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Coccolith

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1096: 921:, arranged in continuous arrays. The two coccolith types were originally thought to be produced by different families of coccolithophores. Now, however, it is known through a mix of observations on field samples and laboratory cultures, that the two coccolith types are produced by the same species but at different life cycle phases. Heterococcoliths are produced in the diploid life-cycle phase and holococcoliths in the haploid phase. Both in field samples and laboratory cultures, there is the possibility of observing a cell covered by a combination of heterococcoliths and holococcoliths. This indicates the transition from the diploid to the haploid phase of the species. Such combination of coccoliths has been observed in field samples, with many of them coming from the Mediterranean. 109: 97: 889: 1043: 1059: 1095: 1028: 215: 797: 20: 925: 183:. These are estimated to consume about two-thirds of the primary production in the ocean  and microzooplankton can exert a strong grazing pressure on coccolithophore populations. Although calcification does not prevent predation, it has been argued that the coccosphere reduces the grazing efficiency by making it more difficult for the predator to utilise the organic content of coccolithophores. 2567: 1382: 1331: 1072: 170:
As of 2021, it is not known why coccolithophores calcify and how their ability to produce coccoliths is associated with their ecological success. The most plausible benefit of having a coccosphere seems to be a protection against predators or viruses. Viral infection is an important cause of
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Monteiro, Fanny M.; Bach, Lennart T.; Brownlee, Colin; Bown, Paul; Rickaby, Rosalind E. M.; Poulton, Alex J.; Tyrrell, Toby; Beaufort, Luc; Dutkiewicz, Stephanie; Gibbs, Samantha; Gutowska, Magdalena A.; Lee, Renee; Riebesell, Ulf; Young, Jeremy; Ridgwell, Andy (2016).
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Honjo, Susumu; Manganini, Steven J.; Krishfield, Richard A.; Francois, Roger (2008). "Particulate organic carbon fluxes to the ocean interior and factors controlling the biological pump: A synthesis of global sediment trap programs since 1983".
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and incorporated in the coccosphere. The coccoliths are either dispersed following death and breakup of the coccosphere, or are shed continually by some species. They sink through the water column to form an important part of the deep-sea
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Because coccoliths are formed of low-Mg calcite, the most stable form of calcium carbonate, they are readily fossilised. They are found in sediments together with similar microfossils of uncertain affinities (nanoliths) from the
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Moheimani, N.R.; Webb, J.P.; Borowitzka, M.A. (2012), "Bioremediation and other potential applications of coccolithophorid algae: A review. . Bioremediation and other potential applications of coccolithophorid algae: A review",
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Poulton, Alex J.; Adey, Tim R.; Balch, William M.; Holligan, Patrick M. (2007). "Relating coccolithophore calcification rates to phytoplankton community dynamics: Regional differences and implications for carbon export".
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phytoplankton death in the oceans, and it has recently been shown that calcification can influence the interaction between a coccolithophore and its virus. The major predators of marine phytoplankton are
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was the first person to observe these forms in modern marine sediments and he gave them the name 'coccoliths' in a report published in 1858. Coccoliths are composed of calcium carbonate as the mineral
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to recent. They are widely used as biostratigraphic markers and as paleoclimatic proxies. Stable oxygen and carbon isotope data from coccoliths is used to reconstruct estimates of oceanic CO
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Although coccoliths are remarkably elaborate structures whose formation is a complex product of cellular processes, their function is unclear. Hypotheses include defence against grazing by
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are able to selectively choose prey on the basis of its size or shape and through chemical signals  and may thus favor other prey that is available and not protected by coccoliths.
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Johns, Christopher T.; Grubb, Austin R.; Nissimov, Jozef I.; Natale, Frank; Knapp, Viki; Mui, Alwin; Fredricks, Helen F.; Van Mooy, Benjamin A. S.; Bidle, Kay D. (2019).
78:, which are about 2–25 micrometres across. Coccolithophores are an important group of about 200 marine phytoplankton species  which cover themselves with a 2183:
Mayers, K.M.J.; Poulton, A.J.; Daniels, C.J.; Wells, S.R.; Woodward, E.M.S.; Tarran, G.A.; Widdicombe, C.E.; Mayor, D.J.; Atkinson, A.; Giering, S.L.C. (2019).
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Deep Sea Soundings in the North Atlantic Ocean between Ireland and Newfoundland, made in H.M.S. Cyclops, Lieut.-Commander Joseph Dayman, in June and July 1857
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Young, J. R. (1994). "Functions of coccoliths," in Coccolithophores, eds A. Winter and W. G. Siesser (Cambridge: Cambridge University Press), 63–82.
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Henderiks, Jorijntje (2008). "Coccolithophore size rules — Reconstructing ancient cell geometry and cellular calcite quota from fossil coccoliths".
917:. Heterococcoliths are formed of a radial array of elaborately shaped crystal units. Holococcoliths are formed of minute (~0.1 micrometre) calcite 559: 2654: 2533:
Bendif, El Mahdi; Probert, Ian; Díaz-Rosas, Francisco; Thomas, Daniela; van den Engh, Ger; Young, Jeremy R.; von Dassow, Peter (2016-05-24).
1350:"Increasing Costs Due to Ocean Acidification Drives Phytoplankton to Be More Heavily Calcified: Optimal Growth Strategy of Coccolithophores" 2681: 780: 2586:"Isotopic record of Pleistocene glacial/interglacial cycles in pelagic carbonates: Revisiting historical data from the Caribbean Sea" 2130:
Calbet, Albert; Landry, Michael R. (2004). "Phytoplankton growth, microzooplankton grazing, and carbon cycling in marine systems".
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RCC4033; (E) Reticulofenestra parvular RCC4034; (F) Reticulofenestra parvular RCC4035; (G) Reticulofenestra parvular RCC4036; (H)
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Bown, Paul R.; Lees, Jackie A.; Young, Jeremy R. (2004). "Calcareous nannoplankton evolution and diversity through time".
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precipitation of calcium carbonate during coccolith formation reduces the total alkalinity of seawater and releases CO
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Young, J. R. (1987). Possible Functional Interpretations of Coccolith Morphology. New York: Springer-Verlag, 305–313.
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Rost, Björn; Riebesell, Ulf (2004). "Coccolithophores and the biological pump: Responses to environmental changes".
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shell called a "coccosphere". They are ecologically and biogeochemically important but the reason why they
24: 2185:"Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)" 74:
Coccolithophores are spherical cells about 5–100 micrometres across, enclosed by calcareous plates called
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Coccolithophore cell surrounded by its shield of coccoliths. The coccolith-bearing cell is called the
1729:"Marine calcification as a source of carbon dioxide: Positive feedback of increasing atmospheric CO2" 1297:
Haunost, Mathias; Riebesell, Ulf; D'Amore, Francesco; Kelting, Ole; Bach, Lennart T. (30 June 2021).
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sediments. Their calcareous shell increases the sinking velocity of photosynthetically fixed CO
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Tillmann, Urban (2004). "Interactions between Planktonic Microalgae and Protozoan Grazers1".
1116: 871: 849:. When the coccolith is complete these vesicles fuse with the cell wall and the coccolith is 276: 2196: 2139: 1895: 1840: 1740: 1701: 1655: 1614: 1575: 1465: 1412: 1042: 308: 3476: 2627: 8: 3701: 2650:
Cocco Express - Coccolithophorids Expressed Sequence Tags (EST) & Microarray Database
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Brussaard, Corina P. D. (2004). "Viral Control of Phytoplankton Populations-a Review1".
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remains elusive. One key function may be that the coccosphere offers protection against
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concentrations in the geologic past. Coccoliths and related fossils are referred to as
763: 753: 604: 570: 487: 480: 391: 240: 2535:"Recent Reticulate Evolution in the Ecologically Dominant Lineage of Coccolithophores" 1058: 3747: 3706: 3059: 2772: 2640:- includes an illustrated guide to coccolith terminology and several image galleries. 2556: 2513: 2486: 2459: 2436: 2262: 2218: 2079: 2022: 1985: 1921: 1789: 1545: 1512: 1481: 1430: 1371: 1320: 1259: 1129: 768: 678: 347: 281: 206: 79: 56: 43: 2603: 2456:
Coccolithophores of the Eastern Mediterranean sea: A look into the marine microworld
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Malinverno, E; Dimiza, MD; Triantaphyllou, MV; Dermitzakis, MD; Corselli, C (2008).
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Irie, Takahiro; Bessho, Kazuhiro; Findlay, Helen S.; Calosi, Piero (2010-10-15).
594: 470: 384: 352: 124: 60:) and cover the cell surface arranged in the form of a spherical shell, called a 47: 2373: 2347: 1785: 1278: 1245: 90:
predation, which is one of the main causes of phytoplankton death in the ocean.
3681: 3522: 3432: 3418: 3219: 3075: 3023: 2880: 2799: 2646:- illustrated guide to the taxonomy of coccolithophores and other nannofossils. 2571: 2288:
Breckels, M. N.; Roberts, E. C.; Archer, S. D.; Malin, G.; Steinke, M. (2011).
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light; or in deep-dwelling species, to concentrate light for photosynthesis.
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Modified material was copied from this source, which is available under a
2050:"The mutual interplay between calcification and coccolithovirus infection" 1385:
Modified material was copied from this source, which is available under a
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Coccoliths are also classified depending on shape. Common shapes include:
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Hay, William W. (2004). "Carbonate fluxes and calcareous nannoplankton".
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Marine Phytoplankton: A Guide to Naked Flagellates and Coccolithophorids
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Hamm, Christian; Smetacek, Victor (2007). "Armor: Why, when, and How".
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Frankignoulle, Michel; Canon, Christine; Gattuso, Jean-Pierre (1994).
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Partial cross section of a coccolithophore with coccolith layer 
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SEM images correspond to coccolith drawings in the previous diagram
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Material was copied from this source, which is available under a
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Calcium carbonate scales covering some phytoplankton species
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Haunost, Mathias; Riebesell, Ulf; Bach, Lennart T. (2020).
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Deep Sea Research Part II: Topical Studies in Oceanography
1000:– rim composed of two plates stacked on top of one another 2182: 1829:"Environmental controls on coccolithophore calcification" 1726: 2689: 2047: 1880: 1564: 988:– basket or cup-shaped with a high rim, opening distally 2637: 2634:, containing essays on blooms, coccolith function, etc. 1347: 1228: 1176: 994:– pentagonal shape composed of five four-sided crystals 155:. Thus, coccolithophores play an important role in the 2572:
Creative Commons Attribution 4.0 International License
2405:"Atlas of Northwestern Mediterranean Coccolithophores" 1387:
Creative Commons Attribution 4.0 International License
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Creative Commons Attribution 4.0 International License
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in the surface ocean  and ~50% to pelagic CaCO
2231:Young, J. R. (1994) "Functions of coccoliths". In: 1012:– a single plate with a club-shaped central process 1257: 1087:is associated with a SEM image in the next diagram 3316: 2472: 2403:Fortuño, José Manuel; Cros, Lluïsa (2002-03-30). 3855: 1360:(10). Public Library of Science (PLoS): e13436. 2505: 1876: 1874: 131:. Today, coccolithophores contribute ~1–10% to 123:Coccolithophores have been an integral part of 2485:. Cambridge University Press. pp. 54–58. 1258:Young, J. R.; Geisen, M.; Probert, I. (2005). 878:times), in which they were first described by 3277: 2675: 2332:. London: British Admiralty. pp. 63–68 . 1826: 1775: 1600: 1498: 1292: 1290: 1288: 1018:– rhombohedral, with parallel lines in center 822: 2638:International Nannoplankton Association site 2129: 1971: 1871: 1601:Broecker, Wallace; Clark, Elizabeth (2009). 1222: 193: 2402: 1642:Klaas, Christine; Archer, David E. (2002). 1641: 946:– basket-shaped with openings near the base 115: 2682: 2668: 2526: 2352:Quarterly Journal of Microscopical Science 1285: 829: 815: 3184: 2550: 2499: 2430: 2420: 2305: 2208: 2159: 2114: 2073: 2004: 1974:Evolution of Primary Producers in the Sea 1956: 1915: 1862: 1852: 1752: 1667: 1626: 1455: 1424: 1365: 1314: 841:Coccoliths are formed within the cell in 2479:Amos Winter; William G. Siesser (2006). 2244: 923: 909:There are two main types of coccoliths, 163:and the oceanic uptake of atmospheric CO 18: 2583: 1077:Coccolith structures of representative 3856: 2378:Annals and Magazine of Natural History 2342: 2320: 2247:The Journal of Eukaryotic Microbiology 2007:The Journal of Eukaryotic Microbiology 1938: 1398: 147:organic matter. At the same time, the 2663: 2368: 159:by influencing the efficiency of the 2613:– via Elsevier Science Direct. 2512:. Academic Press. pp. 161–165. 1052:consisting of overlapping placoliths 1531: 42:are individual plates or scales of 13: 2458:. 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(2012). 1380: 1329: 1182: 1094: 1070: 1057: 1041: 1026: 887: 866:and are the main constituent of 858:(depending on the water depth). 796: 795: 213: 107: 95: 2604:10.1016/j.quascirev.2016.02.003 2577: 2447: 2396: 2362: 2336: 2314: 2281: 2238: 2225: 2176: 2123: 2090: 2041: 1998: 1932: 1820: 1811: 1802: 1769: 1720: 1684: 1635: 1594: 560:microbial calcite precipitation 3429:Membranous/thalloid/foliaceous 1833:Marine Ecology Progress Series 1558: 1525: 1492: 1478:10.1016/j.marmicro.2008.01.005 1449: 1392: 1341: 1251: 958:– horseshoe or wishbone shaped 1: 3399:Tetrasporal/capsal/palmelloid 1464:(1–2). Elsevier BV: 143–154. 1215: 520:marine biogenic calcification 2294:Journal of Plankton Research 2210:10.1016/j.pocean.2018.02.024 1714:10.1016/j.pocean.2007.11.003 1648:Global Biogeochemical Cycles 1542:10.1007/978-3-662-06278-4_19 1509:10.1007/978-3-662-06278-4_18 1399:Aloisi, G. 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Frontiers Media SA. 1908:10.1126/sciadv.1501822 1309:. Frontiers Media SA. 1199:calcareous nanofossils 1112:Gephyrocapsa muellerae 952:– disc- or bowl-shaped 931: 897:Pleurochrysis carterae 714:Magnetotactic bacteria 539:oolitic aragonite sand 397:scaly-foot snail shell 161:biological carbon pump 127:communities since the 36: 2773:Fungus-like organisms 2307:10.1093/plankt/fbq114 1117:Gephyrocapsa oceanica 1035:Helicosphaera carteri 927: 872:white cliffs of Dover 870:deposits such as the 22: 3488:Simple cell membrane 3424:Pseudoparenchymatous 3412:Filamentous/trichal/ 3396:Colonial flagellated 3329:Collar of microvilli 2630:- site dedicated to 2370:Sorby, Henry Clifton 2344:Huxley, Thomas Henry 2322:Huxley, Thomas Henry 1976:. pp. 311–332. 1669:10.1029/2001GB001765 1628:10.1029/2009PA001731 1536:. pp. 509–528. 1503:. pp. 481–508. 3702:Contractile vacuole 3653:Meiosis in protists 3634:Mitosis in protists 3612:Multinucleate cells 2834:Flagellar apparatus 2783:Ambiregnal protists 2725:Sarcodina/Amoeboids 2201:2019PrOce.17701928M 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2465:978-960-411-660-7 1854:10.3354/meps09993 1795:978-3-642-06016-8 1551:978-3-642-06016-8 1518:978-3-642-06016-8 1267:Micropaleontology 1163:; maintenance of 1139:Emiliania huxleyi 1135:Emiliania huxleyi 1130:Emiliania huxleyi 1050:Emiliania huxleyi 845:derived from the 839: 838: 769:permineralization 754:Mineral nutrients 679:Mineral evolution 348:foraminifera test 207:Biomineralization 80:calcium carbonate 57:Emiliania huxleyi 44:calcium carbonate 3871: 3722:Mastigont system 3675: 3600: 3577: 3566:Gliding motility 3544: 3481: 3426:/plektenchymatic 3382:Amoeboflagellate 3355: 3346: 3345: 3322: 3314: 3313: 3283: 3275: 3274: 3249: 3223: 3215: 3214: 3189: 3182: 3181: 3146: 3110: 3079: 3053: 3048: 3047: 3017: 2996: 2975: 2970: 2969: 2939: 2913: 2905: 2904: 2874: 2848: 2812: 2805: 2804: 2796: 2795: 2684: 2677: 2670: 2661: 2660: 2628:The EHUX website 2615: 2614: 2612: 2610: 2581: 2575: 2569: 2564: 2554: 2530: 2524: 2523: 2503: 2497: 2496: 2482:Coccolithophores 2476: 2470: 2469: 2451: 2445: 2444: 2434: 2424: 2400: 2394: 2393: 2366: 2360: 2359: 2340: 2334: 2333: 2318: 2312: 2311: 2309: 2285: 2279: 2278: 2242: 2236: 2233:Coccolithophores 2229: 2223: 2222: 2212: 2180: 2174: 2173: 2163: 2127: 2121: 2120: 2118: 2094: 2088: 2087: 2077: 2060:(6): 1896–1915. 2045: 2039: 2038: 2002: 1996: 1995: 1969: 1963: 1962: 1960: 1936: 1930: 1929: 1919: 1888:Science Advances 1878: 1869: 1868: 1866: 1856: 1824: 1818: 1815: 1809: 1806: 1800: 1799: 1778:Coccolithophores 1773: 1767: 1766: 1756: 1724: 1718: 1717: 1688: 1682: 1681: 1671: 1639: 1633: 1632: 1630: 1607:Paleoceanography 1598: 1592: 1591: 1574:(5–7): 538–557. 1562: 1556: 1555: 1534:Coccolithophores 1529: 1523: 1522: 1501:Coccolithophores 1496: 1490: 1489: 1453: 1447: 1446: 1428: 1396: 1390: 1384: 1379: 1369: 1345: 1339: 1333: 1328: 1318: 1294: 1283: 1282: 1264: 1255: 1249: 1248: 1226: 1155:or infection by 1123:Reticulofenestra 1098: 1079:Noelaerhabdaceae 1074: 1061: 1045: 1033:Helicoliths of 1030: 911:heterococcoliths 891: 831: 824: 817: 804: 799: 798: 719:Magnetoreception 699:Ballast minerals 294:Cephalopod shell 289:Brachiopod shell 236:Remineralisation 217: 198: 197: 173:microzooplankton 111: 99: 88:microzooplankton 48:coccolithophores 3879: 3878: 3874: 3873: 3872: 3870: 3869: 3868: 3854: 3853: 3852: 3847: 3740: 3726: 3673: 3668: 3598: 3593: 3575: 3570: 3542: 3537: 3478: 3475: 3470: 3352: 3349: 3333: 3317: 3299: 3278: 3260: 3244: 3239: 3218: 3200: 3185: 3167: 3141: 3136: 3105: 3100: 3076:Dinoflagellates 3074: 3069: 3051: 3033: 3012: 3007: 2991: 2986: 2973: 2955: 2934: 2929: 2908: 2890: 2869: 2864: 2843: 2838: 2808: 2787: 2708:classifications 2707: 2701: 2688: 2624: 2619: 2618: 2608: 2606: 2582: 2578: 2531: 2527: 2520: 2504: 2500: 2493: 2477: 2473: 2466: 2452: 2448: 2409:Scientia Marina 2401: 2397: 2384:(45): 193–200. 2367: 2363: 2341: 2337: 2319: 2315: 2286: 2282: 2243: 2239: 2230: 2226: 2181: 2177: 2128: 2124: 2095: 2091: 2046: 2042: 2003: 1999: 1992: 1970: 1966: 1937: 1933: 1894:(7): e1501822. 1879: 1872: 1825: 1821: 1816: 1812: 1807: 1803: 1796: 1774: 1770: 1725: 1721: 1689: 1685: 1640: 1636: 1599: 1595: 1563: 1559: 1552: 1530: 1526: 1519: 1497: 1493: 1454: 1450: 1397: 1393: 1346: 1342: 1295: 1286: 1262: 1256: 1252: 1227: 1223: 1218: 1212: 1195: 1185: 1149: 1142: 1102: 1099: 1088: 1082: 1075: 1066: 1065: 1064: 1063: 1062: 1054: 1053: 1048:Coccosphere of 1046: 1038: 1037: 1031: 937: 930: 907: 900: 892: 835: 794: 787: 786: 785: 673: 665: 664: 663: 619: 611: 610: 609: 595:biogenic silica 589: 579: 578: 577: 562: 550: 529: 509: 499: 498: 497: 465: 457: 456: 455: 435: 420: 419: 418: 385:gastropod shell 353:testate amoebae 343:diatom frustule 268: 257: 256: 255: 225: 196: 181:dinoflagellates 166: 154: 142: 138: 125:marine plankton 119: 112: 103: 100: 72: 50:(single-celled 35: 17: 12: 11: 5: 3877: 3867: 3866: 3849: 3848: 3846: 3845: 3840: 3835: 3834: 3833: 3832: 3831: 3826: 3816: 3811: 3806: 3796: 3795: 3794: 3783: 3782: 3781: 3776: 3762: 3761: 3760: 3755: 3744: 3742: 3736: 3735: 3732: 3731: 3728: 3727: 3725: 3724: 3719: 3714: 3709: 3704: 3699: 3694: 3689: 3684: 3678: 3676: 3670: 3669: 3667: 3666: 3665: 3664: 3661: 3658: 3650: 3649: 3648: 3645: 3642: 3639: 3631: 3626: 3621: 3620: 3619: 3609: 3603: 3601: 3595: 3594: 3592: 3591: 3586: 3580: 3578: 3572: 3571: 3569: 3568: 3563: 3558: 3553: 3547: 3545: 3539: 3538: 3536: 3535: 3530: 3525: 3520: 3515: 3510: 3505: 3500: 3495: 3490: 3484: 3482: 3472: 3471: 3469: 3468: 3455: 3454: 3453: 3452: 3451: 3450:Siphonocladous 3448: 3440: 3433:Multinucleated 3430: 3427: 3421: 3419:Parenchymatous 3416: 3410: 3409: 3408: 3403: 3400: 3397: 3386: 3385: 3384: 3379: 3374: 3369: 3358: 3356: 3343: 3339: 3338: 3335: 3334: 3332: 3331: 3325: 3323: 3311: 3305: 3304: 3301: 3300: 3298: 3297: 3292: 3286: 3284: 3272: 3266: 3265: 3262: 3261: 3259: 3258: 3252: 3250: 3241: 3240: 3238: 3237: 3232: 3226: 3224: 3220:Kinetoplastids 3212: 3206: 3205: 3202: 3201: 3199: 3198: 3192: 3190: 3179: 3173: 3172: 3169: 3168: 3166: 3165: 3160: 3155: 3149: 3147: 3138: 3137: 3135: 3134: 3129: 3124: 3119: 3113: 3111: 3102: 3101: 3099: 3098: 3093: 3088: 3082: 3080: 3071: 3070: 3068: 3067: 3062: 3056: 3054: 3045: 3039: 3038: 3035: 3034: 3032: 3031: 3026: 3020: 3018: 3009: 3008: 3006: 3005: 2999: 2997: 2988: 2987: 2985: 2984: 2978: 2976: 2967: 2961: 2960: 2957: 2956: 2954: 2953: 2948: 2942: 2940: 2931: 2930: 2928: 2927: 2922: 2916: 2914: 2902: 2896: 2895: 2892: 2891: 2889: 2888: 2886:Phycobilisomes 2883: 2881:Pit connection 2877: 2875: 2866: 2865: 2863: 2862: 2860:Phycobilisomes 2857: 2851: 2849: 2840: 2839: 2837: 2836: 2831: 2826: 2815: 2813: 2810:Chloroplastida 2802: 2800:Archaeplastida 2793: 2789: 2788: 2786: 2785: 2780: 2775: 2770: 2765: 2760: 2755: 2754: 2753: 2748: 2739: 2738: 2737: 2732: 2722: 2711: 2709: 2703: 2702: 2687: 2686: 2679: 2672: 2664: 2658: 2657: 2652: 2647: 2641: 2635: 2623: 2622:External links 2620: 2617: 2616: 2576: 2525: 2518: 2498: 2491: 2471: 2464: 2446: 2395: 2361: 2354:. New series. 2335: 2313: 2300:(4): 629–639. 2280: 2253:(2): 156–168. 2237: 2224: 2175: 2122: 2089: 2040: 2013:(2): 125–138. 1997: 1990: 1964: 1931: 1870: 1819: 1810: 1801: 1794: 1768: 1739:(2): 458–462. 1719: 1700:(3): 217–285. 1683: 1634: 1593: 1557: 1550: 1524: 1517: 1491: 1448: 1405:Biogeosciences 1391: 1340: 1284: 1273:(4): 267–288. 1250: 1240:(2): 120–133, 1233:Algal Research 1220: 1219: 1217: 1214: 1193: 1190:Upper Triassic 1184: 1181: 1173:photosynthesis 1169:carbon dioxide 1148: 1145: 1144: 1143: 1100: 1093: 1090: 1089: 1076: 1069: 1056: 1055: 1047: 1040: 1039: 1032: 1025: 1024: 1023: 1022: 1021: 1020: 1019: 1013: 1007: 1001: 995: 989: 983: 977: 971: 965: 959: 953: 947: 936: 933: 928: 915:holococcoliths 906: 903: 902: 901: 893: 886: 874:(deposited in 837: 836: 834: 833: 826: 819: 811: 808: 807: 806: 805: 789: 788: 784: 783: 778: 777: 776: 771: 761: 756: 751: 746: 739: 734: 726: 721: 716: 711: 706: 701: 696: 695: 694: 692:immobilization 689: 687:mineralization 681: 675: 674: 671: 670: 667: 666: 662: 661: 656: 655: 654: 644: 643: 642: 637: 627: 621: 620: 617: 616: 613: 612: 608: 607: 602: 600:siliceous ooze 597: 591: 590: 587:Silicification 585: 584: 581: 580: 576: 575: 574: 573: 568: 563: 551: 549: 548: 547: 546: 541: 530: 528: 527: 522: 517: 511: 510: 505: 504: 501: 500: 496: 495: 490: 485: 484: 483: 473: 467: 466: 463: 462: 459: 458: 454: 453: 448: 443: 437: 436: 426: 425: 422: 421: 417: 416: 411: 409:Sponge spicule 406: 405: 404: 402:estuary shells 399: 394: 389: 388: 387: 382: 377: 367: 357: 356: 355: 350: 345: 340: 335: 325: 324: 323: 313: 312: 311: 306: 301: 291: 286: 285: 284: 279: 270: 269: 263: 262: 259: 258: 254: 253: 248: 243: 238: 233: 227: 226: 223: 222: 219: 218: 210: 209: 203: 202: 195: 192: 164: 152: 140: 136: 121: 120: 113: 106: 104: 101: 94: 71: 68: 23: 15: 9: 6: 4: 3: 2: 3876: 3865: 3862: 3861: 3859: 3844: 3841: 3839: 3836: 3830: 3827: 3825: 3822: 3821: 3820: 3817: 3815: 3812: 3810: 3807: 3805: 3802: 3801: 3800: 3797: 3793: 3790: 3789: 3788: 3784: 3780: 3779:Fertilization 3777: 3775: 3772: 3771: 3770: 3766: 3763: 3759: 3756: 3754: 3751: 3750: 3749: 3746: 3745: 3743: 3737: 3723: 3720: 3718: 3715: 3713: 3710: 3708: 3705: 3703: 3700: 3698: 3695: 3693: 3690: 3688: 3685: 3683: 3680: 3679: 3677: 3671: 3662: 3659: 3656: 3655: 3654: 3651: 3647:Pleuromitosis 3646: 3643: 3640: 3637: 3636: 3635: 3632: 3630: 3627: 3625: 3622: 3618: 3615: 3614: 3613: 3610: 3608: 3605: 3604: 3602: 3596: 3590: 3587: 3585: 3584:Hydrogenosome 3582: 3581: 3579: 3573: 3567: 3564: 3562: 3559: 3557: 3554: 3552: 3549: 3548: 3546: 3540: 3534: 3531: 3529: 3526: 3524: 3521: 3519: 3516: 3514: 3511: 3509: 3506: 3504: 3501: 3499: 3496: 3494: 3491: 3489: 3486: 3485: 3483: 3480: 3473: 3466: 3462: 3459: 3458:Multicellular 3456: 3449: 3446: 3445: 3444: 3441: 3439: 3436: 3435: 3434: 3431: 3428: 3425: 3422: 3420: 3417: 3415: 3411: 3407: 3404: 3401: 3398: 3395: 3394: 3393: 3390: 3387: 3383: 3380: 3378: 3375: 3373: 3370: 3368: 3365: 3364: 3363: 3360: 3359: 3357: 3354: 3347: 3344: 3340: 3330: 3327: 3326: 3324: 3320: 3315: 3312: 3310: 3306: 3296: 3293: 3291: 3288: 3287: 3285: 3281: 3280:Dictyostelids 3276: 3273: 3271: 3267: 3257: 3254: 3253: 3251: 3247: 3242: 3236: 3233: 3231: 3228: 3227: 3225: 3221: 3216: 3213: 3211: 3207: 3197: 3194: 3193: 3191: 3188: 3183: 3180: 3178: 3174: 3164: 3161: 3159: 3156: 3154: 3151: 3150: 3148: 3144: 3143:Apicomplexans 3139: 3133: 3130: 3128: 3125: 3123: 3120: 3118: 3115: 3114: 3112: 3108: 3103: 3097: 3094: 3092: 3089: 3087: 3084: 3083: 3081: 3077: 3072: 3066: 3063: 3061: 3058: 3057: 3055: 3049: 3046: 3044: 3040: 3030: 3027: 3025: 3022: 3021: 3019: 3015: 3010: 3004: 3001: 3000: 2998: 2994: 2989: 2983: 2980: 2979: 2977: 2971: 2968: 2966: 2965:Stramenopiles 2962: 2952: 2949: 2947: 2944: 2943: 2941: 2937: 2932: 2926: 2923: 2921: 2918: 2917: 2915: 2911: 2906: 2903: 2901: 2897: 2887: 2884: 2882: 2879: 2878: 2876: 2872: 2867: 2861: 2858: 2856: 2853: 2852: 2850: 2846: 2841: 2835: 2832: 2830: 2827: 2825: 2821: 2817: 2816: 2814: 2811: 2806: 2803: 2801: 2797: 2794: 2790: 2784: 2781: 2779: 2776: 2774: 2771: 2769: 2766: 2764: 2761: 2759: 2756: 2752: 2749: 2747: 2743: 2740: 2736: 2733: 2731: 2728: 2727: 2726: 2723: 2721: 2718: 2717: 2716: 2713: 2712: 2710: 2704: 2700: 2696: 2692: 2685: 2680: 2678: 2673: 2671: 2666: 2665: 2662: 2656: 2653: 2651: 2648: 2645: 2642: 2639: 2636: 2633: 2629: 2626: 2625: 2605: 2601: 2597: 2593: 2592: 2587: 2580: 2573: 2568: 2562: 2558: 2553: 2548: 2544: 2540: 2536: 2529: 2521: 2515: 2511: 2510: 2502: 2494: 2488: 2484: 2483: 2475: 2467: 2461: 2457: 2450: 2442: 2438: 2433: 2428: 2423: 2418: 2415:(S1): 1–182. 2414: 2410: 2406: 2399: 2391: 2387: 2383: 2379: 2375: 2371: 2365: 2357: 2353: 2349: 2345: 2339: 2331: 2327: 2323: 2317: 2308: 2303: 2299: 2295: 2291: 2284: 2276: 2272: 2268: 2264: 2260: 2256: 2252: 2248: 2241: 2234: 2228: 2220: 2216: 2211: 2206: 2202: 2198: 2194: 2190: 2186: 2179: 2171: 2167: 2162: 2157: 2153: 2149: 2145: 2141: 2137: 2133: 2126: 2117: 2112: 2108: 2104: 2100: 2093: 2085: 2081: 2076: 2071: 2067: 2063: 2059: 2055: 2051: 2044: 2036: 2032: 2028: 2024: 2020: 2016: 2012: 2008: 2001: 1993: 1991:9780123705181 1987: 1983: 1979: 1975: 1968: 1959: 1954: 1950: 1946: 1942: 1935: 1927: 1923: 1918: 1913: 1909: 1905: 1901: 1897: 1893: 1889: 1885: 1877: 1875: 1865: 1860: 1855: 1850: 1846: 1842: 1838: 1834: 1830: 1823: 1814: 1805: 1797: 1791: 1787: 1783: 1779: 1772: 1764: 1760: 1755: 1750: 1746: 1742: 1738: 1734: 1730: 1723: 1715: 1711: 1707: 1703: 1699: 1695: 1687: 1679: 1675: 1670: 1665: 1661: 1657: 1653: 1649: 1645: 1638: 1629: 1624: 1620: 1616: 1612: 1608: 1604: 1597: 1589: 1585: 1581: 1577: 1573: 1569: 1561: 1553: 1547: 1543: 1539: 1535: 1528: 1520: 1514: 1510: 1506: 1502: 1495: 1487: 1483: 1479: 1475: 1471: 1467: 1463: 1459: 1452: 1444: 1440: 1436: 1432: 1427: 1422: 1418: 1414: 1410: 1406: 1402: 1395: 1388: 1383: 1377: 1373: 1368: 1363: 1359: 1355: 1351: 1344: 1337: 1332: 1326: 1322: 1317: 1312: 1308: 1304: 1300: 1293: 1291: 1289: 1280: 1276: 1272: 1268: 1261: 1254: 1247: 1243: 1239: 1235: 1234: 1225: 1221: 1213: 1210: 1208: 1207: 1201: 1200: 1191: 1183:Fossil record 1180: 1178: 1174: 1170: 1167:; release of 1166: 1162: 1158: 1154: 1141:morphotype B. 1140: 1136: 1132: 1131: 1126: 1124: 1119: 1118: 1113: 1109: 1107: 1097: 1092: 1091: 1086: 1085:morphospecies 1080: 1073: 1068: 1067: 1060: 1051: 1044: 1036: 1029: 1017: 1014: 1011: 1008: 1005: 1002: 999: 996: 993: 990: 987: 984: 981: 978: 975: 972: 969: 966: 963: 960: 957: 954: 951: 948: 945: 942: 941: 940: 926: 922: 920: 916: 912: 899: 898: 890: 885: 884: 883: 881: 877: 873: 869: 865: 861: 860:Thomas Huxley 857: 852: 848: 844: 832: 827: 825: 820: 818: 813: 812: 810: 809: 803: 793: 792: 791: 790: 782: 779: 775: 772: 770: 767: 766: 765: 764:Fossilization 762: 760: 759:Microbial mat 757: 755: 752: 750: 747: 745: 744: 740: 738: 735: 733: 731: 727: 725: 722: 720: 717: 715: 712: 710: 707: 705: 704:Magnetofossil 702: 700: 697: 693: 690: 688: 685: 684: 682: 680: 677: 676: 669: 668: 660: 657: 653: 650: 649: 648: 645: 641: 638: 636: 633: 632: 631: 628: 626: 623: 622: 615: 614: 606: 603: 601: 598: 596: 593: 592: 588: 583: 582: 572: 569: 567: 564: 561: 558: 557: 556: 553: 552: 545: 544:aragonite sea 542: 540: 537: 536: 535: 532: 531: 526: 523: 521: 518: 516: 513: 512: 508: 507:Calcification 503: 502: 494: 491: 489: 486: 482: 479: 478: 477: 474: 472: 469: 468: 461: 460: 452: 449: 447: 444: 442: 439: 438: 433: 429: 428:Endoskeletons 424: 423: 415: 412: 410: 407: 403: 400: 398: 395: 393: 390: 386: 383: 381: 378: 376: 373: 372: 371: 370:mollusc shell 368: 366: 363: 362: 361: 358: 354: 351: 349: 346: 344: 341: 339: 336: 334: 331: 330: 329: 328:Protist shell 326: 322: 319: 318: 317: 314: 310: 307: 305: 302: 300: 299:cirrate shell 297: 296: 295: 292: 290: 287: 283: 280: 278: 275: 274: 272: 271: 266: 261: 260: 252: 249: 247: 244: 242: 239: 237: 234: 232: 229: 228: 221: 220: 216: 212: 211: 208: 205: 204: 200: 199: 191: 189: 186: 185:Heterotrophic 182: 178: 174: 168: 162: 158: 150: 146: 134: 130: 126: 117: 110: 105: 98: 93: 92: 91: 89: 85: 81: 77: 67: 65: 64: 59: 58: 53: 52:phytoplankton 49: 45: 41: 33: 31: 26: 21: 3799:Heterotrophy 3769:Reproduction 3644:Orthomitosis 3629:Heterokaryon 3576:Mitochondria 3477:Cell surface 3353:organization 3309:Opisthokonta 3132:Micronucleus 3127:Macronucleus 3029:Pneumatocyst 2945: 2845:Glaucophytes 2829:Phragmoplast 2768:Thallophytes 2695:Protistology 2691:Microbiology 2631: 2609:17 September 2607:. Retrieved 2595: 2589: 2579: 2542: 2538: 2528: 2508: 2501: 2481: 2474: 2455: 2449: 2412: 2408: 2398: 2381: 2377: 2364: 2355: 2351: 2338: 2329: 2326:"Appendix A" 2316: 2297: 2293: 2283: 2250: 2246: 2240: 2232: 2227: 2192: 2188: 2178: 2161:10261/134985 2138:(1): 51–57. 2135: 2131: 2125: 2106: 2102: 2092: 2057: 2053: 2043: 2010: 2006: 2000: 1973: 1967: 1948: 1944: 1934: 1891: 1887: 1864:10453/114799 1836: 1832: 1822: 1813: 1804: 1777: 1771: 1736: 1732: 1722: 1697: 1693: 1686: 1651: 1647: 1637: 1610: 1606: 1596: 1571: 1567: 1560: 1533: 1527: 1500: 1494: 1461: 1457: 1451: 1408: 1404: 1394: 1357: 1353: 1343: 1306: 1302: 1270: 1266: 1253: 1237: 1231: 1224: 1211: 1203: 1197: 1186: 1150: 1138: 1134: 1128: 1121: 1115: 1111: 1110:RCC4032 (B) 1106:Gephyrocapsa 1104: 1049: 1034: 1015: 1009: 1004:Prismatolith 1003: 997: 991: 985: 979: 973: 967: 961: 955: 949: 944:Calyptrolith 943: 938: 914: 910: 908: 895: 840: 774:petrifaction 741: 729: 724:Microfossils 471:Limpet teeth 451:Ossification 446:Bone mineral 380:chiton shell 337: 265:Exoskeletons 246:Biointerface 169: 122: 73: 61: 55: 39: 38: 28: 3829:Necrotrophy 3814:Saprotrophy 3804:Phagotrophy 3792:Phototrophy 3785:Nutrition: 3774:Life cycles 3765:Development 3739:Ecology and 3607:Nucleomorph 3561:Pseudopodia 3362:Unicellular 3246:Euglenoidea 3230:Kinetoplast 3014:Brown algae 2982:Mastigoneme 2920:Mastigoneme 2910:Cryptophyta 2820:green algae 2778:Slime molds 1839:: 137–166. 1763:2268/246251 1654:(4): 1116. 1204:calcareous 1153:zooplankton 919:rhombohedra 709:Magnetosome 652:phosphorite 618:Other forms 566:calcite sea 333:coccosphere 277:exoskeleton 116:coccosphere 63:coccosphere 30:Coccolithus 3843:Auxotrophy 3838:Mixotrophy 3819:Parasitism 3809:Osmotrophy 3787:Autotrophy 3741:physiology 3617:Plasmodium 3543:Locomotion 3479:structures 3443:Coenocytic 3187:Phytomyxea 3158:Apicoplast 3086:Dinokaryon 3065:Trichocyst 2936:Haptophyte 2824:Phycoplast 2792:Morphology 2763:Cryptogams 2432:10261/5441 2380:. Ser. 3. 2358:: 203–212. 2195:: 101928. 1216:References 1016:Scapholith 1010:Rhabdolith 986:Lopadolith 980:Helicolith 962:Cribrilith 956:Ceratolith 876:Cretaceous 851:exocytosed 847:golgi body 304:cuttlebone 273:Arthropod 145:ballasting 76:coccoliths 46:formed by 40:Coccoliths 3824:Biotrophy 3697:Extrusome 3687:Cytostome 3551:Flagellum 3508:Cell wall 3467:/histonal 3447:Siphonous 3438:Syncytial 3406:Coenobial 3402:Sarcinoid 3351:Levels of 3290:Macrocyst 3270:Amoebozoa 3235:Glycosome 3163:Microneme 3043:Alveolata 2951:Haptonema 2946:Coccolith 2925:Periplast 2871:Red algae 2855:Cyanelles 2742:Infusoria 2598:: 69–78. 2561:1664-302X 2441:1886-8134 2219:135347218 1486:0377-8398 1435:1726-4189 1376:1932-6203 1325:2296-7745 1108:ericsonii 998:Placolith 992:Pentalith 974:Discolith 968:Cyrtolith 950:Caneolith 882:in 1861. 856:sediments 730:engrailed 647:Phosphate 640:oil shale 534:Aragonite 338:coccolith 32:pelagicus 3858:Category 3717:Axostyle 3712:Pyrenoid 3692:Fimbriae 3624:Dikaryon 3589:Mitosome 3518:Skeleton 3503:Frustule 3493:Mucilage 3465:tissular 3389:Colonial 3372:Amoeboid 3367:Monadoid 3295:Sorocarp 3210:Excavate 3177:Rhizaria 3107:Ciliates 3091:Dinocyst 3052:General: 3003:Frustule 2974:General: 2900:Hacrobia 2751:Sporozoa 2746:Ciliates 2735:Heliozoa 2715:Protozoa 2699:Protists 2644:Nannotax 2372:(1861). 2346:(1868). 2324:(1858). 2275:36526359 2267:15134250 2170:22995996 2084:30043404 2035:21017882 2027:15134247 1926:27453937 1678:34159028 1354:PLOS ONE 1165:buoyancy 1157:bacteria 1147:Function 1125:parvular 843:vesicles 802:Category 683:In soil 635:alginite 625:Bone bed 360:Seashell 267:(shells) 188:protists 177:ciliates 149:biogenic 129:Jurassic 70:Overview 54:such as 3660:Zygotic 3657:Gametic 3599:Nucleus 3377:Coccoid 3342:General 3153:Rhoptry 3060:Alveoli 2993:Diatoms 2730:Testate 2197:Bibcode 2140:Bibcode 2075:7379532 1917:4956192 1896:Bibcode 1841:Bibcode 1741:Bibcode 1702:Bibcode 1656:Bibcode 1615:Bibcode 1576:Bibcode 1466:Bibcode 1443:6227548 1413:Bibcode 1161:viruses 864:calcite 672:Related 630:Kerogen 555:Calcite 476:Otolith 309:gladius 282:cuticle 251:Biofilm 224:General 84:calcify 3663:Sporic 3641:Closed 3556:Cilium 3513:Lorica 3414:hyphal 3122:Cirrus 3117:Cilium 3024:Lamina 2706:Former 2559:  2516:  2489:  2462:  2439:  2273:  2265:  2217:  2168:  2082:  2072:  2033:  2025:  1988:  1924:  1914:  1792:  1676:  1548:  1515:  1484:  1441:  1433:  1374:  1323:  800:  659:Pyrena 316:Lorica 3674:Other 3528:Theca 3498:Scale 3096:Theca 2758:Algae 2271:S2CID 2215:S2CID 2166:S2CID 2031:S2CID 1674:S2CID 1613:(3). 1439:S2CID 1263:(PDF) 1083:Each 935:Shape 905:Types 868:chalk 737:Druse 432:bones 375:nacre 175:like 3682:Cyst 3638:Open 3523:Test 3461:s.s. 3392:s.s. 2611:2024 2557:ISSN 2514:ISBN 2487:ISBN 2460:ISBN 2437:ISSN 2263:PMID 2080:PMID 2023:PMID 1986:ISBN 1922:PMID 1790:ISBN 1546:ISBN 1513:ISBN 1482:ISSN 1431:ISSN 1372:ISSN 1321:ISSN 1171:for 1120:(D) 1114:(C) 1103:(A) 913:and 732:gene 493:Tusk 414:Test 179:and 2822:": 2600:doi 2596:137 2547:doi 2427:hdl 2417:doi 2386:doi 2302:doi 2255:doi 2205:doi 2193:177 2156:hdl 2148:doi 2111:doi 2070:PMC 2062:doi 2015:doi 1978:doi 1953:doi 1912:PMC 1904:doi 1859:hdl 1849:doi 1837:470 1782:doi 1759:hdl 1749:doi 1710:doi 1664:doi 1623:doi 1584:doi 1538:doi 1505:doi 1474:doi 1421:doi 1362:doi 1311:doi 1275:doi 1242:doi 1202:or 1159:or 27:of 3860:: 2697:: 2693:: 2594:. 2588:. 2555:. 2541:. 2537:. 2435:. 2425:. 2413:66 2411:. 2407:. 2376:. 2350:. 2328:. 2298:33 2296:. 2292:. 2269:. 2261:. 2251:51 2249:. 2213:. 2203:. 2191:. 2187:. 2164:. 2154:. 2146:. 2136:49 2134:. 2109:. 2105:. 2101:. 2078:. 2068:. 2058:21 2056:. 2052:. 2029:. 2021:. 2011:51 2009:. 1984:. 1951:. 1947:. 1943:. 1920:. 1910:. 1902:. 1890:. 1886:. 1873:^ 1857:. 1847:. 1835:. 1831:. 1788:. 1757:. 1747:. 1737:39 1735:. 1731:. 1708:. 1698:76 1696:. 1672:. 1662:. 1652:16 1650:. 1646:. 1621:. 1611:24 1609:. 1605:. 1582:. 1572:54 1570:. 1544:. 1511:. 1480:. 1472:. 1462:67 1460:. 1437:. 1429:. 1419:. 1409:12 1407:. 1403:. 1370:. 1356:. 1352:. 1319:. 1305:. 1301:. 1287:^ 1271:51 1269:. 1265:. 1236:, 1209:. 1177:UV 167:. 66:. 3767:/ 3463:/ 3321:: 3282:: 3248:: 3222:: 3145:: 3109:: 3078:: 3016:: 2995:: 2938:: 2912:: 2873:: 2847:: 2818:" 2744:/ 2683:e 2676:t 2669:v 2602:: 2574:. 2563:. 2549:: 2543:7 2522:. 2495:. 2468:. 2443:. 2429:: 2419:: 2392:. 2388:: 2382:8 2356:8 2310:. 2304:: 2277:. 2257:: 2221:. 2207:: 2199:: 2172:. 2158:: 2150:: 2142:: 2119:. 2113:: 2107:7 2086:. 2064:: 2037:. 2017:: 1994:. 1980:: 1961:. 1955:: 1949:6 1928:. 1906:: 1898:: 1892:2 1867:. 1861:: 1851:: 1843:: 1798:. 1784:: 1765:. 1761:: 1751:: 1743:: 1716:. 1712:: 1704:: 1680:. 1666:: 1658:: 1631:. 1625:: 1617:: 1590:. 1586:: 1578:: 1554:. 1540:: 1521:. 1507:: 1488:. 1476:: 1468:: 1445:. 1423:: 1415:: 1389:. 1378:. 1364:: 1358:5 1338:. 1327:. 1313:: 1307:8 1281:. 1277:: 1244:: 1238:1 1194:2 1081:. 830:e 823:t 816:v 434:) 430:( 165:2 153:2 141:2 137:3 118:.

Index


Scanning electron micrograph
Coccolithus
calcium carbonate
coccolithophores
phytoplankton
Emiliania huxleyi
coccosphere
coccoliths
calcium carbonate
calcify
microzooplankton
Partial cross section of a coccolithophore with coccolith layer
Coccolithophore cell surrounded by its shield of coccoliths. The coccolith-bearing cell is called the coccosphere.
coccosphere
marine plankton
Jurassic
primary production
ballasting
biogenic
marine carbon cycle
biological carbon pump
microzooplankton
ciliates
dinoflagellates
Heterotrophic
protists
Biomineralization

Mineralized tissues

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