Dinocyst
Dinocysts or dinoflagellate cysts are typically 15 to 100 µm in diameter and produced by around 15–20% 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. Many books provide overviews on dinocysts.[1]
History
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.[2]
A first relation between dinoflagellate thecae and cysts was made through morphological comparison of both by Bill Evitt and Susan E. Davidson.[3] Further evidence came from detailed culture studies of dinoflagellate cysts by David Wall and Barrie Dale at Woods Hole Oceanographic Institution in the sixties.[4][5]
Types of cysts
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.[6] 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 .[7][8]
Division cysts refer to non-motile division stages wherein asexual reproduction takes place through division.[9] 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.[10]
Taxonomy
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.[11][12][13][14] 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.[15] Another method, rarely used, uses a sucrose gradient.[16] Recent times have brought about the possibility to get molecular sequences from single cysts or single cells.[17][18][19] The proportion of cyst-forming species for marine dinoflagellates is between 15 and 20%[20] and for freshwater dinoflagellates 24%.[21] 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.[22] It has previously been suggested that morphological characters from the cyst stage may be phylogenetically important in marine species[23] and this may to an even greater extent be the case for freshwater dinoflagellates,[24] confirmed by new observations[25][26] and recently reviewed.[21] Several books document general cyst taxonomy.[22][27] There are few guides for determination of marine Quaternary dinocysts.[28][29] Many new species are still being described for the Neogene,[30] which covers the Miocene,[31][32] the Pliocene[33][34][35][36] and the Quaternary, which covers the Pleistocene[37] and recent.[38][39][40]
Size
Quaternary dinocysts are typically between 15 and 100 µm in diameter.[41] One of the smallest recent cysts is the cyst of Pentapharsodinium dalei, which can be as small as 19 µm in length.[42] One of the largest recent cysts is the cyst of Protoperidinium latissimum, which can be as large as 100 µm in length.[5]
Composition
The walls of organic-walled dinocysts are composed of the resistant biopolymer called dinosporin.[43] 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
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.[44] 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.[22]
TEM studies (e.g.[45]) suggest that endophragm and periphragm are not morphologically separable. Therefore the use of the terms pedium and luxuria are suggested instead.[46] Within the cyst wall, a thick cellulose-like layer called the endospore is present which is birefringent under crossed nichols.[47] 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 it should be noted that one large group of dinoflagellates (athecate - or naked dinoflagellates) do not have thecal plates and therefore produce cysts lacking all forms of reflected tabulation.[48]
Cyst ultrastructure
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[45][49][50] and more recent work on Lingulodinium machaerophorum[51] and Alexandrium.[52]
Some freshwater cysts have been investigated with TEM, such as Ceratium hirundinella.[53]
Relation to life cycle
Resting cysts are traditionally associated with the sexual cycle of dinoflagellates.[54] Induced by particular triggers such as changes in temperature, nutrients,[55] 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.[56] Resting stages also constitute a reservoir of genetic diversity, which increases the survival potential of the populations.[57] 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.[58] Sediment can be stored with live Lingulodinium cysts for at least 18 months.[59] Cysts often need triggers to germinate ('excyst'), such as changes in temperature, nutrients, etc. Some cysts, such as Scrippsiella trochoidea, require light to germinate.[60]
Distribution, biogeography and ecology of organic-walled dinocysts
Dinocyst distribution is mainly studied through studies of surface sediments.[61] Many studies are regional, such as the Iberian Margin[62] the North Sea,[63] Kiel bight,[64] Celtic Sea,[65] Norwegian Sea,[66] around Iceland,[67] the Southeast Pacific,[68] the Arctic,[69][70] Equatorial Atlantic,[71] South and Equatorial Atlantic,[72] off West Africa,[73] the Southern Ocean,[74] Benguela upwelling,[75] in the Mediterranean Sea,[76] Caspian Sea,[77] British Columbia,[78] The Northeastern Pacific,[79] Florida,[80] Mexico[81] and Barends Sea.[82]
Such surface sediment studies show that dinoflagellate cyst distribution is controlled by ranges of temperature, salinity and nutrients.[83] This often poses biogeographical boundaries, more particularly temperature.[84] Some species can be clearly related to cold waters.[85] 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.[86] Other species are thermophilic, such as the "living fossil" Dapsilidinium pastielsii currently found in the Indo-Pacific Warm Pool only.[87]
Eutrophication can also be reflected in dinocyst assemblages.[88][89][90]
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.[61] Cyst are also often transported from the inner shelf to the outer shelf or slope.[61]
Another problem with cysts is that they also get transported with ballast water, which can cause introduction of invasive species.[91]
Seasonality and fluxes are studied through sediment trap studies, which help to understand ecological signals.[92][93][94][95][96][97]
Palaeoecology of organic-walled dinocysts
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,[98][99][100][101][102] temperature,[103][104][105] salinity[106][107][108] and ice cover.[109][110][111]
Such reconstructions can be done via semi-quantitative techniques, such as ordination techniques,[48] which can indicate trends in environmental parameters.
A quantitative method is the use of transfer functions,[112][113][114][115][116] although these have been heavily debated.[117][118]
Another late Quaternary application is for environmental goals, more particularly the study of eutrophication[119][120][121] .[122]
An interval of particular interest during the late Quaternary is the Eemian.[123][124][125][126][127]
Also during the Neogene, dinocysts have shown to be useful in the Miocene[128] and particularly the Messinian.[129] Also the paleoclimate of the Pliocene has been investigated.[130][131][132] Transfer functions have also been attempted during the Pliocene.[133] Some species have been suggested to have different environmental preferences during the Neogene.[134]
During the Paleogene dinocysts also are particularly useful,[135] and then more particularly the Eocene.[136][137]
The palaeoecology of freshwater dinoflagellate cysts is relatively unexplored, though several recent studies have shown the relation to changes in nutrients, pH and temperature[138][139][140][141] and is recently reviewed.[21]
Morphological variation of organic-walled dinocysts
There is little known about how organic-walled dinocysts are formed except from culture experiments.[142] Cyst formation is suggested to happen through self-assembly processes.[143]
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,[144] and study of surface sediments.[145] Also variations in the morphology of the species Operculodinium centrocarpum [146][147] can be related to salinity and/or temperature. Also cysts of the species Gonyaulax baltica shows morphological variations in culture,[148] as well as Gonyaulax spinifera.[149] Cyst formed by other species such as Pyrophacus steinii (cyst is called Tuberculodinium vancampoae) do not show a clear relation to variations in salinity.[150]
The morphological variation can be applied for the reconstruction of salinity, in a semi-quantitative[151] or quantitative way.[146] Process length variation of Lingulodinium machaerophorum has been used to reconstruct Black Sea salinity variation.[152]
Biostratigraphy and evolution of organic-walled dinocysts
Organic-walled dinoflagellate cysts have a long geological record with lowest occurrences during the mid Triassic,[153] whilst geochemical markers suggest a presence to the Early Cambrian.[154] 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,[155] but this palynomorph is now considered probably an arthropod remain.[156] Another enigmatic form with possible early dinoflagellate affinity is Palaeodinophysis altaica, which was found in the Devonian of Kazakhstan.[157]
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.[22] The biostratigraphical application of dinoflagellate cysts has been thoroughly studied.[158][159] The Pliocene has been recently investigated[160][161] and also the Miocene.[162] Also for the Quaternary there have been further studies.[163]
Palynological methods
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.[164][165][166][167] 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.[168] The concentration of Dinocysts can be quantified by adding an exotic spike or marker such as Lycopodium clavatum spores.[169][170][171]
Biological functions
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.[172][173][174] These functions have implications for the population dynamics, seasonal succession, genetic diversity, and biogeography of dinoflagellates.
References
- ↑ Evitt, W. R. 1985. Sporopollenin Dinoflagellate Cysts: Their Morphology and Interpretation. American Association Stratigraphic Palynologists Monograph Ser. 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.
- ↑ Evitt, W.R. and Davidson, S.E. 1964. Dinoflagellate studies. 1. Dinoflagellate cysts and thecae. Stanford university publications X (1), pp. 3–12.
- ↑ Wall, D.; Dale, B. (1966). "Living" fossils in western Atlantic plankton". Nature. 211 (5053): 1025–1026. doi:10.1038/2111025a0.
- 1 2 Wall, D.; Dale, B. (1968). "Modern dinoflagellate cysts and evolution of the Peridiniales". Micropaleontology. 14 (3): 265–304. doi:10.2307/1484690.
- ↑ 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.
- ↑ 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: 1–36. doi:10.2307/1485525.
- ↑ 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.
- ↑ 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.
- ↑ 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.
- ↑ Wall, D.; Dale, B. (1966). "Living fossils" in Atlantic plankton". Nature. 211 (5053): 1025–1026. doi:10.1038/2111025a0.
- ↑ Wall, D.; Dale, B. (1968). "Modern dinoflagellate cysts and evolution of the Peridiniales". Micropaleontology. 14: 265–304. doi:10.2307/1484690.
- ↑ 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: 149–177. doi:10.1515/botm.1997.40.1-6.149.
- ↑ 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. doi:10.1016/j.marmicro.2012.08.002.
- ↑ Bolch, C.J.S. (1997). "The use of polytungstate for the separation and concentration of living dinoflagellate cysts from marine sediments". Phycologia. 37: 472–478.
- ↑ 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: 588–592. doi:10.4319/lo.1991.36.3.0588.
- ↑ 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.
- ↑ 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.
- ↑ 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: 259–267. doi:10.1111/j.1440-1835.2009.00545.x.
- ↑ 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.
- 1 2 3 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.
- 1 2 3 4 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.
- ↑ Harland, R (1982). "A review of Recent and Quaternary organic-walled dinoflagellate cysts of the genus Protoperidinium". Palaeontology. 25: 369–397.
- ↑ Schilling, A.J. (1891). "Die Süsswasser-Peridineen". Flora oder Allgemeine botanische Zeitung. 74: 220–299.
- ↑ 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: 1–10.
- ↑ Moestrup, Ø.; Lindberg, K.; Daugbjerg, N. (2009). "Studies on woloszynskioid dinoflagellates IV: The genus Biecheleria gen. nov". Phycological Research. 57: 203–220. doi:10.1111/j.1440-1835.2009.00540.x.
- ↑ 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.
- ↑ 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.
- ↑ 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.
- ↑ 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.
- ↑ 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.
- ↑ Soliman, A., Head, M.J., and Louwye, S. In press. Morphology and distribution of the Miocene dinoflagellate cyst Operculodinium? borgerholtense Louwye 2001, emend. Palynology.
- ↑ 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.
- ↑ 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.
- ↑ 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: 625–644. doi:10.1666/0022-3360(2004)078<0625:ndcais>2.0.co;2.
- ↑ 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.
- ↑ Head, M.J. (2002). "Echinidinium zonneveldiae sp. nov., a new dinoflagellate cyst from the Late Pleistocene of the Baltic region". Journal of Micropalaeontology. 21: 169–173. doi:10.1144/jm.21.2.169.
- ↑ 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: 65–83. doi:10.1016/j.marmicro.2010.10.001.
- ↑ 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: 593–601. doi:10.1046/j.1529-8817.2002.01206.x.
- ↑ 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. doi:10.1016/j.marmicro.2012.08.002.
- ↑ De Vernal, A.; Marret, F. (2007). "Organic-Walled Dinoflagellate Cysts: Tracers of Sea-Surface Conditions". Developments in Marine Geology: 371–408. doi:10.1016/S1572-5480(07)01014-7.
- ↑ Dale, B (1977). "New observations on Peridinium faeroense Paulsen (1905), and classification of small orthoperidinioid dinoflagellates". Br. Phycol. J. 12: 241–253. doi:10.1080/00071617700650261.
- ↑ 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. .
- ↑ DE, VERTEUIL L.; Norris, G. (1996). "Part 2. Homology and structure in dinoflagellate cyst terminology". Micropaleontology. S42: 83–172.
- 1 2 Jux, U (1968). "Über den feinbau der wandung bei Hystrichosphaera bentori Rossignol 1961". Palaeontographica Abt. B. 123 (1–6): 147–152.
- ↑ 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.
- ↑ 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.
- 1 2 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.
- ↑ 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.
- ↑ 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.
- ↑ 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: 265–288. doi:10.1016/s0146-6380(97)00134-4.
- ↑ Kennaway, Gay; Lewis, Jane (2004). "An ultrastructural study of hypnozygotes of Alexandrium species (Dinophyceae)". Phycologia. 43: 353–363.
- ↑ Chapman, D. V.; Dodge, J. D.; Heaney, S. I. (1982). "CYST FORMATION IN THE FRESHWATER DINOFLAGELLATE CERATIUM HIRUNDINELLA (DINOPHYCEAE)". Journal of Phycology. 18: 121–129. doi:10.1111/j.1529-8817.1982.tb03165.x.
- ↑ Stosch, H.A. VON (1965). "Sexualität bei Ceratium cornutum (Dinophyta)". Die Naturwissenschaften. 52: 112–113. doi:10.1007/bf00626331.
- ↑ Pfiester, L. A. & Anderson, D. M. 1987. Dinoflagellate reproduction. In: The biology of dinoflagellates (ed. F. J. R. Taylor), pp. 611–648. - Blackwell, Oxford.
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- ↑ 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: D22108. Bibcode:2009JGRD..11422108C. doi:10.1029/2009JD012077.
- ↑ 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: 315–318. doi:10.1007/s10933-006-9001-1.
- ↑ 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.
- ↑ Hemsley, A.R.; Lewis, J.; Griffiths, P.C. (2004). "Soft and sticky development : some underlying reasons for microarchitectural pattern convergence". Review Palaeobotany & Palynology. 130: 105–119. doi:10.1016/j.revpalbo.2003.12.004.
- ↑ Hallett, R.I., 1999. Consequences of environmental change on the growth and morphology of Lingulodinium polyedrum (Dinophyceae) in culture. Ph.D. thesis. University of Westminster, 109 pp.
- ↑ 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: 54–69. doi:10.1016/j.marmicro.2008.10.004.
- 1 2 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: 242–255. doi:10.1111/j.1502-3885.2010.00193.x.
- ↑ 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: 734–744. doi:10.1002/jqs.2564.
- ↑ 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: 775–789. doi:10.1046/j.1529-8817.2002.01062.x.
- ↑ Rochon, A; Lewis, J; Ellegaard; Harding, IC (2009). "The Gonyaulax spinifera (Dinophyceae) "complex" : perpetuating the paradox ?". Review of Palaeobotany and Palynology. 155: 52–60. doi:10.1016/j.revpalbo.2008.12.017.
- ↑ 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.
- ↑ 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". Palynology. 33: 77–100. doi:10.2113/gspalynol.33.1.77.
- ↑ 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
- ↑ MacRae, R.A.; Fensome, R.A.; Williams, G.L. (1996). "Fossil dinoflagellate diversity, originations, and extinctions and their significance". Can. J. Bot. 74: 1687–1694. doi:10.1139/b96-205.
- ↑ Moldowan, J.M. and Talyzina, N.M., Biogeochemical evidence for dinoflagellate ancestors in the Early Cambrian. Science 281, 1168-1170.
- ↑ 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.
- ↑ 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.
- ↑ 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).
- ↑ Powell, A. J. (ed.), 1992: A Stratigraphic Index of Dinoflagellate Cysts. London: Chapman & Hall, 300 pp.
- ↑ 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.
- ↑ 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.
- ↑ 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: 353–378. doi:10.1017/s0016756804009136.
- ↑ Jiménez-Moreno, G.; Head, M.J.; Harzhauser, M. (2006). "Early and Middle Miocene dinoflagellate cyst stratigraphy of the central Paratethys, central Europe". Journal of Micropalaeontology. 25: 113–139. doi:10.1144/jm.25.2.113.
- ↑ Matthießen, J.; Knies, J.; Nowaczyk, N.; Stein, R. (2001). "Late Quaternary dinoflagellate cyst stratigraphy at the Eurasian continental margin, Arctic Ocean: Indications for Atlantic water inflow in the past 150,000 years". Global and Planetary Change. 31 (1): 65–86. doi:10.1016/S0921-8181(01)00113-8.
- ↑ 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.
- ↑ Riding, J.B.; Kyffin-Hughes, J.E.; Owens, B. (2007). "An effective palynological preparation procedure using hydrogen peroxide". Palynology. 31: 19–36. doi:10.2113/gspalynol.31.1.19.
- ↑ Riding, J.B.; Kyffin-Hughes, J.E. (2009). "The use of pre-treatments in palynological processing". Review of Palaeobotany and Palynology. 158 (3–4): 281–290. doi:10.1016/j.revpalbo.2009.09.009.
- ↑ Riding, J.B.; Kyffin-Hughes, J.E. (2011). "A direct comparison of three palynological preparation techniques". Review of Palaeobotany and Palynology. 167 (3–4): 212–221. doi:10.1016/j.revpalbo.2011.07.008.
- ↑ 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.
- ↑ Stockmarr, J (1971). "Tablets with spores used in absolute pollen analysis". Pollen et Spores. 13: 615–621.
- ↑ 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". Review of Palaeobotany and Palynology. 157: 238–252. doi:10.1016/j.revpalbo.2009.05.004.
- ↑ 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.
- ↑ Wall, D (1971). "Biological problems concerning fossilizable dinoflagellates". Geoscience and Man. 3: 1–15. doi:10.1080/00721395.1971.9989704.
- ↑ 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: 224–234. doi:10.1111/j.1529-8817.1978.tb02452.x.
- ↑ 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
- AASP — THE PALYNOLOGICAL SOCIETY
- L'Association des Palynologues de Langue Française
- Canadian Association of Palynologists
- The Micropalaeontological Society, Palynology group
- Quaternary Dinoflagellate Cyst Association
- Dinoflaj2 Rob Fensome's and Graham Williams' Database on cysts
- Yasuo Fukuyo's site on motile stages and their cysts
- Dino6 meeting
- Dino8 meeting, Montreal
- Dino9 meeting, Liverpool
- Marine Micropaleontology (Journal)
- Review of Palaeobotany and Palynology (Journal)
- Journal of Micropalaeontology (Journal)
- Micropaleontology (Journal)