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In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called Phylum (singular phylum).
Traditionally, textbooks from Canada and the United States have used a system of six kingdoms (, , Fungus, , Archaea/Archaebacteria, and Bacteria or Eubacteria), while textbooks in other parts of the world, such as Bangladesh, Brazil, Greece, India, Pakistan, Spain, and the United Kingdom have used five kingdoms (Animalia, Plantae, Fungi, Protista and Monera).
Some recent classifications based on modern cladistics have explicitly abandoned the term kingdom, noting that some traditional kingdoms are not Monophyly, meaning that they do not consist of all the descendants of a common ancestor. The terms flora (for plants), fauna (for animals), and, in the 21st century, funga (for fungi) are also used for life present in a particular region or time.
Prefixes can be added so subkingdom ( subregnum) and infrakingdom (also known as infraregnum) are the two ranks immediately below kingdom. Superkingdom may be considered as an equivalent of domain or empire or as an independent rank between kingdom and domain or subdomain. In some classification systems the additional rank branch (Latin: ramus) can be inserted between subkingdom and infrakingdom, e.g., Protostomia and Deuterostomia in the classification of Cavalier-Smith.
Carl Linnaeus (1707–1778) laid the foundations for modern biological nomenclature, now regulated by the Nomenclature Codes, in 1735. He distinguished two kingdoms of living things: Regnum Animale ('animal kingdom') and Regnum Vegetabile ('vegetable kingdom', for ). Linnaeus also included in his classification system, placing them in a third kingdom, Mineralia.
At first, microscopic organisms were classified within the animal and plant kingdoms. However, by the mid–19th century, it had become clear to many that "the existing dichotomy of the plant and animal kingdoms had rapidly blurred at its boundaries and outmoded".
In 1860 John Hogg proposed the Protoctista, a third kingdom of life composed of "all the lower creatures, or the primary organic beings"; he retained Regnum Lapideum as a fourth kingdom of minerals. In 1866, Ernst Haeckel also proposed a third kingdom of life, the Protista, for "neutral organisms" or "the kingdom of primitive forms", which were neither animal nor plant; he did not include the Regnum Lapideum in his scheme. Haeckel revised the content of this kingdom a number of times before settling on a division based on whether organisms were unicellular (Protista) or multicellular (animals and plants).
In 1938, Herbert F. Copeland proposed a four-kingdom classification by creating the novel Kingdom Monera of prokaryotic organisms; as a revised phylum Monera of the Protista, it included organisms now classified as Bacteria and Archaea. Ernst Haeckel, in his 1904 book The Wonders of Life, had placed the blue-green algae (or Phycochromacea) in Monera; this would gradually gain acceptance, and the blue-green algae would become classified as bacteria in the phylum Cyanobacteria.
In the 1960s, Roger Stanier and C. B. van Niel promoted and popularized Édouard Chatton's earlier work, particularly in their paper of 1962, "The Concept of a Bacterium"; this created, for the first time, a rank above kingdom—a superkingdom or empire—with the two-empire system of prokaryotes and eukaryotes. The two-empire system would later be expanded to the three-domain system of Archaea, Bacteria, and Eukaryota.
The remaining two kingdoms, Protista and Monera, included unicellular and simple cellular colonies. The five kingdom system may be combined with the two empire system. In the Whittaker system, Plantae included some algae. In other systems, such as Lynn Margulis's system of five kingdoms, the plants included just the land plants (Embryophyte), and Protoctista has a broader definition.
Following publication of Whittaker's system, the five-kingdom model began to be commonly used in high school biology textbooks. But despite the development from two kingdoms to five among most scientists, some authors as late as 1975 continued to employ a traditional two-kingdom system of animals and plants, dividing the plant kingdom into subkingdoms Prokaryota (bacteria and cyanobacteria), Mycota (fungi and supposed relatives), and Chlorota (algae and land plants).
Finally, some protists lacking mitochondria were discovered. As mitochondria were known to be the result of the endosymbiosis of a proteobacterium, it was thought that these amitochondriate eukaryotes were primitively so, marking an important step in eukaryogenesis. As a result, these amitochondriate protists were separated from the protist kingdom, giving rise to the, at the same time, superkingdom and kingdom Archezoa. This superkingdom was opposed to the Metakaryota superkingdom, grouping together the five other eukaryotic kingdoms (Animalia, Protozoa, Fungi, Plantae and Chromista). This was known as the Archezoa hypothesis, which has since been abandoned; later schemes did not include the Archezoa–Metakaryota divide.
‡ No longer recognized by taxonomists.
Cavalier-Smith does not accept the requirement for taxa to be Monophyly ("holophyletic" in his terminology) to be valid. He defines Prokaryota, Bacteria, Negibacteria, Unibacteria, and Posibacteria as valid Paraphyly (therefore "monophyletic" in the sense he uses this term) taxa, marking important innovations of biological significance (in regard of the concept of biological Ecological niche).
In the same way, his paraphyletic kingdom Protozoa includes the ancestors of Animalia, Fungi, Plantae, and Chromista. The advances of phylogenetic studies allowed Cavalier-Smith to realize that all the phyla thought to be (i.e. primitively amitochondriate eukaryotes) had in fact secondarily lost their mitochondria, typically by transforming them into new organelles: . This means that all living eukaryotes are in fact , according to the significance of the term given by Cavalier-Smith. Some of the members of the defunct kingdom Archezoa, like the phylum Microsporidia, were reclassified into kingdom Fungi. Others were reclassified in kingdom Protozoa, like Metamonada which is now part of infrakingdom Excavata.
Because Cavalier-Smith allows paraphyly, the diagram below is an "organization chart", not an "ancestor chart", and does not represent an evolutionary tree.
In 1990, the name "domain" was proposed for the highest rank. This term represents a synonym for the category of dominion (lat. dominium), introduced by Moore in 1974. Unlike Moore, Woese et al. (1990) did not suggest a Latin term for this category, which represents a further argument supporting the accurately introduced term dominion.
Woese divided the prokaryotes (previously classified as the Kingdom Monera) into two groups, called bacteria and Archaea, stressing that there was as much genetic difference between these two groups as between either of them and all eukaryotes.
According to genetic data, although eukaryote groups such as plants, fungi, and animals may look different, they are more closely related to each other than they are to either the Eubacteria or Archaea. It was also found that the eukaryotes are more closely related to the Archaea than they are to the Eubacteria. Although the primacy of the Eubacteria-Archaea divide has been questioned, it has been upheld by subsequent research. There is no consensus on how many kingdoms exist in the classification scheme proposed by Woese.
On this basis, the diagram opposite (redrawn from their article) showed the real "kingdoms" (their quotation marks) of the eukaryotes. A classification which followed this approach was produced in 2005 for the International Society of Protistologists, by a committee which "worked in collaboration with specialists from many societies". It divided the eukaryotes into the same six "supergroups". The published classification deliberately did not use formal taxonomic ranks, including that of "kingdom".
this system the multicellular animals (Metazoa) are descended from the same ancestor as both the unicellular and the fungi which form the Opisthokonta. Plants are thought to be more distantly related to animals and fungi.
However, in the same year as the International Society of Protistologists' classification was published (2005), doubts were being expressed as to whether some of these supergroups were monophyletic, particularly the Chromalveolata, and a review in 2006 noted the lack of evidence for several of the six proposed supergroups.
, there is widespread agreement that the Rhizaria belong with the Stramenopiles and the Alveolata, in a clade dubbed the SAR supergroup, so that Rhizaria is not one of the main eukaryote groups.
The eocyte hypothesis proposes that the emerged from a phylum within the archaea called the Thermoproteota (formerly known as eocytes or Crenarchaeota).
There is ongoing debate as to whether viruses can be included in the tree of life. The arguments against include the fact that they are obligate intracellular parasites that lack metabolism and are not capable of Self-replication outside of a host cell. Another argument is that their placement in the tree would be problematic, since it is suspected that viruses have various evolutionary origins, and they have a penchant for harvesting nucleotide sequences from their hosts.
Four kingdoms
The development of microscopy revealed important distinctions between those organisms whose cells do not have a distinct Cell nucleus () and organisms whose cells do have a distinct nucleus (). In 1937 Édouard Chatton introduced the terms "prokaryote" and "eukaryote" to differentiate these organisms.
Five kingdoms
The differences between fungi and other organisms regarded as plants had long been recognised by some; Haeckel had moved the fungi out of Plantae into Protista after his original classification, but was largely ignored in this separation by scientists of his time. Robert Whittaker recognized an additional kingdom for the Fungus. The resulting five-kingdom system, proposed in 1969 by Whittaker, has become a popular standard and with some refinement is still used in many works and forms the basis for new multi-kingdom systems. It is based mainly upon differences in nutrition; his Plantae were mostly multicellular , his Animalia multicellular , and his Fungi multicellular .
+ Kingdom Monera
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Kingdom Protista
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Kingdom Fungi
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Kingdom Animalia
Six kingdoms
In 1977, Carl Woese and colleagues proposed the fundamental subdivision of the prokaryotes into the Eubacteria (later called the Bacteria) and Archaebacteria (later called the Archaea), based on ribosomal RNA structure; this would later lead to the proposal of three "domains" of life, of Bacteria, Archaea, and Eukaryota. Combined with the five-kingdom model, this created a six-kingdom model, where the kingdom Monera is replaced by the kingdoms Bacteria and Archaea. This six-kingdom model is commonly used in recent US high school biology textbooks, but has received criticism for compromising the current scientific consensus. But the division of prokaryotes into two kingdoms remains in use with the recent seven kingdoms scheme of Thomas Cavalier-Smith, although it primarily differs in that Protista is replaced by Protozoa and Chromista.
Eight kingdoms
Thomas Cavalier-Smith supported the consensus at that time, that the difference between Eubacteria and Archaebacteria was so great (particularly considering the genetic distance of ribosomal genes) that the prokaryotes needed to be separated into two different kingdoms. He then divided Eubacteria into two subkingdoms: Negibacteria (Gram-negative bacteria) and Posibacteria (Gram-positive bacteria). Technological advances in electron microscopy allowed the separation of the Chromista from the Plantae kingdom. Indeed, the chloroplast of the chromists is located in the lumen of the endoplasmic reticulum instead of in the cytosol. Moreover, only chromists contain chlorophyll c. Since then, many non-photosynthetic phyla of protists, thought to have secondarily lost their chloroplasts, were integrated into the kingdom Chromista.
Six kingdoms (1998)
In 1998, Cavalier-Smith published a six-kingdom model, which has been revised in subsequent papers. The version published in 2009 is shown below.Compared to the version Cavalier-Smith published in 2004, the Alveolata and the have been moved from Kingdom Protozoa to Kingdom Chromista. Cavalier-Smith no longer accepted the importance of the fundamental Eubacteria–Archaebacteria divide put forward by Woese and others and supported by recent research. The kingdom Bacteria (sole kingdom of empire Prokaryota) was subdivided into two sub-kingdoms according to their membrane topologies: Unibacteria and Negibacteria. Unibacteria was divided into phyla Archaebacteria and Posibacteria; the bimembranous-unimembranous transition was thought to be far more fundamental than the long branch of genetic distance of Archaebacteria, viewed as having no particular biological significance.
+ Cavalier-Smith's six kingdom system (1998)
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Seven kingdoms
Cavalier-Smith and his collaborators revised their classification in 2015. In this scheme they introduced two superkingdoms of Prokaryota and Eukaryota and seven kingdoms. Prokaryota have two kingdoms: Bacteria and Archaea. (This was based on the consensus in the Taxonomic Outline of Bacteria and Archaea, and the Catalogue of Life). The Eukaryota have five kingdoms: Protozoa, Chromista, Plantae, Fungi, and Animalia. In this classification a protist is any of the eukaryotic unicellular organisms.
Summary
The kingdom-level classification of life is still widely employed as a useful way of grouping organisms, notwithstanding some problems with this approach:
Beyond traditional kingdoms
While the concept of kingdoms continues to be used by some taxonomists, there has been a movement away from traditional kingdoms, as they are no longer seen as providing a Cladistics classification, where there is emphasis in arranging organisms into Clade.
Three domains of life
Based on RNA studies, Carl Woese thought life could be divided into three large divisions and referred to them as the "three primary kingdom" model or "urkingdom" model.
Eukaryotic supergroups
In 2004, a review article by Simpson and Roger noted that the Protista were "a grab-bag for all that are not animals, plants or fungi". They held that only monophyletic groups should be accepted as formal ranks in a classification and that – while this approach had been impractical previously (necessitating "literally dozens of eukaryotic 'kingdoms) – it had now become possible to divide the eukaryotes into "just a few major groups that are probably all monophyletic".
Prokaryotic kingdoms
The Prokaryotic Code treats Bacteria and Archaea each as a domain. Since 2024, each domain each contains 4 kingdoms, for a total of 8.
Comparison of top level classification
Some authors have added non-cellular life to their classifications. This can create a "superdomain" called "Acytota", also called "Aphanobionta", of non-cellular life; with the other superdomain being "cytota" or cellular life.<" title="https://pubmed.ncbi.nlm.nih.gov/26305806/"><">
/ref> (1993). 9780412364402, Chapman & Hall. ISBN 9780412364402 (see section below for further discussion)
Viruses
The International Committee on Taxonomy of Viruses uses the taxonomic rank "kingdom" in the classification of viruses (with the suffix -virae); but this is beneath the top level classifications of realm and subrealm.
See also
Notes
Further reading
External links
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