Phylum

The hierarchy of biological classification's eight major taxonomic ranks. A kingdom contains one or more phyla. Intermediate minor rankings are not shown.

In biology, a phylum (/ˈfaɪləm/; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature for algae, fungi, and plants accepts the terms as equivalent. Depending on definitions, the animal kingdom Animalia contains about 31 phyla, the plant kingdom Plantae contains about 14 phyla, and the fungus kingdom Fungi contains about 8 phyla. Current research in phylogenetics is uncovering the relationships between phyla, which are contained in larger clades, like Ecdysozoa and Embryophyta.

General description

The term phylum was coined in 1866 by Ernst Haeckel from the Greek phylon (φῦλον, "race, stock"), related to phyle (φυλή, "tribe, clan"). Haeckel noted that species constantly evolved into new species that seemed to retain few consistent features among themselves and therefore few features that distinguished them as a group ("a self-contained unity"). "Wohl aber ist eine solche reale und vollkommen abgeschlossene Einheit die Summe aller Species, welche aus einer und derselben gemeinschaftlichen Stammform allmählig sich entwickelt haben, wie z. B. alle Wirbelthiere. Diese Summe nennen wir Stamm (Phylon)." which translates as: However, perhaps such a real and completely self-contained unity is the aggregate of all species which have gradually evolved from one and the same common original form, as, for example, all vertebrates. We name this aggregate [a] Stamm [i.e., race] (Phylon). In plant taxonomy, August W. Eichler (1883) classified plants into five groups named divisions, a term that remains in use today for groups of plants, algae and fungi. The definitions of zoological phyla have changed from their origins in the six Linnaean classes and the four embranchements of Georges Cuvier.

Informally, phyla can be thought of as groupings of organisms based on general specialization of body plan. At its most basic, a phylum can be defined in two ways: as a group of organisms with a certain degree of morphological or developmental similarity (the phenetic definition), or a group of organisms with a certain degree of evolutionary relatedness (the phylogenetic definition). Attempting to define a level of the Linnean hierarchy without referring to (evolutionary) relatedness is unsatisfactory, but a phenetic definition is useful when addressing questions of a morphological nature—such as how successful different body plans were.

Definition based on genetic relation

The most important objective measure in the above definitions is the "certain degree" that defines how different organisms need to be members of different phyla. The minimal requirement is that all organisms in a phylum should be clearly more closely related to one another than to any other group. Even this is problematic because the requirement depends on knowledge of organisms' relationships: as more data become available, particularly from molecular studies, we are better able to determine the relationships between groups. So phyla can be merged or split if it becomes apparent that they are related to one another or not. For example, the bearded worms were described as a new phylum (the Pogonophora) in the middle of the 20th century, but molecular work almost half a century later found them to be a group of annelids, so the phyla were merged (the bearded worms are now an annelid family). On the other hand, the highly parasitic phylum Mesozoa was divided into two phyla (Orthonectida and Rhombozoa) when it was discovered the Orthonectida are probably deuterostomes and the Rhombozoa protostomes.

This changeability of phyla has led some biologists to call for the concept of a phylum to be abandoned in favour of placing taxa in clades without any formal ranking of group size.

Definition based on body plan

A definition of a phylum based on body plan has been proposed by paleontologists Graham Budd and Sören Jensen (as Haeckel had done a century earlier). The definition was posited because extinct organisms are hardest to classify: they can be offshoots that diverged from a phylum's line before the characters that define the modern phylum were all acquired. By Budd and Jensen's definition, a phylum is defined by a set of characters shared by all its living representatives.

This approach brings some small problems—for instance, ancestral characters common to most members of a phylum may have been lost by some members. Also, this definition is based on an arbitrary point of time: the present. However, as it is character based, it is easy to apply to the fossil record. A greater problem is that it relies on a subjective decision about which groups of organisms should be considered as phyla.

The approach is useful because it makes it easy to classify extinct organisms as "stem groups" to the phyla with which they bear the most resemblance, based only on the taxonomically important similarities. However, proving that a fossil belongs to the crown group of a phylum is difficult, as it must display a character unique to a sub-set of the crown group. Furthermore, organisms in the stem group of a phylum can possess the "body plan" of the phylum without all the characteristics necessary to fall within it. This weakens the idea that each of the phyla represents a distinct body plan.

A classification using this definition may be strongly affected by the chance survival of rare groups, which can make a phylum much more diverse than it would be otherwise.

Known phyla

Animals

Total numbers are estimates; figures from different authors vary wildly, not least because some are based on described species, some on extrapolations to numbers of undescribed species. For instance, around 25,000–27,000 species of nematodes have been described, while published estimates of the total number of nematode species include 10,000–20,000; 500,000; 10 million; and 100 million.

Plants

The kingdom Plantae is defined in various ways by different biologists (see Current definitions of Plantae). All definitions include the living embryophytes (land plants), to which may be added the two green algae divisions, Chlorophyta and Charophyta, to form the clade Viridiplantae. The table below follows the influential (though contentious) Cavalier-Smith system in equating "Plantae" with Archaeplastida, a group containing Viridiplantae and the algal Rhodophyta and Glaucophyta divisions.

The definition and classification of plants at the division level also varies from source to source, and has changed progressively in recent years. Thus some sources place horsetails in division Arthrophyta and ferns in division Monilophyta, while others place them both in Monilophyta, as shown below. The division Pinophyta may be used for all gymnosperms (i.e. including cycads, ginkgos and gnetophytes), or for conifers alone as below.

Since the first publication of the APG system in 1998, which proposed a classification of angiosperms up to the level of orders, many sources have preferred to treat ranks higher than orders as informal clades. Where formal ranks have been provided, the traditional divisions listed below have been reduced to a very much lower level, e.g. subclasses.

Fungi

Phylum Microsporidia is generally included in kingdom Fungi, though its exact relations remain uncertain, and it is considered a protozoan by the International Society of Protistologists (see Protista, below). Molecular analysis of Zygomycota has found it to be polyphyletic (its members do not share an immediate ancestor), which is considered undesirable by many biologists. Accordingly, there is a proposal to abolish the Zygomycota phylum. Its members would be divided between phylum Glomeromycota and four new subphyla incertae sedis (of uncertain placement): Entomophthoromycotina, Kickxellomycotina, Mucoromycotina, and Zoopagomycotina.

Protista

Kingdom Protista (or Protoctista) is included in the traditional five- or six-kingdom model, where it can be defined as containing all eukaryotes that are not plants, animals, or fungi. Protista is a paraphyletic taxon, which is less acceptable to present-day biologists than in the past. Proposals have been made to divide it among several new kingdoms, such as Protozoa and Chromista in the Cavalier-Smith system.

Protist taxonomy has long been unstable, with different approaches and definitions resulting in many competing classification schemes. The phyla listed here are used for Chromista and Protozoa by the Catalogue of Life, adapted from the system used by the International Society of Protistologists (ISP), with updates from the latest (2022) publication by Cavalier-Smith. Some of the descriptions are based on the 2019 revision of eukaryotes by the ISP.

Phylum	Meaning	Common name	Distinguishing characteristics	Example	Species	Image
Amoebozoa	Amorphous animal	Amoebas, amoebozoans	Presence of pseudopodia for amoeboid movement, tubular cristae.	Amoeba	2400
Apusozoa (P)			Gliding biciliates with two or three connectors between centrioles	Podomonas	32
Bigyra	Two rings		Stramenopiles with a double helix in ciliary transition zone	Aplanochytrium
Cercozoa	Flagellated animal		Widely diverse morphological groups (flagellates, amoebae, heliozoans...) united only by phylogeny	Euglypha
Choanozoa (P)	Funnel animal		Filose pseudopods; some with a colar of microvilli surrounding a flagellum	Desmarella	125
Ciliophora	Cilia bearer	Ciliates	Presence of multiple cilia and a cytostome	Paramecium	4,500
Cryptista	Hidden		Different morphological groups (Palpitomonas, cryptophytes, endoheleans...) united by phylogeny	Rhodomonas
Eolouka (P)	Early groove		Biciliates with ventral feeding groove	Stygiella
Euglenozoa	True eye animal		One of the two cilia inserted into an apical or subapical ciliary pocket	Euglena	800
Gyrista	Ring		Helical or double helix system in ciliary transition zone; contains ochrophytes and pseudofungi	Diatoms	1,500+
Haptista			Thin microtubule-based haptonema or axopodia for feeding, complex mineralized scales	Coccolithus
Malawimonada			Small free-living bicilates with two kinetosomes, one or two vanes in posterior cilium	Malawimonas
Metamonada	Middle monads		anaerobic or microaerophilic, some without mitochondria; four kinetosomes per kinetid	Giardia
Miozoa	Suckling animal		One-step meiosis, non-tubular ciliary hairs, usually haploid	Dinoflagellates	1,555+
Opisthosporidia (now part of fungi)	Opisthokont spores		Parasites with chitinous spores and extrusive host-invasion apparatus	Microsporidia
Percolozoa			Complex life cycle containing amoebae, flagellates and cysts	Naegleria
Retaria			Reticulopodia or axopodia, and usually various types of skeleton	Foraminifera	10,000 (50,000 extinct)
Total: 17

The Catalogue of Life includes Rhodophyta and Glaucophyta in kingdom Plantae, but other systems consider these phyla part of Protista.

Bacteria

Currently there are 40 bacterial phyla (not including "Cyanobacteria") that have been validly published according to the Bacteriological Code

1. Acidobacteriota, phenotypically diverse and mostly uncultured
2. Actinomycetota, High-G+C Gram positive species
3. Aquificota, deep-branching
4. Armatimonadota
5. Atribacterota
6. Bacillota, Low-G+C Gram positive species, such as the spore-formers Bacilli (aerobic) and Clostridia (anaerobic)
7. Bacteroidota
8. Balneolota
9. Bdellovibrionota
10. Caldisericota, formerly candidate division OP5, Caldisericum exile is the sole representative
11. Calditrichota
12. Campylobacterota
13. Chlamydiota
14. Chlorobiota, green sulphur bacteria
15. Chloroflexota, green non-sulphur bacteria
16. Chrysiogenota, only 3 genera (Chrysiogenes arsenatis, Desulfurispira natronophila, Desulfurispirillum alkaliphilum)
17. Coprothermobacterota
18. Deferribacterota
19. Deinococcota, Deinococcus radiodurans and Thermus aquaticus are "commonly known" species of this phyla
20. Dictyoglomota
21. Elusimicrobiota, formerly candidate division Thermite Group 1
22. Fibrobacterota
23. Fusobacteriota
24. Gemmatimonadota
25. Ignavibacteriota
26. Kiritimatiellota
27. Lentisphaerota, formerly clade VadinBE97
28. Mycoplasmatota, notable genus: Mycoplasma
29. Myxococcota
30. Nitrospinota
31. Nitrospirota
32. Planctomycetota
33. Pseudomonadota, the most well-known phylum, containing species such as Escherichia coli or Pseudomonas aeruginosa
34. Rhodothermota
35. Spirochaetota, species include Borrelia burgdorferi, which causes Lyme disease
36. Synergistota
37. Thermodesulfobacteriota
38. Thermomicrobiota
39. Thermotogota, deep-branching
40. Verrucomicrobiota

Archaea

Currently there are 2 phyla that have been validly published according to the Bacteriological Code

1. Nitrososphaerota
2. Thermoproteota, second most common archaeal phylum

Other phyla that have been proposed, but not validly named, include:

1. "Euryarchaeota", most common archaeal phylum
2. "Korarchaeota"
3. "Nanoarchaeota", ultra-small symbiotes, single known species

See also

• Cladistics
• Phylogenetics
• Systematics
• Taxonomy

External links

Wikispecies has information related to Phylum.

Look up Phylum in Wiktionary, the free dictionary.

• Are phyla "real"? Is there really a well-defined "number of animal phyla" extant and in the fossil record?
• Major Phyla Of Animals Archived 16 July 2006 at the Wayback Machine