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The apusomonads (family Apusomonadidae) are a group of
protozoan Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
zooflagellates that glide on surfaces, and mostly consume
prokaryotes A prokaryote (; less commonly spelled procaryote) is a single-celled organism whose cell lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Greek (), meaning 'before', and (), meaning 'nut' ...
. They are of particular evolutionary interest because they appear to be the sister group to the Opisthokonts, the clade that includes both
animals Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a ...
and
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
. Together with the Breviatea, these form the
Obazoa Obazoa is a proposed sister clade of Amoebozoa (which together form Amorphea). The term Obazoa is based on the OBA acronym for Opisthokonta, Breviatea, and Apusomonadidae, the group's three constituent clades. Determining the placement of Bre ...
clade.


Characteristics

Apusomonads are small
gliding Gliding is a recreational activity and competitive air sports, air sport in which pilots fly glider aircraft, unpowered aircraft known as Glider (sailplane), gliders or sailplanes using naturally occurring currents of rising air in the atmospher ...
heterotrophic A heterotroph (; ) is an organism that cannot produce its own food, instead taking nutrition from other sources of organic carbon, mainly plant or animal matter. In the food chain, heterotrophs are primary, secondary and tertiary consumers, but ...
biflagellates (i.e. with two
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
) that possess a
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
, formed partly or entirely by the anterior flagellum surrounded by a membranous sleeve. There is a pellicle under the dorsal cell membrane that extends into the proboscis sleeve and into a skirt that covers the sides of the cell. Apusomonads present two different cell plans: *Derived cell plan, represented by '' Apusomonas'', with a round cell body and a mastigophore, a projection of the cell containing both
basal bodies A basal body (synonymous with basal granule, kinetosome, and in older cytological literature with blepharoplast) is a protein structure found at the base of a eukaryotic undulipodium (cilium or flagellum). The basal body was named by Theodor ...
at its end. *"''Amastigomonas''-like" cell plan, with an oval or oblong cell that generally forms
pseudopodia A pseudopod or pseudopodium (: pseudopods or pseudopodia) is a temporary arm-like projection of a eukaryotic cell membrane that is emerged in the direction of movement. Filled with cytoplasm, pseudopodia primarily consist of actin filaments and ...
from the ventral surface, with no mastigophore, and the proboscis comprising solely the flagellum and the sleeve. These characteristics are considered 'primitive' or 'ancestral' in comparison with '' Apusomonas''. Organisms with this body plan, although historically assigned to the same genus '' Amastigomonas'', are a
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
group from which '' Apusomonas'' has evolved.


Evolution


External relationships

The apusomonads are the sister group to Opisthokonta, the lineage that includes
animal Animals are multicellular, eukaryotic organisms in the Biology, biological Kingdom (biology), kingdom Animalia (). With few exceptions, animals heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, ...
s,
fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...
and an array of related
protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...
. Because of this, apusomonads occupy an important
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
position to understand eukaryotic evolution. They retain ancestral characteristics, such as the biflagellate
body plan A body plan, (), or ground plan is a set of morphology (biology), morphological phenotypic trait, features common to many members of a phylum of animals. The vertebrates share one body plan, while invertebrates have many. This term, usually app ...
, which in opisthokonts evolves into a uniflagellate plan. Apusomonads are vital to understanding
multicellularity A multicellular organism is an organism that consists of more than one cell, unlike unicellular organisms. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and pa ...
. Genes involved in multicellularity have been found in the apusomonad '' Thecamonas'', such as adhesion proteins, calcium-signaling genes and types of sodium channels characteristic of animals. The
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
of the strain "'' Amastigomonas'' sp." presents the
integrin Integrins are transmembrane receptors that help cell–cell and cell–extracellular matrix (ECM) adhesion. Upon ligand binding, integrins activate signal transduction pathways that mediate cellular signals such as regulation of the cell cycle, o ...
-mediated
adhesion Adhesion is the tendency of dissimilar particles or interface (matter), surfaces to cling to one another. (Cohesion (chemistry), Cohesion refers to the tendency of similar or identical particles and surfaces to cling to one another.) The ...
machinery, the primary cell-matrix adhesion mechanism seen in
Metazoa Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia (). With few exceptions, animals consume organic material, breathe oxygen, have myocytes and are able to move, can reproduce sexually, and grow from a hol ...
(animals).


Internal relationships

Apusomonads are a poorly and narrowly studied group. Currently, the diversity of described apusomonads consists of the round '' Apusomonas'' and a wide array of "'' Amastigomonas''-type" organisms that have been reclassified into the genera '' Thecamonas'', '' Manchomonas'', '' Podomonas'', '' Multimonas'', '' Chelonemonas'' and, most recently, '' Catacumbia'', '' Cavaliersmithia'', '' Karpovia'', '' Mylnikovia'' and '' Singekia''. The relationships between these genera are depicted by the cladogram below.


Taxonomy


History

Apusomonads were first described in 1989 as one
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
Apusomonadidae inside the monotypic
order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...
Apusomonadida, as a group of flagellates containing the genera '' Apusomonas'' and '' Amastigomonas''. Later, British protozoologist
Thomas Cavalier-Smith Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow (21 October 1942 – 19 March 2021), was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford. His research has led to discov ...
classified them within the monotypic
class Class, Classes, or The Class may refer to: Common uses not otherwise categorized * Class (biology), a taxonomic rank * Class (knowledge representation), a collection of individuals or objects * Class (philosophy), an analytical concept used d ...
Thecomonadea as part of the
paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...
phylum In biology, a phylum (; : phyla) is a level of classification, or taxonomic rank, that is below Kingdom (biology), kingdom and above Class (biology), class. Traditionally, in botany the term division (taxonomy), division has been used instead ...
Apusozoa. Modern
cladistic Cladistics ( ; from Ancient Greek 'branch') is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is ...
approaches to
eukaryotic The eukaryotes ( ) constitute the Domain (biology), domain of Eukaryota or Eukarya, organisms whose Cell (biology), cells have a membrane-bound cell nucleus, nucleus. All animals, plants, Fungus, fungi, seaweeds, and many unicellular organisms ...
classification refer to apusomonads by their order-level name alone.


Classification

There are 10 recognized genera, as well as the "'' Amastigomonas''-like"
archetype The concept of an archetype ( ) appears in areas relating to behavior, historical psychology, philosophy and literary analysis. An archetype can be any of the following: # a statement, pattern of behavior, prototype, "first" form, or a main mo ...
that includes primitive forms not yet transferred to new genera. *'' Amastigomonas'' **''A. caudata'' 'Amastigomonas borokensis'' **''A. debruynei'' **''A. marisrubri'' *'' Catacumbia'' **''C. lutetiensis'' *'' Cavaliersmithia'' **''C. chaoae'' *'' Multimonas'' **''M. koreensis'' **''M. marina'' 'Cercomonas marina'' ; ''Amastigomonas marina'' **''M. media'' *'' Mylnikovia'' **''M. oxoniensis'' 'Thecamonas oxoniensis'' *'' Podomonas'' **''P. capensis'' **''P. gigantea'' 'Amastigomonas gigantea'' **''P. griebenis'' 'Amastigomonas griebenis'' **''P. kaiyoae'' **''P. klosteris'' 'Amastigomonas klosteris'' **''P. magna'' *Apusomonadinae Cavalier-Smith 2010 **'' Apusomonas'' 'Rostromonas'' ">Rostromonas.html" ;"title="'Rostromonas">'Rostromonas'' ***''A. australiensis'' ***''A. proboscidea'' [''Rostromonas applanata'' ] **'' Manchomonas'' ***''M. bermudensis'' [''Amastigomonas bermudensis'' ] *Thecamonadinae [''Thecamonas''/''Chelomonas'' clade] **'' Chelonemonas'' ***''C. dolani'' ***''C. geobuk'' ***''C. masanensis'' **'' Karpovia'' ***''K. croatica'' **'' Singekia'' ***''S. franciliensis'' ***''S. montserratensis'' **'' Thecamonas'' ***''T. filosa'' 'Amastigomonas filosa'' ***''T. muscula'' 'Amastigomonas muscula'' ***''T. mutabilis'' 'Rhynchomonas mutabilis'' ; ''Amastigomonas mutabilis'' ***''T. trahens'' 'Amastigomonas trahens''


References


External links

{{Taxonbar, from=Q21224721, from2=Q25365870, from3=Q4782334