1. Brusca And Brusca Invertebrates

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Transmissionelectron micrograph of mesenchyme displaying the ultrastructure of a typical cell and matrix.
Carnegie stage6b
PrecursorPrimarily mesoderm
Anatomical terminology
[edit on Wikidata]

Mesenchyme, in vertebrate embryology, is a type of connective tissue found mostly during the development of the embryo.[1] It is composed mainly of ground substance with few cells or fibers. It can also refer to a group of mucoproteins found in certain types of cysts (etc.), resembling mucus. It is most easily found as a component of Wharton's jelly.

The vitreous of the eyeball is a similar tissue.[2]

In invertebrate zoology, the term refers to free cells loosely arranged in a matrix.[3]

  • 2In vertebrate embryology
    • 2.2Development


'Mesenchyme' is a term introduced by Oscar Hertwig in 1881.[3]

In order to differentiate the use of the word mesenchyme in invertebrate zoology (an ecto- or entomesodermal middle layer of some invertebrates) and the use in vertebrate embryology (that is, undifferentiated tissue found in embryonic true mesoderm - entomesoderm - from which all connective tissues like blood vessels, blood cells, the lymphatic system, and the heart are derived.), some authors prefer to use the term mesoglea (in wider sense) in lieu of mesenchyme when referring to the middle layers of sponges and diploblasts, reserving the term mesenchyme for the embryological sense. However, Brusca & Brusca discourage this usage, using mesoglea in its strict sense (noncellular mesenchyme), and preferring to maintain both the embryological and zoological senses for the term mesenchyme.[4]

Finally, some similar terms used in botany generally are differentiated by the suffix 'a': mesenchyma (a tissue between xylem and phloem in roots), collenchyma (primordial leaf tissues) and parenchyma (supportive tissues).[4]

In vertebrate embryology[edit]


Mesenchyme is characterized morphologically by a prominent ground substancematrix containing a loose aggregate of reticular fibrils and unspecialized cells.[5] Mesenchymal cells can migrate easily, in contrast to epithelial cells, which lack mobility and are organized into closely adherent sheets, and are polarized in an apical-basal orientation.

Brusca And Brusca Invertebrates Pdf Creator


The mesenchyme originates from the mesoderm.[6] From the mesoderm, the mesenchyme appears as an embryologically primitive 'soup'. This 'soup' exists as a combination of the mesenchymal cells plus serous fluid plus the many different tissue proteins. Serous fluid is typically stocked with the many serous elements, such as sodium and chloride. The mesenchyme develops into the tissues of the lymphatic and circulatory systems, as well as the musculoskeletal system. This latter system is characterized as connective tissues throughout the body, such as bone, muscle and cartilage. A malignant cancer of mesenchymal cells is a type of sarcoma.[7][8]

Epithelial to mesenchymal transition[edit]

The first emergence of mesenchyme occurs during gastrulation from the epithelial–mesenchymal transition (EMT) process. This transition occurs through the loss of epithelial cadherin, tight junctions, and adherens junctions on the cell membranes of epithelial cells.[9] The surface molecules undergo endocytosis and the microtubule cytoskeleton loses shape, enabling mesenchyme to migrate along the extracellular matrix (ECM). Epithelial–mesenchymal transition occurs in embryonic cells that require migration through or over tissue, and can be followed with a mesenchymal–epithelial transition to produce secondary epithelial tissues. Embryological mesenchymal cells expresses fibroblast-specific protein (Fsp1), which is indicative of their shared properties with the migratory adult fibroblasts, and c-Fos, an oncogene associated with the down-regulation of epithelial cadherin.[10][11] Both formation of the primitive streak and mesenchymal tissue is dependent on the Wnt/β-catenin pathway.[12] Specific markers of mesenchymal tissue include the additional expression of ECM factors such as fibronectin and vitronectin.[13]


The first cells of the embryo to undergo EMT and form mesenchyme are the extra-embryonic cells of the trophectoderm. These migrate from the body of the blastocyst into the endometrial layer of the uterus in order to contribute to the formation of the anchored placenta.[14]

Primary mesenchyme[edit]

Primary mesenchyme is the first embryonic mesenchymal tissue to emerge, and it is produced from EMT in epiblast cells. In the epiblast, it is induced by the primitive streak through Wnt signaling, and produces endoderm and mesoderm from a transitory tissue called mesendoderm during the process of gastrulation.[15]

The formation of primary mesenchyme depends on the expression of WNT3. Other deficiencies in signaling pathways, such as in Nodal (a TGF-beta protein), will lead to defective mesoderm formation.[9]

The tissue layers formed from the primitive streak invaginate together into the embryo and the induced mesenchymal stem cells will ingress and form the mesoderm. Mesodermal tissue will continue to differentiate and/or migrate throughout the embryo to ultimately form most connective tissue layers of the body.[16]

Neural mesenchyme[edit]

Embryological mesenchyme is particularly transitory and soon differentiates after migration. Neural mesenchyme forms soon after primary mesenchyme formation.[17]

The interaction with ectoderm and somite-forming morphogenic factors cause some primary mesenchyme to form neural mesenchyme, or paraxial mesoderm, and contribute to somite formation. Neural mesenchyme soon undergoes a mesenchymal–epithelial transition under the influence of WNT6 produced by ectoderm to form somites.[18] These structures will undergo a secondary EMT as the somite tissue migrates later in development to form structural connective tissue such as cartilage and skeletal muscle.[19]

Deer hunter 3 download full game. Neural crest cells (NCCs) form from neuroectoderm, instead of the primary mesenchyme, from morphogenic signals of the neural crest. The EMT occurs as a result of Wnt signaling, the influence of Sox genes and the loss of E-cadherin from the cell surface. NCCs additionally require the repression of N-cadherin, and neural cell adhesion molecule. NCCs ingress into the embryo from the epithelial neuroectodermal layer and migrate throughout the body in order form multiple peripheral nervous system (PNS) cells and melanocytes. Migration of NCCs is primarily induced by BMP signaling and its inhibitor, Noggin.[20][21]

In invertebrate zoology[edit]

In some invertebrates, e.g., Porifera, Cnidaria, Ctenophora and some triploblasts (the acoelomates), the term 'mesenchyme' refers to a more-or-less solid but looselly organized tissue consisting of a gel matrix (the mesoglea) with various cellular and fibrous inclusions, located between epidermis and gastrodermis. In some cases, the mesoglea is noncellular.[22]

  • In sponges, the mesenchyme is called mesohyl.[4]
  • In diploblasts (Cnidaria and Ctenophora), the mesenchyme is fully ectodermally derived. This kind of mesenchyme is called ectomesodermal, and is not considered true mesoderm.
  • In triploblastic acoelomates (e.g., flatworms), the term parenchyme is sometimes used for the middle (mesenchymal) layer, in which the dense layer includes tissues derived from both ecto- and entomesoderm (true mesoderm, derived from entoderm).

When cellular material is sparse or densely packed, the mesenchyme may be sometimes called collenchyme (e.g., cnidarians) or parenchyme (e.g., flatworms), respectively.[4] When no cellular material is present (e.g., in Hydrozoa), the layer is properly called mesoglea.[4]

In some colonial cnidarians, the mesenchyme is perforated by gastrovascular channels continuous among colony members. This entire matrix of common basal material is called coenenchyme.[4]


  1. ^'Blue Histology - Connective Tissues'. Retrieved 2008-12-05.
  2. ^Singh, Inderbir (April 15, 2007). Textbook of Human Histology With Colour Atlas. Jaypee Brothers Medical Publishers.[page needed]. ISBN978-81-7179-967-1.
  3. ^ abCollins, P.; Billett, F. S. (1995). 'The terminology of early development: History, concepts, and current usage'. Clinical Anatomy. 8 (6): 418–425. doi:10.1002/ca.980080610. PMID8713164.
  4. ^ abcdefBrusca, R.C.; Brusca, G.J. (2003). Invertebrates (2nd ed.). Sunderland, Massachusetts. p. 220.
  5. ^Mesenchymal tissue
  6. ^Kierszenbaum, Abraham L.; Tres, Laura (2015). Histology and Cell Biology: An Introduction to Pathology E-Book (4 ed.). Elsevier Health Sciences. p. 123. ISBN9780323313353.
  7. ^Strum, Judy M.; Gartner, Leslie P.; Hiatt, James L. (2007). Cell biology and histology. Hagerstown, MD: Lippincott Williams & Wilkins. p. 83. ISBN978-0-7817-8577-8.
  8. ^Sadler, T.W. (2006). Langman's Medical Embryology. Lippincott Williams & Wilkins. pp. 68–70. ISBN978-0-7817-9485-5.
  9. ^ abKalluri, Raghu; Weinberg, Robert A. (2009). 'The basics of epithelial-mesenchymal transition'. Journal of Clinical Investigation. 119 (6): 1420–8. doi:10.1172/JCI39104. PMC2689101. PMID19487818.
  10. ^Okada, H; Danoff, T. M.; Kalluri, R; Neilson, E. G. (1997). 'Early role of Fsp1 in epithelial-mesenchymal transformation'. The American Journal of Physiology. 273 (4 Pt 2): F563–74. PMID9362334.
  11. ^Eger, A; Stockinger, A; Schaffhauser, B; Beug, H; Foisner, R (2000). 'Epithelial mesenchymal transition by c-Fos estrogen receptor activation involves nuclear translocation of beta-catenin and upregulation of beta-catenin/lymphoid enhancer binding factor-1 transcriptional activity'. The Journal of Cell Biology. 148 (1): 173–88. doi:10.1083/jcb.148.1.173. PMC3207144. PMID10629227.
  12. ^Mohamed, O. A.; Clarke, H. J.; Dufort, D (2004). 'Beta-catenin signaling marks the prospective site of primitive streak formation in the mouse embryo'. Developmental Dynamics. 231 (2): 416–24. doi:10.1002/dvdy.20135. PMID15366019.
  13. ^Thiery, J. P.; Sleeman, J. P. (2006). 'Complex networks orchestrate epithelial-mesenchymal transitions'. Nature Reviews Molecular Cell Biology. 7 (2): 131–42. doi:10.1038/nrm1835. PMID16493418.
  14. ^Yamakoshi, S; Bai, R; Chaen, T; Ideta, A; Aoyagi, Y; Sakurai, T; Konno, T; Imakawa, K (2012). 'Expression of mesenchymal-related genes by the bovine trophectoderm following conceptus attachment to the endometrial epithelium'. Reproduction (Cambridge, England). 143 (3): 377–87. doi:10.1530/REP-11-0364. PMID22157247.
  15. ^Bellairs, R (1986). 'The primitive streak'. Anatomy and Embryology. 174 (1): 1–14. doi:10.1007/bf00318331. PMID3518538.
  16. ^Hay, E. D. (2005). 'The mesenchymal cell, its role in the embryo, and the remarkable signaling mechanisms that create it'. Developmental Dynamics. 233 (3): 706–20. doi:10.1002/dvdy.20345. PMID15937929.
  17. ^Mareschi, K; Novara, M; Rustichelli, D; Ferrero, I; Guido, D; Carbone, E; Medico, E; Madon, E; Vercelli, A; Fagioli, F (2006). 'Neural differentiation of human mesenchymal stem cells: Evidence for expression of neural markers and eag K+ channel types'. Experimental Hematology. 34 (11): 1563–72. doi:10.1016/j.exphem.2006.06.020. PMID17046576.
  18. ^Schmidt, C; Stoeckelhuber, M; McKinnell, I; Putz, R; Christ, B; Patel, K (2004). 'Wnt 6 regulates the epithelialisation process of the segmental plate mesoderm leading to somite formation'. Developmental Biology. 271 (1): 198–209. doi:10.1016/j.ydbio.2004.03.016. PMID15196961.
  19. ^Stockdale, F. E.; Nikovits Jr, W; Christ, B (2000). 'Molecular and cellular biology of avian somite development'. Developmental Dynamics. 219 (3): 304–21. doi:10.1002/1097-0177(2000)9999:9999<::AID-DVDY1057>3.0.CO;2-5. PMID11066088.
  20. ^Bronner-Fraser, M (1994). 'Neural crest cell formation and migration in the developing embryo'. FASEB Journal. 8 (10): 699–706. doi:10.1096/fasebj.8.10.8050668. PMID8050668.
  21. ^Trainor, P. A. (2005). 'Specification of neural crest cell formation and migration in mouse embryos'. Seminars in Cell & Developmental Biology. 16 (6): 683–93. doi:10.1016/j.semcdb.2005.06.007. PMID16043371.
  22. ^Brusca, R.C.; Brusca, G.J. (2003). Invertebrates (2nd ed.). Sunderland, Massachusetts. p. 101.

External links[edit]

Look up mesenchyme in Wiktionary, the free dictionary.
Retrieved from 'https://en.wikipedia.org/w/index.php?title=Mesenchyme&oldid=914702214'

The taxonomy of invertebrates as proposed by Richard C. Brusca and Gary J. Brusca in 2003 is a system of classification with emphasis on the invertebrates, in other words, a way to classify animals, primarily those which have no backbone.[1]

  • 2Eukaryotes (Eukaryota, or Eukarya)
    • 2.1Kingdom Animalia (Metazoa)
      • 2.1.1Parazoa
      • 2.1.2Mesozoa
      • 2.1.3Eumetazoa


  • Kingdom Eubacteria (Bacteria)
  • Kingdom Archaea (Archaebacteria)

Eukaryotes (Eukaryota, or Eukarya)[edit]

  • Kingdom Fungi
  • Kingdom Plantae (= Metaphyta)
  • Kingdom Protista
  • Phylum Euglenida
  • Phylum Kinetoplastida
  • Alveolata phyla
  • Phylum Ciliophora
  • Phylum Apicomplexa
  • Phylum Dinoflagellata
  • Phylum Stramenopila
  • Phylum Rhizopoda
  • Phylum Actinopoda
  • Phylum Granuloreticulosa
  • Phylum Diplomonadida
  • Phylum Parabasilida
  • Phylum Cryptomonada
  • Phylum Microspora
  • Phylum Ascetospora
  • Phylum Choanoflagellata
  • Phylum Chlorophyta
  • Phylum Opalinida
  • Incertae sedis: Genus Stephanopogon

Kingdom Animalia (Metazoa)[edit]


Phylum Porifera[edit]
  • Phylum Porifera
  • Class Calcarea
  • Subclass Calcinea
  • Subclass Calcaronea
  • Class Hexactinellida
  • Subclass Amphidiscophora
  • Subclass Hexasterophora
  • Class Demospongiae
  • Subclass Homoscleromorpha
  • Subclass Tetractinomorpha
  • Subclass Ceractinomorpha


Phylum Placozoa[edit]
  • Phylum Placozoa
Phylum Monoblastozoa[edit]
  • Phylum Monoblastozoa
Phylum Rhombozoa[edit]
  • Phylum Rhombozoa
  • Order Dicyemida
  • Order Heterocyemida
Phylum Orthonectida[edit]
  • Phylum Orthonectida


Phylum Cnidaria[edit]
  • Phylum Cnidaria
  • Class Hydrozoa
  • Order Hydroida
  • Suborder Anthomedusae (= Gymnoblastea or Athecata)
  • Suborder Leptomedusae (= Calyptoblastea or Thecata)
  • Order Trachylina
  • Order Siphonophora
  • Order Chondrophora
  • Order Actinulida
  • Class Anthozoa
  • Subclass Octocorallia (= Alcyonaria)
  • Order Alcyonacea
  • Order Gastraxonacea
  • Order Gorgonacea
  • Order Helioporacea
  • Order Pennatulacea
  • Order Protoalcyonaria
  • Order Stolonifera
  • Order Telestacea
  • Subclass Hexacorallia (= Zoantharia)
  • Order Actiniaria
  • Order Scleractinia (= Madreporaria)
  • Order Zoanthidea
  • Order Corallimorpharia
  • Subclass Ceriantipatharia
  • Order Antipatharia
  • Order Ceriantharia
  • Class Cubozoa
  • Class Scyphozoa
  • Order Stauromedusae
  • Order Coronatae
  • Order Semaeostomae
  • Order Rhizostomae
Phylum Ctenophora[edit]
  • Phylum Ctenophora
  • Order Beroida
  • Order Cestida
  • Order Cydippida
  • Order Ganeshida
  • Order Lobata
  • Order Platyctenida
  • Order Thalassocalycida

The authors divide the bilaterians in three informal groups:

  • acoelomates (phyla Platyhelminthes, Gastrotricha, Entoprocta, Gnathostomulida, Cycliophora)
  • blastocoelomate (or pseducoelomate, phyla Rotifera, Kinorhyncha, Nematoda, Nematomorpha, Acanthocephala, Loricifera)
  • coelomates (or eucoelomates, phyla Nemertea, Priapula, Annelida, Sipuncula, Echiura, Onychophora, Tardigrada, Arthropoda, Mollusca, Phoronida, Ectoprocta, Brachiopoda, Echinodermata, Chaetognatha, Hemichordata, Chordata).

Several groups traditionally viewed as having a blastocoelomate condition are viewed here as acoelomates (e.g., Gastrotricha, Entoprocta, Gnathostomulida).

Some of the coelomates groups (e.g., Arthropoda, Mollusca) have greatly reduced celomic spaces; often the main body cavity is a bloodfilled space called a hemocoel, and is associated with an open circulatory system.

The Brachiopoda, Ectoprocta and Phoronida are viewed as lophophorates.

In a phylogeny,[2] the bilaterians are divided in:

Phylum Platyhelminthes[edit]
  • Phylum Platyhelminthes
  • Class Turbellaria
  • Order Acoela
  • Order Catenulida
  • Order Haplopharyngida
  • Order Lecithoepitheliata
  • Order Macrostomida
  • Order Nemertodermatida
  • Order Polycladida
  • Order Prolecithophora
  • Order Proplicastomata
  • Order Proseriata
  • Order Rhabdocoela
  • Suborder Dalyellioida
  • Suborder Thyphloplanoida
  • Suborder Kalyptorhynchia
  • Suborder Temnocephalida
  • Order Tricladida
  • Class Monogenea
  • Subclass Monopisthocotylea
  • Subclass Polyopisthocotylea
  • Class Trematoda
  • Subclass Digenea
  • Subclass Aspidogastrea
  • Class Cestoda
  • Subclass Cestodaria
  • Subclass Eucestoda
Phylum Nemertea[edit]
  • Phylum Nemertea
  • Class Anopla
  • Class Enopla
Phylum Rotifera[edit]
  • Phylum Rotifera
  • Class Digonata
  • Order Seisonidea
  • Order Bdelloidea
  • Class Monogononta
Phylum Gastrotricha[edit]
  • Phylum Gastrotricha
  • Order Macrodasyida
  • Order Chaetonotida
  • Suborder Multitubulatina
  • Suborder Paucitubulatina
Phylum Kinorhyncha[edit]
  • Phylum Kinorhyncha
  • Order Cyclorhagida
  • Order Homalorhagida
Phylum Nematoda (= Nemata)[edit]
  • Phylum Nematoda (= Nemata)
  • Class Adenophorea (= Aphasmida)
  • Subclass Enoplia
  • Subclass Chromadoria
  • Class Secernentea (= Phasmida)
  • Subclass Rhabditia
  • Subclass Spiruria
  • Subclass Diplogasteria
Phylum Nematomorpha[edit]
  • Phylum Nematomorpha
  • Order Nectonematoidea
  • Order Gordioidea
Phylum Priapula[edit]
  • Phylum Priapula
  • Family Priapulidae
  • Family Tubiluchidae
  • Family Maccabeidae (= Chaetostephanidae)
Phylum Acanthocephala[edit]
  • Phylum Acanthocephala
  • Class Palaeacanthocephala
  • Class Archiacanthocephala
  • Class Eoacanthocephala
Phylum Entoprocta (= Kamptozoa)[edit]
  • Phylum Entoprocta (= Kamptozoa)
  • Family Loxosomatidae
  • Family Loxokalypodidae
  • Family Pedicellinidae
  • Family Barentsiidae
Phylum Gnathostomulida[edit]
  • Phylum Gnathostomulida
  • Order Filospermoidea
  • Order Bursovaginoidea
Phylum Loricifera[edit]
  • Phylum Loricifera
  • Order Nanaloricida
  • Family Nanaloricidae
  • Family Pliciloricidae
Phylum Cycliophora[edit]
  • Phylum Cycliophora
Phylum Annelida[edit]
  • Phylum Annelida
  • Class Polychaeta, with 25 orders and 87 families (not all are listed)
  • Order Capitellida
  • Order Chaetopterida
  • Order Cirratulida
  • Order Eunicida
  • Order Myzostomida
  • Order Opheliida
  • Order Spionida
  • Order Orbiniida
  • Order Oweniida
  • Order Phyllodocida
  • Order Sabellida
  • Order Terebellida
  • Class Clitellata
  • Subclass Oligochaeta
  • Order Lumbriculida
  • Family Lumbriculidae
  • Order Moniligastrida
  • Family Moniligastridae
  • Order Haplotaxida, with 25 families (not all are listed)
  • Family Almidae
  • Family Megascolecidae
  • Family Tubificidae
  • Family Naididae
  • Family Lumbricidae
  • Subclass Hirudinoidea
  • Order Acanthobdellida
  • Order Branchiobdellida
  • Order Hirudinida
Phylum Sipuncula[edit]
  • Phylum Sipuncula
  • Class Phascolosomida
  • Order Aspidosiphoniformes
  • Order Phascolosomiformes
  • Class Sipunculida
  • Order Golfingiaformes
  • Family Themistidae
  • Family Phascolionidae
  • Family Golfingiidae
  • Order Sipunculiformes
Phylum Echiura[edit]
  • Phylum Echiura
  • Order Echiuroinea
  • Order Xenopneusta
  • Order Heteromyota
Phylum Onychophora[edit]
  • Phylum Onychophora
  • Family Peripatidae
  • Family Peripatopsidae
Phylum Tardigrada[edit]
  • Phylum Tardigrada
  • Order Heterotardigrada
  • Order Mesotardigrada
  • Order Eutardigrada
Phylum Arthropoda[edit]
  • Phylum Arthropoda
  • Subphylum Trilobitomorpha
  • Subphylum Crustacea
  • Class Remipedia
  • Class Cephalocarida
  • Class Branchiopoda
  • Order Anostraca
  • Order Notostraca
  • Order Diplostraca
  • Suborder Laevicaudata
  • Suborder Spinicaudata
  • Suborder Cyclestherida
  • Suborder Cladocera
  • Class Malacostraca
  • Subclass Phyllocarida
  • Order Lepstostraca
  • Subclass Eumalacostraca
  • Superorder Hoplocarida
  • Order Stomatopoda
  • Superorder Syncarida
  • Order Bathynellacea
  • Order Anaspidacea
  • Superorder Eucarida
  • Order Euphausiacea
  • Order Amphionidacea
  • Order Decapoda
  • Suborder Dendrobranchiata
  • Suborder Pleocyemata
  • Infraorder Caridea
  • Infraorder Stenopodidea
  • Infraorder Brachyura
  • Infraorder Anomura
  • Infraorder Astacidea
  • Infraorder Palinura
  • Infraorder Thalassinidea
  • Superorder Peracarida
  • Order Mysida
  • Order Lophogastrida
  • Order Cumacea
  • Order Tanaidacea
  • Order Mictacea
  • Order Spelaeogriphacea
  • Order Thermosbaenacea
  • Order Isopoda
  • Suborder Anthuridea
  • Suborder Asellota
  • Suborder Calabozoidea
  • Suborder Epicaridea
  • Suborder Flabellifera
  • Suborder Gnathiidea
  • Suborder Oniscidea
  • Suborder Phreatoicidea
  • Suborder Valvifera
  • Order Amphipoda
  • Suborder Gammaridea
  • Suborder Hyperiidea
  • Suborder Caprellidea
  • Suborder Ingolfiellidea
  • Class Maxillopoda
  • Subclass Thecostraca

Brusca And Brusca Invertebrates

  • Infraclass Facetotecta
  • Infraclass Ascothoracida
  • Infraclass Cirripedia
  • Superorder Acrothoracica
  • Superorder Rhizocephala
  • Superorder Thoracica
  • Subclass Tantulocarida
  • Subclass Branchiura
  • Subclass Pentastomida
  • Subclass Mystacocarida
  • Subclass Copepoda
  • Infraclass Progymnoplea
  • Order Platycopioida
  • Infraclass Neocopepoda
  • Order Calanoida
  • Order Cyclopoida
  • Order Gelyelloida
  • Order Harpacticoida
  • Order Misophrioida
  • Order Monstrilloida
  • Order Mormonilloida
  • Order Poecilostomatoida
  • Order Siphonostomatoida
  • Subclass Ostracoda
  • Superorder Myodocopa
  • Order Myodocopida
  • Order Halocyprida
  • Superorder Podocopa
  • Order Podocopida
  • Order Platycopida
  • Order Palaeocopida
  • Subphylum Hexapoda
  • Class Entognatha
  • Order Collembola
  • Order Protura
  • Order Diplura
  • Class Insecta
  • Subclass Archaeognatha
  • Order Archaeognatha (= Microcoryphia)
  • Subclass Zygentoma
  • Order Thysanura
  • Subclass Pterygota
  • Infraclass Palaeoptera
  • Order Ephemeroptera
  • Order Odonata
  • Infraclass Neoptera
  • Order Plecoptera
  • Order Blattodea
  • Order Isoptera
  • Order Mantodea
  • Order Phasmida (= Phasmatoptera)
  • Order Grylloblattodea
  • Order Dermaptera
  • Order Orthoptera
  • Order Mantophasmatodea
  • Order Embioptera
  • Order Zoraptera
  • Order Psocoptera
  • Order Phthiraptera
  • Order Thysanoptera
  • Order Hemiptera
  • Order Strepsiptera
  • Order Megaloptera
  • Order Raphidioptera
  • Order Neuroptera
  • Order Coleoptera
  • Order Mecoptera
  • Order Siphonaptera
  • Order Diptera
  • Order Trichoptera
  • Order Lepidoptera
  • Order Hymenoptera
  • Subphylum Myriapoda
  • Class Diplopoda
  • Subclass Penicillata
  • Order Polyxenida
  • Subclass Chilognatha
  • Order Callipodida
  • Order Chordeumatida
  • Order Glomerida
  • Order Glomeridesmida
  • Order Julida
  • Order Platydesmida
  • Order Polydesmida
  • Order Polyzoniida
  • Order Siphonophorida
  • Order Siphoniulida
  • Order Sphaerotheriida
  • Order Spirobolida
  • Order Spirostreptida
  • Order Stemmiulida
  • Class Chilopoda
  • Subclass Notostigmophora
  • Order Scutigeromorpha
  • Subclass Pleurostigmophora
  • Order Craterostigmomorpha
  • Order Geophilomorpha
  • Order Lithobiomorpha
  • Order Scolopendromorpha
  • Class Pauropoda
  • Class Symphyla
  • Subphylum Cheliceriformes
  • Class Pycnogonida
  • Class Chelicerata
  • Subclass Merostomata
  • Order Eurypterida
  • Order Xiphosura
  • Subclass Arachnida
  • Order Acari
  • Suborder Opiloacarifomes
  • Suborder Parasitiformes
  • Suborder Acariformes
  • Order Amblypygi
  • Order Araneae
  • Suborder Mesothelae
  • Suborder Opisthothelae
  • Superfamily Mygalomorpha
  • Family Ctenizidae
  • Family Atypicae
  • Family Theraphosidae
  • Family Dipluridae
  • Superfamily Araneomorphae
  • Family Loxoscelidae
  • Family Theridiidae
  • Family Uloboridae
  • Family Araneidae
  • Family Tetragnathidae
  • Family Clubionidae
  • Family Linyphiidae
  • Family Agelenidae
  • Family Argyronetidae
  • Family Lycosidae
  • Family Pisauridae
  • Family Oxyopidae
  • Family Thomisidae
  • Family Heteropodidae
  • Family Salticidae
  • Family Dinopidae
  • Family Scytodidae
  • Order Opiliones
  • Order Palpigradi
  • Order Pseudoscorpionida
  • Order Ricinulei
  • Order Schizomida
  • Order Scorpiones
  • Order Solpugida
  • Order Uropygi
Phylum Mollusca[edit]
  • Phylum Mollusca
  • Class Aplacophora
  • Subclass Chaetodermomorpha (= Caudofoveata)
  • Subclass Neomeniomorpha (= Solenogastres)
  • Class Monoplacophora
  • Class Polyplacophora
  • Order Lepidopleurida
  • Order Ischnochitonida
  • Order Acanthochitonida
  • Class Gastropoda
  • Subclass Prosobranchia
  • Order Archaeogastropoda
  • Order Mesogastropoda
  • Order Neogastropoda
  • Subclass Opisthobranchia
  • Order Acochlidioidea
  • Order Cephalaspidea
  • Order Runcinoidea
  • Order Sacoglossa
  • Order Anaspidea
  • Order Thecosomata
  • Order Gymnosomata
  • Order Notaspidea
  • Order Nudibranchia
  • Subclass Pulmonata
  • Order Archaeopulmonata
  • Order Basommatophora
  • Order Stylommatophora
  • Order Systellommatophora
  • Class Bivalvia (=Pelecypoda, or Lamellibranchiata)
  • Subclass Protobranchia
  • Order Nuculida (= Palaeotaxodonta)
  • Order Solemyida (= Cryptodonta)
  • Subclass Lamellibranchia
  • Superorder Filibranchia (= Pteriomorpha)
  • Superorder Eulamellibranchia (= Heterodonta)
  • Order Palaeoheterodonta
  • Order Veneroida
  • Order Myoida
  • Subclass Anomalodesmata
  • Class Scaphopoda
  • Class Cephalopoda (= Siphonopoda)
  • Subclass Nautiloidea (= Tetrabranchiata)
  • Subclass Coleoidea (= Dibranchiata)
  • Order Sepioida
  • Order Teuthoida (= Decapoda)
  • Order Octopoda
  • Order Vampyromorpha
Phylum Phoronida[edit]
  • Phylum Phoronida
Phylum Ectoprocta (= Bryozoa)[edit]
  • Phylum Ectoprocta (= Bryozoa)
  • Class Phylactolaemata
  • Class Stenolaemata
  • Class Gymnolaemata
  • Order Ctenostomata
  • Order Cheilostomata
Phylum Brachiopoda[edit]
  • Phylum Brachiopoda
  • Class Inarticulata
  • Class Articulata
Phylum Echinodermata[edit]
  • Phylum Echinodermata
  • Class Crinoidea
  • Class Asteroidea
  • Order Platysterida
  • Order Paxillosida
  • Order Valvatida
  • Order Spinulosida
  • Order Forcipulatida
  • ”Sea daisies” (previously the class Concentricycloidea, but assigned by many authorities to the Spinulosida)
  • Class Ophiuroidea
  • Order Oegophiurida
  • Order Phrynophiurida
  • Order Ophiurida
  • Class Echinoidea
  • Subclass Cidaroidea
  • Subclass Euechinoidea
  • Infraclass Echinothurioidea
  • Infraclass Acroechinoidea
  • Cohort Diadematacea
  • Cohort Echinacea
  • Cohort Irregularia
  • Class Holothuroidea
  • Subclass Dendrochirotacea
  • Order Dactylochirotida
  • Order Dendrochirotida
  • Subclass Aspidochirotacea
  • Order Aspidochirotida
  • Order Elasipodida
  • Subclass Apodacea
  • Order Molpadida
  • Order Apodida
Phylum Chaetognatha[edit]
  • Phylum Chaetognatha
  • Order Phragmophora
  • Order Aphragmophora
Phylum Hemichordata[edit]
  • Phylum Hemichordata
  • Class Enteropneusta
  • Class Pterobranchia
  • Class Planctosphaeroidea
Phylum Chordata[edit]
  • Phylum Chordata
  • Subphylum Urochordata (= Tunicata)
  • Class Ascidiacea
  • Class Thaliacea
  • Class Appendicularia (= Larvacea)
  • Class Sorberacea
  • Subphylum Cephalochordata (= Acrania)
  • Subphylum Vertebrata
  • Class Myxini
  • Class Cephalaspidomorphi
  • Class Chondrichthyes
  • Class Osteichthyes
  • Class Amphibia
  • Class Reptiliomorpha (= Sauropsida)
  • Class Mammalia


  1. ^Richard C. Brusca & Gary J. Brusca (2003). Invertebrates (2nd ed.). Sunderland, Massachusetts: Sinauer Associates. ISBN978-0-87893-097-5.
  2. ^Richard C. Brusca & Gary J. Brusca (2003), p. 875.
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