The initial version of a classification system of life by British zoologist
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 ...
appeared in 1978. This initial system continued to be modified in subsequent versions that were published until he died in 2021. As with classifications of others, such as
Carl Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
,
Ernst Haeckel
Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, natural history, naturalist, eugenics, eugenicist, Philosophy, philosopher, physician, professor, marine biology, marine biologist and artist ...
,
Robert Whittaker, and
Carl Woese
Carl Richard Woese ( ; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal ...
, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy., Cavalier-Smith used his classifications to convey his opinions about the evolutionary relationships among various organisms, principally microbial. His classifications complemented his ideas communicated in scientific publications, talks, and diagrams. Different iterations might have a wider or narrow scope, include different groupings, provide greater or lesser detail, and place groups in different arrangements as his thinking changed. His classifications has been a major influence in the modern taxonomy, particularly of protists.
Cavalier-Smith has published extensively on the
classification
Classification is the activity of assigning objects to some pre-existing classes or categories. This is distinct from the task of establishing the classes themselves (for example through cluster analysis). Examples include diagnostic tests, identif ...
of
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 ...
. One of his major contributions to biology was his proposal of a new
kingdom of life: the
Chromista
Chromista is a proposed but polyphyletic obsolete Biology, biological Kingdom (biology), kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their Photosynthesi ...
, although the usefulness of the grouping is questionable given that it is generally agreed to be an arbitrary (polyphyletic) grouping of taxa. He also proposed that all chromista and
alveolata share the same common ancestor, a claim later refuted by studies of morphological and molecular evidence by other labs. He named this new group the
Chromalveolates. He also proposed and named many other high-rank taxa, like
Opisthokonta (1987),
Rhizaria
The Rhizaria are a diverse and species-rich clade 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 ...
(2002), and
Excavata
Excavata is an obsolete, extensive and diverse 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 ...
(2002), though he himself consistently does not include Opisthonkonta as a formal taxon in his schemes. Together with Chromalveolata,
Amoebozoa
Amoebozoa is a major Taxonomy (biology), taxonomic group containing about 2,400 described species of Amoeba, amoeboid protists, often possessing blunt, fingerlike, Pseudopod#Morphology, lobose pseudopods and tubular mitochondrial cristae. In trad ...
(he amended their description in 1998), and
Archaeplastida
The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced ) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It als ...
(which he called Plantae since 1981) the six formed the basis of the taxonomy of
eukaryote
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 ...
s in the middle 2000s. He has also published prodigiously on issues such as the origin of various
cellular organelles
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,'' th ...
(including the
nucleus,
mitochondria
A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
),
genome size
Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms (trillionths or 10−12 of a gram, abbreviated pg) or less frequently in daltons, or as the tot ...
evolution
Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
, and
endosymbiosis
An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
. Though fairly well known, many of his claims have been controversial and have not gained widespread acceptance in the
scientific community
The scientific community is a diverse network of interacting scientists. It includes many "working group, sub-communities" working on particular scientific fields, and within particular institutions; interdisciplinary and cross-institutional acti ...
to date. Most recently, he has published a paper citing the
paraphyly
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 ...
of his
bacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
l kingdom, the origin of
Neomura Neomura (from Ancient Greek ''neo-'' "new", and Latin ''-murus'' "wall") is a proposed clade of life composed of the two domains Archaea and Eukaryota, coined by Thomas Cavalier-Smith in 2002. Its name reflects the hypothesis that both archaea and ...
from
Actinobacteria and taxonomy of
prokaryote
A prokaryote (; less commonly spelled procaryote) is a unicellular organism, single-celled organism whose cell (biology), cell lacks a cell nucleus, nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Ancient Gree ...
s.
According to
Palaeos Palaeos.com is a web site on biology, paleontology, phylogeny and geology and which covers the history of Earth. The site is well respected and has been used as a reference by professional paleontologists such as Michael J. Benton, the professor of ...
.com:
Prof. Cavalier-Smith of Oxford University has produced a large body of work which is well regarded. Still, he is controversial in a way that is a bit difficult to describe. The issue may be one of writing style. Cavalier-Smith has a tendency to make pronouncements where others would use declarative sentences, to use declarative sentences where others would express an opinion, and to express opinions where angels would fear to tread. In addition, he can sound arrogant, reactionary, and even perverse. On the other and he has a long history of being right when everyone else was wrong. To our way of thinking, all of this is overshadowed by one incomparable virtue: the fact that he ''will'' grapple with the details. This makes for very long, very complex papers and causes all manner of dark murmuring, tearing of hair, and gnashing of teeth among those tasked with trying to explain his views of early life. See, or example Zrzavý (2001) ndPatterson (1999). Nevertheless, he deals with all of the relevant facts.
Eight kingdoms model
The first two kingdoms of life: Plantae and Animalia
The use of the word "
kingdom" to describe a major branch of the living world dates as far back as
Linnaeus
Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...
(1707–1778) who divided the natural world into three kingdoms:
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, ...
,
vegetable
Vegetables are edible parts of plants that are consumed by humans or other animals as food. This original meaning is still commonly used, and is applied to plants collectively to refer to all edible plant matter, including edible flower, flo ...
, and
mineral
In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid substance with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2011): Mi ...
.
[Dan H. Nicolson]
Animal, Vegetable or Mineral?
Proceedings of a Mini-Symposium on Biological Nomenclature in the 21st Century held at the University of Maryland on 4 November 1996. Edited by James L. Reveal The taxa "animal kingdom" (or kingdom
Animalia
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, ...
) and "plant kingdom" (or kingdom
Plantae) remain in use by some modern
evolutionary biologists
Evolutionary biology is the subfield of biology that studies the evolutionary processes such as natural selection, common descent, and speciation that produced the diversity of life on Earth. In the 1930s, the discipline of evolutionary bi ...
. The initial targets of Cavalier-Smith's classification, the protozoa were classified as members of the animal kingdom,
and many algae were regarded as part of the plant kingdom. With growing awareness that the animals and plants embraced unrelated taxa, the use of the two kingdom system was rejected by specialists. It remains in use in less critical circles.
The third kingdom: Protista
250px, The is an animal that resembles a plant.">sea anemone is an animal that resembles a plant.
By mid-nineteenth century,
microscopic organisms were generally classified into four groups:
#Protozoa (primitive animals),
#
Protophyta (primitive plants),
#
Phytozoa (animal-like plants & plant-like animals), and
In 1858,
Richard Owen
Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
(1804–1892) proposed that the animal phylum Protozoa be elevated to the status of kingdom.
In 1860,
John Hogg (1800–1869) proposed that protozoa and protophyta be grouped together into a new kingdom which he called "Regnum Primigenum" (kingdom primitive). According to Hogg, this new classification scheme prevented "the unnecessary trouble of contending about their supposed natures, and of uselessly trying to distinguish the Protozoa from the Protophyta". In 1866,
Ernst Haeckel
Ernst Heinrich Philipp August Haeckel (; ; 16 February 1834 – 9 August 1919) was a German zoologist, natural history, naturalist, eugenics, eugenicist, Philosophy, philosopher, physician, professor, marine biology, marine biologist and artist ...
(1834–1919) proposed the name "
Protista
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 ...
" for the primigenial kingdom and included bacteria in this third kingdom of life.
)
The fourth kingdom: Fungi
By 1959,
Robert Whittaker proposed that fungi, which were formerly classified as plants, be given their own kingdom. Therefore, he divided life into four kingdoms such as:
#Protista, (or unicellular organisms);
#
Plantae, (or multicellular plants);
#Fungi; and
#Animalia (or
multicellular
A multicellular organism is an organism that consists of more than one cell (biology), cell, unlike unicellular organisms. All species of animals, Embryophyte, land plants and most fungi are multicellular, as are many algae, whereas a few organism ...
animals).
Whittaker subdivided the Protista into two subkingdoms:
#
Monera (
bacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of Prokaryote, prokaryotic microorganisms. Typically a few micr ...
) and
# Eunucleata (single celled
eukaryotes
The eukaryotes ( ) constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, seaweeds, and many unicellular organisms are eukaryotes. They constitute a major group of ...
).
The fifth kingdom: Bacteria (Monera)
Bacteria are fundamentally different from the
eukaryote
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 ...
s (plants, animals, fungi,
amebas, protozoa, and
chromista
Chromista is a proposed but polyphyletic obsolete Biology, biological Kingdom (biology), kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their Photosynthesi ...
). Eukaryotes have
cell nuclei
The cell nucleus (; : nuclei) is a membrane-bound organelle found in eukaryotic cells. Eukaryotic cells usually have a single nucleus, but a few cell types, such as mammalian red blood cells, have no nuclei, and a few others including osteoc ...
, bacteria do not. In 1969, Whittaker elevated the bacteria to the status of kingdom. His new classification system divided the living world into five kingdoms:
#Plantae,
#Animalia,
#Protista (Eunucleata),
#Fungi, and
#Monera (the kingdom bacteria).
The sixth kingdom: Archaebacteria
300px, Phylogenetic tree based on Woese ''et al.'' rRNA analysis in 1990
The kingdom Monera can be divided into two distinct groups:
eubacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the ...
(true bacteria) and
archaebacteria
Archaea ( ) is a domain of organisms. Traditionally, Archaea only included its prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even though the domain Archaea cladis ...
(
archaea
Archaea ( ) is a Domain (biology), domain of organisms. Traditionally, Archaea only included its Prokaryote, prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even thou ...
). In 1977
Carl Woese
Carl Richard Woese ( ; July 15, 1928 – December 30, 2012) was an American microbiologist and biophysicist. Woese is famous for defining the Archaea (a new domain of life) in 1977 through a pioneering phylogenetic taxonomy of 16S ribosomal ...
and
George E. Fox established that archaebacteria (methanogens in their case) were genetically different (based on their ribosomal RNA genes) from bacteria so that life could be divided into three principle lineages, namely:
#Eubacteria (all typical bacteria),
#Archaebacteria (methanogens), and
#Urkaryotes (all eukaryotes).
In 1990, Woese introduced domain above kingdom by creating
three-domain system
The three-domain system is a taxonomic classification system that groups all cellular life into three domains, namely Archaea, Bacteria and Eukarya, introduced by Carl Woese, Otto Kandler and Mark Wheelis in 1990. The key difference from ea ...
such as:
#Bacteria,
#Archaea, and
#Eucarya.
But Cavalier-Smith considered Archaebacteria as a kingdom.
The seventh kingdom: Chromista
right , 350px, The are a member of the kingdom Chromista. ">brown algae are a member of the kingdom Chromista.
By 1981, Cavalier-Smith had divided all the eukaryotes into nine kingdoms.
In it, he created Chromista for a separate kingdom of some protists.
Most chromists are
photosynthetic
Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
. This distinguishes them from most other protists which lack photosynthesis. In both plants and chromists photosynthesis takes place in
chloroplasts
A chloroplast () is a type of membrane-bound organelle, organelle known as a plastid that conducts photosynthesis mostly in plant cell, plant and algae, algal cells. Chloroplasts have a high concentration of chlorophyll pigments which captur ...
. In plants, however, the chloroplasts are located in the
cytosol
The cytosol, also known as cytoplasmic matrix or groundplasm, is one of the liquids found inside cells ( intracellular fluid (ICF)). It is separated into compartments by membranes. For example, the mitochondrial matrix separates the mitochondri ...
while in chromists the chloroplasts are located in the
lumen of their
rough endoplasmic reticulum
The endoplasmic reticulum (ER) is a part of a transportation system of the eukaryotic cell, and has many other important functions such as protein folding. The word endoplasmic means "within the cytoplasm", and reticulum is Latin for "little n ...
. This distinguishes chromists from plants.
Based on the addition of Chromista as a kingdom, he suggested that even with his nine kingdoms of eukaryotes, "the best one for general scientific use is a system of seven kingdoms",
[ which includes:
# Plantae,
#]Animalia
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, ...
,
#Protozoa
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 ...
,
#Chromista
Chromista is a proposed but polyphyletic obsolete Biology, biological Kingdom (biology), kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their Photosynthesi ...
,
#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 ...
,
#Eubacteria
Bacteria (; : bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were among the ...
, and
#Archaebacteria
Archaea ( ) is a domain of organisms. Traditionally, Archaea only included its prokaryotic members, but this has since been found to be paraphyletic, as eukaryotes are known to have evolved from archaea. Even though the domain Archaea cladis ...
.
The eighth kingdom: Archezoa
In 1983, Cavalier-Smith adapted the term Archezoa, previously used by Haeckel, Perty, and for protists that lack mitochondria
A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is us ...
. Initially, the taxon included the quite unrelated metamonads and microscopridia, and expanded to include other unrelated taxa such that the term referred to many different groupings of protists. He originally considered it as a subkingdom, but by 1989, with the establishment of Chromista as separate kingdom, he treated it as a kingdom.[
Archezoa is now defunct. He assigned former members of the kingdom Archezoa to the phylum Amoebozoa. Today, its former members belong to the Metamonada (e. g. ''Trimastix''), fungi (Microsporidia), Amoebozoa (e. g. '' Pelomyxa'').]
Kingdom Protozoa ''sensu'' Cavalier-Smith
Cavalier-Smith referred to what remained of the protist kingdom, after he removed the kingdoms Archezoa and Chromista, as the "kingdom Protozoa". In 1993, this kingdom contained 18 phyla as summarized in the following table. The first major division relied on the basis of presence or absence of dictyosomes, although a Golgi Apparatus was subsequently demonstrated in the 'Adictyozoa'.
The phylum Opalozoa was established by Cavalier-Smith in 1991.
Six kingdoms models
By 1998, Cavalier-Smith had reduced the total number of kingdoms from eight to six: Animalia, Protozoa, Fungi, Plantae (including red and green algae
The green algae (: green alga) are a group of chlorophyll-containing autotrophic eukaryotes consisting of the phylum Prasinodermophyta and its unnamed sister group that contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ...
), Chromista, and Bacteria.
Five of Cavalier-Smith's kingdoms are classified as eukaryotes
The eukaryotes ( ) constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, seaweeds, and many unicellular organisms are eukaryotes. They constitute a major group of ...
as shown in the following scheme:
*Eubacteria
*Neomura Neomura (from Ancient Greek ''neo-'' "new", and Latin ''-murus'' "wall") is a proposed clade of life composed of the two domains Archaea and Eukaryota, coined by Thomas Cavalier-Smith in 2002. Its name reflects the hypothesis that both archaea and ...
**Archaebacteria
**Eukaryotes
***Kingdom Protozoa
*** Unikonts (heterotrophs
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 ...
)
****Kingdom Animalia
****Kingdom Fungi
*** Bikonts (primarily photosynthetic
Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
)
****Kingdom Plantae (including red and green algae)
****Kingdom Chromista
Eukaryotes are divided into two major groups: Unikont and Bikont. Uniciliates are cells with only one flagellum
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 ...
and unikonts are descended from uniciliates. Unikont cells often have only one centriole
In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers ( Pinophyta), flowering plants ( angiosperms) and most fungi, an ...
as well. Biciliate cells have two flagella and bikonts are descended from biciliates. Biciliates undergo ciliary transformation by converting a younger anterior flagellum into a dissimilar older posterior flagellum. Animals and fungi are unikonts while plants and chromists are bikonts. Some protozoa are unikonts while others are bikonts.
The Bacteria (= prokaryotes) are subdivided into Eubacteria and Archaebacteria. According to Cavalier-Smith, Eubacteria is the oldest group of terrestrial organisms still living. He classifies the groups which he believes are younger (archaebacteria and eukaryotes) as Neomura.
The 1998 model
Kingdom Animalia
In 1993, Cavalier-Smith classified Myxozoa
Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. ...
as a protozoan parvkingdom. By 1998, he had reclassified it as an animal subkingdom. Myxozoa contains three phyla, Myxosporidia, Haplosporidia, and Paramyxia, which were reclassified as animals along with Myxozoa. Likewise, Cavalier-Smith reclassified the protozoan phylum Mesozoa
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm (outer layer) of ciliated cells surrounding one or more reproductive cells.
A 2017 study recovered Mesozoa ...
as an animal subkingdom.
In his 1998 scheme, the animal kingdom was divided into four subkingdoms:
* Radiata
Radiata or Radiates is a historical taxonomic rank that was used to classify animals with Symmetry (biology)#Radial symmetry, radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of anim ...
(phyla Porifera
Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a Basal (phylogenetics) , basal clade and a sister taxon of the Eumetazoa , diploblasts. They are sessility (motility) , sessile ...
, Cnidaria
Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
, Placozoa
Placozoa ( ; ) is a phylum of free-living (non-parasitic) marine invertebrates. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by Phagocytosis, engulfment, reproducing by Fission (biol ...
, and Ctenophora
Ctenophora (; : ctenophore ) is a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and they are ...
),
* Myxozoa
Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. ...
,
* Mesozoa
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm (outer layer) of ciliated cells surrounding one or more reproductive cells.
A 2017 study recovered Mesozoa ...
, and
* Bilateria
Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left� ...
(all other animal phyla).
He created five new animal phyla:
* Acanthognatha (rotifers
The rotifers (, from Latin 'wheel' and 'bearing'), sometimes called wheel animals or wheel animalcules, make up a phylum (Rotifera ) of microscopic and near-microscopic pseudocoelomate animals.
They were first described by Rev. John Harris ...
, acanthocephalans, gastrotrichs, and gnathostomulids),
* Brachiozoa ( brachiopods and phoronids),
* Lobopoda (onychophorans
Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (for their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus''), is a phylum of el ...
and tardigrades
Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged Segmentation (biology), segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who calle ...
),
* Kamptozoa (Entoprocta
Entoprocta (), or Kamptozoa , is a phylum (biology), phylum of mostly Sessility (zoology), sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whos ...
and '' Symbion''), and
* Nemathelminthes
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (helm ...
(Nematoda
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitism, parasitic. Parasitic ...
, Nematomorpha
Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms, hairsnakes, or Gordian worms) are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size f ...
, Loricifera
Loricifera (from Latin, ''wikt:lorica, lorica'', corselet (armour) + ''ferre'', to bear) is a phylum of very small to microscopic marine cycloneuralian sediment-dwelling animals with 43 described species and approximately 100 more that hav ...
, Priapulida
Priapulida (priapulid worms, from Gr. πριάπος, ''priāpos'' ' Priapus' + Lat. ''-ul-'', diminutive), sometimes referred to as penis worms, is a phylum of unsegmented marine worms. The name of the phylum relates to the Greek god of fertilit ...
, and Kinorhyncha
Kinorhyncha (, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are commonly called mud dragons. Modern species are or less, but Cambrian forms could reac ...
)
and recognized a total of 23 animal phyla, as shown here:
*Kingdom Animaia
**Subkingdom Radiata
Radiata or Radiates is a historical taxonomic rank that was used to classify animals with Symmetry (biology)#Radial symmetry, radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of anim ...
***Infrakingdom Spongiaria
****Phylum Porifera
Sponges or sea sponges are primarily marine invertebrates of the animal phylum Porifera (; meaning 'pore bearer'), a Basal (phylogenetics) , basal clade and a sister taxon of the Eumetazoa , diploblasts. They are sessility (motility) , sessile ...
***Infrakingdom Coelenterata
Coelenterata is a term encompassing the animal phyla Cnidaria (corals, true jellies, sea anemones, sea pens, and their relatives) and Ctenophora (comb jellies). The name comes , referring to the hollow body cavity common to these two phyla. Th ...
****Phylum Cnidaria
Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
****Phylum Ctenophora
Ctenophora (; : ctenophore ) is a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and they are ...
***Infrakingdom Placozoa
Placozoa ( ; ) is a phylum of free-living (non-parasitic) marine invertebrates. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by Phagocytosis, engulfment, reproducing by Fission (biol ...
****Phylum Placozoa
Placozoa ( ; ) is a phylum of free-living (non-parasitic) marine invertebrates. They are blob-like animals composed of aggregations of cells. Moving in water by ciliary motion, eating food by Phagocytosis, engulfment, reproducing by Fission (biol ...
**Subkingdom Myxozoa
Myxozoa (etymology: Greek: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic cnidarian animals – all obligate parasites. It contains the smallest animals ever known to have lived. ...
***Phylum Myxosporidia
**Subkingdom Bilateria
Bilateria () is a large clade of animals characterised by bilateral symmetry during embryonic development. This means their body plans are laid around a longitudinal axis with a front (or "head") and a rear (or "tail") end, as well as a left� ...
***Branch Protostomia
Protostomia () is the clade of animals once thought to be characterized by the formation of the organism's mouth before its anus during embryonic development. This nature has since been discovered to be extremely variable among Protostomia's memb ...
****Infrakingdom Lophozoa
*****Superphylum Polyzoa
******Phylum Bryozoa
Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic animal, aquatic invertebrate animals, nearly all living in sedentary Colony (biology), colonies. Typically about long, they have a spe ...
******Phylum Kamptozoa (Entoprocta
Entoprocta (), or Kamptozoa , is a phylum (biology), phylum of mostly Sessility (zoology), sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whos ...
and Cycliophora)
*****Superphylum Conchozoa
******Phylum Mollusca
Mollusca is a phylum of protostome, protostomic invertebrate animals, whose members are known as molluscs or mollusks (). Around 76,000 extant taxon, extant species of molluscs are recognized, making it the second-largest animal phylum ...
******Phylum Brachiozoa ''sensu lato'' ( Brachiozoa and Phoronida)
*****Superphylum Sipuncula
The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a class containing about 162 species of marine annelid worms, that have secondarily lost their segmentation. Sipuncula was once considered a phylum of unsegmented ...
******Phylum Sipuncula
The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a class containing about 162 species of marine annelid worms, that have secondarily lost their segmentation. Sipuncula was once considered a phylum of unsegmented ...
*****Superphylum Vermizoa
******Phylum Nemertina
******Phylum Annelida
The annelids (), also known as the segmented worms, are animals that comprise the phylum Annelida (; ). The phylum contains over 22,000 extant species, including ragworms, earthworms, and leeches. The species exist in and have adapted to variou ...
****Infrakingdom Chaetognathi
*****Phylum Chaetognatha
The Chaetognatha or chaetognaths (meaning ''bristle-jaws'') are a phylum of predatory marine worms that are a major component of plankton worldwide. Commonly known as arrow worms, they are mostly pelagic; however about 20% of the known species ...
****Infrakingdom Ecdysozoa
Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerates (including arachnids), crustaceans, and myriapods), Nematoda, and several smaller phylum (biology), phyla. The grouping of these animal phyla into a single ...
*****Superphylum Nemathelminthes
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (helm ...
******Phylum Nemathelminthes
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (helm ...
(Nematoda
The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitism, parasitic. Parasitic ...
and Nematomorpha
Nematomorpha (sometimes called Gordiacea, and commonly known as horsehair worms, hairsnakes, or Gordian worms) are a phylum of parasitoid animals superficially similar to nematode worms in morphology, hence the name. Most species range in size f ...
; Priapozoa, Kinorhyncha
Kinorhyncha (, ' "snout") is a phylum of small marine invertebrates that are widespread in mud or sand at all depths as part of the meiobenthos. They are commonly called mud dragons. Modern species are or less, but Cambrian forms could reac ...
and Loricifera
Loricifera (from Latin, ''wikt:lorica, lorica'', corselet (armour) + ''ferre'', to bear) is a phylum of very small to microscopic marine cycloneuralian sediment-dwelling animals with 43 described species and approximately 100 more that hav ...
)
*****Superphylum Haemopoda
******Phylum Lobopodia
Lobopodians are members of the informal group Lobopodia (), or the formally erected phylum Lobopoda Cavalier-Smith (1998). They are panarthropods with stubby legs called lobopods, a term which may also be used as a common name of this group as ...
(Onychophora
Onychophora (from , , "claws"; and , , "to carry"), commonly known as velvet worms (for their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus''), is a phylum of el ...
and Tardigrada
Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged Segmentation (biology), segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who calle ...
)
******Phylum Arthropoda
Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an exoskeleton with a cuticle made of chitin, often mineralised with calcium carbonate, a body with differentiated ( metameric) segments, and paired jointed appendages. ...
****Infrakingdom Platyzoa
*****Phylum Platyhelminthes
Platyhelminthes (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") is a phylum of relatively simple bilaterian, unsegmented, soft-bodied invertebrates commonly called f ...
(incl. Xenacoelomorpha
Xenacoelomorpha () is a small phylum of bilaterian invertebrate animals, consisting of two sister groups: xenoturbellids and acoelomorphs. This new phylum was named in February 2011 and suggested based on morphological synapomorphies (physica ...
)
*****Phylum Acanthognatha (incl. Rotifera, Acanthocephala, Gnathostomulida
Gnathostomulids, or jaw worms, are a small phylum of nearly microscopic marine animals. They inhabit sand and mud beneath shallow coastal waters and can survive in relatively anoxic environments. They were first recognised and described in 1956.
...
, Gastrotricha
The gastrotrichs (phylum Gastrotricha), common name, commonly referred to as hairybellies or hairybacks, are a group of Microscopic scale, microscopic (0.06–3.0 mm), cylindrical, acoelomate animals, and are widely distributed and abundant ...
)
***Branch Deuterostomia
Deuterostomes (from Ancient Greek, Greek: ) are bilaterian animals of the superphylum Deuterostomia (), typically characterized by their anus forming before the mouth during embryogenesis, embryonic development. Deuterostomia comprises three Phyl ...
****Infrakingdom Coelomopora
*****Phylum Echinoderm
An echinoderm () is any animal of the phylum Echinodermata (), which includes starfish, brittle stars, sea urchins, sand dollars and sea cucumbers, as well as the sessile sea lilies or "stone lilies". While bilaterally symmetrical as ...
ata
*****Phylum Hemichordata
****Infrakingdom Chordonia
*****Phylum Urochorda
*****Phylum Chordata
A chordate ( ) is a bilaterian animal belonging to the phylum Chordata ( ). All chordates possess, at some point during their larval or adult stages, five distinctive physical characteristics (Apomorphy and synapomorphy, synapomorphies) th ...
**Subkingdom Mesozoa
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm (outer layer) of ciliated cells surrounding one or more reproductive cells.
A 2017 study recovered Mesozoa ...
***Phylum Mesozoa
The Mesozoa are minuscule, worm-like parasites of marine invertebrates. Generally, these tiny, elusive creatures consist of a somatoderm (outer layer) of ciliated cells surrounding one or more reproductive cells.
A 2017 study recovered Mesozoa ...
Kingdom Protozoa
Under Cavalier-Smith's proposed classification system, protozoa share the following traits:
* they have or are descended from organisms with mitochondria
* they have or are descended from organisms with peroxisomes
A peroxisome () is a membrane-bound organelle, a type of microbody, found in the cytoplasm of virtually all eukaryotic cells. Peroxisomes are oxidative organelles. Frequently, molecular oxygen serves as a co-substrate, from which hydrogen pe ...
* they lack collagen
Collagen () is the main structural protein in the extracellular matrix of the connective tissues of many animals. It is the most abundant protein in mammals, making up 25% to 35% of protein content. Amino acids are bound together to form a trip ...
ous connective tissue
* they lack epiciliary retronemes (rigid thrust-reversing tubular ciliary hairs)
* they lack two additional membranes outside their chloroplast envelope
Organisms that do not meet these criteria were reassigned to other kingdoms by Cavalier-Smith.
The 2003 model
Kingdom Protozoa
In 1993, Cavalier-Smith divided the kingdom Protozoa into two subkingdoms and 18 phyla. By 2003 he used phylogenic evidence to revise the total number of proposed phyla down to 11: Amoebozoa, Choanozoa, Cercozoa, Retaria, Loukozoa, Metamonada, Euglenozoa, Percolozoa, Apusozoa, Alveolata, Ciliophora, and Miozoa.
Unikonts and bikonts
Amoebozoa
Amoebozoa is a major Taxonomy (biology), taxonomic group containing about 2,400 described species of Amoeba, amoeboid protists, often possessing blunt, fingerlike, Pseudopod#Morphology, lobose pseudopods and tubular mitochondrial cristae. In trad ...
do not have flagella and are difficult to classify as unikont or bikont based on morphology
Morphology, from the Greek and meaning "study of shape", may refer to:
Disciplines
*Morphology (archaeology), study of the shapes or forms of artifacts
*Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
. In his 1993 classification scheme, Cavalier-Smith incorrectly classified amoebas as bikonts. Gene fusion research later revealed that the clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
Amoebozoa, was ancestrally uniciliate. In his 2003 classification scheme, Cavalier-Smith reassigned Amoebozoa to the unikont clade along with animals, fungi, and the protozoan phylum Choanozoa. Plants and all other protists were assigned to the clade Bikont by Cavalier-Smith.
Cavalier-Smith's 2003 classification scheme:
* Unikonts
** protozoan phylum Amoebozoa (ancestrally uniciliate)
** opisthokonts
*** uniciliate protozoan phylum Choanozoa
*** kingdom Fungi
*** kingdom Animalia
* Bikonts
** protozoan infrakingdom Rhizaria
The Rhizaria are a diverse and species-rich clade 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 ...
*** phylum Cercozoa
Cercozoa (now synonymised with Filosa) is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead united by phylogeny, molecular phylogenies of rRNA and actin or Ubiqu ...
*** phylum Retaria
Retaria is a clade within the supergroup Rhizaria containing the Foraminifera and the Radiolaria. In 2019, the Retaria were recognized as a basal Rhizaria group, as sister of the Cercozoa
Cercozoa (now synonymised with Filosa) is a phylum of ...
( Radiozoa and Foraminifera
Foraminifera ( ; Latin for "hole bearers"; informally called "forams") are unicellular organism, single-celled organisms, members of a phylum or class (biology), class of Rhizarian protists characterized by streaming granular Ectoplasm (cell bio ...
)
** protozoan infrakingdom Excavata
Excavata is an obsolete, extensive and diverse 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 ...
*** phylum Loukozoa
*** phylum Metamonada
*** phylum Euglenozoa
Euglenozoa are a large group of flagellate Discoba. They include a variety of common free-living species, as well as a few important parasites, some of which infect humans. Euglenozoa are represented by four major groups, ''i.e.,'' Kinetoplastea, ...
*** phylum Percolozoa
** protozoan phylum Apusozoa ( Thecomonadea and Diphylleida
Collodictyonidae (also Diphylleidae) is a group of aquatic, unicellular Eukaryote, eukaryotic organisms with two to four terminal Flagellum, flagella. They feed by phagocytosis, ingesting other unicellular organisms like algae and bacteria.
Rec ...
)
** the chromalveolate clade
*** kingdom Chromista
Chromista is a proposed but polyphyletic obsolete Biology, biological Kingdom (biology), kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their Photosynthesi ...
(Cryptista
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.
Other characteristic features of cryptophyte mtDNAs include large syntenic ...
, Heterokonta, and Haptophyta
The haptophytes, classified either as the Haptophyta, Haptophytina or Prymnesiophyta (named for '' Prymnesium''), are a clade of algae.
The names Haptophyceae or Prymnesiophyceae are sometimes used instead. This ending implies classification at ...
)
*** protozoan infrakingdom Alveolata
**** phylum Ciliophora
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different ...
**** phylum Miozoa ( Protalveolata, Dinozoa, and Apicomplexa
The Apicomplexa (also called Apicomplexia; single: apicomplexan) are organisms of a large phylum of mainly parasitic alveolates. Most possess a unique form of organelle structure that comprises a type of non-photosynthetic plastid called an ap ...
)
** kingdom Plantae ( Viridaeplantae, Rhodophyta
Red algae, or Rhodophyta (, ; ), make up one of the oldest groups of eukaryotic algae. The Rhodophyta comprises one of the largest phyla of algae, containing over 7,000 recognized species within over 900 genera amidst ongoing taxonomic revisions. ...
and Glaucophyta
The glaucophytes, also known as glaucocystophytes or glaucocystids, are a small group of unicellular algae found in freshwater and moist terrestrial environments, less common today than they were during the Proterozoic. The stated number of spec ...
)
Cladogram of life
By September 2003, Cavalier-Smith's tree of life looked like this:
In the above tree, the traditional plant, animal, and fungal kingdoms, as well as Cavalier-Smith's proposed kingdom Chromista, are shown as leaves. The leaves Eubacteria and Archaebacteria together make up the kingdom Bacteria. All remaining leaves together make up the kingdom Protozoa.
By 2006, Cavalier-Smith's microbial tree look like this:
By 2010 new data emerged that showed that Unikonts and Bikonts, originally considered to be separate because of an apparently different organization of cilia and cytoskeleton, are in reality more similar than previously thought. As a consequence, Cavalier-Smith revised the above tree and proposed to move its root to reside in between the Excavata and Euglenozoa kingdoms.
Seven kingdoms model
In 1987, Cavalier-Smith introduced a classification divided into two superkingdoms (Prokaryota and Eukaryota) and seven kingdoms, two prokaryotic kingdoms (Eubacteria and Archaebacteria) and five eukaryotic kingdoms (Protozoa, Chromista, Fungi, Plantae and Animalia).
Cavalier-Smith and his collaborators revised the classification in 2015, and published it in '' PLOS ONE''. In this scheme they reintroduced the classification with the division of prokaryotes superkingdom into two kingdoms, Bacteria (=Eubacteria) and Archaea (=Archaebacteria). This is based on the consensus in the Taxonomic Outline of Bacteria and Archaea (TOBA) and the Catalogue of Life
The Catalogue of Life (CoL) is an online database that provides an index of known species of animals, plants, fungi, and microorganisms. It was created in 2001 as a partnership between the global Species 2000 and the American Integrated Taxono ...
.
References
{{reflist
Taxonomy (biology)
High-level systems of taxonomy