Free-living Gymnamoebia (Rhizopoda, Lobosea): a checklist of species,
with illustrations, advices on identification and guide to the literature
Appeared online: 09.10.2000. Last revised: 02.04.2003
- Development of amoebae systematics produced lots of species names and
many different systems, based on various background. The result is a a huge
number of synonims and certain mess with the names and position of genera and
families. Since F.C. Page published his revised classification of gymnamoebae
in 1987, no real attempt was done to deal with the accumulating changes in
amoebae systematics (long expected second edition of the "Illustrated Guide to
Protozoa" is a special matter that I'll discuss later). The need in the list
like this thus seems to be evident.
- This list includes all valid and recognisable (i.e. species which
we can identify with sufficient level of confidence) species of Gymnamoebia
(Rhizopoda, Lobosea). The criteria of inclusion are:
- Species must be recognisable with sufficient level of confidence.
For example, "Amoeba annulata" Penard 1902 was documented with
few line drawings, but the description contained several unique
features of this species, that finally allowed to recognise and
re-describe it (Smirnov & Goodkov, 1998). In contrast,
"Amoeba elongata" Korotneff, 1880 - the "type" species of the
genus Dactylamoeba sensu F.C. Page (listed as Korotnevella
in the present list, see Smirnov, 1997 for details) has no distinctiv
features leaving any chance to re-isolate it and thus is not listed
here.
- Species description must be valid, i.e. to fit the basic taxonomical
requirements (see International Code of Zoological Nomenclature).
It may look old fashioned, but I do call to keep within the
frames of the code. With all its disadvantages, it accumulates
a lot of experience of taxonomists and allow certain standartisation of
the usage and creation of names, so important in systematics.
I do not include species and genera established solely basing on
biochemical (e.g. "Metamoeba" Friz 1999) and molecular data,
without proper diagnoses.
The reason is that the recognition of these genera and species
among usual morphospecies will evidently produce a mess related with
the unequal level of taxonomical resolution through the taxa.
- Identification of an isolate as a species, not listed here, requires
special motivation and respective taxonomically valid re-description of
a species prior to use its name.
- I will develop this list as a guide to Gymnamoebia, incorporating
illustrations, brief species descriptions, advices on identification and
basic references and sources of strains for each species. This is a lasting
work, so I will add new materials step by step. If the species name is a
hypertext reference - this means that there are materials on the species.
Class Lobosea Carpenter, 1861
Subclass Gymnamoebia Haeckel, 1866
Order Euamoebida Lepsi, 1960
Family Amoebidae Ehrenberg, 1838
- Amoeba proteus (Pallas, 1766) Leidy, 1878
- Amoeba borokensis Kalinina, Afon'kin, Gromov, Khrebtukova & Page, 1987
- Amoeba leningradensis Page & Kalinina, 1984
- Amoeba amazonas Flickinger, 1974
- Chaos carolinense (Willson, 1900) King & Jahn, 1948
- Chaos illinoisense (Kudo, 1950) Bovee & Jahn, 1973
- Chaos nobile (Penard, 1902) Bovee & Jahn, 1973
- Chaos glabrum Smirnov & Goodkov, 1997
- Polychaos fasciculatum (Penard, 1902) Schaeffer, 1926
- Polychaos annulatum (Penard, 1902) Smirnov & Goodkov, 1998
- Polychaos dubium (Schaeffer, 1916) Schaeffer, 1926
- Polychaos nitidubium Bovee, 1970 ultrastructure unknown, needs re-isolation
- Deuteramoeba algonquinensis (Baldock, Rogerson & Berger, 1983) Page, 1987
- Deuteramoeba mycophaga (Pussard, Alabouvette & Pons, 1980) Page, 1988
- Trichamoeba sinuosa Siemensma & Page, 1986
- Trichamoeba myakka Bovee, 1972 ultrastructure unknown, needs re-isolation
- Trichamoeba cloaca Bovee, 1972 ultrastructure unknown, needs re-isolation
- Trichamoeba osseosaccus Schaeffer, 1926 ultrastructure unknown, needs re-isolation
- Parachaos zoochlorellae (Willumsen, 1982) Willumsen, Siemensma & Suhr-Jessen, 1987
- Hydramoeba hydroxena (Entz, 1912) Reynolds & Looper, 1928
Family Hartmannellidae (Volkonsky, 1931) Page, 1974
- Saccamoeba stagnicola Page, 1974
- Saccamoeba limax (Dujardin, 1841) Page, 1974
- Saccamoeba lucens Frenzel, 1892 ultrastructure unknown, needs re-isolation
- Saccamoeba wakulla Bovee, 1972 ultrastructure unknown, needs re-isolation
- Saccamoeba limna Bovee, 1972 ultrastructure unknown, needs re-isolation
- Saccamoeba wellneri Siemensma, 1987 ultrastructure unknown, needs re-isolation
- Saccamoeba marina Anderson, Rogerson & Hannah, 1997
- Cashia limacoides (Page, 1967) Page, 1974
- Glaeseria mira (Glaeser, 1912) Volkonsky, 1931
- Hartmannella cantabrigiensis Page, 1974
- Hartmannella vermiformis Page, 1967
- Hartmannella lobifera Smirnov, 1997
- Hartmannella abertawensis Page, 1980
- Hartmannella vacuolata Anderson, Rogerson & Hannah, 1997
- Nolandella hibernica (Page, 1980) Page, 1983
Family Thecamoebidae (Schaeffer, 1926) Smirnov & Goodkov, 1994
- Thecamoeba striata (Penard, 1890) Schaeffer, 1926
- Thecamoeba quadrilineata (Carter, 1856) Lepsi, 1960
- Thecamoeba munda (Schaeffer, 1926) Smirnov, 1999
- Thecamoeba orbis Schaeffer, 1926
- Thecamoeba hilla Schaeffer, 1926
- Thecamoeba sparolata Fishbeck & Bovee, 1993
- Thecamoeba sphaeronucleolus (Greef, 1891) Schaeffer, 1926
- Thecamoeba verrucosa (Ehrenberg, 1838) Schaeffer, 1926
- Thecamoeba terricola (Greef, 1866) Lepsi, 1960
- Thecamoeba similis (Greef, 1891) Lepsi, 1960
- Sappinia diploidea (Hartmann & Naegler, 1908) Alexieff, 1912
- Dermamoeba granifera (Greef, 1866) Page & Blakey, 1979
- Dermamoeba minor (Pussard, Alabouvette & Pons, 1979) Page, 1988
- Paradermamoeba valamo Smirnov & Goodkov, 1993
- Paradermamoeba levis Smirnov & Goodkov, 1994
- Pseudothecamoeba proteoides (Page, 1976) Page, 1988
- Thecochaos album (Greef, 1891) Page, 1981 known from stained preparations only, needs re-isolation
- Thecochaos fibrillosum (Greef, 1891) Page, 1981 known from stained preparations only, needs re-isolation
Family Vannellidae (Bovee, 1970) Page, 1987
- Vannella simplex (Wohlfarth-Bottermann, 1960) Smirnov, Nassonova, Holzmann and Pawlowski, 2002
- Vannella platypodia (Glaeser, 1912) Page, 1976
- Vannella lata Page, 1988
- Vannella cirifera (Frenzel, 1892) Page, 1988
- Vannella miroides Bovee, 1965
- Vannella persistens Smirnov & Brown, 2000
- Vannella peregrinia Smirnov & Fenchel, 1996
- Vannella devonica Page, 1979
- Vannella aberdonica Page, 1980
- Vannella caledonica Page, 1979
- Vannella arabica Page, 1980
- Vannella anglica Page, 1980
- Vannella serpentrionalis Page, 1980
- Vannella sensilis Bovee & Sawyer, 1979 ultrastructure unknown, needs re-isolation
- Vannella mira (Schaeffer, 1926) Smirnov, 2002
- Vannella ebro Smirnov, 2001
- Platyamoeba placida (Page, 1968) Page, 1969
- Platyamoeba stenopodia Page, 1969
- Platyamoeba schaefferi Singh & Hanumaiah, 1979
- Platyamoeba plurinucleolus Page, 1974
- Platyamoeba calycinucleolus Page, 1974
- Platyamoeba bursella Page, 1974
- Platyamoeba australis Page, 1983
- Platyamoeba mainensis Page, 1971
- Platyamoeba flabellata Page, 1974
- Platyamoeba pseudovannellida Hauger, Rogerson & Anderson, 2001
- Platyamoeba murchelanoi Sawyer, 1975 ultrastructure unknown, generic position questionable
- Platyamoeba langae Sawyer, 1975 ultrastructure unknown, generic position questionable
- Platyamoeba weinsteini Sawyer, 1975 ultrastructure unknown, generic position questionable
- Platyamoeba douvresi Sawyer, 1975 ultrastructure unknown, generic position questionable
- "Clydonella" rosenfeldi Sawyer, 1975 ultrastructure unknown, generic position questionable
- "Clydonella" sindermanni Sawyer, 1975 ultrastructure unknown, generic position questionable
- "Clydonella" wardi Sawyer, 1975 ultrastructure unknown, generic position questionable
- "Lingulamoeba" leei Sawyer, 1975 ultrastructure unknown, generic position questionable
- "Pessonella" marginata Pussard, 1973 ultrastructure unknown, generic position questionable
The genera Clydonella and Lingulamoeba (Sawyer, 1975-Trans. Amer. Microsc. Soc.
94:305-323) and Pessonella (Pussard, 1973-Protistologica 9:175-185) are
established basing on LM data only. Their representatives needs EM study to
confirm the validity of mentioned genera. I have TEM indications that
the cell coat of Pessonella is identical to that of Vannella
and the cell coat of Clydonella is organised like in Platyamoeba
(Smirnov, unpubl.). Both mentioned genera may be closed in future, and their
species will be distributed between Vannella and Platyamoeba, respectively.
Family Paramoebidae (Poche, 1913) Page, 1987
- Mayorella viridis (Leidy, 1874) Harnisch, 1968
- Mayorella cantabrigiensis Page, 1983
- Mayorella vespertilioides Page, 1983
- Mayorella penardi Page, 1972
- Mayorella bigemma (Schaeffer, 1926) Schaeffer, 1926 ultrastructure unknown, needs re-isolation
- Mayorella augusta Schaeffer, 1926 ultrastructure unknown, needs re-isolation
- Mayorella pussardi Hollande, Nicolas & Escaig, 1981
- Mayorella kuwaitensis (Page, 1982) Page, 1983
- Mayorella gemmifera Schaeffer, 1926
- Mayorella dactylifera Goodkov & Buryakov, 1986
- Korotnevella stella (Schaeffer, 1926) Goodkov, 1988
- Korotnevella bulla (Schaeffer, 1926) Goodkov, 1988
- Korotnevella diskophora Smirnov, 1999
- Korotnevella nivo Smirnov, 1997
- Korotnevella hemistilolepis O'Kelly, Peglar, Black, Sawyer & Nerad, 2001
- Korotnevella monacantholepis O'Kelly, Peglar, Black, Sawyer & Nerad, 2001
- Paramoeba eilhardi Schaudinn, 1896
E.Bovee (1951-Proc. Iowa Acad. Sci. 58: 415-421; 1961-Trans. Amer. Microsc.
Soc. 80: 54-62; 1970-Arch. Protistenk. 112: 178-227) lists 19 more Mayorella-like and
Korotnevella-like amoebae combined in the genus Mayorella and 4 more
genera (Flagellipodium Bovee, 1961; Oscillosignum Bovee, 1951;
Subulamoeba Bovee, 1951 and Trienamoeba Bovee, 1970); all
these species need re-isolation and study prior to be recognised as
a valid ones and assigned to any genus; the same is true for all 4 mentioned
genera.
F.C. Page in his keys use the generic name "Dactylamoeba
Korotneff, 1880" (Page, 1981-Protistologica 17: 543-562)
which was replaced with the name "Korotnevella" by
Goodkov (1988-Zoologicheskij journal 67: 1728-1730). The generic name
"Dactylamoeba" is incorrect and should not be used. See Smirnov
(1997-Arch. Protistenk. 147: 283-292) for details.
Family Vexilliferidae Page, 1987
- Vexillifera bacillipedes Page, 1969
- Vexillifera granatensis Mascaro, Osuna & Mascaro, 1986
- Vexillifera lemani Page, 1976 ultrastructure unknown, needs re-isolation
- Vexillifera minutissima Bovee & Sawyer, 1979 ultrastructure unknown, needs re-isolation
- Vexillifera armata Page, 1979
- Vexillifera telmathalassa Bovee, 1956 ultrastructure unknown, needs re-isolation
- Vexillifera aurea Schaeffer, 1926 ultrastructure unknown, needs re-isolation
- Vexillifera browni Sawyer, 1975 ultrastructure unknown, needs re-isolation
- Vexillifera ottoi Sawyer, 1975 ultrastructure unknown, needs re-isolation
- Pseudoparamoeba pagei (Sawyer, 1975) Page, 1983
- Neoparamoeba pemaquidensis Page, 1970
- Neoparamoeba aestuarina Page, 1970
There are little-known genera of Vexillifera-like amoebae Boveella
Sawyer (1975-Trans. Amer. Microsc. Soc. 94:71-92) Trienamoeba (Bovee,
1970 - Arch.Protistenk. 112:178-227). Representatives of both must be studied
with EM to make a desision about the validity and position of these genera.
Order Leptomyxida (Pussard & Pons, 1976) Page, 1987
Suborder Rhizoflabellina Page, 1987
Family Flabellulidae Bovee, 1970
- Flabellula citata Schaeffer, 1926
- Flabellula calkinsi (Hogue, 1914) Page, 1983
- Flabellula demetica Page, 1980
- Flabellula trinovanica Page, 1980
- Flabellula baltica Smirnov, 1999
- Paraflabellula kudoi (Singh & Hanumaiah, 1979) Page, 1983
- Paraflabellula reniformis (Schmoller, 1964) Page, 1983
- Paraflabellula hoguae (Sawyer, 1975) Page, 1983
- Flamella citrensis Bovee, 1956
- Flamella magnifica Schaeffer, 1926
- Flamella tiara Fishbeck & Bovee, 1993
- Flamella lacustris Michel & Smirnov, 1999
- Flamella aegyptia Michel & Smirnov, 1999
Family Leptomyxidae (Pussard & Pons, 1976) Page, 1987
- Leptomyxa reticulata Goodey, 1914
- Leptomyxa fragilis (Penard, 1904) Siemensma, 1987
- Rhizamoeba polyura Page, 1972
- Rhizamoeba saxonica Page, 1974
- Rhizamoeba australiensis (Chakraborty & Pussard, 1985) Page, 1988
- Rhizamoeba schnepfi
- Rhizamoeba clavarioides (Penard, 1902) Siemensma, 1980
- Rhizamoeba flabellata (Goodey, 1914) Cann, 1984
- Rhizamoeba caerulea (Schaeffer, 1926) Siemensma, 1980
Suborder Leptoramosina Page, 1987
Family Gephyramoebidae Pussard & Pons, 1976
- Gephyramoeba delicatula Goodey, 1914
Family Stereomyxidae Grell, 1966
- Stereomyxa ramosa Grell, 1966
- Stereomyxa angulosa Grell, 1966
Order Loboreticulatida Page, 1987
Family Corallomyxidae Page, 1987
- Corallomyxa chatonni Grell & Benwitz, 1978
- Corallomyxa mutabilis Grell, 1966
- Corallomyxa multipara Grell, 1988
Order Acanthopodida Page, 1976
Family Acanthamoebidae Sawyer & Griffin, 1975
Species composition of the genus Acanthamoeba is questionable, and
non-morphological methods require to distinguish some species. There are
serious disagreements between authors about the validity of species
and sinonymics. In contrast with other sections of this checklist, here
only approximate species composition is listed - namely,
species, recognisable at the morphological level and accepted by most
of scientists. I would be grateful for any comments on the composition of the
genus Acanthamoebae.
- Acanthamoeba tubiashi Lewis & Sawyer, 1979
- Acanthamoeba astronixis (Ray & Hayes, 1954) Page, 1967
- Acanthamoeba comandoni Pussard, 1964
- Acanthamoeba lugdunensis Pussard & pons, 1977
- Acanthamoeba castellanii (Douglas, 1930) Volkonski, 1931
- Acanthamoeba rhysodes (Singh, 1952) Griffin, 1972
- Acanthamoeba mauritaniensis Pussard & Pons, 1977
- Acanthamoeba polyphaga (Pushkarew, 1913) Volkonsky, 1931
- Acanthamoeba griffini Sawyer, 1971
- Acanthamoeba quina Pussard & Pons, 1977
- Acanthamoeba divionensis Pussard & Pons, 1977
- Acanthamoeba triangularis Pussard & Pons, 1977
- Acanthamoeba hatchetti Sawyer, Visvesvara & Harke, 1977
- Acanthamoeba palestinensis (Reich, 1933) Page, 1977 the species Acanthamoeba pustulosa we accept to be a synonyme of A. palestinensis
- Acanthamoeba royreba Willaert, Stevens & Tyndall, 1978
- Acanthamoeba culbertsoni (Singh & Das, 1970) Griffin, 1972
- Acanthamoeba lenticulata Molet & Ermolieff-Braun, 1976
- Acanthamoeba gigantea Schmoller, 1964
- Acanthamoeba jacobsi Sawyer, Nerad & Visvesvara, 1992
- Acanthamoeba stevensoni Sawyer, Nerad, Lewis & McLaughlin, 1993
- Acanthamoeba pearcei Nerad, Sawyer, Levis & McLaughlin, 1995
- Acanthamoeba healyi Moura, Wallace & Visvesvara, 1992
- Protacanthamoeba caledonica Page, 1981
- Protacanthamoeba invadens (Singh & Hanumaiah, 1979) Page, 1981
Gymnamoebia incertae sedis
- Family Echinamoebidae Page, 1975
- Echinamoeba exudans Page, 1975
- Echinamoeba silvestris Page, 1975
- Filamoeba nolandi Page, 1967
- Comandonia operculata Pernin & Pussard, 1979
Rhizopoda incertae sedis
- Genus Stygamoeba (Sawyer, 1975) Smirnov, 1995
- Stygamoeba polymorpha Sawyer, 1975
- Stygamoeba regulata Smirnov, 1995
Family Entamoebidae Chatonn, 1925
the only probably free-living member of this family is
Entamoeba moshkovskii Chalaja, 1941
All other species are obligate parasites