Dinoflagellata phylogeny

From Palaeos

Eukarya
  |--+--Fungi
  |  `--Metazoa
  |--Amoebozoa
  `--+--Rhizaria
     |--Metamonada
     `--o--Plantae
        `--o--Chromista
           `--Alveolata
                |--Ciliophora
                `--Miozoa
                     |--Dinozoa
                     |    `--Dinoflagellata
                     `--Apicomplexa

[edit] Phylogeny and Evolution

[edit] Affinities

...

Some dinoflagellates photosynthesise; they generally possess chlorophyll a and variants of c, and other pigments including carotenes and xanthins. Others, however, are heterotrophic.

Two other phyla thought to be closely related to dinoflagellates are the Ciliophora and the Apicomplexa.

"Ideas on dinoflagellate evolution have been developed by, or summarized in, Taylor (1980), Tappan (1980), Bujak & Williams (1981), Loeblich III (1984) and Goodman (1987). A possible scenario for dinoflagellates, proposed by Fensome, Taylor et al. (1993) is shown in Text-Figure 60.

"From cytological and biochemical evidence, dinoflagellates appear to be an ancient group of protists, most authorities now believing them to have originated in the Late Precambrian (Taylor 1978, 1980; Loeblich III 1984). These earliest dinoflagellates either produced no preservable cysts or generated cysts (acritarchs) whose morphology does not demonstrate their affinity (see Downie 1973; Sarjeant 1974). A study of openings and process distribution in Early Paleozoic acritarchs led Lister (1970) to conclude that some may be the cysts of thecate dinoflagellates. However, the tabulations produced by Lister were speculative, and not convincingly similar to any Mesozoic-Cenozoic or modem tabulations.

"Most workers have accepted that the Late Silurian genus Arpylorus is a dinoflagellate cyst (Calandra 1964; Evitt in van Oyen 1964; Sarjeant 1978b; Stover & Evitt 1978; Lentin & Williams 1981; for a contrary view, see Bujak & Williams 1981). It clearly has plates that can reasonably be interpreted as thecal. However, like Lister’s tabulations, they do not closely resemble any Mesozoic-Cenozoic or modem tabulations’ and the cingulum and sulcus are not prominent as they are in later dinoflagellates. Perhaps Arpylorus offers a fleeting glimpse of an earlier, Paleozoic radiation of dinoflagellates. Possibly more closely comparable with a group of modem dinoflagellates is the Devonian genus Palaeodinophysis (Vozzhennikova & Sheshegova 1989). There is at least a superficial similarity between Palaeodinophysis and living dinophysialeans (as well as fossil nannoceratopsialeans) and, if its dinophysialean affinity and stratigraphic distribution are confirmed by future studies, the evolutionary scenario for Mesozoic to Recent dinoflagellates as provided below will require modification;

"Early dinoflagellates may have had a temporary dinokaryon (see Fensome, Taylor et al. 1993), but there is, of course, no proof of this in the fossil record. The temporary dinokaryon of blastodinialeans (e.g. Text-Fig. 3K) and noctilucaleans (e.g. Text-Fig. 3Q, R, T, U) is possibly a relict feature, but living blastodinialeans and noctilucaleans are highly specialized and, apart from their nucleus, are not good models for primitive dinoflagellates" (Fensome et al. 1996, p. 157).

[edit] Fossil Record

Dinoflagellates have left a rich, if taxonomically selective, fossil record of organic-walled, calcareous and rare siliceous forms, almost exclusively cysts. Insofar as body fossils are concerned, the record begins with the single occurence of Arpylorus antiquua, in the Silurian of Tunisia. After that, there is nothing until the Triassic, when fossils begin to become common. By the Jurassic, the group is well-known, well-established, and morphologically diverse.

"Fossil dinoflagellates occur primarily in strata of Late Triassic to Recent age. They are mostly of marine origin, but some fresh water fossils are known. As already noted, most fossil dinoflagellates appear to represent resting cysts or hypnozygotes (termed dinocysts by some workers and, in this work, hereafter referred to as cysts). A cyst becomes fossilizable if one or more wall layers are impregnated with a resistant organic or inorganic substance. Most fossil dinoflagellate cysts have organic walls comprising dinosporin. Calcareous and siliceous cysts may have a fossilizable organic component in their wall, and some "organic-walled" fossil cysts in palynological preparations may represent the organic linings of calcareous cysts (Lentin 1985; Hultberg 1985). Such fossils are thus somewhat analogous to the organic linings of foraminifera" (Fensome et al. 1996, p. 124).

"Cysts are produced inside the dinoflagellate theca (with one possible partial exception, Palaeoperidinium, which is discussed below). Cyst shape may approximate that of the motile cell, involving no long protrusions unrelated to thecal shape; such cysts are termed proximate (see Sarjeant 1982c; Text-Fig. 22; PI. 1, Fig. 1-5). Alternatively, the cyst may comprise a more or less spherical central body with processes or crests (PI. 1, Fig. 6-16); such cysts are termed chorate or proximochorate, depending upon the height of the extensions relative to the central body. Although there is a morphological gradation between proximate, proximochorate and chorate cysts, these terms are useful in descriptions" (Fensome et al. 1996, p. 124).

"The Late Silurian species Arpylorus antiquus provides further evidence of cyst formation during only a limited interval of geologic time. Alone in all the Paleozoic, Arpylorus appears to this author to be a very dinoflagellate-like dinoflagellate - so much so, in fact, that were it to be found in a Mesozoic assemblage, it might attract no more attention than any other distinctive species. Unlike other Paleozoic microfossils discussed later in this chapter that may also represent dinoflagellate cysts but lack a minimum of features that would establish their affinity with relative certainty, A. antiquus was described (Calandra, 1964) as a dinoflagellate because it looks like one. Restudy of the type material from subsurface Algeria by Evitt (1967) and Sarjeant (1978) led these authors to reaffirm the basic identification, although the original material is not ideal, and their interpretations of it are not identical. However, the dinoflagellate nature of this fossil is not unquestioned. Bujak and Williams (1981) and Bujak and Davies (1983) have urged an open mind on its identification and suggested it may not be a dinoflagellate. Beyond that, they discount its bearing on the matter of a selective dinoflagellate fossil record. It is clear that one occurrence of such a potentially important fossil, consisting of less than perfectly preserved specimens, is insufficient to resolve the matter satisfactorily. The need is for new material, including better, or at least differently, preserved specimens and coming from another area, which will enable the characters of this fossil species to be determined afresh.

"But it is not the recovery of a dinoflagellate from Silurian strata that is surprising, for we have already considered the biological reasons to believe that dinoflagellates were probably present in the Precambrian. What is spectacular in this case is the absence of fossil dinoflagellates from younger Paleozoic strata. After the Silurian, there are no other fossils definitely identifiable as dinoflagellates for about 200 million years, through all the rest of the Paleozoic and part of the Triassic. This is a span of time approximately equal to the entire subsequent and essentially continuous fossil record of dinoflagellates from the Carnian to the present. In light of what we now know about the production of preservable cysts among modern dinoflagellates, we can probably best regard A. antiquus as an especially "precocious" species, which carried out a successful early experiment in sporopollenin production long before that technique really "caught on" as the "fashionable" dinoflagellate thing to do in the early Mesozoic" (Evitt 1985, p. 38).

[edit] Origins

Although body fossils of dinoflagellates are not recognised until the Silurian, several lines of evidence have indicated that dinoflagellates originated in the Neoproterozoic (Knoll 1996).

RNA molecular sequencing and examination of mitochondrial cristae of modern organisms (Lipps 1993) suggest that dinoflagellates are older than Foraminifera and Radiolaria, which have been found in Cambrian rocks.

Biochemical studies confirm the presence of dinoflagellate-specific biomarkers (dinosteranes and 4a-methyl-24-ethylcholestane) at least as early as the earliest Cambrian. Reports include:

Proterozoic Bitter Springs and Pertatataka Formations, central Australia (Summons & Walter 1990);

Nonesuch Formation, North American mid-continent rift (Pratt et al. 1991);

lower part of the Upper Riphean (Neoproterozoic) Visingsö Beds, Sweden (ref?);

Atdabanian (Early Cambrian), in glauconitic clays from the Lükati Formation of Estonia (Moldowan & Talyzina 1998);

Buen Formation in northern Greenland (ref?).

(After Moldowan & Talyzina 1998.)

"Biomarkers are organic molecules that are stable at moderate temperatures, which can be preserved in rocks even when recognizable fossils are absent" (Moldowan & Talyzina 1998, p. 1169). The dinosterane biomarkers have a carbon structure which occurs in sterols found in high concentrations in numerous modern dinoflagellate species, but has rarely been found in other taxa.

From the Emsian age (late Early Devonian) Battery Point Formation, Cap-aux-Os Member exposed at Gaspé Bay, Quebec, approximately ten species of acritarchs have been recovered, including Veryhachium, Helosphaeridium, Micrhystridium, Multiplicisphaeridium and Gorgonosphaeridium. "Most are thought to represent cysts of marine phytoplankton (Strother 1996); recent geochemical analyses suggest that many may represent dinoflagellates (Moldowan and Talyzina 1998)" (Hotton et al. 2001, p. 195b).

"The presence of dinoflagellate relatives among acritarchs explains the continuous record of dinosteroids from Precambrian to Cenozoic source rocks from numerous localities world-wide" (Moldowan & Talyzina 1998, p. 1170).

"Some acritarchs resemble dinoflagellate cysts (Margulis & Schwartz 1982; Tappan 1980; Mendelson 1993), but they do not show paratabulation and they have excystments that are different from classical archeopyles of recognised Mesozoic and younger dinocysts. Many acritarch specimens have no excystment structure. However, most modern dinocysts reach sediments before germination (Anderson et al. 1985), and some of these can fossilize without excystment structure formation. Some Ordovician acanthomorphic acritarchs have a double-wall structure (Martin & Kjellström 1973) comparable with that of dinoflagellate cysts. Certain cysts of living dinoflagellates from the order Gymnodiniales lack clearly defined archeopyles or reflected tabulation (Wall & Dale 1968). ... [But, on balance,] the morphological evidence has not been sufficient to establish links between acritarchs and dinoflagellates" (Moldowan & Talyzina 1998, pp. 1168-1169).

[edit] Evolution

"The fossilized matter available for paleontological investigation represents less than 1% of organisms that once existed on Earth. A high abundance of related specimens in a particular age suggests that there was an earlier radiation. Various kinds of simply structured, rounded acritarchs and dinoflagellate biomarkers coexist in upper Riphean rocks, although the dinoflagellate affinity of any particular Proterozoic genus requires further investigation.

"Dinosterane-containing acanthomorphic acritarchs are widespread in Lower Cambrian sediments. These results suggest the evolutionary sequence in which dinoflagellate ancestry originated by the Late Riphean (~800 million years ago); specimens with processes became abundant in the Early Cambrian; and finally, the branch of dinoflagellates with classical archeopyles and paratabulation developed in the Middle Triassic" (Moldowan & Talyzina 1998, p. 1170).

"The fossil record of dinoflagellates appears to show evolutionary patterns similar to those of other groups, such as a major adaptive radiation, which occurred in dinoflagellates in the Late Triassic and Early Jurassic. Should these patterns in the dinoflagellate record be taken as normal, or as curious coincidences? The initial Triassic-Jurassic rapid increase of diversity and its subsequent stability, as indicated by fossils, could be explained by the random or environmentally induced "switching on" and "switching off" of the ability to produce fossilizable cysts by long-ranging Phanerozoic taxa. Furthermore, the observed record does not include important taxa such as the Gymnodiniphycidae (except for Suessia and Dinogymnium, the latter appearing clearly to be a "switched on"-"switched off" ptychodiscalean), Dinophysiales (except possibly for Ternia and Palaeodinophysis), Prorocentrales, Noctilucales, Blastodiniales and Phytodiniales. However, the Mesozoic-Cenozoic fossil record shows a pattern that would be expected of a group undergoing an initial adaptive radiation and subsequent stabilization. It is, therefore, reasonable to believe that the observed pattern reflects a real phenomenon. The isolated Paleozoic occurrences of two possible dinoflagellate genera need to be considered in the context of dinoflagellate phylogeny (see below), but their existence does not diminish the striking nature, or disrupt the general pattern, of the Mesozoic-Cenozoic dinoflagellate fossil record.

"Within the dinoflagellate fossil record, examples of adaptive radiations or episodes of "experimentation" at lower taxonomic levels can be recognized. For example, in the early and middle Cretaceous, peridiniaceans had an "experimental" variety of mostly combination archeopyles; in contrast, most later Cretaceous peridiniaceans had a single plate archeopyle comprising the middorsal intercalary. In a second example, the archeopyle of Middle Jurassic gonyaulacaceans also appears to have undergone a period of experimentation. In the Aalenian and early Bajocian, many of the gonyaulacacean genera possessed multiplate precingular archeopyles: e.g. Durotrigia has a 1-5P archeopyle and Dissiliodinium has a 1-6P archeopyle. From late Bajocian onwards, gonyaulacaceans tended to have apical, single plate precingular, or epitractal archeopyles, the last of these being especially common in the Bathonian to early Oxfordian interval.

"The fossil record of dinoflagellates also reveals excellent examples of morphological stasis. For instance, the tabulation among fossil peridiniaceans shows great stability .The earliest known peridiniaceans have a bipesioid tabulation (Text-Fig. 52C’). The vast majority of Cretaceous and Cenozoic fossil organic-walled and calcareous peridiniaceans show not only the bipesioid stacking of the three middorsal plates, but also the same shapes and interrelationships of these plates. For example, the middorsal anterior intercalary plate (2a) is six-sided (hexa), except in the subfamily Wetzelielloideae, in which it is four-sided (quadra). This stability would perhaps not be so remarkable were it not for the great variation in the episomal tabulations of extant peridiniaceans.

"The question thus arises as to whether the stability in tabulation observed among fossil peridiniacean cysts is real or apparent. Is there more consistency in the tabulation of the cyst than of the theca? Did only past peridiniaceans with a bipesioid tabulation produce cysts (Goodman 1987), with the exception of the siliceous, cinctioid lithoperidinioideans? Or are extant peridiniaceans currently undergoing an episode of experimentation in their tabulation, perhaps stimulated by the environmental rigors or opportunities associated with the Quaternary glaciations? The family Congruentidiaceae, which includes Protoperidinium, and which appears to have arisen from peridiniaceans in Late Cretaceous or earliest Cenozoic times, also shows variation in episomal tabulation in the present day. However, the asymmetry of the archeopyle in such fossil genera as Selenopemphix (Text-Fig. 56K, L) indicates that this family has not had a stable bipesioid fossil history.

"Morphological stasis among fossil dinoflagellates is also exemplified by Gonyaulacysta jurassica, which maintained the same tabulation and general shape within a single species throughout the Middle and Late Jurassic. The related cyst Spiniferites ramosus endured even longer, from the Early Cretaceous to the present. Students of evolutionary theory (e.g. Vrba 1980; Eldredge 1985) have suggested that species with long histories are generalists, whereas those with short histories are more specifically tuned to their environment. Thus, Gonyaulacysta jurassica and Spiniferites ramosus could be visualized as generalists of Middle to Late Jurassic and Cretaceous to Recent seas respectively, whereas species with shorter histories, such as Spiniferites septatus and Alisocysta circumtabulata, may have been more specialized" (Fensome et al. 1996, pp. 156-157).

[edit] Systematics

Relationships within the dinoflagellates are ...

So, what are the apomorphies which we might use to classify fossil cysts?

We cannot say, for sure, though there are some characteristics which we can confidently say are not apomorphies. The nature of the ornamentation – whether chorate or whatever – has long been, for convenience, used to define form taxa at the generic level. Yet we see these characteristics recurring again and again, in lineages which are patently far removed. Intuitively, we realise that gross features like this, which clearly exercise a considerable effect on the life functions (e.g. flotation characteristics) of the organisms, are highly sensitive to evolutionary pressures, and are therefore likely to evolve quickly and repeatedly. Thus it is that the convenient, gross morphological features beloved of stratigraphers, and for so many years the underpinning of dinoflagellate taxonomy, are quite useless indicators of phylogeny.

This may seem obvious today, when words like 'apomorphy' are a standard part of any taxonomists vocabulary, but it was not always so. The writer once ventured the observation that "I consider such features as the clarity with which the cingulum is delimited by sutural or penitabular septa, and indeed the distinction between these two types of ornament, to be relatively unimportant; of infrageneric significance only" (Clowes 1984, p. 29), only to be pilloried by the journal's anonymous referee. Mercifully, the then editor, Doug Nichols, was made of sterner stuff and sought a second opinion. I am grateful to him to this day. Although the paper missed the publication deadline for that volume, and so was delayed by a year, the quoted passage finally appeared without amendment.

... associations of characteristics ...

"Although it is a worthy objective, a widely accepted classification of fossil dinoflagellates at the family level has yet to be devised. Currently, divergent views on principles and criteria are more evident than is any general agreement on results. A comprehensive classification of fossil cysts that originated conceptually with Eisenack (1961) and was elaborated by Sarjeant and Downie (1966) has now been modified by them (Sarjeant and Downie, 1974; Sarjeant, 1974) and by others (Norris, 1978; Tappan, 1980) into several similar arrangements by which fossil cysts are distributed among about 40 families. While based mos.tly on cyst morphology, these families are regarded by Norris, at least, as approaching phylogenetically defensible entities. In contrast, Evitt (1975b) contended that a few modern genera collectively encompass the affinities of a majority of fossil cysts. In line with that view, but with modifications reflecting more recent interpretations, Table 1 .1 lists 13 families, including nine from the hierarchy of modern taxa, that would appear to accommodate the great majority of fossil cysts (admitting that considerable uncertainty must attach to many fossils with highly "generalized" morphology). However, it is not the intent in this volume to pursue the problems of a phylogenetic classification. Instead, with obvious philosophical allegiance to the second approach mentioned above, we will focus attention in Chapter 8 on 17 morphological categories. While they will be defined without strict regard to family boundaries and will include cysts with "generalized" as well as "distinctive" morphology, their approximate correspondence to the families listed at the left in Table 1.1 is shown at the right" (Evitt 1985, p. 27).

[edit] Kingdom? Alveoles [Authority?]

[edit] Phylum* Dinoflagellata Bütschli 1885

1885 Dinoflagellata Bütschli

1914 Pyrrhophyta Pascher

1985 Pyrrhophyta, Evitt, p. 26

1993 Dinoflagellata (Bütschli 1885) Fensome et al., p. ??

• Dinoflagellates are protists - neither plants nor animals. Mercifully, taxonomy has not yet been cursed with an International Code of Protistan Nomenclature (given that the objective is the same, and the issues to be overcome nearly so, it is quite bad enough that there exists separate botanical and zoological codes) so it is necessary to treat dinoflagellates as one or the other, for the purposes of nomenclature. The botanical code has been settled upon, more or less by historical accident. Botanists frequently refer to the phylum-level taxonomic rank as a "division" - another absurd terminological distinction where there is no difference.

Type: [?] [Authority]

Original Diagnosis: xxx

Description: xxx

Habit: xxx

Distribution Occurrence: xxx

Discussion: xxx

Review of sub-ranks, if appropriate...

[edit] Class Dinophyceae [Authority]

Type: [?] [Authority]

Original Diagnosis: xxx

Description: xxx

Habit: xxx

Distribution Occurrence: xxx

Discussion: xxx

Evitt pp. 26-27:

Class DINOPHYCEAE -pyrrhophytes in which one flagellum is whiplike and extends longitudinally, while the second is ribbon-like and follows a circular path in a plane about at right angles to the first. .

Order PROROCENTRALES -dinoflagellates in which the flagella are inserted terminally (desmokont condition), the longitudinal one extending in advance of the cell, and the transverse one encircling the other anterior to the cell body. Some forms have a cellulosic theca of distinctive structure. Preservable resting cysts are unknown and there is no certain fossil record, although the order is conceivably represented by some of the organic-walled fossils currently regarded as acritarchs. Representative genera: Exuviaella (nonthecate), Prorocentrum (thecate).

Note: In all three of the following orders for which the living cell is known, the flagella are inserted laterally (dinokont condition), the longitudinal one extends posteriorly, and both normally lie, at least in part, within channels (the so-called flagellar furrows) defined by various features on the cell surface. Fossil cysts of the extinct fourth order appear to record a similar organization.

Order DINOPHYSIALES - dinoflagellates having the transverse flagellar furrow near the anterior limit of the cell; cell normally shows moderate to strong lateral compression; two lateral plates of the cellulosic theca are much larger than any others. Preservable resting cysts are unknown and there is no unequivocal fossil record. Representative genera: Dinophysis, Ornithocercus.

Order PERIDINIALES - dinoflagellates having the transverse flagellar furrow normally located within the medial third of cell length; theca composed of several tens of cellulosic plates organized in several series paralleling the transverse furrow. Preservable resting cysts are found in some living species and there is an extensive fossil record. Representative living genera: Peridinium, Gonyaulax, Ceratium. Representative fossil genera: Deflandrea, [[Gonyaulacysta]], Odontochitina. Silurian, Triassic-Holocene.

Order GYMNODINIALES - dinoflagellates having the transverse flagellar furrow usually located within the medial third of cell length; cellulosic thecal plates lacking or (rarely) thin, but corresponding vesicles more numerous than typical for Peridiniales, small, and all of about similar size. Preservable resting cysts are known in a few living species. Moderate fossil record of cysts and distinctive sporopollenin coverings of possibly motile cells. Representative living genera: Gymnodinium, Polykrikos. Representative fossil genera: Dinogymnium, ?Distatodinium, ?Suessia. Triassic-Holocene.

Order NANNOCERATOPSIALES - dinoflagellates having the transverse flagellar furrow near anterior extremity of cell; cell compressed laterally as in Dinophysiales; tabulation of inferred theca similar to Peridiniales in anterior part, similar to Dinophysiales in posterior part. Fossil; sole genus: Nannoceratopsis. Jurassic.

Class EBRIOPHYCEAE -nonphotosynthetic, biflagellate, free-Iiving pyrrhophytes, lacking a resistant external covering but having a fossilizable internal siliceous skeleton. Representative genus: Ebria. Geologic range: Cretaceous to Holocene.

Class ELLOBIOPHYCEAE- attached parasitic pyrrhophytes without known fossil record.

Class SYNDINIOPHYCEAE -intracellular parasites without known fossil record.

[edit] Conclusion

xxx

[edit] Further Information

"Significant works on living dinoflagellates include books edited by Spector (1984) and Taylor (1987a) and monographs by Sournia (1986; an overview of marine taxa) and Popovsky & Pfiester (1990; an overview of nonmarine taxa). Dodge (1985) published an atlas of scanning electron photomicrographs of extant dinoflagellates. Fossil dinoflagellates have been discussed in detail by Evitt (1985). Sarjeant (1974) and Edwards (1993) provided overviews of living and fossil dinoflagellates, Williams (1977, 1978) surveyed fossil dinoflagellates, Dale (1983) and Sarjeant et al. (1987) reviewed the morphology and ecology of dinoflagellate cysts with emphasis on the fossil record, and Fensome, Taylor et al. (1993) treated the classification and evolution of both fossil and living dinoflagellates. Several catalogs and indices produced in recent decades include: the catalog series initiated by Eisenack & Klement (1964) , with subsequent issues by Eisenack (1967), Eisenack & Kjellström (1971, 1972, 1975a, b, 1981a, b) and Fensome, Gocht et al. (1991, 1993); the indexes of Lentin & Williams (1973, 1975, 1977, 1981,1985, 1989, 1993); and several compendia of genera, including Stover & Evitt (1978), Artzner et al. (1979), Wilson & Clowes (1980) and Stover & Williams (1987)" (Fensome et al. 1996, p. 107).

[edit] Dendrogram

The following tree differs in some regards from the classification above - Ellobiopsida are not included in Dinoflagellata, but are closely related within the Alveolata. Ebriophyceae are not regarded as related to dinoflagellates, but are tentatively included in the Apusozoa. Oxyrrhinales are here included as basal dinoflagellates, but some authors would exclude them from Dinoflagellata (Fensome et al., 1993).

<==Dinoflagellata (see below for synonymy)
   |  i. s.: Yalkalpodium scutum Morgan 1980
   |         Downiesphaeridium Islam 1993
   |           `--D. armatum (Deflandre) Islam 1993
   |         Disphaeria Cookson & Eisenack 1960
   |           `--D. macropyla Cookson & Eisenack 1960
   |         Sepispinula ancorifera (Cookson & Eisenack) Islam 1993
   |         Peridiniopsis polonicum
   |         Cysta moebii (Jörgensen) Loeblich & Loeblich 1970 [=Pterosphaera moebii]
   |         Entomosigma Schiller 1925
   |           `--E. simplicius Conrad 1939
   |         Halophilodinium Loeblich & Loeblich 1966 (see below for synonymy)
   |           `--H. gessneri (Schiller) Loeblich & Loeblich 1968 (see below for synonymy)
   |         Roscoffia capitata Balech 1956
   |         Albertidinium acutulum
   |         Agrosphaera Lo Bianco 1903
   |         Geodinium Chodat 1921 [incl. Chlorodinium Chodat 1921 (n. n.)]
   |         Gleba Bruguière 1791
   |         Glenoaulax Diesing 1866
   |         Parapodinium Chatton 1920
   |         Proaulax Diesing 1866
   |         Radiozoum Mingazzini 1904
   |         ‘Diplocystis’ Cleve 1901 nec Trevisan 1848 nec Berkeley & Curtis 1869 nec Agardh 1896
   |         Karlodinium micrum
   |--Oxyrrhinales
   |    |--Glyphidium Fresenius 1865
   |    `--Oxyrrhis Dujardin 1841
   |         |--O. marina Dujardin 1841
   |         |--O. maritima Van Meel 1969
   |         `--O. tentaculifera Conrad 1939
   `--+--Arpylorus Calandra 1964 [Arpyloraceae]
      |    `--A. antiquus Calandra 1964
      `--+--Peridinea
         `--+--Noctilucales
            `--Syndiniales (see below for synonymy)
                 |  i. s.: Coccidinium Chatton & Biecheler 1934 [Coccidiniaceae]
                 |         Atelodinium Chatton 1920
                 |--Amoebophrya Koeppen 1894 [Amnoebophryaceae, Amoebophryidae]
                 |    `--A. ceratii (Koeppen 1899) Cachon 1964
                 |--Sphaeriparaceae [Blastuloidae, Sphaeriparidae]
                 |    |--Atlanticellodinium Cachon & Cachon-Enjumet 1965
                 |    `--Sphaeripara Poche 1911 (see below for synonymy)
                 |         `--S. catenata (Neresheimer 1903) Loeblich & Loeblich 1966
                 |--Duboscquellaceae [Duboscquellidae]
                 |    |--Dogelodinium Loeblich & Loeblich 1966 (see below for synonymy)
                 |    |--Duboscquodinium Grassé in Chatton 1952
                 |    |--Keppenodinium Loeblich & Loeblich 1966 (see below for synonymy)
                 |    `--Duboscquella Chatton 1920
                 |         |--*D. tintinnicola (Lohmann 1908) Chatton 1920
                 |         |--D. anisospora Grassé in Chatton 1952
                 |         `--D. aspida Cachon 1964
                 `--Syndiniaceae [Syndinidae]
                      |--Hematodinium Chatton & Poisson 1930
                      |--Ichthyodinium Hollande & Cachon 1952
                      |--Merodinium Chatton 1923
                      |--Trypanodinium Chatton 1912
                      |--Syndinium Chatton 1910
                      |    `--S. belarii (Holland & Enjumet 1953) Hollande & Enjument 1955
                      `--Solenodinium (Chatton 1923) Chatton 1938
                           `--S. fallax (Chatton 1923) Chatton 1952

Dinoflagellata [Adiniferae, Adinophycidae, Amphilothales, Athecatales, Cilioflagellata, Desmocapsineae, Desmokontae, Desmomonadineae, Desmophyceae, Dinifera, Diniferae, Diniferida, Diniferidea, Diniferina, Diniferophycidae, Dinocapsales, Dinocapsineae, Dinococcales, Dinococcineae, Dinococcophycidae, Dinoflagellatae, Dinoflagellatophycidae, Dinoflagelliae, Dinoflagellida, Dinoflagellidea, Dinoflagellidia, Dinokaryota, Dinokontae, Dinomastigota, Dinomonadea, Dinophyceae, Dinophycidae, Dinophyta, Dinotrichales, Dinotrichineae, Endoflagellatophycidae, Kolkwitziellales, Mesocaryota, Peridineae, Peridiniaea, Peridinieae, Peridinina, Peridiniophyceae, Peridinophyceae, Phytodinierae, Phytodinozoa, Pyrrhophycophyta, Pyrrhophyta, Pyrrophyta, Rhizodineae, Rhizodininae, Thecatales]

Dogelodinium Loeblich & Loeblich 1966 [=Collinella Cachon 1964 nec Schmidt 1879 nec Duda 1918 nec Chatton & Pérard 1919]

Halophilodinium Loeblich & Loeblich 1966 [=Haematodinium Schiller 1956 non Hematodinium Chatton & Poisson 1930]

Halophilodinium gessneri (Schiller) Loeblich & Loeblich 1968 [=Haematodinium gessneri Schiller 1956]

Keppenodinium Loeblich & Loeblich 1966 [=Hollandella Cachon 1964 non Gill 1901]

Phytodiniaceae [Amoebodiniaceae, Amoebodinidae, Cystodiniaceae, Cystodiniidae, Dinamoebaceae, Dinamoebidiaceae, Dinocapsaceae, Dinococcaceae, Dinococcidae, Gloeodiniaceae, Gloeodinidae, Hemidiniaceae, Hypnodiniaceae, Stylodiniaceae]

Phytodiniales [Dinamoebales, Dinamoebidiales, Dinocapsida, Dinocloniales, Dinococcida, Dinotrichida, Gloeodiniales, Rhizodiniales, Rhizodinida]

Sphaeripara Poche 1911 (nom. cons. prop.) [=Lohmannella Neresheimer 1904 non Trouessart 1901 (ICZN), Lohmannia Neresheimer 1903 non Michael 1898 (ICZN), Neresheimeria Uebel 1912]

Syndiniales [Coccidinea, Coccidiniales, Coelomastigina, Duboscquodinida, Syndina, Syndinea, Syndinida, Syndinina, Syndiniophyceae, Syndiniophycidae]

* Type species of genus indicated

[edit] References

Anderson, DM, JJ Lively, EM Reardon, & CA Price (1985), Sinking characteristics of dinoflagellate cysts. Limnol. Oceanogr. 30: 1000.

Brugerolle, G., G. Bricheux, H. Philippe & G. Coffe. 2002. Collodictyon triciliatum and Diphylleia rotans (=Aulacomonas submarina) form a new family of flagellates (Collodictyonidae) with tubular mitochondrial cristae that is phylogenetically distinct from other flagellate groups. Protist 153: 59-70.

Cavalier-Smith, T. 1998. A revised six-kingdom system of life. Biological Reviews 73: 203-266.

Cavalier-Smith, T. 2002. The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa. International Journal of Systematic and Evolutionary Microbiology 52: 297-354.

Cavalier-Smith, T., & E. E.-Y. Chao. 2003. Molecular phylogeny of centrohelid heliozoa, a novel lineage of bikont eukaryotes that arose by ciliary loss. Journal of Molecular Evolution 56: 387-396.

Clowes, CD (1985), Stoveracysta, a new gonyaulacacean dinoflagellate genus from the upper Eocene and lower Oligocene of New Zealand. Palynology 9: 27-36.

Cruickshank, A. R. I., & R. E. Fordyce. 2003. A new marine reptile (Sauropterygia) from New Zealand: Further evidence for a Late Cretaceous austral radiation of cryptoclidid plesiosaurs. Palaeontology 45 (3): 557-575.

Curds, C. R. 1979. Group phenomena in the phylum Protozoa. In Biology and Systematics of Colonial Organisms (G. Larwood & B. R. Rosen, eds.) pp. 29-37. Academic Press: London.

Dale, B (1996), Dinoflagellate cyst ecology: modelling and geological applications, in J Jansonius & DC McGregor [eds.], Palynology: Principles and Applications. Am. Assoc. Stratigr. Palynologists Found., 3: 1249–1275.

Evitt, WR (1985), Sporopollenin dinoflagellate cysts -- their morphology and interpretation. Am. Assoc. Strat. Palynologists Found., 333 pp.

Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i-viii, 1-351.

Friedrich, O., & K. J. S. Meier. 2003. Stable isotope indication for the cyst formation depth of Campanian/Maastrichtian calcareous dinoflagellates. Micropaleontology 49 (4): 375-380.

Gottschling, M., H. Keupp, J. Plötner, R. Knop, H. Willems & M. Kirsch. 2005. Phylogeny of calcareous dinoflagellates as inferred from ITS and ribosomal sequence data. Molecular Phylogenetics and Evolution 36: 444-455.

Hotton, CL, FM Hueber, DH Griffing, & JS Bridge (2001), Early Terrestrial Plant Environments: An Example from the Emsian of Gaspé, Canada, in PG Gensel & D Edwards [eds.] Plants Invade the Land. Columbia Univ. Press: pp. 179-203.

Ibrahim, M. I. A. 2002. Late Albian-Middle Cenomanian palynofacies and palynostratigraphy, Abu Gharadig-5 well, Western Desert, Egypt. Cretaceous Research 23: 775-788.

Jacobsen, S., K. Lüning & F. Goulard. 2003. Circadian changes in relative abundance of two photosynthetic transcripts in the marine macroalga Kappaphycus alvarezii (Rhodophyta). Journal of Phycology 39: 888-896.

Klein-MacPhee, G. 2002a. Smelts. Family Osmeridae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds.) 3rd ed. pp. 162-170. Smithsonian Institute Press: Washington.

Klein-MacPhee, G. 2002b. Porcupinefishes. Family Diodontidae. In Bigelow and Schroeder’s Fishes of the Gulf of Maine (B. B. Collette & G. Klein-MacPhee, eds.) 3rd ed. pp. 601-602. Smithsonian Institute Press: Washington.

Knoll, AH (1996), Archean and proterozoic paleontology, in J Jansonius & DC McGregor [eds.], Palynology: Principles and Applications. Am. Assoc. Stratigr. Palynologists Found. 1: 51-80.

Leander, B. S., R. E. Clopton & P. J. Keeling. 2003. Phylogeny of gregarines (Apicomplexa) as inferred from small-subunit rDNA and β-tubulin. International Journal of Systematic and Evolutionary Microbiology 53: 345-354.

Lee, J. J., R. Olea, M. Cevasco, X. Pochon, M. Correia, M. Shpigel & J. Pawlowski. 2003. A marine dinoflagellate, Amphidinium eilatiensis n. sp., from the benthos of a mariculture sedimentation pond in Eilat, Israel. Journal of Eukaryotic Microbiology 50 (6): 439-448.

Margulis, L & KV Schwartz (1982), Five Kingdoms: An Illustrated Guide to the Phyla of Life. Freeman.

Martin, F & G Kjellström (1973), Ultrastructural study of some Ordovician acritarchs from Gotland, Sweden. Neues Jahrb. Geol. Paläont. Monatsh. 1973: 44.

Mendelson, CV (1993), Acritarchs and prasinophytes, in JH Lipps [ed.], Fossil Prokaryotes and Protists. Blackwell Scientific, pp 77-104.

Marret, F., & J. Scourse. 2002. Control of modern dinoflagellate cyst distribution in the Irish and Celtic seas by seasonal stratification dynamics. Marine Micropaleontology 47: 101-116.

Melkonian, M., & D. Mollenhauer. 2005. Robert Lauterborn (1869 – 1952) and his Paulinella chromatophora. Protist 156: 253-262.

Moldowan, JM & NM Talyzina (1998), Biogeochemical evidence for dinoflagellate ancestors in the Early Cambrian. Science 281: 1168-1169.

Moore, R. B., K. M. Ferguson, W. K. W. Loh, O. Hoegh-Guldberg & D. A. Carter. 2003. Highly organized structure in the non-coding region of the psbA minicircle from clade C Symbiodinium. International Journal of Systematic and Evolutionary Microbiology 53: 1725-1734.

Okamoto, N., & I. Inouye. 2005. The katablepharids are a distant sister group of the Cryptophyta: A proposal for Katablepharidophyta divisio nova / Kathablepharida phylum novum based on SSU rDNA and beta-tubulin phylogeny. Protist 156: 163-179.

Pollingher, U (1987), Ecology: freshwater systems, in FJR Taylor [ed.], The Biology of Dinoflagellates. Blackwell Scientific, pp. 502-529.

Pratt, LM, RE Summons & GB Hieshima (1991), Sterane and triterpane biomarkers in the Precambrian Nonesuch Formation, North American Midcontinent Rift. Geochim. Cosmochim. Acta 55: 911-916.

Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology (3rd ed.) Wm. C. Brown Publishers: Dubuque (Iowa).

Quattrocchio, M. E., & W. A. S. Sarjeant. 2003. Dinoflagellates from the Chorrillo Chico Formation (Paleocene) of southern Chile. Ameghiniana 40 (2): 129-153.

Quinn, P., H. R. Thierstein, L. Brand & A. Winter. 2003. Experimental evidence for the species character of Calcidiscus leptoporus morphotypes. Journal of Paleontology 77 (5): 825-830.

Riding, J. B., & J. A. Crame. 2002. Aptian to Coniacian (Early – Late Cretaceous) palynostratigraphy of the Gustav Group, James Ross Basin, Antarctica. Cretaceous Research 23: 739-760.

Saldarriaga, J. F., M. L. McEwan, N. M. Fast, F. J. R. Taylor & P. J. Keeling. 2003. Multiple protein phylogenies show that Oxyrrhis marina and Perkinsus marinus are early branches of the dinoflagellate lineage. International Journal of Systematic and Evolutionary Microbiology 53: 355-365.

Shimuzu, Y. 1987. Dinoflagellate toxins. In The Biology of Dinoflagellates (F. J. R. Taylor, ed.) Botanical Monographs 21: 282-315.

Sournia, A. 1973. Catalogue des espèces et taxons infraspécifiques de Dinoflagellés marins actuels publiés depuis la révision de J. Schiller. I. Dinoflagellés libres. Beihefte zur Nova Hedwigia 48: 1-92.

Strother, PK (1996), Acritarchs, in J Jansonius & DC McGregor [eds.], Palynology: Principles and Applications. Am. Assoc. Stratigr. Palynologists Found. 1: 81-106.

Summons, RE & MR Walter (1990), Molecular fossils and microfossils of procaryotes and protists from Proterozoic sediments. Amer. J. Sci. 290-A: 212-244.

Tappan, H (1980), The Paleobiology of Plant Protists. Freeman, 1028 pp.

Taylor, F. J. R. 1987. General and marine ecosystems. In F. J. R. Taylor & U. Pollingher. Ecology of dinoflagellates. In F. J. R. Taylor [ed.], The Biology of Dinoflagellates. Botanical Monographs 21: 399-502.

Taylor, F. J. R. 2003. The collapse of the two-kingdom system, the rise of protistology and the founding of the International Society for Evolutionary Protistology (ISEP). International Journal of Systematic and Evolutionary Microbiology 53: 1707-1714.

Vallejo, C., P. A. Hochuli, W. Winkler & K. von Salis. 2002. Palynological and sequence stratigraphic analysis of the Napo Group in the Pungarayacu 30 well, sub-Andean zone, Ecuador. Cretaceous Research 23: 845-859.

Wall, D & B Dale (1968), Modern dinoflagellate cysts and evolution of the Peridiniales. Micropaleontology 14: 265-304.

Wright, A.-D. G., & A. Colorni. 2002. Taxonomic re-assignment of Cryptocaryon irritans, a marine fish parasite. European Journal of Protistology 37: 375-378.

Zhukova, N. V. 2003. A biochemical approach for assessment of the diversity of symbiotic dinoflagellates. Russian Journal of Marine Biology 29 (5): 328-332.

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