Pleurobranchus Areolatus
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''Pleurobranchus areolatus'' is a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of pleurobranchid
sea slug 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 (marine gastropod mollusks) that over evolutionary t ...
, a type of
marine Marine is an adjective meaning of or pertaining to the sea or ocean. Marine or marines may refer to: Ocean * Maritime (disambiguation) * Marine art * Marine biology * Marine debris * Marine habitats * Marine life * Marine pollution Military * ...
gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...
mollusc Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000  extant species of molluscs are recognized. The number of fossil species is esti ...
, commonly found in the
Caribbean Sea The Caribbean Sea ( es, Mar Caribe; french: Mer des Caraïbes; ht, Lanmè Karayib; jam, Kiaribiyan Sii; nl, Caraïbische Zee; pap, Laman Karibe) is a sea of the Atlantic Ocean in the tropics of the Western Hemisphere. It is bounded by Mexico ...
. It is up to 15 cm long and it feeds on ascidians.


Taxonomy

Although there were believed to occur six species of ''Pleurobranchus'' in the Caribbean Sea, the other five (''P. atlanticus'' Abbott, 1949, ''P. evelinae'' Thompson, 1977, ''P. crossei'' Vayssière, 1896, ''Susania gardineri'' White, 1952, ''P. reesi'' White, 1952 and ''P. emys'' Ev. Marcus, 1984) were synonymized with ''P. areolatus'' in 2015, based on molecular and morphological evidence.Goodheart J., Camacho-García Y., Padula V., Schrödl M., Cervera J. L., Gosliner T. M. & Valdés Á. (2015). "Systematics and biogeography of ''Pleurobranchus'' Cuvier, 1804, sea slugs (Heterobranchia: Nudipleura: Pleurobranchidae)". ''
Zoological Journal of the Linnean Society The ''Zoological Journal of the Linnean Society'' is a monthly peer-reviewed scientific journal covering zoology published by Oxford University Press on behalf of the Linnean Society. The editor-in-chief is Maarten Christenhusz (Linnean Society). ...
'' 174(2): 322–362. .
The specific names ''P. areolatus'' and ''P. evelinae'' were also commonly in use in literature, while other names were not. There is no known
type material In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes the ...
of ''Pleurobranchus areolatus'', ''Susania gardineri'' and ''P. reesi'', but the anatomy of Mörch's type specimen was illustrated by
Rudolph Bergh Rudolph Bergh (15 October 1824 – 20 July 1909), full name Ludvig Sophus Rudolph Bergh, was a Danish physician and malacologist. He worked in Copenhagen. As a doctor his speciality was sexually transmitted diseases. In Copenhagen a hospital and ...
in 1897. Type material of ''P. atlanticus'' is stored at the National Museum of Natural History (Smithsonian Institution), the Academy of Natural Sciences of Drexel University and the Museum of Comparative Zoology. Those of ''P. crossei'' and ''P. evelinae'' are at the National Museum of Natural History (France) and the
Natural History Museum, London The Natural History Museum in London is a museum that exhibits a vast range of specimens from various segments of natural history. It is one of three major museums on Exhibition Road in South Kensington, the others being the Science Museum an ...
, respectively. The sister species (the closest relative) is ''
Pleurobranchus varians ''Pleurobranchus varians'' is a species of sea slug, a sidegill slug, a marine gastropod mollusc in the family Pleurobranchidae. Taxonomy The sister species (the closest relative) is '' Pleurobranchus areolatus'' from Caribbean.Goodheart J ...
'' from the Central Pacific. Those two species split 3.10 million years ago (the
Isthmus of Panama The Isthmus of Panama ( es, Istmo de Panamá), also historically known as the Isthmus of Darien (), is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North and South America. It contains the country ...
formed 3.1–3.4 Mya). Both species have color morphs and for their proper determination the knowledge of their locality is needed.


Distribution

''Pleurobranchus areolatus'' occurs off Mexico, Costa Rica, Venezuela, Brazil, Jamaica, Puerto Rico, St. Thomas, Aruba, St. Maarten/St Martin, Bahamas, Bermuda, Panama, Canary Islands and Madeira. This was the only species of ''Pleurobranchus'' considered to occur in the
Caribbean Sea The Caribbean Sea ( es, Mar Caribe; french: Mer des Caraïbes; ht, Lanmè Karayib; jam, Kiaribiyan Sii; nl, Caraïbische Zee; pap, Laman Karibe) is a sea of the Atlantic Ocean in the tropics of the Western Hemisphere. It is bounded by Mexico ...
until 2016, when ''
Pleurobranchus iouspi ''Pleurobranchus'' is a genus of sea slugs, specifically side-gill slugs, marine gastropod mollusc in the family Pleurobranchidae. They typically occur in intertidal and shallow subtidal waters of tropical and subtropical seas worldwide. ...
'' was reported from
Santa Marta Santa Marta (), officially Distrito Turístico, Cultural e Histórico de Santa Marta ("Touristic, Cultural and Historic District of Santa Marta"), is a city on the coast of the Caribbean Sea in northern Colombia. It is the capital of Magdalena ...
, Colombia. The type locality is
Saint Thomas, U.S. Virgin Islands Saint Thomas ( da, Sankt Thomas) is one of the Virgin Islands in the Caribbean Sea which, together with Saint John, Water Island, Hassel Island, and Saint Croix, form a county-equivalent and constituent district of the United States Virgin ...
. ''P. areolatus'' was also reported from the eastern Pacific, but those records were identified as ''
Pleurobranchus digueti ''Pleurobranchus'' is a genus of sea slugs, specifically side-gill slugs, marine gastropod mollusc in the family Pleurobranchidae. They typically occur in intertidal and shallow subtidal waters of tropical and subtropical seas worldwide. ...
'' in 2015.


Description

The shape of the body is oval. The dorsum contains numerous small, polygonal and flat tubercles. The background color ranges from light brown to deep violet, with varying degrees of opaque white pigment on the tubercles. In some cases the opaque white pigment is arranged in a symmetrical pattern across the body. It is up to 150 mm long. Rhinophores are rolled and fused at the base, with horizontal striations from base to tip. The color pattern of rhinophores is the same as color pattern of the body.Alvim J. & Pimenta A. D. (2016). "Comparative Morphology and Redescription of ''Pleurobranchus'' Species (Gastropoda, Pleurobranchoidea) from Brazil". '' Zoological Studies'' 55(15): . A well marked pedal gland is visible at the posterior end of the foot in some preserved specimens. The shell is reduced and internal (as in all other ''Pleurobranchus'' species). The shell itself does not allow proper species identification within the genus ''Pleurobranchus''. The shape of the shell is oval. The
protoconch A protoconch (meaning first or earliest or original shell) is an embryonic or larval shell which occurs in some classes of molluscs, e.g., the initial chamber of an ammonite or the larval shell of a gastropod. In older texts it is also called ...
has one
whorl A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs). Whorls in nature File:Photograph and axial plane floral ...
and the size of the protoconch is about 400 μm. The most recent drawing of the reproductive system was published by Goodheart et al. (2015) and by Alvim & Pimenta (2016). The genital cup is just before the gill on the right side of the body. The
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
is triaulic (it has two female openings and one male opening). The shape of the prostate is elongated and the frontal part of the prostate is rounded. The
radula The radula (, ; plural radulae or radulas) is an anatomical structure used by molluscs for feeding, sometimes compared to a tongue. It is a minutely toothed, chitinous ribbon, which is typically used for scraping or cutting food before the food ...
has 73 rows, it has no central tooth and it has 115 lateral teeth on both sides (radular formula 73 × 115.0.115), while the radular formula reported from smaller specimens (27 mm) is 66 × 90.0.90. The gill has the size of 2/3 of the body in living specimens of 27 mm in length. The
circulatory system The blood circulatory system is a system of organs that includes the heart, blood vessels, and blood which is circulated throughout the entire body of a human or other vertebrate. It includes the cardiovascular system, or vascular system, tha ...
was described by Alvim & Pimenta in detail in 2016, but it is the same as in ''
Pleurobranchus reticulatus ''Pleurobranchus'' is a genus of sea slugs, specifically side-gill slugs, marine gastropod mollusc in the family Pleurobranchidae. They typically occur in intertidal and shallow subtidal waters of tropical and subtropical seas worldwide. ...
''. They also described the
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
, which is very similar to that of ''P. reticulatus''.


Ecology

''Pleurobranchus areolatus'' was found to be abundant in Biscayne Bay, Florida in 1946. Minimum recorded depth is 0 m. Maximum recorded depth is 70 m.Welch J. J. (2010). "The “Island Rule” and Deep-Sea Gastropods: Re-Examining the Evidence". '' PLoS ONE'' 5(1): e8776. doi:10.1371/journal.pone.0008776. This species is found under rocks and coral rubble. For example in Florida it is sometimes found among ''
Porites porites ''Porites porites'', commonly known as hump coral or finger coral, is a species of stony coral in the genus ''Porites''. It is found in the Caribbean Sea and western Atlantic Ocean and also along the coast of West Africa. Description ''Porites p ...
'' corals. They are laying large, translucent and gelatinous egg masses during spring in Florida. All species in the genus ''Pleurobranchus'' are carnivorous. ''P. areolatus'' probably feeds on ascidians, for example, ''
Didemnum ''Didemnum'' is a genus of colonial tunicates in the family Didemnidae. It is the most speciose genus in the didemnid family. Species in this genus often have small calcareous spicules embedded in the tunic and form irregular or lobed colonies. S ...
'' sp. Its ectoparasites include copepod '' Anthessius ovalipes''.Walter T. Chad (2015). ''Anthessius ovalipes'' Stock, Humes & Gooding, 1963. In: Walter, T.C. & Boxshall, G. (2015). World of Copepods database. Accessed through: World Register of Marine Species at http://www.marinespecies.org/aphia.php?p=taxdetails&id=348850 on 2016-10-15 Rodriguesic acids and its esters (they are modified
diketopiperazine A diketopiperazine (DKP), also known as a ''dioxopiperazine'' or ''piperazinedione'', is a class of organic compounds related to piperazine but containing two amide linkages. DKP's are the smallest known class of cyclic peptide. Despite their name, ...
s) were isolated from ''P. areolatus'' in 2014.Pereira F. R., Santos M. F., Williams D. E., Andersen R. J., Padula V., Ferreira A. G. & Berlinck R. G. (2014). "Rodriguesic acids, modified diketopiperazines from the gastropod mollusc ''Pleurobranchus areolatus''". ''
Journal of the Brazilian Chemical Society The Journal of the Brazilian Chemical Society (print , eISSN , CODEN JOCSET) is a Brazilian scientific journal in chemistry. It was founded in 1990 and is published by the '' Brazilian Society of Chemistry'' (''Sociedade Brasileira de Química'') ...
'' 25(4): 788–794. .
These are the first diketopiperazine derivatives isolated from a mollusc. Similar diketopiperazines were isolated also from ascidians ''
Didemnum ''Didemnum'' is a genus of colonial tunicates in the family Didemnidae. It is the most speciose genus in the didemnid family. Species in this genus often have small calcareous spicules embedded in the tunic and form irregular or lobed colonies. S ...
'' sp. These derivatives may either result from a ''P. areolatus'' symbiont or may come from feeding on the ascidian. These chemical compounds may be used by ''P. areolatus'' as a chemical defense.


Gallery

Pleurobranchus areolatus map.png, Distribution of ''P. areolatus'' Pleurobranchus areolatus shell.png, Drawing of dorsal view of the internal shell. Pleurobranchus areolatus shell 2.png, Drawing of ventral view of the internal shell including the hyaline sheath. Pleurobranchus areolatus reproductive system.png,
Reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are als ...
of ''Pleurobranchus areolatus'' by R. Tucker Abbott (1949). Pleurobranchus crossei.jpg, Left side view. Pleurobranchus areolatus (Warty Sidegill Slug).jpg, Left side view. Pleurobranchus areolatus 2.png, Drawing of right side view showing the position of the internal shell, the gill, anus behind the gill, and genital pore in front of the gill. Pleurobranchus areolatus 3.png, Drawing ventral view shows rhinophores, anterior sinus, velum, foot, gill and posterior pedal gland.


References

This article incorporates Creative Commons (CC-BY-4.0) text from the referenceGoodheart J. A., Ellingson R. A., Vital X. G., Galvão Filho H. C., McCarthy J. B., Medrano S. M., Bhave V. J., García-Méndez K., Jiménez L. M., López G. & Hoover C. A. (2016). "Identification guide to the heterobranch sea slugs (Mollusca: Gastropoda) from Bocas del Toro, Panama". ''Marine Biodiversity Records'' 9(1), p.56. and public domain text from the reference {{taxonbar, from=Q5048433 Pleurobranchidae Gastropods described in 1863