Eukaryota

Spironema multiciliatum, from Lemmermann (1910).

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).

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.

Phylogenetic distribution of species historically assigned to Monas, from Boenigk (2008).

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 (from 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 1975FT93Ollula’ 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
  Prionognathodus Fay 1959 [=Prionognathus Pander 1856 non Ferté-Sénèctere 1851]Has62
  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.)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
  Anthophysa vegetansH04
  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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