Nebela

Nebela Temporal range: Pleistocene–Recent 2.58–0  Ma [1] PreꞒ Ꞓ O S D C P T J K Pg N ↓
Nebela flabellulum
Scientific classification
Domain:	Eukaryota
Clade:	Amorphea
Phylum:	Amoebozoa
Class:	Tubulinea
Order:	Arcellinida
Family:	Hyalospheniidae
Genus:	Nebela Leidy 1874
Type species
Nebela collaris (Ehrenberg 1848) Leidy 1879
Species
13 species
Synonyms
Difflugia (Reticella) Ehrenberg 1872 CyphoderiopsisPlayfair 1918 ArgynniaJung 1942 nom. nud. LeidyellaJung 1942 nom. nud. UmbonariaJung 1942 nom. nud. PterygiaJung 1942 nom. nud. SchaudinniaJung 1942 nom. nud. DeflandriaJung 1942 nom. nud.

Nebela is a diverse genus of testate amoebae of cosmopolitan distribution, belonging to the family Hyalospheniidae. ^[1] They are "prey agglutinated" or "kleptosquamic" organisms, meaning they take the inorganic plates from their prey to construct their test. ^[2]

Morphology

Members of this genus have a thin, transparent, pseudochitinous, flattened test that can be ovate, pyriform or elongate, with a length of around 180 microns. The surface of the test has numerous oval or circular scales of variable size, or in rare occasions rectangular or rod-like scales. The protoplasm is granular and colorless but can contain food vacuoles that show color. They have a single nucleus and a variable number of pseudopodia that are blunt in shape. The cell body is attached to the test's interior by strands of ectoplasm. ^[1]

Classification

Nebela originally belonged to the family Nebelidae, but phylogenetic analyses showed that the genus was paraphyletic and the genera Hyalosphenia and Quadrulella , which belong to Hyalospheniidae, branch within Nebela. Accordingly, the family Nebelidae was synonymised with Hyalospheniidae. ^[3]

Species

There are at least 13 remaining species in the genus:

• Nebela acolla Cash 1909 ^[4]
• Nebela aliciae Mitchell & Lara 2013 ^[4]
• Nebela barbata Leidy 1874 ^[5]
• Nebela carinatella Beyens & Chardez 1982 ^[6]
• Nebela collaris (Ehrenberg 1848) Leidy 1879 sensu Kosakyan & Gomaa 2013 (= Difflugia collarisEhrenberg 1848; =D. cancellataEhrenberg 1848; =D. reticulataEhrenberg 1848; =D. carpioEhrenberg 1854; =D. laxaEhrenberg 1871; =D. celluliferaEhrenberg 1874; =N. numataLeidy 1874; =N. bohemicaTaranek 1882; =N. sphagnophila(Steinecke) Van Oye 1933; =N. tincta var. majorDeflandre 1936; =N. tincta f. stenostomaJung 1936) ^[4]
• Nebela cylindrica Bonnet, 1979 ^{[7] [8]}
• Nebela equicalceus Leidy 1874 ^[9]
• Nebela flabellulum Leidy 1874
• Nebela guttata Kosakyan & Lara 2013 ^[4]
• Nebela pechorensis Kosakyan & Lara 2013 ^[4]
• Nebela rotunda Penard 1890 ^[4]
• Nebela tenella Penard 1893
• Nebela tincta (Leidy 1879) Awerintzew 1906 sensu Kosakyan & Lara 2013 (= Hyalosphenia tinctaLeidy 1879; =N. bursellaVejdovsky 1882; =N. parvulaCash & Hopkinson 1909; =N. minorPenard 1902) ^[4]

Former species

The paraphyly of Nebela is slowly being resolved by transferring species from Nebela to other genera through phylogenetic analyses. ^[3] The following species were previously considered Nebela but have been moved accordingly:

• Alabasta Duckert, Blandenier, Kosakyan & Singer 2018 ^[10]
  • Alabasta kivuense (Gauthier-Lievre & Thomas 1961) Duckert, Blandenier, Kosakyan & Singer 2018 (=N. kivuenseGauthier-Lievre & Thomas 1961 ^[8] )
  • Alabasta longicollis (Penard 1890) Duckert, Blandenier, Kosakyan & Singer 2018 (=N. longicollisPenard 1890)
  • Alabasta militaris (Penard 1890) Duckert, Blandenier, Kosakyan & Singer 2018 (=N. militarisPenard 1890; ^[11] =N. bursellaTaranek 1881; =N. americana var. bryophilaVan Oye 1933)
• Cornutheca Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 ^[12]
  • Cornutheca ansata (Leidy 1879) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. ansataLeidy 1879)
  • Cornutheca saccifera (Wailes 1913) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. sacciferaWailes 1913)
  • Cornutheca hippocrepis (=N. hippocrepis)
  • Cornutheca jiuhuensis ^[13] (=N. jiuhuensisQin, Mitchell & Lara 2016 ^[9] )
• Gibbocarina Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 ^[12]
  • Gibbocarina galeata (Penard 1890) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. galeataPenard 1890)
  • Gibbocarina gracilis (Penard 1910) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. gracilisPenard 1890)
• Longinebela Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 ^[12]
  • Longinebela golemanskyi (Todorov 2010) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. golemanskyiTodorov 2010 ^[14] )
  • Longinebela meisterfeldi (Heger & Mitchell 2012) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 ^[12] (=N. meisterfeldiHeger & Mitchell 2012) ^[3]
  • Longinebela penardiana (Deflandre 1936) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. penardianaDeflandre 1936)
  • Longinebela speciosa (Deflandre 1936) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. speciosaDeflandre 1936)
  • Longinebela tubulosa (Penard 1902) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. tubulosaPenard 1902)
• Netzelia Ogden 1979
  • Netzelia tuberculata (Wallich 1864) Netzel 1983 ^[15] (=N. tuberculata(Wallich 1864) Owen III & Jones 1976 ^[16] )
• Padaungiella Lara & Todorov 2012 ^[3]
  • Padaungiella lageniformis (Penard 1890) Lara & Todorov 2012 (=N. lageniformisPenard 1890)
  • Padaungiella wailesi (Deflandre 1936) Lara & Todorov 2012 (=N. wailesiDeflandre 1936)
  • Padaungiella wetekampi (Jung 1942) Lara & Todorov 2012 (=N. wetekampiJung 1942)
  • Padaungiella tubulata (Brown 1911) Lara & Todorov 2012 (=N. tubulataBrown 1911)
  • Padaungiella nebeloides (Gauthier-Lièvre & Thomas 1958) Lara & Todorov 2012 (=N. nebeloides (Gauthier-Lièvre & Thomas 1958) Todorov, Golemansky & Meisterfeld 2010 ^[17] )
• Planocarina Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 ^[12]
  • Planocarina carinata (Archer 1867) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. carinata(Archer 1867) Leidy 1879)
  • Planocarina marginata (Penard 1902) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. marginataPenard 1902)
  • Planocarina maxima (Awerintzew 1907) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. maximaAwerintzew 1907)
  • Planocarina spumosa (Awerintzew 1907) Kosakyan, Lahr, Mulot, Meisterfeld, Mitchell & Lara 2016 (=N. spumosaAwerintzew 1907)
• Physochila Jung 1942
  • Physochila griseola (Wailes & Penard 1911) (=N. griseolaWailes & Penard 1911)

References

1. Loeblich, Alfred Richard; Tappan, Helen (1964). Moore, Raymond C. (ed.). Treatise on Invertebrate Paleontology, Part C: Protista 2, Sarcodina, Chiefly "Thecamoebians" and Foraminiferida. Vol. 1. The Geological Society of America & The University of Kansas Press. ISBN   978-0813730035.
2. González-Miguéns, Rubén; Todorov, Milcho; Blandenier, Quentin; Duckert, Clément; Porfirio-Sousa, Alfredo L.; Ribeiro, Giulia M.; Ramos, Diana; Lahr, Daniel J.G.; Buckley, David; Lara, Enrique (2022). "Deconstructing Difflugia: The tangled evolution of lobose testate amoebae shells (Amoebozoa: Arcellinida) illustrates the importance of convergent evolution in protist phylogeny". Molecular Phylogenetics and Evolution. 175 (107557): 107557. Bibcode:2022MolPE.17507557G. doi: 10.1016/j.ympev.2022.107557 . hdl: 10261/281619 . ISSN   1055-7903. PMID   35777650.
3. Kosakyan A, Heger TJ, Leander BS, Todorov M, Mitchell EAD, Lara E (2012). "COI Barcoding of Nebelid Testate Amoebae (Amoebozoa: Arcellinida): Extensive Cryptic Diversity and Redefinition of the Hyalospheniidae Schultze" . Protist. 163 (3): 415–434. doi:10.1016/j.protis.2011.10.003. ISSN   1434-4610. PMID   22130576.
4. Kosakyan A, Gomaa F, Mitchell EAD, Heger TJ, Lara E (2013). "Using DNA-barcoding for sorting out protist species complexes: A case study of the Nebela tincta–collaris–bohemica group (Amoebozoa; Arcellinida, Hyalospheniidae)" . European Journal of Protistology. 49 (2): 222–237. doi:10.1016/j.ejop.2012.08.006. hdl:11380/1302341. ISSN   0932-4739. PMID   23092639.
5. Payne RJ, Bobrov AA, Tsyganov AN, Babeshko KV, Sloan TJ, Kay M, Kupriyanov DA, Surkov NV, Novenko EY, Andreev AA, Mazei YA (10 August 2020). "First records of contemporary testate amoeba assemblages from the Kamchatka Peninsula, Russia and potential for palaeoenvironmental reconstruction". Boreas. 50 (4): 998–1010. doi: 10.1111/bor.12469 .
6. Beyens L, Chardez D (1982). "Cryptodifflugia angustastoma et Nebela carinatella, nouveaux Thécamoebiens des tourbières dans la Campine beige". Archiv für Protistenkunde. 126 (2): 169–172. doi:10.1016/S0003-9365(82)80047-X. ISSN   0003-9365.
7. Bonnet L (1979). "Nouveaux thécamoebiens du sol X". Bulletin de la Société d'Histoire Naturelle de Toulouse. 115: 106–118.
8. Nicholls KH (2015). "Nebela kivuense Gauthier-Lievre et Thomas, 1961 (Amoebozoa, Arcellinida), Missing for a Half-century; Found 11,500 km from "home"". Acta Protozoologica. 54 (4): 283–288. doi:10.4467/16890027AP.15.023.3537. ISSN   0065-1583.
9. Qin Y, Man B, Kosakyan A, Lara E, Gu Y, Wang H, Mitchell EAD (2016). "Nebela jiuhuensis nov. sp. (Amoebozoa; Arcellinida; Hyalospheniidae): A New Member of the Nebela saccifera - equicalceus - ansata Group Described from Sphagnum Peatlands in South-Central China". The Journal of Eukaryotic Microbiology. 63 (5): 558–566. doi:10.1111/jeu.12300. hdl:11380/1302145. PMID   27593700. S2CID   24354110.
10. Duckert C, Blandenier Q, Kupferschmid FAL, Kosakyan A, Mitchell EAD, Lara E, et al. (2018). "En garde! Redefinition of Nebela militaris (Arcellinida, Hyalospheniidae) and erection of Alabasta gen. nov" (PDF). Eur J Protistol. 66: 156–165. doi:10.1016/j.ejop.2018.08.005. hdl:11380/1301652. PMID   30366198. S2CID   53116247.
11. Duckert C, Blandenier Q, Kupferschmid FAL, Kosakyan A, Mitchell EAD, Lara E, Singer D (30 April 2020). "Case 3782 – Nebela militaris Penard, 1890 (Arcellinida, Hyalospheniidae): proposed conservation of the specific name by giving it precedence over Nebela bursella Taranek, 1881". The Bulletin of Zoological Nomenclature. 77 (1): 22–28. doi:10.21805/bzn.v77.a007. hdl: 11380/1301647 . S2CID   218783898.
12. Kosakyan A, Lahr DJG, Mulot M, Meisterfeld R, Mitchell EAD, Lara E (2016). "Phylogenetic reconstruction based on COI reshuffles the taxonomy of hyalosphenid shelled (testate) amoebae and reveals the convoluted evolution of shell plate shapes". Cladistics. 32 (6): 606–623. doi: 10.1111/cla.12167 . PMID   34727671. S2CID   89592346.
13. Qin Y, Puppe D, Zhang L, Sun R, Li P, Xie S (August 2021). "How does Sphagnum growing affect testate Amoeba communities and corresponding protozoic Si pools? Results from field analyses in SW China". Microbial Ecology. 82 (2): 459–469. Bibcode:2021MicEc..82..459Q. doi:10.1007/s00248-020-01668-6. PMID   33442762. S2CID   231605855.
14. Todorov, Milcho (2010). "Nebela golemanskyi sp. nov., a New Sphagnicolous Testate Amoeba from Bulgaria (Amoebozoa: Arcellinida, Nebelidae)". Acta Protozoologica. 49 (1).
15. Bankov N, Todorov M, Ganeva A (22 May 2018). "Checklist of Sphagnum-dwelling testate amoebae in Bulgaria". Biodivers Data J. 6 (e25295): e25295. doi: 10.3897/BDJ.6.e25295 . PMC   5974006 . PMID   29861653.
16. Owen III, Goronwy; Jones, E. E. (1976). "Nebela tuberculata comb. nov. (Arcellinida), its History and Ultrastructure". The Journal of Eukaryotic Microbiology. 23 (4): 485–487. doi:10.1111/j.1550-7408.1976.tb03819.x.
17. Todorov M, Golemansky V, Meisterfeld R (2010). "Is Difflugia nebeloides (Amoebozoa: Arcellinida) really a Difflugia? Re-description and new combination". Acta Zoologica Bulgarica. 62 (1): 13–20.