Cavalier-Smith's system of classification
The initial version of a classification system of life by British zoologist Thomas Cavalier-Smith appeared in 1978.[1][2] 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, Ernst Haeckel, Robert Whittaker, and Carl Woese, Cavalier-Smith's classification attempts to incorporate the latest developments in taxonomy.[3][4], 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. [citation needed]
Cavalier-Smith has published extensively on the
According to Palaeos.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 [hand], 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, [for example], Zrzavý (2001)[5] [and] Patterson (1999).[6][7][8] Nevertheless, he deals with all of the relevant facts.[9]
Eight kingdoms model
The first two kingdoms of life: Plantae and Animalia
The use of the word "
The third kingdom: Protista
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,
)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 multicellularanimals).
Whittaker subdivided the Protista into two subkingdoms:
The fifth kingdom: Bacteria (Monera)
Bacteria are fundamentally different from the
- Plantae,
- Animalia,
- Protista (Eunucleata),
- Fungi, and
- Monera (the kingdom bacteria).[16]
The sixth kingdom: Archaebacteria
The kingdom Monera can be divided into two distinct groups:
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).[18]
In 1990, Woese introduced domain above kingdom by creating three-domain system such as:
- Bacteria,
- Archaea, and
- Eucarya.[17]
But Cavalier-Smith considered Archaebacteria as a kingdom.[19]
The seventh kingdom: Chromista
By 1981, Cavalier-Smith had divided all the eukaryotes into nine kingdoms.[20] In it, he created Chromista for a separate kingdom of some protists.[21]
Most chromists are
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",[20] which includes:
The eighth kingdom: Archezoa
In 1983, Cavalier-Smith adapted the term Archezoa, previously used by Haeckel, Perty, and for protists that lack
1983a | 1983b | 1987 | 1989 | 1990 | 1991 | 1993 | 1995 | 1996 | 1997 | 1998 | 1999 | |
---|---|---|---|---|---|---|---|---|---|---|---|---|
Diplomonads | + | + | + | + | + | + | + | + | + | + | + | + |
Retortamonads | + | + | + | + | + | + | + | + | + | + | + | + |
Oxymonads | + | + | + | + | + | + | + | + | + | + | + | + |
Microspora | + | + | + | + | + | + | + | + | + | |||
Hypermastigids | + | + | + | |||||||||
Trichomonads | + | + | + | |||||||||
Mastigamoebids | + | + | + | + | + | + | ||||||
Pelomyxa | + | + | + | + | + | |||||||
Entamoeba | + | + | + | + | + | |||||||
Phreatamoeba | + | + | ||||||||||
Trimastix | + |
Archezoa is now defunct.[40] He now assigns former members of the kingdom Archezoa to the phylum Amoebozoa.[41]
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.[13] The first major division relied on the basis of presence or absence of dictyosomes, although a Gogli Apparatus was subsequently demonstrated in the 'Adictyozoa'.[42]
# | Phylum | Assigned to: | Characteristics | Fate |
---|---|---|---|---|
1 | Percolozoa | subkingdom Adictyozoa | lacks Golgi dictyosomes | |
2 | Parabasalia
|
subkingdom Dictyozoa branch Parabasalia |
has Golgi dictyosomes lacks mitochondria |
|
3 | Euglenozoa | subkingdom Dictyozoa branch Bikonta infrakingdom Euglenozoa |
has mitochondria of
with trans-splicing |
|
4 | flagellates )
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Ciliomyxa superphylum Opalomyxa |
has cortical alveoli
|
|
5 | Mycetozoa (slime molds) | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Ciliomyxa superphylum Opalomyxa |
has introns predominantly cortical alveoli
|
|
6 | Choanozoa (choanoflagellates) | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Ciliomyxa superphylum Choanozoa |
has cortical alveoli
|
|
7 | Dinoflagellata and Protalveolata )
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Alveolata superphylum Miozoa |
has cortical alveoli
|
Reassigned to Miozoa in Alveolata.[43] |
8 | Apicomplexa | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Alveolata superphylum Miozoa |
has cortical alveoli
|
Reassigned to Miozoa in Alveolata.[43] |
9 | Ciliophora
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Alveolata superphylum Heterokaryota |
has cortical alveoli
|
Reassigned to Alveolata.[43] |
10 | amoebae )
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Neosarcodina |
has introns
|
|
11 | Reticulosa (foraminifera; reticulopodial amoebae) | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Neosarcodina |
has introns
|
|
12 | Heliozoa | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Actinopoda |
has axopodia
|
|
13 | Radiozoa
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Actinopoda |
has axopodia
|
|
14 | Entamoebia | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Entamoebia |
has introns no mitochondria, peroxisomes, hydrogenosomes or cilia transient intranuclear centrosomes |
|
15 | Myxosporidia
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Myxozoa |
has introns endoparasitic, multicellular spores, mitochondria, and no cilia |
Reclassified as animals in 1998.[44] |
16 | Haplosporidia
|
subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Myxozoa |
has introns endoparasitic, multicellular spores, mitochondria, and no cilia |
Reclassified as animals in 1998.[44][clarification needed] |
17 | Paramyxia | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Myxozoa |
has introns endoparasitic, multicellular spores, mitochondria, and no cilia |
Reclassified as animals in 1998.[44] |
18 | Mesozoa | subkingdom Dictyozoa branch Bikonta infrakingdom Neozoa parvkingdom Mesozoa |
has introns tubular mitochondrial cristae multicellular with no collagenous connective tissue |
Reclassified as animals in 1998.[44] |
The phylum Opalozoa was established by Cavalier-Smith in 1991.[45]
Six kingdoms models
By 1998, Cavalier-Smith had reduced the total number of kingdoms from eight to six: Animalia, Protozoa, Fungi, Plantae (including
Five of Cavalier-Smith's kingdoms are classified as
- Eubacteria
- Neomura
- Archaebacteria
- Eukaryotes
- Kingdom Protozoa
- heterotrophs)
- Kingdom Animalia
- Kingdom Fungi
- photosynthetic)
- 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 and unikonts are descended from uniciliates. Unikont cells often have only one centriole 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
In his 1998 scheme, the animal kingdom was divided into four subkingdoms:
He created five new animal phyla:
- gnathostomulids),
- phoronids),
- tardigrades),
- Kamptozoa (Entoprocta and Symbion), and
- Nemathelminthes (Nematoda, Nematomorpha, Loricifera, Priapulida, and Kinorhyncha)
and recognized a total of 23 animal phyla, as shown here:
- Kingdom Animaia
- Subkingdom Radiata
- Infrakingdom Spongiaria
- Phylum Porifera
- Phylum
- Infrakingdom Coelenterata
- Phylum Cnidaria
- Phylum Ctenophora
- Infrakingdom Placozoa
- Phylum Placozoa
- Infrakingdom
- Subkingdom Myxozoa
- Phylum Myxosporidia
- Subkingdom Bilateria
- Branch Protostomia
- Infrakingdom Lophozoa
- Infrakingdom Chaetognathi
- Phylum Chaetognatha
- Infrakingdom Ecdysozoa
- Superphylum Nemathelminthes
- Phylum )
- Superphylum Haemopoda
- Phylum Tardigrada)
- Phylum Arthropoda
- Phylum
- Superphylum Nemathelminthes
- Infrakingdom Platyzoa
- Phylum Platyhelminthes (incl. Xenacoelomorpha)
- Phylum Gastrotricha)
- Phylum
- Branch Deuterostomia
- Infrakingdom Coelomopora
- Phylum Echinodermata
- Phylum Hemichordata
- Infrakingdom Chordonia
- Phylum Urochorda
- Phylum Chordata
- Phylum
- Infrakingdom Coelomopora
- Branch
- Subkingdom Mesozoa
- Subkingdom Radiata
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
- 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.
Unikonts and bikonts
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
- phylum Cercozoa
- phylum Radiozoa and Foraminifera)
- protozoan infrakingdom Excavata
- phylum Loukozoa
- phylum Metamonada
- phylum Euglenozoa
- phylum Percolozoa
- protozoan phylum Diphylleida)
- the chromalveolateclade
- kingdom Haptophyta)
- protozoan infrakingdom Alveolata
- phylum Ciliophora
- phylum Dinozoa, and Apicomplexa)
- phylum
- kingdom
- kingdom Glaucophyta)
- protozoan infrakingdom Rhizaria
Cladogram of life
By September 2003, Cavalier-Smith's tree of life looked like this:[46]
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: Cavalier-Smith's Tree of Life, 2006[cstol 1]
[A]
|
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Legend:
[A]
[B] Oxygenic Photosynthesis, Omp85 and four new catalases.
[C]
[D]
[E] Flagella.
[F] Four sections: an amino acid in HSP60 and FtsZ and a domain in RNA polymerases β and σ.
[G]
[H]
[I] Glycerol 1-P dehydrogenase.
[J] Proteasome and phosphatidylinositol.
[K] Neomura revolution: Replacement of peptidoglycan by glycoproteins and lipoproteins.
[L] Reverse
[M] Phagocytosis.
- PMID 16754610.
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.[47]
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).[48]
Cavalier-Smith and his collaborators revised the classification in 2015, and published it in
References
- ^ Cavalier-Smith, T. 1978. The evolutionary origin and phylogeny of microtubules, mitotic spindles and eucaryotic flagella. BioSystems 10: 93-114.
- ^ C. Jeffrey. 1982. Kingdoms, Codes and Classification. Kew Bulletin: 37: 403-416
- JSTOR 4451669.
- S2CID 16939886.
- PMID 11437135.
- S2CID 4367158.
- ^ "Apusomonadida". Archived from the original on 2008-09-07. Retrieved 2016-02-11.
- ^ Eukarya Archived 2010-12-20 at the Wayback Machine.
- ^ "Origins of the Eukarya". Archived from the original on December 20, 2010. Retrieved February 9, 2009.
- ^ 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
- ^ PMID 10943416.
- ^ Penny, Douglas A.; Waern, Regina (1965). Biology. An Introduction to aspects of Modern Biological Science. Vancouver Calgary Toronto Montreal: Pitman Publishing. pp. 626–40.
- ^ PMID 8302218.
- ^ Kutschera, U. Haeckel's 1866 tree of life and the origin of eukaryotes. Nat Microbiol 1, 16114 (2016). https://doi.org/10.1038/nmicrobiol.2016.114
- S2CID 28836075.
- ISBN 978-0-7637-4649-0.
Robert Whittaker (1969) five kingdoms.
- ^ PMID 2112744.
- PMID 270744.
- ^ S2CID 30405691.
- ^ PMID 7337818.
- ^ Cavalier-Smith, T. (1986). "The kingdom Chromista : Origin and systematics". Progress in Phycological Research. 4: 309–347.
- PMID 3561476.
- ^ Cavalier Smith, T. 1983. A six-kingdom classification and a unified phylogeny. Endocytobiology 2:1027–1034.
- ^ Cavalier Smith, T. 1987. Eukaryotes with no mitochondria. Nature (London) 326:332–333.
- ^ Cavalier Smith, T. 1987 The origin of eukaryote and archaebacterial cells. Annals of the New York Academy of Sciences 503:17–54.
- ^ Cavalier Smith, T. 1989. Archaebacteria and Archezoa. Nature (London) 339:100–101.
- ^ Cavalier Smith, T. 1990. Microorganism megaevolution: integrating the fossil and living evidence. Revue de Micropalêontologie 33:145–154.
- ^ Cavalier Smith, T. 1991. Archamoebae: the ancestral eukaryotes? Biosystems 25:25–38.
- ^ Cavalier Smith, T. . The evolution of cells. pp 271–304 in S. Osawa and T. Honjo, eds. Evolution of life. Springer, Berlin.
- ^ Cavalier Smith, T. 1992. Percolozoa and the symbiotic origin of the metakaryote cell. Endocytobiology 5:399–406.
- ^ Cavalier Smith, T. Kingdom Protozoa and its 18 Phyla. Microbiological Reviews 57:953–994.
- ^ Cavalier Smith, T. 1995. Zooflagellate phylogeny and classification. Cytology 37:1010–1029.
- ^ Cavalier Smith, T. 1997. Amoeboflagellates and mitochondrial cristae in eukaryote evolution: megasystematics of the new protozoan subkingdoms Eozoa and Neozoa. Archiv für Protistenkunde 147:237–258.
- ^ Cavalier Smith, T. 1997. Zooflagellate phylogeny and the evolution and classification of protozoa. Page 65 in Programme and abstracts, Tenth International Congress of Protozoology, Sydney.
- ^ Cavalier Smith, T. 1998. A revised six–kingdom system of life. Biological Reviews of the Cambridge Philosophical Society 73:203–266.
- ^ Cavalier Smith, T. 1999. Zooflagellate phylogeny and the systematics of protozoa. Biological Bulletin (Woods Hole) 196: 393–395.
- ^ Cavalier-Smith, T.and Chao, E. 1995. The opalozoan Apusomonas is related to the common ancestor of animals, fungi and choanoflagellates. Proceedings of the Royal Society of London B, Biological Sciences 261:1–6.
- ^ Cavalier-Smith, T.and Chao, E. 1996. Molecular diversity of the free-living archezoan Trepomonas agilis and the nature of the first eukaryote. Journal of Molecular Evolution 43:551–563.
- ^ Patterson, D. J. 1999. The diversity of eukaryotes. American Naturalist, Supplement 65: 96-124.
- S2CID 28992966.
- PMID 15306349.
- PMID 29535209.
- ^ .
- ^ S2CID 6557779.
- S2CID 84129692.
- PMID 12956967.
- PMID 20132544.
- S2CID 38405158.
- PMID 25923521.