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22. Annelida

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Annelida, or segmented worms, is a morphologically diverse group of animals, with a large number of species (around 22,000) that inhabit practically all marine habitats, as well as many terrestrial and freshwater environments. The placement of annelids in the tree of life is still unclear, although most recent hypotheses relate them to molluscs, nemerteans, phoronids, and brachiopods. The phylogeny within this group also presents majar challenges. Annelida has traditionally been split into three classes: "Polychaeta" (bristleworms), "Oligochaeta" (earthworms), and Hirudinea (leeches); however, recent phylogenetic analyses have shown that Hirudinea is nested within a paraphyletic "Oiigochaeta" forming a clade now referred to as Clitellata. Furthermore, Clitellata also appears to be nested within a paraphyletic "Polychaeta." Additionally, other groups displaying a morphology far from the typical annelid and considered as separate phyla in the past, such as Echiura (spoon worms), Sipuncula (peanut worms), Siboglinidae (pogonophores and vestimentiferans), and Myzostomida are now also considered to be annelids, though there have been some controversies about their specific position. This chapter offers an updated synthesis of the evolution of segmented worms and particular characteristics that were key to their evolution since the Cambrian.
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Annelida is one of three animal groups possessing segmentation and is central in considerations about the evolution of different character traits. It has even been proposed that the bilaterian ancestor resembled an annelid. However, a robust phylogeny of Annelida, especially with respect to the basal relationships, has been lacking. Our study based on transcriptomic data comprising 68,750 - 170,497 amino acid sites from 305 - 622 proteins resolves annelid relationships, including Chaetopteridae, Amphinomidae, Sipuncula, Oweniidae, Magelonidae in the basal part of the tree. Myzostomida, which have been indicated to belong to the basal radiation as well, are now found deeply nested within Annelida as sister group to Errantia in most analyses. Based on our reconstruction of a robust annelid phylogeny, we show that the basal branching taxa include a huge variety of life-styles such as tube-dwelling and deposit-feeding, endobenthic and burrowing, tubicolous and filter-feeding, as well as errant and carnivorous forms. Ancestral character state reconstruction suggests that the ancestral annelid possessed a pair of either sensory or grooved palps, bicellular eyes, biramous parapodia bearing simple chaeta and lacked nuchal organs. Since the oldest fossil of Annelida is reported for Sipuncula (520 Mya), we infer that the early diversification of annelids took place at least in the Lower Cambrian.
Polychaetes and Allies: The Southern Synthesis
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Beesley, P. L., Ross, G. J. B., and Glasby, C. J. 2000. Polychaetes and Allies: The Southern Synthesis. Fauna of Australia. Vol. 4 Polychaeta, Myzostomida, Pogonophora, Echiura, Sipuncula. Melbourne, Australia: CSIRO Publishing.
Towards a Tree of Life for the Annelida
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Siddall, M. E., Borda, E., and Rouse, G. W. 2004. Towards a Tree of Life for the Annelida. In, Assembling the Tree of Life (Cracraft, J. and Donoghue, M., eds.), pp. 237-251. New York, NY: Oxford University Press.
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Rouse, G. W. and Pleijel, F. 2001. Polychaetes. Oxford, UK: Oxford University Press.
  • G W Rouse
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Rouse, G. W. and Pleijel, F. 2001. Polychaetes. Oxford, UK: Oxford University Press.