Phylum

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.[1][2][3] Depending on definitions, the animal kingdom Animalia or Metazoa contains approximately 35 phyla; the plant kingdom Plantae contains about 14, 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.

For other uses, see Phyla.

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

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").[4][5] 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.[1][6] The definitions of zoological phyla have changed from their origins in the six Linnaean classes and the four embranchements of Georges Cuvier.[7]

Informally, phyla can be thought of as groupings of organisms based on general specialization of body plan.[8] 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).[9] 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.[9] 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).[10] 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.[11]

This changeability of phyla has led some biologists to call for the concept of a phylum to be abandoned in favour of cladistics, a method in which groups are placed on a "family tree" without any formal ranking of group size.[9]

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.[9] 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.[9] 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.[12]

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.[13]

Known phyla

Animals
Main article: Animal

Total numbers are estimates; figures from different authors vary wildly, not least because some are based on described species,[14] 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.[15]

Protostome Bilateria
Deuterostome
Basal/disputed
Others
PhylumMeaningCommon nameDistinguishing characteristicSpecies described
Acanthocephala Thorny head Thorny-headed worms[16]:278 Reversible spiny proboscis that bears many rows of hooked spines 1,420
Annelida Little ring [16]:306 Segmented worms Multiple circular segment 17,000 + extant
Arthropoda Jointed foot Arthropods Segmented bodies and jointed limbs, with Chitin exoskeleton 1,250,000+ extant;[14] 20,000+ extinct
Brachiopoda Arm foot[16]:336 Lampshells[16]:336 Lophophore and pedicle 300-500 extant; 12,000+ extinct
Bryozoa Moss animals Moss animals, sea mats, ectoprocts[16]:332 Lophophore, no pedicle, ciliated tentacles, anus outside ring of cilia 6,000 extant[14]
Chaetognatha Longhair jaw Arrow worms[16]:342 Chitinous spines either side of head, fins approx. 100 extant
Chordata With a cord Chordates Hollow dorsal nerve cord, notochord, pharyngeal slits, endostyle, post-anal tail approx. 55,000+[14]
Cnidaria Stinging nettle Cnidarians Nematocysts (stinging cells) approx. 16,000[14]
Ctenophora Comb bearer Comb jellies[16]:256 Eight "comb rows" of fused cilia approx. 100-150 extant
Cycliophora Wheel carrying Symbion Circular mouth surrounded by small cilia, sac-like bodies 3+
Echinodermata Spiny skin Echinoderms[16]:348 Fivefold radial symmetry in living forms, mesodermal calcified spines approx. 7,500 extant;[14] approx. 13,000 extinct
Entoprocta Inside anus[16]:292 Goblet worms Anus inside ring of cilia approx. 150
Gastrotricha Hairy stomach[16]:288 Gastrotrich worms Two terminal adhesive tubes approx. 690
Gnathostomulida Jaw orifice Jaw worms[16]:260 approx. 100
Hemichordata Half cord[16]:344 Acorn worms, hemichordates Stomochord in collar, pharyngeal slits approx. 130 extant
Kinorhyncha Motion snout Mud dragons Eleven segments, each with a dorsal plate approx. 150
Loricifera Corset bearer Brush heads Umbrella-like scales at each end approx. 122
Micrognathozoa Tiny jaw animals Limnognathia Accordion-like extensible thorax 1
Mollusca Soft[16]:320 Mollusks / molluscs Muscular foot and mantle round shell 85,000+ extant;[14] 80,000+ extinct[17]
Nematoda Thread like Round worms, thread worms[16]:274 Round cross section, keratin cuticle 25,000[14]
Nematomorpha Thread form[16]:276 Horsehair worms, gordian worms[16]:276 approx. 320
Nemertea A sea nymph[16]:270 Ribbon worms, rhynchocoela[16]:270 approx. 1,200
Onychophora Claw bearer Velvet worms[16]:328 Legs tipped by chitinous claws approx. 200 extant
Orthonectida Straight swimming[16]:268 Orthonectids[16]:268 Single layer of ciliated cells surrounding a mass of sex cells approx. 26
Phoronida Zeus's mistress Horseshoe worms U-shaped gut 11
Placozoa Plate animals Trichoplaxes[16]:242 Differentiated top and bottom surfaces, two ciliated cell layers, amoeboid fiber cells in between 3
Platyhelminthes Flat worm[16]:262 Flatworms[16]:262 approx. 29,500[14]
Porifera [lower-alpha 1] Pore bearer Sponges[16]:246 Perforated interior wall 10,800 extant[14]
Priapulida Little Priapus Penis worms approx. 20
Rhombozoa Lozenge animal Rhombozoans[16]:264 Single anteroposterior axial cell surrounded by ciliated cells 100+
Rotifera Wheel bearer Rotifers[16]:282 Anterior crown of cilia approx. 2,000[14]
Sipuncula Small tube Peanut worms Mouth surrounded by invertible tentacles 144-320
Tardigrada Slow step Water bears, Moss piglets Four segmented body and head 1,000
Xenacoelomorpha Strange hollow form Acoels, xenoturbellids Bilaterian, but lacking typical bilaterian structures such as gut cavities, anuses, and circulatory systems[18] 400+
Total: 34 1,525,000[14]

Plants
Main article: Plant

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,[19] 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 Pteridophyta,[20] while others place them both in Pteridophyta, as shown below. The division Pinophyta may be used for all gymnosperms (i.e. including cycads, ginkgos and gnetophytes),[21] 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.[22]

Land plants Viridiplantae
Green algae
Other algae (Biliphyta)[19]
DivisionMeaningCommon nameDistinguishing characteristicsSpecies described
Anthocerotophyta[23] Anthoceros-like plants Hornworts Horn-shaped sporophytes, no vascular system 100-300+
Bryophyta[23] Bryum-like plants, moss plants Mosses Persistent unbranched sporophytes, no vascular system approx. 12,000
Charophyta Chara-like plants Charophytes approx. 1,000
Chlorophyta (Yellow-)green plants[16]:200 Chlorophytes approx. 7,000
Cycadophyta[24] Cycas-like plants, palm-like plants Cycads Seeds, crown of compound leaves approx. 100-200
Ginkgophyta[25] Ginkgo-like plants Ginkgo, maidenhair tree Seeds not protected by fruit (single living species) only 1 extant; 50+ extinct
Glaucophyta Blue-green plants Glaucophytes 15
Gnetophyta[26] Gnetum-like plants Gnetophytes Seeds and woody vascular system with vessels approx. 70
Lycopodiophyta,[21]

Lycophyta[27]

 Lycopodium-like plants

Wolf plants

 Clubmosses & spikemosses Microphyll leaves, vascular system 1,290 extant
Magnoliophyta Magnolia-like plants Flowering plants, angiosperms Flowers and fruit, vascular system with vessels 300,000
Marchantiophyta,[28]

Hepatophyta[23]

 Marchantia-like plants

Liver plants

 Liverworts Ephemeral unbranched sporophytes, no vascular system approx. 9,000
Polypodiopsida Ferns approx. 10,560
Pinophyta,[21]

Coniferophyta[29]

 Pinus-like plants

Cone-bearing plant

 Conifers Cones containing seeds and wood composed of tracheids 629 extant
Rhodophyta Rose plants Red algae Use phycobiliproteins as accessory pigments. approx. 7,000
Total: 13

Fungi
Main article: Fungi
DivisionMeaningCommon nameDistinguishing characteristicsSpecies described
Ascomycota Bladder fungus[16]:396 Ascomycetes,[16]:396 sac fungi Tend to have fruiting bodies (ascocarp).[30] Filamentous, producing hyphae separated by septa. Can reproduce asexually.[31]
Basidiomycota Small base fungus[16]:402 Basidiomycetes[16]:402 Bracket fungi, toadstools, smuts and rust. Sexual reproduction.[32]
Blastocladiomycota Offshoot branch fungus[33] Blastoclads
Chytridiomycota Little cooking pot fungus[34] Chytrids Predominantly Aquatic saprotrophic or parasitic. Have a posterior flagellum. Tend to be single celled but can also be multicellular.[35][36][37]
Glomeromycota Ball of yarn fungus[16]:394 Glomeromycetes, AM fungi[16]:394 Mainly arbuscular mycorrhizae present, terrestrial with a small presence on wetlands. Reproduction is asexual but requires plant roots.[32]
Microsporidia Small seeds[38] Microsporans[16]:390
Neocallimastigomycota New beautiful whip fungus[39] Neocallimastigomycetes Predominantly located in digestive tract of herbivorus animals. Anaerobic, terrestrial and aquatic.[40]
Zygomycota Pair fungus[16]:392 Zygomycetes[16]:392 Most are saprobes and reproduce sexually and asexually.[40]
Total: 8

Phylum Microsporidia is generally included in kingdom Fungi, though its exact relations remain uncertain,[41] and it is considered a protozoan by the International Society of Protistologists[42] (see Protista, below). Molecular analysis of Zygomycota has found it to be polyphyletic (its members do not share an immediate ancestor),[43] 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.[41]

Protista
Main article: Protista taxonomy

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.[16]:120 Protista is a polyphyletic taxon,[44] 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.[45]

Protist taxonomy has long been unstable,[46] 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,[47] adapted from the system used by the International Society of Protistologists.[42]

Harosa
Protozoa
Phylum/DivisionMeaningCommon nameDistinguishing characteristicsExampleSpecies described
Amoebozoa Amorphous animal Amoebas Amoeba 2400
Bigyra Two ring
Cercozoa
Choanozoa Funnel animal 125
Ciliophora Cilia bearer Ciliates Paramecium 4500
Cryptista
Euglenozoa True eye animal Euglena 800
Foraminifera Hole bearers Forams Complex shells with one or more chambers Forams 10000, 50000 extinct
Haptophyta
Loukozoa Groove animal
Metamonada Giardia
Microsporidia Small spore
Myzozoa Suckling animal 1555+
Ochrophyta Yellow plant Diatoms
Oomycota Egg fungus[16]:184 Oomycetes
Percolozoa
Radiozoa Ray animal Radiolarians
Sarcomastigophora
Sulcozoa
Total: 20

The Catalogue of Life includes Rhodophyta and Glaucophyta in kingdom Plantae,[47] but other systems consider these phyla part of Protista.[48]

Bacteria
Main article: Bacterial phyla

Currently there are 29 phyla accepted by List of Prokaryotic names with Standing in Nomenclature (LPSN)[49]

  1. Acidobacteria, phenotypically diverse and mostly uncultured
  2. Actinobacteria, High-G+C Gram positive species
  3. Aquificae, only 14 thermophilic genera, deep branching
  4. Armatimonadetes
  5. Bacteroidetes
  6. Caldiserica, formerly candidate division OP5, Caldisericum exile is the sole representative
  7. Chlamydiae, only 6 genera
  8. Chlorobi, only 7 genera, green sulphur bacteria
  9. Chloroflexi, green non-sulphur bacteria
  10. Chrysiogenetes, only 3 genera (Chrysiogenes arsenatis, Desulfurispira natronophila, Desulfurispirillum alkaliphilum)
  11. Cyanobacteria, also known as the blue-green algae
  12. Deferribacteres
  13. Deinococcus-Thermus, Deinococcus radiodurans and Thermus aquaticus are "commonly known" species of this phyla
  14. Dictyoglomi
  15. Elusimicrobia, formerly candidate division Thermite Group 1
  16. Fibrobacteres
  17. Firmicutes, Low-G+C Gram positive species, such as the spore-formers Bacilli (aerobic) and Clostridia (anaerobic)
  18. Fusobacteria
  19. Gemmatimonadetes
  20. Lentisphaerae, formerly clade VadinBE97
  21. Nitrospira
  22. Planctomycetes
  23. Proteobacteria, the most known phyla, containing species such as Escherichia coli or Pseudomonas aeruginosa
  24. Spirochaetes, species include Borrelia burgdorferi, which causes Lyme disease
  25. Synergistetes
  26. Tenericutes, alternatively class Mollicutes in phylum Firmicutes (notable genus: Mycoplasma)
  27. Thermodesulfobacteria
  28. Thermotogae, deep branching
  29. Verrucomicrobia

Archaea
Main article: Archaea

Currently there are five phyla accepted by List of Prokaryotic names with Standing in Nomenclature (LPSN).[49]

  1. Crenarchaeota, second most common archaeal phylum
  2. Euryarchaeota, most common archaeal phylum
  3. Korarchaeota
  4. Nanoarchaeota, ultra-small symbiotes, single known species
  5. Thaumarchaeota

See also

Notes
  1. Paraphyletic

References
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