Pinna (bivalve)

{{Short description|Genus of bivalves}}

{{Automatic taxobox

| fossil_range = {{fossil range|298|0|Permian–Recent|ref= }}

| image = Pinnidae - Pinna nobilis-001.jpg

| image_caption = Pinna nobilis

| taxon = Pinna

| authority = Linnaeus, 1758

| type_species = Pinna rudis Linnaeus, 1758

| synonyms_ref =

| synonyms = * Exitopinna Iredale, 1939

  • Pinna (Abyssopinna) P. W. Schultz & M. Huber, 2013· accepted, alternate representation
  • Pinna (Cyrtopinna) Mörch, 1853· accepted, alternate representation
  • Pinna (Exitopinna) Iredale, 1939· accepted, alternate representation
  • Pinna (Pinna) Linnaeus, 1758· accepted, alternate representation
  • Pinna (Quantulopinna) Iredale, 1939· accepted, alternate representation
  • Pinna (Subitopinna) Iredale, 1939· accepted, alternate representation
  • Pinnarius Duméril, 1805
  • Pinnula Rafinesque, 1815
  • Quantulopinna Iredale, 1939
  • Subitopinna Iredale, 1939

| display_parents = 3

}}File:Pinna internal and external shell anatomy.jpgPinna is a genus of bivalve molluscs belonging to the family Pinnidae. The type species of the genus is Pinna rudis.

These bivalves are sessile suspension feeders that live in shallow water, fixed to the substrate with a large, silky byssus. There are 32 different species in the genus Pinna, accounting for around 40% of the diversity in the family Pinnidae, and members of the genus are present almost globally. The most extensively studied species in the genus is the critically endangered P. nobilis, a Mediterranean pen shell which was historically important as the principal source of sea silk.{{Cite news |last=Stein |first=Eliot |date=2017 |title=The last surviving sea silk seamstress |url=https://www.bbc.com/travel/article/20170906-the-last-surviving-sea-silk-seamstress |access-date=March 25, 2025 |work=BBC}} Members of Pinna are also valued as sources of food, pearls and for the aesthetic value of their shells.

Description

These pen shells can reach a length of about {{convert|80|-|90|cm|abbr=on}}. They are characterized by thin, elongated, wedge-shaped, and almost triangular shells with long, toothless edges. The surface of the shells shows radial ribs over their entire length.

Pinna is distinguished from its sibling genus Atrina by the presence of a sulcus dividing the nacreous region of the valves, and the positioning of the adductor scar on the dorsal side of shells. Pinna can also be distinguished from another of its relatives Streptopinna by being larger and having a more uniform shell shape.{{Cite web |title=Streptopinna saccata |url=https://seashellsofnsw.org.au/Pinnidae/Pages/Streptopinna_saccata.htm |access-date=2025-03-26 |website=seashellsofnsw.org.au}}

The internal anatomy is consistent with that of a typical mussel, and includes adductor muscles, the mantle and gut, the foot, and the byssal glands.{{Cite journal |last1=Simone |first1=Luiz |last2=Mikkelsen |first2=Paula M |last3=Bieler |first3=Rüdiger |date=May 2015 |title=Comparative Anatomy of Selected Marine Bivalves from the Florida Keys, with Notes on Brazilian Congeners (Mollusca: Bivalvia) |url=https://www.researchgate.net/publication/276268059 |journal=Malacologia |volume=58 |issue=1–2 |pages=1–127 |doi=10.4002/040.058.0201 |via=ResearchGate}}

The main muscles consists of an anterior and a posterior adductor muscle, a posterior retractor muscle, and dorsal and ventral pallial retractor muscles. With the exception of the ventral pallial retractor muscle, everything else is located on the dorsal valve. The anterior adductor muscle is very small, and it is located close to the anterior point, while the posterior adductor muscle is much larger, and is located about a third of the length along the shell, near the hinge plate. The latter is attached to the valve alongside the posterior pedal retractor muscle. The foot has two parts: a more slender anterior part that projects out and a posterior part that wraps around the byssus. The byssus is composed of fibrous, iridescent, brown threads and surrounds the pair of byssal glands. The mantle is mostly translucent, except near the posterior edge, which is spotted.

Distribution

Species in the genus Pinna are geographically widespread and is known to occur in tropical and subtropical seas around the world.{{Cite journal |last1=Lemer |first1=Sara |last2=Buge |first2=Barbara |last3=Bemis |first3=Amanda |last4=Giribet |first4=Gonzalo |date=2014 |title=First molecular phylogeny of the circumtropical bivalve family Pinnidae (Mollusca, Bivalvia): Evidence for high levels of cryptic species diversity". Molecular Phylogenetics and Evolution. |url=https://www.sciencedirect.com/science/article/abs/pii/S1055790314000591 |journal=Molecular Phylogenetics and Evolution |volume=75 |pages=11–23 |doi=10.1016/j.ympev.2014.02.008 |pmid=24569016 |via=Elsevier Science Direct|url-access=subscription }}

Pinna species are widespread in the Indo-Pacific, ranging as far south as South Africa and New Zealand and as far north as the Persian Gulf and Japan.{{Cite book |last=Strack |first=Elisabeth |title=Perlen |publisher=Rühle-Diebener-Verlag |year=2006 |isbn=9783981084801 |location=Stuttgart, Germany |language=German |trans-title=Pearls}} On the west coast of North America, Pinna members are known from North Carolina to Argentina, and are abundant in the Caribbean and Gulf of Mexico. This genus is also historically very well known from the Mediterranean and Red Seas.{{Cite journal |last1=Davenport |first1=John |last2=Ezgeta-Balić |first2=Daria |last3=Peharda |first3=Melita |last4=Skejić |first4=Sanda |last5=Ninčević-Gladan |first5=Živana |last6=Matijević |first6=Slavica |date=2011 |title=Size-differential feeding in Pinna nobilis L. (Mollusca: Bivalvia): Exploitation of detritus, phytoplankton and zooplankton |url=https://www.sciencedirect.com/science/article/abs/pii/S0272771410004579 |journal=Estuarine, Coastal and Shelf Science |volume=92 |issue=2 |pages=246–254 |doi=10.1016/j.ecss.2010.12.033 |bibcode=2011ECSS...92..246D |via=Elsevier Science Direct|url-access=subscription }}

Fossil members of this genus have been found globally, date back to the late Permian period{{Cite journal |last1=Shilekhin |first1=Lev |last2=Mazaev |first2=Alexey |last3=Biakov |first3=Alexander |date=2023 |title=The Earliest Representatives of the Genus Pinna (Bivalvia), from the Early Permian Reef of Shakhtau (Southern Cisuralia, Russia)". Paleontological Journal. 57: 375–379 – via SpringerNature. |url=https://www.researchgate.net/publication/373410808 |journal=Paleontological Journal |volume=57 |pages=375–379 |via=ResearchGate}} and are especially well-represented and widespread in Jurassic and Cretaceous fossils.

Ecology

= Life habit =

Pinna species live in coastal and marine waters, as well as in transitional inlets partially influenced by freshwater flow. Member of this genus are known to inhabit soft, muddy substrates, sandy seagrass meadows, coral reefs, and fields of coral rubble in relatively shallow water.{{Cite journal |last1=Hanafi Idris |first1=Mohd |last2=Bin Arshad |first2=Aziz |last3=Sidik Bujang |first3=Japar |last4=Abd. Ghaffar |first4=Mazlan |last5=Khalijah Daud |first5=Siti |date=2009 |title=Morphological Characteristics of Pinna bicolor Gmelin and Pinna deltodes Menke from the Seagrass Bed of Sungai Pulai, Johor, Peninsular Malaysia |url=https://www.researchgate.net/publication/289185813 |journal=Sains Malaysiana |volume=38 |issue=3 |pages=333–339 |via=ResearchGate}}

All members of the genus Pinna are sessile, and orient themselves vertically relative to the substrate with the thin, tapered end pointing downwards and the wide end open upwards.{{Cite journal |last=Aucoin |first=Serge |date=2010 |title=A first report on the shrimp Pontonia sp. and other potential symbionts in the mantle cavity of the penshell Pinna carnea in the Dominican Republic |url=https://www.researchgate.net/publication/225983991 |journal=Symbiosis |volume=50 |issue=3 |pages=130–131 |doi=10.1007/s13199-010-0050-x |bibcode=2010Symbi..50..135A |via=ResearchGate}} In muddy or soft, sandy conditions, Pinna will bury 50-95% of its body in substrate and will use tough byssal threads to keep them fixed in the soft substrate around them with the aperture elevated from, or flush with, the surface of the sediment.{{Cite journal |last1=Kiyotaka |first1=Chinzei |last2=Savazzi |first2=Enrico |last3=Seilacher |first3=Adolf |author-link3=Adolf Seilacher |date=1982 |title=Adaptational strategies of bivalves living as infaunal secondary soft bottom dwellers |journal=Neues Jahrbuch für Geologie und Paläontologie |volume=164 |issue=1–2 |pages=229–244 |doi=10.1127/njgpa/164/1982/229 |bibcode=1982NJGPA.164..229C |via=ResearchGate}} On more solid, rockier substrates, some Pinna can anchor themselves directly to hard substrates similarly to other common byssate molluscs.

Members of the genus Pinna often arrange themselve in clusters or loosely spaced colonies, and can provide an important hard substrate for boring and encrusting organisms in otherwise inhospitable sandy or muddy settings, and may promote ecological diversity in environments they inhabit.{{Cite journal |last1=Tiller |first1=Georgia |last2=Martin |first2=Bradley |last3=Baring |first3=Ryan |date=2024 |title=Razor clam (Pinna bicolor) structural mimics as drivers of epibenthic biodiversity; a manipulative experiment |url=https://www.sciencedirect.com/science/article/pii/S0141113624003192 |journal=Marine Environmental Research |volume=200 |doi=10.1016/j.marenvres.2024.106658 |pmid=39088890 |bibcode=2024MarER.20006658T |via=Elsevier Science Direct|doi-access=free }} Pinna are often well known hosts of Pontonia shrimp and pea crabs, as well as of cardinalfish, amphipods, isopods and sea anemones, which can live inside of their mantle cavities.{{Cite journal |last1=Chávez-Villalba |first1=Jorge |last2=de Jesús Reynaga-Franco |first2=Felipe |last3=Hoyos-Chairez |first3=Francisco |date=2022 |title=Worldwide overview of reproduction, juvenile collection, spat production and cultivation of pen shells |journal=Reviews in Aquaculture |volume=14 |issue=3 |pages=1371–1388|doi=10.1111/raq.12654 |bibcode=2022RvAq...14.1371C }}

= Feeding =

Members of the genus Pinna are generally filter feeders, however, some soft-bottom taxa, especially ones that are deeply buried in sediment, are likely deposit feeders. Pinna-genus bivalves consume a wide variety of prey, and includes zooplankton, phytoplankton, and organic detritus.{{Cite journal |last=Vicente |first=Nardo |date=2014 |title=Utilization of muddy detritus as organic matter source by the fan mussel Pinna nobilis |url=https://www.academia.edu/116584605 |journal=Mediterranean Marine Science |volume=15 |issue=3 |page=667 |doi=10.12681/mms.836 |bibcode=2014MedMS..15..667T |via=Academia|doi-access=free }}

In Pinna nobilis, the size of the shell, and, as a result, the proportion of the shell that is above the surface of the substrate, are directly related with where nutrients were sourced. Larger P. nobilis showed a preference for prey higher in the water column, such as Calanoid copepods and diatoms, while smaller P. nobilis preferred more benthic prey, like Harpacticoid copepods and consumed a higher proportion of organic detritus, which is denser on the bottom.

= Reproduction and development =

Pinna bivalves are iteroparous broadcast spawners, and release male and female gametes into the water column separately to allow for external fertilization. Reproduction across the genus Pinna beyond this is diverse, some members of the genus, such as Pinna nobilis, P. rugosa, and P. bicolor have defined spawning periods, which generally occur during the summer, while others, like P. carnea that live in more tropical climates spawn year-round.

Fertilized gametes form trochophores, then veliger larvae, which are planktonic can drift for many days in the water column before settling onto substrate.{{Cite journal |last1=Trigos |first1=Sergio |last2=Vicente |first2=Nardo |last3=Prado |first3=Patricia |last4=Espinós |first4=Francisco |date=2018 |title=Adult spawning and early larval development of the endangered bivalve Pinna nobilis |url=https://www.sciencedirect.com/science/article/abs/pii/S0044848617314515 |journal=Aquaculture |volume=483 |pages=102–110 |doi=10.1016/j.aquaculture.2017.10.015 |bibcode=2018Aquac.483..102T |via=Elsevier Science Direct|hdl=10251/103987 |hdl-access=free }} The veliger of the genus Pinna are stubby, triangular in outline, transparent and already contains the heteromyarian, or anisomyarian musculature seen in adult Pinna, where one adductor muscle is much reduced in size compared to the other.{{Cite journal |last=Allen |first=John |date=2011 |title=On the functional morphology of Pinna and Atrina larvae (Bivalvia: Pinnidae) from the Atlantic |journal=Journal of the Marine Biological Association of the United Kingdom |volume=91 |issue=4 |pages=823–829 |doi=10.1017/S0025315410001694 |bibcode=2011JMBUK..91..823A |via=Cambridge University Press}}

Upon reaching an appropriate substrate, the larva develops quickly, rapidly becoming sessile, and adult characteristics such as the ctenidium, mantle, and shell ornament take shape. Interestingly, all shell growth only occurs along the dorsal (pointed), ventral (wedge), and posterior margins, while no growth occurs anteriorly. The veliger shell quickly erodes, and is uncommon to see in adult forms.

Human uses

= Sea silk =

{{Main|Sea silk}}

= Pearls =

File:Pinna 03.jpg covering with sulcus]]

Members of the genus Pinna are known to produce pearls. Due to the partial coverage of the insides of their shells with nacre, they are capable of producing both nacreous and non-nacreous pearls.{{Cite journal |last=Karampelas |first=Stefanos |date=2009 |title=Characterization of Some Pearls of the Pinnidae Family |url=https://www.gia.edu/gems-gemology/fall-2009-gem-news-international |journal=Gems & Gemology |volume=45 |issue=3 |pages=221–223 |via=Gemological Institute of America}} Uniquely, members of Pinna appear to be unique in their capacity to produce gem-quality pearls that are made of calcite and contain carotenoids which can lead to colouration that ranges from off-white, to vivid orange, brown and black. Pinna pearls may also be elongate and teardrop-shaped, mirroring the shape of the host shell. The nacreous shell itself is also valuable as a collector's item.{{Cite journal |last1=Katsanevakis |first1=Stelios |last2=Poursanidis |first2=Dimitris |last3=Issaris |first3=Yiannis |last4=Panou |first4=Aliki |date=2011 |title="Protected" marine shelled molluscs: Thriving in Greek seafood restaurants |url=https://www.researchgate.net/publication/230729383 |journal=Mediterranean Marine Science |volume=12 |issue=2 |pages=429–438 |doi=10.12681/mms.42 |bibcode=2011MedMS..12..429K |via=ResearchGate|doi-access=free }}

= Food =

Members of the genus Pinna, as well as its sibling genus Atrina, are prized food sources around the Indo-pacific and Mediterranean.{{Cite journal |last1=Beer |first1=Andrew |last2=Southgate |first2=Paul |date=2006 |title=Spat collection, growth and meat yield of Pinna bicolor (Gmelin) in suspended culture in northern Australia |url=https://www.sciencedirect.com/science/article/abs/pii/S0044848606002833 |journal=Aquaculture |volume=258 |issue=1 |pages=424–429 |doi=10.1016/j.aquaculture.2006.04.014 |bibcode=2006Aquac.258..424B |via=Elsevier Science Direct|url-access=subscription }} Members of the genus are both harvested in the wild and are cultured, within the genus Pinna, most aquaculture efforts are concentrated on Pinna rugosa in Mexico and Pinna nobilis, the noble pen shell, in the Mediterranean. Aquaculture from the harvesting and growing of spats has seen success in some members of Pinna, but is not performed commercially, and the majority of Pinna catch remains harvested wild. P. nobilis remains a delicacy in parts of the Mediterranean, and is still served in restaurants, despite its critically endangered status and the fact it is illegal to harvest.

Conservation

= Human efforts =

As of 2019, Pinna nobilis is classified as a critically endangered species by the IUCN. Mass mortality events have caused nearly all Mediterranean populations to die out.{{Cite journal |last1=Nebot-Colomer |first1=Elisabet |last2=Hernandis |first2=Sebastián |last3=Mourre |first3=Baptiste |last4=Fraile-Nuez |first4=Eugenio |last5=Álvarez |first5=Elvira |last6=Deudero |first6=Salud |last7=Albentosa |first7=Marina |last8=Vázquez-Luis |first8=Maite |date=June 2024 |title=No recruits for an ageing population: First signs of probable population extinction in one of the last reservoirs of the Critically Endangered species Pinna nobilis |url=https://www.sciencedirect.com/science/article/pii/S1617138124000499#b0170 |journal=Journal for Nature Conservation |volume=79 |doi=10.1016/j.jnc.2024.126600 |bibcode=2024JNatC..7926600N |via=Elsevier Science Direct|hdl=10261/359843 |hdl-access=free }} Since then, conservation efforts such as transporting individuals to safer habitats, population monitoring, and captive breeding efforts have been made to preserve the species.{{Cite journal |last1=Paola Ferranti |first1=Maria |last2=Azzena |first2=Ilenia |last3=Batistini |first3=Edoardo |last4=Caracciolo |first4=Daniela |last5=Casu |first5=Marco |last6=Chiantore |first6=Mariachiara |last7=Ciriaco |first7=Saul |last8=Firpo |first8=Valerio |last9=Intini |first9=Luca |last10=Locci |first10=Chiara |last11=Montefalcone |first11=Monica |last12=Oprandi |first12=Alice |last13=Sanna |first13=Daria |last14=Scarpa |first14=Fabio |last15=Segarchi |first15=Marco |date=December 15, 2024 |title=Handling of the Bivalve Pinna nobilis, Endangered and Pathogen-Affected Species, for Controlled Reproduction: Precautions Taken |journal=Ecology and Evolution |volume=14 |issue=12 |pages=e70565 |doi=10.1002/ece3.70565 |pmid=39687581 |pmc=11646934 |bibcode=2024EcoEv..1470565F }} In December 2022, the European Life Pinna Project set out to test their conservation protocols using the related Atrina genus as an experimental population because Atrina was not critically endangered or a protected species.

= Parasites =

Parasites such as the protozoan Haplosporidium pinnae, Myobacterium species and more, have been theorized as a potential cause of mortality for Pinna nobilis. In 2019, an event termed the "cold drop" resulted in high volumes of nutrients, sediments, and fresh water to flood Pinna habitats and allow the introduction of H. pinnae. The presence of parasites, along with mass mortality events, only contributed to the decline of P. nobilis populations following 2019.

Species

According to the World Register of Marine Species, extant species in the genus Pinna are:

{{div col|colwidth=20em}}

{{div col end}}

The following species are only known from the fossil record:

{{div col|colwidth=20em}}

{{div col end}}

Nomen nudum

  • Pinna inflata Röding, 1798
  • Pinna lubrica Lightfoot, 1786
  • Pinna nebulosa Lightfoot, 1786
  • Pinna nigricans Lightfoot, 1786
  • Pinna striata Röding, 1798
  • Pinna tenera Lightfoot, 1786
  • Pinna violacea Röding, 1798

Nomen dubium

  • Pinna atrata Clessin, 1891
  • Pinna bullata Gmelin, 1791
  • Pinna marginata Lamarck, 1819
  • Pinna minax Hanley, 1858
  • Pinna rollei Clessin, 1891
  • Pinna rostellum Hanley, 1858
  • Pinna rotundata Linnaeus, 1758
  • Pinna sanguinea Gmelin, 1791
  • Pinna virgata Menke, 1843

Synonyms

{{clear}}

Gallery

File:Naturalis Biodiversity Center - RMNH.MOL.318944 - Pinna angustana Lamarck, 1819 - Pinnidae - Mollusc shell.jpeg|Pinna angustana

File:Naturalis Biodiversity Center - ZMA.MOLL.414274 - Pinna atropurpurea Sowerby , 1825 - Pinnidae - Mollusc shell.jpeg|Pinna atropurpurea

File:Naturalis Biodiversity Center - RMNH.MOL.318930 - Pinna attenuata Reeve, 1858 - Pinnidae - Mollusc shell.jpeg|Pinna attenuata

File:Pinna bicolor Razor clam P1182475.JPG|Pinna bicolor

File:Pinna carnea no.1.jpg|Pinna carnea

File:Naturalis Biodiversity Center - RMNH.MOL.318959 - Pinna deltodes Menke, 1843 - Pinnidae - Mollusc shell.jpeg|Pinna deltodes

File:Naturalis Biodiversity Center - RMNH.MOL.318949 - Pinna electrina Reeve, 1858 - Pinnidae - Mollusc shell.jpeg|Pinna electrina

File:Naturalis Biodiversity Center - ZMA.MOLL.414491 - Pinna incurva Gmelin, 1791 - Pinnidae - Mollusc shell.jpeg|Pinna incurva

File:Naturalis Biodiversity Center - RMNH.MOL.318885 - Pinna muricata Linnaeus, 1758 - Pinnidae - Mollusc shell.jpeg|Pinna muricata

File:Pinnidae - Pinna rudis.JPG|Pinna rudis

File:Naturalis Biodiversity Center - ZMA.MOLL.414295 - Pinna rugosa Sowerby, 1835 - Pinnidae - Mollusc shell.jpeg|Pinna rugosa

File:Naturalis Biodiversity Center - RMNH.MOL.318953 - Pinna trigonium Dunker, 1852 - Pinnidae - Mollusc shell.jpeg|Pinna trigonium

File:Pinna noblis shell & byssus.JPG|Pinna nobilis shell with byssus

References

{{Reflist|refs=

{{GBIF |id=2285348 |taxon=Pinna Linnaeus, 1758 |access-date=17 September 2024}}

{{cite web |url=https://paleobiodb.org/classic/basicTaxonInfo?taxon_no=16400 |title=Pinna Linnaeus, 1758 |work=Paleobiology Database |access-date=17 September 2024}}

{{cite journal|last1=Shilekhin |first1=L. E. |last2=Mazaev |first2=A. V. |last3=Biakov |first3=A. S. |year=2023 |title=The most ancient representatives of the genus Pinna (Bivalvia) in the Early Permian reef of Shakhtau (southern Cis-Urals, Russia) |journal=Paleontological Journal |volume=57 |issue=4 |pages=21–25 |doi=10.1134/S0031030123040111 |url=https://www.elibrary.ru/item.asp?id=54096158 |url-access=subscription }}

{{cite WoRMS |title=Pinna Linnaeus, 1758 |year=2024 |id=138352 |db=MolluscaBase |access-date=17 September 2024}}

}}

Further reading

  • Frank H.T. Rodes, Herbert S. Zim en Paul R. Shaffer (1993) - Natuurgids Fossielen (het ontstaan, prepareren en rangschikken van fossielen), Zuidnederlandse Uitgeverij N.V., Aartselaar. ISBN D-1993-0001-361
  • Cyril Walker & David Ward (1993) - Fossielen: Sesam Natuur Handboeken, Bosch & Keuning, Baarn. {{ISBN|90-246-4924-2}}
  • {{Cite journal|

last=Packard |

first=Earl |author2=Jones, David L.|

title=Cretaceous Pelecypods of the Genus Pinna from the West Coast of North America |

journal=Journal of Paleontology |

volume=39 |

issue=1 |

pages=910–915 |

date=Sep 1965 }}

  • {{Cite web |

url=http://www.manandmollusc.net/glossary.html |

title=Glossary |

publisher=Man and Mollusc |

access-date=2008-01-30 }}

  • Coan, E. V.; Valentich-Scott, P. (2012). Bivalve seashells of tropical West America. Marine bivalve mollusks from Baja California to northern Peru. 2 vols, 1258 pp.
  • Schultz, P. W.; Huber, M. (2013). Revision of the worldwide Recent Pinnidae and some remarks of fossil European Pinnidae. Acta Conchyliorum. 13: 1–164.