Parabasalia
Taxonavigation
[edit]Superregnum: Eukaryota
Regnum: Protozoa
Subregnum: Eozoa
Infraregnum: Excavata
Phylum: Metamonada
Subphylum: Trichozoa
Superclassis: Parabasalia
Classes: Trichomonadea - Trichonymphea
Genera (incertae sedis):
Coelotrichomastix –
Cthulhu –
Cthylla –
Mesojoenia
Name
[edit]Parabasalia Honigberg, 1973: 187.
Similar groups:
- Parabasalea Hausmann & Hülsmann, 1996
- Parabasala Cavalier-Smith, 1997
References
[edit]- Cavalier-Smith, T. 2003. The excavate protozoan phyla Metamonada Grassé emend. (Anaeromonadea, Parabasalia, Carpediemonas, Eopharyngia) and Loukozoa emend. (Jakobea, Malawimonas): their evolutionary affinities and new higher taxa. International Journal of Systematic and Evolutionary Microbiology 53: 1741–1758. DOI: 10.1099/ijs.0.02548-0 Reference page.
- Cavalier-Smith T. 2003. Protist phylogeny and the high-level classification of Protozoa. European Journal of Protistology 39: 338–348. DOI: 10.1078/0932-4739-00002 Reference page.
- Čepička, I, Dolan, M.F., Gile, G.H. (2016). Parabasalia. In: Archibald, J.M., Simpson, A.G.B., Slamovits, C. (Eds.). Handbook of the Protists. Springer, pp. 1-44. DOI: 10.1007/978-3-319-32669-6_9-1
- Honigberg, B. M. 1973. Remarks upon trichomonad affinities of certain parasitic protozoa. In: Progress in Protozoology: Abstracts of Papers Read at the 4th International Congress of Protozoology, Clermont-Ferrand, 2–10 September 1973, p. 187. Edited by P. De Puytorac & J. Grain. Clermont-Ferrand: Université de Clermont.
Alternative classifications
[edit]John O. Corliss (1984)
[edit]- Corliss, J.O. 1984. The kingdom Protista and its 45 phyla. Biosystems 17(2): 87–126. DOI: 10.1016/0303-2647(84)90003-0 Reference page.
In Kingdom Protista, Assemblage "Polymastigotes"
Phylum Parabasalia Honigberg, 1973
Dyer in Margulis et al. (1990)
[edit]Dyer, B.D., 1990. 14i. Phylum Zoomastigina, Class Parabasalia. In: Margulis, L., J.O. Corliss, M. Melkonian, D.J. Chapman (ed.). Handbook of Protoctista. Jones and Bartlett Publishers, Boston, p. 252-258. See Brands, S.J. (1989-2015), [1].
Kingdom Protoctista
- Section III: Phyla with flagellated stages, but without complex sexual cycles
- Phylum 14. Zoomastigina
- Class Parabasalia
- Phylum 14. Zoomastigina
Cavalier-Smith (1993)
[edit]From Kingdom Protozoa and its 18 phyla
In Kingdom Protozoa, Subkingdom Dictyozoa
Branch Parabasalia new branch
Phylum Parabasalia Honigberg, 1973 stat. nov. Cavalier-Smith, 1981
- Class Trichomonadea
- Class Hypermastigea
Cavalier-Smith (1998)
[edit]From A revised six-kingdom system of life
In Phylum Trichozoa
Subphylum Parabasala Honigberg 1973 stat. nov. Cavalier-Smith 1997
David J. Patterson (1999)
[edit]From The Diversity of Eukaryotes
In Eukaryotes
Clade Parabasalids
Brugerolle & Lee, in Lee et al. (2000)
[edit]Brugerolle, G. & Lee, J.J., 2000. Phylum Parabasalia. In: Lee, J.J., Leedale, G.F. & Bradbury, P. An Illustrated Guide to the Protozoa. Society of Protozoologists/Allen Press: Lawrence, Kansas, U.S.A, 2nd ed., vol. 2, p. 1196-1250, [2]. See Brands, S.J. (1989-2015), [3].
In “protozoa”, “flagellated protists”
- Phylum Parabasalia
- Class Trichomonada
- 1 Family Monocercomonadidae Kirby, 1944
- 1 Subfamily Monocercomonadinae Kirby, 1944 [see Family Monocercomonadidae Kirby, 1944]
- 1 Genus Monocercomonas Grassi, 1879
- 2 Genus Eutrichomastix Kofoid & Swezy, 1915 [see Genus Monocercomonas Grassi, 1879]
- 3 Genus Tricercomitus Kirby, 1930
- 4 Genus Tetratrichomastix Mackinnon, 1913 [see Genus Hexamastix Alexeieff, 1912]
- 5 Genus Trichomastix Blochmann, 1884 [see Genus Tetratrichomastix Mackinnon, 1913]
- 6 Genus Hexamastix Alexeieff, 1912
- 2 Subfamily Chilomitinae Honigberg, 1963
- 1 Genus Chilomitus de Fonseca, 1915
- 3 Subfamily Protrichomonadinae
- 1 Genus Protrichomonas Alexeieff, 1911
- 2 Genus Parahistomonas Lund, 1963
- 3 Genus Histomonas Tyzzer, 1920
- 4 Subfamily Dientamoebinae Grassé, 1953
- 1 Genus Dientamoeba Dobell, 1940
- 5 Subfamily Hypotrichomonadinae Honigberg, 1963 [see Family Hypotrichomonadidae (Honigberg, 1963) Cepicka et al., 2010]
- 1 Genus Hypotrichomonas Lee, 1960
- 2 Genus Pseudotrichomonas Bishop, 1939
- 3 Genus Ditrichomonas Farmer, 1993
- 1 Subfamily Monocercomonadinae Kirby, 1944 [see Family Monocercomonadidae Kirby, 1944]
- 2 Family Trichomonadidae Chalmers & Pekkola, 1918
- 1 Subfamily Tritrichomonadinae Honigberg, 1963 [see Family Tritrichomonadidae Honigberg, 1963]
- 1 Genus Tritrichomonas Kofoid, 1920
- 2 Subfamily Trichomonadinae Chalmers & Pekkola, 1918
- 1 Genus Trichomitus Swezy, 1915
- 2 Genus Tetratrichomonas Parisi, 1910
- 3 Genus Trichomonas Donné, 1836
- 4 Genus Pentatrichomonas Mesnil, 1914
- 3 Subfamily Trichomitopsinae Brugerolle, 1977
- 1 Genus Trichomitopsis Kofoid & Swezy, 1919
- 2 Genus Pseudotrypanosoma Grassi, 1917
- 4 Subfamily Pentatrichomonoidinae Honigberg, 1963
- 1 Genus Pentatrichomonoides Kirby, 1931
- 1 Subfamily Tritrichomonadinae Honigberg, 1963 [see Family Tritrichomonadidae Honigberg, 1963]
- 3 Family Devescovinidae Doflein, 1911 [see Family Lophomonadidae Stein]
- 1 Subfamily Devescovininae Doflein, 1911
- 01 Genus Foaina Janicki, 1915
- 02 Genus Crucinympha Kirby, 1939 [see Genus Foaina Janicki, 1915]
- 03 Genus Janickiella Duboscq & Grassé [see Genus Foaina Janicki, 1915]
- 04 Genus Paradevescovina Kirby [see Genus Foaina Janicki, 1915]
- 05 Genus Parajoenia Janicki, 1911
- 06 Genus Devescovina Foà, 1905
- 07 Genus Metadevescovina Light, 1926
- 08 Genus Caduceia França, 1918
- 09 Genus Bullanympha Kirby, 1939
- 10 Genus Hyperdevescovina Kirby, 1947
- 11 Genus Pseudodevescovina Sutherland, 1933
- 12 Genus Polymastigoides Grassé & Hollande, 1951
- 13 Genus Achemon Grassé & Hollande, 1950
- 14 Genus Astronympha Grassé, 1952
- 15 Genus Evemonia Grassé & Hollande, 1950
- 16 Genus Mixotricha Sutherland, 1933
- 17 Genus Kirbynia Grassé & Hollande, 1950
- 2 Subfamily Gigantomonadinae H. Kirby, 1946
- 1 Genus Gigantomonas Dogiel, 1916
- 2 Genus Macrotrichomonas Grassi, 1917
- 1 Subfamily Devescovininae Doflein, 1911
- 4 Family Calonymphidae Grassi & Foa, 1911 [see Family Lophomonadidae Stein]
- 1 Genus Coronympha Kirby, 1929
- 2 Genus Metacoronympha Kirby, 1939
- 3 Genus Stephanonympha Janicki, 1911
- 4 Genus Diplonympha Grassi, 1920
- 5 Genus Calonympha Foà, 1905
- 6 Genus Snyderella Kirby, 1929
- 5 Family Cochlosomatidae E.E. Tyzzer, 1930 [see Family Trichomonadidae Chalmers & Pekkola, 1918]
- 1 Genus Cochlosoma Kotlán, 1923
- 2 Genus Cyathosoma Tyzzer, 1930
- 3 Genus Ptychostoma Tyzzer, 1930
- 1 Family Monocercomonadidae Kirby, 1944
- Class Hypermastigia
- 1 Order Lophomonadida Light, 1927 [see Order Cristamonadida Brugerolle & Patterson, 2001]
- 1 Family Joeniidae Janicki [see Family Lophomonadidae Stein]
- 1 Genus Joenia Grassi, 1917
- 2 Genus Placojoenia Radek & Hausmann, 1994
- 3 Genus Joenina Grassi, 1885
- 4 Genus Joenoides Grassé, 1952
- 5 Genus Joenopsis Cutler, 1920
- 6 ?Genus Cyclojoenia Nurse, 1945
- 7 Genus Projoenia Lavette, 1970
- 2 Family Lophomonadidae Stein
- 1 Genus Lophomonas Stein, 1860
- 3 Family Rhizonymphidae [see Family Lophomonadidae Stein]
- 1 Genus Rhizonympha Grassé & Hollande, 1951
- 4 Family Microjoeniidae Grassé, 1952 [see Family Holomastigotoididae Grassi & Foà, 1911]
- 1 Genus Microjoenia Grassi, 1892
- 2 Genus Torquenympha Brown, 1930 [see Genus Microjoenia Grassi, 1892]
- 5 Family Kofoidiidae [see Family Lophomonadidae Stein]
- 1 Genus Kofoidia Light, 1927
- 6 Family Deltotrichonymphidae [see Family Lophomonadidae Stein]
- 1 Genus Deltotrichonympha Sutherland, 1933
- 2 Genus Koruga Cleveland, 1966
- 1 Family Joeniidae Janicki [see Family Lophomonadidae Stein]
- 2 Order Trichonymphida Poche, 1913
- 1 Family Hoplonymphidae
- 1 Genus Barbulanympha Cleveland et al., 1934
- 2 Genus Hoplonympha Light, 1926
- 3 Genus Urinympha Cleveland, Hall, Sanders & Collier, 1934
- 4 Genus Rhynchonympha Cleveland, Hall, Sanders & Collier, 1934
- 2 Family Staurojoeninidae
- 1 Genus Staurojoenina Kirby, 1926
- 2 Genus Idionympha Cleveland, Hall, Sanders & Collier, 1934
- 3 Family Trichonymphidae Kent, 1880
- 1 Genus Trichonympha Leidy, 1877
- 4 Family Eucomonymphidae [see Family Teranymphidae]
- 1 Genus Eucomonympha Cleveland et al., 1934
- 2 Genus Pseudotrichonympha Grassi & Foà, 1911
- 5 Family Teranymphidae
- 1 Genus Teranympha Koidzumi, 1917
- 2 Genus Teratonympha Koidzumi, 1921 [see Genus Teranympha Koidzumi, 1917]
- 3 Genus Cyclonympha Dogiel, 1917 [see Genus Teranympha Koidzumi, 1917]
- 6 Family Spirotrichosomidae
- 1 Genus Macrospironympha Cleveland et al., 1934
- 2 Genus Leptospironympha Cleveland et al., 1934
- 3 Genus Spirotrichosoma Sutherland, 1933
- 4 Genus Apospironympha Cleveland & Day, 1958
- 5 Genus Colospironympha Cleveland & Day, 1958
- 1 Family Hoplonymphidae
- 3 Order Spirotrichonymphida Grassé, 1952
- 1 Family Spirotrichonymphidae Grassi, 1917 [see Family Holomastigotoididae Grassi & Foà, 1911]
- 1 Genus Spirotrichonympha Grassi & Foà, 1911
- 2 Genus Spirotrichonymphella Grassi, 1917
- 3 ?Genus Spironympha Koidzumi, 1921
- 4 Genus Microspironympha Koidzumi, 1921 [see Genus Spironympha Koidzumi, 1921]
- 5 ?Genus Micromastigotes Hollande & Carruette-Valentin, 1971
- 2 Family Holomastigotoididae Grassi & Foà, 1911
- 1 Genus Holomastigotoides Grassi & Foà, 1911
- 2 Genus Rostronympha Duboscq, Grassé & Rose, 1937
- 3 Family Holomastigotidae Janicky, 1915 [see Family Holomastigotoididae Grassi & Foà, 1911]
- 1 Genus Holomastigotes Grassi, 1892
- 2 ?Genus Spiromastigotes Duboscq & Grassé, 1943
- 1 Family Spirotrichonymphidae Grassi, 1917 [see Family Holomastigotoididae Grassi & Foà, 1911]
- 1 Order Lophomonadida Light, 1927 [see Order Cristamonadida Brugerolle & Patterson, 2001]
- Class Trichomonada
Cavalier-Smith (2002)
[edit]From The phagotrophic origin of eukaryotes and phylogenetic classification of Protozoa
In Kingdom Protozoa, Subkingdom Corticata, Infrakingdom Excavata, Superphylum Archezoa
Phylum Parabasalia
Alastair G. B. Simpson (2003)
[edit]From Cytoskeletal organization, phylogenetic affinities and systematics in the contentious taxon Excavata (Eukaryota)
In Excavata
Clade Parabasalia
Cepicka et al. (2010)
[edit]Cepicka, I., Hampl, V., & Kulda, J. (2010). Critical taxonomic revision of parabasalids with description of one new genus and three new species. Protist 161(3), 400-433, [4], [5].
Parabasalia Honigberg, 1973
- 1 Class Hypotrichomonadea Čepička et al., 2010
- Family Hypotrichomonadidae Honigberg, 1963 (= Trichomitidae Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006)
- 1 Genus Hypotrichomonas Lee, 1960
- 2 Genus Trichomitus Swezy, 1915
- Family Hypotrichomonadidae Honigberg, 1963 (= Trichomitidae Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006)
- 2 Class Trichomonadea Kirby, 1947 sensu Margulis, 1974
- Order Trichomonadida Kirby, 1947
- Family Trichomonadidae Chalmers & Pekkola, 1918 sensu Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006
- Genus Trichomonas Donné, 1836
- Genus Tetratrichomonas Parisi, 1910
- Genus Pentatrichomonas Mesnil, 1914
- Genus Pseudotrypanosoma Grassi, 1917
- Genus Trichomitopsis Kofoid & Swezy, 1919
- Genus Cochlosoma Kotlán, 1923
- Genus Pentatrichomonoides Kirby, 1931
- Genus Trichomonoides Brugerolle & Bordereau, 2004
- Family Trichomonadidae Chalmers & Pekkola, 1918 sensu Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006
- 2 Order Honigbergiellida Čepička et al., 2010
- 1 Family Honigbergiellidae Čepička et al., 2010
- Genus Honigbergiella Hampl, Čepička & Kulda, 2007
- Genus Pseudotrichomonas Bishop, 1939
- Genus Ditrichomonas Farmer, 1993
- Genus Monotrichomonas Bernard, Simpson & Patterson, 2000
- 2 Family Hexamastigidae Čepička et al., 2010
- Genus Hexamastix Alexeieff, 1912
- Genus Tetratrichomastix Mackinnon, 1913
- 3 Family Tricercomitidae Čepička et al., 2010
- 1 Genus Tricercomitus Kirby, 1930
- 1 Family Honigbergiellidae Čepička et al., 2010
- Order Trichomonadida Kirby, 1947
- 3 Class Tritrichomonadea Cepicka et al., 2010
- Order Tritrichomonadida Čepička et al., 2010
- Family Tritrichomonadidae Honigberg, 1963 sensu Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006
- 1 Genus Tritrichomonas Kofoid, 1920
- Family Simplicimonadidae Čepička et al., 2010
- 1 Genus Simplicimonas Čepička et al., 2010
- Family Monocercomonadidae Kirby, 1944
- 1 Genus Monocercomonas Grassi, 1879
- Family Dientamoebidae Grassé, 1953 (= Protrichomonadinae Honigberg & Kuldová, 1969)
- Genus Dientamoeba Dobell, 1940
- Genus Protrichomonas Alexeieff, 1911
- Genus Histomonas Tyzzer, 1920
- Genus Parahistomonas Honigberg & Kuldová, 1969
- Family Tritrichomonadidae Honigberg, 1963 sensu Hampl, Vrlík, Cepicka, Pecka, Kulda & Tachezy, 2006
- Order Tritrichomonadida Čepička et al., 2010
- 4 Class Cristamonadea (Brugerolle & Patterson, 2001) Cepicka et al., 2010
- Order Cristamonadida Brugerolle & Patterson, 2001
- Family Lophomonadidae Kent, 1880 (= Calonymphidae Grassi, 1911; Devescovinidae Doflein, 1911; Joeniidae Janicki, 1915; Kofoidiidae Light, 1927; Deltotrichonymphidae; Gigantomonadinae Kirby, 1944; Rhizonymphidae Grassé & Hollande, 1951)
- Genus Lophomonas Grassi, 1885 [Stein, 1860?]
- Genus Joenia Grassi, 1885
- Genus Calonympha Foà, 1905
- Genus Devescovina Foà, 1905
- Genus Stephanonympha Janicki, 1911
- Genus Foaina Janicki, 1915
- Genus Gigantomonas Dogiel, 1916
- Genus Caduceia França, 1918
- Genus Diplonympha Grassi, 1920
- Genus Joenina Grassi, 1920
- Genus Joenopsis Cutler, 1920
- Genus Macrotrichomonas Grassi, 1920
- Genus Metadevescovina Light, 1926
- Genus Kofoidia Light, 1927
- Genus Coronympha Kirby, 1929
- Genus Snyderella Kirby, 1929
- Genus Deltotrichonympha Sutherland, 1933
- Genus Mixotricha Sutherland, 1933
- Genus Pseudodevescovina Sutherland, 1933
- Genus Prolophomonas Cleveland, Hall, Sanders & Collier, 1934
- Genus Bullanympha Kirby, 1939
- Genus Metacoronympha Kirby, 1939
- Genus Cyclojoenia Nurse, 1945
- Genus Hyperdevescovina Kirby, 1947
- Genus Achemon Grassé & Hollande, 1950
- Genus Evemonia Grassé & Hollande, 1950
- Genus Kirbynia Grassé & Hollande, 1950
- Genus Polymastigoides Grassé & Hollande, 1951
- Genus Rhizonympha Grassé & Hollande, 1951
- Genus Astronympha Grassé, 1952
- Genus Joenoides Grassé, 1952
- Genus Koruga Cleveland, 1966
- Genus Projoenia Lavette, 1970
- Genus Placojoenia Radek & Hausmann, 1994
- Genus Criconympha Dolan & Kirby, 2002
- Genus Gyronympha Dolan & Kirby, 2002
- Genus Prosnyderella Dolan & Kirby, 2002
- Genus Pachyjoenia Brugerolle & Bordereau, 2005
- Family Lophomonadidae Kent, 1880 (= Calonymphidae Grassi, 1911; Devescovinidae Doflein, 1911; Joeniidae Janicki, 1915; Kofoidiidae Light, 1927; Deltotrichonymphidae; Gigantomonadinae Kirby, 1944; Rhizonymphidae Grassé & Hollande, 1951)
- Order Cristamonadida Brugerolle & Patterson, 2001
- 5 Class Trichonymphea Cavalier-Smith, 2003
- Order Trichonymphida Poche, 1913 sensu Cavalier-Smith,2003
- Family Trichonymphidae Kent, 1880
- Genus Trichonympha Leidy, 1877
- Family Teranymphidae Koidzumi, 1921 (= Eucomonymphidae Cleveland, Hall, Sanders & Collier, 1934)
- Genus Teranympha Koidzumi, 1917
- Genus Pseudotrichonympha Grassi & Foà, 1911
- Genus Eucomonympha Cleveland, Hall, Sanders & Collier, 1934
- Family Spirotrichosomidae Hollande & Caruette-Valentin, 1971
- Genus Spirotrichosoma Sutherland, 1933
- Genus Macrospironympha Cleveland, Hall, Sanders & Collier, 1934
- Genus Leptospironympha Cleveland, Hall, Sanders & Collier, 1934
- Genus Apospironympha Cleveland & Day, 1958
- Genus Colospironympha Cleveland & Day, 1958
- Genus Bispironympha Bobyleva, 1969
- Family Hoplonymphidae Light, 1926
- Genus Hoplonympha Light, 1926
- Genus Barbulanympha Cleveland, Hall, Sanders & Collier, 1934
- Genus Rhynchonympha Cleveland, Hall, Sanders & Collier, 1934
- Genus Urinympha Cleveland, Hall, Sanders & Collier, 1934
- Family Staurojoeninidae Grassi, 1917
- Genus Staurojoenina Kirby, 1926
- Genus Idionympha Cleveland, Hall, Sanders & Collier, 1934
- Family Trichonymphidae Kent, 1880
- Order Trichonymphida Poche, 1913 sensu Cavalier-Smith,2003
- 6 Class Spirotrichonymphea Grassé, 1952 stat. nov.
- Order Spirotrichonymphida Grassé, 1952
- Family Holomastigotoididae Grassi & Foà, 1911 (= Holomastigotidae Janicky, 1915; Spirotrichonymphidae Grassi, 1917; Microjoeniidae Grassé, 1952)
- Genus Holomastigotoides Grassi & Foà, 1911
- Genus Holomastigotes Grassi, 1892
- Genus Microjoenia Grassi, 1892
- Genus Spirotrichonympha Grassi & Foà, 1911
- Genus Spironympha Koidzumi, 1917
- Genus Spirotrichonymphella Grassi, 1917
- Genus Rostronympha Duboscq, Grassé & Rose, 1937
- Genus Spiromastigotes Duboscq & Grassé, 1943
- Genus Micromastigotes Hollande & Carruette-Valentin, 1971
- Genus Uteronympha Brugerolle, 2006
- Family Holomastigotoididae Grassi & Foà, 1911 (= Holomastigotidae Janicky, 1915; Spirotrichonymphidae Grassi, 1917; Microjoeniidae Grassé, 1952)
- Order Spirotrichonymphida Grassé, 1952
- ?Genus Trichocovina Maaß & Radek, 2006
- ?Genus Chilomitus de Fonseca, 1915
Adl et al. (2012)
[edit]- Adl, S.M., Simpson, A.G.B., Lane, C.E., Lukeš, J., Bass, D., Bowser, S.S., Brown, M.W., Burki, F., Dunthorn, M., Hampl, V., Heiss, A., Hoppenrath, M., Lara, E., Le Gall, L., Lynn, D.H., McManus, H., Mitchell, E.A.D., Mozley-Stanridge, S.E., Parfrey, L.W., Pawlowski, J., Rueckert, S., Shadwick, L., Schoch, C.L., Smirnow, A. & Spiegel, F.W. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59(5): 429–514. DOI: 10.1111/j.1550-7408.2012.00644.x . [Erratum: 60(3): 321. DOI: 10.1111/jeu.12033 .] Reference page.
- Excavata Cavalier-Smith 2002, emend. Simpson 2003
- Metamonada Cavalier-Smith 1987 (= Metamonadina Grassé 1952), emend. Cavalier-Smith 2003
- Parabasalia Honigberg 1973
- Trichomonadea Kirby 1947, sensu Margulis 1974, emend. Čepička et al. 2010 (P?)
- Hypotrichomonadea Čepička et al. 2010
- Tritrichomonadea Čepička et al. 2010 (P?)
- Cristamonadea Brugerolle & Patterson 2001, sensu Čepička et al. 2010
- Trichonymphea Poche 1913, sensu Cavalier-Smith 2003
- Spirotrichonymphea Grassé 1952, sensu Čepička et al. 2010
- Parabasalia Honigberg 1973
- Metamonada Cavalier-Smith 1987 (= Metamonadina Grassé 1952), emend. Cavalier-Smith 2003