suite. Hoboken, NJ: John Wiley & Sons. ISBN   0-471-39378-9. OCLC   53287676.
  18. December 2011, Joseph Castro 30 (30 December 2011). "Sea Slug Offers Clues to Improving Long-Term Memory". livescience.com. Retrieved 2020-10-31.{{cite web}}: CS1 maint: numeric names: authors list (link)
  19. Approved Sequencing Targets Archived 2012-07-27 at the Wayback Machine . Last updated 14 September 2009. Accessed 24 November 2009
  20. National Human Genome Research Institute (1 March 2005) "NHGRI Targets 12 More Organisms for Genome Sequencing". NIH new Releases, Last Updated: 12 June 2009.
  21. aplCal1 at UCSC Genome Browser Gateway

Further reading