Brachiopoda

A phylum of predominantly sessile, filter-feeding marine invertebrates, with a ciliated, tentaculate feeding organ or lophophore and a bivalved shell. The two main living types (with either articulated or inarticulated valves) first appeared in the Lower Cambrian. Articulated forms give the phylum its common name: ‘lamp shells’. In recent molecular analyses, brachiopods and phoronids form a single clade.

Keywords: Brachiopoda; lophophore; inarticulate brachiopod; articulate brachiopod; Phoronida

Figure 1. Comparison of the body plans of the principal lineages of living brachiopods. Mouth (m) and anus (a) are marked. (After Nielsen, 1995.) (Not to scale)
Figure 2. The principal organs of a generalized rhynchonelliform brachiopod. (After Williams and Rowell, 1965.)
Figure 3. Close-up view of the rhynchonelliform, articulate brachiopod Liothyrella neozelanica in feeding position with its gape fully open, showing a conspicuous fringe of marginal setae and the curtain-like lophophore forming inhalant and exhalant compartments within the mantle cavity. The inhalant current enters at the sides and the exhalant current leaves centrally. In many fossil and extant deep-water forms, the inhalant and exhalant currents are offset by a median fold or groove in the shell, presumably to minimze recirculation of filtered water. This animal's shell is about 40 mm across. (From a photograph provided by L. Shackleton, Wellington, New Zealand.)
Figure 4. A group of the rhynchonelliform, articulate brachiopod Megathiris detruncata on the wall of a small cave, south coast of Madeira, depth c. 10 m. The largest shell is c. 10 mm wide. There are no setae. (Photo credit: Peter Wirtz.)
Figure 5. A group of the thecideidine, articulate brachiopod Pajaudina atlantica on the roof of a small cave, La Palma, Canary Islands, depth c. 15 m. The complexly folded lophophore is supported by bas-relief ridges on the brachial valve and the shell gapes very widely. Each shell is c. 5 mm wide. As in Megathiris, there are no setae. (Photo credit: Peter Wirtz.)
Figure 6. Time ranges of principal brachiopod lineages. (Modified from Williams A, Carlson SJ, Brunton CHC, Holmer L and Popov L (1996) A supra-ordinal classification of Brachiopoda. Philosophical Transactions of the Royal Society, B 351: 1171–1193). Lineages surviving to the present day are coloured yellow. The background curve gives an impression of changing family-level diversity through geological time. (After Rudwick, 1970.)
Figure 7. Phylogeny of brachiopods and phoronids, summarizing recent gene sequence evidence and showing the distribution of key characters mentioned in the text. The uncertain placement of the thecideidine lineage within rhynchonelliforms is indicated by a dashed line.
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 Further Reading
    Cohen BL, Gawthrop AB and Cavalier-Smith T (1998) Molecular phylogeny of brachiopods and phoronids based on nuclear-encoded small subunit ribosomal RNA gene sequences. Philosophical Transactions of the Royal Society, B 353: 2039–2061.
    Cohen BL, Stark S, Gawthrop AB, Burke ME and Thayer CW (1998) Comparison of articulate brachiopod nuclear and mitochondrial gene trees leads to a clade-based redefinition of protostomes (Protostomozoa) and deuterostomes (Deuterostomozoa). Proceedings of the Royal Society, London, Series B 265: 475–482.
    Cohen BL and Weydmann A (2005) Molecular evidence that phoronids are a subtaxon of brachiopods (Brachiopoda: Phoronata) and that genetic divergence of metazoan phyla began long before the Early Cambrian. Organisms, Diversity & Evolution 5: 253–273.
    book Hyman LH (1959) The Invertebrates: Smaller Coelomate Groups, pp. 1–783. London: McGraw-Hill.
    other Kaesler R (ed.) (1997–2002 and in press). Treatise on Invertebrate Paleontology, Part H, Brachiopoda (revised). Boulder, CO: Geological Society of America and Lawrence, KA: University of Kansas Press. [This treatise is appearing in six volumes, of which four were published and two were in press at the time of writing.].
    book Nielsen C (1995) Animal Evolution: Interrelationships of the Living Phyla, pp. 1–467. Oxford: Oxford University Press.
    Richardson JL (1986) Brachiopods. Scientific American 255: 96–102.
    book Rudwick MJS (1970) Living and Fossil Brachiopods. London: Hutchinson.
    book Williams A and Rowell AJ (1965) "Brachiopod Anatomy". Moore RC (ed.) Treatise on Invertebrate Paleontology, Part H, Brachiopoda, pp. 6–57. Boulder, CO: Geological Society of America and Lawrence, KA: University of Kansas Press.
    book Willmer P (1990) Invertebrate Relationships, pp. 1–400. Cambridge: Cambridge University Press.
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Cohen, Bernard L(Jan 2007) Brachiopoda. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001614.pub2]