The alveolates (meaning "pitted like a honeycomb") are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria, the group being referred to as SAR.

Characteristics

The most notable shared characteristic is the presence of cortical (near the surface) alveoli (sacs). These are flattened vesicles (sacs) arranged as a layer just under the

membrane A membrane is a selective barrier; it allows some things to pass through but stops others. Such things may be molecules, ions, or other small particles. Membranes can be generally classified into synthetic membranes and biological membranes. ...

and supporting it, typically contributing to a flexible pellicle (thin skin). In armored dinoflagellates they may contain stiff plates. Alveolates have mitochondria with tubular cristae (invaginations), and cells often have pore-like intrusions through the cell surface. The group contains free-living and parasitic organisms, predatory flagellates, and photosynthetic organisms. EM alveoli ciliate paramecium putrinum

Almost all sequenced mitochondrial genomes of ciliates and apicomplexa are linear. The mitochondria almost all carry

mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondrion, mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mit ...

of their own but with greatly reduced genome sizes. Exceptions are '' Cryptosporidium'' which are left with only a

mitosome A mitosome is an organelle found in some unicellular eukaryotic organisms, like in members of the supergroup Excavata. The mitosome was found and named in 1999, and its function has not yet been well characterized. It was termed a ''crypton'' by ...

; ciliates; and Janouškovec et al 2013 demonstrated that '' Acavomonas'' diverged early and thus have retained some gene-encoding mtDNA. The mitochondrial genome of '' Babesia microti'' is circular. This species is also now known not to belong to either of the genera '' Babesia'' or ''

Theileria ''Theileria'' is a genus of parasites that belongs to the phylum Apicomplexa, and is closely related to '' Plasmodium''. Two ''Theileria'' species, ''T. annulata'' and ''T. parva'', are important cattle parasites. ''T. annulata'' causes tropical ...

'' and a new genus will have to be created for it.

History

The relationship of apicomplexa, dinoflagellates and ciliates had been suggested during the 1980s, and this was confirmed in the early 1990s by comparisons of ribosomal RNA sequences, most notably by Gajadhar ''et al''. Cavalier-Smith introduced the formal name Alveolata in 1991, although at the time he was unconvinced that the group was monophyletic, and considered the grouping to be a

paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

assemblage. Many biologists prefer the use of the colloquial name 'alveolate'.

Classification

Alveolata include around nine major and minor groups. They are diverse in form, and are known to be related by various ultrastructural and genetic similarities: * Ciliates – very common protozoa with many short cilia arranged in rows, and two nuclei * Acavomonidia * Colponemidia * Dinoflagellates s.l. – mostly marine flagellates many of which have

chloroplast A chloroplast () is a type of membrane-bound organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. The photosynthetic pigment chlorophyll captures the energy from sunlight, converts it, and stores it ...

s * Perkinsozoa *

Chromerida Chromerida is a phylum of unicellular alveolates, which includes photosynthetic species '' Chromera velia'' and '' Vitrella brassicaformis''. General features of the phylum include spherical cells each with a thick cell wall, chloroplast present ...

– a marine phylum of photosynthetic protozoa *

Colpodellida Colpodellida is an order of alveolate The alveolates (meaning "pitted like a honeycomb") are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among ...

* Voromonadida * Apicomplexa – parasitic and secondary non-photosynthetic protozoa that lack axonemal locomotive structures except in

gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce ...

s The Acavomonidia and Colponemidia were previously grouped together as colponemids, a taxon now split because each has a distinctive organization or

ultrastructural identity Ultrastructural identity is a concept in biology. It asserts that evolutionary lineages of eukaryotes in general and protists in particular can be distinguished by complements and arrangements of cellular organelles. These ultrastructural component ...

. The Acavomonidia are closer to the dinoflagellate/perkinsid group than the Colponemidia are. As such, the informal term "colponemids", as it stands currently, covers two non-sister groups within Alveolata: the Acavomonidia and the Colponemidia. The Apicomplexa and dinoflagellates may be more closely related to each other than to the ciliates. Both have plastids, and most share a bundle or cone of microtubules at the top of the cell. In apicomplexans this forms part of a complex used to enter host cells, while in some colorless dinoflagellates it forms a peduncle used to ingest prey. Various other genera are closely related to these two groups, mostly flagellates with a similar apical structure. These include free-living members in '' Oxyrrhis'' and ''

Colponema ''Colponema'' is a genus of single-celled flagellates that feed on eukaryotes in aquatic environments and soil. The genus contains 6 known species and has not been thoroughly studied. ''Colponema'' has two flagella which originate just below the ...

'', and parasites in '' Perkinsus'', '' Parvilucifera'', '' Rastrimonas'' and the ellobiopsids. In 2001, direct amplification of the rRNA gene in marine picoplankton samples revealed the presence of two novel alveolate linages, called group I and II. Group I has no cultivated relatives, while group II is related to the dinoflagellate parasite ''

Amoebophrya ''Amoebophyra'' (or ''Amoebophrya'') is a genus of dinoflagellates. ''Amoebophyra'' is a syndinian parasite that infects free-living dinoflagellates that are attributed to a single species by using several host-specific parasites. It acts as ...

'', which was classified until now in the Syndiniales dinoflagellate order. Some studies suggested the haplosporids, mostly parasites of marine invertebrates, might belong here, but they lack alveoli and are now placed among the Cercozoa. The ellobiopsids are of uncertain relation within the alveolates. Silberman et al 2004 establish that the '' Thalassomyces'' genus of ellobiopsids are alveolates using

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

analysis, however no more certainty exists on their place.

Phylogeny

Based on a compilation of the following works. Recent research indicates that the Dinoflagelllata actually emerged in the Perkinsea. Furthermore, the Perkinsea emerged in the Apicomplexa.

Taxonomy

Alveolata Cavalier-Smith 1991 lveolatobiontes* Phylum Ciliophora Doflein 1901 stat. n. Copeland 1956 iliata Perty 1852; Infusoria Bütschli 1887; Ciliae, Ciliozoa, Cytoidea, Eozoa, Heterocaryota, Heterokaryota** Subphylum

Postciliodesmatophora Postciliodesmatophora is a subphylum of ciliates. Members of this subphylum share stacks of postciliary microtubular ribbons associated with somatic kinetosome A basal body (synonymous with basal granule, kinetosome, and in older cytologic ...

Gerassimova & Seravin 1976 *** Class Heterotrichea Stein 1859 *** Class Karyorelictea Corliss 1974 ** Subphylum Intramacronucleata Lynn 1996 *** Class ? Mesodiniea Chen et al. 2015 *** Infraphylum Lamellicorticata **** Class Litostomatea Small & Lynn 1981 **** Class Armophorea Lynn 2004 **** Class Cariacotrichea Orsi et al. 2011 **** Class Spirotrichea Bütschli 1889 **** Infraphylum Ventrata Cavalier-Smith 2004 onthreep Lynn 2012**** Order ? Discotrichida Chen et al. 2015 **** Class Protocruziea Chen et al. 2015 rotocruziidia de Puytorac, Grain & Mignot 1987**** Class Colpodea Small & Lynn 1981 **** Class Nassophorea Small & Lynn 1981 **** Class Phyllopharyngea de Puytorac et al. 1974 **** Class Prostomatea Schewiakoff 1896 **** Class Plagiopylea Small & Lynn 1985 sensu Lynn 2008 **** Class Oligohymenophorea de Puytorac et al. 1974 * Phylum Miozoa Cavalier-Smith 1987 ** Subphylum Colponemidia Tikhonenkov, Mylnikov & Keeling 2013 *** Class Colponemea Cavalier-Smith 1993 ** Subphylum Acavomonadia Tikhonenkov et al. 2014 *** Class Acavomonadea Tikhonenkov et al. 2014 ** Subphylum Myzozoa Cavalier-Smith 2004 *** Infraphylum Apicomplexa Levine 1970 emend. Adl et al. 2005 **** Order ? Vitrellida Cavalier-Smith 2017 **** Class ? Myzomonadea Cavalier-Smith & Chao 2004 sensu Ruggiero et al. 2015 **** Class Chromerea **** Order

Patterson & Zölffel 1991 piromonadida Krylov & Mylnikov 1986**** Superclass Sporozoa Leuckart 1879 stat. nov. Cavalier-Smith 2013 amontozoa***** Class Blastogregarinida Chatton & Villeneuve 1936 lastogregarinina; Blastogregarinorina Chatton & Villeneuve 1936***** Class

Paragregarea The gregarines are a group of Apicomplexan alveolates, classified as the Gregarinasina or Gregarinia. The large (roughly half a millimeter) parasites inhabit the intestines of many invertebrates. They are not found in any vertebrates. However, gr ...

Cavalier-Smith 2014 ***** Class

Gregarinomorphea The gregarines are a group of Apicomplexan alveolates, classified as the Gregarinasina or Gregarinia. The large (roughly half a millimeter) parasites inhabit the intestines of many invertebrates. They are not found in any vertebrates. However, gr ...

Grassé 1953 ***** Class

Coccidiomorphea The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. Th ...

Doflein 1901 *** Infraphylum Dinozoa Cavalier-Smith 1981 emend. 2003 **** Order ? Acrocoelida Cavalier-Smith & Chao 2004 **** Order ? Rastromonadida Cavalier-Smith & Chao 2004 **** Class Squirmidea Norén 1999 stat. nov. Cavalier-Smith 2014 **** Superclass Perkinsozoa Norén et al. 1999 s.s. ***** Class Perkinsea Levine 1978 erkinsasida Levine 1978**** Superclass Dinoflagellata Butschli 1885 stat. nov. Cavalier-Smith 1999 sensu Cavalier-Smith 2013 inozoa Cavalier-Smith 1981***** Class Pronoctilucea ***** Class Ellobiopsea Cavalier-Smith 1993 llobiophyceae Loeblich III 1970; Ellobiopsida Whisler 1990***** Class Myzodinea Cavalier-Smith 2017 ***** Class Oxyrrhea Cavalier-Smith 1987 ***** Class Syndinea Chatton 1920 s.l. yndiniophyceae Loeblich III 1970 s.s.; Syndina Cavalier-Smith***** Class Endodinea Cavalier-Smith 2017 ***** Class Noctiluciphyceae Fensome et al. 1993 octilucae Haeckel 1866; Noctilucea Haeckel 1866 stat. nov.; Cystoflagellata Haeckel 1873 stat. nov. Butschli 1887***** Class Dinophyceae Pascher 1914 eridinea Ehrenberg 1830 stat. nov. Wettstein

Development

The development of plastids among the alveolates is intriguing. Cavalier-Smith proposed the alveolates developed from a chloroplast-containing ancestor, which also gave rise to the Chromista (the chromalveolate hypothesis). Other researchers have speculated that the alveolates originally lacked plastids and possibly the dinoflagellates and Apicomplexa acquired them separately. However, it now appears that the alveolates, the dinoflagellates, the Chromerida and the heterokont algae acquired their plastids from a red alga with evidence of a common origin of this organelle in all these four clades.

Evolution

A Bayesian estimate places the evolution of the alveolate group at ~. The Alveolata consist of Myzozoa, Ciliates, and Colponemids. In other words, the term Myzozoa, meaning "to siphon the contents from prey", may be applied informally to the common ancestor of the subset of alveolates that are neither ciliates nor colponemids. Predation upon algae is an important driver in alveolate evolution, as it can provide sources for endosymbiosis of novel plastids. The term Myzozoa is therefore a handy concept for tracking the history of the alveolate phylum. The ancestors of the alveolate group may have been photosynthetic. The ancestral alveolate probably possessed a plastid. Chromerids, apicomplexans, and peridinin dinoflagellates have retained this

organelle In cell biology, an organelle is a specialized subunit, usually within a cell, that has a specific function. The name ''organelle'' comes from the idea that these structures are parts of cells, as organs are to the body, hence ''organelle,'' t ...

. Going one step even further back, the chromerids, the peridinin dinoflagellates and the heterokont algae have been argued to possess a monophyletic plastid lineage in common, i.e. acquired their plastids from a red alga, and so it seems likely that the common ancestor of alveolates and heterokonts was also photosynthetic. In one school of thought the common ancestor of the dinoflagellates, apicomplexans, '' Colpodella'',

, and '' Voromonas'' was a myzocytotic predator with two heterodynamic flagella, micropores, trichocysts,

rhoptries A rhoptry is a specialized secretory organelle. They are club-shaped organelles connected by thin necks to the extreme apical pole of the parasite. These organelles, like micronemes, are characteristic of the motile stages of Apicomplexa protoz ...

, micronemes, a polar ring and a coiled open sided conoid. While the common ancestor of alveolates may also have possessed some of these characteristics, it has been argued that Myzocytosis was not one of these characteristics, as ciliates ingest prey by a different mechanism. An ongoing debate concerns the number of membranes surrounding the plastid across apicomplexans and certain dinoflagellates, and the origin of these membranes. This ultrastructural character can be used to group organisms and if the character is in common, it can imply that phyla had a common photosynthetic ancestor. On the basis that apicomplexans possess a plastid surrounded by four membranes, and that peridinin dinoflagellates possess a plastid surrounded by three membranes, Petersen et al. have been unable to rule out that the shared stramenopile-alveolate plastid could have been recycled multiple times in the alveolate phylum, the source being stramenopile-alveolate donors, through the mechanism of ingestion and

endosymbiosis An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "withi ...

. Ciliates are a model alveolate, having been genetically studied in great depth over the longest period of any alveolate lineage. They are unusual among eukaryotes in that reproduction involves a micronucleus and a macronucleus. Their reproduction is easily studied in the lab, and made them a model eukaryote historically. Being entirely predatory and lacking any remnant plastid, their development as a phylum illustrates how predation and autotrophy are in dynamic balance and that the balance can swing one way or other at the point of origin of a new phylum from mixotrophic ancestors, causing one ability to be lost. File:Paramecium caudatum Ehrenberg, 1833.jpg, '' Paramecium caudatum'' ( Ciliophora) File:Mikrofoto.de-Glockentierchen-1.jpg, '' Vorticella'' ( Ciliophora) (left) File:Plasmodium.jpg, '' Plasmodium falciparum'' ( Apicomplexa) in blood File:Emaxima oocysts usda.jpg, '' Eimeria maxima'' ( Apicomplexa) File:Dinophysis acuminata.jpg, '' Dinophysis acuminata'' ( Dinoflagellata)

References

External links

Tree of Life: Alveolates
{{Taxonbar, from=Q448746 Infrakingdoms Taxa named by Thomas Cavalier-Smith