Belongs within: Life.
Contains: Chitinozoa, Acritarcha, Microrhabdulaceae, Palaeopascichnida, Discoba, Amorphea, Diaphoretickes.
The Eukaryota is the group of organisms in which the chromosomes are contained within a nuclear membrane that is separate from the external cell membrane. In the majority of cases, cells of eukaryotes are much larger than those of non-eukaryotes. Movement of cells is commonly via the beating of one or more cilia (alternatively referred to as ‘flagella’ when relatively long compared to the remainder of the cell) but numerous lineages have lost cilia for at least part of the life cycle.
Basal relationships within the eukaryotes have been subject to much argument in the past and even now are far from settled. The major clades Amorphea and Diaphoretickes may be ancestrally united in a clade by a derived cytochrome c lyase biogenesis as opposed to the bacterial pathway found in more basal taxa (Cavalier-Smith 2018). Conversely, other authors unite the clades Discoba and Metamonada in a clade ‘Excavata’ of flagellates ancestrally possessing a ventral feeding groove (Adl et al. 2019). The Hemimastigophora are ellipsoid to vermiform cells with cilia arranged in two lateral rows that may or may not run the whole length of the cell (Adl et al. 2019).
The oldest fossil eukaryotes, recognised by their size and complexity relative to prokaryotes, may be known from the latest Palaeoproterozoic era. However, biomarkers of sterol production (thought to have been present in the last common ancestor of modern eukaryotes) are not known until the late Mesoproterozoic, suggesting that earlier fossils may be representatives of the stem group (Porter 2020). The Early Mesoproterozoic Tappania plana bears irregularly distributed hollow processes over the cell surface (Porter 2020).
Naming the monad
Published 28 March 2008
Know you not that around the animalcule that sports in the water there shines a halo, as around the star (The Monas mica, found in the purest pools, is encompassed with a halo. And this is frequent amongst many other species of animalcule.) that revolves in bright pastime through the space? True art finds beauty everywhere.—Zanoni, Edward Bulwer-Lytton (1842).
The classification of microscopic organisms long lagged behind that of other organisms in attention and reliability, for the simple reason that the technology to distinguish a significant amount of features in said microscopic organisms took a lot longer to develop. The really minute flagellates posed a particular challenge due to their particularly small size, as well as a shortage of distinguishing features between taxa. A review by Boenigk (2008) looks at the history of classification of the nanoflagellates through the lens of the history of the first genus named, Monas Müller 1773. Monas was long a receptacle for almost all minute flagellates (and even if not specifically referred to Monas, such organisms were still referred to as ‘monads’), only to eventually slip into disuse as it became increasingly unclear what the name Monas had originally referred to.
When Monas was first named in 1773, life was still firmly divided between ‘plants’ and ‘animals’, and the concept of organisms falling outside these classes had not yet developed (Hogg proposed ‘Protoctista’ in 1860). The two kingdoms were distinguished by the motility of animals, making most protists such as Monas animals (and fungi into plants). It was only later that some protists were reclassified as plants due to their autotrophy (or more accurately, lack of particulate food uptake—fungi were still plants). Perty summed up the problem when establishing his order Phytozoidia in 1852 (as quoted in Boenigk 2008):
Die zweite Ordnung [der Infusorien] kann den Namen Phytozoidia erhalten, weil unter ihnen sehr viele Formen sich befinden, welche in ihrem Lebenscyklus in Wahrheit bald dem Thier- bald dem Pflanzenreiche angehören, zwischen beiden oscilliren, während andere, bei denen dies nicht der Fall ist, so sehr in Gestalt, Bau, Bewegung und sonstigem Verhalten mit ihnen übereinstimmen, dass an eine völlige Trennung nicht zu denken ist.
[The second order [of the Infusoria] may get the name Phytozoidia as it contains many forms, which in their life cycle truly belong at times to the animal—at times to the plant kingdom; they either oscillate between both, or, when this is not the case, are that similar to the former in shape, structure, motion, and other behavior, that they cannot be separated.]
The genus Monas was first established with three species, M. termo, M. lens and M. mica. Monas termo has since been reclassified to the chrysophyte genus Oikomonas while M. lens is now in the kinetoplastid genus Bodo, leaving M. mica as the effective type species of Monas* (Silva 1960). Over the years, more than 100 species have been assigned to Monas with little more to unite them than being small, fairly nondescript and (usually) with flagella. Different authors have applied different definitions of the genus, often in complete contradiction to each other (for instance, Bory [1824] defined Monas as lacking caudiform appendages, while Dujardin [1841] defined it as possessing a single flagellum). As recounted by Boenigk, species attributed to Monas eventually turned out to belong to the entire range of protist groups—amoebozoans, fungi, chlorophytes, excavates, even a few bacteria.
*As with many if not most protists, it is a little unclear whether Monas should be treated as a name under the Zoological or Botanical Code. The typification of the genus by M. mica through removing the other two species is valid under the ICBN, but not necessarily so under the ICZN. Nevertheless, we’ll accept M. mica as the type species of Monas because there’s no reason to unnecessarily tick off the botanists.
So what does the name Monas ultimately belong to? Unfortunately, that’s where we must draw a blank. Boenigk (2008) gives us the original description of Monas mica (as well as the original illustration, shown above):
Lenticula 3. microsc. simplicis puctulum lucidem conspicitur, aucta vero magnitudine animalculum ovale vel sphaericum, nam utramque figuram pro lubitu assumit, exhibetur. Hyalina est, circulo ovali intus inscripta; hic mobilis est, & in medio, vel versus antica vel versus postica videtur. Motus vacillatorius; saepe eodem in loco, assumata figura sphaerica, diu gyratur, tumque impressio reniformis oculo in medio corporis sistitur, animalculumque halone, absque dubio e ciliis vibrantibus invisibilibus orto, pulchre cingitur. In aquis purioribus passim.
[Under the lens 3 of a simple microscope it is seen as a bright dot, yet at higher magnification as oval or spherical animalcule, because it adopts at times the one, and at times the other shape. It is hyaline, with an oval shape inside, here it is motile and seen either in the centre or anteriorly or posteriorly. With trembling movement; often rotating in one place, then with a spherical shape but with a persisting kidney shaped depression near the centre of the body, and the animalcule is surrounded by a nice halo, which arises without doubt from trembling invisible cilia. In pure water.]
There’s only a few thousand taxa to which that description could apply to choose between. In fact, the description is a little vague as to whether the organism in question was even a flagellate—the presence of flagella was only inferred from the ‘halo’, not demonstrated. Later descriptions of Monas mica in Pritchard (1834) and Hogg (1854) add little to the original description, though both repeat the description of a halo, as for that matter does the passage quoted at the top of this post*.
*Edward Bulwer-Lytton was an immensely popular Victorian author whom Mary Shelley described as the “first author of the age”. Modern audiences have a slightly different opinion of him. San Jose University in California established its Bulwer-Lytton Fiction Contest for the worst beginning to an imaginary novel in memory of the opening words of Bulwer-Lytton’s Paul Clifford—”It was a dark and stormy night…”.
A general assumption has evolved that Monas is vaguely equivalent to the chrysophyte genus Spumella. However, Silva (1960) demonstrated that this association was not connected with any of the original species of Monas but due to the species described as M. vivipara (now Spumella vivipara) by Ehrenberg in 1835. As a result, there is no reason to associate Monas with Spumella any more than with any other flagellate genus, and the suggestion by Cavalier-Smith & Chao (2006) that Monas could be used for one of the clades of the also polyphyletic Spumella lacks any taxonomic justification. Despite having a long and exalted history, the name Monas has fallen into obscurity, and there it must stay.
More things in heaven and ocean
Published 8 July 2009
Following on from the post just put up on xenophyophores, I couldn’t pass on without mentioning these fellows:
This is Haeckel’s (1889) plate of members of the Ammoclathrinidae, taken from here. Haeckel described members of this family* among his “Deep-Sea Keratosa”, the other examples of which have since been identified as xenophyophores (and not, as Haeckel thought they were, sponges). Like xenophyophores, ammoclathrinids had coverings constructed from foreign objects, and when I wrote the page on xenophyophores for Palaeos.com, I interpreted ammoclathrinids as being xenophyophores. True, Haeckel had originally identified separate cells inside his ammoclathrinids, but I suggested that this could have been an artefact caused by his use of acid to prepare the specimens, and they might have originally been coenocytic. However, when I wrote that I was working from Tendal’s (1972) recounting of Haeckel’s work, and hadn’t seen the original description. Looking at it now, I have to admit that the illustrations in Haeckel’s plate look very different from xenophyophores.
*Which he called Ammoconidae, but this name was changed by Tendal (1972) for reasons of preoccupation.
Unfortunately, Haeckel’s original material no longer exists (it seems likely that he pulled it apart in the process of describing it), and no-one has ever laid eyes on a recognisable ammoclathrinid since. The nature of these organisms, therefore, is a complete mystery. Such a situation is more common than you may think, particularly in microbiology. The protistological literature is littered with examples of organisms seen and described by someone looking down a microscope in the 1800s, and never recorded since. It might be tempting to wonder whether the original describer ever really saw what they thought they saw at all, but that may be ignoring one of the major issues in microbiology itself—the world is very, very large, and the things being looked for a very, very small. If comparatively large mammal species can elude attention for numbers of years, how much more so can tiny micro-organisms? Or something like Ammoclathrinidae, lurking somewhere in the little-explored deep sea?
Systematics of Eukaryota
Characters (Cavalier-Smith 2002): Chromosomes and ribosomes attached to endoplasmic reticulum/nuclear envelope, which is topologically within, and separate from, plasma membrane; endoplasmic reticulum growing by direct insertion of individual lipid molecules synthesised by proteins embedded in same membrane; plasma membrane growing by fusion of vesicles budded from endomembranes. Complex endoskeleton (cytoskeleton) present of microtubules and actin filaments using attached molecular motors to mediate chromosome segregation and cell division, respectively; cells ancestrally phagotrophic. Cell motile organs (when present) cilia.
<==Eukaryota (see below for synonymy)
|--Tappania planaP20
|--Dictyosphaera macroreticulataP20
|--GerminosphaeraP20
| |--G. alveolataP20
| `--G. bispinosaP20
`--+--DiscobaAB19
`--Neozoa (see below for synonymy)C-S18
| i. s.: Oscana Bosc 1797B97
| `--*O. astacaria Bosc 1797B97
|--AmorpheaC-S18
`--+--DiaphoretickesC-SCL16
`--Hemimastigophora [Hemimastigea, Hemimastigida]AB19
| i. s.: ParamastixAB19
`--SpironemidaeC-S93
|--Hemimastix Foissner, Blatterer & Foissner 1988LE18, C-S93
| |--H. amphikinetaLE18
| `--H. kukwesjijk Eglit & Simpson in Lax, Eglit et al. 2018LE18
`--Spironema Klebs 1922 non Lindley 1840 (ICBN)C-S93
|--S. goodeyiLE18
|--S. multiciliatumLE18
`--S. terricolaLE18
Eukaryota incertae sedis:
MelanocyrilliumC-S10
Trachyhystrichosphaera aimikaP20
ChitinozoaWBN02
Corollolithion kennedyiGT02
Helenea chiastiaGT02
Marthasterites Deflandre 1959FT93
|--M. contortusP79
|--M. furcatusVH02
`--M. tribrachiatusP79
Lithastrinus septenariusVH02
EprolithusVH02
|--E. moratusVH02
`--E. octopetalusVH02
Quadrum gartneriVH02
Calculites anfractusVH02
Reinhardites antophorusVH02
Lucianorhabdus Deflandre 1959H75
`--*L. cayeuxi Deflandre 1959H75
Lithocodium aggregatumHS02
Cinetidomyxa chattoniT72
Lagenidiopsis Golemansky 1974 non De Wild. 1896 (ICBN)G86
|--L. ancora Sudzuki 1979G86
|--L. elegans (Gruber 1884)G86
`--L. valkanovi Golemansky 1974G86
Pseudolagenidiopsis Sudzuki 1979G86
`--P. golemanskyi Sudzuki 1979G86
‘Voluta’ Chardez 1972 non Linnaeus 1758G86
`--V. hemispiralis Chardez 1972G86
Bicorium Maslov 1956BB05
`--*B. kusbassense Maslov 1956BB05
ArchaeolithothamniumBB05
|--A. irinae Maslov 1956BB05
`--A. saipanense [=Sporolithon saipanense]A04
Wengania Zhang 1989X04
|--*W. globosa Zhang 1989X04
|--W. exquisita Zhang, Yin et al. 1998X04
`--W. minuta Xiao 2004X04
Paratetraphycus giganteus Zhang 1985X04
Meghystricosphaeridium Zhang, Yin et al. 1998X04
|--*M. chadianensis (Chen & Liu 1986) (see below for synonymy)X04
|--M. densum (Kolosova 1991) Zhang, Yin et al. 1998X04
|--M. gracilentum (Yin in Yin & Liu 1988) Zhang, Yin et al. 1998X04
|--M. magnificum Zhang, Yin et al. 1998X04
|--M. perfectum (Kolosova 1991) Zhang, Yin et al. 1998X04
`--M. reticulatumX04
Dicrospinasphaera Yuan & Hofmann 1998X04
`--*D. zhangii Yuan & Hofmann 1998X04
Apodostoides basileus Zhang et al. 1998X04
Tianzhushania spinosa Yin & Li 1978X04
EriciasphaeraX04
|--E. magna (Zhang 1984)X04
|--E. rigidaX04
|--E. sparsaX04
`--E. triangulaP20
Asterocapsoides sinensisX04
Cerionopora ordinata [incl. Bacatisphaera baokangensis]X04
Briareus borealis Knoll 1992X04
Castaneasphaera speciosaX04
Cymatiosphaeroides yiniiX04
Distosphaera speciosa Zhang et al. 1998X04
Echinosphaeridium maximumX04
EotylotopallaX04
|--E. dactylosX04
`--E. delicataX04
FilisphaeridiumX04
GoniosphaeridiumX04
|--G. acuminatumX04
|--G. conoideumX04
`--G. cratumX04
LophosphaeridiumX04
Mastosphaera changyangensisX04
MicrhystridiumX04
Pustulisphaera membranaceaX04
Sinosphaera rupinaX04
Vulcanisphaera phacelosaX04
Gremiphyca corymbiata Zhang et al. 1998X04
Paramecia incognata Zhang & Yuan 1992X04
Thallophycoides phloeatus Zhang & Yuan 1992X04
MonasLB90
|--M. arhabdomonasV63
|--M. guttulaV63
|--M. lensG20
|--M. minimaLB90
|--M. socianilisSX97
|--M. stigmaticaPHK96
|--M. termoG20
`--M. vulgarisV63
Radiosus Zhuravleva 1964T84, G79
|--*R. limpidus Zhuravleva 1964G79
`--R. radialisT84
GranomarginataT84
|--G. dhalliT84
|--G. primitivaT84
`--G. simlaensisT84
Griffonia simplicifoliaWN00
MizziaVH01
|--M. velebitana Schubert 1908VH01
`--M. yabeiVH01
Pseudoepimastopora likana Kochansky-Devidé & Herak 1960VH01
Arabicodium [incl. Siamporidium Endo 1969]VH01
OrtonellaVH01
|--O. latifibrosa Endo in Endo & Kanuma 1954VH01
`--O. morikawai Endo 1958VH01
NanoporaVH01
|--‘Anthracoporella’ fragilissima Maslov 1939VH01
`--‘Anthracoporella’ kasachiensis Maslov 1939VH01
Roquesselsia Termier et al. 1977VH01
`--‘Anthracoporella’ insolita Ivanova 1988VH01
Oligoporella Pia 1912VH01
Clavaphysoporella Endo 1958VH01
Clavaporella [incl. Eoclypeina Poncet 1987 non Emberger in Vachard 1985]VH01
`--C. reinae Racz 1966 [incl. *Eoclypeina crassa Poncet 1987]VH01
Neoanchicodium Endo in Endo & Kanuma 1954VH01
Nostocites Maslov 1929VH01
`--*N. vesiculosa Maslov 1929VH01
Sphairionia Tien 1989 [incl. Pseudosphairionia Pronina 1996]VH01
|--*S. sikuoides Tien 1989 (see below for synonymy)VH01
`--S. gallinae Vachard, Hauser et al. 2001VH01
Marpolia spissaCM98
Polysphaerinella bullaCM01
Luffisphaera [Holosea]C-S03
KamaenaE03
|--K. delicataE03
`--K. tatarstanicaE03
Izhella nubiformisE03
Bosea Srivastava 1975FT93
Evittia Brito 1967FT93
Anthosphaeridium Cookson & Eisenack 1968FT93
Archaeodinium Shu 1979FT93
Artemisiocysta Benedek 1972FT93
Ascostomocystis Drugg & Loeblich 1967FT93
`--A. hydria Drugg & Loeblich 1967E77
Beringiella Bujak 1984FT93
Caddasphaera Fenton et al. 1980FT93
Catillopsis Drugg 1970FT93
Ceratocystidiopsis Deflandre 1937FT93
Chonesphaera Klumpp 1953FT93
Cirrifera Cookson & Eisenack 1960FT93
Collumosphaera Jain & Dutta in Dutta & Jain 1980FT93
Conicoidium Jiabo 1978FT93
Cupritia Courtinat & Gaillard 1980FT93
‘Cyclodictyon’ Cookson & Eisenack 1958 non Mitt. 1864FT93
Cyclopsiella Drugg & Loeblich 1967FT93
Cystidiopsis Nagy 1965FT93
Deflandridium Nagy 1969FT93
Dictyosphaeridium Wetzel 1952FT93
Diplofusa Cookson & Eisenack 1960FT93
Wallodinium Loeblich & Loeblich 1968 [=Diplotesta Cookson & Eisenack 1960 (preoc.)]FT93
Domasiella Eisenack 1969FT93
Enigmadinium Wrenn & Hart 1988FT93
AcritarchaE77
Halodinium Bujak 1984FT93
Halophoridia Cookson & Eisenack 1962FT93
Korojonia Cookson & Eisenack 1958FT93
Lophodictyotidium Pocock 1972FT93
Mathurosphaera Varma & Dangwal 1964FT93
Membranosphaera Samoilovich in Samoilovich & Mtchedlishvili 1961FT93
Mirificysta Yu 1982FT93
Nummus Morgan 1975FT93
‘Ollula’ Góczán 1962 non Lév. 1863FT93
Palaeohystrichosphaeridium Eiserhardt 1986FT93
Palaeostomocystis Deflandre 1937FT93
Palhistiodinia Deflandre 1939FT93
Piperodinium Mudie in Scott et al. 1984FT93
Platycystidia Cookson & Eisenack 1960FT93
Polyasterias Meunier 1910 non Perrier 1894 (ICZN)FT93
Prismatocystis Habib 1969FT93
Pseudofromea Burger 1980FT93
Pterocystidiopsis Deflandre 1937FT93
Quadrina Bujak 1980FT93
Rhombodella Cookson & Eisenack 1962FT93
Schizocystia Cookson & Eisenack 1962FT93
Sicyoideum He 1980FT93
Spicadinium Batten & Lister 1988FT93
Spinuliferites Huang 1981FT93
Stylodiniopsis Eisenack 1954FT93
`--*S. maculatum Eisenack 1954H75
Tetragonidium Pascher 1914FT93
‘Tubulifera’ Schumacker-Lambry 1978 non Müll. ex Jacq. 1779FT93
Sinosabellidites Zheng 1980 [incl. Huainanella Wang 1982]SWZ86
`--*S. huainanensis Zheng 1980 [incl. Huainanella cylindrica Wang 1982]SWZ86
Vindhyania Mathur 1983SWZ86
Phimopollenites pannosus (Dettmann & Playford 1968) Dettmann 1974RC02
Sarjeantia Horowitz 1975FT93
Ammoclathrinidae [Ammoconidae]T72
|--Ammosolenia Haeckel 1889T72
| `--A. rhizammina Haeckel 1889T72
|--Ammoclathrina Laubenfels 1948 [=Ammoconia Haeckel 1889 (preoc.)]T72
| |--*A. aulopegma (Haeckel 1889) [=Ammoconia aulopegma]T72
| `--A. sagenella (Haeckel 1889) [=Ammoconia sagenella]T72
`--Ammolynthus Haeckel 1889T72
|--*A. haliphysema Haeckel 1889T72
`--A. prototypus Haeckel 1889T72
Morania Walcott 1919H85
|--*M. globosa Walcott 1919H85
`--M. antiqua Fenton & Fenton 1939H85
Longfengshania Du 1982H85
`--*L. stipitata Du 1982H85
Daltaenia Hofmann 1985H85
`--*D. mackenziensis Hofmann 1985H85
Globochaete Lombard 1945MS99
`--G. alpinaF71
OvummuridaeMSV00
|--Ovummurus Minoura & Chitoku 1979MSV00
| `--*O. duoportius Minoura & Chitoku 1979 [=O. duoportio]MSV00
|--Minourella Munnecke, Servais & Vachard 2000MSV00
| `--*M. gotlandica Munnecke, Servais & Vachard 2000MSV00
|--Arouxina Munnecke, Servais & Vachard 2000MSV00
| `--*A. pluricellata Munnecke, Servais & Vachard 2000MSV00
`--Samtlebenella Munnecke, Servais & Vachard 2000MSV00
`--*S. circumcamerata Munnecke, Servais & Vachard 2000MSV00
CalcisphaeridaeMSV00
|--Polyderma Derville 1950MSV00
|--Asterosphaerella Villain 1975MSV00
|--Eovolvox Kazmierczak 1975MSV00
`--Calcisphaera Williamson 1880MSV00 [incl. Granulosphaera Derville 1931H75]
|--*C. robusta Williamson 1881H75
`--C. leavis Williamson 1881M98 [=*Granulosphaera laevisH75]
Pleuromonas jaculansLB90
Thylacomonas compressSX97
Nebula carinatallaSX97
Rosenia Waagen & Wentzel 1886BR05
Pseudodimorpha Mikrjukov 2000M00
`--*P. marina (Fenchel et al. 1995) [=Tetradimorpha marina]M00
Moncharmontia apeninicaLSM-CG00
Disphaeria Cookson & Eisenack 1960FT93
`--D. macropyla Cookson & Eisenack 1960RC02
Nanogyra nanaYZ83
Thurmanella pentaptyctaYZ83
Paleonuculus makitoensisYZ83
TriflagellumS03
|--T. diaphanum Ruinen 1938S03
`--T. hardyi Ruinen 1938S03
Eothrix alpinaF71
Sphenolithus [Sphenolithaceae]VC88
|--S. abies Deflandre in Deflandre & Fert 1954VC88
|--S. belemnosP79
|--S. ciperoensisP79
|--S. distentusP79
|--S. heteromorphusP79
|--S. neoabies Bukry & Bramlette 1969EB93
|--S. predistentusP79
|--S. primus Perch-Nielsen 1971EB93
`--S. pseudoradiansP79
Heteroderma Fitzinger 1843LT64a
Astacoderma Harley 1861Has62
Bransonella Harlton 1933Has62
Dermatolithis Ehrenberg 1854Has62
Fortscottella Gunnell 1931Has62
Icthyodus Harris & Hollingsworth 1933Has62
Lepodus Branson & Mehl 1933Has62
Lepognathodus Mehl in Fay 1959Has62
Multidentodus Harlton 1933Has62
Scolopodella Stauffer & Plummer 1932Has62
Stephanodella Matern 1933Has62
Halysium Swidzinski 1934 non Corda 1837 (ICBN) (see below for synonymy)H75
|--*H. problematicum Swidzinski 1934H75
|--‘Hormosira’ moniliformis Heer 1877H75
`--‘Halimeda’ saportae Fuchs 1894H75
Hensonella Elliott 1960H75
`--*H. cylindrica Elliott 1960H75
Margaretia Walcott 1931H75
`--*M. dorus Walcott 1931H75
Nannoconus Kamptner 1931 [Nannoconaceae]H75
|--*N. steinmanni Kamptner 1931H75
|--N. colomi (de Lapparent 1931) [=Lagena colomi]Han62
|--N. compressus Bralower & Thierstein in Bralower et al. 1989EB93
`--N. elongatus Bronnimann 1955EB93
Porocystis Cragin 1893H75
`--P. globularis (Giebel 1853)Han62 (see below for synonymy)
Taitia Crookall 1931H75
`--*T. catena Crookall 1931H75
Tubulites Bein 1932 (ICBN) non Gesner 1758 (ICZN)H75
Ampelitocystis Deunff 1957H75
`--*A. feuguerollensis Deunff 1957H75
Ampulites Flower 1961H75
`--*A. vasiformis Flower 1961H75
Ancestrulites Flower 1961H75
`--*A. tubiformis Flower 1961H75
Ancientia Ross 1967H75
`--*A. ohioensis Ross 1967H75
Birrimarnoldia Hovasse & Couture 1961 (see below for synonymy)H75
`--*B. antiqua (Hovasse 1956) [=*Arnoldia antiqua]H75
Cayeuxipora Graindor 1957H75
Cayeuxistylus Graindor 1957H75
Cheneyella Flower 1961H75
`--*C. clausa Flower 1961H75
Chisibyllites Deflandre 1961H75
`--*C. kerguelenensis Deflandre 1961H75
Chotecella Obrhel 1964H75
`--*C. leiotheca Obrhel 1964H75
Claviradix Ferguson 1961H75
`--*C. ashi Ferguson 1961H75
Palaeocoryne Duncan & Jenkins 1869H75
Coptocampylodon Elliott 1963H75
`--*C. lineolatus Elliott 1963H75
Cystosphaera Flower 1961 non Skottsberg 1907 (ICBN)H75
`--*C. rotunda Flower 1961H75
Dicasignetella Keij 1969H75
`--*D. eocaenica Keij 1969H75
Draffania Cummings 1957H75
`--*D. biloba Cummings 1957H75
Eliasites Flower 1961H75
`--*E. pedunculatus Flower 1961H75
Eotaeniopsis Eisenack 1955H75
`--*E. articulata Eisenack 1955H75
Fentonites Flower 1961H75
`--*F. irregularis Flower 1961H75
Gochtia Eisenack 1968H75
`--*G. rete Eisenack 1968H75
Goldringella Flower 1961H75
`--*G. plana Flower 1961H75
Harjesia Flower 1961H75
`--*H. anomala Flower 1961H75
Hikorocodium Endo 1951 [=Hicorocodium (l. c.)]H75
`--*H. elegantae Endo 1951H75
Hydrocytium silicula Matthew 1890H75
Induropilarius Deflandre & Ters in Ters & Deflandre 1966H75
`--*I. aenigmaticus Deflandre & Ters in Ters & Deflandre 1966H75
Ivesella Flower 1961H75
`--*I. adnata Flower 1961H75
Kockelites Alberti 1968H75
`--*K. longus Alberti 1968H75
Kruschevia Flower 1961H75
`--*K. verruca Flower 1961H75
Lacrymorphus Elliott 1958H75
`--*L. perplexus Elliott 1958H75
Ladinella Ott in Kraus & Ott 1968H75
`--*L. porata Ott in Kraus & Ott 1968H75
Ladinosphaera Oberhauser 1940H75
`--*L. geometrica Oberhauser 1940H75
Lamellitubus Ott in Kraus & Ott 1968H75
`--*L. cauticus Ott in Kraus & Ott 1968H75
Linotolypa Eisenack 1962H75
`--*L. arcuata Eisenack 1962H75
Lombardia Brönnimann 1955H75
|--*L. arachnoidea Brönnimann 1955H75
|--L. angulataH75
`--L. perplexaH75
MicrorhabdulaceaeEB93
Microrhabduloidus Deflandre 1963H75
`--*M. rugosus (Bouché 1962) [=Microrhabdulus rugosus]H75
Microtubus Flügel 1964H75
`--*M. communis Flügle 1964H75
Mooreopsis Flower 1961H75
`--*M. rotundus Flower 1961H75
Moundia Flower 1961H75
`--*M. fibrosa Flower 1961H75
Niccumites Flower 1961H75
`--*N. oculatus Flower 1961H75
Palamphimorphium Deflandre & Ters in Ters & Deflandre 1966H75
`--*P. speciosum Deflandre & Ters in Ters & Deflandre 1966H75
Paleocryptidium Deflandre 1955H75
`--*P. cayeuxi Deflandre 1955H75
Papinochium Deflandre & Ters in Ters & Deflandre 1966H75
`--*P. dubium Deflandre & Ters in Ters & Deflandre 1966H75
Pedicellaria Flower 1961H75
`--*P. bifurcata Flower 1961H75
Pictonicopila Deflandre & Ters 1966H75
`--*P. polymorpha Deflandre & Ters 1966H75
Plutoneptunites Deflandre 1961H75
`--*P. antarcticus Deflandre 1961H75
Polysiphonidia Eisenack 1971H75
`--*P. enigmatica Eisenack 1971H75
Pyritella Love 1958H75
`--*P. polygonalis Love 1958H75
Pyritosphaera Love 1958H75
`--*P. barbaria Love 1958H75
Slocomia Flower 1961H75
`--*S. quadrata Flower 1961H75
Spinophenia Szczechuria 1969H75
`--*S. multituba Szczechuria 1969H75
Tetrabrachiophora Eisenack 1954H75
`--*T. natans Eisenack 1954H75
Tholella Flower 1961H75
`--*T. idiotica Flower 1961H75
Triangulina Quilty 1970H75
`--*T. aequilateralis Quilty 1970H75
Urnulella Szczechura 1969H75
`--*U. costata Szczechura 1969H75
Vallenia Raunsgaard Pedersen 1967H75, G79
`--*V. erlingi Raunsgaard Pedersen 1967G79
Voorthuyseniella Szczechura 1969H75
`--*V. lageniformae Szczechura 1969H75
Warthinites Flower 1961H75
`--*W. adhaerens Flower 1961H75
Xenokalymma Eisenack 1968H75
`--*X. trematophora Eisenack 1968H75
Argonevis Kozur & Mostler 1972H75
`--*A. nuda Kozur & Mostler 1972H75
Limolepis Kozur & Mostler 1972H75
`--*L. interruptus Kozur & Mostler 1972H75
Erinea Kozur & Mostler 1972H75
`--*E. triassica Kozur & Mostler 1972H75
Venerella Kozur & Mostler 1972H75
Nemotapis Kozur & Mostler 1972H75
Cornuvacites Kozur & Mostler 1972H75
Irinella Kozur & Mostler 1972H75
`--*I. canalifera Kozur & Mostler 1972H75
Bogschites Kozur & Mostler 1972H75
`--*B. carnicus Kozur & Mostler 1972H75
Havinellites Kozur & Mostler 1972H75
`--*H. spinosus Kozur & Mostler 1972H75
Strechoritina Kozur & Mostler 1972H75
`--*S. radiata Kozur & Mostler 1972H75
Uvanogelia Kozur & Mostler 1972H75
`--*U. incurvata Kozur & Mostler 1972H75
Radimonis Kozur & Mostler 1972H75
`--*R. foliacea Kozur & Mostler 1972H75
Placerotapis Kozur & Mostler 1972H75
`--*P. subplanus Kozur & Mostler 1972H75
Fanerocoelia Kozur & Mostler 1972H75
`--*F. pennata Kozur & Mostler 1972H75
Bavlinella Shepeleva 1962 [incl. Sphaerocongregus Moorman 1974]G79
`--‘*Sphaerocongregus’ variabilis Moorman 1974G79
Beltanelloides Sokolov 1965G79
`--*B. sorichevae Sokolov 1965G79
Funisia dorotheaEL11
Hadrynichorde catalenensisEL11
Hadryniscala avalonicaEL11
Lomosovis malusEL11
PalaeopascichnidaG14
Paracharina dengyingensisEL11
Parviscopa bonavistensisEL11
Primocandelabrum hemialoranumEL11
Somatohelix sinuosusEL11
Yangtziramulus zhangiEL11
Anabylia improvisaEL11
Archangelia valdaicaEL11
Barmia lobatusEL11
Blackbrookia oaksiEL11
ElaseniaEL11
|--E. aseevaeEL11
`--E. uralicaEL11
Garania petaliEL11
Gehlingia dibrachidaEL11
Glaessneria imperfectaEL11
Irridinitus multiradiatusEL11
Ivesheadia lobataEL11
Jampolium wyrzhykoowskiiEL11
Kubisia glabraEL11
Nadalina yukonensisEL11
Namiana bakeeviEL11
Platyopholinia pholataEL11
Ramellina pennataEL11
Sekwia excentricaEL11
Shepshedia palmataEL11
Stauinidia cruciculaEL11
VendellaEL11
|--V. haelenicaeEL11
`--V. sokoloviEL11
Veprina undosaEL11
Annulina Terquem 1862 (n. d.) non Link 1820 (ICBN)LT64b
`--*A. metensis Terquem 1862LT64b
Cylidria Desvoidy 1830 [=Cylindria (l. c.) non de Gregorio 1930]LT64b
Balanulina Rzehak 1888 (n. d.)LT64b
`--*B. kittlii Rzehak 1888 (n. d.)LT64b
Millarella Carter 1888LT64b
`--*M. cantabrigiensis Carter 1888LT64b
Spongina de Gregorio 1930LT64b
`--*S. permica (de Gregorio 1930) [=Globigerina (*Spongina) permica]LT64b
Terquemina Galloway 1933LT64b
`--*T. devonica Galloway 1933LT64b
Callimastix Weissenb. 1912KC01
`--C. cyclopsisHAJ00
Adelphamoeba galeacystisLB90
Poteriodendron petiolatumH04
Uvella glaucomaH04
ActinocomaAS12
AdinomonasAS12
AletiumAS12
AmphimonasAS12
Amylophagus Scherff. 1925KC01
AphelidiopsisAS12
AsterocaelumAS12
AsthmatosAS12
AurosporaAS12
BertarelliaAS12
BodopsisAS12
BoekeloviaAS12
BranchipocolaAS12
CamptoptycheAS12
ChalarodoraAS12
CichkoviaAS12
CladomonasAS12
CodonoecaAS12
CopromonasAS12
CyanomastixAS12
CyclomonasAS12
CytamoebaAS12
DallingeriaAS12
DictyomyxaAS12
DimastigamoebaAS12
DinemulaAS12
DiplocaliumAS12
DiplomitaAS12
DiplophysalisAS12
DiploselmisAS12
DobellinaAS12
DucelleriaAS12
EctobiellaAS12
Elaeorhanis cinctaG84
EmbryocolaAS12
EndobiellaAS12
EndomonasAS12
Enteromyxa Cienk. 1884 non Ces. 1879 (ICBN)KC01
EperythrocytozoonAS12
ErreraAS12
FromentellaAS12
Gymnococcus Zopf 1884 [Gymnococcaceae, Gymnococcidae, Gymnococcinae]KC01
GymnophrydiumAS12
HaematotractidiumAS12
HartmanninaAS12
HeliobodoAS12
HeliomonasAS12
HermisenellaAS12
HilleaAS12
HyalodaktylethraAS12
Immanoplasma Neumann 1909AB19
IsoselmisAS12
KameraAS12
LiegeosiaAS12
LymphocytozoonAS12
LymphosporidiumAS12
MacappellaAS12
MagosphaeraAS12
MalpighiellaAS12
MegamoebomyxaAS12
MeringosphaeraAS12
MyrmicisporidiumAS12
NaupliicolaAS12
NephrodiniumAS12
NeurosporidiumAS12
OvicolaAS12
PalisporomonasAS12
ParadinemulaAS12
ParaplasmaAS12
ParastasiellaAS12
PeliainiaAS12
PeltomonasAS12
PetasariaAS12
PhloxamoebaAS12
PhyllomonasAS12
PhyscosporidiumAS12
PiridiumAS12
PleuromastixAS12
ProtenterosporaAS12
RhynchodiniumAS12
SchewiakoffiaAS12
SergentellaAS12
SphaerasuctansAS12
StephanomonasAS12
StrobilomonasAS12
ThaulirensAS12
TopsentellaAS12
ToshibaAS12
Concentricisporites bianulatus (Neves 1961)DV06
Ovalipollis pseudoalatusDV06
Pityosporites neomundanusDV06
Kyrtomisporis ervii Van der Eem 1983DV06
Cotyledion tyloides Luo & Hu in Luo et al. 1999AH07
Caudina cauda Gnilovskaya 1979G14
Pleraplysilla minchiniPP64
Biflabellum anthophyllumPP64
MyxastrumG84
|--M. liguricum Gruber 1884G84
`--M. radiansG84
Dorimorphe Audouin & Milne-Edwards 1832 [=Dorimorpha Herrmannsen 1852]BR17
ConusphaeraceaeEB93
|--Eoconusphaera zlambachensis Jafar 1983EB93
`--Conusphaera rothiiEB93
Fasciculithus [Fasciculithaceae]EB93
|--F. magnus Bukry & Percival 1971EB93
`--F. thomasii Perch-Nielsen 1971EB93
Goniolithus [Goniolithaceae]EB93
`--G. fluckigeri Deflandre 1957EB93
Heliolithus [Heliolithaceae]EB93
|--H. elegansEB93
`--H. megastypusEB93
Trochoaster [Lithostromationaceae]EB93
|--T. deflandreiEB93
`--T. simplex Klumpp 1953EB93
Prinsiosphaera Jafar 1983EB93
`--P. triassica Jafar 1983EB93
Scampanella Forchheimer & Stradner 1973EB93
`--S. bispinosa Perch-Nielsen 1977EB93
Lapideacassis Black 1971EB93
`--L. mariae Black 1971EB93
Triquetrorhabdulus [Triquetrorhabdulaceae]EB93
|--T. carinatus Martini 1965EB93
`--T. striatus Muller 1974EB93
CalpionellideaHW93
|--Colomiella Bonet 1956 [Colomiellidae]HW93
`--CalpionellidaeHW93
|--ChitinoidellaHW93
`--Tintinnopsella Colom 1948HW93
Meteora Hausmann et al. 2002AB19
`--M. sporadicaAB19
AcinetactisAB19
AnaeramoebaAB19
OrbulinellaAB19
ParaluffisphaeraAB19
Shuiyousphaeridium maeroreticulatumCKK09
Amoebosporus Ivimey Cook 1933KC01
Ancistrospora Menéndez & Azcuy 1972KC01
Antlea Dang. 1890 (n. d.)KC01
Bactriexta Preuss 1852 (n. d.)KC01
Fictoderma Preuss 1852KC01
Kuhneria Dang. 1933 (n. d.)KC01
Labyrinthorhiza Chadef. 1956 (n. d.)KC01
Mesenterica Tode 1790 (n. d.)KC01
Minutularia (n. d.)KC01
Myrioblepharis Thaxt. 1895 (n. d.)KC01
PolycarpumKC01
Protochytrium Borzí 1884 (n. d.)KC01
Siphopodium Reinsch 1875 (n. d.)KC01
‘Spirospora’ Scherff. 1926 non Mangin & Vincens 1920KC01
Sporoclema Tiesenh. 1912 (n. d.)KC01
Vampyrelloides Schepotieff 1911KC01
Myxotricha paradoxaSBM11
Kaibabia gemmulellaP20
Lanulatisphaera laufeldiiP20
Galerosphaera walcottiiP20
Cheilofilum hysteriopsisP20
Plicatosphaeridum impostorP20
Comasphaeridium toniumP20
Hysterolenus asiaticusCT83
Diceratopyge mobergiCT83
Milleporites flustriformisJ23
Acetabulum mediterraneumR26
DactyloporaR26
|--D. cervicornis Risso 1826R26
|--D. cylindracea Bosc 1806G20
`--D. irregularis Risso 1826R26
Birrimarnoldia Hovasse & Couture 1961 [=Arnoldia Hovasse 1956 nec Cass. 1824 (ICBN) nec Massal. 1856 (ICBN) nec Mayer-Eymar 1887 nec Stolley 1937]H75
Eukaryota [Actinopodea, Acyttaria, Alternatia, Anisomonadea, Anisomonadia, Aphrotoraca, Archaeprotista, Archimycetes, Autotractea, Axostylaria, Biciliata, Bikonta, Cabozoa, Cnidosporidia, Corallineae, Cyathobodonea, Cyclia, Eolouka, Eosarcodina, Eozoa, Euglenobiontes, Eukarya, Eukaryotae, Eurymonada, Eustomata, Excavata, Extrumonada, Flagellata, Fungilli, Gymnica, Gymnomycota, Gymnomyxa, Heliozoia, Heterozooflagellata, Histionia, Histionida, Hydraulea, Infusoria, Isomita, Jakobidea, Kinetomonada, Kinetomonadea, Lithozoa, Mastigogenina, Mastigophora, Metakaryota, Monocyphia, Mycetozoia, Myxothallophyta, Neomonada, Neosarcodina, Neosporidia, Panstomata, Pantostomata, Phycomycetes, Phytomastigophorea, Plasmodroma, Plasmodromata, Polystomata, Proterozoa, Protista, Protoctista, Protomonadina, Protomycota, Protoneozoa, Protophyta, Protoplasta, Protozoa, Pseudoplasmodida, Rhizoflagellata, Rhizomastigina, Rhizopoda, Rhizopodea, Rhizopodes, Rhizopodia, Sarcodea, Sarcodina, Sarcomastigophora, Sarcomastigota, Sarkodina, Sorophora, Sorophorae, Sorophoreae, Sorophorina, Symplectomeres, Tetramitacea, Thecomonadia, Varicristia, Vermes, Zoomastigina, Zoomastigota, Zoomastigophorea, Zoophyta]
Halysium Swidzinski 1934 non Corda 1837 (ICBN) [incl. Arthrodendron Ulrich 1904 (n. n.) nec Seward 1898 nec Scott 1900]H75
*Meghystrichosphaeridium chadianensis (Chen & Liu 1986) [incl. M. acrotrichum Yin in Yin & Liu 1988, Tianzhushania tuberifera]X04
Neozoa [Adjungentiidae, Athalamia, Athalamida, Homogenea, Mastigiomycota, Mastigomycotina, Phycomycotera, Protrudentiidae, Reticulolobosa, Testacea, Testareticulosina]C-S10
Porocystis globularis (Giebel 1853)Han62 [=Siphonia globularisH75; incl. Araucarites wardi Hill 1893H75, *Porocystis pruniformis Cragin 1893H75]
*Sphairionia sikuoides Tien 1989 [incl. Soroorina ghezelbashi Baghbani 1993 (n. n.), Sp. (*Pseudosphairionia) tienii Pronina 1996]VH01
*Type species of generic name indicated
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