From Wikipedia, the free encyclopedia
Jump to navigation Jump to search

Dinocysts or dinoflagellate cysts are typically 15 to 100 µm in diameter and produced by around 15–20%[citation needed]of living dinoflagellates as a dormant, zygotic stage of their lifecycle, which can accumulate in the sediments as microfossils. Organic-walled dinocysts are often resistant and made out of dinosporin. There are also calcareous dinoflagellate cysts and siliceous dinoflagellate cysts.


Dinocyst drawn by Ehrenberg in 1837

The first person to recognize fossil dinoflagellates was Christian Gottfried Ehrenberg, who reported his discovery in a paper presented to the Berlin Academy of Sciences in July 1836. He had observed clearly tabulate dinoflagellates in thin flakes of Cretaceous flint and considered those dinoflagellates to have been silicified. Along with them, and of comparable size, were spheroidal to ovoidal bodies bearing an array of spines or tubes of variable character. Ehrenberg interpreted these as being originally siliceous and thought them to be desmids (freshwater conjugating algae), placing them within his own Recent desmid genus Xanthidium. Though summaries of Ehrenberg's work appeared earlier, it was not published in full until 1837 or 1838; the date is uncertain.[1]

A first relation between dinoflagellate thecae and cysts was made through morphological comparison of both by Bill Evitt and Susan E. Davidson.[2] Further evidence came from detailed culture studies of dinoflagellate cysts by David Wall and Barrie Dale at Woods Hole Oceanographic Institution in the sixties.[3][4]

Types of cysts[edit]

Ontologically, the term cyst can apply to (1) a temporary resting state (pellicle, temporary or ecdysal cyst), (2) a dormant zygote (resting cysts or hypnozygotes) or (3) a coccoid condition in which the cells are still photosynthetically active.[5] For example for this last special case, all cysts described from species of the order Phytodiniales (e.g. Cystodinium, Stylodinium, Hypnodinium, Tetradinium, Dinococcus, Gloeodinium), are coccoid stages.

Digestive cyst or digestion cysts denote pellicle cysts formed after feeding by phagocytosis as in Katodinium fungiforme .[6][7]

Division cysts refer to non-motile division stages wherein asexual reproduction takes place through division.[8] These are not pellicle or resting cysts since they are not dormant. Similarly, palmelloid or mucilage stages are not pellicle or resting cysts, but stages in which the monad loses its flagella and becomes enveloped in multilayered mucilage wherein division takes place.[9]


Dinoflagellate cysts described in the literature have been linked to a particular motile stage through morphological similarities and/or co-occurrence in the same population/culture or through the technique of establishing the so-called cyst-theca relation by incubation of the cysts.[10][4][11][12] Geologists use a cyst-based taxonomy, whilst biologists use a motile-stage based taxonomy. Therefore cysts can have different names than the corresponding motile stages. Living cysts can be easily isolated from the sediment using sodium polytungstate, a heavy liquid.[13] Another method, rarely used, uses a sucrose gradient.[14] Recent times have brought about the possibility to get molecular sequences from single cysts or single cells.[15][16][17] The proportion of cyst-forming species for marine dinoflagellates is between 15 and 20%[18] and for freshwater dinoflagellates 24%.[19] The tabulation of the Dinoflagellate is sometimes mirrored in the tabulation (previously called paratabulation) of the dinocyst, allowing species to be deduced from the cyst.[20] It has previously been suggested that morphological characters from the cyst stage may be phylogenetically important in marine species[21] and this may to an even greater extent be the case for freshwater dinoflagellates,[22] confirmed by new observations[23][24] and recently reviewed.[19] Several books document general cyst taxonomy.[20][25] There are few guides for determination of marine Quaternary dinocysts.[26][27] Many new species are still being described for the Neogene,[28] which covers the Miocene,[29][30] the Pliocene[31][32][33][34] and the Quaternary, which covers the Pleistocene[35] and recent.[36][37][38]


Quaternary dinocysts are typically between 15 and 100 µm in diameter.[39] One of the smallest recent cysts is the cyst of Pentapharsodinium dalei, which can be as small as 19 µm in length.[40] One of the largest recent cysts is the cyst of Protoperidinium latissimum, which can be as large as 100 µm in length.[4]


The walls of organic-walled dinocysts are composed of the resistant biopolymer called dinosporin.[41] This organic compound has similarities to sporopollenin, but is unique to dinoflagellates.

In addition to organic-walled cysts, there are also calcareous dinoflagellate cysts and siliceous dinoflagellate cysts.

Morphological terms[edit]

In pure morphological terms, a dinocyst can be described as the body formed by the cyst wall, as well as the space it encloses and all the spaces within it.[42] Cysts may develop their wall immediately within the theca, and such cysts are called proximate. Alternatively, the cyst may comprise a more or less spherical central body with processes or crests, and such cysts are termed chorate or proximochorate. Cysts may have a single-layered wall (autophragm), a two-layered wall (comprising an outer periphragm and an inner endophragm) or a three-layered wall (ectophragm, periphragm and endophragm if the outer wall is structurally supported, or otherwise periphragm, mesophragm and endophragm). Cysts with two or more wall layers that define a cavity are termed cavate. Excystment usually results in loss of part of, or an opening in, the cyst wall, termed archeopyle, the shape and position of which may indicate the position and/or shape of one or more thecal plates.[20]

Transmission electron microscopy (TEM) studies (e.g.[43]) suggest that endophragm and periphragm are not morphologically separable. Therefore the use of the terms pedium and luxuria are suggested instead.[44] Within the cyst wall, a thick cellulose-like layer called the endospore is present which is birefringent under crossed nichols.[45] Cysts may be identified using the overal body shape but more often based on the characteristic furrows housing the flagella (cingulum and sulcus) or details of the patterns of plates covering many motiles (thecal tabulation). The one distinctive feature common to all cysts is the excystment opening (archaeopyle) through which the emerging new motile stage exits. In many cases this reflects a recognizable part of the tabulation (one or more plates). However, one large group of dinoflagellates (athecate - or naked dinoflagellates) do not have thecal plates and therefore produce cysts lacking all forms of reflected tabulation.[46]

Cyst ultrastructure[edit]

There have been very few ultrastructural studies of marine cysts with TEM, except for early on Hystrichosphaea bentorii, on Hystrichosphaeridium, Impletosphaeridium, Lingulodinium machaerophorum and Operculodinium centrocarpum and Bitectatodinium tepikiense[43][47][48] and more recent work on Lingulodinium machaerophorum[49] and Alexandrium.[50]

Some freshwater cysts have been investigated with TEM, such as Ceratium hirundinella.[51]

Relation to life cycle[edit]

Resting cysts are traditionally associated with the sexual cycle of dinoflagellates.[52] Induced by particular triggers such as changes in temperature, nutrients,[53] etc., dinoflagellates undergo gamete formation. The gametes fuse to form the planozygote and undergo encystment: they form cysts within the thecae of the planozygote. These rapidly sink to the sediment. Many species may spend longer periods resting in the sediment than active in the water column.[54] Resting stages also constitute a reservoir of genetic diversity, which increases the survival potential of the populations.[55] Thus, dinoflagellate cysts have great ecological importance and act as "seed banks", comparable to those found in terrestrial ecosystems. The encysted forms may remain viable for up to 100 years.[56] Sediment can be stored with live Lingulodinium cysts for at least 18 months.[57] Cysts often need triggers to germinate ('excyst'), such as changes in temperature, nutrients, etc. Some cysts, such as Scrippsiella acuminata, require light to germinate.[58]

Distribution and ecology of organic-walled dinocysts[edit]

Dinocyst distribution is mainly studied through studies of surface sediments.[59] Many studies are regional, such as the Iberian Margin[60] the North Sea,[61] Kiel bight,[62] Celtic Sea,[63] Norwegian Sea,[64] around Iceland,[65] the Southeast Pacific,[66] the Arctic,[67][68] Equatorial Atlantic,[69] South and Equatorial Atlantic,[70] off West Africa,[71] the Southern Ocean,[72] Benguela upwelling,[73] in the Mediterranean Sea,[74] Caspian Sea,[75] British Columbia,[76] The Northeastern Pacific,[77] Florida,[78] Mexico[79] and Barends Sea.[80]

Such surface sediment studies show that dinoflagellate cyst distribution is controlled by ranges of temperature, salinity and nutrients.[81] This often poses biogeographical boundaries, more particularly temperature.[82] Some species can be clearly related to cold waters.[83] Recent molecular work has shown the presence of such cold-water indicator, a life-stage of Islandinium sp. in Canadian sea-ice for the first time.[84] Other species are thermophilic, such as the "living fossil" Dapsilidinium pastielsii currently found in the Indo-Pacific Warm Pool only.[85]

Eutrophication can also be reflected in dinocyst assemblages.[86][87][88]

Cysts can be transported via ocean-currents, which can distort ecological signals. This has been documented for the warm water species Operculodinium israelianum and Polysphaeridium zoharyi which were interpreted to have been transported along the Southern coast of the United States.[59] Cyst are also often transported from the inner shelf to the outer shelf or slope.[59]

Another problem with cysts is that they also get transported with ballast water, which can cause introduction of invasive species.[89]

Seasonality and fluxes are studied through sediment trap studies, which help to understand ecological signals.[90][91][92][93][94][95]

Palaeoecology of organic-walled dinocysts[edit]

The palaeoecology of marine organic-walled dinoflagellate cysts has been extensively studied, more particularly in the Quaternary. Changes in Quaternary dinocyst assemblages reflect the palaeoceanography through variations in productivity,[96][97][98][99][100] temperature,[101][102][103] salinity[104][105][106] and ice cover.[107][108][109]

Palynodinium, a fossil species of dinoflagellate cyst, is used to demarcate the K/Pg boundary, which marks the terminal Cretaceous and the extinction of the dinosaurs.[110]

Such reconstructions can be done via semi-quantitative techniques, such as ordination techniques,[46] which can indicate trends in environmental parameters.

A quantitative method is the use of transfer functions,[111][112][113][114][115] although these have been heavily debated.[116][117]

Another late Quaternary application is for environmental goals, more particularly the study of eutrophication[118][119][120] .[121]

An interval of particular interest during the late Quaternary is the Eemian.[122][123][124][125][126]

Also during the Neogene, dinocysts have shown to be useful in the Miocene[127] and particularly the Messinian.[128] Also the paleoclimate of the Pliocene has been investigated.[129][130][131] Transfer functions have also been attempted during the Pliocene.[132] Some species have been suggested to have different environmental preferences during the Neogene.[133]

The palaeoecology of freshwater dinoflagellate cysts is relatively unexplored, though several recent studies have shown the relation to changes in nutrients, pH and temperature[134][135][136][137]

Morphological variation of organic-walled dinocysts[edit]

There is little known about how organic-walled dinocysts are formed except from culture experiments.[138] Cyst formation is suggested to happen through self-assembly processes.[139]

Organic-walled dinocyst morphology is shown to be controlled by changes in salinity and temperature in some species, more particularly process length variation. This is known to be the case for Lingulodinium machaerophorum from culture experiments,[140] and study of surface sediments.[141] Also variations in the morphology of the species Operculodinium centrocarpum [142][143] can be related to salinity and/or temperature. Also cysts of the species Gonyaulax baltica shows morphological variations in culture,[144] as well as Gonyaulax spinifera.[145] Cyst formed by other species such as Pyrophacus steinii (cyst is called Tuberculodinium vancampoae) do not show a clear relation to variations in salinity.[146]

The morphological variation can be applied for the reconstruction of salinity, in a semi-quantitative[147] or quantitative way.[142] Process length variation of Lingulodinium machaerophorum has been used to reconstruct Black Sea salinity variation.[148]

Biostratigraphy and evolution of organic-walled dinocysts[edit]

Organic-walled dinoflagellate cysts have a long geological record with lowest occurrences during the mid Triassic,[149] whilst geochemical markers suggest a presence to the Early Cambrian.[150] Some of the Paleozoic acritarchs possibly are related to dinoflagellate cysts. Arpylorus, from the Silurian of North Africa, was at one time considered to be a dinoflagellate cyst,[151] but this palynomorph is now considered probably an arthropod remain.[152] Another enigmatic form with possible early dinoflagellate affinity is Palaeodinophysis altaica, which was found in the Devonian of Kazakhstan,[153] however Fensome et al. (1999) consider its dinoflagellate affinity (and also supposed age) unlikely.[154]

The fossil record supports a major adaptive radiation of dinoflagellates during later Triassic and earlier Jurassic times. The majority of living thecate dinoflagellates can be interpreted as having either a peridinalean or gonyaulacalean tabulation, and that these tabulations, and hence the orders Gonyaulacales and Peridiniales, have been separate since at least the Early Jurassic.[20] The biostratigraphical application of dinoflagellate cysts has been thoroughly studied.[155][156] The Pliocene has been recently investigated[157][158] and also the Miocene.[159]

Palynological methods[edit]

Organic-walled dinoflagellate cysts are extracted using palynological methods, which can be highly variable between different palynological laboratories, and often involve use of hydrochloric acid (HCl), hydrofluoric acid (HF) and/or alternative acids at different temperatures.[160][161][162][163] The use of KOH or acetolysis is not advised in dinocyst studies, because this causes swelling and/or destruction of dinocysts. The palynological method can cause difficulty in identification of certain species: it has been shown that cysts of Alexandrium tamarense and of Scrippsiella trifida are difficult to discriminate in samples that have been treated with the palynological method.[164] The concentration of Dinocysts can be quantified by adding an exotic spike or marker such as Lycopodium clavatum spores.[165][166][167]

Biological functions[edit]

Dinocysts are suggested to have a number of adaptive functions including survival during adverse conditions, bloom initiation and termination, dispersal in time, a seed bank for genetic diversity and dispersal in space.[168][169][170]


  1. ^ W.A.S. Sarjeant, 2002. 'As chimney-sweeps, come to dust': a history of palynology to 1970. pp. 273–327 In: Oldroyd, D. R. The earth inside and out: some major contributions to geology in the twentieth century. Geological Society (London) Special Publication no. 192.
  2. ^ Evitt, W.R. and Davidson, S.E. 1964. Dinoflagellate studies. 1. Dinoflagellate cysts and thecae. Stanford university publications X (1), pp. 3–12.
  3. ^ Wall, D.; Dale, B. (1966). "Living" fossils in western Atlantic plankton". Nature. 211 (5053): 1025–1026. Bibcode:1966Natur.211.1025W. doi:10.1038/2111025a0.
  4. ^ a b c Wall, D.; Dale, B. (1968). "Modern dinoflagellate cysts and evolution of the Peridiniales". Micropaleontology. 14 (3): 265–304. doi:10.2307/1484690. JSTOR 1484690.
  5. ^ Pfiester L.A. & Anderson D.M. 1987. Dinoflagellate reproduction. In: The biology of dinoflagellates. Botanical monographs 21 (Ed. by F.J.R. Taylor), pp. 611–648., Blackwell Scientific Publications.
  6. ^ Sarjeant, W.A.S.; Lacalli, T.; Gaines, G. (1987). "The cysts and skeletal elements of dinoflagellates: speculations on the ecological causes for their morphology and development". Micropaleontology. 33 (1): 1–36. doi:10.2307/1485525. JSTOR 1485525.
  7. ^ Spero, H.J.; Moree, M.D. (1981). "Phagotrophic feeding and its importance to the life cycle of the holozoic dinoflagellate Gymnodinium fungiforme". Journal of Phycology. 17: 43–51. doi:10.1111/j.1529-8817.1981.tb00817.x.
  8. ^ BRAVO I., FIGUEROA R.I., GARCÉS E., FRAGA S. & MASSANET A. 2010. The intricacies of dinoflagellate pellicle cysts: the example of Alexandrium minutum cysts from a bloom-recurrent area (Bay of Baiona, NW Spain). Deep-Sea Research Part II: Topical Studies in Oceanography 57: 166–174.
  9. ^ POPOVSKÝ J. & PFIESTER L.A. 1990. Dinophyceae (Dinoflagellida). In: Süßwasserflora von Mitteleuropa. Begründet von A. Pascher. Band 6 (Ed. by H. Ettl,J. Gerloff,H. Heynig. & D. Mollenhauer). Gustav Fischer Verlag, Jena, 272 pp.
  10. ^ Wall, D.; Dale, B. (1966). "Living fossils" in Atlantic plankton". Nature. 211 (5053): 1025–1026. doi:10.1038/2111025a0.
  11. ^ Sonneman, J.A.; Hill, D.R.A. (1997). "A taxonomic survey of cyst-producing dinoflagellates from recent sediments of Victorian coastal waters, Australia". Botanica Marina. 40 (1–6): 149–177. doi:10.1515/botm.1997.40.1-6.149.
  12. ^ Mertens, K.N.; Yamaguchi, A.; Kawami, H.; Ribeiro, S.; Leander, B.S.; Price, A.M.; Pospelova, V.; Ellegaard, M.; Matsuoka, K. (2012). "Archaeperidinium saanichi sp. nov.: a new species based on morphological variation of cyst and theca within the Archaeperidinium minutum Jörgensen 1912 species complex". Marine Micropaleontology. 96–97: 48–62. Bibcode:2012MarMP..96...48M. doi:10.1016/j.marmicro.2012.08.002.
  13. ^ Bolch, C.J.S. (1997). "The use of polytungstate for the separation and concentration of living dinoflagellate cysts from marine sediments". Phycologia. 36 (6): 472–478. doi:10.2216/i0031-8884-36-6-472.1.
  14. ^ Schwinghamer, P.; Anderson, D.M.; Kulis, D.M. (1991). "Separation and concentration of living dinoflagellate resting cysts from marine sediments via density-gradient centrifugation;". Limnology and Oceanography. 36 (3): 588–592. Bibcode:1991LimOc..36..588S. doi:10.4319/lo.1991.36.3.0588.
  15. ^ Bolch, C.J.S. (2001). "PCR protocols for genetic identification of dinoflagellates directly from single cysts and plankton cells". Phycologia. 40 (2): 162–167. doi:10.2216/i0031-8884-40-2-162.1.
  16. ^ Takano, Y.; Horiguchi, T. (2006). "Acquiring scanning electron microscopical, light microscopical and multiple gene sequence data from a single dinoflagellate cell". Journal of Phycology. 42: 251–256. doi:10.1111/j.1529-8817.2006.00177.x.
  17. ^ Kawami, H.; Van Wezel, R.; Koeman, R.P.; Matsuoka, K. (2009). "Protoperidinium tricingulatum sp. nov. (Dinophyceae), a new motile form of a round, brown, and spiny dinoflagellate cyst". Phycological Research. 57 (4): 259–267. doi:10.1111/j.1440-1835.2009.00545.x.
  18. ^ HEAD M.J. 1996. Modern dinoflagellate cysts and their biological affinities. In: Palynology: principles and applications (Ed. by J. Jansonius & D. C. McGregor), pp. 1197–1248. American Association of Stratigraphic Palynologists Foundation, Dallas, Texas.
  19. ^ a b Neil Mertens, Kenneth; Rengefors, Karin; Moestrup, Øjvind; Ellegaard, Marianne (2012). "A review of recent freshwater dinoflagellate cysts: taxonomy, phylogeny, ecology and palaeocology". Phycologia. 51 (6): 612–619. doi:10.2216/11-89.1.
  20. ^ a b c d Fensome, R.A.; Taylor, F.J.R.; Norris, G.; Sarjeant, W.A.S.; Wharton, D.I.; Williams, G.L. (1993). "A classification of living and fossil dinoflagellates". American Museum of Natural History, Micropaleontology, Special Publication. 7: 1–351.
  21. ^ Harland, R (1982). "A review of Recent and Quaternary organic-walled dinoflagellate cysts of the genus Protoperidinium". Palaeontology. 25: 369–397.
  22. ^ Schilling, A.J. (1891). "Die Süsswasser-Peridineen". Flora Oder Allgemeine Botanische Zeitung. 74: 220–299.
  23. ^ Tardio, M.; Ellegaard, M.; Lundholm, N.; Sangiorgi, F.; DI Giuseppe, D. (2009). "A hypocystal archeopyle in a freshwater dinoflagellate from the Peridinium umbonatum group (Dinophyceae) from Lake Nero di Cornisello, South-East Alps, Italy". European Journal of Phycology. 44 (2): 1–10. doi:10.1080/09670260802588442.
  24. ^ Moestrup, Ø.; Lindberg, K.; Daugbjerg, N. (2009). "Studies on woloszynskioid dinoflagellates IV: The genus Biecheleria gen. nov". Phycological Research. 57 (3): 203–220. doi:10.1111/j.1440-1835.2009.00540.x.
  25. ^ Evitt, W.R., Lentin, J.K., Millioud, M.E., Stover, L.E. and Williams, G.L., 1977. Dinoflagellate cyst terminology. Geological survey of Canada, Paper 76-24, 1-11.
  26. ^ Rochon, A., de Vernal, A., Turon, J.-L., Matthiessen, J., and Head, M.J., 1999. Distribution of recent dinoflagellate cysts in surface sediments from the North Atlantic Ocean and adjacent seas in relation to sea-surface parameters. AASP Contribution Series, 35, 146 pp.
  27. ^ MATSUOKA, K. & FUKUYO, Y. 2000. Technical guide for modern dinoflagellate cyst study. WESTPAC-HAB/WESTPAC/IOC, Japan Society of the Promotion Science, Tokyo, 29 pp.
  28. ^ Head, M.J.; Norris, G. (2003). "New species of dinoflagellate cysts and other palynomorphs from the late Neogene of the western North Atlantic, DSDP Hole 603C". Journal of Paleontology. 77: 1–15. doi:10.1666/0022-3360(2003)077<0001:nsodca>2.0.co;2.
  29. ^ Louwye, S.; Mertens, K.N.; Vercauteren, D. (2008). "New dinoflagellate cysts species from the Miocene of Porcupine Basin, off Southwest Ireland". Palynology. 32: 131–142. doi:10.2113/gspalynol.32.1.131.
  30. ^ Soliman, A., Head, M.J., and Louwye, S. In press. Morphology and distribution of the Miocene dinoflagellate cyst Operculodinium? borgerholtense Louwye 2001, emend. Palynology.
  31. ^ Head, M.J., 1999. The Late Pliocene St. Erth Beds of Cornwall: a review of the palynology and reappraisal of the dinoflagellates. In: Scource, J. and Furze, M.F.A. (eds.), The Quaternary of West Cornwall. Field Guide, Quaternary Research Association, Durham, U.K., p. 88–92.
  32. ^ Head, M.J. 2000. Geonettia waltonensis, a new goniodomacean dinoflagellate from the Pliocene of the North Atlantic region, and its evolutionary implications" Journal of Paleontology 74(5): 812–827, 6 pls.
  33. ^ De Schepper, S.; Head, M.J.; Louwye, S. (2004). "New dinoflagellate cyst and incertae sedis taxa from the Pliocene of northern Belgium, southern North Sea Basin". Journal of Paleontology. 78 (4): 625–644. doi:10.1666/0022-3360(2004)078<0625:ndcais>2.0.co;2.
  34. ^ De Schepper, S.; Head, M.J. (2008). "New dinoflagellate cyst and acritarch taxa from the Pliocene and Pleistocene of the eastern North Atlantic (DSDP Site 610)". Journal of Systematic Palaeontology. 6: 101–117. doi:10.1017/s1477201907002167.
  35. ^ Head, M.J. (2002). "Echinidinium zonneveldiae sp. nov., a new dinoflagellate cyst from the Late Pleistocene of the Baltic region". Journal of Micropalaeontology. 21 (2): 169–173. doi:10.1144/jm.21.2.169.
  36. ^ Verleye, T.; Pospelova, V.; Mertens, K.N.; Louwye, S. (2011). "The geographical distribution and (palaeo)ecology of Selenopemphix undulata sp. nov., a new late Quaternary dinoflagellate cyst from the Pacific Ocean". Marine Micropaleontology. 78 (3–4): 65–83. Bibcode:2011MarMP..78...65V. doi:10.1016/j.marmicro.2010.10.001.
  37. ^ Pospelova, V.; Head, M.J. (2002). "Islandinium brevispinosum sp. nov. (Dinoflagellata), a new organic-walled dinoflagellate cyst from modern estuarine sediments of New England (USA)". Journal of Phycology. 38 (3): 593–601. doi:10.1046/j.1529-8817.2002.01206.x.
  38. ^ Mertens, K.N.; Yamaguchi, A.; Kawami, H.; Ribeiro, S.; Leander, B.S.; Price, A.M.; Pospelova, V.; Ellegaard, M.; Matsuoka, K. (2012). "Archaeperidinium saanichi sp. nov.: a new species based on morphological variation of cyst and theca within the Archaeperidinium minutum Jörgensen 1912 species complex". Marine Micropaleontology. 96–97: 48–62. Bibcode:2012MarMP..96...48M. doi:10.1016/j.marmicro.2012.08.002.
  39. ^ De Vernal, A.; Marret, F. (2007). Organic-Walled Dinoflagellate Cysts: Tracers of Sea-Surface Conditions. Developments in Marine Geology. Vol. 1. pp. 371–408. doi:10.1016/S1572-5480(07)01014-7. ISBN 9780444527554.
  40. ^ Dale, B (1977). "New observations on Peridinium faeroense Paulsen (1905), and classification of small orthoperidinioid dinoflagellates". Br. Phycol. J. 12 (3): 241–253. doi:10.1080/00071617700650261.
  41. ^ Fensome, R.A., Taylor, F.J.R., Norris, G., Sarjeant, W.A.S., Wharton, D.I., and Williams, G.L., 1993. A classification of modern and fossil dinoflagellates, Sheridan Press, Hanover. .
  42. ^ DE, VERTEUIL L.; Norris, G. (1996). "Part 2. Homology and structure in dinoflagellate cyst terminology". Micropaleontology. S42: 83–172.
  43. ^ a b Jux, U (1968). "Über den feinbau der wandung bei Hystrichosphaera bentori Rossignol 1961". Palaeontographica Abteilung B. 123 (1–6): 147–152.
  44. ^ Head, M.J. (1994). "Morphology and paleoenvironmental significance of the Cenozoic dinoflagellate genera Tectatodinium and Habibacysta". Micropaleontology. 40 (4): 289–321. doi:10.2307/1485937. JSTOR 1485937.
  45. ^ Reid, P.C.; Boalch, G.T. (1987). "A new method for the identification of dinoflagellate cysts". Journal of Plankton Research. 9: 249–253. doi:10.1093/plankt/9.1.249.
  46. ^ a b Dale, B. & Dale, A.L. 2002. Environmental applications of dinoflagellate cysts and acritarchs . In Quaternary environmental micropalaeontology (Haslett, S.K., editor), 207-240. Arnold, London.
  47. ^ Jux, U (1971). "Über den feinbau der wandungen einiger Tertiärer Dinophyceen-zysten und Acritarcha Hystrichosphaeridium, Impletosphaeridium, Lingulodinium". Palaeontographica, Abt. B. 132 (5–6): 165–174.
  48. ^ Jux, U (1976). "Über den feinbau der wandungen bei Operculodinium centrocarpum (Deflandre & Cookson) Wall 1967 und Bitectatodinium tepikiense Wilson 1973". Palaeontographica, Abt. B. 155 (5–6): 149–156.
  49. ^ Kokinos, J. P.; Eglinton, T.I.; Goñi, M.A.; Boon, J.J.; Martoglio, P.A.; Anderson, D.M. (1998). "Characterisation of a highly resistant biomacromolecular material in the cellwall of a marine dinoflagellate resting cyst". Organic Geochemistry. 28 (5): 265–288. doi:10.1016/s0146-6380(97)00134-4.
  50. ^ Kennaway, Gay; Lewis, Jane (2004). "An ultrastructural study of hypnozygotes of Alexandrium species (Dinophyceae)". Phycologia. 43 (4): 355–363. doi:10.2216/i0031-8884-43-4-355.1.
  51. ^ Chapman, D. V.; Dodge, J. D.; Heaney, S. I. (1982). "Cyst Formation in the Freshwater Dinoflagellaie Ceratium Hirundinella (Dinophyceae)". Journal of Phycology. 18: 121–129. doi:10.1111/j.1529-8817.1982.tb03165.x.
  52. ^ Stosch, H.A. VON (1965). "Sexualität bei Ceratium cornutum (Dinophyta)". Die Naturwissenschaften. 52 (5): 112–113. doi:10.1007/bf00626331.
  53. ^ Pfiester, L. A. & Anderson, D. M. 1987. Dinoflagellate reproduction. In: The biology of dinoflagellates (ed. F. J. R. Taylor), pp. 611–648. - Blackwell, Oxford.
  54. ^ RENGEFORS K. 1998. Seasonal succession of dinoflagellates coupled to the benthic cyst dynamics in Lake Erken, Sweden. Archiv für Hydrobiologie, Special Issues, Advances in Limnology 51: 123–141.
  55. ^ Alpermann, T.J.; Beszteri, B.; John, U.; Tillmann, U.; Cembella, A.D. (2009). "Implications of life history transitions on the population genetic structure of the toxigenic marine dinoflagellate Alexandrium tamarense". Molecular Ecology. 18 (10): 2122–2133. doi:10.1111/j.1365-294x.2009.04165.x. PMID 19389181.
  56. ^ Ribeiro, S.; Berge, T.; Lundholm, N.; Andersen, T.J.; Abrantes, F.; Ellegaard, M. (2011). "Phytoplankton growth after a century of dormancy illuminates past resilience to catastrophic darkness". Nature Communications. 2: 311. Bibcode:2011NatCo...2..311R. doi:10.1038/ncomms1314. PMC 3113231. PMID 21587228.
  57. ^ Lewis, Jane; Harris, A.; Jones, K.; Edmonds, R. (1999). "Long-term survival of marine planktonic diatoms and dinoflagellates in stored sediment samples". Journal of Plankton Research. 21 (2): 343–354. doi:10.1093/plankt/21.2.343.
  58. ^ Binder, B. J.; Anderson, D. M. (1986). "Green light-mediated photomorphogenesis in a dinoflagellate resting cyst". Nature. 322 (6080): 659–661. Bibcode:1986Natur.322..659B. doi:10.1038/322659a0.
  59. ^ a b c Wall, D.; Dale, B.; Lohman, G.P.; Smith, W.K. (1977). "The environmental and climatic distribution of dinoflagellate cysts in modern sediments from regions in the North and South Atlantic oceans and adjacent seas". Marine Micropaleontology. 2: 121–200. Bibcode:1977MarMP...2..121W. doi:10.1016/0377-8398(77)90008-1.
  60. ^ Sprangers, M.; Dammers, N.; Brinkhuis, H.; van Weering, T.C.E.; Lotter, A.F. (2004). "Modern organic-walled dinoflagellate cyst distribution offshore NW Iberia; tracing the upwelling system". Review of Palaeobotany and Palynology. 128 (1–2): 97–106. doi:10.1016/s0034-6667(03)00114-3. hdl:1874/19661.
  61. ^ Nehring, S (1995). "Dinoflagellate resting cysts as factors in phytoplankton ecology of the North Sea". Helgoländer Meeresuntersuchungen. 49 (1–4): 375–392. Bibcode:1995HM.....49..375N. doi:10.1007/bf02368363.
  62. ^ Nehring, S (1994). "Spatial distribution of dinoflagellate resting cysts in Recent sediments of Kiel Bight, Germany (Baltic Sea)". Ophelia. 39 (2): 137–158. doi:10.1080/00785326.1994.10429540.
  63. ^ Marret, F.; Scource, J. (2002). "Control of modern dinoflagellate cyst distribution in the Irish and Celtic seas by seasonal stratification dynamics". Marine Micropaleontology. 47 (1–2): 101–116. Bibcode:2003MarMP..47..101M. doi:10.1016/s0377-8398(02)00095-6.
  64. ^ Matthießen, J. (1995) Distribution patterns of dinoflagellate cysts and other organic-walled microfossils in recent Norwegian-Greenland Sea sediments , Marine Micropaleontology
  65. ^ Marret, F.; Eiriksson, J.; Knudsen, K-L.; Scourse, J. (2004). "Distribution of dinoflagellate cyst assemblages in surface sediments from the northern and western shelf of Iceland". Review of Palaeobotany and Palynology. 128 (1–2): 35–54. doi:10.1016/s0034-6667(03)00111-8.
  66. ^ Verleye, T.J.; Louwye, S. (2010). "Recent geographical distribution of organic-walled dinoflagellate cysts in the southeast Pacific (25-53ºS) and their relation to the prevailing hydrographical conditions". Palaeogeography, Palaeoclimatology, Palaeoecology. 298 (3–4): 319–340. Bibcode:2010PPP...298..319V. doi:10.1016/j.palaeo.2010.10.006.
  67. ^ Richerol, T; Rochon, A; Blasco, S; Scott, DB; Schell; Bennett, R (2008). "Distribution of dinoflagellate cysts in surface sediments of the Mackenzie Shelf and Amundsen Gulf, Beaufort Sea (Canada)". Journal of Marine Systems. 74 (3): 825–839. Bibcode:2008JMS....74..825R. doi:10.1016/j.jmarsys.2007.11.003.
  68. ^ Matthiessen, J., De Vernal, A., Head, M., Okolodkov, Y., Ángel, P., Zonneveld, K. and Harland, R. Modern organic-walled dinoflagellate cysts in Arctic marine environments and their (paleo-) environmental significance. Paläontologische Zeitschrift 79(1): 3-51.
  69. ^ Vink, A.; Zonneveld, K.A.F.; Willems, H. (2000). "Organic-walled dinoflagellate cysts in western equatorial Atlantic surface sediments: distribution and their relation to environment". Review of Palaeobotany and Palynology. 112 (4): 247–286. doi:10.1016/s0034-6667(00)00046-4. PMID 11134709.
  70. ^ Vink, A., Baumann, K-H., Böckel, B., Esper, O., Kinkel, H., Volbers, A., Willems, H., Zonneveld, K.A.F. Coccolithophorid and dinoflagellate synecology in the South and Equatorial Atlantic: Improving the paleoecological significance of phytoplankton microfossils. In: Wefer, G., Mulitza, S. and Ratmeyer, V. (eds.) The South Atlantic in the Late Quaternary: reconstruction of material budgets and current systems. Springer, Berlin: 121-142.
  71. ^ Bouimetarhan, I.; Marret, F.; Dupont, L.; Zonneveld, K.A.F. (2009). "Dinoflagellate cyst distribution in marine surface sediments off West Africa (17 – 6°N) in relation to sea-surface conditions, freshwater input and seasonal coastal upwelling". Marine Micropaleontology. 71 (3–4): 113–130. Bibcode:2009MarMP..71..113B. doi:10.1016/j.marmicro.2009.02.001.
  72. ^ Oliver Esper, Karin Zonneveld. The potential of organic-walled dinoflagellate cysts to reconstruct past sea-surface conditions in the Southern Ocean" Marine Micropaleontology 63 (3/4): 185-212.
  73. ^ Holzwarth, Ulrike; Esper, Oliver; Zonneveld, Karin A.F. (2007). "Distribution of organic-walled dinoflagellate cysts in shelf surface sediments of the Benguela upwelling system in relationship to environmental conditions". Marine Micropaleontology. 64 (1–2): 91–119. Bibcode:2007MarMP..64...91H. doi:10.1016/j.marmicro.2007.04.001.
  74. ^ Elshanawany, R., Zonneveld, K.A.F., Ibrahim, M.I. and Kholeif, S.E.A. (2010). Distribution patterns of recent organic-walled dinoflagellate cysts in relation to environmental parameters in the Mediterranean Sea. Palynology
  75. ^ Marret, F.; Leroy, S.A.G.; ChaliÉ, F.; Gasse, F. (2004). "New organic-walled dinoflagellate cysts from recent sediments of Central Asian seas". Review of Palaeobotany and Palynology. 129 (1–2): 1–20. doi:10.1016/j.revpalbo.2003.10.002.
  76. ^ Radi, T.; Pospelova, V.; de Vernal, A.; Barrie, J.V. (2007). "Dinoflagellate cysts as indicators of water quality and productivity in British Columbia estuarine environments". Marine Micropaleontology. 62 (4): 296–297. Bibcode:2007MarMP..62..269R. doi:10.1016/j.marmicro.2006.09.002.
  77. ^ Pospelova, V; de Vernal, A; Pedersen, TF (2008). "Distribution of dinoflagellate cysts in surface sediments from the northeastern Pacific Ocean (43-25°N) in relation to sea-surface temperature, salinity, productivity and coastal upwelling". Marine Micropaleontology. 68 (1–2): 21–48. Bibcode:2008MarMP..68...21P. doi:10.1016/j.marmicro.2008.01.008.
  78. ^ Cremer, H.; Sangiorgi, F.; Wagner, F.; McGee, V.; Lotter, A.F.; Visscher, H. (2007). "Marine Littoral Diatoms (Bacillariophyceae) and Dinoflagellates cysts (Dinophyceae) from Rookery Bay, Florida, U.S.A.". Caribbean Journal of Science. 43 (1): 23–58. doi:10.18475/cjos.v43i1.a4.
  79. ^ Limoges, A.; Kielt, J.-F.; Radi, T.; Ruíz-Fernandez, A.C.; de Vernal, A. (2010). "Dinoflagellate cyst distribution in surface sediments along the south-western Mexican coast (14.76° N to 24.75°N)". Marine Micropaleontology. 76 (3–4): 104–123. Bibcode:2010MarMP..76..104L. doi:10.1016/j.marmicro.2010.06.003.
  80. ^ Solignac, S.; Grøsfjeld, K.; Giraudeau, J.; de Vernal, A. (2009). "Distribution of recent dinocyst assemblages in the western Barents Sea". Norwegian Journal of Geology. 89 (1–2): 109–119.
  81. ^ Zonneveld, Karin A.F.; Marret, Fabienne; Versteegh, Gerard; Bogus, Kara; Bonnet, Sophie; Bouimetarhan, Ilham; Crouch, Erica; de Vernal, Anne; Elshanawany, Rehab; Edwards, Lucy; Esper, Oliver; Forke, Sven; Grøsfjeld, Kari; Henry, Maryse; Holzwarth, Ulrike; Kielt, Jean-François; Kim, So-Young; Ladouceur, Stéphanie; Ledu, David; Liang, Chen; Limoges, Audrey; Londeix, Laurent; Lu, S.-H.; Mahmoud, Magdy S.; Marino, Gianluca; Matsouka, Kazumi; Matthiessen, Jens; Mildenhal, D.C.; Mudie, Peta; Neil, H.L.; Pospelova, Vera; Qi, Yuzao; Radi, Taoufik; Richerol, Thomas; Rochon, André; Sangiorgi, Francesca; Solignac, Sandrine; Turon, Jean-Louis; Verleye, Thomas; Wang, Yan; Wang, Zhaohui; Young, Marty (2013). "Atlas of modern dinoflagellate cyst distribution based on 2405 datapoints". Review of Palaeobotany and Palynology. 191: 1–197. doi:10.1016/j.revpalbo.2012.08.003. hdl:1854/LU-3226112.
  82. ^ Dale, B., 1996. Dinoflagellate cyst ecology: modelling and geological applications. In Jansonius, J. & McGregor, D.C. (eds.): Palynology: Principles and Applications, volume 3, 1249-1275, AASP Foundation, Dallas.
  83. ^ Head, M.J., Harland, R., and Matthiessen, J. 2001. Cold marine indicators of the late Quaternary: the new dinoflagellate cyst genus Islandinium and related morphotypes. Journal of Quaternary Science, 16(7): 621–636, 3 pls.
  84. ^ Comeau, A. M.; Philippe, B.; Thaler, M.; Gosselin, M.; Poulin, M.; Lovejoy, C. (2013). "Protists in Arctic Drift and Land-Fast Sea Ice". Journal of Phycology. 49 (2): 229–240. doi:10.1111/jpy.12026. PMID 27008512.
  85. ^ Mertens, K.N.; Takano, Y.; Head, M.J.; Matsuoka, K. (2014). "Living fossils in the Indo-Pacific warm pool: A refuge for thermophilic dinoflagellates during glaciations". Geology. 42 (6): 531–534. Bibcode:2014Geo....42..531M. doi:10.1130/G35456.1.
  86. ^ Matsuoka, K; Joyce, LB; Kotani, Y; Matsuyama, Y (2003). "Modern dinoflagellate cysts in hypertrophic coastal waters of Tokyo Bay, Japan". Journal of Plankton Research. 25 (12): 1461–1470. doi:10.1093/plankt/fbg111.
  87. ^ Pospelova, V.; Chmura, G.L.; Boothman, W.; Latimer, J.S. (2005). "Spatial distribution of modern dinoflagellate cysts in polluted estuarine sediments from Buzzards Bay (Massachusetts, USA) embayments". Marine Ecology Progress Series. 292: 23–40. Bibcode:2005MEPS..292...23P. doi:10.3354/meps292023.
  88. ^ Krepakevich, A.; Pospelova, V. (2010). "Anthropogenic impact on coastal bays of Southern Vancouver Island (BC, Canada) as reflected in phytoplankton sedimentary records". Continental Shelf Research. 30 (18): 1924–1940. Bibcode:2010CSR....30.1924K. doi:10.1016/j.csr.2010.09.002.
  89. ^ Hallegraeff, GM (1998). "Transport of toxic dinoflagellates via ships' ballast water: bioeconomic risk assessment and efficacy of possible ballast water management strategies". Marine Ecology Progress Series. 168: 297–309. Bibcode:1998MEPS..168..297H. doi:10.3354/meps168297.
  90. ^ Susek, E.; Zonneveld, K.A.F.; Fischer, G.; Versteegh, G.J.M.; Willems, H. (2005). "Organic walled dinoflagellate cyst production in relation to upwelling intensity and lithogenic influx in the Cape Blanc region (off north-west Africa)". Phycological Research. 53 (2): 97–112. doi:10.1111/j.1440-1835.2005.tb00362.x.
  91. ^ Price, AM; Pospelova, V (2011). "High-resolution sediment trap study of organic-walled dinoflagellate cyst production and biogenic silica flux in Saanich Inlet (BC, Canada)". Marine Micropaleontology. 80 (1–2): 18–43. Bibcode:2011MarMP..80...18P. doi:10.1016/j.marmicro.2011.03.003.
  92. ^ Pospelova, V.; Esenkulova; Johannessen, S.C.; O'Brien, M.C.; Macdonald, R.W. (2010). "Organic-walled dinoflagellate cyst production, composition and flux from 1996 to 1998 in the central Strait of Georgia (BC, Canada): a sediment trap study". Marine Micropaleontology. 75 (1–4): 17–37. Bibcode:2010MarMP..75...17P. doi:10.1016/j.marmicro.2010.02.003.
  93. ^ Fujii, R.; Matsuoka, K. (2005). "Seasonal change of dinoflagellates cyst flux collected in a sediment trap in Omura Bay, West Japan". Journal of Plankton Research. 28 (2): 131–147. doi:10.1093/plankt/fbi106.
  94. ^ Zonneveld, K. A. F. Susek E.; Fischer, G. (2010). "Interannual and seasonal variability of the organic-walled dinoflagellate cyst production in the coastal upwelling region off Cape Blanc (Mauritania)". Journal of Phycology. 46 (1): 202–215. doi:10.1111/j.1529-8817.2009.00799.x.
  95. ^ Bringué, Manuel; Pospelova, Vera; Pak, Dorothy (2013). "Seasonal production of organic-walled dinoflagellate cysts in an upwelling system: A sediment trap study from the Santa Barbara Basin, California". Marine Micropaleontology. 100: 34–51. Bibcode:2013MarMP.100...34B. doi:10.1016/j.marmicro.2013.03.007.
  96. ^ Pospelova, V.; Pedersen, T F.; DE Vernal, A. (2006). "Dinoflagellate cysts as indicators of climatic and oceanographic changes during the past 40 kyr in the Santa Barbara Basin, southern California". Paleoceanography. 21 (2): 2010. Bibcode:2006PalOc..21.2010P. doi:10.1029/2005PA001251.
  97. ^ González, C.; Dupont, L.M.; Mertens, K.; Wefer, G. (2008b). "Reconstructing marine productivity of the Cariaco Basin during marine isotope stage 3 and 4 using organic-walled dinoflagellate cysts". Paleoceanography. 23 (3): 3215. Bibcode:2008PalOc..23.3215G. doi:10.1029/2008PA001602. hdl:1854/LU-592855.
  98. ^ Mertens, K. N.; González, C.; Delusina, I.; Louwye, S. (2009b). "30 000 years of productivity and salinity variations in the late Quaternary Cariaco Basin revealed by dinoflagellate cysts". Boreas. 38 (4): 647–662. doi:10.1111/j.1502-3885.2009.00095.x.
  99. ^ Verleye, T.J.; Louwye, S. (2010). "Late Quaternary environmental changes and latitudinal shifts of the Antarctic Circumpolar Current as recorded by dinoflagellate cysts from offshore Chile (41ºS)". Quaternary Science Reviews. 29 (7): 1025–1039. Bibcode:2010QSRv...29.1025V. doi:10.1016/j.quascirev.2010.01.009.
  100. ^ Price, A.M.; Mertens, K.N.M.; Pospelova, V.; Pedersen, T.F.; Ganeshram, R.S. (2013). "Late Quaternary climatic and oceanographic changes in the Northeast Pacific as recorded by dinoflagellate cysts from Guaymas Basin, Gulf of California (Mexico)". Paleoceanography. 28 (1): 200–212. Bibcode:2013PalOc..28..200P. doi:10.1002/palo.20019. hdl:1854/LU-3131952.
  101. ^ Londeix, L.; Herreyre, Y.; Turon, J.L.; Fletcher, W. (2009). "Last Glacial to Holocene hydrology of the Marmara Sea inferred from a dinoflagellate cyst record". Review of Palaeobotany and Palynology. 158 (1–2): 52–71. doi:10.1016/j.revpalbo.2009.07.004.
  102. ^ Chen, L.; Zonneveld, K.A.F.; Versteegh, G.J.M. (2011). "Short term climate variability during the "Roman Classical Period" in the Eastern Mediterranean". Quaternary Science Reviews. 30 (27): 3880–3891. Bibcode:2011QSRv...30.3880C. doi:10.1016/j.quascirev.2011.09.024.
  103. ^ Kunz-Pirrung, M.; Matthießen, J.; Vernal, A. (2001). "Late Holocene dinoflagellate cysts as indicators for short-term climate variability in the eastern Laptev Sea (Arctic Ocean)". Journal of Quaternary Science. 16 (7): 711–716. Bibcode:2001JQS....16..711K. doi:10.1002/jqs.649.
  104. ^ Sorrel, P.; Popescu, S.-M.; Head, M.J.; Suc, J.P.; Klotz, S.; Oberhänsli, H. (2006). "Hydrographic development of the Aral Sea during the last 2000 years based on a quantitative analysis of dinoflagellate cysts". Palaeogeography, Palaeoclimatology, Palaeoecology. 234 (2–4): 304–327. Bibcode:2006PPP...234..304S. doi:10.1016/j.palaeo.2005.10.012.
  105. ^ Ellegaard, M (2000). "Variations in dinoflagellate cyst morphology under conditions of changing salinity during the last 2000 years in the Limfjord, Denmark". Review of Palaeobotany and Palynology. 109 (1): 65–81. doi:10.1016/s0034-6667(99)00045-7. PMID 10708791.
  106. ^ Mudie, P.J.; Aksu, A.E.; Yaşar, D. (2001). "Late Quaternary dinoflagellate cysts from the Black, Marmara and Aegean seas: variations in assemblages, morphology and paleosalinity". Marine Micropaleontology. 43 (1–2): 155–178. Bibcode:2001MarMP..43..155M. doi:10.1016/s0377-8398(01)00006-8.
  107. ^ DE, VERNAL; Eynaud, F.; Henry, M.; Hillaire-MARCEL, C.; Londeix, L.; Mangin, S.; Matthiessen, J.; Marret, F.; Radi, T.; Rochon, A.; Solignac, S.; Turon, J.-L. (2005). "Reconstruction of sea-surface conditions at middle to high latitudes of the Northern Hemisphere during the Last Glacial Maximum (LGM) based on dinoflagellate cyst assemblages". Quaternary Science Reviews. 24 (7–9): 897–924. Bibcode:2005QSRv...24..897D. doi:10.1016/j.quascirev.2004.06.014.
  108. ^ Bonnet, S.; de Vernal, A.; Hillaire-Marcel, C.; Radi, T.; Husum, K. (2010). "Variability of seasurface temperature and sea-ice cover in the Fram Strait over the last two millennia". Mar. Micropaleontol. 74 (3–4): 59–74. Bibcode:2010MarMP..74...59B. doi:10.1016/j.marmicro.2009.12.001.
  109. ^ de Vernal, A., Rochon, A., 2011. Dinocysts as tracers of sea-surface conditions and sea-ice cover in polar and subpolar environments, IOP Conference Series: Earth and Environmental Science, 14, 012007.
  110. ^ Beaudoin, Alwynne Bowyer; Head, M. J.; Head, Martin J. (2004). The Palynology and Micropalaeontology of Boundaries. p. 261. ISBN 9781862391604.
  111. ^ De Vernal, A.; Henry, M.; Matthiessen, J.; Mudie, P.J.; Rochon, A.; Boessenkool, K.P.; Eynaud, F.; Grøsfjeld, K.; Guiot, J.; Hamel, D.; Harland, R.; Head, M.J.; Kunz-Pirrung, M.; Levac, E.; Loucheur, V.; Peyron, O.; Pospelova, V.; Radi, T.; Turon, J.-L.; Voronina, E. (2001). "Dinoflagellate cyst assemblages as tracers of sea-surface conditions in the northern North Atlantic, Arctic and sub-Arctic seas: the new 'n = 677' data base and its application for quantitative palaeoceanographic reconstruction". Journal of Quaternary Science. 16 (7): 681–698. Bibcode:2001JQS....16..681D. doi:10.1002/jqs.659.
  112. ^ DE, VERNAL A.; Eynaud, F.; Henry, M.; Hillaire-MARCEL, C.; Londeix, L.; Mangin, S.; Matthiessen, J.; Marret, F.; Radi, T.; Rochon, A.; Solignac, S.; Turon, J.-L. (2005). "Reconstruction of sea-surface conditions at middle to high latitudes of the Northern Hemisphere during the Last Glacial Maximum (LGM) based on dinoflagellate cyst assemblages". Quaternary Science Reviews. 24 (7–9): 897–924. Bibcode:2005QSRv...24..897D. doi:10.1016/j.quascirev.2004.06.014.
  113. ^ Guiot, J., de Vernal, A., 2007. Transfer functions: methods for quantitative paleoceanography based on microfossils, In Hillaire-Marcel and de Vernal (eds.) Proxies in Late Cenozoic Paleoceanography, Elsevier, pp. 523–563.
  114. ^ Waelbroeck, C.; Paul, A.; Kucera, M.; Rosell-Melé, A.; Weinelt, M.; Schneider, R.; Mix, A.; Abelmann-Gersonde, A.; Armand, L.; Barker, S.; Barrows, T.; Benway, H.; Cacho, I.; Chen, M.; Cortijo, E.; Crosta, X.; de Vernal, A.; Dokken, T.; Duprat, J.; Elderfield, H.; Eynaud, F.; Gersonde, R.; Hayes, A.; Henry, M.; Hillaire-Marcel, C.; Huang, C.; Jansen, E.; Juggins, S.; Kallel, N.; Kiefer, T.; Kienast, M.; Labeyrie, L.; Leclaire, H.; Londeix, L.; Mangin, S.; Matthießen, J.; Marret, F.; Meland, M.; Morey, A.; Mulitza, S.; Pflaumann, U.; Pisias, N.; Radi, T.; Rochon, A.; Rohling, E.; Sbaffi, L.; Schäfer-Neth, C.; Solignac, S.; Spero, H.; Tachikawa, K.; Turon, J. (2009). "Constraints on the magnitude and patterns of ocean cooling at the Last Glacial Maximum". Nature Geoscience. 2 (2): 127–132. Bibcode:2009NatGe...2..127M. doi:10.1038/NGEO411. hdl:1885/52333.
  115. ^ Eynaud, F.; Turon, J.; Matthießen, J.; Peypouquet, F.; Vernal, A.; Henry, M. (2002). "Norwegian sea-surface palaeoenvironments of marine oxygen-isotope stage 3: the paradoxical response of dinoflagellate cysts". Journal of Quaternary Science. 17 (4): 349–359. Bibcode:2002JQS....17..349E. doi:10.1002/jqs.676.
  116. ^ Telford, R.J., 2006. Limitations of dinoflagellate cyst transfer functions. Quaternary Science Reviews 25 : 1375-1382.
  117. ^ Guiot, J.; DE Vernal, A. (2011). "Is spatial autocorrelation introducing biases in the apparent accuracy of paleoclimatic reconstructions". Quaternary Science Reviews. 30 (15): 1965–1972. Bibcode:2011QSRv...30.1965G. doi:10.1016/j.quascirev.2011.04.022.
  118. ^ Dale, B (2001). "Marine dinoflagellate cysts as indicators of eutrophication and pollution:a discussion". Sci. Total Environ. 264 (3): 235–240. Bibcode:2001ScTEn.264..235D. doi:10.1016/s0048-9697(00)00719-1. PMID 11213194.
  119. ^ Dale, B (2009). "Eutrophication signals in the sedimentary record of dinoflagellate cysts in coastal waters". Journal of Sea Research. 61 (1): 103–113. Bibcode:2009JSR....61..103D. doi:10.1016/j.seares.2008.06.007.
  120. ^ Zonneveld, K.A.F.; Chen, L.; El-Shanawany, R.; Fischer, H.W.; Hoins, M.; Pittaurova, D. (2012). "The use of dinoflagellate cysts to separate human and natural variability in the trophic state of the Po River discharge plume during the last two centuries". Marine Pollution Bulletin. 64 (1): 114–132. doi:10.1016/j.marpolbul.2011.10.012. PMID 22118910.
  121. ^ Donders, T.H.; Gorissen, P.M.; Sangiorgi, F.; Cremer, H.; Wagner-Cremer, F.; McGee, V. (2008). "Three-hundred-year hydrological changes in a subtropical estuary, Rookery Bay (Florida): Human impact versus natural variability". Geochemistry, Geophysics, Geosystems. 9 (7): Q07V06. Bibcode:2008GGG.....9.7V06D. doi:10.1029/2008GC001980.
  122. ^ Head, M.J.; Seidenkrantz, M.-S.; Janczyk-Kopikowa, Z.; Marks, L.; Gibbard, P.L. (2005). "Last Interglacial (Eemian) hydrographic conditions in the southeastern Baltic Sea, NE Europe, based on dinoflagellate cysts". Quaternary International. 130 (1): 3–30. Bibcode:2005QuInt.130....3H. doi:10.1016/j.quaint.2004.04.027.
  123. ^ Head, M.J. (2007). "Last Interglacial (Eemian) hydrographic conditions in the southwestern Baltic Sea based on dinoflagellate cysts from Ristinge Klint, Denmark". Geological Magazine. 144 (6): 987–1013. Bibcode:2007GeoM..144..987H. doi:10.1017/s0016756807003780.
  124. ^ Van Nieuwenhove, N.; Bauch, H.A. (2008). "Last Interglacial (MIS 5e) surface water conditions at the Vøring Plateau (Norwegian Sea), based on dinoflagellate cysts". Polar Research. 27 (2): 175–186. doi:10.3402/polar.v27i2.6175.
  125. ^ Van Nieuwenhove, N., Bauch, H.A., Matthiessen, J., 2008. Last Interglacial surface water conditions in the eastern Nordic Seas inferred from dinocyst
  126. ^ Matthießen, J.; Knies, J. (2001). "Dinoflagellate cyst evidence for warm interglacial conditions at the northern Barents Sea margin, during marine isotope stage 5". Journal of Quaternary Science. 16 (7): 727–737. Bibcode:2001JQS....16..727M. doi:10.1002/jqs.656.
  127. ^ Louwye, S., Foubert, A., Mertens, K.N., Van Rooij, D. & IODP Expedition 307 scientific party (2007). Integrated stratigraphy and palaeoecology of the Lower and Middle Miocene of the Porcupine Basin. Geological Magazine 145, 321-344.
  128. ^ Popescu, S.-M.; Dalesme, F.; Jouannic, G.; Escarguel, G.; Head, M.J.; Melinte-Dobrinescu, M.C.; Sütö-Szentai, M.; Bakrac, K.; Clauzon, G.; Suc, J.-P. "Galeacysta etrusca Corradini & Biffi 1988, dinoflagellate cyst marker of Paratethyan influxes into the Mediterranean Sea before and after the peak of the Messinian Salinity Crisis". Palynology.
  129. ^ Head, M.J. and Westphal, H. 1999. Palynology and paleoenvironments of a Pliocene carbonate platform: the Clino Core, Bahamas" Journal of Paleontology 73(1): 1–25, 8 pls.
  130. ^ De Schepper, S.; Head, M.J.; Louwye, S. (2009). "Pliocene dinoflagellate cyst stratigraphy, palaeoecology and sequence stratigraphy of the Tunnel-Canal Dock, Belgium". Geological Magazine. 146 (1): 92–112. Bibcode:2009GeoM..146...92D. doi:10.1017/s0016756808005438.
  131. ^ De Schepper S, Head MJ, Groeneveld J (2009) North Atlantic Current variability through marine isotope stage M2 (circa 3.3 Ma) during the mid-Pliocene. Paleoceanography 24:PA4206
  132. ^ Edwards, L.E.; Mudie, P.J.; de Vernal, A. (1991). "Pliocene paleoclimatic reconstruction using dinoflagellate cysts: comparison of methods". Quat. Sci. Rev. 10 (2): 259–274. Bibcode:1991QSRv...10..259E. doi:10.1016/0277-3791(91)90024-o.
  133. ^ De Schepper, S.; Fischer, E.; Groeneveld, J.; Head, M.; Matthießen, J. (2011). "Deciphering the palaeoecology of Late Pliocene and Early Pleistocene dinoflagellate cysts". Palaeogeography, Palaeoclimatology, Palaeoecology. 309 (1–2): 17–32. doi:10.1016/j.palaeo.2011.04.020.
  134. ^ Mccarthy, F.M.G.; Mertens, K.N.; Ellegaard, M.; Sherman, K.; Pospelova, V.; Ribeiro, S.; Blasco, S.; Vercauteren, D. (2011). "Resting cysts of freshwater dinoflagellates in southeastern Georgian Bay (Lake Huron) as proxies of cultural eutrophication". Review of Palaeobotany and Palynology. 166 (1–2): 46–62. doi:10.1016/j.revpalbo.2011.04.008.
  135. ^ CHU, G.; SUN, Q.; Rioual, P.; Boltovskoy, A.; LIU, Q.; SUN, P.; HAN, J.; LIU, J. (2008). "Distinct microlaminations and freshwater "red tides" recorded in Lake Xiaolongwan, northeastern, China". Journal of Paleolimnology. 39 (3): 319–333. Bibcode:2008JPall..39..319C. doi:10.1007/s10933-007-9106-1.
  136. ^ CHU, G; SUN, Q.; Wang, X.; LI, D.; Rioual, P.; Qiang, L.; HAN, J.; LIU, J. (2009). "A 1600 year multiproxy record of paleoclimatic change from varved sediments in Lake Xiaolongwan, northeastern China". Journal of Geophysical Research. 114 (D22): D22108. Bibcode:2009JGRD..11422108C. doi:10.1029/2009JD012077.
  137. ^ Tardio, M.; Sangiorgi, F.; Brinkhuis, H.; Filippi, M.L.; Cantonati, M.; Lotter, A.F. (2006). "Peridinioid dinoflagellate cysts in a Holocene high-mountain lake deposits in Italy". Journal of Paleolimnology. 36 (3): 315–318. Bibcode:2006JPall..36..315T. doi:10.1007/s10933-006-9001-1.
  138. ^ Kokinos, J.P.; Anderson, D.M. (1995). "Morphological development of resting cysts in cultures of the marine dinoflagellate Lingulodinium polyedrum (= L. machaerophorum)". Palynology. 19: 143–166. doi:10.1080/01916122.1995.9989457.
  139. ^ Hemsley, A.R.; Lewis, J.; Griffiths, P.C. (2004). "Soft and sticky development : some underlying reasons for microarchitectural pattern convergence". Review of Palaeobotany and Palynology. 130 (1–4): 105–119. doi:10.1016/j.revpalbo.2003.12.004.
  140. ^ Hallett, R.I., 1999. Consequences of environmental change on the growth and morphology of Lingulodinium polyedrum (Dinophyceae) in culture. PhD thesis. University of Westminster, 109 pp.
  141. ^ Mertens, K. N.; Ribeiro, S. Bouimetarhan; Caner, H.; Combourieu-Nebout, N. Dale; de Vernal, M. Filipova; Ellegaard, A.; Godhe, U. Leroy; Grøsfjeld, A.; Holzwarth, K.; Kotthoff, U.; Londeix, L.; Marret, F.; Matsuoka, K.; Mudie, P.; Naudts, L.; Peña-manjarrez, J.; Persson, A.; Popescu, S.; Sangiorgi, F.; van der Meer, M.; Vink, A.; Zonneveld, K.; Vercauteren, D.; Vlassenbroeck, J.; Louwye, S. (2009a). "Process length variation in cysts of a dinoflagellate, Lingulodinium machaerophorum, in surface sediments investigating its potential as salinity proxy". Marine Micropaleontology. 70 (1–2): 54–69. Bibcode:2009MarMP..70...54M. doi:10.1016/j.marmicro.2008.10.004.
  142. ^ a b Mertens, K.N.; Dale, B.; Ellegaard, M.; Jansson, I.-M.; Godhe, A.; Kremp, A.; Louwye, S. (2010). "Process length variation in cysts of the dinoflagellate Protoceratium reticulatum from surface sediments of the Baltic-Kattegat-Skaggerak estuarine system: a regional salinity proxy". Boreas. 40 (2): 242–255. doi:10.1111/j.1502-3885.2010.00193.x.
  143. ^ Mertens, K.N.; Bringué, M.; Van Nieuwenhove, N.; Takano, Y.; Pospelova, V.; Rochon, A.; de Vernal, A.; Radi, T.; Dale, B.; Patterson, R.T.; Weckström, K.; Andrén, E.; Louwye, S.; Matsuoka, K. (2012). "Process length variation of the cyst of the dinoflagellate Protoceratium reticulatum in the North Pacific and Baltic-Skagerrak region: calibration as annual density proxy and first evidence of pseudo-cryptic speciation". Journal of Quaternary Science. 27 (7): 734–744. Bibcode:2012JQS....27..734M. doi:10.1002/jqs.2564.
  144. ^ Ellegaard, M; Lewis, J; Harding, I (2002). "Cyst-theca relationship, life cycle, and effects of temperature and salinity on the cyst morphology of Gonyaulax baltica sp. nov. (Dinophyceae) from the Baltic Sea area". Journal of Phycology. 38 (4): 775–789. doi:10.1046/j.1529-8817.2002.01062.x.
  145. ^ Rochon, A; Lewis, J; Ellegaard; Harding, IC (2009). "The Gonyaulax spinifera (Dinophyceae) "complex" : perpetuating the paradox ?". Review of Palaeobotany and Palynology. 155 (1–2): 52–60. doi:10.1016/j.revpalbo.2008.12.017.
  146. ^ Zonneveld, Karin A.F.; Susek, Ewa (2007). "Effect of temperature, light and salinity on cyst production and morphology of Tuberculodinium vancampoae (Rossignol 1962) Wall 1967 (Pyrophacus steinii (Schiller 1935) Wall et Dale 1971)". Review of Palaeobotany and Palynology. 145 (1–2): 77–88. doi:10.1016/j.revpalbo.2006.09.001.
  147. ^ Verleye, T.J.; Mertens, K.N.; Louwye, S.; Arz, H.W. (2009). "Holocene salinity changes in the southwestern black sea: a reconstruction based on dinoflagellate cysts" (PDF). Palynology. 33: 77–100. doi:10.2113/gspalynol.33.1.77.
  148. ^ Mertens, K.N., Bradley, L.R., Takano, Y., Mudie, P.J., Marret, F., Aksu, A.E., Hiscott, R.N., Verleye, T.J., Mousing, E.A., Smyrnova, L.L., Bagheri, S., Mansor, M., Pospelova, V. & Matsuoka, K. (in press). Quantitative estimation of Holocene surface salinity variation in the Black Sea using dinoflagellate cyst process length. Quaternary Science Reviews
  149. ^ MacRae, R.A.; Fensome, R.A.; Williams, G.L. (1996). "Fossil dinoflagellate diversity, originations, and extinctions and their significance". Can. J. Bot. 74 (11): 1687–1694. doi:10.1139/b96-205.
  150. ^ Moldowan, J.M. and Talyzina, N.M., Biogeochemical evidence for dinoflagellate ancestors in the Early Cambrian. Science 281, 1168-1170.
  151. ^ Sarjeant, W.A.S. (1978). "Arpylorus antiquus Calandra emend., a dinoflagellate cyst from the upper Silurian". Palynology. 2: 167–179. doi:10.1080/01916122.1978.9989171.
  152. ^ LeHerissé, A., Masure, E., Al Ruwaili, M., Massa, D., 2000. Revision of Arpylorus antiquus from the Silurian: the end of a myth. In: Wang, W., Quyang, S., Sun, X., Yu, G. (Eds.), Abstracts 10th International Palynological Congress, Nanjing. National Natural Science Foundation of China, p. 88.
  153. ^ Vozzhennikova, T.F., Shegeshova, L.I., 1989. Palaeodinophysis gen. et sp. N. from the Devonian of the Rudnyy Altay (a unique find of dinoflagellate fossils), Doklady Akademii Nauk SSSR 307, 442–445 (in Russian).
  154. ^ Fensome, Robert A.; Saldarriaga, Juan F.; Taylor, "Max" F. J. R. (1999). "Dinoflagellate phylogeny revisited: Reconciling morphological and molecular based phylogenies". Grana. 38 (2–3): 66–80. doi:10.1080/00173139908559216.
  155. ^ Powell, A. J. (ed.), 1992: A Stratigraphic Index of Dinoflagellate Cysts. London: Chapman & Hall, 300 pp.
  156. ^ Williams, G.L., Stover, L.E., & Kidson, E.J., 1993: Morphology and stratigraphic ranges of selected Mesozoic-Cenozoic dinoflagellate taxa in the northern hemisphere. Geological Survey of Canada, Paper. 92-10 , 137 pp., 2 pl.
  157. ^ De Schepper, S.; Head, M.J. (2008). "Age calibration of dinoflagellate cyst and acritarch events in the Pliocene–Pleistocene of the eastern North Atlantic (DSDP Hole 610A)". Stratigraphy. 5 (2): 137–161.
  158. ^ Louwye, S.; Head, M.J.; De Schepper, S. (2004). "Dinoflagellate cyst stratigraphy and palaeoecology of the Pliocene in northern Belgium, southern North Sea Basin". Geological Magazine. 141 (3): 353–378. Bibcode:2004GeoM..141..353L. doi:10.1017/s0016756804009136.
  159. ^ Jiménez-Moreno, G.; Head, M.J.; Harzhauser, M. (2006). "Early and Middle Miocene dinoflagellate cyst stratigraphy of the central Paratethys, central Europe" (PDF). Journal of Micropalaeontology. 25 (2): 113–139. doi:10.1144/jm.25.2.113.
  160. ^ Riding, J.B.; Kyffin-Hughes, J.E. (2004). "A review of the laboratory preparation of palynomorphs with a description of an effective non-acid technique". Revista Brasileira de Paleontologia. 7 (1): 13–44. doi:10.4072/rbp.2004.1.02.
  161. ^ Riding, J.B.; Kyffin-Hughes, J.E.; Owens, B. (2007). "An effective palynological preparation procedure using hydrogen peroxide" (PDF). Palynology. 31: 19–36. doi:10.2113/gspalynol.31.1.19.
  162. ^ Riding, J.B.; Kyffin-Hughes, J.E. (2009). "The use of pre-treatments in palynological processing" (PDF). Review of Palaeobotany and Palynology. 158 (3–4): 281–290. doi:10.1016/j.revpalbo.2009.09.009.
  163. ^ Riding, J.B.; Kyffin-Hughes, J.E. (2011). "A direct comparison of three palynological preparation techniques" (PDF). Review of Palaeobotany and Palynology. 167 (3–4): 212–221. doi:10.1016/j.revpalbo.2011.07.008.
  164. ^ Head, M.J.; Lewis, J.; de Vernal, A. (2006). "The cyst of the calcareous dinoflagellate Scrippsiella trifida: resolving the fossil record of its organic wall with that of Alexandrium tamarense". Journal of Paleontology. 80 (1): 1–18. doi:10.1666/0022-3360(2006)080[0001:tcotcd]2.0.co;2.
  165. ^ Stockmarr, J (1971). "Tablets with spores used in absolute pollen analysis". Pollen et Spores. 13: 615–621.
  166. ^ Mertens, K.N.; Verhoeven, K.; Verleye, T.; Louwye, S.; Amorim, A.; Ribeiro, S.; Deaf, A.S.; Harding, I.C.; DE Schepper, S.; GonzÁLEZ, C.; Kodrans-NSIAH, M.; DE Vernal, A.; Henry, M.; Radi, T.; Dybkjaer, K.; Poulsen, N.E.; Feist-BURKHARDT, S.; Chitolie, J.; Heilmann-CLAUSEN, C.; Londeix, L.; Turon, J.-L.; Marret, F.; Matthiessen, J.; Mccarthy, F.M.G.; Prasad, V.; Pospelova, V.; Hughes, J.E.K.; Riding, J.B.; Rochon, A.; Sangiorgi, F.; Welters, N.; Sinclair, N.; Thun, C.; Soliman, A.; VAN Nieuwenhove, N.; Vink, A.; Young, M. (2009). "The absolute abundance calibration project: the Lycopodium marker-grain method put to the test" (PDF). Review of Palaeobotany and Palynology. 157 (3–4): 238–252. doi:10.1016/j.revpalbo.2009.05.004.
  167. ^ Mertens, K.N.; et al. (2012). "Determining the absolute abundance of dinoflagellate cysts in recent marine sediments II: Further tests of the Lycopodium…". Review of Palaeobotany and Palynology. 184: 74–81. doi:10.1016/j.revpalbo.2012.06.012.
  168. ^ Wall, D (1971). "Biological problems concerning fossilizable dinoflagellates". Geoscience and Man. 3: 1–15. doi:10.1080/00721395.1971.9989704.
  169. ^ Anderson, D.M.; Wall, D. (1978). "Potential importance of benthic cysts of Gonyaulax tamarensis and G. excavata in initiating toxic dinoflagellate blooms". Journal of Phycology. 14 (2): 224–234. doi:10.1111/j.1529-8817.1978.tb02452.x.
  170. ^ FRYXELL G.A. 1983. Introduction. In: Survival strategies of the algae (Ed. by A. Fryxell), pp. 1–22, Cambridge University Press, Cambridge, U.K.

External links[edit]