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''Spondylus'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of bivalve molluscs, the only genus in the
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
Spondylidae.MolluscaBase (2019). MolluscaBase. Spondylus Linnaeus, 1758. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=taxdetails&id=138518 on 2019-03-04 They are known in English as spiny oysters or thorny oysters (though they are not, in fact,
true oysters The Ostreidae, the true oysters, include most species of molluscs commonly consumed as oysters. Pearl oysters are not true oysters, and belong to the order Pteriida. Like scallops, true oysters have a central adductor muscle, which means the ...
).


Description

The many species of ''Spondylus'' vary considerably in appearance. They are grouped in the same superfamily as the scallops. They are not closely related to true oysters (family
Ostreidae The Ostreidae, the true oysters, include most species of molluscs commonly consumed as oysters. Pearl oysters are not true oysters, and belong to the order Pteriida. Like scallops, true oysters have a central adductor muscle, which means the ...
); however, they do share some habits such as cementing themselves to rocks rather than attaching themselves by a
byssus A byssus () is a bundle of filaments secreted by many species of bivalve mollusc that function to attach the mollusc to a solid surface. Species from several families of clams have a byssus, including pen shells ( Pinnidae), true mussels (Mytil ...
. The two halves of their shells are joined with a ball-and-socket type of hinge, rather than with a toothed hinge as is more common in other bivalves. They also still retain vestigial anterior and posterior ''auricles'' ("ears", triangular shell flaps) along the hinge line, a characteristic feature of scallops, though not of oysters. As is the case in all scallops, ''Spondylus'' spp. have multiple eyes around the edges of their mantle, and they have relatively well-developed
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 ...
s. Their nervous ganglia are concentrated in the
viscera In biology, an organ is a collection of tissues joined in a structural unit to serve a common function. In the hierarchy of life, an organ lies between tissue and an organ system. Tissues are formed from same type cells to act together in a f ...
l region, with recognisable optic lobes connected to the eyes.


Evolutionary history

The genus ''Spondylus'' appeared in the Mesozoic era, and is known in the fossil records from the
Triassic The Triassic ( ) is a geologic period and system (stratigraphy), system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Year#Abbreviations yr and ya, Mya), to the beginning of the Jurassic Period 201.36 ...
Cassian beds in Italy (235 to 232 million years ago) onwards. About 40 extinct species are known. Fossils of these molluscs can be found in fossiliferous marine strata all over the world. For example, they are present in Cretaceaous rocks in the
Fort Worth Formation The Fort Worth Formation is a geologic formation in Texas. It preserves fossils dating back to the Cretaceous period. See also * List of fossiliferous stratigraphic units in Texas * Paleontology in Texas Paleontology in Texas refers to paleo ...
of
Texas Texas (, ; Spanish: ''Texas'', ''Tejas'') is a state in the South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2020, it is the second-largest U.S. state by ...
, and in the
Trent River Formation Trent may refer to: Places Italy * Trento in northern Italy, site of the Council of Trent United Kingdom * Trent, Dorset, England, United Kingdom Germany * Trent, Germany, a municipality on the island of Rügen United States * Trent, California, ...
of
Vancouver Island Vancouver Island is an island in the northeastern Pacific Ocean and part of the Canadian province of British Columbia. The island is in length, in width at its widest point, and in total area, while are of land. The island is the largest by ...
, as well as in other parts of North America.Finsley, Chalres. 1999. A Field Guide to the Fossils of Texas. Gulf Publishing. Lanham, Maryland. plate 55.Ludvigsen, Rolf & Beard, Graham. 1997. West Coast Fossils: A Guide to the Ancient Life of Vancouver Island. pg. 104


Distribution

Spiny oysters are found in all subtropical and (especially) tropical seas, usually close to the coasts.


Ecology

Spondylus are filter feeders. The adults live cemented to hard substrates, a characteristic they share, by convergent evolution, with
true oysters The Ostreidae, the true oysters, include most species of molluscs commonly consumed as oysters. Pearl oysters are not true oysters, and belong to the order Pteriida. Like scallops, true oysters have a central adductor muscle, which means the ...
and jewel boxes. Like the latter, they are protected by spines and a layer of
epibiont An epibiont (from the Ancient Greek meaning "living on top of") is an organism that lives on the surface of another living organism, called the basibiont ("living underneath"). The interaction between the two organisms is called epibiosis. An ep ...
s and, like the former, they can produce pearls. The type of substrate they use depends on the species: many only attach to coral, and the largest diversity of species is found in tropical coral reefs; others, (particularly S. spinosus) however, easily adapt to man-made structures, and have become important invasive species. Others still are often found attached to other shells, perhaps the most common belonging to the genus
Malleus The malleus, or hammer, is a hammer-shaped small bone or ossicle of the middle ear. It connects with the incus, and is attached to the inner surface of the eardrum. The word is Latin for 'hammer' or 'mallet'. It transmits the sound vibrations f ...
.


Uses

Archaeological evidence indicates that people in
Neolithic The Neolithic period, or New Stone Age, is an Old World archaeological period and the final division of the Stone Age. It saw the Neolithic Revolution, a wide-ranging set of developments that appear to have arisen independently in several p ...
Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a subcontinent of Eurasia and it is located entirel ...
were trading the shells of ''S. gaederopus'' to make bangles and other ornaments throughout much of the Neolithic period. The main use period appears to have been from around 5350 to 4200 BC. The shells were harvested from the
Aegean Sea The Aegean Sea ; tr, Ege Denizi (Greek: Αιγαίο Πέλαγος: "Egéo Pélagos", Turkish: "Ege Denizi" or "Adalar Denizi") is an elongated embayment of the Mediterranean Sea between Europe and Asia. It is located between the Balkans ...
, but were transported far into the center of the continent. In the LBK and
Lengyel culture __NOTOC__ The Lengyel culture is an archaeological culture of the European Neolithic, centered on the Middle Danube in Central Europe. It flourished from 5000 to 4000 BC, ending with phase IV, e.g., in Bohemia represented by the ' Jordanow/Jorda ...
s, ''Spondylus'' shells from the Aegean Sea were worked into bracelets and belt buckles. Over time styles changed with the middle neolithic favouring generally larger barrel-shaped beads and the late neolithic smaller flatter and disk shaped beads. Significant finds of jewelry made from Spondylus shells were made at the
Varna Necropolis The Varna Necropolis ( bg, Варненски некропол), or Varna Cemetery, is a burial site in the western industrial zone of Varna (approximately half a kilometre from Lake Varna and 4 km from the city centre), internationally c ...
. During the late Neolithic the use of Spondylus in grave goods appears to have been limited to women and children. ''S. crassisquama'' is found off the coast of Colombia and
Ecuador Ecuador ( ; ; Quechua: ''Ikwayur''; Shuar: ''Ecuador'' or ''Ekuatur''), officially the Republic of Ecuador ( es, República del Ecuador, which literally translates as "Republic of the Equator"; Quechua: ''Ikwadur Ripuwlika''; Shuar: ' ...
and has been important to
Andean The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S l ...
peoples since
pre-Columbian In the history of the Americas, the pre-Columbian era spans from the original settlement of North and South America in the Upper Paleolithic period through European colonization, which began with Christopher Columbus's voyage of 1492. Usually, ...
times, serving as both an offering to the
Pachamama Pachamama is a goddess revered by the indigenous peoples of the Andes. In Inca mythology she is an "Earth Mother" type goddess, Dransart, Penny. (1992) "Pachamama: The Inka Earth Mother of the Long Sweeping Garment." ''Dress and Gender: Making ...
and as currency. In fact, much like in Europe, the ''Spondylus'' shells also reached far and wide, as pre-Hispanic Ecuadorian peoples traded them with peoples as far north as present-day
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
and as far south as the central
Andes The Andes, Andes Mountains or Andean Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
. The Moche people of ancient
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = National seal , national_motto = "Firm and Happy f ...
regarded the sea and animals as sacred; they used ''Spondylus'' shells in their art and depicted ''Spondylus'' in effigy pots. Spondylus were also harvested from the
Gulf of California The Gulf of California ( es, Golfo de California), also known as the Sea of Cortés (''Mar de Cortés'') or Sea of Cortez, or less commonly as the Vermilion Sea (''Mar Bermejo''), is a marginal sea of the Pacific Ocean that separates the Baja C ...
and traded to tribes through Mexico and the American Southwest. Spondylus shells were the driving factor of trade within the Central Andes and were used in a similar manner to gold nuggets, copper hatches, coca, salt, red pepper, and cotton cloth. The use of Spondylus shells is what led to an economy of sorts in the Central Andes and led to the development of a merchant class, "mercardes", in different cultures within the Central Andes. This caused the development of different styles of trade that went through evolutionary changes throughout pre-Columbian times. These are reciprocity (home based), reciprocity (boundary), down-the-line trade, central place (redistribution), central place (market exchange), emissary trading, and port of trade. These modes of trade dictate the way that the Spondylus shells are traded, as well as who is benefiting the most from the trades. Modes such as central place (redistribution) require the entity that is the central place to be the one that gains the most benefit from the trade, and modes such as emissary trading and port of trade are the modes that started the "mercardes" class within the Central Andes. The value of Spondylus shells in the Central Andes stems from supply and demand. There was a lot of demand for Spondylus shells due to the "fetishistic needs to the south." Even today, there are collectors of ''Spondylus'' shells, and a commercial market exists for them. Additionally, some species (especially ''S. americanus'') are sometimes found in the saltwater aquariums. ''S. limbatus'' was commonly ground for mortar in Central America, giving raise to its junior synonym, "''S. calcifer"''. Some Mediterranean species are edible and are commonly consumed, with ''S. gaederopus'' in particular being popular in
Sardinia Sardinia ( ; it, Sardegna, label=Italian, Corsican and Tabarchino ; sc, Sardigna , sdc, Sardhigna; french: Sardaigne; sdn, Saldigna; ca, Sardenya, label=Algherese and Catalan) is the second-largest island in the Mediterranean Sea, after ...
. Tropical species, however, tend to bioaccumulate
saxitoxin Saxitoxin (STX) is a potent neurotoxin and the best-known paralytic shellfish toxin (PST). Ingestion of saxitoxin by humans, usually by consumption of shellfish contaminated by toxic algal blooms, is responsible for the illness known as paralyt ...
. The Romans ate them. Macrobius in Saturnalia III.13 describes a dinner party in 63 BCE in which there were two courses of spondylus.


Species

Spondylidae taxonomy has undergone many revisions, mostly due to the fact that identification is traditionally based on the shell only, and this is highly variable. To add to this, while some shallow-water species are extremely common, at least two deep-water ones are known from a single specimen, while a third (S. gravis) was only rediscovered after 77 years. At least another common species (S. regius) has a different shell when it grows in deep water. *'' Spondylus americanus'' Hermann, 1781 - Atlantic thorny oyster *'' Spondylus anacanthus'' Mawe, 1823 - nude thorny oyster *'' Spondylus aonis'' d'Orbigny, 1850 *'' Spondylus asiaticus'' Chenu, 1844 *'' Spondylus asperrimus'' G. B. Sowerby II, 1847 *'' Spondylus aucklandicus'' P. Marshall, 1918 *'' Spondylus avramsingeri'' Kovalis, 2010 * † '' Spondylus bostrychites'' Guppy, 1867 *'' Spondylus butleri'' Reeve, 1856 *'' Spondylus candidus''
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1819
*'' Spondylus clarksoni'' Lamprell, 1992 *'' Spondylus concavus'' Deshayes in Maillard, 1863 *'' Spondylus crassisquama''
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1819
*'' Spondylus croceus'' Schreibers, 1793 *'' Spondylus darwini'' Jousseaume, 1882 *'' Spondylus deforgesi'' Lamprell & Healy, 2001 *'' Spondylus depressus'' Fulton, 1915 *'' Spondylus eastae'' Lamprell, 1992 *'' Spondylus echinatus'' Schreibers, 1793 *'' Spondylus erectospinosus'' Habe, 1973 *'' Spondylus exiguus'' Lamprell & Healy, 2001 *'' Spondylus exilis'' G. B. Sowerby III, 1895 *'' Spondylus fauroti'' Jousseaume, 1888 *'' Spondylus foliaceus'' Schreibers, 1793 *''
Spondylus gaederopus ''Spondylus gaederopus'' is a species of marine bivalve mollusc, a thorny oyster in the family Spondylidae. This species is endemic to the Mediterranean Sea. Description ''Spondylus gaederopus'' attaches itself to the substrate with its lower ...
''
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, 1758
- European thorny oyster *'' Spondylus gloriandus'' Melvill & Standen, 1907 *'' Spondylus gloriosus'' Dall, Bartsch & Rehder, 1938 * † '' Spondylus granulosus'' Deshayes, 1830 *'' Spondylus gravis'' Fulton, 1915 *'' Spondylus groschi'' Lamprell & Kilburn, 1995 *'' Spondylus gussonii'' O. G. Costa, 1830 *'' Spondylus heidkeae'' Lamprell & Healy, 2001 *'' Spondylus imperialis'' Chenu, 1844 * '' Spondylus jamarci'' Okutani, 1983 *''
Spondylus Lamarckii ''Spondylus'' is a genus of bivalve molluscs, the only genus in the family Spondylidae.MolluscaBase (2019). MolluscaBase. Spondylus Linnaeus, 1758. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p ...
'' Chenu, 1845 *'' Spondylus layardi'' Reeve, 1856 *'' Spondylus leucacanthus'' Broderip, 1833 *'' Spondylus limbatus'' G. B. Sowerby II, 1847 *'' Spondylus linguafelis'' G. B. Sowerby II, 1847 *'' Spondylus maestratii'' Lamprell & Healy, 2001 *'' Spondylus marinensis'' Cossignani & Allary, 2018 *'' Spondylus mimus'' Dall, Bartsch & Rehder, 1938 *'' Spondylus morrisoni'' Damarco, 2015 *'' Spondylus multimuricatus'' Reeve, 1856 *'' Spondylus multisetosus'' Reeve, 1856 * † '' Spondylus multistriatus'' Deshayes, 1830 *'' Spondylus nicobaricus'' Schreibers, 1793 *'' Spondylus occidens'' G. B. Sowerby III, 1903 *'' Spondylus ocellatus'' Reeve, 1856 *'' Spondylus orstomi'' Lamprell & Healy, 2001 *'' Spondylus ostreoides'' E. A. Smith, 1885 *'' Spondylus pratii'' Parth, 1990 *'' Spondylus proneri'' Lamprell & Healy, 2001 * '' Spondylus pseudogaederopus'' T. Cossignani, 2022 * '' Spondylus purpurascens'' T. Cossignani, 2022 * † '' Spondylus radula'' Lamarck, 1806 *'' Spondylus raoulensis'' W. R. B. Oliver, 1915 * † '' Spondylus rarispina'' Deshayes, 1830 *'' Spondylus reesianus'' G. B. Sowerby III, 1903 *''
Spondylus regius ''Spondylus regius'', the regal thorny oyster, is a species of bivalve mollusc in the family Spondylidae. It can be found in the Western Pacific, and can grow 156 mm in length. Distribution and habitat ''Spondylus regius'' is found in the Red S ...
''
Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, the ...
, 1758
- regal thorny oyster *'' Spondylus rippingalei'' Lamprell & Healy, 2001 *'' Spondylus rubicundus'' Reeve, 1856 *'' Spondylus senegalensis'' Schreibers, 1793 *'' Spondylus sinensis'' Schreibers, 1793 *'' Spondylus spinosus'' Schreibers, 1793 *'' Spondylus squamosus'' Schreibers, 1793 *'' Spondylus tenellus'' Reeve, 1856 *'' Spondylus tenuis'' Schreibers, 1793 *'' Spondylus tenuispinosus'' G. B. Sowerby II, 1847 *'' Spondylus tenuitas'' Garrard, 1966 *'' Spondylus variegatus'' Schreibers, 1793 *''
Spondylus varius ''Spondylus varius'', is a species of large marine bivalve mollusc in the family Spondylidae, the spiny oysters. Description ''Spondylus varius'' is the largest of the spiny oysters, reaching a maximum size of about 20 cm. Aside from the s ...
'' G. B. Sowerby I, 1827 * † '' Spondylus vaudini'' Deshayes, 1858 *'' Spondylus versicolor'' Schreibers, 1793 *'' Spondylus victoriae'' G. B. Sowerby II, 1860 *'' Spondylus violacescens''
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1819
*'' Spondylus virgineus'' Reeve, 1856 *'' Spondylus visayensis'' Poppe & Tagaro, 2010 *'' Spondylus zonalis''
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1819
*'' Spondylus echinus'' Jousseaume in Lamy, 1927 (
taxon inquirendum In biological classification, a ''species inquirenda'' is a species of doubtful identity requiring further investigation. The use of the term in English-language biological literature dates back to at least the early nineteenth century. The term t ...
) *'' Spondylus imbricatus'' Perry, 1811 ( nomen dubium) *'' Spondylus microlepos''
Lamarck Jean-Baptiste Pierre Antoine de Monet, chevalier de Lamarck (1 August 1744 – 18 December 1829), often known simply as Lamarck (; ), was a French naturalist, biologist, academic, and soldier. He was an early proponent of the idea that biolo ...
, 1819
( nomen dubium) *'' Spondylus unicolor'' G. B. Sowerby II, 1847 ( nomen dubium) See also: Tikod amo, an undescribed species


References


External links


Spondylidae
pictures of the shells of most extant species.


Session Abstracts
on ''Spondylus'' research at the 13th Meeting of the European Association of Archaeologists at Zadar, Croatia, September 2007

from the website of the Gladys Archerd Shell Collection at Washington State University Tri-Cities Natural History Museum
Article on "notched" ''Spondylus''
Neolithic artifacts in Europe


Bibliography


A full and constantly updated bibliography
on ''Spondylus'' spp. in Aegean, Balkan, European and American contexts * Lamprell, Kevin L.: ''Spondylus: Spiny Oyster Shells of the World'', E. J. Brill, Leiden, 1987 {{Commons category Spondylidae Bivalve genera Extant Triassic first appearances