Understanding Kingdoms in Biology
Understanding Kingdoms in Biology
In biology, a kingdom is the second highest taxonomic rank, just below domain. Kingdoms are divided into smaller groups called
phyla (singular phylum).
Traditionally, textbooks from Canada and the United States have used a system of six kingdoms (Animalia, Plantae, Fungi, Protista,
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 monophyletic, 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.[1][2]
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 plants). Linnaeus also included minerals in his
classification system, placing them in a third kingdom, Regnum Lapideum.
In 1674, Antonie van Leeuwenhoek, often called the "father of microscopy", sent the Royal Society of London a copy of his first observations of microscopic
single-celled organisms. Until then, the existence of such microscopic organisms was entirely unknown. Despite this, Linnaeus did not include any microscopic
creatures in his original taxonomy.
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 become] rapidly blurred at its boundaries and outmoded".[8]
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.[8] 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.[8] 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).[8]
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.[8][9]
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.[9] The two-empire system would later be expanded to the three-domain system of Archaea,
Bacteria, and Eukaryota.[10]
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,[8] but was largely ignored in this separation by scientists of his time. Robert Whittaker recognized an additional
kingdom for the Fungi.[11] 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
autotrophs, his Animalia multicellular heterotrophs, and his Fungi multicellular saprotrophs.
The remaining two kingdoms, Protista and Monera, included unicellular and simple cellular colonies.[11] 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 (Embryophyta), and Protoctista has a broader definition.[12]
Following publication of Whittaker's system, the five-kingdom model began to be commonly used in high school biology textbooks.[13] 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).[14]
Kingdom Monera Kingdom Protista Kingdom Plantae Kingdom Fungi Kingdom Animalia
Phylum Acanthocephala
Phylum Aschelminthes
Phylum Entoprocta or Kamptozoa
Grade Coelomata
Subgrade Schizocoela
Phylum Brachiata or
Pogonophora
Phylum Chaetognatha
Phylum Echinodermata
Phylum Hemichordata
Phylum Chordata
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;[15] this would later lead to the proposal of three "domains" of life, of Bacteria, Archaea, and
Eukaryota.[5] Combined with the five-kingdom model, this created a six-kingdom model, where the kingdom Monera is replaced by the kingdoms Bacteria and
Archaea.[16] This six-kingdom model is commonly used in recent US high school biology textbooks, but has received criticism for compromising the current
scientific consensus.[13] 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.[17]
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.
Finally, some protists lacking mitochondria were discovered.[18] 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;[19] later schemes did not include the Archezoa–Metakaryota divide.[6][17]
Kingdom Eubacteria
Superkingdom Prokaryota
Kingdom Archaebacteria
Superkingdom Archezoa‡ Kingdom Archezoa‡
Kingdom Protozoa
Life
Kingdom Chromista
Superkingdom Metakaryota‡ Kingdom Plantae
Kingdom Fungi
Kingdom Animalia
‡ No longer recognized by taxonomists.
Cavalier-Smith does not accept the requirement for taxa to be monophyletic ("holophyletic" in his terminology) to be valid. He defines Prokaryota, Bacteria,
Negibacteria, Unibacteria, and Posibacteria as valid paraphyla (therefore "monophyletic" in the sense he uses this term) taxa, marking important innovations of
biological significance (in regard of the concept of biological 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 archezoans (i.e. primitively amitochondriate eukaryotes) had in fact secondarily lost their
mitochondria, typically by transforming them into new organelles: Hydrogenosomes. This means that all living eukaryotes are in fact metakaryotes, 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.
Kingdom Chromista — e.g. Alveolata, cryptophytes, Heterokonta (Brown Algae, Diatoms etc.), Haptophyta,
Life Rhizaria
Empire Eukaryota
Kingdom Plantae — e.g. glaucophytes, red and green algae, land plants
Kingdom Fungi
Kingdom Animalia
Cavalier-Smith's six kingdom system (1998)[6]
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.[17]
Kingdom Bacteria
Superkingdom Prokaryota
Kingdom Archaea
Kingdom Protozoa — e.g. Amoebozoa, Choanozoa, Excavata
Life Kingdom Chromista — e.g. Alveolata, cryptophytes, Heterokonta (Brown Algae, Diatoms etc.), Haptophyta,
Rhizaria
Superkingdom Eukaryota
Kingdom Plantae — e.g. glaucophytes, red and green algae, land plants
Kingdom Fungi
Kingdom Animalia
Summary
Woese et Cavalier-
Linnaeus Haeckel Chatton Copeland Whittaker Woese et al. Cavalier-Smith Ruggiero et al.
al. Smith
1735[23] 1866[24] 1925[25][26] 1938[27][28] 1969[29] 1977[30][31] 1993[33][34][35] 2015[39]
1990[32] 1998[36][37][38]
3 2
— — 2 empires 2 empires 2 empires 2 empires 3 domains 2 empires
superkingdoms superkingdoms
2 3 4 5
— 6 kingdoms — 8 kingdoms 6 kingdoms 7 kingdoms
kingdoms kingdoms kingdoms kingdoms
Eubacteria Bacteria Eubacteria Bacteria
Prokaryota Monera Monera Bacteria
Archaebacteria Archaea Archaebacteria Archaea
— Protista Archezoa
Protozoa Protozoa
Protista Protista Protista Protozoa
Chromista Chromista Chromista
Eukaryota Eucarya
Plantae Plantae Plantae Plantae Plantae
Vegetabilia Plantae Plantae
Fungi Fungi Fungi Fungi Fungi
Animalia Animalia Animalia Animalia Animalia Animalia Animalia Animalia
The kingdom-level classification of life is still widely employed as a useful way of grouping organisms, notwithstanding some problems with this approach:
Kingdoms such as Protozoa represent grades rather than clades, and so are rejected by phylogenetic classification systems.
The most recent research does not support the classification of the eukaryotes into any of the standard systems. In 2009, Andrew Roger and
Alastair Simpson emphasized the need for diligence in analyzing new discoveries: "With the current pace of change in our understanding of
the eukaryote tree of life, we should proceed with caution."[40] Kingdoms are rarely used in academic phylogeny and are more common in
introductory education, where 5-6 kingdom models are preferred.[41]
In 1990, the name "domain" was proposed for the highest rank.[5] This term represents
a synonym for the category of dominion (lat. dominium), introduced by Moore in
1974.[43] 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.[44]
Woese divided the prokaryotes (previously classified as the Kingdom Monera) into
two groups, called Eubacteria and Archaebacteria, stressing that there was as much
genetic difference between these two groups as between either of them and all A phylogenetic tree based on rRNA data showing Woese's three-domain
eukaryotes. system. All smaller branches can be considered kingdoms.
Eukaryotic supergroups
In 2004, a review article by Simpson and Roger noted that the Protista were "a grab-bag for all eukaryotes 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".[42]
On this basis, the diagram opposite (redrawn from their article) showed the real "kingdoms" (their quotation marks) of the eukaryotes.[42] 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".[45] The published classification deliberately did not use formal taxonomic
ranks, including that of "kingdom".
Archaeplastida (or Primoplantae) Land plants, green algae, red algae, and glaucophytes
In this system the multicellular animals (Metazoa) are descended from the same ancestor as
both the unicellular choanoflagellates and the fungi which form the Opisthokonta.[45] 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,[46] and a review in 2006 noted the lack
of evidence for several of the six proposed supergroups.[47]
As of 2019, there is widespread agreement that the Rhizaria belong with the Stramenopiles
and the Alveolata, in a clade dubbed the SAR supergroup,[48] so that Rhizaria is not one of
the main eukaryote groups.[20][49][50][51][52] One hypothesis of eukaryotic relationships depicted by Alastair
Simpson
Virusobiota (Viruses,
Acytota / Aphanobionta Viroids)
non-cellular life
Prionobiota (Prions)
Prokaryota / Bacteria Bacteria Eubacteria Bacteria
Procarya Monera
Biota / Vitae / Life (Monera) Archaea Archaea Archaebacteria
Cytota Protista
cellular life Archaea including
Fungi eukaryotes
Eukaryota / Eukarya
Plantae
Animalia
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.[58]
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 replication outside of a host cell.[59][60] Another argument is that their placement in the tree would be
problematic, since it is suspected that viruses have various evolutionary origins,[59] and they have a penchant for harvesting nucleotide sequences from their
hosts.
On the other hand, there are arguments in favor of their inclusion.[61] One of these comes from the discovery of unusually large and complex viruses, such as
Mimivirus, that possess typical cellular genes.[62]
See also
Biology portal
Cladistics
Phylogenetics
Systematics
Taxonomy
Notes
a. Compared to the version Cavalier-Smith published in 2004, the alveolates and the rhizarians have been moved from Kingdom Protozoa to
Kingdom Chromista.
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Further reading
Pelentier, B. (2007-2015). Empire Biota: a comprehensive taxonomy, [1] ([Link]
[Link]/). [Historical overview.]
Peter H. Raven and Helena Curtis (1970), Biology of Plants, New York: Worth Publishers. [Early presentation of five-kingdom system.]
External links
A Brief History of the Kingdoms of Life ([Link] at Earthling
Nature
The five kingdom concept ([Link] Archived ([Link]
[Link] 2021-11-07 at the Wayback Machine
Whittaker's classification ([Link]