Malawimonas | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Malawimonada |
Phylum: | Malawimonada |
Class: | Malawimonadea |
Order: | Malawimonadida |
Family: | Malawimonadidae |
Genus: | Malawimonas O’Kelly & Nerad 1999 |
Type species | |
Malawimonas jakobiformis O’Kelly & Nerad 1999 | |
Species [1] | |
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Malawimonas is genus of unicellular, heterotrophic flagellates with uncertain phylogenetic affinities. They have variably being assigned to Excavata and Loukozoa. [2] [3] Recent studies suggest they may be closely related to the Podiata. [4] [5]
In 1993, Charles J O’ Kelly studied the jakobid groups flagellates and implications for the early diversification of eukaryotes and recognized that Jakoba, Reclimonas, and Histonia or often refer as “core jakobids” were morphologically somewhat similar. Interestingly, they included an unnamed and undescribed free-swimming, flagellate, and also groove- bearing cell. [6] [7]
During the early study, these cells were thought to be a member from Jakoba due to the external morphology features that resembles Jakoba libera in terms of lack of cell covering, sessile trophic stages, swimming in a similar manner and sharing the tendency for the anterior flagellum to form a “crook. However, later discovery found that this species seems to not fit and can not be assigned to the genus Jakoba, nor to any other genus of Jakobids, because of their discoidal mitochondrial cristae, which is different from Jakoba that have irregularly flattened, and the other Jakobids member have tubular mitochondrial cristae. [8]
This undescribed organism was later described formally as Malawimonas jakobiformis and placed in its own new family Malawimonadidae, a bacterivorous heterotrophic isolated from the Malawi shore of Lake Nyasa (eastern Africa). [8]
Only more than a decade later, a related organism has been studied under several studies that revolved around the phylogenetic positions of the Jakobids and Cercozoans group under the names Malawimonas californiana, but there are no formal descriptions included. [9] [10] [11]
Malawimonas share some common features with the jakobids and other Excavata group members by having a conspicuous feeding grove on the ventral side and two flagellates. The overall cytoskeleton of Malawimonas resembles Carpediemonas, typical Excavata belonging to the anaerobic Metamonada clade and closely related to Diplomonads and Retormonads. [3] [11]
Malawimonas jakobiformis characterized by a uninucleate, biflagellate, heterotrophic, “naked” cell, where neither scales nor lorica was present. Observation of the ultrastructure revealed a substantial glycocalyx as a surface coat. The cells of Malawimonas are usually slender, and the shape is plastic and often deformed by influences such as coverslip pressure or food they digest. The two flagella are approximately 1- 1.5 times as long as the cell body and are more or less equal length. The anterior flagellum has a “crook” shape where the posterior flagellum is appresed but not attached to the ventral cell surface. Cells swim in straight lines a and turn longitudinal axes as they move. The posterior flagellar vane of Malawimonas jakobiformis arises from a clearly defined point on the ventral surface of the flagellum. In contrast, the other jakobids’ vane has a diffuse origin along the dorsal surface. [8] [12]
The early study of molecular investigations revealed that the genome of Reclinomonas americana, Jakoba libera and Malawimonas jakobiformis collectively represent the most eubacterial-like mitochondrial DNAs yet discovered among all eukaryotes. [9] Jakobids group have all the basic forms of mitochondrial cristae known in eukaryotes. The mitochondrial shape has been seen as a strongly conserved character and is used to delimit the deepest evolutionary division within eukaryotes. [13]
A study based on alpha and beta- tubulin phylogenies found that Malawimonas jakobiformis occupied a relatively basal position in the plant-protists superclade and showed the distinctness from the “core jakobids”. [13]
Molecular phylogenies have not resolved the position of malawimonads within eukaryotes. Analyses of the small number of nucleus-encoded genes often place malawimonads as close relatives of metamonads. Since 2016, most phylogenomic shows place malawimonads separately from other excavates. Phylogenesis of SSU rDNA, tubulins, 5-7 nucleus-encoded proteins, and phylogenetic analyses conclude that malawimonads are not specifically related to jakobids. [14] [15] [10] [16] The similarities between jakobids and malawimonads apparently reflect the ancestral morphology of excavate protists, or perhaps convergence, and not a close phylogenetic relationship. [11]
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.
The Stramenopiles, also called Heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost. Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.
Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.
The opisthokonts are a broad group of eukaryotes, including both the animal and fungus kingdoms. The opisthokonts, previously called the "Fungi/Metazoa group", are generally recognized as a clade. Opisthokonts together with Apusomonadida and Breviata comprise the larger clade Obazoa.
Telonemia is a phylum of microscopic eukaryotes commonly known as telonemids. They are unicellular free-living flagellates with a unique combination of cell structures, including a highly complex cytoskeleton unseen in other eukaryotes.
Reclinomonas is a monotypic genus of jakobid eukaryotes containing the single species Reclinomonas americana.
Malawimonadidae is a family of unicellular eukaryotes of outsize importance in understanding eukaryote phylogeny.
Breviata anathema is a single-celled flagellate amoeboid eukaryote, previously studied under the name Mastigamoeba invertens. The cell lacks mitochondria, much like the pelobionts to which the species was previously assigned, but has remnant mitochondrial genes, and possesses an organelle believed to be a modified anaerobic mitochondrion, similar to the mitosomes and hydrogenosomes found in other eukaryotes that live in low-oxygen environments.
Carpediemonas is genus of Metamonada, and belongs to the group Excavata. This organism is a unicellular flagellated eukaryote that was first discovered in substrate samples from the Great Barrier Reef. Carpediemonas can be found in anaerobic intertidal sediment, where it feeds on bacteria. A feature of this species is the presence of a feeding groove, a characteristic of the excavates. Like most other metamonads, Carpediemonas does not rely on an aerobic mitochondrion to produce energy. Instead, it contains hydrogenosomes that are used to produce ATP. This organism has two flagella: a posterior one used for feeding on the substrate, and an anterior one that moves in a slower sweeping motion. Carpediemonas is assigned to the fornicates, where similar Carpediemonas-like organisms are used in researching the evolution within excavates. Although Carpediemonas is a member of the metamonads, it is unusual in the sense that it is free-living and has three basal bodies.
Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small, and can be found in aerobic and anaerobic environments. The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993. There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.
Jakoba is a genus in the taxon Excavata, and currently has a single described species, Jakoba libera described by Patterson in 1990, and named in honour of Dutch botanist Jakoba Ruinen.
Andalucia is a genus of jakobids, currently containing the sole species A. godoyi.
Monocercomonoides is a genus of flagellate Excavata belonging to the order Oxymonadida. It was established by Bernard V. Travis and was first described as those with "polymastiginid flagellates having three anterior flagella and a trailing one originating at a single basal granule located in front of the anteriorly positioned nucleus, and a more or less well-defined axostyle". It is the first eukaryotic genus to be found to completely lack mitochondria, and all hallmark proteins responsible for mitochondrial function. The genus also lacks any other mitochondria related organelles (MROs) such as hydrogenosomes or mitosomes. Data suggests that the absence of mitochondria is not an ancestral feature, but rather due to secondary loss. Monocercomonoides sp. was found to obtain energy through an enzymatic action of nutrients absorbed from the environment. The genus has replaced the iron-sulfur cluster assembly pathway with a cytosolic sulfur mobilization system, likely acquired by horizontal gene transfer from a eubacterium of a common ancestor of oxymonads. These organisms are significant because they undermine assumptions that eukaryotes must have mitochondria to properly function. The genome of Monocercomonoides exilis has approximately 82 million base pairs, with 18 152 predicted protein-coding genes.
Stygiella /ˌstɪ.d͡ʒiˈɛ.lə/ is a genus of free-living marine flagellates belonging to the family Stygiellidae in the jakobids (excavata).
Helkesida 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.
Stygiellidae is a family of free-living marine flagellates belonging to the order Jakobida, a deep-branching lineage within the eukaryotic supergroup Discoba. They are unicellular organisms that commonly inhabit anoxic, sulfide-rich and ammonium-rich marine habitats worldwide.
Imasa is a genus of marine flagellates containing the single species Imasa heleensis, discovered in 2020 in the Solomon Islands. It is the first marine member of a basal group of eukaryotes known as Malawimonadida. It is the only member of the family Imasidae.
Urceolus is a genus of heterotrophic flagellates belonging to the Euglenozoa, a phylum of single-celled eukaryotes or protists. Described by Russian biologist Konstantin Mereschkowsky in 1877, its type species is Urceolus alenizini. Species of this genus are characterized by deformable flask-shaped cells that exhibit at least one flagellum that is active at the tip, arising from a neck-like structure that also hosts the feeding apparatus. They are found in a variety of water body sediments across the globe. According to evolutionary studies, Urceolus belongs to a group of Euglenozoa known as peranemids, closely related to the euglenophyte algae.
Colponemids are free-living alveolates, unicellular flagellates related to dinoflagellates, apicomplexans and ciliates. They are predators of other small eukaryotes, found in freshwater, marine and soil environments. They do not form a solid clade, but a sparse group of deep-branching alveolate lineages.
Paratrimastix pyriformis is a species of free-living anaerobic freshwater bacteriovorous flagellated protists formerly known as Trimastix pyriformis and Tetramitus pyriformis.