View source for Biology:Limnofila

{{Short description|Genus of amoebae}}
{{Automatic taxobox
| taxon = Limnofila
| authority = Cavalier-Smith & Bass, 2009<ref name="novel naked filose"/>
| parent_authority = Cavalier-Smith & Bass, 2009
| grandparent_authority = Cavalier-Smith & Bass, 2009
| subdivision_ranks = Species
| subdivision =
* ''L. anglica''
* ''L. borokensis''
* ''L. longa''
* ''L. oxoniensis''
* ''L. mylnikovi''
| type_species = ''Limnofila borokensis''
| type_species_authority = Cavalier-Smith & Bass, 2009
}}

'''''Limnofila''''' ({{etymology|gre|limnos|marsh||fila|threads}}) is a genus of heterotrophic protists that live in freshwater habitats and feed on [[Biology:Bacteria|bacteria]]. They are also present in the [[Chemistry:Soil|soil]] ecosystem, where they play an important role as predators of [[Biology:Bacteria|bacteria]]. They are classified as a single family '''Limnofilidae''' and order '''Limnofilida'''.

== Characteristics ==

Limnofilida is [[Biology:Phylogenetic nomenclature|phylogenetically defined]] as the largest clade that includes ''Limnofila'', its only current member, but excludes ''[[Biology:Massisteria|Massisteria]]'', which belongs to order Leucodictyida. It excludes the phylogenetically distant
marine Nanofila which is morphologically similar but with unbranched filopodia. They are small freshwater heterotrophic protists with very slender, branching granular filopodia (or granulopodia), adhered to the substrate during feeding. Their trophic (feeding) phase is a small globular cell. They can present flagella, but are generally not visible under a light microscope.<ref name="novel naked filose"/>

The cellular [[Physics:Ultrastructure|ultrastructure]] of two species of Limnofilida has been studied (''L. borokensis'' and ''L. mylnikovi''). They present flat [[Biology:Crista|mitochondrial cristae]] and complex concentric-structured [[Biology:Extrusome|extrusome]]s. Their [[Biology:Filopodia|filopodia]] (filose, or thread-like, pseudopodia) are supported by bundles of 2–6 microtubules. Unlike [[Biology:Heliomonadida|Heliomonadida]], they have no prominent [[Biology:Centrosome|centrosome]].<ref name="novel naked filose"/>

== Ecology ==

Limnofilida are bacterivorous eukaryotes, they feed on [[Biology:Bacteria|bacteria]] through [[Biology:Phagocytosis|phagocytosis]]. As such, they are important soil predators of microbes and belong to a diverse ecological community of soil protists.<ref name="Degrune"/> Although they are described as a freshwater group,<ref name="novel naked filose"/> they are present in [[Chemistry:Soil|soil]] microbial communities. They appear as a minority of cercozoan [[Biology:Environmental DNA|environmental DNA sequences]] or [[Operational taxonomic unit|OTUs]] in temperate agricultural fields of {{wipe|Germany}} (1.3%).<ref name="Degrune">{{cite journal|first1=Florine |last1=Degrune |first2=Kenneth |last2=Dumack |first3=Anna Maria |last3=Fiore-Donno |first4=Michael |last4=Bonkowski |first5=Moisés A. |last5=Sosa-Hernández |first6=Michael |last6=Schloter |first7=Timo |last7=Kautz |first8=Doreen |last8=Fischer |first9=Matthias C. |last9=Rillig |title=Distinct communities of Cercozoa at different soil depths in a temperate agricultural field |journal=FEMS Microbiology Ecology |volume=95 |issue=4 |date=April 2019 |pages=fiz041|doi=10.1093/femsec/fiz041 |url=https://academic.oup.com/femsec/article/95/4/fiz041/5420472}}</ref> They also compose a similar minority of the cercozoan OTUs found in two separate [[Earth:Forest|forest]] soils of {{wipe|Norway}} and the [[Place:Czech Republic|Czech Republic]] (2%), but are present in a consistent manner across all levels: leaf litter, pure soil, [[Earth:Rhizosphere|rhizosphere]], and in association with plant roots.<ref name="Fiore-Donno 2022">{{cite journal|first1=Anna Maria|last1=Fiore-Donno|first2=Zander R.|last2=Human|first3=Martina|last3=Štursová|first4=Sunil|last4=Mundra|first5=Luis|last5=Morgado|first6=Håvard|last6=Kauserud|first7=Petr|last7=Baldrian|first8=Michael|last8=Bonkowski|title=Soil compartments (bulk soil, litter, root and rhizosphere) as main drivers of soil protistan communities distribution in forests with different nitrogen deposition|journal=Soil Biology and Biochemistry|volume=168|date=2022|pages=108628|doi=10.1016/j.soilbio.2022.108628|doi-access=free}}</ref>

== Systematics ==

{{Location map+ | Europe
|caption    = [[Biology:Type#type locality|Type locality]] of all five ''Limnofila'' species<ref name="novel naked filose"/>
|width      = 215
|float      = right
| places =
  {{Location map~ | Europe
    | label = ''L. anglica''<br/>''L. longa''
    | lat_deg = 54.4
    | lon_deg = -3.0 
    | position = top
    | background = white
    | mark = Cyan pog.svg
 }}
  {{Location map~ | Europe
    | label = {{nowrap|''L. oxoniensis''}}
    | coordinates = {{coord|51.7520|1.2}}
    | position = bottom
    | background = white
    | mark = Cyan pog.svg
  }}
  {{Location map~ | Europe
    | label = {{nowrap|''L. borokensis''}}<br/>{{nowrap|''L. mylnikovi''}}
    | coordinates = {{coord|57.7746|38.4938}}
    | position = top
    | background = white
    | mark = Cyan pog.svg
  }}
}}

=== Origin and etymology ===

The [[Biology:Genus|genus]] ''Limnofila'' was created in 2009 by [[Biography:Protistologist|protistologist]]s [[Biography:Thomas Cavalier-Smith|Thomas Cavalier-Smith]] and David Bass, through a research article published in the journal ''Protist''. It was described to encompass several naked amoebae with granular filose [[Biology:Pseudopodia|pseudopodia]] (or [[Biology:Filopodia|filopodia]]) that did not belong in phylogenetic analyses to either [[Biology:Endomyxa|Endomyxa]] or [[Biology:Foraminifera|Foraminifera]]. On the basis of phylogenetic analyses based on 18S rRNA [[Biology:Gene|gene]]s, they were assigned to a [[Biology:Clade|clade]] of [[Biology:Cercozoa|Cercozoa]] known as [[Biology:Granofilosea|Granofilosea]], which was newly described in the same article. In particular, the genus was assigned to a monotypic order '''Limnofilida''' and family '''Limnofilidae'''. Limnofilida was defined as the most inclusive clade containing ''Limnofila'' but excluding ''[[Biology:Massisteria|Massisteria]]'', which belongs to Leucodictyida. The name ''Limnofila'' comes {{etymology|gre|limnos|marsh||fila|threads}}, to emphasise both their exclusively freshwater habitat and their exceedingly thin thread-like filopodia.<ref name="novel naked filose"/>

=== Molecular phylogeny ===

''Limnofila'', and Limnofilida as a whole, are the earliest [[Biology:Divergent evolution|diverging]] [[Biology:Clade|clade]] within [[Biology:Granofilosea|Granofilosea]], followed by Leucodictyida, Cryptofilida and [[Biology:Desmothoracida|Desmothoracida]]. Granofilosea is, in turn, the [[Biology:Sister group|sister group]] of [[Biology:Monadofilosa|Monadofilosa]], which contains the remaining [[Biology:Filopodia|filose]] cercozoan amoebae. Below is a simplified [[Biology:Cladogram|cladogram]] of [[Biology:Granofilosea|Granofilosea]] from a 2011 phylogenetic analysis based on 18S rRNA [[Biology:Gene|gene]]s, where the clades of [[Biology:Environmental DNA|environmental DNA]] are not shown.<ref name="novel naked filose"/><ref name="novel cultured protists"/>

{{clade|style=font-size:90%;|2=[[Biology:Monadofilosa|Monadofilosa]]|label1=[[Biology:Granofilosea|Granofilosea]]|1={{clade|2={{clade|label1='''Limnofilida'''&nbsp;|1='''''Limnofila'''''}}|1={{clade|2={{clade|label1=Leucodictyida&nbsp;|1={{clade|1=Leucodictyidae|2=[[Biology:Massisteriidae|Massisteriidae]]}}}}|1={{clade|label1=[[Biology:Desmothoracida|Desmothoracida]]|1=Clathrulinidae|label2=Cryptofilida|2={{clade|1=''[[Biology:Mesofila|Mesofila]]''|2=''[[Biology:Nanofila|Nanofila]]''}}}}}}}}}}

=== Species ===

There are 5 species of ''Limnofila'', all described in the same 2009 paper where the genus was first described. Two species, ''L. anglica'' and ''L. longa'', were isolated from a freshwater lake known as Priest Pot in Cumbria, {{wipe|England}}.<ref>{{citation|vauthors=Calabuig I|date=2016|title=Priest Pot species list, Cumbria, Britain|publisher=Department of Biology, University of Copenhagen|doi=10.15468/lih6qc|access-date=2023-09-22|url=https://www.gbif.org/es/dataset/84d4a910-f762-11e1-a439-00145eb45e9a}}</ref> One species, ''L. oxoniensis'', was isolated from garden soil in Oxford. The remaining two species, ''L. borokensis'' and ''L. mylnikovi'', were isolated from samples collected in waste treatment plants of Borok, Yaroslavskaya oblast, {{wipe|Russia}}, and were originally misidentified as ''Gymnophrys cometa''.<ref name = "Nikolaev2003"/><ref name="novel naked filose"/>

# ''Limnofila anglica'' — Cumbria, {{wipe|England}}.
# ''Limnofila borokensis'' — Borok, {{wipe|Russia}}.
# ''Limnofila longa'' — Cumbria, England.
# ''Limnofila oxoniensis'' — Oxford, England.
# ''Limnofila mylnikovi'' — Borok, Russia ([[Biology:DNA sequencing|unsequenced]]).

== References ==
{{reflist|refs=

<ref name="novel naked filose">{{cite journal |vauthors=Bass D, Chao EE, Nikolaev S, Yabuki A, Ishida K, Berney C, Pakzad U, Wylezich C, Cavalier-Smith T |title=Phylogeny of novel naked Filose and Reticulose Cercozoa: Granofilosea cl. n. and Proteomyxidea revised |journal=Protist |volume=160 |issue=1 |pages=75–109 |year=2009 |doi=10.1016/j.protis.2008.07.002 |pmid=18952499 }}</ref>

<ref name="novel cultured protists">{{cite journal|vauthors=Howe AT, Bass D, Scoble JM, Lewis R, Vickerman K, Arndt H, Cavalier-Smith T|title=Novel Cultured Protists Identify Deep-branching Environmental DNA Clades of Cercozoa: New Genera Tremula, Micrometopion, Minimassisteria, Nudifila, Peregrinia|journal=Protist|volume=162|issue=2|date=2011|pages=332–372|issn=1434-4610|doi=10.1016/j.protis.2010.10.002}}</ref>

<ref name = "Nikolaev2003">{{cite journal | author = Nikolaev, S.I. | title = ''Gymnophrys cometa'' and ''Lecythium'' sp. are Core Cercozoa: Evolutionary Implications | journal = Acta Protozoologica | year = 2003 | volume = 42 | pages = 183–190 | url = http://www.nencki.gov.pl/pdf/ap/ap696.pdf | display-authors = etal | url-status = dead | archiveurl = https://web.archive.org/web/20090617090207/http://www.nencki.gov.pl/pdf/ap/ap696.pdf | archivedate = 2009-06-17 }}</ref>

}}

{{taxonbar|from1=Q1825529|from2=Q19833394|from3=Q19833395}}

[[Category:Cercozoa genera]]

{{Sourceattribution|Limnofila}}