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Although a phylum is often spoken of as if it were a hard and fast entity, no satisfactory definition of a phylum exists. Consequently the number of phyla varies from author to author. The relationship of phyla is increasingly well known, and larger clades can be erected to contain many of the phyla.
Informally, phyla can be thought of as grouping animals based on general body plan, developmental or internal organizations. For example, though seemingly divergent, spiders and crabs both belong to Arthropoda, whereas earthworms and tapeworms, similar in shape, are from Annelida and Platyhelminthes, respectively.
The best known animal phyla are the Mollusca, Porifera, Cnidaria, Platyhelminthes, Nematoda, Annelida, Arthropoda, Echinodermata, and Chordata, the phylum to which humans belong. Although there are approximately 35 phyla, these nine include over 96% of animal species. Many phyla are exclusively marine, and only one phylum, the Onychophora (velvet worms) is entirely absent from the world's oceans–although ancestral oncyophorans were marine.
The origin of phyla has traditionally been interpreted as a sudden and rapid event early in the Cambrian period, known as the Cambrian explosion. However, this interpretation stemmed from an incomplete knowledge of the fossil record, and a circular definition of a phylum; organisms of the time were mainly similar to, but not strictly members of, modern phyla. The significance of this event depends on (1) for how long modern phyla had existed prior to the Cambrian–fossil embryos and new interpretations of the Ediacara biota suggest that there may be a fair Precambrian root to the phyla; and (2) how soon phyla appeared in the Cambrian, a factor that depends on both the definition of a phylum and our interpretation of early fossils which may not display vital characteristics–for instance, non-mineralised parts of organisms rarely preserve.
The magnitude of the event was also overestimated as early authors felt it necessary to erect a new phylum for any organism that could not be accommodated in modern phyla. This approach is misleading and unhelpful; by one definition, such organisms do not fall into any phylum, but are classified as "aunts" of a phylum.
Defining a phylum
At the most basic level, 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 relatedness (the phylogenetic definition). Attempting to define a level of the Linnean hierarchy without referring to relatedness is an unsatisfactory approach, but the phenetic definition is more useful when addressing questions of a morphological nature–such as how successful different body plans were.
The largest objective measure in the above definitions is the "certain degree"–how unrelated do organisms need to be to be members of different phyla? Implicitly, the general definition is that they should be closely enough related that it is clear that they are more closely related to one another than to any other group. However, this too is problematic, as the definition is a function of our current knowledge about relationships. As more data becomes available, particularly from molecular studies, we are better able to judge the relationships between groups; therefore the phyla will eventually be merged as it becomes apparent that they are related to one another; for instance, the onychophora and tardigrada are now accepted as stem group arthropods; by the general definition, these three phyla should be combined.
This has led to calls for the concept of a phylum to be abandoned in favour of cladistics, a method in groups are placed on a "family tree" without any formal ranking of group size. So as to provide a handle on the size and significance of groups, a "body-plan" based definition of a phylum has been proposed by paleontologists Graham Budd an Sören Jensen. The definition was posited by paleontologists because it is extinct organisms that are typically hardest to classify, because they can be extinct off-shoots that diverged from a phylum's history before the characters that define the modern phylum were all acquired.
By Budd and Jensen's definition, phyla are defined by a set of characters shared by all their living representatives. This has a couple of small problems – for instance, characters common to most members of a phylum may be secondarily lost by some members. It is also defined 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 more major problem is that it relies on an objective decision of which group of organisms should be considered a phylum.
Its utility is that 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. Further, organisms in the stem group to a phylum can bear all the aspects of the "body plan" of the phylum without all the characters necessary to fall within it. This weakens the idea that each of the phyla represents a distinct body plan.
Based upon this definition, which some say is unreasonably affected by the chance survival of rare groups, which vastly increase the size of phyla, representatives of many modern phyla did not appear until long after the Cambrian–as late as the Carboniferous in the case of the Priapulids.
|Acanthocephala||Thorny head||Thorny-headed worms||Reversible spiny proboscis||about 750|
|Acoelomorpha||Without gut||Acoels||No mouth or alimentary canal|
|Annelida||Little ring||Segmented worms||Multiple circular segments||about 15,300 modern|
|Arthropoda||Jointed foot||Arthropods||Chitin exoskeleton||1,134,000+|
|Brachiopoda||Arm foot||Lamp shells||Lophophore and pedicle||between 300 and 500 extant|
|Bryozoa||Moss animals||Moss animals, sea mats||Lophophore, no pedicle, ciliated tentacles||about 5,000 living species|
|Chaetognatha||Longhair jaw||Arrow worms||Chitinous spines either side of head, fins||about 100 modern species|
|Chordata||Cord||Chordates||Hollow dorsal nervous chord, notochord, pharyngeal slits, endostyle, post-anal tail||about 100,000+|
|Cnidaria||Coelenterates||Nematocysts (stinging cells)||about 11,000|
|Ctenophora||Comb bearer||Comb jellies||Eight "comb rows" of fused cilia||about 100 modern species|
|Cycliophora||Wheel carrying||Symbion||Circular mouth surrounded by small cilia||at least 3|
|Echinodermata||Spiny skin||Echinoderms||Five-fold radial symmetry in living forms, mesodermal calcified spines||about 7,000 extant and 13,000 extinct species|
|Echiura||Spine tail||Spoon worms||Set of hooks at posterior end||about 140|
|Entoprocta||Inside anus||Goblet worm||Anus inside ring of cilia||about 150|
|Gastrotricha||Hair stomach||Meiofauna||Two terminal adhesive tubes||about 690|
|Gnathostomulida||Jaw orifice||Jaw worms||about 100|
|Hemichordata||Half cord||Acorn worms, pterobranchs||Stomochord in collar, pharyngeal slits||about 100 living species|
|Kinorhyncha||Motion snout||Mud dragons||Eleven segments, each with a dorsal plate||about 150|
|Loricifera||Corset bearer||Brush heads||Umbrella-like scales at each end||about 122|
|Micrognathozoa||Tiny jaw animals||—||Accordion like extensible thorax||1|
|Mollusca||Thin shell||Mollusks / molluscs||Muscular foot and mantle round shell||112,000|
|Nematoda||Thread like||Round worms||Round cross section, keratin cuticle||80 000 – 1 million|
|Nematomorpha||Thread form||Horsehair worms||about 320|
|Nemertea||A sea nymph||Ribbon worms||about 1200|
|Onychophora||Claw bearer||Velvet worms||Legs tipped by chitinous claws||about 200 modern|
|Orthonectida||Straight swim||Single layer of ciliated cells surrounding a mass of sex cells||about 20|
|Phoronida||Zeus' mistress||Horseshoe worms||U-shaped gut||20|
|Platyhelminthes||Flat worms||Flat worms||about 25,000|
|Porifera||Pore bearer||Sponges||Perforated interior wall||over 5,000 modern|
|Priapulida||Penis||Priapulid worms||Retractable proboscis surrounded by papillae||17|
|Rhombozoa||Lozenge animal||—||Single axial cell surrounded by ciliated cells||75|
|Rotifera||Wheel bearer||Rotifers||Anterior crown of cilia||about 2000|
|Sipuncula||Small tube||Peanut worms||Mouth surrounded by invertible tentacles||144–320|
|Tardigrada||Slow step||Water bears||Four segmented body and head||1,000+|
|Xenoturbellida||Strange flatworm||—||Ciliated deuterostome||2|
Groups formerly ranked as phyla
|Name as phylum||Common name||Current consensus|
|Aschelminthes||Pseudocoelomates||Divided into several pseudocoelomate phyla.|
|Craniata||—||Subgroup of phylum Chordata; perhaps synonymous with Vertebrata.|
|Cephalochordata||Lancelets||Subphylum of phylum Chordata.|
|Enterepneusta||Acorn worms||Class of phylum Hemichordata.|
|Gephyra||Peanut worms and spoon worms||Divided into phyla Sipuncula and Echiura.|
|Mesozoa||Mesozoans||Divided into phyla Orthonectida and Rhombozoa.|
|Myxozoa||—||Severely modified Cnidarians.|
|Pentastomida||Tongue worms||Subclass of Maxillopoda of phylum Arthropoda.|
|Pogonophora||Beard worms||Part of family Siboglinidae of phylum Annelida.|
|Pterobranchia||—||Class of phylum Hemichordata.|
|Symplasma||Glass sponges||Class Hexactinellida of phylum Porifera.|
|Urochordata||Tunicates||Subphylum of phylum Chordata.|
|Vestimentifera||Vent worms||Part of family Siboglinidae of phylum Annelida.|
|Division||Meaning||Common name||Distinguishing characteristics|
|Anthocerotophyta||Flower-horn plants||Hornworts||Horn-shaped sporophytes, no vascular system|
|Bryophyta||Moss plants||Mosses||Persistent unbranched sporophytes, no vascular system|
|Marchantiophyta||Marchantia plants||Liverworts||Ephemeral unbranched sporophytes, no vascular system|
|Lycopodiophyta||Wolf foot plants||Clubmosses & Spikemosses||Microphyll leaves, vascular system|
|Pteridophyta||Fern plants||Ferns & Horsetails||Prothallus gametophytes, vascular system|
|Pteridospermatophyta||Fern with seeds plant||Seed ferns||Only known from fossils, mostly Devonian, ranking in dispute|
|Pinophyta||Sap/pitch plants||Conifers||Cones containing seeds and wood composed of tracheids|
|Cycadophyta||Palm plants||Cycads||Seeds, crown of compound leaves|
|Ginkgophyta||Ginkgo plants||Ginkgo, Maidenhair||Seeds not protected by fruit (single species)|
|Gnetophyta||Gnetophytes||Seeds and woody vascular system with vessels|
|Anthophyta (or Magnoliophyta)||Flower plant||Flowering plants||Flowers and fruit, vascular system with vessels|
|Phylum||Meaning||Common name||Distinguishing characteristics|
|Chytridiomycota||Little pot mushroom||Chytrids||Cellulose in cell walls, flagellated gametes|
|Deuteromycota||Second mushroom||Imperfect fungi||Only reproduce asexually|
|Zygomycota||Yoke mushroom||Zygomycetes||Blend gametangia to form a zygosporangium|
|Glomeromycota||Ball mushroom||None||Form arbuscular mycorrhizae with plants|
|Ascomycota||Bag/Wineskin Mushroom||Sac fungi||Produce spores in an 'ascus'|
|Basidiomycota||Basidium Mushroom||Club Fungi||Produce spores from a 'basidium'|
- ↑ "Phylum" is adopted from the Greek φυλαί phylai, the clan-based voting groups in Greek city-states.
- ↑ (2005) "Life sciences" The American Heritage New Dictionary of Cultural Literacy, third edition, Houghton Mifflin Company. URL accessed 2008-10-04. "Phyla in the plant kingdom are frequently called divisions."
- ↑ Valentine, James W. (2004). On the Origin of Phyla, 7, Chicago: University Of Chicago Press."Classifications of organisms in hierarchical systems were in use by the seventeenth and eighteenth centuries. Usually organisms were grouped according to their morphological similarities as perceived by those early workers, and those groups were then grouped according to their similarities, and so on, to form a hierarchy."
- ↑ Parker, Andrew (2003). In the blink of an eye: How vision kick-started the big bang of evolution, 1–4, Sydney: Free Press."The job of an evolutionary biologist is to make sense of the conflicting diversity of form – there is not always a relationship between internal and external parts. Early in the history of the subject, it became obvious that internal organisations were generally more important to the higher classification of animals than are external shapes. The internal organisation puts general restrictions on how an animal can exchange gases, obtain nutrients and reproduce."
- ↑ Davidson, E. H.; Erwin, D. H. (2006), "Gene Regulatory Networks and the Evolution of Animal Body Plans", Science (American Association for the Advancement of Science) 311 (5762): 796–800, doi:10.1126/science.1113832, PMID 16469913, http://www.sciencemag.org/cgi/content/abstract/311/5762/796
- ↑ Template:Wonderful life
- ↑ 6.0 6.1 6.2 6.3 6.4 6.5 6.6 Budd, G.E., Jensen, S. (2000). A critical reappraisal of the fossil record of the bilaterian phyla. Biological Reviews 75 (02): 253–295.
- ↑ 7.0 7.1 Briggs, D. E. G.; Fortey, R. A. (2005), "Wonderful strife: systematics, stem groups, and the phylogenetic signal of the Cambrian radiation", Paleobiology 31 (2 (Suppl)): 94–112, doi:10.1666/0094-8373(2005)031[[0094:WSSSGA]]2.0.CO;2 (inactive 2008-10-21)
- ↑ Budd, G.E. (1998), Lethaia (Blackwell Synergy) 31 (3): 197–210, doi:10.1111/j.1502-3931.1998.tb00508.x (inactive 2008-10-21), http://www.blackwell-synergy.com/doi/abs/10.1111/j.1502-3931.1998.tb00508.x
- ↑ Feldkamp, S. (2002) Modern Biology. Holt, Rinehart, and Winston, USA. (pp. 725)
- ↑ Species Register Flatworms — Phylum Platyhelminthes. Marine Discovery Centres. URL accessed on 2007-04-09.
- ↑ "Kingdon Plantae Tree of Life".
- Are phyla "real"? Is there really a well-defined "number of animal phyla" extant and in the fossil record?
- Major Phyla Of Animals
- American Heritage Dictionary: New Latin phylum, from Greek phūlon, class.
- Online Etymological Dictionary: from Gk. phylon "race, stock," related to phyle "tribe, clan," and phylein "bring forth" of physikos "pertaining to nature," from physis "nature"
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