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'''''Leukoma staminea''''', commonly known as the '''Pacific littleneck clam''', the '''littleneck clam''', the '''rock cockle''', the '''hardshell clam''', the '''Tomales Bay cockle''', the '''rock clam''' or the '''ribbed carpet shell''',<ref name=WallaWalla/> is a species of [[bivalve]] [[mollusc]] in the [[family (biology)|family]] [[Veneridae]].<ref>{{cite web |url=http://shells.tricity.wsu.edu/ArcherdShellCollection/Bivalvia/Veneridae.html |title=Family: Veneridae (Venus Clams) |publisher=Archerd Shell Collection |accessdate=29 December 2018}}</ref> This species of mollusc was exploited by early humans in [[North America]]; for example, the [[Chumash people|Chumash]] peoples of [[Central California]] harvested these clams in [[Morro Bay]] approximately 1,000 years ago.<ref>{{cite web |url=http://www.megalithic.co.uk/article.php?sid=18353 |title=Los Osos Back Bay |author=Burnham, C. Michael |year=2008 |publisher=The Megalithic Portal |accessdate=29 December 2018}}</ref> and the distinctive shells form middens near their settlements,<ref>{{cite book|author1=Lightfoot, Kent |author2=Parrish, Otis |title=California Indians and Their Environment: An Introduction |url=https://books.google.com/books?id=bdAKPOEfrooC&pg=PA271 |year=2009 |publisher=University of California Press|isbn=978-0-520-94228-8 |page=271}}</ref>
'''''Leukoma staminea''''', commonly known as the '''Pacific littleneck clam''', the '''littleneck clam''', the '''rock cockle''', the '''hardshell clam''', the '''Tomales Bay cockle''', the '''rock clam''' or the '''ribbed carpet shell''',<ref name=WallaWalla/> is a species of [[bivalve]] [[mollusc]] in the [[family (biology)|family]] [[Veneridae]].<ref>{{cite web |url=http://shells.tricity.wsu.edu/ArcherdShellCollection/Bivalvia/Veneridae.html |title=Family: Veneridae (Venus Clams) |work=Gladys Archerd Shell Collection |publisher=[[Washington State University Tri-Cities]] Natural History Museum |accessdate=29 December 2018}}</ref> This species of mollusc was exploited by early humans in [[North America]]; for example, the [[Chumash people|Chumash]] peoples of [[Central California]] harvested these clams in [[Morro Bay]] approximately 1,000 years ago.<ref>{{cite web |url=http://www.megalithic.co.uk/article.php?sid=18353 |title=Los Osos Back Bay |author=Burnham, C. Michael |year=2008 |publisher=The Megalithic Portal |accessdate=29 December 2018}}</ref> and the distinctive shells form middens near their settlements,<ref>{{cite book|author1=Lightfoot, Kent |author2=Parrish, Otis |title=California Indians and Their Environment: An Introduction |url=https://books.google.com/books?id=bdAKPOEfrooC&pg=PA271 |year=2009 |publisher=University of California Press|isbn=978-0-520-94228-8 |page=271}}</ref>


==Description==
==Description==
Like other members of the [[Veneridae]] family, this species has a chalky shell, the umbo being anterior to the midline of the shell, but closer to the midline than to the anterior end of the shell. The two equal-sized valves are oval or heart-shaped. The width of the shell is greater than a quarter of its length, and the shell seldom exceeds {{convert|6|cm|in|1|abbr=on}} in length. The umbones point towards the anterior end of the shell. The hinge has three cardinal teeth in each valve, and a row of small teeth along the ventral margins of the valves. There are numerous concentric ridges, more clearly demarcked at the anterior end, but the radial ridges are often more clearly sculpted. The foot is large and there is a clearcut pallial sinus.<ref name=WallaWalla>{{cite web |url=https://inverts.wallawalla.edu/Mollusca/Bivalvia/Veneroida/Veneridae/Protothaca_staminea.html |title=''Leukoma staminea'' (Conrad, 1857) |author=Cowles, Dave |date= |work= |publisher= |accessdate=29 December 2018}}</ref>
Like other members of the [[Veneridae]] family, this species has a chalky shell, the umbo being anterior to the midline of the shell, but closer to the midline than to the anterior end of the shell. The two equal-sized valves are oval or heart-shaped. The width of the shell is greater than a quarter of its length, and the shell seldom exceeds {{convert|6|cm|in|1|abbr=on}} in length. The umbones point towards the anterior end of the shell. The hinge has three cardinal teeth in each valve, and a row of small teeth along the ventral margins of the valves. There are numerous concentric ridges, more clearly demarcked at the anterior end, but the radial ridges are often more clearly sculpted. The foot is large and there is a clearcut pallial sinus.<ref name=WallaWalla>{{cite web |url=https://inverts.wallawalla.edu/Mollusca/Bivalvia/Veneroida/Veneridae/Protothaca_staminea.html |title=''Leukoma staminea'' (Conrad, 1857) |work=Invertebrates of the Salish Sea |last=Cowles |first=Dave |year=2007 |publisher=Rosario Beach Marine Laboratory (a campus of [[Walla Walla University]], Washington) |accessdate=29 December 2018}}</ref>


==Distribution and habitat==
==Distribution and habitat==
Line 40: Line 40:


==Ecology==
==Ecology==
This clam is a [[filter feeder]] and consumes microscopic [[alga]]e such as [[dinoflagellate]]s, [[diatom]]s, and [[cyanobacteria]].<ref name = WallaWalla /><ref name = ClarkEtAl>{{cite journal |pmid=10485519 |year=1999 |last1=Clark |first1=R. F. |last2=Williams |first2=S. R. |last3=Nordt |first3=S. P. |last4=Manoguerra |first4=A. S. |title=A review of selected seafood poisonings |volume=26 |issue=3 |pages=175–184 |journal=[[Undersea & Hyperbaric Medicine]]|url=http://archive.rubicon-foundation.org/xmlui/bitstream/handle/123456789/2314/10485519.pdf}}</ref> Some dinoflagellates produce [[neurotoxin]]s, such as [[saxitoxin]] and its derivates, that [[bioaccumulate]] in the clams and other [[Bivalvia|bivalve mollusks]] and can cause [[paralytic shellfish poisoning]] (PSP) when the clams are eaten.<ref name = WallaWalla /><ref name = ClarkEtAl /> Despite this fact, the clam was eaten by Native Americans and is still used as a food for humans.<ref name = WallaWalla /> According to a 1996 report from the Marine Advisory Program at the [[University of Alaska]], the [[United States Food and Drug Administration]] (FDA) considers seafood unsafe if it contains more than 80&nbsp;[[micrograms|&mu;g]] of PSP-causing toxins per 100&nbsp;g of tissue of the seafood.<ref name = MAP1>{{cite web|url = https://seagrant.uaf.edu/features/PSP/PSP.pdf|title = Paralytic Shellfish Poisoning: The Alaska Problem|first = Raymond|last = RaLonde|work = Alaska's Maritime Resources|publisher = Marine Advisory Board, the [[University of Alaska]]|volume = 8|issue = 2|date = October 1996|pages = 1-7}}</ref><ref name = MAP2>{{cite web|url = https://seagrant.uaf.edu/features/PSP/PSP.pdf|title = How Toxic Are Alaska's Most Common Shellfish?|work = Alaska's Maritime Resources|publisher = Marine Advisory Board, the [[University of Alaska]]|volume = 8|issue = 2|date = October 1996|pages = 10-11}}</ref> PSP is caused by a mixture of at least 21 different chemical species, some of which undergo chemical transformations within the dinoflagellates or within the animals the acquire the saxitoxins, and which are retained by different animals for different lengths of time.<ref name = MAP1 /> It is clear that PSP-causing toxin levels are typically much higher in the summer months<ref name = MAP3>{{cite web|url = https://seagrant.uaf.edu/features/PSP/PSP.pdf|title = Epidemiology of Paralytic Shellfish Poisoning Outbreaks in Alaska|first = Brad|last = Gessner|work = Alaska's Maritime Resources|publisher = Marine Advisory Board, the [[University of Alaska]]|volume = 8|issue = 2|date = October 1996|pages = 16-17, 19}}</ref> though this does not mean the seafood is necessarily safe at other times.<ref name = MAP4>{{cite web|url = https://seagrant.uaf.edu/features/PSP/PSP.pdf|title = Truths and Myths about PSP|work = Alaska's Maritime Resources|publisher = Marine Advisory Board, the [[University of Alaska]]|volume = 8|issue = 2|date = October 1996|pages = 14-15}}</ref> Risks also vary based on species<ref name = MAP1 /><ref name = MAP2 /> but seafood available for retail sale is required to meet the FDA standards.<ref name = MAP4 />
This clam was eaten by native Americans and still is used for human food despite the fact that it can cause [[paralytic shellfish poisoning]].<ref name=WallaWalla/> [[Predation|Predators]] include such molluscs as the [[Ceratostoma foliatum|leafy hornmouth snail]] (''Ceratostoma foliatum'') and [[Neverita lewisii|Lewis's moon snail]] (''Neverita lewisii''), the crabs ''[[Metacarcinus magister]]'' and ''[[Cancer productus]]'', the [[giant Pacific octopus]] (''Enteroctopus dofleini'') and the [[sea otter]].<ref name=WallaWalla/> Fish such as the [[Pacific staghorn sculpin]] sometimes nips off the [[Siphon_(mollusc)#Paired_siphons_of_bivalves|siphons]] when they are extended to feed. This clam spawns during the summer in the north of its range. It is a slow-growing species and may live for up to sixteen years.<ref name=WallaWalla/>

In PSP poisonings that occurred in the summer of 1993 in [[Kodiak, Alaska]], saxitoxin levels as high as 19,600&nbsp;&mu;g&nbsp;/&nbsp;100&nbsp;g were measured in the Alaksa blue mussel ''[[Mytilus edulis]]'' &ndash; sufficient to provide a lethal dose in a single 2.5&nbsp;g mussel.<ref name = MAP1 /> By contrast, the highest measured level of saxitoxin in the Pacific Littleneck Clam was 580&nbsp;&mu;g&nbsp;/&nbsp;100&nbsp;g according to this 1996 report.<ref name = MAP2 /> Littleneck clams are "typically less toxic and retain their toxins for a shorter amount of time than the other species" &nbsp; such as "butter clams and blue mussels [that] tend to accumulate the highest levels of PSP toxins ... [and] geoducks and scallops [that] tend to be toxic for longer periods of time" &nbsp; but this does not mean that these clams "are always safe to eat [as] you can get PSP from littleneck clams."<ref>{{cite web|title = Paralytic Shellfish Poisoning in Alaska Facts and Discussion|url = https://seagrant.uaf.edu/topics/environmental-hazards-alaskas-coasts/paralytic-shellfish-poisoning/psp-facts.php|accessdate = 22 January 2019|date = 13 August 2015|author = Alaska Sea Grant Program|publisher = College of Fisheries and Ocean Sciences at the [[University of Alaska Fairbanks]]}}</ref> The reason for the difference is likely that the littleneck clam has an [[enzyme]] that converts saxitoxin into [[decarbamoylsaxitoxin]], a capability not shared by the blue mussel nor by the butter clam ''[[Saxidomus gigantea]]'' (in which levels of saxitoxin as high as 7,750&nbsp;&mu;g&nbsp;/&nbsp;100&nbsp;g have been reported<ref name = MAP2 />).<ref>{{cite journal|title = Enzymatic transformation of PSP toxins in the littleneck clam (''Protothaca staminea'')|first1 = John J.|last1 = Sullivan|first2 = Wayne T.|last2 = Iwaoka|first3 = John|last3 = Liston|journal = [[Biochemical and Biophysical Research Communications]]|volume = 114|issue = 2|year = 1983|pages = 465-472|doi = 10.1016/0006-291X(83)90803-3|pmid = 6882435}}</ref> This transformation to decarbomyl derivative has been reported in some other clam species and significantly reduces the toxicity of saxitoxins present.<ref>{{cite journal|title = Non-Traditional Vectors for Paralytic Shellfish Poisoning|first1 = Jonathan R.|last1 = Deeds|first2 = Jan H.|last2 = Landsberg|first3 = Stacey M.|last3 = Etheridge|first4 = Grant C.|last4 = Pitcher|first5 = Sara Watt|last5 = Longan|journal = [[Marine Drugs]]|year = 2008|volume = 6|issue = 2|pages = 308–348|doi = 10.3390/md20080015|pmc = 2525492|pmid = 18728730}}</ref>

[[Predation|Predators]] include such molluscs as the [[Ceratostoma foliatum|leafy hornmouth snail]] (''Ceratostoma foliatum'') and [[Neverita lewisii|Lewis's moon snail]] (''Neverita lewisii''), the crabs ''[[Metacarcinus magister]]'' and ''[[Cancer productus]]'', the [[giant Pacific octopus]] (''Enteroctopus dofleini'') and the [[sea otter]].<ref name=WallaWalla/> Fish such as the [[Pacific staghorn sculpin]] sometimes nips off the [[Siphon_(mollusc)#Paired_siphons_of_bivalves|siphons]] when they are extended to feed. This clam spawns during the summer in the north of its range. It is a slow-growing species and may live for up to sixteen years.<ref name=WallaWalla/>


==References==
==References==

Revision as of 23:43, 21 January 2019

Littleneck clam
Scientific classification
Kingdom:
Phylum:
Class:
Order:
Family:
Genus:
Species:
L. staminea
Binomial name
Leukoma staminea
(Conrad, 1837)[1]
Synonyms[1]
  • Chione ruderata Deshayes, 1853
  • Paphia staminea (Conrad, 1837)
  • Paphia staminea orbella Carpenter, 1864
  • Paphia staminea var. sulculosa Dall, 1902
  • Protothaca grewingkii Dall, 1904
  • Protothaca staminea (Conrad, 1837)
  • Tapes diversa G. B. Sowerby II, 1852
  • Tapes tumida G. B. Sowerby II, 1852
  • Tapes tumida Carpenter, 1857
  • Venerupis petiti Deshayes, 1839
  • Venus ampliata Carpenter, 1857
  • Venus conradi Römer, 1867
  • Venus mundulus Reeve, 1863
  • Venus pectunculoides Valenciennes, 1846
  • Venus rigida Gould, 1850
  • Venus staminea Conrad, 1837

Leukoma staminea, commonly known as the Pacific littleneck clam, the littleneck clam, the rock cockle, the hardshell clam, the Tomales Bay cockle, the rock clam or the ribbed carpet shell,[2] is a species of bivalve mollusc in the family Veneridae.[3] This species of mollusc was exploited by early humans in North America; for example, the Chumash peoples of Central California harvested these clams in Morro Bay approximately 1,000 years ago.[4] and the distinctive shells form middens near their settlements,[5]

Description

Like other members of the Veneridae family, this species has a chalky shell, the umbo being anterior to the midline of the shell, but closer to the midline than to the anterior end of the shell. The two equal-sized valves are oval or heart-shaped. The width of the shell is greater than a quarter of its length, and the shell seldom exceeds 6 cm (2.4 in) in length. The umbones point towards the anterior end of the shell. The hinge has three cardinal teeth in each valve, and a row of small teeth along the ventral margins of the valves. There are numerous concentric ridges, more clearly demarcked at the anterior end, but the radial ridges are often more clearly sculpted. The foot is large and there is a clearcut pallial sinus.[2]

Distribution and habitat

Leukoma staminea is native to the eastern Pacific Ocean. Its range extends along the coasts of North America from the Aleutian Islands and Alaska in the north to Baja California in the south. It usually occurs in protected areas on sand, hard mud and clayey-gravel substrates from the mid and lower shore down to depths of about 10 m (33 ft), usually buried less than 8 cm (3 in) beneath the surface of the sediment. Occasionally it is in more exposed locations, in gravel-filled cracks in rocks or in empty burrows of other clams.[2]

Ecology

This clam is a filter feeder and consumes microscopic algae such as dinoflagellates, diatoms, and cyanobacteria.[2][6] Some dinoflagellates produce neurotoxins, such as saxitoxin and its derivates, that bioaccumulate in the clams and other bivalve mollusks and can cause paralytic shellfish poisoning (PSP) when the clams are eaten.[2][6] Despite this fact, the clam was eaten by Native Americans and is still used as a food for humans.[2] According to a 1996 report from the Marine Advisory Program at the University of Alaska, the United States Food and Drug Administration (FDA) considers seafood unsafe if it contains more than 80 μg of PSP-causing toxins per 100 g of tissue of the seafood.[7][8] PSP is caused by a mixture of at least 21 different chemical species, some of which undergo chemical transformations within the dinoflagellates or within the animals the acquire the saxitoxins, and which are retained by different animals for different lengths of time.[7] It is clear that PSP-causing toxin levels are typically much higher in the summer months[9] though this does not mean the seafood is necessarily safe at other times.[10] Risks also vary based on species[7][8] but seafood available for retail sale is required to meet the FDA standards.[10]

In PSP poisonings that occurred in the summer of 1993 in Kodiak, Alaska, saxitoxin levels as high as 19,600 μg / 100 g were measured in the Alaksa blue mussel Mytilus edulis – sufficient to provide a lethal dose in a single 2.5 g mussel.[7] By contrast, the highest measured level of saxitoxin in the Pacific Littleneck Clam was 580 μg / 100 g according to this 1996 report.[8] Littleneck clams are "typically less toxic and retain their toxins for a shorter amount of time than the other species"   such as "butter clams and blue mussels [that] tend to accumulate the highest levels of PSP toxins ... [and] geoducks and scallops [that] tend to be toxic for longer periods of time"   but this does not mean that these clams "are always safe to eat [as] you can get PSP from littleneck clams."[11] The reason for the difference is likely that the littleneck clam has an enzyme that converts saxitoxin into decarbamoylsaxitoxin, a capability not shared by the blue mussel nor by the butter clam Saxidomus gigantea (in which levels of saxitoxin as high as 7,750 μg / 100 g have been reported[8]).[12] This transformation to decarbomyl derivative has been reported in some other clam species and significantly reduces the toxicity of saxitoxins present.[13]

Predators include such molluscs as the leafy hornmouth snail (Ceratostoma foliatum) and Lewis's moon snail (Neverita lewisii), the crabs Metacarcinus magister and Cancer productus, the giant Pacific octopus (Enteroctopus dofleini) and the sea otter.[2] Fish such as the Pacific staghorn sculpin sometimes nips off the siphons when they are extended to feed. This clam spawns during the summer in the north of its range. It is a slow-growing species and may live for up to sixteen years.[2]

References

  1. ^ a b Huber, Markus (2018). "Leukoma staminea (Conrad, 1837)". WoRMS. World Register of Marine Species. Retrieved 29 December 2018.
  2. ^ a b c d e f g h Cowles, Dave (2007). "Leukoma staminea (Conrad, 1857)". Invertebrates of the Salish Sea. Rosario Beach Marine Laboratory (a campus of Walla Walla University, Washington). Retrieved 29 December 2018.
  3. ^ "Family: Veneridae (Venus Clams)". Gladys Archerd Shell Collection. Washington State University Tri-Cities Natural History Museum. Retrieved 29 December 2018.
  4. ^ Burnham, C. Michael (2008). "Los Osos Back Bay". The Megalithic Portal. Retrieved 29 December 2018.
  5. ^ Lightfoot, Kent; Parrish, Otis (2009). California Indians and Their Environment: An Introduction. University of California Press. p. 271. ISBN 978-0-520-94228-8.
  6. ^ a b Clark, R. F.; Williams, S. R.; Nordt, S. P.; Manoguerra, A. S. (1999). "A review of selected seafood poisonings" (PDF). Undersea & Hyperbaric Medicine. 26 (3): 175–184. PMID 10485519.
  7. ^ a b c d RaLonde, Raymond (October 1996). "Paralytic Shellfish Poisoning: The Alaska Problem" (PDF). Alaska's Maritime Resources. Marine Advisory Board, the University of Alaska. pp. 1–7.
  8. ^ a b c d "How Toxic Are Alaska's Most Common Shellfish?" (PDF). Alaska's Maritime Resources. Marine Advisory Board, the University of Alaska. October 1996. pp. 10–11.
  9. ^ Gessner, Brad (October 1996). "Epidemiology of Paralytic Shellfish Poisoning Outbreaks in Alaska" (PDF). Alaska's Maritime Resources. Marine Advisory Board, the University of Alaska. pp. 16–17, 19.
  10. ^ a b "Truths and Myths about PSP" (PDF). Alaska's Maritime Resources. Marine Advisory Board, the University of Alaska. October 1996. pp. 14–15.
  11. ^ Alaska Sea Grant Program (13 August 2015). "Paralytic Shellfish Poisoning in Alaska Facts and Discussion". College of Fisheries and Ocean Sciences at the University of Alaska Fairbanks. Retrieved 22 January 2019.
  12. ^ Sullivan, John J.; Iwaoka, Wayne T.; Liston, John (1983). "Enzymatic transformation of PSP toxins in the littleneck clam (Protothaca staminea)". Biochemical and Biophysical Research Communications. 114 (2): 465–472. doi:10.1016/0006-291X(83)90803-3. PMID 6882435.
  13. ^ Deeds, Jonathan R.; Landsberg, Jan H.; Etheridge, Stacey M.; Pitcher, Grant C.; Longan, Sara Watt (2008). "Non-Traditional Vectors for Paralytic Shellfish Poisoning". Marine Drugs. 6 (2): 308–348. doi:10.3390/md20080015. PMC 2525492. PMID 18728730.{{cite journal}}: CS1 maint: unflagged free DOI (link)