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Annelida: the Metameric Body Form 221 12 Annelida:The Metameric Body Form The Samoan palolo worm (Eunice viridis) is one of approximately15,000 spe­ cies in the phylum Annelida. The palolo's 1·eproductive habits are unusual, but effective.As you will see in this chapter, "unusual" is not the exceptionfor this Chapter Outline taxonomicallychallenging phylum. 12.1 Evolutionary Perspective Relationships to Other Animals Metamerism and Tagmatization 12.2 Annelid Structure and Function 12. 1 EVOLUTIONARY PERSPECTIVE External Structure and Locomotion Feeding and the Digestive System Gas Exchange and Circulation LEARNING OUTCOMES Nervous and SensoryFunctions Excretion 1. Describe the relationships of members of the Annelida to other animal phyla. Regeneration, Reproduction, 2. Explain the benefits of metamerism for an annelid and Development 12.3 Clade (Class) Errantia Nereis (Neanthes, Alitta) At the time of the November full moon on islands near Samoa in the South Glycera Pacific, people rush about preparing for one of their biggest yearly feasts. In Fireworms just one week, the sea will yield a harvest that can be scooped up in nets and 12.4 Clade (Class) Sedentaria Tubeworms buckets (figure 12.1). Worms by the millions transform the ocean into what one Siboglinidae writer called "vermicelli soup!" Celebrants gorge themselves on worms that have Echiura been cooked or wrapped in breadfruit leaves.The Samoan palolo worm (Eunice Clade Clitel/ata 12.5 Basal Annelid Groups viridis, alternatively Palo/a viridis) spends its entire adult life in coral burrows Chaetopteridae at the sea bottom. Each November, one week after the full moon, this worm Sipuncula emerges from its burrow, and specialized body segments devoted to sexual 12.6 Further Phylogenetic Considerations reproduction break free and float to the surface, while the rest of the worm is safe on the ocean floor. The surface water is discolored as gonads release their countless eggs and sperm. The natives' feast is short lived, however; these reproductive swarms last only two days and do not recur for another year. The Samoan palolo worm is a member of the phylum Annelida (ah-nel' i-dah) (L. annellus, ring). Other members of this phylum include countless marine worms, the soil-building earthworms, and predatory leeches (table 12.1). Characteristics of the phylum Annelida include: 1. Body metameric, bilaterally symmetrical, and worm-like 2. Spiral cleavage, trochophore larvae (when larvae are present), and schizocoelous coelom formation 3. Paired, epidermal setae (chaetae) 4. Closed circulatory system 5. Dorsal suprapharyngeal ganglia and ventral nerve cord(s) with ganglia 6. Metanephridia (usually) or protonephridia Relationships to Other Animals It has been clear for many years that Annelida is a monophyletic assemblage of marine, freshwater, and terrestrial worms. They are lophotrochozoans and, thus, share common ancestry with Mollusca, Brachiopoda, Bryozoa, Nemertea, and others Annelida: The Metameric Body Form 221 (a) (b) FIGURE 12.1 Palolo Feasts. (a) Women from the Nggela Group (Florida Islands) of the Solomon Islands use light from a torch to attract reproductive segments of Eunice viridis (family Eunicidae) during collecting. Photoreceptors on the reproductive segments elicit the response of the worms to light. (b) A swarm of worms as viewed through the lens of a diver's camera. The photograph on page 220 shows palolo worms (referred to by islanders as "odu") ready for feasting. (Pbotograpbs courte.,y of Dr. Si111011 Foale, Principle Resectrcb Fe/10111, ARC Ce11tre of Exce/le11cefor Coral Req( Studies.James Cook University, Quee11sla11d, Austmlio.) Basal Protists Phyla FIGURE 12.2 Evolutionary Relationships of Annelids to Other Animals. This figure shows an interpretation of the relationship of the Annelida to other members of the animal kingdom. The relationships depicted here are based on evidence from developmental and molecular biology. Annelids are placed within the Lophotrochozoa along with the Mollusca, Plathyhelminthes, Rotifera, and others (see pages xvi-xvii). The phylum includes approximately 15,000 species of segmented worms. Most of these are marine and traditionally classified into the class Polychaeta. As discussed in the text, this placement is being reevaluated based on evidence from molecular biology. The Christmas-tree worm (Spirohrcmcbus gigan/e11s) (family Serpulidae) shown here is a memher of the clade Senclentaria The spiral fans of this tube-dwelling annelid are derived from prostomial palps that surround the mouth and are specialized for feeding and gas exchange. 222 CHAPTER TWELVE to differentiate groups within the polychaetes have been res­ TABLE 12.1 urrected to represent two major annelid clades: Errantia and CLASSIFICATION OF THE PHYLUM ANNELIDA Sedentaria. Sedentaria now includes some "polychaetes," leeches, earthworms, and even worms that were formerly Phylum Annelida (ah-nel'i-dah) considered separate phyla (Echiura and Pogonophora). Cli­ The phylum of triploblastic, coelomate animals whose members tellata is still a valid clade within the Sendentaria composed are metameric (segmented), elongate, and cylindrical or of earthworms, leeches, and a few smaller taxa. The leeches oval in cross section. Annelids have a complete digestive form a monophyletic clade (Hirudinea) within the Clitellata. tract; paired, epidermal setae; and a ventral nerve cord. The earthworms and their relatives, however, are not mono­ Approximately 15,000 species of annelids have been phyletic. Thus the name "Oligochaeta," which was formerly described. considered a subclass name within the Clitellata, should Clade (Class) Errantia (er-ran'tiah) be abandoned as a taxonomic designation. Two other Marine annelids; parapodia with prominent lobes supported groups lie outside of Sedentaria and Errantia. One of these, by internalized chaetae and ventral cirri; palps well Chaetopteridae, was formerly considered to be a family developed. Nereis, Eunice, Glycera. within the "Polychaeta." The other, Sipuncula, was another Clade (Class) Sedentaria (sed-en-ter'iah) phylum outside of Annelida. The hierarchical taxonomic­ Marine, freshwater, and terrestrial annelids; parapodia with categories associated with these clades have not been estab­ reduced lobes or parapodia lacking; setae associated lished. We treat Errantia and Sedentaria as clades with a with the stiff body wall to facilitate anchoring in tubes and burrows; palps reduced. Includes the clade Clitellata, parenthetical "class" designation, which seems a logical out­ echiruans, and siboglinids. Arenicola, Riftia, Lumbricus, come of ongoing and future research. Hirudo Chapter 12 is now organized to reflect this new phylo­ Other Annelid Taxa genetic work. In the next section, we describe metamerism Sipuncula (si-pun'ku-lah) and tagmatization. This discussion is followed by coverage Marine; unsegmented body; retractable anterior trunk of annelid structure and function. The term "polychaete" is called an introvelt. Formerly phylum Sipuncula. Inclusion used in a nontaxonomic sense to refer to a host of marine of this group within the Annelida remains somewhat annelids (both errantians and sedentarians) when adap­ controversial. Themiste. tations related to their common marine habitat result in Chaetopteridae (ke-top-ter' ida) similar structures and functions. After the basics of anne­ Marine; three-palt body; lives in a U-shaped tube, lid structure and function are discussed, the two major appendages for feeding and creating water currents. annelid clades and the two outlying groups are described. Chaetoptei·us. Since members of the clade Clitellata are different in many respects from other sedentarians, unique aspects of their structure and function are described in the section on Sed­ entaria. As always, this chapter ends with "Further Phyloge­ netic Considerations." (figure 12. 2 and see chapters 10 and 11). Our understanding of phylogeny within the phylum, however, has a history of contentious debate. The application of modern phylogenetic analysis is helping unravel the annelid taxonomic-tangle. Metamerism and Tagmatization Anyone who has followed the debates involved with elu­ Earthworm bodies are organized into a series of ringlike seg­ cidating annelid phylogeny will have a better appreciation ments. What is not externally obvious, however, is that the of the dynamics within the field of animal taxonomy. These body is divided internally as well. Segmental arrangement of debates underscore the importance of taxonomy in helping body parts in an animal is called metamerism (Gr. meta, zoologists understand the evolution of bilateral morphology after + mere, part). (see the discussion of metamerism and tagmatization in the Metamerism profoundly influences virtually eve1y next section). aspect of annelid structure and function, such as the anatomi­ Previous editions of this book described the Annelida cal arrangement of organs that are coincidentally associated as being composed of two classes: Polychaeta (marine with metamerism. For example, the compartmentalization worms) and Clitellata (leeches, earthworms, and others). of the body has resulted in each segment having its own While "Polychaeta" was rightly described as being paraphy­ excreto1y, nervous, and circulato1y structures. In most mod­ letic, recent molecular phylogenetic work has shown that ern annelids two related functions are probably the primary the entire annelid assemblage is encompassed by what was adaptive features of metamerism: flexible support and effi­ being called "Polychaeta." As will
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