Chromalveolata | |
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Clockwise from top-left: a haptophyte (coccolithophore: Emiliania huxleyi), some diatoms, a water mold, a cryptomonad, and Macrocystis , a phaeophyte | |
Scientific classification | |
Domain: | Eukaryota |
(unranked): | Bikonta |
(unranked): | Chromalveolata Adl et al., 2005 (not a monophyletic group) [1] [2] |
Phyla | |
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. [3] It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1981. Chromalveolata was proposed to represent the organisms descended from a single secondary endosymbiosis involving a red alga and a bikont. [4] The plastids in these organisms are those that contain chlorophyll c.
However, the monophyly of the Chromalveolata has been rejected. Thus, two papers published in 2008 have phylogenetic trees in which the chromalveolates are split up, [5] [6] and recent studies continue to support this view. [7] [8]
Historically, many chromalveolates were considered plants, because of their cell walls, photosynthetic ability, and in some cases their morphological resemblance to the land plants (Embryophyta). However, when the five-kingdom system (proposed in 1969) took prevalence over the animal–plant dichotomy, most of what we now call chromalveolates were put into the kingdom Protista, but the water molds and slime nets were put into the kingdom Fungi, while the brown algae stayed in the plant kingdom. These various organisms were later grouped together and given the name Chromalveolata by Cavalier-Smith. He believed them to be a monophyletic group, but this is not the case. [9]
In 2005, in a classification reflecting the consensus at the time, the Chromalveolata were regarded as one of the six major clades of eukaryotes. [3] Although not given a formal taxonomic status in this classification, elsewhere the group had been treated as a Kingdom.[ citation needed ] The Chromalveolata were divided into four major subgroups:
Other groups that may be included within, or related to, chromalveolates, are:
Though several groups, such as the ciliates and the water molds, have lost the ability to photosynthesize, most are autotrophic. All photosynthetic chromalveolates use chlorophylls a and c, and many use accessory pigments. Chromalveolates share similar glyceraldehyde 3-phosphate dehydrogenase proteins. [11]
However, as early as 2005, doubts were being expressed as to whether Chromalveolata was monophyletic, [9] and a review in 2006 noted the lack of evidence for several of the supposed six major eukaryote groups, including the Chromalveolata. [12] In 2012, consensus emerged that the group is not monophyletic. The four original subgroups fall into at least two categories: one comprises the Stramenopiles and the Alveolata, to which the Rhizaria are now usually added to form the SAR group; the other comprises the Cryptophyta and the Haptophyta. [5] [6] A 2010 paper splits the Cryptophyta and Haptophyta; the former are a sister group to the SAR group, the latter cluster with the Archaeplastida (plants in the broad sense). The katablepharids are closely related to the cryptophytes and the telonemids and centrohelids may be related to the haptophytes. [7]
A variety of names have been used for different combinations of the groups formerly thought to make up the Chromalveolata.
Chromalveolates, unlike other groups with multicellular representatives, do not have very many common morphological characteristics. Each major subgroup has certain unique features, including the alveoli of the Alveolata, the haptonema of the Haptophyta, the ejectisome of the Cryptophyta, and the two different flagella of the Heterokontophyta. However, none of these features are present in all of the groups.
The only common chromalveolate features are these:
Since this is such a diverse group, it is difficult to summarize shared chromalveolate characteristics.
Many chromalveolates affect our ecosystem in enormous ways.
Some of these organisms can be very harmful. Dinoflagellates produce red tides, which can devastate fish populations and intoxicate oyster harvests. Apicomplexans are some of the most successful specific parasites to animals (including the genus Plasmodium , the malaria parasites). Water molds cause several plant diseases - it was the water mold Phytophthora infestans that caused the Irish potato blight that led to the Great Irish Famine.
However, many others are vital members of our ecosystem. Diatoms are one of the major photosynthetic producers, and as such produce much of the oxygen that we breathe, and also take in much of the carbon dioxide from the atmosphere. Brown algae, most specifically kelps, create underwater "forest" habitats for many marine creatures, and provide a large portion of the diet of coastal communities.
Chromalveolates also provide many products that we use. The algin in brown algae is used as a food thickener, most famously in ice cream. The siliceous shells of diatoms have many uses, such as in reflective paint, in toothpaste, or as a filter, in what is known as diatomaceous earth.
Like other organisms, chromalveolata have viruses. In the case of Emiliania huxleyi (a common algal bloom chromalveolate), a virus believed to be specific to it causes mass death and the end of the bloom. [15]
The centrohelids or centroheliozoa are a large group of heliozoan protists. They include both mobile and sessile forms, found in freshwater and marine environments, especially at some depth.
Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.
Thomas (Tom) Cavalier-Smith, FRS, FRSC, NERC Professorial Fellow, was a professor of evolutionary biology in the Department of Zoology, at the University of Oxford.
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-photosynthetic, 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.
A bikont is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled and multi-celled organisms are members of the group, and these, as well as the presumed ancestor, have two flagella.
The Archaeplastida are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans. The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by phagocytosis of a cyanobacterium. All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae. Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.
In the classification of eukaryotes, Cabozoa was a taxon proposed by Cavalier-Smith. It was a putative clade comprising the Rhizaria and Excavata. More recent research places the Rhizaria with the Alveolata and Stramenopiles instead of the Excavata, however, so "Cabozoa" is polyphyletic.
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.
SAR or Harosa is a highly diverse clade of eukaryotes, often considered a supergroup, that includes stramenopiles (heterokonts), alveolates, and rhizarians. It is a node-based taxon, including all descendants of the three groups' last common ancestor, and comprises most of the now-rejected Chromalveolata. Their sister group has been found to be telonemids, with which they make up the TSAR clade.
The cryptomonads-haptophytes assemblage is a proposed but disputed monophyletic grouping of unicellular eukaryotes that are not included in the SAR supergroup. Several alternative names have been used for the group, including Hacrobia ; CCTH ; and "Eukaryomonadae".
The kathablepharids or katablepharids are a group of heterotrophic flagellates closely related to cryptomonads. First described by Heinrich Leonhards Skuja in 1939, kathablepharids were named after the genus Kathablepharis. This genus is corrected to Katablepharis under botanical nomenclature, but the original spelling is maintained under zoological nomenclature. They are single-celled protists with two anteriorly directed flagella, an anterior cytostome for ingesting eukaryotic prey, and a sheath that covers the cell membrane. They have extrusomes known as ejectisomes, as well as tubular mitochondrial cristae.
Telonema is a genus of single-celled organisms.
Diaphoretickes is a major group of eukaryotic organisms, with over 400,000 species. The majority of the earth's biomass that carries out photosynthesis belongs to Diaphoretickes.
Plants+HC clade is a group of eukaryotes proposed by Burki et al. (2008).
Halvaria is a taxonomic grouping of protists that includes Alveolata and Stramenopiles (Heterokonta).
Cryptista is a clade of alga-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes.
Haptista is a proposed group of protists made up of centrohelids and haptophytes. Phylogenomic studies indicate that Haptista, together with Ancoracysta twista, forms a sister clade to the SAR+Telonemia supergroup, but it may also be sister to the Cryptista (+Archaeplastida). It is thus one of the earliest diverging Diaphoretickes.
Endohelea is a proposed clade of eukaryotes that are related to Archaeplastida and the SAR supergroup. They used to be considered heliozoans, but phylogenetically they belong to a group of microorganisms known as Cryptista.
The cortical alveolum is a cellular organelle consisting of a vesicle located under the cytoplasmic membrane, to which they give support. The term "corticate" comes from an evolutionary hypothesis about the common origin of kingdoms Plantae and Chromista, because both kingdoms have cortical alveoli in at least one phylum. At least three protist lineages exhibit these structures: Telonemia, Alveolata and Glaucophyta.
A supergroup, in evolutionary biology, is a large group of organisms that share one common ancestor and have important defining characteristics. It is an informal, mostly arbitrary rank in biological taxonomy that is often greater than phylum or kingdom, although some supergroups are also treated as phyla.
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