Overview

Comprehensive Description

At hot vents only. The life habit of the adult is semiepibiotic on bare basalt, living in cracks and crevices from which hydrothermal fluids emanate; early life history stages are often encountered living within basaltic rubble associated with such warm cracks and crevices. The presence of this species is correlated with elevated levels of hydrogen sulphide. The soft tissue is coloured dark red when retrieved living due to the presence of intracellular hemoglobin. The thick and large gills contain sulfur oxidizing chemoautotrophic symbionts. The large rugose foot is often seen protruding when the clams are viewed in life position.
  • BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.

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Distribution

Discovered in 1977, the vesicomyid clam Calyptogena magnifica populates the deep-sea hydrothermal vent areas of the East Pacific Rise and the Galapagos Rift. Only a handful of scientists have studied this species, and it has yet to be found elsewhere.

Biogeographic Regions: pacific ocean (Native )

  • Boss, K., R. Turner. 1980. The giant white clam from the Galapagos Rift, Calyptogena magnifica species novum. Malacologia, 20(1): 161-194.
  • Terwilliger, R., N. Terwilliger, A. Arp. 1983. Thermal vent clam (Calyptogena magnifica) hemoglobin. Science, 219(4587): 981-983.
  • Van Dover, C. 2000. The ecology of deep-sea hydrothermal vents. Princeton, NJ: Princeton University Press.
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Entire Northern East Pacific Rise and Southern East Pacific Rise: 21°N to 22°S; Galapagos Spreading Center. ARP A. J., CHILDRESS J.J. & C.J. FISHER (1984) Physiol. l Zool. 57: 648-662.
  • BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.

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Physical Description

Morphology

Calyptogena magnifica is a heterodont bivalve that can reach 26 cm in length. The valves are white and fairly elliptical in shape, and individuals grow an average of 4 cm per year. This species has the basic bivalvian body plan, with a few unique features. Most notable are the communities of sulfur-oxidizing chemolithotrophic bacterial symbionts within the gills and other tissues. The bacteria produce organic carbon and nitrogen, which serve as nutrition for the clam. As a result, the digestive system and labial palps of the bivalve are extremely reduced, and the foot and gills are highly vascularized to better facilitate gas exchange and hydrogen sulfide uptake. The visceral mass is a conspicuous red color due to intracellular hemoglobin, and the circulatory system is about 44% of the clam’s weight.

Range length: 10 to 26 cm.

Other Physical Features: ectothermic ; heterothermic ; bilateral symmetry

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Boss and Turner (1980) described the anatomy of the species. Below is a summary of their observations:

Mantle: Pedal aperture large. Middle mantle fold with short papillae. Short, separate incurrent and excurrent siphons formed by fusion of the inner mantle folds.

Foot: Large, with byssal groove and gland, but no byssal threads produced by adult specimens.

Gills: Large, flat (non-plicate), with inner and outer demibranchs displaying both lamellae. Formed by filaments of identical morphology (homorhabdic).

Digestive tract: Labial palps very small. Rectum passes through the pericardial cavity and the ventricle of the heart.

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Shell very large, variable, oval to elongate-oval in outline, inequilateral and equivalve, usually without gape; valves white, entirely aragonitic; periostracum present, but frequently eroded away on older portions of shell; umbos prosogyrous, sometimes partially enrolled; lunule and escutcheon variable, present or absent; ligament external, opisthodetic, and parivincular; foot strong and rugose, with byssal gland.
  • BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.

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Size

Shell length up to 263 mm.
  • BOSS K. & R. TURNER (1980) Malacologia 20: 161-194.

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Diagnostic Description

Shell white, brittle, elongate-ovate, with the right and left valves equal in size, outline and inflation (equivalve). Umbones at the anterior third, with the beaks poorly defined. Periostracum brown, dehiscent (absent from the older portions of the shell). External sculpture of irregular commarginal growth increments generally restricted to the margins of the shell flank. Internal surface and margins smooth. Pallial sinus absent. Ventral surface of the hinge plate with a series of small scars. Ligament external, supported by nymphal ridges, extending along most of the postero-dorsal margin of the shell. Hinge teeth represented by irregular cardinals; lateral teeth absent.

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Ecology

Habitat

Calyptogena magnifica thrives in the lush areas surrounding hydrothermal vents. Using the foot and byssal threads as an anchor, these organisms pack, or clump, themselves in the crevices of basalt on the ocean floor. These clumps are termed “clambakes.” Within the cracks, there is a low flow of warm, carbon dioxide and hydrogen sulfide rich vent water, no sunlight, over a thousand atmospheres of pressure, and a temperature of about 10-15 degrees Celsius (C). The visceral mass of the animal experiences a temperature of about 2-4 degrees C, due to its location above the foot. The siphons of the clam are used to tap into the ambient oxygen and carbon dioxide above the valves. Population density increases with increasing concentrations of hydrogen sulfide.

Average depth: 2000 m.

Habitat Regions: saltwater or marine

Aquatic Biomes: oceanic vent

  • Cary, S., S. Giovannoni. 1993. Transovarial inheritance of endosymbiotic bacteria in clams inhabiting deep-sea hydrothermal vents and cold seeps. Proceedings of the National Academy of Sciences of the United States of America, 90: 5695-5699. Accessed October 19, 2012 at http://www.pnas.org/content/90/12/5695.full.pdf+html.
  • Powell, M., G. Somero. 1986. Adaptations to sulfide by hydrothermal vent animals: sites and mechanisms of detoxification and metabolism. The Biological Bulletin, 171: 274-290. Accessed October 20, 2012 at http://www.biolbull.org/content/171/1/274.full.pdf+html.
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Depth range based on 21 specimens in 1 taxon.
Water temperature and chemistry ranges based on 21 samples.

Environmental ranges
  Depth range (m): 2451 - 2632.5
  Temperature range (°C): 1.823 - 2.075
  Nitrate (umol/L): 35.653 - 41.112
  Salinity (PPS): 34.663 - 34.674
  Oxygen (ml/l): 2.461 - 3.601
  Phosphate (umol/l): 2.483 - 2.812
  Silicate (umol/l): 126.871 - 161.139

Graphical representation

Depth range (m): 2451 - 2632.5

Temperature range (°C): 1.823 - 2.075

Nitrate (umol/L): 35.653 - 41.112

Salinity (PPS): 34.663 - 34.674

Oxygen (ml/l): 2.461 - 3.601

Phosphate (umol/l): 2.483 - 2.812

Silicate (umol/l): 126.871 - 161.139
 
Note: this information has not been validated. Check this *note*. Your feedback is most welcome.

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Trophic Strategy

Calyptogena magnifica suspension-feeds on particles rich in nitrogen and lipid compounds present in the hydrothermal fluid. It also receives nutrients through a symbiotic relationship with sulfur-metabolizing bacteria that are located on the outer region of its gill tissue.

Other Foods: microbes

Foraging Behavior: filter-feeding

  • Grassle, J. 1985. Hydrothermal vent animals: distribution and biology. Science, 229(4715): 713-717.
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Associations

Calyptogena magnifica has a mutualistic relationship with sulfur-metabolizing bacteria located on its gill tissue, foot, and mantle. It depends on these sulfur-metabolizing bacteria to receive its nutrients from the rich hydrogen sulfide environment of the hydrothermal vent. The bacteria located on the outer layer of the foot and mantle also provide peripheral defense by detoxifying the sulfide as soon as it enters the body. High molecular weight factors in the blood bind the sulfide tightly to extract the sulfide from the environment. The sulfide is then transported to the bacterial symbiont via circulation. As a result, low concentration of free sulfide is found in the blood, protecting the aerobic respiration of the organisms from poisoning by sulfide due to its sensitivity to cytochrome-c oxidase and the enzymes involved in carbon fixation in the symbiont.

Mutualist Species:

  • sulfur-metabolizing bacteria

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Direct predation of Calyptogena magnifica has been observed by mobile grazers such as small gastropods, amphipods, and crabs. These predators consume newly settled larvae and juveniles. At the same time, indirect predation has also been observed by the removal of bacterial film on rocks by these mobile grazers. Without the bacterial film on rocks, larvae lose the marker that indicates where to settle during development. Large epibenthic predators such as zoarcid fish, Thermarces cerberus, indirectly aid in decreasing the mortality of sessile invertebrates, such as C. magnifica, by feeding on small mobile grazers. Also, the tissue in C. magnifica is considered unpalatable to predators when hydrogen sulfide is released from the sulfide-metabolizing bacterial symbiont on the gill tissue. Once an adult, these clams have a thick shell that provides a structural defense against predators.

Known Predators:

  • Deep sea vent gastropods
  • Deep sea vent amphipods
  • Brachyuran crab, Bythograea thermidron

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Life History and Behavior

Behavior

Calyptogena magnifica has a synchronized but not always consistent release of sperm and eggs. The males detect an increase in water temperature and release sperm through their exhalant siphons. In response, the females release eggs from their exhalant siphons when a threshold of sperm or associated chemicals is detected. However, the water current must be slow for the females to detect the high concentration of chemical cues. If either or both conditions are not met, the females will not release their eggs. The neurotransmitter serotonin is commonly responsible for the stimulation of the release and re-initiation of meiosis in the oocyctes and may be one of the chemical cues responsible for egg release in C. magnifica. Calyptogena magnifica has an inhalant siphon that is used to sense the chemical environment from the incoming flow of water.

Communication Channels: chemical

Perception Channels: infrared/heat ; tactile ; chemical

  • Fujikura, K., K. Amaki, J. Barry, Y. Fujiwara, V. Furushima, R. Iwase, H. Yamamoto, T. Maruyama. 2007. Long-term in situ monitoring of spawining behavior and fecundity in Calyptogena spp. Marine Ecology Progress Series, 333: 185-193.
  • Krylova, E., H. Sahling. 2005. Recent bivalve molluscs of the genus Calyptogena (Vesicomyidae). Journal of MolluscanStudies, 72(4): 359-395. Accessed October 20, 2012 at http://mollus.oxfordjournals.org/content/72/4/359.full.pdf+html.
  • Micheli, F., C. Peterson, L. Mullineaux, C. Fisher, S. Mills, G. Sancho, G. Johnson, H. Lenihan. 2002. Predation structures communities at deep-sea hydrothermal vents. Ecological Monographs, 72(3): 365–382. Accessed January 03, 2013 at http://micheli.stanford.edu/pdf/69-Michelietal2002EcolMonog.pdf.
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Life Cycle

Calyptogena magnifica larvae are lecithotrophic, and nonplanktonic. Larvae are free-swimming and rely solely on their stored yolk reserves for energy. Once contacting a substrate, the larvae metamorphoses into the adult form. This species has indeterminate growth, as the shell of the bivalve grows in annual, evident increments.

Development - Life Cycle: metamorphosis ; indeterminate growth

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Life Expectancy

Calyptogena magnifica individuals can survive anywhere from 3.5 to approximately 25 years. The hydrothermal vents where they anchor themselves last only tens of years, so strong colonization abilities are favored over longevity since adults are non-motile. The shell of the bivalve grows in annual increments, which makes it possible to determine the age.

Average lifespan

Status: wild:
25 years.

Typical lifespan

Status: wild:
3.5 to 25 years.

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Reproduction

Once sexually mature, Calyptogena magnifica gametes are released into the environment continuously and in small numbers by all individuals. Egg cells range from 105-195 micrometers, and the heads of sperm cells are about 3 micrometers in diameter. Fertilization results from any successful union of an egg cell and a sperm cell. Therefore, mating is random and results in high gene flow and genetic variability.

Mating System: polygynandrous (promiscuous)

Calyptogena magnifica is a sessile bivalve species. Sexes are separate (dioecious) and fertilization is external and non-specific. Once clams are greater than 6 cm in length (between 1 and 4 years old) they begin maturing sexually, and by 9-10 cm long they are considered ripe. The gonads then start filling, and complete sexual maturity is reached at 12-14 cm. Since environmental stimuli are largely absent, spawning is continuous and takes place between the ages of 3 and 15. Thus, breeding occurs year round. Large clams remain almost fully ripe, as only a small proportion of gametes are released at any one time. Although dispersal distance is limited, this species has no problem effectively dispersing gametes from individuals throughout its range.

Breeding interval: Calyptogena magnifica spawns year round.

Range age at sexual or reproductive maturity (female): 1 to 4 years.

Range age at sexual or reproductive maturity (male): 1 to 4 years.

Key Reproductive Features: year-round breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; fertilization (Internal )

Due to the newness of vent invertebrate studies, very little is known about the degree of parental involvement in Calyptogena magnifica. The yolk reserves of the lecithotrophic larvae are the only known maternal support the offspring receive.

Parental Investment: pre-fertilization (Provisioning)

  • Berg, C. 1985. Reproductive strategies of mollusks from abyssal hydrothermal vent communities. Bulletin of the Biological Society of Washington, 6: 185-197.
  • Boss, K., R. Turner. 1980. The giant white clam from the Galapagos Rift, Calyptogena magnifica species novum. Malacologia, 20(1): 161-194.
  • Cary, S., S. Giovannoni. 1993. Transovarial inheritance of endosymbiotic bacteria in clams inhabiting deep-sea hydrothermal vents and cold seeps. Proceedings of the National Academy of Sciences of the United States of America, 90: 5695-5699. Accessed October 19, 2012 at http://www.pnas.org/content/90/12/5695.full.pdf+html.
  • Van Dover, C. 2000. The ecology of deep-sea hydrothermal vents. Princeton, NJ: Princeton University Press.
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Molecular Biology and Genetics

Molecular Biology

Barcode data: Calyptogena magnifica

The following is a representative barcode sequence, the centroid of all available sequences for this species.


There are 12 barcode sequences available from BOLD and GenBank.

Below is a sequence of the barcode region Cytochrome oxidase subunit 1 (COI or COX1) from a member of the species.

See the BOLD taxonomy browser for more complete information about this specimen and other sequences.

GGAGTTTTAGGAGAT---AGTCATTTGTACCAGGTAGTGGTTACTGCACATGGTTTGTTGATGATTTTCTTCTTAGTAATGCCGATGATGATTGGGGGTTTTGGGAATTGATTAGTACCTTTAATG---TTGCAAATTCCTGATATGGCTTTTCCACGGATGAATAATTTAAGATTTTGGTTAATACCTAGGTCGGTTTTAATGCTTCTTGGCTCTGCTTATGTAGAGAGTGGGGCAGGTACTGGTTGAACTATTTATCCCCCATTGTCTAGTATTATAGGGCATGCTGGTCCATCTGTAGATTTT---GTAATTTTGTCTCTTCATTTAGGTGGCGTGTCTTCTATTTTGGCTTCAATTAATTTTGTTGTTACTTCTCTTTGTATGCGTGTTGAAGCTTTATCGTTATTGCGTGTGACTATGTTTGTTTGATGTGTTGCTGTCACTGGATTTTTACTGATTATTGCTATACCTGTCTTGGCTGGGAGA---TTAACAATACTATTAACTGATCGTAATTTTAATACTAGTTTTTTTGATCCTGCTGGGTTGGGGGATCCTATTCTCTTTGGGCATTTGGTTTGATTTTTTGGACACCCTGAGGTTTATATTTTGATTTTACCTGGATTTGGGATTATTTCCCACGTGATTAAAGTTGGGAGGAGAAAGCTT---GAATTGTTTGGAAAAGTTCCTATAATTTATGCTGTTTTATCTATTGGATTTTTAGGGTTTATTGTTTGG
-- end --

Download FASTA File

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Statistics of barcoding coverage: Calyptogena magnifica

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 12
Specimens with Barcodes: 12
Species With Barcodes: 1
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Conservation

Conservation Status

This species is not under any protection status.

US Federal List: no special status

CITES: no special status

State of Michigan List: no special status

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Relevance to Humans and Ecosystems

Benefits

There are no known adverse effects of Calyptogena magnifica on humans.

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The shells of Calyptogena magnifica can be used to study the thermal and chemical history of hydrothermal vent systems in the mid-ocean ridge and volcanic activity through a time series. By measuring the strontium/calcium ratio and the annual growth rate observed on the shell, the hydrothermal vent temperature can be studied over time and used to evaluate eruptions that may have occurred. Also, its shells are composed of calcium carbonate and will dissolve at a rate dependent on shell mineralogy, shell microstructure, and proximity to hydrothermal vent fluids. The rate of shell dissolution provides information in determining the longevity of hydrothermal vent activity along the rise axis.

Positive Impacts: research and education

  • Kennish, M., R. Lutz. 1999. Calcium carbonate dissolution rates in deep-sea bivalve shells on the East Pacific Rise at 21°N: results of an 8-year in-situ experiment. Palaeogeography, Palaeoclimatology, Palaeoecology, 154(4): 293-299. Accessed October 20, 2012 at http://www.sciencedirect.com/science/article/pii/S0031018299001029.
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Wikipedia

Calyptogena magnifica

Calyptogena magnifica is a species of giant white clam found clustered around hydrothermal vents at abyssal depths in the Pacific Ocean.

Contents

Description

The systematics of the family Vesicomyidae is unclear because of the small number of specimens collected, the variability between specimens of the same species and their wide dispersal in isolated, deep water locations. The morphology of Calyptogena magnifica resembles another member of the genus, Calyptogena elongata, the type locality of which is several hundred miles further north. C elongata is only known from three small specimens and the size of mature individuals is unknown.[2]

The two valves of Calyptogena magnifica are oval or slightly kidney-shaped and about two times as long as they are high. The umbones are towards the anterior end of the valve and the growth rings are most noticeable near the margins. The shell material is thick and the exterior is white and usually chalky in appearance. The periostracum is yellowish brown, wrinkled and loose. The ligament is external and there are several U-shaped cardinal hinge teeth on each valve. The largest specimen so far collected has a valve length of 263 millimetres (10.4 in). The mantle is an iridescent purplish pink and there is a large pink protrudible foot divided into two portions. The two separate siphons are short and do not extend beyond the edge of the valves. The pallial sinus is small. The gills are large and thick and the visceral mass is red due to the haemoglobin in the blood.[2]

Distribution

Calyptogena magnifica was first described by Kenneth Boss and Ruth Turner of Harvard's Museum of Comparative Zoology, in 1980, following its discovery during research dives by the submersible vehicle DSV Alvin to the floor of the Pacific Ocean in 1977 and 1979. The location of the thermal vent where it was found was approximately 200 miles (320 km) west of Punta Mita, Mexico at a depth of 2,645 metres (8,678 ft).[2] Further deep water exploration shows that it is present at other thermal vents on the East Pacific Rise between 21°N and 22°S as well as in the Galapagos Rift. In some locations it is plentiful while in other, apparently suitable habitats, it is not present at all.[3]

Biology

Calyptogena magnifica is assumed to burrow and it is thought the divided foot may be specially adapted for insertion into cracks in hard substrates or among mussels (Bathymodiolus thermophilus). The animal can move around on the sea floor with its muscular foot and usually takes up a vertical position rather than lying flat.[2]

Calyptogena magnifica is specially adapted to life round hydrothermal vents by the chemosymbiotic bacteria it harbours in its gills which oxidize hydrogen sulfide seeping from the vents. The clam absorbs nutrients produced by these bacteria rather than photosynthetically derived products and no longer has guts.[4]

Little is known of the reproduction and life cycle of Calyptogena magnifica but examination of specimens brought up from the deep showed numerous large oocytes with yolks in various stages of development among the visceral mass.[2] Researchers thought this might mean that the clam had poor dispersal abilities but a study using rDNA analysis showed that larvae did in fact disperse to other vents throughout its range.[3] Hydrothermal vents emit hot, sulfur-rich water for several years and then cease to flow. This results in the death of the community surrounding them, and for the continuing existence of their species, there is a need for the larvae of these animals to have dispersed to other existing vents and for them to exploit new vents when they open up.[4]

Ecology

Calyptogena magnifica was found near thermal vents in the deep sea floor where it was part of a rich benthic community. There were considerable numbers of empty shells and a few live individuals in the small area studied. The clams were lodged in crevices among a large number of mussels and some large galatheid crabs were observed walking over the bed of bivalves. Shrimps and octopuses were also observed in the vicinity.[2]

References

  1. ^ Tran, Bastien (2010). "Calyptogena magnifica Boss & Turner, 1980". World Register of Marine Species. http://www.marinespecies.org/aphia.php?p=taxdetails&id=464366. Retrieved 2012-04-14.
  2. ^ a b c d e f Boss, K. J.; Turner, R. D. (1980). "The giant white clam from the Galapagos Rift, Calyptogena magnifica species novum". Malacologia 20 (1): 161–194. 
  3. ^ a b Deep-Sea Vent Clams Monterey Bay Aquarium Research Institute. Retrieved 2012-04-15.
  4. ^ a b Hydrothermal vents Deep Ocean. Retrieved 2012-04-15.
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