Arcella

Dorsal view of Arcella vulgaris, from Microworld.

Belongs within: Arcellinida.

Arcella is a genus of amoeboids with a rigid, chitinous test found in brackish to fresh water (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Test membranous, transparent, chitinoid, punctate or minutely cancellate, rarely spinose, hemispherical, recurved at circular to crenulate aperture (av. diam. 70µ), young tests hyaline, adult brown; protoplasm centrally placed, attached to interior of test by threads of ectoplasm, may secrete gas vacuoles, to serve hydrostatic function, allowing individual to float, and later resorb vacuoles in order to settle; pseudopodia few, lobose, blunt; commonly binucleate, but in some species as many as 200 nuclei occur; 4 or more small contractile vacuoles; reproduction by "budding".

<==Arcella Ehrenberg 1832 (see below for synonymy) LT64
    |--*A. vulgaris Ehrenberg 1832 [=A. (*Euarcella) vulgaris, A. (*Sticholepis) vulgaris] LT64
    |--A. arcelloides (Playfair 1918) [=*Leptocystis arcelloides] LT64
    |--A. arenaria LB90
    |--A. artocrea Deflandre 1928 NM05
    |--A. atava Collin 1914 [=A. (*Antarcella) atava] LT64
    |--A. aureola (Ehrenberg 1837) [=*Cyphidium aureolum, A. (*Cyphidium) aureola] LT64
    |--A. catinus LB90
    |--A. dentata Ehr. 1830 W86
    |--A. discoides Ehr. 1843 W86 [incl. A. discoides var. scutelliformis TG03]
    |--A. excavata Cunningham 1919 TG03
    |--A. hemisphaerica C-SCL16
    |--A. peristicta Ehrenberg 1854 [=A. (*Heterocosmia) peristicta] LT64
    |--A. mitrata D01
    `--A. polypora [incl. A. polypora var. curvata Wailes 1913] TG03

Arcella Ehrenberg 1832 [=Arcellina Carter 1856, Euarcella Deflandre 1928, Sticholepis Ehrenberg 1872; incl. Antarcella Deflandre 1928, Cyphidium Ehrenberg 1837, Heterocosmia Ehrenberg 1872, Leptocystis Playfair 1918 non Jaeckel 1899; Arcellida, Arcellidae, Arcellinae, Arcellinea, Arcellineae, Arcellinina] LT64

*Type species of generic name indicated

REFERENCES

[C-SCL16] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2016. 187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution. Molecular Phylogenetics and Evolution 99: 275–296.

[D01] Daday, E. 1901. Édesvizi mikroszkópi állatok [Mikroskopische Süsswasserthiere]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 375–470. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[NM05] Nikolaev, S. I., E. A. D. Mitchell, N. B. Petrov, C. Berney, J. Fahrni & J. Pawlowski. 2005. The testate lobose amoebae (order Arcellinida Kent, 1880) finally find theit home within Amoebozoa. Protist 156: 191–202.

[TG03] Todorov, M., & V. Golemansky. 2003. Morphology, biometry and ecology of Arcella excavata Cunningham, 1919 (Rhizopoda: Arcellinida). Acta Protozoologica 42: 105–111.

[W86] Whitelegge, T. 1886. List of the freshwater Rhizopoda of N. S. Wales. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 497–504.

Tubulinea

Amoeba proteus, copyright Wim van Egmond.

Belongs within: Amoebozoa.
Contains: Arcellinida.

The Tubulinea are a group of strictly amoeboid amoebozoans that are capable of producing tubular pseudopodia.

See also: Amoeba: much weirder than you think; Tubulinea: the paragons of amoeboids.

Characters (from Smirnov et al. 2005): Naked or testate amoebae producing tubular, subcylindrical pseudopodia or capable of altering locomotive form from flattened, expanded one to subcylindrical one. Monoaxial flow of cytoplasm in every pseudopodium or in entire cell. No cytoplasmic microtubule-organising centres; no flagellate stage in life cycle.

<==Tubulinea [Hartmannellina, Lobosea, Tubulina]
    |--Leptomyxida [Leptomyxidae, Leptomyxoidea] SN05
    |    |  i. s.: Gephyramoeba [Gephyramoebidae] AS12
    |    |           `--G. delicatula LB90
    |    |--Leptomyxa reticulata SN05
    |    `--+--Rhizamoeba saxonica SN05
    |       `--Flabellulidae [Flabellulina] C-SCO04
    |            |--Flabellula SN05
    |            `--Paraflabellula SN05
    |                 |--P. hoguae SN05
    |                 `--P. reniformis SN05
    `--+--Arcellinida SN05
       `--Euamoebida AS12
            |--Copromyxidae [Copromyxida] C-SCO04
            |    |--Copromyxa C-SCO04
            |    `--Copromyxella C-SCO04
            `--Tubulinida [Amoebidae, Amoeboidea, Chaidae, Chaosidae, Hartmannellidae, Hartmannellinae] SN05
                 |  i. s.: Cashia SN05
                 |         Deuteramoeba SN05
                 |         Hydramoeba SN05
                 |         Parachaos SN05
                 |         Polychaos SN05
                 |         Trichamoeba villosa SN05, LB90
                 |--Saccamoeba SN05
                 |    |--S. limax SN05
                 |    `--S. stagnicola LB90
                 `--+--+--Glaeseria mira SN05
                    |  `--Hartmannella cantabrigiensis SN05
                    `--+--Chaos Linné 1767 SN05, LT64 [Chaosina]
                       |    |--C. carolinense SN05
                       |    |--C. diffluens LT64
                       |    `--C. nobile SN05
                       `--Amoeba Ehrenberg 1830 SN05, LT64 [=Amiba Bory de St. Vincent 1822 LT64]
                            |--A. dubia LG91
                            |--A. leningradensis SN05
                            |--A. limax SX97
                            |--A. proteus SN05
                            `--A. vespertilis SX97

*Type species of generic name indicated

REFERENCES

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[C-SCO04] Cavalier-Smith, T., E. E.-Y. Chao & B. Oates. 2004. Molecular phylogeny of Amoebozoa and the evolutionary significance of the unikont Phalansterium. European Journal of Protistology 40: 21–48.

[LG91] Li, W.-H., & D. Graur. 1991. Fundamentals of Molecular Evolution. Sinauer: Sunderland (MA).

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[SN05] Smirnov, A., E. Nassonova, C. Berney, J. Fahrni, I. Bolivar & J. Pawlowski. 2005. Molecular phylogeny and classification of the lobose amoebae. Protist 156: 129–142.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl): 60–68.

Amoebozoa

Cochliopodium sp., copyright Proyecto Agua.

Belongs within: Amorphea.
Contains: Variosea, Eumycetozoa, Tubulinea, Archamoebae, Centramoebia, Thecamoebida, Vannellida, Paramoebidae.

The Amoebozoa is a clade of primarily unicellular eukaryotes in which most members have an amoeboid stage in their life cycle; some are exclusively amoeboid. Members of the clade Eudiscosea produce flattened pseudopodia with a polyaxial cytoplasmic flow or lacking a pronounced axis, and so far as known lack a ciliated stage (Adl et al. 2019).

See also: Amoebozoan classification: putting the formless in formation; Amoebozoan oddments.

Characters (from Adl et al. 2012): Cells “naked” or testate; tubular mitochondrial cristae, often branched (ramicristate), secondarily lost in some; uninucleate, binucleate or multinucleate; cysts common, morphologically variable; sexual or asexual; many taxa exhibit either sporocarpic (single amoeboid cell differentiates into a usually stalked, subaerial structure that supports one to many propagules termed spores) or sorocarpic (amoebae aggregate into a multicellular mass that develops into a multicellular fruiting body) fruiting; or myxogastroid ciliated stages; when amoeboid, locomotion with noneruptive morphologically variable pseudopodia; ancestrally bikont with many taxa exhibiting reduction of the bikinetid.

Amoebozoa (see below for synonymy)
    |--Evosea AB19
    |    |--Semiconosia (see below for synonymy) C-SCL16
    |    |    |--Variosea TA16
    |    |    `--Eumycetozoa TA16
    |    `--+--+--Tubulinea TA16
    |       |  `--Archamoebae TA16
    |       `--+--+--Pessonella marginata TA16, SX97
    |          |  `--Cutosa [Cutosea] C-SCL16
    |          |       |--Armaparvus AB19
    |          |       |--Squamamoeba Kudryavtsev & Pawlowski 2013 [Squamamoebidae] C-SCL16
    |          |       |    `--S. japonica C-SCL16
    |          |       `--Sapocribrum Lahr et al. 2015 TA16, C-SCL16 [Sapocribridae]
    |          |            `--S. chincoteaguense C-SCL16
    |          `--Himatismenida [Cochliopodidae, Cochliopodiidae, Cochliopodiinae, Pseudonebelinae, Tectiferina] TA16
    |               |  i. s.: Chlamydamoeba Collin 1912 LT64
    |               |           `--*C. tentaculifera Collin 1912 LT64
    |               |         Coenopodium C-SCL16
    |               |--Ovalopodium desertum TA16
    |               `--+--Parvamoeba [Parvamoebidae, Parvamoebina] TA16
    |                  |    |--P. monoura TA16
    |                  |    `--P. rugata C-SCL16
    |                  `--Cochliopodium Hertwig & Lesser 1874 TA16, KB05 [=Kochliopodium (l. c.) LT64]
    |                       |--C. bilimbosum (Auerbach 1856) (see below for synonymy) LT64
    |                       |--C. gulosum C-SCO04
    |                       |--C. longispinum SX97
    |                       |--C. minus Page 1976 KB05
    |                       |--C. minutoidum C-SCL15
    |                       |--C. minutum West 1901 KB05
    |                       `--C. spiniferum Kudryatsev 2004 KB05
    `--Eudiscosea [Longamoebia] C-SCL16
         |--Centramoebia AS12
         `--Discosea AB19
              |  i. s.: Mycamoeba Blandenier et al. 2017 AB19
              |           `--M. gemmipara AB19
              |--Thecamoebida AS12
              `--Flabellinea [Flabellina, Flabellinia, Glycostylida, Vanelloidea] TA16
                   |--Vannellida C-SCL16
                   `--Dactylopodida [Conopodina, Dactylopodina, Paramoeboidea] TA16
                        |  i. s.: Boveella AS12
                        |         Dactylosphaerium AS12
                        |         Oscillodignum AS12
                        |         Podostoma filigerum AS12, G89
                        |         Strioluatus AS12
                        |         Subulamoeba saphirina AS12, SX97
                        |         Trienamoeba AS12
                        |--Paramoebidae C-SCL16
                        `--+--Vexillifera [Vexilliferidae] C-SCL16
                           |    |--V. armata SN05
                           |    |--V. bacillipedes SN05
                           |    `--V. minutissima SN05
                           `--Dermamoebida C-SCL16
                                |--Dermamoebidae C-SCL16
                                |    |--Dermamoeba algensis C-SCL16
                                |    `--Paradermamoeba C-SCL16
                                `--Mayorella [Mayorellida, Mayorellidae, Mayorellina] TA16
                                     |--M. gemmifera C-SCL16
                                     |--M. inquisita SX97
                                     |--M. penardi MS98
                                     |--M. riparia LB90
                                     `--M. spatula SX97

Amoebozoa incertae sedis:
  Malamoeba locustae SN05, R91
  Malpighamoeba SN05
  Pansporella AS12 [Pansporellidae LT64, Sporamoebidae]
  Gibbodiscus AS12
  Protamoeba vorax Gruber 1884 G84
  Belonocystis AB19
  Gymnophryidae [Araudia, Biomyxida, Gymnophrea, Reticulosida] C-SC03
    |--Gymnophrys cometa NB04
    |--Borkovia desaedeleeri C-SC03, M00
    `--Biomyxa Leidy 1875 AB19, LT64 [Biomyxidae]
         `--*B. vagans Leidy 1875 LT64 [incl. Amoeba porrecta G84]
  Oscillosignum AB19
  Schoutedamoeba minuta Van Vichelin et al. 2016 AB19

Amoebozoa [Acanthopodina, Addiffluentia, Amastigogeninia, Amebea, Amoebaea, Amoebeae, Amoebiae, Amoebida, Amoeboea, Chaetoproteida, Chaetoproteidae, Chaoinea, Conosa, Conosea, Diffluentia, Glycopoda, Gymnamoebaea, Gymnamoebia, Gymnamoebida, Gymnamoebina, Lobosa, Lobosia, Monamoebidae, Monamoebina, Monostega, Monostegia, Pelomyxacea, Pharopodida, Protamoebae, Proteina, Protomyxidae, Protosteliales, Protosteliomycetes, Testacealobosea, Testamoebida, Thecamoebina, Thecina]

Cochliopodium bilimbosum (Auerbach 1856) [=Amoeba bilimbosa; incl. *C. pellucidum Hertwig & Lesser 1874] LT64

Semiconosia [Ceratiomyxacea, Ceratiomyxaceae, Ceratiomyxales, Ceratiomyxioidea, Ectosporeae, Eumyxa, Exosporae, Exosporales, Exosporea, Exosporeen, Exosporei, Exosporinei, Parastelida] C-SCL16

*Type species of generic name indicated

REFERENCES

[AB19] Adl, S. M., D. Bass, C. E. Lane, J. Lukeš, C. L. Schoch, A. Smirnov, S. Agatha, C. Berney, M. W. Brown, F. Burki, P. Cárdenas, I. Čepička, L. Chistyakova, J. del Campo, M. Dunthorn, B. Edvardsen, Y. Eglit, L. Guillou, V. Hampl, A. A. Heiss, M. Hoppenrath, T. Y. James, A. Karnkowska, S. Karpov, E. Kim, M. Kolisko, A. Kudryavtsev, D. J. G. Lahr, E. Lara, L. Le Gall, D. H. Lynn, D. G. Mann, R. Massana, E. A. D. Mitchell, C. Morrow, J. S. Park, J. W. Pawlowski, M. J. Powell, D. J. Richter, S. Rueckert, L. Shadwick, S. Shimano, F. W. Spiegel, G. Torruella, N. Youssef, V. Zlatogursky & Q. Zhang. 2019. Revisions to the classification, nomenclature, and diversity of eukaryotes. Journal of Eukaryotic Microbiology 66: 4–119.

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[C-SC03] Cavalier-Smith, T., & E. E.-Y. Chao. 2003. Phylogeny and classification of phylum Cercozoa. Protist 154: 341–358.

[C-SCL15] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2015. Multiple origins of Heliozoa from flagellate ancestors: new cryptist subphylum Corbihelia, superclass Corbistoma, and monophyly of Haptista, Cryptista, Hacrobia and Chromista. Molecular Phylogenetics and Evolution 93: 331–362.

[C-SCL16] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2016. 187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution. Molecular Phylogenetics and Evolution 99: 275–296.

[C-SCO04] Cavalier-Smith, T., E. E.-Y. Chao & B. Oates. 2004. Molecular phylogeny of Amoebozoa and the evolutionary significance of the unikont Phalansterium. European Journal of Protistology 40: 21–48.

[G84] Gruber, A. 1884. Die Protozoen des Hafens von Genua. Verhandlungen der Kaiserlichen Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher [Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum] 46 (4): 473–539, pls 7–11.

[G89] Gruber, A. 1889. Ueber einige Rhizopoden aus dem Genueser Hafen. Berichte der Naturforschenden Gesellschaft zu Freiburg I. B. 4: 33–44, pl. 31.

[KB05] Kudryavtsev, A., D. Bernhard, M. Schlegel, E. E-Y. Chao & T. Cavalier-Smith. 2005. 18S ribosomal RNA gene sequences of Cochliopodium (Himatismenida) and the phylogeny of Amoebozoa. Protist 156: 215–224.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[MS98] Margulis, L., & K. V. Schwartz. 1998. Five Kingdoms: An Illustrated Guide to the Phyla of Life on Earth 3^rd ed. W. H. Freeman and Company: New York.

[M00] Mikrjukov, K. A. 2000. Taxonomy and phylogeny of Heliozoa: should this taxon exist in modern classification of Protista? Zoologicheskii Zhurnal 79 (8): 883–897 (translated: Entomological Review 80 (Suppl. 1): S35–S50).

[NB04] Nikolaev, S. I., C. Berney, J. F. Fahrni, I. Bolivar, S. Polet, A. P. Mylnikov, V. V. Aleshin, N. B. Petrov & J. Pawlowski. 2004. The twilight of Heliozoa and rise of Rhizaria, an emerging supergroup of amoeboid eukaryotes. Proceedings of the National Academy of Sciences of the USA 101 (21): 8066–8071.

[R91] Rentz, D. C. F. 1991. Orthoptera (grasshoppers, locusts, katydids, crickets). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 369–393. Melbourne University Press: Carlton (Victoria).

[SN05] Smirnov, A., E. Nassonova, C. Berney, J. Fahrni, I. Bolivar & J. Pawlowski. 2005. Molecular phylogeny and classification of the lobose amoebae. Protist 156: 129–142.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl.): 60–68.

[TA16] Tekle, Y. I., O. R. Anderson, L. A. Katz, X. X. Maurer-Alcalá, M. A. Cerón Romero & R. Molestina. 2016. Phylogenomics of 'Discosea': a new molecular phylogenetic perspective on Amoebozoa with flat body forms. Molecular Phylogenetics and Evolution 99: 144–154.

Last updated: 4 June 2022.

Difflugia

Pair of Difflugia lucida, copyright Angie Opitz.

Belongs within: Arcellinida.

Difflugia is a genus of testate amoebozoans found in freshwater lakes and ponds, with a test composed of a combination of pseudochitin and agglutinated foreign material. They are known in the fossil record from the Eocene (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Test globular, elongate, pyriform, or acuminate, typically circular in section (length 15–450 µm, average approx. 100 µm); wall with pseudochitinous base and variable amounts of agglutinated material; aperture rounded , may be somewhat produced; protoplasm occupying most of test, attached internally by threads of ectoplasm; nucleus large, single; one or more contractile vacuoles; pseudopodia few, simple or bifid, of colourless ectoplasm, more rarely including some granular endoplasm.

<==Difflugia Leclerc in Lamarck 1816 (see below for synonymy) LT64
    |--*D. protoeiformis Lamarck 1816 [=D. (*Corticella) proteiformis, D. (*Eudifflugia) proteiformis] LT64
    |--D. acuminata Ehrenberg 1838 N03, LT64
    |    |--D. a. acuminata N03
    |    `--D. a. curvata N03
    |--D. constricta D01
    |--D. elegans N03
    |    |--D. e. elegans N03
    |    `--D. e. bicornis N03
    |--D. globularis Wallich 1864 [=D. (Reticella) globularis, *Odontodictya globularis] LT64
    |--D. globulosa Dujardin 1837 W86
    |--D. globulus (Ehrenberg 1848) [=Arcella globulus] LT64
    |--D. lanceolata SX97
    |--D. longum SX97
    |--D. lucida G86
    |--D. maroccana (Gauthier-Lièvre & Thomas 1958) [=*Pentagonia maroccana] LT64
    |--D. oblonga Ehrenberg 1838 LT64
    |--D. oviformis LT64
    |--D. penardi LNB03
    |--D. pristis LNB03
    |--D. pulex SX97
    |--D. pyriformis Perty 1848 LT64
    |--D. regularis (Jung 1942) [=*Schwabia regularis] LT64
    |--D. submarina Chardez & Thomas 1980 G86
    |--D. subterranea Golemansky 1970 [incl. D. subterranea var. inflata Decloître 1975] G86
    `--D. urceolata Carter 1864 W86

Difflugia Leclerc in Lamarck 1816 [=Corticella Ehrenberg 1872, Eudifflugia Diesing 1848; incl. Acipyxis Jung 1942 (n. n.), Globonota Jung 1942 (n. n.), Odontodictya Ehrenberg 1872, Pentagonia Gauthier-Lièvre & Thomas 1958 non Cozzens 1846, Planodifflugia Jung 1942 (n. n.), Pycnochila Jung 1942 (n. n.) non Horn 1905, Schwabia Jung 1942] LT64

*Type species of generic name indicated

REFERENCES

[D01] Daday, E. 1901. Édesvizi mikroszkópi állatok [Mikroskopische Süsswasserthiere]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 375–470. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[G86] Golemansky, V. G. 1986. Rhizopoda: Testacea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 5–16. E. J. Brill/Dr W. Backhuys: Leiden.

[LNB03] Ledeganck, P., I. Nijs & L. Beyens. 2003. Plant functional group diversity promotes soil protist diversity. Protist 154 (2): 239–249.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[N03] Nicholls, K. H. 2003. Form variation in Campascus minutus and a description of Campascus simcoei sp. n. (Testaceafilosea, Psammonobiotidae). European Journal of Protistology 39 (1): 103–112.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl.): 60–68.

[W86] Whitelegge, T. 1886. List of the freshwater Rhizopoda of N. S. Wales. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 497–504.

Last updated: 29 November 2017.

Arcellinida

Test of Cyclopyxis eurystoma, copyright Ferry Siemensma.

Belongs within: Tubulinea.
Contains: Difflugia, Hyalospheniidae, Centropyxiidae, Arcella, Microcoryciidae, Plagiopyxidae.

The Arcellinida, testate amoebae, are a group of amoebozoans which contain themselves inside an organic or mineral extracellular test of either self-secreted elements (calcareous, siliceous, or chitinoid) or recycled mineral particles bound together, with a single main opening. Pseudopodia are conical and pointed in species of Phryganellina, and digitate and granular in Difflugina and Arcellina (Adl et al. 2012).

<==Arcellinida [Lampramoebae, Testacealobosia, Testacida, Testalobosina, Thecamoebaea]
    |  i. s.: Paleoarcella athanata SM03
    |         Geamphorella Bonnet 1959 AS12, LT64
    |           `--*G. lucida Bonnet 1959 [=Tracheleuglypha lucida] LT64
    |         Pyxidicula Ehrenberg 1838 C-SCL16, FT93
    |           `--*P. operculata (Agardh 1827) [=Frustrulia operculata, Arcella operculata] LT64
    |--Phryganellina AS12
    |    |--Wailesella Deflandre 1928 AS12, LT64
    |    |    `--*W. eboracensis (Wailes in Wailes & Penard 1911) [=Cryptodifflugia eboracensis] LT64
    |    |--Phryganella Penard 1902 G86 (see below for synonymy)
    |    |    |--*P. nidulus Penard 1902 LT64
    |    |    |--P. acropodia LB90
    |    |    |--P. hemisphaerica LNB03
    |    |    |--P. marinus Chardez 1971 G86
    |    |    `--P. sylvicola (Bonnet & Thomas 1955) [=*Geopyxella sylvicola] LT64
    |    `--Cryptodifflugia Penard 1890 AS12, G86 [incl. Geococcus Francé 1913 non Green 1902 LT64; Cryptodifflugiidae]
    |         |--*C. oviformis Penard 1890 LT64
    |         |--C. brevicolla Golemansky 1979 G86
    |         |--C. lanceolata Golemansky 1970 G86
    |         |--C. operculata C-SCL16
    |         |--C. paludosa Golemansky 1981 G86
    |         `--C. vulgaris (Francé 1913) [=*Geococcus vulgaris] LT64
    `--Difflugina [Arcellacea, Eulobosa] AS12
         |  i. s.: Difflugiidae (see below for synonymy) LT64
         |           |--Difflugia AS12
         |           |--Cucurbitella Penard 1902 (see below for synonymy) LT64
         |           |    `--*C. mespiliformis Penard 1902 LT64
         |           |--Maghrebia Gauthier-Lièvre & Thomas 1958 LT64
         |           |    `--*M. spatulata Gauthier-Lièvre & Thomas 1958 LT64
         |           |--Protocucurbitella Gauthier-Lièvre & Thomas 1960 LT64
         |           |    `--*P. coroniformis Gauthier-Lièvre & Thomas 1960 LT64
         |           |--Pseudopontigulasia Van Oye 1956 LT64
         |           |    `--*P. gessneri Van Oye 1956 LT64
         |           |--Sexangularia Averintsev 1906 LT64
         |           |    `--*S. parvula Averintsev 1906 LT64
         |           |--Loboforamina Jung 1942 (see below for synonymy) LT64
         |           |    |--L. playfairi Loeblich & Tappan 1964 (see below for synonymy) LT64
         |           |    `--‘*Corona’ cornula Vejdovský 1881 (see below for synonymy) LT64
         |           |--Pontigulasia Rhumbler 1895 LT64
         |           |    |--P. rhumbleri Hopkinson in Cash & Hopkinson 1909 (see below for synonymy) LT64
         |           |    `--P. compressa (Carter 1864) LT64 (see below for synonymy)
         |           `--Prantlitina Vašíček & Růžička 1957 [incl. Prantlitinopsis Vašíček & Růžička 1957] LT64
         |                |--*P. prantli Vašíček & Růžička 1957 LT64
         |                `--P. sturi Vašíček & Růžička 1957 [=P. (*Prantlitinopsis) sturi] LT64
         |         Paraquadrula Deflandre 1932 LT64
         |           `--*P. irregularis (Archer 1877) [=Quadrula irregularis] LT64
         |         Distomatopyxis [Distomatopyxidae] AS12
         |           `--D. couillardi LB90
         |         Bullinaria indica C-SCL16
         |--Heleopera Leidy 1879 NM05, LT64 [Heleoperidae, Heleoperinae]
         |    |--H. sphagni (Leidy 1874) NM05 [=Difflugia (Nebela) sphagni LT64; incl. *H. picta Leidy 1879 LT64]
         |    |--H. petricola LB90
         |    `--H. rosea LB90
         `--+--Hyalospheniidae NM05
            `--+--+--Centropyxiidae NM05
               |  `--Arcellina AS12
               |       |  i. s.: Spumochlamys AS12
               |       |--Arcella NM05
               |       `--Microcoryciidae AS12
               `--+--Plagiopyxidae NM05
                  `--Trigonopyxidae LT64
                       |--Trigonopyxis Penard 1912 NM05, FT93 (see below for synonymy)
                       |    `--*T. arcula (Leidy 1879) [=Difflugia arcula, *Cystidina arcula] LT64
                       |--Silicoplacentina Köváry 1956 LT64
                       |    `--*S. hungarica Köváry 1956 LT64
                       `--Cyclopyxis Deflandre 1929 (see below for synonymy) LT64
                            |--*C. arcelloides (Penard 1902) [=Centropyxis arcelloides] LT64
                            |--C. ambigua LB90
                            |--C. arenata (Cushman 1930) [=Pseudarcella arenata, *Leptodermella arenata] LT64
                            |--C. cylindrica LT64
                            |--C. eurystoma [incl. C. eurystoma var. parvula] LB90
                            `--C. puteus LB90

‘*Corona’ cornula Vejdovský 1881 [incl. Difflugia proteiformis globularis var. corona Wallich 1864] LT64

Cucurbitella Penard 1902 [incl. Cingodifflugia Jung 1942 (n. n.), Eustoma Jung 1942 (n. n.) nec Piette 1855 nec Beneden 1871 nec Girard 1893] LT64

Cyclopyxis Deflandre 1929 [incl. Cylindropyxis Kufferath 1932 (n. n.), Leptodermella Rhumbler 1935] LT64

Difflugiidae [Cingodifflugiidae, Difflugiida, Difflugiina, Difflugiinae, Diffluginae, Planodifflugiidae, Pontigulasiidae] LT64

Loboforamina Jung 1942 [incl. Corona Vejdovský 1881 nec Albers 1850 nec Récluz 1850, Pseudocucurbitella Gauthier-Lièvre & Thomas 1960 (n. n.)] LT64

Loboforamina playfairi Loeblich & Tappan 1964 [=Difflugia lobostoma var. globulus Playfair 1918 non Arcella globulus Enrenberg 1848] LT64

Phryganella Penard 1902 G86 [=Phryanella (l. c.) LT64; incl. Geopyxella Bonnet & Thomas 1955 LT64; Phryganellidae LT64]

Pontigulasia compressa (Carter 1864) LT64 [=Difflugia compressa LT64, D. pyriformis var. compressa W86]

Pontigulasia rhumbleri Hopkinson in Cash & Hopkinson 1909 [=*P. compressa Rhumbler 1905 non Difflugia compressa Carter 1864] LT64

Trigonopyxis Penard 1912 NM05, FT93 [=Cystidina Volz 1929 LT64, Trigonopsis (l. c.) non Perty 1833 LT64]

*Type species of generic name indicated

REFERENCES

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[C-SCL16] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2016. 187-gene phylogeny of protozoan phylum Amoebozoa reveals a new class (Cutosea) of deep-branching, ultrastructurally unique, enveloped marine Lobosa and clarifies amoeba evolution. Molecular Phylogenetics and Evolution 99: 275–296.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[G86] Golemansky, V. G. 1986. Rhizopoda: Testacea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 5–16. E. J. Brill/Dr W. Backhuys: Leiden.

[LNB03] Ledeganck, P., I. Nijs & L. Beyens. 2003. Plant functional group diversity promotes soil protist diversity. Protist 154 (2): 239–249.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[NM05] Nikolaev, S. I., E. A. D. Mitchell, N. B. Petrov, C. Berney, J. Fahrni & J. Pawlowski. 2005. The testate lobose amoebae (order Arcellinida Kent, 1880) finally find theit home within Amoebozoa. Protist 156: 191–202.

[SM03] Schlegel, M., & R. Meisterfeld. 2003. The species problem in protozoa revisited. European Journal of Protistology 39 (4): 349–355.

[W86] Whitelegge, T. 1886. List of the freshwater Rhizopoda of N. S. Wales. Part I. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 497–504.

Last updated: 29 November 2017.

Centropyxiidae

Individual of Centropyxis among filamentous algae, copyright John C. Walsh.

Belongs within: Arcellinida.

The Centropyxiidae are a group of testate amoebozoans with a chitinous test into which sand or other foreign matter has been incorporated (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Test chitinous, enclosing some foreign material or with sandy layer, generally with dorsoventral symmetry; aperture ventral, with recurved margin, eccentric; single nucleus.

<==Centropyxiidae [Conchulina]
    |--Centropyxiella Valkanov 1970 G86
    |    |--C. arenaria Valkanov 1970 G86
    |    |--C. elegans Valkanov 1970 G86
    |    |--C. gibbula Valkanov 1970 G86
    |    |--C. gibbulina Chardez & Thomas 1980 G86
    |    |--C. golemanskyi Chardez 1977 G86
    |    |--C. lucida Golemansky 1971 G86
    |    |--C. oopyxiformis Chardez 1977 G86
    |    `--C. platystoma Golemansky 1981 G86
    `--Centropyxis Stein 1859 (see below for synonymy) LT64
         |--*C. aculeata (Ehrenberg 1832) [=Arcella aculeata, *Echinopyxis aculeata] LT64
         |--C. aerophila [incl. C. aerophila var. sphagnicola] LNB03
         |--C. cassis LNB03
         |--C. constricta [incl. C. constricta var. minima] LB90
         |--C. ecornis SX97
         |--C. gasparella LNB03
         |--C. globulosa LB90
         |--C. hemisphaerica SX97
         |--C. hirsuta SR06
         |--C. kahli LNB03
         |--C. laevigata Penard 1902 NM05
         |--C. marsupiformis N03
         |--C. orbicularis SX97
         |--C. plagiostoma LB90
         |--C. platystoma LNB03
         |--C. pleurostomelloides (Millett 1899) (see below for synonymy) LT64
         |--C. pontigulasiformis LNB03
         `--C. sylvatica [incl. C. sylvatica var. minor] LB90

Centropyxis Stein 1859 [=Echinopyxis Claparède & Lachmann 1859; incl. Armillettum Rhumbler 1913, Millettella Rhumbler 1904] LT64

Centropyxis pleurostomelloides (Millett 1899) [=Reophax pleurostomelloides, *Armillettum pleurostomelloides, *Millettella pleurostomoides] LT64

*Type species of generic name indicated

REFERENCES

[G86] Golemansky, V. G. 1986. Rhizopoda: Testacea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 5–16. E. J. Brill/Dr. W. Backhuys: Leiden.

[LNB03] Ledeganck, P., I. Nijs & L. Beyens. 2003. Plant functional group diversity promotes soil protist diversity. Protist 154 (2): 239–249.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[N03] Nicholls, K. H. 2003. Form variation in Campascus minutus and a description of Campascus simcoei sp. n. (Testaceafilosea, Psammonobiotidae). European Journal of Protistology 39 (1): 103–112.

[NM05] Nikolaev, S. I., E. A. D. Mitchell, N. B. Petrov, C. Berney, J. Fahrni & J. Pawlowski. 2005. The testate lobose amoebae (order Arcellinida Kent, 1880) finally find theit home within Amoebozoa. Protist 156: 191–202.

[SR06] Schmidt, A. R., E. Ragazzi, O. Coppellotti & G. Roghi. 2006. A microworld in Triassic amber. Nature 444: 835.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl): 60–68.

Hyalospheniidae

Live Nebela tincta scavenging scales from an Euglypha test, copyright Ferry Siemensma.

Belongs within: Arcellinida.

The Hyalospheniidae are a group of testate amoebozoans in which the test is composed of pseudochitin, and which have a distinctive pseudopodium conformation in which retreating pseudopodia leave a long thin pellicle that is slowly retracted (Loeblich & Tappan 1964).

Characters (from Loeblich & Tappan 1964): Test pseudochitinous, with siliceous plates or scales, rounded or angular, may have added foreign matter; aperture elongate or rounded; single nucleus; pseudopodia largely composed of ectoplasm, may have slight penetration of endoplasm near base (endolobopodia), rounded terminally during advance and retraction, but when retracting temporarily leave behind fine, structureless, hyaline pellicle which tapers to point, resembles ectoplasm in appearance, later pulls away from substratum to be resorbed.

<==Hyalospheniidae (see below for synonymy)
    |--Pomoriella Golemansky 1970 G86
    |    `--P. valkanovi Golemansky 1970 G86
    |--Awerintzewia Schouteden 1906 (see below for synonymy) LT64
    |    `--*A. cyclostoma (Penard 1902) (see below for synonymy) LT64
    |--Certesella Loeblich & Tappan 1961 [incl. Penardiella Jung 1942 (n. n.) non Kahl 1930] LT64
    |    `--*C. martiali (Certes 1891) [=Nebela martiali] LT64
    |--Jungia Loeblich & Tappan 1961 LT64
    |    `--*J. sundanensis Van Oye 1949 LT64
    |--Leptochlamys West 1901 LT64
    |    `--*L. ampullacea West 1901 LT64
    |--Pseudawerintzewia Bonnet 1959 LT64
    |    `--*P. calcicola Bonnet 1959 LT64
    |--Pseudonebela Gauthier-Lièvre 1954 LT64
    |    `--*P. africana Gauthier-Lièvre 1954 LT64
    |--Porosia Jung 1942 [incl. Alocodera Jung 1942] LT64
    |    |--*P. bigibbosa (Penard 1890) [=Nebela bigibbosa] LT64
    |    `--‘Hyalosphenia’ cockayni Penard 1910 [=*Alocodera (Nebela) cockayni] LT64
    |--Quadrulella Cockerell 1909 [incl. Quadrula Schulze 1875 non Rafinesque 1820] LT64
    |    |--*Q. symmetrica (Wallich 1863) [=Difflugia proteiformis var. symmetrica, *Quadrula symmetrica] LT64
    |    |--Q. globulosa SX97
    |    `--Q. marina (Bailey 1856) [=Difflugia marina] LT64
    |--Lesquereusia Schlumberger 1845 [=Lecquereusia] LT64
    |    |--*L. jurassica Schlumberger 1845 LT64
    |    |--L. modesta Rhumbler 1895 LT64
    |    `--L. spiralis (Ehrenberg 1840) [=Difflugia spiralis] LT64
    |--Hyalosphenia Stein 1857 G86 [incl. Catharia Leidy 1874 non Lederer 1863 LT64]
    |    |--H. ligata (Tatem 1870) (see below for synonymy) LT64
    |    |--H. minuta LB90
    |    |--H. papilio Leidy 1875 NM05
    |    |--H. savoiei Chardez 1978 G86
    |    `--H. subflava LB90
    |--Apodera Loeblich & Tappan 1961 LT64
    |    `--*A. vas (Certes 1891) LT64 (see below for synonymy)
    `--Nebela Leidy 1875 (see below for synonymy) LT64
         |--N. collaris (Ehrenberg 1848) (see below for synonymy) LT64
         |--N. ansata LT64
         |--N. bohemica GM03
         |--‘*Pterygia’ carinulata Jung 1942 LT64
         |--N. dentistoma LB90
         |--N. lageniformsi LB90
         |--N. longicollis Penard 1890 [=*Cyphoderiopsis longicollis] LT64
         |--N. militaris NM05
         |--N. parvula LB90
         |--N. tabulata LB90
         `--N. tincta [incl. N. tincta var. major Deflandre 1936] NM05

*Apodera vas (Certes 1891) LT64 [=Nebela vas LT64, Difflugia pyriformis var. vas FEF04, Pontigulasia vas FEF04; incl. P. bryophila FEF04, P. compressa var. flexa FEF04, Nebela vas var. longicollis Grospietsch 1971 non N. longicollis Penard 1890 FEF04, N. vas var. obliqua Grospietsch 1971 FEF04, Apodera (Nebela) vas f. recticollis Jung 1942 FEF04]

Awerintzewia Schouteden 1906 [=Averincevia Jírovec 1953, Averintzia Wailes in Cash, Wailes & Hopkinson 1919, Averinzia Jung 1942, Awerintzevia Neave 1939, Awerintzia Calkins 1926; incl. Physochila Jung 1942 (n. n.)] LT64

*Awerintzewia cyclostoma (Penard 1902) [=Heleopera cyclostoma, *Averincevia cyclostoma, *Averintzia cyclostoma, *Averinzia cyclostoma, *Awerintzevia cyclostoma, *Awerintzia cyclostoma] LT64

Hyalosphenia ligata (Tatem 1870) [=Difflugia ligata, *Catharia ligata; incl. *Hyalosphenia cuneata Stein in Schulze 1875, H. lata Schulze 1875] LT64

Hyalospheniidae [Hyalosphenidae, Lecquereusiidae, Nebelida, Nebelidae, Nebelina, Nebelinae, Nebelini, Physochilini, Quadrulellinae, Quadrulidae, Quadrulina]

Nebela Leidy 1875 [=Nebella (l. c.); incl. Argynnia Jung 1942 (n. n.), Cyphoderiopsis Playfair 1918, Deflandria Jung 1942 (n. n.), Leidyella Jung 1942 (n. n.), Pterygia Jung 1942 nec Röding 1798 nec Laporte 1832, Reticella Ehrenberg 1872 non Gray 1870, Schaudinnia Jung 1942 (n. n.) non Schulze 1900, Umbonaria Jung 1942 (n. n.)] LT64

Nebela collaris (Ehrenberg 1848) [=Difflugia collaris, D. (*Reticella) collaris; incl. D. (*Nebela) numata Leidy 1875] LT64

*Type species of generic name indicated

REFERENCES

[FEF04] Finlay, B. J., G. F. Esteban & T. Fenchel. 2004. Protist diversity is different? Protist 155: 15–22.

[GM03] Gilbert, D., E. A. D. Mitchell, C. Amblard, G. Bourdier & A.-J. Francez. 2003. Population dynamics and food preferences of the testate amoeba Nebela tincta major-bohemica-collaris complex (Protozoa) in a Sphagnum peatland. Acta Protozoologica 42: 99–104.

[G86] Golemansky, V. G. 1986. Rhizopoda: Testacea. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 5–16. E. J. Brill / Dr. W. Backhuys: Leiden.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[NM05] Nikolaev, S. I., E. A. D. Mitchell, N. B. Petrov, C. Berney, J. Fahrni & J. Pawlowski. 2005. The testate lobose amoebae (order Arcellinida Kent, 1880) finally find theit home within Amoebozoa. Protist 156: 191–202.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl): 60–68.

Archamoebae

Stained trophozoite of Entamoeba histolytica, from Centers for Disease Control and Prevention. The solid purple circle is an ingested red blood cell.

Belongs within: Amoebozoa.

The Archamoebae are a group of amoebozoans in which the mitochondria have been converted into nonaerobic organelles (Adl et al. 2012).

See also: Archamoebae: the apogee (or nadir) or amoebozoan evolution.

<==Archamoebae [Archamoebea, Entamoebea, Pelobiontea, Pelomyxidae, Testamoebidae]
    |--Pelobiontida KB05
    |    |--Pelomyxa Greef 1874 KB05, AS12 [Caryoblastea, Peloflagellata]
    |    |    `--P. palustris KB05
    |    `--Entamoeba Casagrandi & Barbagallo 1895 KB05, LT64 [Entamoebida, Entamoebidae, Entamoebinae]
    |         |  i. s.: E. coli KB05
    |         |         E. dispar PHK96
    |         |         E. invadens BL01
    |         |         E. terrapinae SS02
    |         |--E. polecki HA01
    |         `--+--E. gingivalis HA01
    |            `--+--E. hartmanni HA01
    |               `--+--E. histolytica HA01
    |                  `--E. moshkovskii HA01
    `--Mastigamoebaea [Mastigamoebida, Mastigamoebidae, Rhizoflagellida, Rhizomastigidae] KB05
         |  i. s.: Tricholimax WDE06
         |         Mastigina hylae AS12, MM98
         |--+--Phreatamoeba Chàvez et al. 1986 KB05, WDE06 [Phreatamoebida]
         |  |    `--P. balamuthi KB05 [=Mastigamoeba balamuthi C-SCO04]
         |  `--Mastigella Frenzel 1897 KB05, WDE06
         |       |--M. commutans WDE06
         |       |--M. simplex LB90
         |       `--M. unica SX97
         `--+--Endolimacidae C-SCO04
            |    |--Endolimax Kuenen & Swellengrebel 1913 KB05, C-SCO04
            |    |    `--E. nana WDE06
            |    `--Endamoeba Leidy 1879 C-SCO04, LT64 [Endamoebida, Endamoebidae, Endamoebinae]
            |         `--E. blattae PHK96
            `--Mastigamoeba Schulze 1875 KB05, WDE06
                 |--*M. aspera Schulze 1875 WDE06
                 |--M. commutans OS03
                 |--M. invertens Klebs 1892 (n. d.) WDE06
                 |--M. psammobia WDE06
                 |--M. punctachlora C-SCO04
                 |--M. reptans LB90
                 |--M. schizophrenia C-SCO04
                 `--M. simplex KB05

*Type species of generic name indicated

REFERENCES

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[BL01] Biron, D., P. Libros, D. Sagi, D. Mirelman & E. Moses. 2001. ‘Midwives’ assist dividing amoebae. Nature 410: 430.

[C-SCO04] Cavalier-Smith, T., E. E.-Y. Chao & B. Oates. 2004. Molecular phylogeny of Amoebozoa and the evolutionary significance of the unikont Phalansterium. European Journal of Protistology 40: 21–48.

[HA01] Hackstein, J. H. P., A. Akhmanova, F. Voncken, A. van Hoek, T. van Alen, B. Boxma, S. Y. Moon-van der Staay, G. van der Staay, J. Leunissen, M. Huynen, J. Rosenberg & M. Veenhuis. 2001. Hydrogenosomes: convergent adaptations of mitochondria to anaerobic environments. Zoology 104: 290–302.

[KB05] Kudryavtsev, A., D. Bernhard, M. Schlegel, E. E-Y. Chao & T. Cavalier-Smith. 2005. 18S ribosomal RNA gene sequences of Cochliopodium (Himatismenida) and the phylogeny of Amoebozoa. Protist 156: 215–224.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[OS03] O’Kelly, C. J., J. D. Silberman, L. A. Amaral Zettler, T. A. Nerad & M. L. Sogin. 2003. Monopylocystis visvesvarai n. gen., n. sp. and Sawyeria marylandensis n. gen., n. sp.: two new amitochondrial heterolobosean amoebae from anoxic environments. Protist 154 (2): 281–290.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3^rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[SS02] Silberman, J. D., A. G. B. Simpson, J. Kulda, I. Cepicka, V. Hampl, P. J. Johnson & A. J. Roger. 2002. Retortamonad flagellates are closely related to diplomonads—implications for the history of mitochondrial function in eukaryote evolution. Molecular Biology and Evolution 19 (5): 777–786.

[SX97] Song B. & Xie P. 1997. Preliminary studies on the community structure of the planktonic protozoa from the outlet of Lake Dongting. Acta Hydrobiologica Sinica 21 (Suppl): 60–68.

[WDE06] Walker, G., J. B. Dacks & T. M. Embley. 2006. Ultrastructural description of Breviata anathema n. gen., n. sp., the organism previously studied as “Mastigamoeba invertens”. Journal of Eukaryotic Microbiology 53: 65–78.

Myxogastrea

Physarum polycephalum, copyright Stephanie Davidson.

Belongs within: Mycetozoa.
Contains: Trichiacea.

The Myxogastrea include the plasmodial slime moulds, amoeboid organisms that form a dispersed plasmodium in the nutritive phase of their life cycle, aggregating to form fruiting bodies in the reproductive phase.

Characters (from Cavalier-Smith et al. 2004): Life cycle with lamellipodial/filopodial amoeba, flagellate, and (usually) plasmodial phases; flagellate phase with two centrioles, each with two microtubular bands, biciliate or sometimes uniciliate; mitochondria typically with central rod-shaped nucleoid; with multilayered cysts or with spores (developing within plasmodia) borne on fruiting bodies; flagellate state with inner microtubular cone attached apically to microtubule nucleating centre at end of fibrillar root emanating from anterior centriolar base.

Myxogastrea (see below for synonymy)
    |  i. s.: Hyperamoeba flagellata KB05
    |         Cladostelium bisporum [=Echinostelium bisporum] C-SCO04
    |         Craterium FRS83
    |           |--C. leucocephalum FRS83
    |           `--C. minutum FRS83
    |         Protophysarum phloiogenum FRS83
    |--Protosporangiida AS12
    |    |--Protosporangiidae AS12
    |    |    |--Clastostelium AS12
    |    |    `--Protosporangium AS12
    |    `--Ceratiomyxa Schröter 1889 AS12
    |         `--C. fructiculosa LB90
    `--Myxogastria (see below for synonymy) AS12
         |  i. s.: Barbyella AS12
         |         Calomyxa metallica AS12, SK03
         |         Comatricha AS12
         |           |--C. acanthodes SK03
         |           |--C. ellae SK03
         |           `--C. nigra SK03
         |         Diderma AS12
         |           |--D. chondrioderma SK03
         |           |--D. effusum SK03
         |           `--D. hemisphaericum SK03
         |         Leocarpus AS12
         |         Lepidoderma tigrinum AS12, H04
         |         Macbrideola AS12
         |           |--M. cornea SK03
         |           `--M. scintillans SK03
         |         Oligonema AS12
         |         Willkommlangea AS12
         |--Trichiacea LT64
         `--Stemonitacea (see below for synonymy) LT64
              |--Echinosteliida C-SCO04
              |    |--Echinostelium SK03 [Echinosteliaceae, Echinosteliidae LT64]
              |    |    `--E. minutum SK03
              |    `--Clastodermidae C-SCO04
              |         |--Barbeyella C-SCO04
              |         `--Clastoderma C-SCO04
              |--Stemonitidae (see below for synonymy) LT64
              |    |  i. s.: Enerthenema papillatum SK03
              |    |         Raciborskia Berlese in Saccardo 1888 FT93
              |    |         Diachea leucopodia LB90
              |    |         Lamproderma AS12
              |    |--Colloderminae LT64
              |    |--Amaurochaetinae LT64
              |    `--Stemonitinae [Brefeldiei] LT64
              |         |--Stemonitis S03
              |         |    |--S. flavogenita KB05
              |         |    |--S. fusca SK03
              |         |    `--S. smithii SK03
              |         `--Brefeldia AS12
              `--Physarida (see below for synonymy) C-SCO04
                   |--Elaeomyxidae [Elaeomyxaceae] C-SCO04
                   |--+--Hyperamoeba dachnaya KB05
                   |  `--Didymium (see below for synonymy) C-SCO04
                   |       |--D. farinaceum H04
                   |       |--D. iridis FF02
                   |       `--D. nigripes KB05
                   `--Physaridae [Physarinae] LT64
                        |--Fuligo septica C-SCO04, BL04
                        |--Physarella LT64
                        |--Badhamia C-SCO04
                        |    |--B. panicea H04
                        |    `--B. rugulosa SK03
                        `--Physarum SK03
                             |--P. crateriforme SK03
                             |--P. didermoides H04
                             |--P. flavicomum PHK96
                             |--P. nutans SK03
                             |--P. plumbeum H04
                             `--P. polycephalum SK03

Didymium [Didymiacea, Didymiaceae, Didymeae, Didymidae, Didymieae, Didymiei, Didymiinae, Didymiidae, Spumariei] C-SCO04

Myxogastrea [Enteridiea, Eumycetozoina, Myxomycetales, Myxogasteres, Myxogastrales, Myxogastres, Myxomycetes, Myxomycota, Trichospermi]

Myxogastria [Atrichae, Columelliferae, Endosporea, Endosporeae, Endosporei, Endosporinei, Endotrichea, Euplasmodida] AS12

Physarida [Aethalini, Calcareae, Calcarinea, Calcarineae, Cienkowskiaceae, Lithodermeae, Physaraceae, Physaracei, Physarae, Physarales, Physarea, Physareae, Physarei, Physariineae, Physarina, Physaroidea, Physaroinae, Spumariaceae]

Stemonitacea [Amaurochaetides, Amaurosporae, Amaurosporales, Amaurosporeae, Amaurosporei, Stemonitei, Stemonitidides, Stemonitioidea] LT64

Stemonitidae [Amaurochaetacea, Amaurochaetaceae, Amaurochaeteae, Amaurochaetene, Amaurochaetinea, Amaurochaetineae, Amaurochaetoinae, Brefeldiaceae, Brefeldiidae, Collodermaceae, Collodermataceae, Enerthenemaceae, Enerthenemea, Lamprodermaceae, Lamprodermae, Lamprodermeae, Leptonemeae, Raciborskiaceae, Stemonitaceae, Stemonitacei, Stemonitae, Stemonitales, Stemonitea, Stemonitida, Stemonitina]

*Type species of generic name indicated

REFERENCES

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[BL04] Bullians, M. S., & R. A. B. Leschen. 2004. Noteucinetus new genus from New Zealand and Chile and notes on Eucinetus stewarti (Broun) (Coleoptera: Eucinetidae). New Zealand Entomologist 27: 29–38.

[C-SCO04] Cavalier-Smith, T., E. E.-Y. Chao & B. Oates. 2004. Molecular phylogeny of Amoebozoa and the evolutionary significance of the unikont Phalansterium. European Journal of Protistology 40: 21–48.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

[FF02] Foster, K. R., A. Fortunato, J. E. Strassmann & D. C. Queller. 2002. The costs and benefits of being a chimera. Proceedings of the Royal Society of London Series B—Biological Sciences 269: 2357–2362.

[FRS83] Frederick, L., I. L. Roth & R. Simons. 1983. SEM studies on operculum structure in the myxomycete Trichia crateriformis. Mycologia 75 (1): 46–52.

[H04] Haeckel, E. 1899–1904. Kunstformen der Natur. Bibliographisches Institut: Leipzig und Wien.

[KB05] Kudryavtsev, A., D. Bernhard, M. Schlegel, E. E-Y. Chao & T. Cavalier-Smith. 2005. 18S ribosomal RNA gene sequences of Cochliopodium (Himatismenida) and the phylogeny of Amoebozoa. Protist 156: 215–224.

[LT64] Loeblich, A. R., Jr & H. Tappan. 1964. Sarcodina: chiefly “thecamoebians” and Foraminiferida. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt C. Protista 2 vol. 1. The Geological Society of America and The University of Kansas Press.

[LB90] Lousier, J. D., & S. S. Bamforth. 1990. Soil protozoa. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 97–136. John Wiley & Sones: New York.

[PHK96] Prescott, L. M., J. P. Harley & D. A. Klein. 1996. Microbiology 3^rd ed. Wm. C. Brown Publishers: Dubuque (Iowa).

[SK03] Snell, K. L., & H. W. Keller. 2003. Vertical distribution and assemblages of corticolous myxomycetes on five tree species in the Great Smoky Mountains National Park. Mycologia 95 (4): 565–576.