Biology:Taxonomy of commonly fossilised invertebrates

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The Ordovician cystoid Echinosphaerites (an extinct echinoderm of the Class Rhombifera) from northeastern Estonia; encrusted by a graptolite (black branches).

Taxonomy of commonly fossilised invertebrates is a complex and evolving field that combines both traditional and modern paleozoological terminology. This article provides an overview of various invertebrate taxa found in the fossil record, ranging from protists to arthropods. The taxonomy discussed is not exhaustive, but rather emphasizes invertebrates that are either frequently collected as fossils or are extinct. This includes groups that are significant in paleontological contexts, notably abundant in the fossil record, or have a high proportion of extinct species. For better understanding, special notations used in this context are explained below:

  • [ ! ]: Indicates clades that are important as fossils or very abundant in the fossil record.
  • [ – ]: Indicates clades that contain a large proportion of extinct species.
  • [ † ]: Indicates clades that are completely extinct.

The paleobiologic systematics that follow are not intended to be comprehensive, rather encompass invertebrates that (a) are popularly collected as fossils and/or (b) extinct. As a result, some groups of invertebrates are not listed.[1]

If an invertebrate animal is mentioned below using its common (vernacular) name, it is an extant (living) taxon, but if it is cited by its scientific genus, then it is typically an extinct invertebrate known only from the fossil record.[2]

Invertebrate clades that are important fossils (e.g. ostracods, frequently used as index fossils), and/or clades that are very abundant as fossils (e.g. crinoids, easily found in crinoidal limestone),[3] are highlighted with a bracketed exclamation mark [ ! ].

Domain of Eukaryota/Eukarya

Quinqueloculina, a foraminiferan (a type of protist) from Donegal Bay, Ireland.

Eukaryotes; eukaryotes are cellular organisms bearing a central, organized nucleus with DNA.

Sub-domain of Opisthokonta

Opisthokonts; the animal-related kingdoms. Include: proto-spongal choanoflagellates; proto-fungal microsporidians; true fungi; true animals.

  • most life forms documented, extinct or extant.
    • excludes: many molds; all one-celled protists (protoctists); all algae; all green plants.

Kingdom of Animalia / Metazoa - All Invertebrates and Vertebrates

Metazoans; multicellular "true" animals (multicellular creatures that capture and ingest their organic food).

  • comprises most living and deceased species which have ever been recorded, extinct or extant.
    • excludes all unicellular and fungal opisthokonts.

Sub-kingdom of Parazoa

Parazoans; typically sessile, basal non-eumetazoans. They are the most-primitive animals, comprising simple, colonial, attached, bottom-dwelling marine invertebrates.

Phylum Archaeocyatha/Archeocyatha/Archaeocyathida/Archeocyathida/Pleospongia [†]

Cone-shaped archaeocyathids/archeocyathids; cup-shaped archaeocyathans/archeocyathans; reef-building pleosponges;calcareous "ancient-cups".

Includes fossil genera such Archaeocyathus, Cambrocyathus, Atikonia, Tumuliolynthus, Kotuyicyathus, Metaldetes, Ajacicyathus and Paranacyathus.

Archaeocyatha is sometimes classified as a class of Porifera below.

Phylum Porifera/Nuda/Spongia

Pattersonia ulrichi Rauff, 1894; an Ordovician hexactinellid sponge from near Cincinnati, Ohio.
Tetractinella trigonella at MUSE - Science Museum in Trento

Quintessential true sponges; marine, colonial, pore-bearing animals; organized collar-flagellates; poriferans - today mostly siliceous – half of all documented species of Porifera are fossils and extinct.[4]

Porifera may eventually be broken up into separate phyla:

  • Sub-phylum Calcarea/Calcispongiae (primitive calcareous poriferans such as yellow lemon sponge, sphinctozoans, pharetronids, Scypha, Leucetta, Gravestockia, Grantia, Astraeospongium, Clathrina, Lelapia, Rhaphidonema, and Girtyocoelia).
    • Class Calcinea
    • Class Calcaronea
    • Class Stromatoporoidea/Stromatoporata/Stromatoporida/Spongliomorphida [†] (lime-layered stromatoporoids/reef-building stromatoporates/button-shaped stromatoporids/disc-shaped spongliomorphids; e.g., Stromatopora, Aulacera, Stromatactis, Actinostroma, Discophyllum, Parallelopora and Amphipora)
    • Class Heteractinida [†] (Paleozoic calcitic heteractinids such as Eiffelia)
  • Sub-phylum Silicea / Silicospongia (siliceous poriferans):
    • Class Demospongea/Demospongiae (most living sponges hardened by opaline silica or spongin; for instance, horny sponge, bath sponge, stove-pipe sponge, yellow boring sponge, carnivorous sponge, bristle sponge, chaetids, lithistids, Astroclera, Ceractinomorpha, Clionoides, Hindia, Ventriculites, Laosoiadia, Clionolithes, Tetractinella, and Astylospongia)
    • Class Hexactinellida/Hyalospongiae/Sclerospongiae (siliceous, deep-sea glass sponges, e.g. glassy-latticed Venus flower basket, bird's nest sponge, cloud sponge, Hexactinella, Hydroceras, Dictyonina, Brachiospongia, Titusvillea, and Rhizopoterion)

Sub-kingdom of Eumetazoa

Eumetazoans; true metazoans (typically mobile, multicellular animals).

Eumetazoa contains most of the living and deceased species of recorded life, including most invertebrates (extinct and extant), as well as all vertebrate animals.

Super-phylum of Radiata

Radiates; non-bilaterian eumetazoans.

Phylum Cnidaria/Coelenterata

Aulopora (a tabulate coral) from the Silica Shale (Middle Devonian), northwestern Ohio.

Cnidarians/coelenterates:

Super-phylum of Lophotrochozoa / Protostomia # 1

Lophotrochozoan bilaterians, such as flatworms, ribbon worms, lophophorates, and molluscs.

Phylum Bryozoa/Ectoprocta/Polyzoa

Heterotrypa, a trepostome bryozoan from the Corryville Formation (Upper Ordovician) in Covington, Kentucky.

Bryozoans – half of all documented species of Bryozoa are fossils and extinct.[5]

  • Class Stenolaemata / Gymnolaemata [!] (mostly marine, calcareous bryozoans):
    • Order Cheilostomata [!] (living, rimmed-mouthed moss animals)
    • Order Cyclostomatida (uncontracted, round-mouthed bryozoans including fossil Stomatopora)
    • Order Cystoporata [†] (extinct, minor group of moss animals)
    • Order Trepostomata [†] [!] (changed-mouthed bryozoans such as extinct Constellaria and Monticulipora)
    • Order Cryptostomata [†] [!] (round hidden-mouthed bryozoans such as Archimedes, Fenestrellina and Rhombopora)
    • Order Ctenostomata [†] (uncommon, comb-mouthed bryozoans)
    • Order Phylactolaemata (living, fresh-water bryozoans)

Phylum Brachiopoda

Rhynchotrema dentatum, a rhynchonellid brachiopod from the Cincinnatian (Upper Ordovician) of southeastern Indiana.

Lampshells, brachiopods or "brachs," (not to be confused with the hard-shelled marine mollusks below) – 99% of all documented species of Brachiopoda are now extinct.

Phylum Annelida

Segmented worms such as earthworms and leeches.

  • Class Polychaeta (marine annelids / polychaetes)
    • Order Scolecodonta [!] (mostly chitinous jaws of scolecodonts)

Phylum Mollusca

Peltoceras solidum ammonite from the Matmor Formation (Jurassic, Callovian) in the Matmor Formation, Makhtesh Gadol, Israel.
Vermetid gastropod Petaloconchus intortus attached to a branch of the coral Cladocora; Pliocene of Cyprus.

Molluscs or mollusks, not to be confused with the hard-shelled marine brachiopods above.

Super-phylum of Ecdysozoa/Protostomia # 2

Ecdysozoans, such as nematodes, horsehair worms, and molting bilaterians/panarthropods

Phylum Tardigrada

Panarthropodic water bears.

Phylum Onychophora

Panarthropodic velvet worms, including proto-arthropodic fossils of Arthropleura and Aysheaia.

Phylum Arthropoda

Elrathia kingii (trilobite) from the Wheeler Shale (Middle Cambrian), Utah.

Arthropods; jointed legged creatures with an exoskeleton.

Super-phylum of Deuterostomia / Enterocoelomata

Second-mouthed bilaterians called deuterostomians, such as chordates and echinoderms.

Phylum Echinodermata

Middle Jurassic (Callovian) crinoid pluricolumnals (Apiocrinites) from the Matmor Formation in Hamakhtesh Hagadol, southern Israel.

Echinoderms – 72% of all documented species of Echinodermata are fossils and extinct.[7]

  • Subphylum Crinozoa (sessile echinoderms) – 91 percent of all documented species of Crinozoa are now extinct
    • Class Crinoidea (crinoids / sea lilies) – See Crinozoa above
  • Subphylum Blastozoa [†] (extinct blastoids)
  • Subphylum Echinozoa (mobile echinoderms) – 89 percent of all documented species of Echinozoa are now extinct
  • Subphylum Asterozoa
    • Class Asteroidea (sea stars / starfish)
    • Class Ophiuroidea

Phylum Hemichordata

Pendeograptus fruticosus graptolites from the Bendigonian Australian Stage (Lower Ordovician) near Bendigo, Victoria, Australia. Two overlapping, three-stiped rhabdosomes.

Hemichordates such as extant acorn worms – Less than half of the documented species of Hemichordata are fossils and extinct.

  • Class Graptoloidea [†] (extinct graptolites)
    • Order Dendroidea [†]
    • Order Graptoloidea [†]
      • Suborder Didymograptina [†]
      • Suborder Diplograptina [†]
      • Suborder Monograptina [†]

Phylum Chordata

Both invertebrate and vertebrate chordates; animals possessing a notochord.

Invertebrate subphyla

Subphylum Vertebrata

Deinosuchus hatcheri at the Natural History Museum of Utah.

See also

  • Invertebrate paleontology

References

  1. ↑ For superb anatomical illustrations and much-more comprehensive information, see Volume E (Archaeocyatha / Porifera) through Volume V (Graptolithina), published 1953 to 2006 (and continuing), of the Treatise on Invertebrate Paleontology, long-edited by Raymond C. Moore and Roger L. Kaesler (Boulder, Colorado: Geological Society of America; and Lawrence, Kansas: University of Kansas Press). But be warned that some terms therein employed – such as supersubphylum – can be unnecessarily wordy or abstruse. Incidentally, revised volumes have been recently published regarding the sponges/archaeocyatha (2004, ISBN:0-8137-3131-3) and the brachiopods (2006, ISBN:0-8137-3135-6).
  2. ↑ The names of genera, orders, classes and phyla have been culled from dozens of sources, both current and decades-old. See the International Code of Zoological Nomenclature (ICZN), as well as Volume 1 and Volume 2 of Grzimek's Animal Life Encyclopedia (Farmington Hills, Michigan: Gale Group), edited by zoologists Michael Hutchin, Dennis A. Thorney and Sean F. Craig (2003).
  3. ↑ For correspondingly ancient ecosystems, see the Treatise on Ecology and Paleoecology, Volume 2: Paleoecology, edited for years by Harry S. Ladd (1957 / 1971), and published by both the Geological Society of America (Boulder, Colorado) and the Waverly Press (Washington, D.C.).
  4. ↑ The rates of extinction for sponges and other phyla are derived from W. H. Easton, 1960, Invertebrate Paleontology (New York: Harper and Brothers) and various modern sources.
  5. ↑ For bryozoans and brachiopods, the same footnote as above.
  6. ↑ For bivalves and cephalopods (both mollusks), see the above notation.
  7. ↑ For the echinoderms, see the above footnote regarding W. E. Easton, 1960, Invertebrate Paleontology, and other sources.



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