Helkesida

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Helkesida
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Phylum: Cercozoa
Superclass: Eoglissa
Class: Helkesea
Cavalier-Smith 2018
Order: Helkesida
Cavalier-Smith 2018 [1]
Families [1] [2]
Diversity
24 species

Helkesida (formerly known as Sainouroidea) [1] is a group of microscopic protists belonging to the supergroup Rhizaria, both discovered through molecular phylogenetic analyses. It contains amoeboid flagellates with two flagella. They are either free-living, mostly on fecal matter, or live inside the gut of animals. Among these amoebae, one lineage has independently evolved aggregative multicellularity similarly to slime moulds. [3]

Contents

Biology

The organisms classified as Helkesida commonly have a gliding motility in which the cells glide on their posterior flagellum. They are ancestrally amoeboid bi-flagellates without scales or theca. [2] Unlike most Cercozoa which have tubular mitochondrial cristae, they can also present flat cristae or discoid cristae. They are the only group within Rhizaria that present discoid mitochondrial cristae. [4]

These organisms have an amorphous apical centrosome attached to the nucleus by a rhizoplast. The kinetid arises from 2–4 very short centrioles with dense fibrous roots that attach them to each other and to the nucleus. Their anterior flagellum is reduced to a stub without its 9+2 axoneme. The centrosome also generates numerous microtubules in larger cells. The Golgi apparatus is seen attached to the nuclear envelope and the anterior rhizoplast. They have a microbody attached to the posterior end of the nucleus. [5]

One helkesid genus, Guttulinopsis , represents an independent lineage in which aggregative multicellularity has evolved to generate "fungi-like" fruiting bodies called sorocarps, similarly to slime moulds such as Dictyostelium . [4]

Ecology

The helkesid amoebae are bacterivores that can be free-living, mostly associated to fecal environments, or endozoic, associated to animals. [6] They thrive in aerobic conditions and the microaerophilic gut environment of animals. Rosculus can thrive in anaerobic culture. It is unknown if their preferred habitat is free-living or endozoic. [3]

Some host species can harbor different helkesid genera and species. One animal can be infected by multiple species simultaneously, and one species can also infect different animal hosts. More sampling of hosts, amoebae and molecular data is needed to better understand the life history and ecology of these protists. [3]

Evolution and systematics

History

Helkesida is a group initially named Sainouroidea. It was discovered in 2009 as a highly divergent clade within Cercozoa through phylogenetic analyses that used the sequencing of 18S ribosomal RNA from Cholamonas cytrodiopsidis , Sainouron acronematica and Helkesimastix marina . It is a molecularly diverse clade that branches within a group of ancestrally amoeboid bi-flagellates that usually lack an outer cell coat, known as Monadofilosa. [5] A 2016 study revealed a previously unknown wide diversity of Sainouroidea in fecal environments. Previous environmental samplings excluded sequences from Sainouroidea due to their highly divergent 18S rDNA sequences. [6] A 2018 study described several new genera and species. [3]

The initial name for this group, Sainouroidea, had the -oidea suffix for superfamilies, but it was not assigned to any existing classes or orders due to the uncertainty of its phylogenetic position. [5] In a 2018 revision, the class Helkesea and order Helkesida were created as a substitute for this name. Sainouroidea was then modified to only include one of the three helkesid families, Sainouridae. A second superfamily, Helkesimastigoidea, was created to host the remaining two families, Helkesimastigidae and Guttulinopsidae. [1]

Classification

Phylogeny of Helkesida

Cholamonas

Sainouron

Acantholus

Homocognata

Helkesimastix

Puppisaman

Olivorum

multicellularity

Guttulinopsis

Rosculus

Cladogram of Helkesida, based on a 18S rDNA phylogenetic analysis within a 2018 study. [3] The study names this group "Sainouroidea" due to being published prior to the taxonomic change to Helkesida. [1]

Currently, Helkesida contains 24 species distributed in 9 genera, 3 families [2] and 2 superfamilies. [1] Additionally, many OTUs found through environmental sequencing may represent undescribed clades. [4] [6]

Related Research Articles

<span class="mw-page-title-main">Flagellate</span> Group of protists with at least one whip-like appendage

A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word flagellate also describes a particular construction characteristic of many prokaryotes and eukaryotes and their means of motion. The term presently does not imply any specific relationship or classification of the organisms that possess flagella. However, the term "flagellate" is included in other terms which are more formally characterized.

<span class="mw-page-title-main">Percolozoa</span> Phylum of Excavata

The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages.

<span class="mw-page-title-main">Nucleariida</span> Order of amoebae

Nucleariida is a group of amoebae with filose pseudopods, known mostly from soils and freshwater. They are distinguished from the superficially similar vampyrellids mainly by having mitochondria with discoid cristae, in the absence of superficial granules, and in the way they consume food.

<span class="mw-page-title-main">Vampyrellidae</span> Family of single-celled organisms

The family Vampyrellidae is a subgroup of the order Vampyrellida within the supergroup Rhizaria. Based on molecular sequence data, the family currently comprises the genus Vampyrella, and maybe several other vampyrellid amoebae. The cells are naked and characterised by radiating, filose pseudopodia and an orange colouration of the main cell body.

<span class="mw-page-title-main">Heliozoa</span> Phylum of protists with spherical bodies

Heliozoa, commonly known as sun-animalcules, are microbial eukaryotes (protists) with stiff arms (axopodia) radiating from their spherical bodies, which are responsible for their common name. The axopodia are microtubule-supported projections from the amoeboid cell body, and are variously used for capturing food, sensation, movement, and attachment. They are similar to Radiolaria, but they are distinguished from them by lacking central capsules and other complex skeletal elements, although some produce simple scales and spines. They may be found in both freshwater and marine environments.

<span class="mw-page-title-main">Cercozoa</span> Group of single-celled organisms

Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major eukaryotic group to be recognized mainly through molecular phylogenies. They are the natural predators of many species of bacteria. They are closely related to the phylum Retaria, comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria.

<span class="mw-page-title-main">Rhizaria</span> Infrakingdom of protists

The Rhizaria are a diverse and species-rich supergroup of mostly unicellular eukaryotes. Except for the Chlorarachniophytes and three species in the genus Paulinella in the phylum Cercozoa, they are all non-photosynthethic, but many foraminifera and radiolaria have a symbiotic relationship with unicellular algae. A multicellular form, Guttulinopsis vulgaris, a cellular slime mold, has been described. This group was used by Cavalier-Smith in 2002, although the term "Rhizaria" had been long used for clades within the currently recognized taxon. Being described mainly from rDNA sequences, they vary considerably in form, having no clear morphological distinctive characters (synapomorphies), but for the most part they are amoeboids with filose, reticulose, or microtubule-supported pseudopods. In the absence of an apomorphy, the group is ill-defined, and its composition has been very fluid. Some Rhizaria possess mineral exoskeletons, which are in different clades within Rhizaria made out of opal, celestite, or calcite. Certain species can attain sizes of more than a centimeter with some species being able to form cylindrical colonies approximately 1 cm in diameter and greater than 1 m in length. They feed by capturing and engulfing prey with the extensions of their pseudopodia; forms that are symbiotic with unicellular algae contribute significantly to the total primary production of the ocean.

<span class="mw-page-title-main">Phaeodarea</span> Class of protists

Phaeodarea or Phaeodaria is a group of amoeboid cercozoan organisms. They are traditionally considered radiolarians, but in molecular trees do not appear to be close relatives of the other groups, and are instead placed among the Cercozoa. They are distinguished by the structure of their central capsule and by the presence of a phaeodium, an aggregate of waste particles within the cell.

<span class="mw-page-title-main">Lobosa</span> Phylum of protozoans

Lobosa is a taxonomic group of amoebae in the phylum Amoebozoa. Most lobosans possess broad, bluntly rounded pseudopods, although one genus in the group, the recently discovered Sapocribrum, has slender and threadlike (filose) pseudopodia. In current classification schemes, Lobosa is a subphylum, composed mainly of amoebae that have lobose pseudopods but lack cilia or flagella.

<span class="mw-page-title-main">Cercomonad</span> Order of single-celled organisms

Cercomonads are small amoeboflagellates, widespread in aqueous habitats and common in soils.

<span class="mw-page-title-main">Tectofilosid</span> Group of protists

The tectofilosids are a group of filose amoebae with shells. These are composed of organic materials and sometimes collected debris, in contrast to the euglyphids, which produce shells from siliceous scales. The shell usually has a single opening, but in Amphitrema and a few other genera it has two on opposite ends. The cell itself occupies most of the shell. They are most often found on marsh plants such as Sphagnum.

<span class="mw-page-title-main">Monadofilosa</span> Group of protists

Monadofilosa is a grouping of Cercozoa. These organisms are single-celled amoeboid protists.

<span class="mw-page-title-main">Thaumatomonadida</span> Order of single-celled organisms

Thaumatomonadida is an order of flagellates.

<span class="mw-page-title-main">Thecofilosea</span> Class of single-celled organisms

Thecofilosea is a class of unicellular testate amoebae belonging to the phylum Cercozoa. They are amoeboflagellates, organisms with flagella and pseudopodia, distinguished from other cercozoa by their scale-lacking test composed of organic material. They are closely related to the Imbricatea, a group of testate amoebae with tests composed of inorganic silica scales.

<span class="mw-page-title-main">Sarcomonadea</span> Class of flagellate protists

The sarcomonads or class Sarcomonadea are a group of amoeboid biciliate protists in the phylum Cercozoa. They are characterized by a propensity to move through gliding on their posterior cilium or through filopodia, a lack of scales or external theca, a soft cell surface without obvious cortical filamentous or membranous skeleton, two cilia without scales or hairs, tubular mitochondrial cristae, near-spherical extrusomes, and a microbody attached to the nucleus.

Leucodictyids are heterotrophic amoeboid protists that comprise the order Leucodictyida in the phylum Cercozoa.

Cryptofilida is an order of small heterotrophic protists in the phylum Cercozoa. They are filose amoebae that lack cilia and gliding, and are instead characterized by movement through branching or unbranched granular filopodia that are appressed to the substrate during their feeding.

<span class="mw-page-title-main">Glissomonadida</span> Order of protists

The glissomonads are a group of bacterivorous gliding flagellated protists that compose the order Glissomonadida, in the amoeboflagellate phylum Cercozoa. They comprise a vast, largely undescribed diversity of soil and freshwater organisms. They are the sister group to cercomonads; the two orders form a solid clade of gliding soil-dwelling flagellates called Pediglissa.

Pediglissa is a subclass of phagotrophic protists that inhabit soil or freshwater habitats. They were defined in 2018 according to phylogenetic analyses that showed a clade containing the orders Cercomonadida and Glissomonadida. They're the sister group of Paracercomonadida.

<span class="mw-page-title-main">Amoeboflagellate</span> Cellular body type

An amoeboflagellate is any eukaryotic organism capable of behaving as an amoeba and as a flagellate at some point during their life cycle. Amoeboflagellates present both pseudopodia and at least one flagellum, often simultaneously.

References

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