The biological classification system of life introduced 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 ...

involves systematic arrangements of all life forms on earth. Following and improving the classification systems introduced by

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, ...

Ernst Haeckel Ernst Heinrich Philipp August Haeckel (; 16 February 1834 – 9 August 1919) was a German zoologist, naturalist, eugenicist, philosopher, physician, professor, marine biologist and artist. He discovered, described and named thousands of new s ...

Robert Whittaker Robert Whitaker or Whittaker may refer to: *Robert Whittaker (fighter) (born 1990), Australian mixed martial artist *Robert Whitaker (equestrian) (born 1983), British showjumper *Robert Whitaker (author) (active since 1989), American author *Robert ...

, 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 RNA, ...

, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy. His classification has been a major foundation in modern taxonomy, particularly with revisions and reorganisations of kingdoms and phyla. Cavalier-Smith has published extensively on the

classification Classification is a process related to categorization, the process in which ideas and objects are recognized, differentiated and understood. Classification is the grouping of related facts into classes. It may also refer to: Business, organizat ...

protists A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the ex ...

. One of his major contributions to biology was his proposal of a new

kingdom Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...

of life: the

Chromista Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all protists whose plastids contain chlorophyll ''c'', suc ...

, 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

Chromalveolate Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in ...

s. He also proposed and named many other high-rank taxa, like

Opisthokont 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 ...

a (1987),

Rhizaria The Rhizaria are an ill-defined but 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-photosynthethic, but many fora ...

(2002), and

Excavata Excavata is a major supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free ...

(2002), though he himself consistently does not include Opisthonkonta as a formal taxon in his schemes. Together with Chromalveolata,

Amoebozoa Amoebozoa is a major taxonomic group containing about 2,400 described species of amoeboid protists, often possessing blunt, fingerlike, lobose pseudopods and tubular mitochondrial cristae. In traditional and currently no longer supported c ...

(he amended their description in 1998), and

Archaeplastida The Archaeplastida (or kingdom Plantae '' sensu lato'' "in a broad sense"; pronounced /ɑːrkɪ'plastɪdə/) are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor grou ...

(which he called Plantae since 1981) the six formed the basis of the taxonomy of

eukaryote Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bact ...

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 Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom * Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...

, mitochondria),

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 (10−12) of a gram, abbreviated pg) or less frequently in daltons, or as the to ...

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

, 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 ...

. 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 " sub-communities" working on particular scientific fields, and within particular institutions; interdisciplinary and cross-institutional activities are als ...

to date. Most recently, he has published a paper citing the paraphyly of his

bacteria Bacteria (; singular: 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 am ...

l kingdom, the origin of

Neomura Neomura is a possible clade composed of the two domains of life of Archaea and Eukaryota. The group was named by Thomas Cavalier-Smith in 2002. Its name means "new walls", reflecting his hypothesis that it evolved from Bacteria, and one of ...

from

Actinobacteria The ''Actinomycetota'' (or ''Actinobacteria'') are a phylum of all gram-positive bacteria. They can be terrestrial or aquatic. They are of great economic importance to humans because agriculture and forests depend on their contributions to soi ...

and taxonomy of

prokaryote A prokaryote () is a single-celled organism that lacks a nucleus and other membrane-bound organelles. The word ''prokaryote'' comes from the Greek πρό (, 'before') and κάρυον (, 'nut' or 'kernel').Campbell, N. "Biology:Concepts & Con ...

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 or AND may refer to: Logic, grammar, and computing * Conjunction (grammar), connecting two words, phrases, or clauses * Logical conjunction in mathematical logic, notated as "∧", "⋅", "&", or simple juxtaposition * Bitwise AND, a bool ...
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 Or or OR may refer to: Arts and entertainment Film and television * "O.R.", a 1974 episode of M*A*S*H * Or (My Treasure), a 2004 movie from Israel (''Or'' means "light" in Hebrew) Music * ''Or'' (album), a 2002 album by Golden Boy with Mis ...
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 "

" to describe the living world dates as far back as

Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his ennoblement in 1761 as Carl von Linné Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalised binomial nomenclature, t ...

(1707–1778) who divided the natural world into three kingdoms:

animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...

vegetable Vegetables are parts of plants that are consumed by humans or other animals as food. The original meaning is still commonly used and is applied to plants collectively to refer to all edible plant matter, including the flowers, fruits, stems ...

, and

mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. ...

.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 classifications "animal kingdom" (or kingdom Animalia) and "plant kingdom" (or kingdom

Plantae Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...

) remain in use by modern

evolutionary biologists Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life form ...

. The protozoa were originally classified as members of the animal kingdom. Now they are classified as multiple separate groups.

The third kingdom: Protista

250px, The sea anemone is an animal that resembles a plant. By mid-nineteenth century,

microscopic organisms A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...

were generally classified into four groups: #Protozoa (primitive animals), #

Protophyta Thallophytes (Thallophyta, Thallophyto or Thallobionta) are a polyphyletic group of non-motile organisms traditionally described as "thalloid plants", "relatively simple plants" or " lower plants". They form a division of kingdom Plantae that inc ...

(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 anatomist and paleontologist. Owen is generally considered to have been an outstanding naturalist with a remarkable gift for interpreting fossils. Ow ...

(1804–1892) proposed that the animal phylum Protozoa be elevated to the status of kingdom. In 1860,

John Hogg John Joseph Hogg (born 19 March 1949) is a former Australian politician who served as a Senator for Queensland from 1996 to 2014, representing the Labor Party. He served as President of the Senate from 2008 to 2014. Early life Hogg was born ...

(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,

(1834–1919) proposed the name "

Protista A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the e ...

" for the primigenial kingdom and included bacteria in this third kingdom of life.

The fourth kingdom: Fungi

By 1959,

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); #

, (or multicellular plants); #Fungi; and #Animalia (or

multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...

animals). Whittaker subdivided the Protista into two subkingdoms: #

Monera Monera (/məˈnɪərə/) (Greek - μονήρης (monḗrēs), "single", "solitary") is a biological kingdom that is made up of prokaryotes. As such, it is composed of single-celled organisms that lack a nucleus. The taxon Monera was first p ...

(

) and # Eunucleata (single celled

eukaryotes Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bact ...

The fifth kingdom: Bacteria (Monera)

Bacteria are fundamentally different from the

s (plants, animals, fungi, amebas, protozoa, and

chromista Chromista is a biological kingdom consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all protists whose plastids contain chlorophyll ''c'', suc ...

). Eukaryotes have

cell nuclei The cell nucleus (pl. nuclei; from Latin or , meaning ''kernel'' or ''seed'') 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, h ...

, 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 (; singular: 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 ...

(true bacteria) and

archaebacteria Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebacte ...

( archaea). In 1977

and

George E. Fox George Edward Fox (born December 17, 1945) is an astrobiologist, a Professor Emeritus and researcher at the University of Houston. He is an elected fellow of the American Academy of Microbiology, the American Association for the Advancement of S ...

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 biological classification introduced by Carl Woese, Otto Kandler, and Mark Wheelis in 1990 that divides cellular life forms into three domains, namely Archaea, Bacteria, and Eukaryota or Eukarya. The key differenc ...

such as: #Bacteria, #Archaea, and #Eucarya. But Cavalier-Smith considered Archaebacteria as a kingdom.

The seventh kingdom: Chromista

right , 350px, The 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 process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored in ...

. 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 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 i ...

. 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, in essence, the transportation system of the eukaryotic cell, and has many other important functions such as protein folding. It is a type of organelle made up of two subunits – rough endoplasmic reticulum ( ...

. 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: #

, # Animalia, #

Protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histor ...

, #

Fungi A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...

, #

Eubacteria Bacteria (; singular: 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 ...

, and #

Archaebacteria Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaebacte ...

The eighth kingdom: Archezoa

In 1983, Cavalier-Smith introduced Archezoa for (which he called) primitive protists that lack mitochondria. 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 now assigns former members of the kingdom Archezoa to the phylum Amoebozoa.

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 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 Red is the color at the long wavelength end of the visible spectrum of light, next to orange and opposite violet. It has a dominant wavelength of approximately 625–740 nanometres. It is a primary color in the RGB color model and a seconda ...

and

green algae The green algae (singular: green alga) are a group consisting of the Prasinodermophyta and its unnamed sister which contains the Chlorophyta and Charophyta/ Streptophyta. The land plants ( Embryophytes) have emerged deep in the Charophyte alg ...

), Chromista, and Bacteria. Five of Cavalier-Smith's kingdoms are classified as

as shown in the following scheme: *Eubacteria *

**Archaebacteria **Eukaryotes ***Kingdom Protozoa ***

Unikont Amorphea are members of a taxonomic supergroup that includes the basal Amoebozoa and Obazoa. That latter contains the Opisthokonta, which includes the Fungi, Animals and the Choanomonada, or Choanoflagellates. The taxonomic affinities of the mem ...

s (

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 ***

Bikont A bikont ("two flagella") is any of the eukaryotic organisms classified in the group Bikonta. Many single-celled members of the group, and the presumed ancestor, have two flagella. Enzymes Another shared trait of bikonts is the fusion of two ge ...

s (primarily

) ****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 (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have f ...

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, a ...

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. O ...

as a protozoan parvkingdom. By 1998, he had reclassified it as an animal subkingdom. Myxozoa contains three phyla,

Myxosporidia Myxosporea is a class of microscopic parasites, belonging to the Myxozoa clade within Cnidaria. They have a complex life cycle which comprises vegetative forms in two hosts, an aquatic invertebrate (generally an annelid but sometimes a bryozoan ...

Haplosporidia The Ascetosporea are a group of eukaryotes that are parasites of animals, especially marine invertebrates. The two groups, the haplosporids and paramyxids, are not particularly similar morphologically, but consistently group together on molecula ...

, 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 recent study recovered Mesoz ...

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 radially symmetric body plans. The term Radiata is no longer accepted, as it united several different groupings of animals that do not form a monophyletic ...

(phyla

Porifera Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throug ...

Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter. Their distinguishing feature is cnidocyt ...

Placozoa The Placozoa are a basal form of marine free-living (non-parasitic) multicellular organism. They are the simplest in structure of all animals. Three genera have been found: the classical '' Trichoplax adhaerens'', '' Hoilungia hongkongensis'', a ...

, and

Ctenophora Ctenophora (; ctenophore ; ) comprise 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 ...

), *

, *

, and *

Bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly an ...

(all other animal phyla). He created five new animal phyla: * Acanthognatha (

rotifers The rotifers (, from the Latin , "wheel", and , "bearing"), commonly 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 Ha ...

acanthocephalans Acanthocephala (Greek , ', thorn + , ', head) is a phylum of parasitic worms known as acanthocephalans, thorny-headed worms, or spiny-headed worms, characterized by the presence of an eversible proboscis, armed with spines, which it uses to p ...

gastrotrichs The gastrotrichs (phylum Gastrotricha), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environme ...

, and gnathostomulids), *

Brachiozoa Brachiozoa is a grouping of lophophorate animals including Brachiopoda and Phoronida. It also includes their ancestors, the extinct tommotiid Tommotiids are an extinct group of Cambrian invertebrates thought to be early lophophorates (the ...

(

brachiopods Brachiopods (), phylum Brachiopoda, are a phylum of trochozoan animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear end, wh ...

and

phoronids Phoronids (scientific name Phoronida, sometimes called horseshoe worms) are a small phylum of marine animals that Filter feeder, filter-feed with a lophophore (a "crown" of tentacles), and build upright tubes of chitin to support and protect thei ...

), *

Lobopoda ''Lobopoda'' is a genus of comb-clawed beetles in the family Tenebrionidae. The type species is '' Lobopoda striata''. The following subgenera In biology, a subgenus (plural: subgenera) is a taxonomic rank directly below genus. In the Interna ...

(

onychophorans Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, '' Peripatus ...

and

tardigrades Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbär ...

), *

Kamptozoa Entoprocta (), or Kamptozoa , is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that ...

(

Entoprocta Entoprocta (), or Kamptozoa , is a phylum of mostly sessile aquatic animals, ranging from long. Mature individuals are goblet-shaped, on relatively long stalks. They have a "crown" of solid tentacles whose cilia generate water currents that ...

and '' Symbion''), and * Nemathelminthes (

Nematoda The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...

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 siz ...

Loricifera Loricifera (from Latin, '' lorica'', corselet (armour) + ''ferre'', to bear) is a phylum of very small to microscopic marine cycloneuralian sediment-dwelling animals that had been determined to be 37 described species, in 9 genera, b ...

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 ( grc, κινέω, kīnéō, I move, ' "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 also called mud dragons. Modern species are or less, ...

) and recognized a total of 23 animal phyla, as shown here: *Kingdom Animaia **Subkingdom

***Infrakingdom Spongiaria ****Phylum

***Infrakingdom

Coelenterata Coelenterata is a term encompassing the animal phyla Cnidaria (coral animals, 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 ph ...

****Phylum

***Infrakingdom

****Phylum

**Subkingdom

***Phylum

**Subkingdom

***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 mem ...

****Infrakingdom Lophozoa *****Superphylum

Polyzoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...

******Phylum

Bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...

******Phylum

(

and

Cycliophora ''Symbion'' is a genus of commensal aquatic animals, less than 0.5 mm wide, found living attached to the mouthparts of cold-water lobsters. They have sac-like bodies, and three distinctly different forms in different parts of their two-sta ...

) *****Superphylum Conchozoa ******Phylum Mollusca ******Phylum

''sensu lato'' (

and Phoronida) *****Superphylum

Sipuncula The Sipuncula or Sipunculida (common names sipunculid worms or peanut worms) is a class containing about 162 species of unsegmented marine annelid worms. The name ''Sipuncula'' is from the genus name '' Sipunculus'', and comes from the Latin ...

******Phylum

*****Superphylum Vermizoa ******Phylum Nemertina ******Phylum Annelida ****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, about 20% of the known Chaetognatha species are benthic, and ca ...

****Infrakingdom

Ecdysozoa Ecdysozoa () is a group of protostome animals, including Arthropoda (insects, chelicerata, crustaceans, and myriapods), Nematoda, and several smaller phyla. They were first defined by Aguinaldo ''et al.'' in 1997, based mainly on phylogeneti ...

*****Superphylum Nemathelminthes ******Phylum Nemathelminthes (

and

; Priapozoa,

and

) *****Superphylum Haemopoda ******Phylum

Lobopodia The lobopodians, members of the informal group Lobopodia (from the Greek, meaning "blunt feet"), or the formally erected phylum Lobopoda Cavalier-Smith (1998), are panarthropods with stubby legs called lobopods, a term which may also be used as ...

(

Onychophora Onychophora (from grc, ονυχής, , "claws"; and , , "to carry"), commonly known as velvet worms (due to their velvety texture and somewhat wormlike appearance) or more ambiguously as peripatus (after the first described genus, ''Peripatus' ...

and

Tardigrada Tardigrades (), known colloquially as water bears or moss piglets, are a phylum of eight-legged segmented micro-animals. They were first described by the German zoologist Johann August Ephraim Goeze in 1773, who called them Kleiner Wasserbär ...

) ******Phylum

Arthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a Segmentation (biology), segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and Arth ...

****Infrakingdom

Platyzoa The paraphyletic "Platyzoa" are a group of protostome unsegmented animals proposed by Thomas Cavalier-Smith in 1998. Cavalier-Smith included in Platyzoa the phylum Platyhelminthes (or flatworms), and a new phylum, the Acanthognatha, into which ...

*****Phylum

Platyhelminthes The flatworms, flat worms, Platyhelminthes, or platyhelminths (from the Greek πλατύ, ''platy'', meaning "flat" and ἕλμινς (root: ἑλμινθ-), ''helminth-'', meaning "worm") are a phylum of relatively simple bilaterian, unsegment ...

(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 (physical a ...

) *****Phylum Acanthognatha (incl.

Rotifera The rotifers (, from the Latin , "wheel", and , "bearing"), commonly 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 H ...

Acanthocephala Acanthocephala (Greek , ', thorn + , ', head) is a phylum of parasitic worms known as acanthocephalans, thorny-headed worms, or spiny-headed worms, characterized by the presence of an eversible proboscis, armed with spines, which it uses to p ...

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), commonly referred to as hairybellies or hairybacks, are a group of microscopic (0.06-3.0 mm), worm-like, acoelomate animals, and are widely distributed and abundant in freshwater and marine environme ...

) ***Branch

Deuterostomia Deuterostomia (; in Greek) are animals typically characterized by their anus forming before their mouth during embryonic development. The group's sister clade is Protostomia, animals whose digestive tract development is more varied. Some examp ...

****Infrakingdom

Coelomopora Ambulacraria , or Coelomopora , is a clade of invertebrate phyla that includes echinoderms and hemichordates; a member of this group is called an ambulacrarian. Phylogenetic analysis suggests the echinoderms and hemichordates separated around ...

*****Phylum

Echinoderm An echinoderm () is any member of the phylum Echinodermata (). The adults are recognisable by their (usually five-point) radial symmetry, and include starfish, brittle stars, sea urchins, sand dollars, and sea cucumbers, as well as the s ...

ata *****Phylum Hemichordata ****Infrakingdom Chordonia *****Phylum Urochorda *****Phylum

Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These five ...

**Subkingdom

***Phylum

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 collagenous 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 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 ...

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

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 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 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 ...

*** uniciliate protozoan phylum Choanozoa *** kingdom Fungi *** kingdom Animalia * Bikonts ** protozoan infrakingdom

*** phylum

Cercozoa Cercozoa is a phylum of diverse single-celled eukaryotes. They lack shared morphological characteristics at the microscopic level, and are instead defined by molecular phylogenies of rRNA and actin or polyubiquitin. They were the first major ...

*** 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 is a phylum of diverse single-celled eukaryot ...

( Radiozoa and

Foraminifera Foraminifera (; Latin for "hole bearers"; informally called "forams") are single-celled organisms, members of a phylum or class of amoeboid protists characterized by streaming granular ectoplasm for catching food and other uses; and commonly ...

) ** protozoan infrakingdom

*** phylum

Loukozoa Loukozoa (+ Ancyromonads) (From Greek ''loukos'': groove) is a proposed taxon used in some classifications of eukaryotes, consisting of the Metamonada and Malawimonadea. Ancyromonads are closely related to this group, as sister of the entire gr ...

*** 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 three major clades, i.e., Kinetoplastea, D ...

*** phylum

Percolozoa The Percolozoa are a group of colourless, non-photosynthetic Excavata, including many that can transform between amoeboid, flagellate, and cyst stages. Characteristics Most Percolozoa are found as bacterivores in soil, fresh water and occasion ...

** protozoan phylum

Apusozoa The Apusozoa are an Obazoa phylum comprising several genera of flagellate eukaryotes. They are usually around 5–20 μm in size, and occur in soils and aquatic habitats, where they feed on bacteria. They are grouped together based on the presen ...

( Thecomonadea and Diphylleida) ** the

chromalveolate Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in ...

clade *** kingdom

(

Cryptista Cryptista is a clade of algae-like eukaryotes. It is most likely related to Archaeplastida which includes plants and many algae, within the larger group Diaphoretickes. Although it has sometimes placed along with Haptista in the group Hacrobia, ...

Heterokonta Heterokonts are a group of protists (formally referred to as Heterokonta, Heterokontae or Heterokontophyta). The group is a major line of eukaryotes. Most are algae, ranging from the giant multicellular kelp to the unicellular diatoms, which ...

, 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) 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. The ...

) ** kingdom

( Viridaeplantae,

Rhodophyta Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...

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 speci ...

)

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 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 Taxonomic I ...

References

{{reflist Taxonomy (biology) High-level systems of taxonomy