Telonemia is a phylum of microscopic commonly known as telonemids. They are unicellular free-living with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.
Telonemia shares several distinctive features with its related group, the SAR supergroup. Among these features are cortical alveoli, small sacs beneath the cell's surface that act as cushions, providing support and helping to maintain the cell's shape. Additionally, they possess tripartite , complex three-part hair-like structures on their flagella, the whip-like tails used for movement. These structures enhance their swimming capabilities by increasing resistance against water. Furthermore, Telonemia is equipped with filopodia, very thin, thread-like projections extending from the cell body. These projections can serve various purposes, such as aiding in movement or capturing food particles by wrapping around them. Together, the two lineages compose the TSAR clade.
This phylum is monotypic, composed of a single class Telonemea, order Telonemida and family Telonemidae. It is classified in three Genus and seven species, although numerous undescribed clades of environmental DNA are known. They are detected in all marine and freshwater environments, where they prey on bacteria and small phytoplankton by engulfing them in their plasma membrane (phagotrophy).
Morphology
The phylum Telonemia comprises microscopic
unicellular , or protists.
Most of the
biodiversity of telonemids is morphologically uncharacterized.
The few described species are free-living predatory
Phagocytosis Flagellate composed of pear-shaped cells with two
Flagellum. These cells measure approximately 5–10 μm in length and 3–7 μm in width. The flagella have different lengths, with the short one measuring up to 12 μm and the long one measuring up to 16 μm. Between the flagella protrudes a short
proboscis-like structure, known as a rostrum. Their
crista are tubular. They have a unique multi-layered
cytoskeleton of high complexity, composed of layers of
and
Microtubule, unseen in any other eukaryote.
They exhibit a unique combination of cell traits that were previously believed to be
synapomorphic to different
Chromalveolata groups, such as complex tripartite mastigonemes (as in
Stramenopile), cortical alveoli-like structures (as in
) and
filopodia (as in
).
Despite their evolutionary proximity to chromalveolates, they lack
chloroplasts.
Ecology and distribution
Telonemids feed on a wide range of organisms, namely
bacteria and
phytoplankton ranging in size between
picoplankton and
nanoplankton. They are widely distributed and are sometimes abundant, implying they may play an important ecological role in aquatic ecosystems.
Around one hundred
clades of environmental sequences from undescribed telonemids have been recovered in a variety of
sea (
Antarctic,
Arctic and
;
Mediterranean,
Baltic Sea,
Kara Sea, Marmara and
White Sea seas), including
deep sea, and
Fresh water from different regions (
Norway,
France,
Antarctica,
Finland,
Canada,
Japan).
Several telonemid
clades favor open waters with lower nutrients, such as the
Canada Basin and offshore the
Mackenzie River, suggesting that they are able to thrive in low-productivity ecosystems (i.e.
oligotrophic).
Systematics
History
The first telonemid
genus and
species,
Telonema subtile, was described by Karl Griessmann in 1913 from crude cultures of the
green alga Ulva and of
red algae off the coast of
Roscoff and
Naples.
Eighty years later, in 1993, American protistologist Thomas Cavalier-Smith created a family
Telonemidae, order
Telonemida and class
Telonemea to contain this
protist. Initially, this group was included within the now obsolete phylum
Opalozoa, along with other unrelated groups of
such as
,
,
,
,
,
,
and so on. In this scheme, the class Telonemea was distinguished by the presence of two posterior
cilia of equal length (isokont cilia). It contained an additional order besides Telonemida, Nephromycida, which comprised the genus
Nephromyces (later treated as an
apicomplexan).
In 2005 a second species of telonemid was described,
T. antarcticum, from the surface waters of the
Oslofjord.
Since 2006, Telonemea was separated into a new eukaryotic phylum Telonemia by protistologist Kamran Shalchian-Tabrizi and coauthors, on the basis of Phylogenetics that placed it near chromalveolate groups such as Haptista and Cryptista. However, in 2015, Cavalier-Smith and coauthors rejected their treatment as an independent phylum and transferred Telonemea to the phylum Cryptista, under the obsolete subphylum Corbihelia. This subphylum included other protists with a pharyngeal basket or radiating axopodia such as Picomonas (later classified as a separate phylum Picozoa closely related to red algae) and Microheliella (now proposed as the sister group to Cryptista). In addition, they transferred T. antarcticum to a new genus Lateronema, on the basis of genetic distance from Telonema.
Numerous phylogenetic analyses in the following years solidified the position of Telonemia as the sister clade to the SAR supergroup, both collectively composing the TSAR clade ( Telonemia, Stramenopila, Alveolata and Rhizaria), which lead Cavalier-Smith to finally consider Telonemia a separate phylum in 2022. In the same year, five more species and a third genus, Arpakorses, were described by protistologist Denis Victorovich Tikhonenkov and coauthors.
Classification
Until 2019, only two species had been formally described,
although DNA sequences collected from seawater suggested there were many more species not yet described.
In 2022, five additional species were described along with a third new genus, bringing the total number of species to seven.
Evolution
Telonemia is a
clade of protists that branch independently from other eukaryotic supergroups groups as their own 'micro-kingdom'.
Early molecular analyses of Telonemia placed them as an independent branch within the
SAR supergroup, a diverse clade of eukaryotes that contain
Rhizaria,
Alveolata and
Stramenopila.
Other analyses proposed a close relationship with
,
,
and
.
At this time, they were suggested to have
evolutionary significance in being a possible transitional form between ecologically important
heterotrophic and
photosynthetic species among chromalveolates.
The present phylogenetic analyses place them as sister to the SAR supergroup in a clade commonly known as TSAR, which is widely accepted by the scientific community. As the sister clade to SAR, Telonemia has a key position in the tree of eukaryotic life. They are morphologically complex organisms that combine characteristics of different SAR lineages. The main trait uniting each SAR lineage has been described in at least one genus of Telonemia: tripartite in the flagella, typical of stramenopiles and described in Lateronema; cortical alveoli underneath the plasma membrane, typical of and described in Lateronema; and fine pseudopodia (filopodia), typical of and described in Telonema. Moreover, Arpakorses presents a kinetid structure similar to that seen in Rhizaria, and Telonema subtile presents microtubules in a formation superficially resembling the apical complex of .
All telonemid genera possess a highly intricate multi-layered cytoskeleton, whose complexity is not found in any other eukaryote. This finding may indicate that telonemids have retained an ancestral cytoskeleton organization that has been lost in other eukaryotes.
Notes
External links
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Taxonomy, from the Taxonomy Browser of the National Center for Biotechnology Information.
-
Images, etc. from iSpecies.org