Clitellata

The Clitellata are a class of annelid worms, characterized by having a clitellum - the 'collar' that forms a reproductive cocoon during part of their life cycles. The clitellates comprise around 8,000 species. Unlike the class of Polychaeta, they do not have parapodia and their heads are less developed.

Clitellata
Temporal range: Triassic–Recent[1]
Earthworm
Scientific classification
Kingdom: Animalia
Phylum: Annelida
Class: Clitellata
Subclasses

Branchiobdellae
Hirudinea
"Oligochaeta" (paraphyletic)
and see text

Characteristics

Clitellate annelids are segmented worms characterised by the clitellum or girdle which is located near the head end of mature individuals. The mouth is on the ventral surface and is overhung by the prostomium (proboscis). The brain is not located in the head but in one of the body segments. The clitellum is formed by a modification of several segments, and either includes the female gonopores or is located just behind them. During copulation, this glandular tissue secretes mucus that keeps the paired individuals together while they exchange sperm. Afterwards it secretes material that forms a cocoon that encircles the animal's body and encloses the eggs and sperm. The animal works this cocoon forward and over its head end, whereupon the ends of the cocoon become sealed, with fertilisation and development taking place inside.[2][3]

Earthworms and their kin, in the subclass Oligochaeta, lack eyes but have photoreceptor cells in the skin, especially in the dorsal portion of the anterior end. They also lack parapodia and appendages on the prostomium, the body and the periproct (terminal segment on which the anus is located). The gonads are located in a few segments near the clitellum, with the testes being anterior to the ovaries. There are four bundles of one to twenty-five chaetae on each segment; these have muscles attached to their bases and can be extended or retracted.[2]

Leeches and their relatives, in the subclass Hirudinea, mostly have flattened bodies, usually tapered at both ends. They have a fixed number of segments, 33, but the segmentation is not visible externally because the cuticle is marked with annulations. Leeches do not bear chaetae. The front few segments or head have been modified into a sucker that usually surrounds the mouth. These segments usually bear several ocelli on the upper side. The clitellum occupies segments 9 to 11 but is only noticeable during breeding periods. The hindermost segments form another, larger, disc-shaped sucker located on the underside of the body. The anus is on the dorsal surface just in front of the posterior sucker. The body wall includes strong transverse, longitudinal and diagonal muscles which give the animal great flexibility and extensibility.[2]

Distribution and habitat

Clitellates live on land, in freshwater or in the ocean. The subclass Branchiobdellae includes tiny species which crawl over the surface of freshwater crustaceans, especially crayfish. The leeches in subclass Hirudinea are mostly aquatic, a few living in the sea but mostly inhabiting freshwater locations, particularly the sediments on the bottom of lakes, ponds and sluggish streams. They thrive in polluted waters and places with high quantities of decaying organic matter and may be very numerous. They are more abundant in temperate waters in the northern hemisphere than elsewhere.[2]

The subclass Oligochaeta, which includes the earthworms as the largest members of the group, mostly live on land, burrowing in damp soil. Smaller freshwater species burrow in mud or live among aquatic vegetation. The marine species are mostly tiny and live in the interstices between sand grains, from the intertidal zone to the deep sea.[2]

Reproduction

All clitellata are hermaphrodites. During copulation, the clitellum produces a mucus that holds worms in place whilst they mate. During reproduction, the clitellum secretes a yolk (albumen) and a proteinaceous sheath which hardens. The worm then creeps out backward from the coat and deposits either fertilized zygotes or both ovae and sperm into the coat, which is then packed into a cocoon. The zygotes then develop directly in the cocoon without passing through a larval stage (as opposed to other annelids, e.g. Polychaeta.) This mechanism is considered to be apomorphic (a newly derived characteristic rather than an evolutionarily ancestral one).[4]

Systematics

According to modern phylogenetic analyses, the Clitellata are considered to be a monophyletic clade embedded deep in the polychaetes.[5][6][7]

Historically, the group was classified into the subclasses Oligochaeta and the Hirudinea. The oligochaetes contained the tubificids (Naididae, Lumbricidae, and Lumbriculidae - commonly the tube worms and the earthworms. The Hirudinea contained the leeches and the branchiobdellids. Modern analysis has revealed Branchiobdella and Hirudinea are two sister groups to the lumbriculids and they are daughter groups to the tree of oligochaetes.

The Acanthobdellidea, a sister group to Hirudinea, are sometimes moved out of the Hirudinea as a distinct subclass, too. Overall, clitellate phylogeny is not well resolved.

Namely, the Acanthobdellidea, Branchiobdella and Hirudinea are monophyletic, but are embedded among the Oligochaeta, which are an evolutionary grade of lineages that are outwardly similar, but not very close relatives. In particular, the leeches and earthworms appear to be very close relatives. Two approaches are possible:[8]

  • abolish Oligochaeta as traditionally delimited in favor of a number of smaller monophyletic lineages[8]
  • treat Oligochaeta and Clitellata as synonymous while splitting up the traditional "oligochaetes" into monophyletic lineages.[8]

Footnotes
  1. Manum, S. B.; Bose, M. N.; Sawyer, R. Y. T. (1991). "Clitellate cocoons in freshwater deposits since the Triassic". Zoologica Scripta. 20 (4): 347. doi:10.1111/j.1463-6409.1991.tb00300.x.
  2. Ruppert, Edward E.; Fox, Richard, S.; Barnes, Robert D. (2004). Invertebrate Zoology, 7th edition. Cengage Learning. pp. 459–482. ISBN 978-81-315-0104-7.
  3. "Animal reproductive system: Mechanisms that aid in the union of gametes: Annelids and mollusks". Encyclopedia Britannica. Retrieved 25 October 2017.
  4. Reichardt (2006): pp.63, 67-68
  5. Zrzavý, Jan; Říha, Pavel; Piálek, Lubomír; Janouškovec, Jan (2009). "Phylogeny of Annelida (Lophotrochozoa): total-evidence analysis of morphology and six genes". BMC Evolutionary Biology. 9 (1): 189. doi:10.1186/1471-2148-9-189. ISSN 1471-2148. PMC 2732625. PMID 19660115.
  6. Struck, Torsten H.; Paul, Christiane; Hill, Natascha; Hartmann, Stefanie; Hösel, Christoph; Kube, Michael; Lieb, Bernhard; Meyer, Achim; Tiedemann, Ralph; Purschke, Günter; Bleidorn, Christoph (2011). "Phylogenomic analyses unravel annelid evolution". Nature. 471 (7336): 95–98. doi:10.1038/nature09864. ISSN 0028-0836. PMID 21368831.
  7. Weigert, Anne; Helm, Conrad; Meyer, Matthias; Nickel, Birgit; Arendt, Detlev; Hausdorf, Bernhard; Santos, Scott R.; Halanych, Kenneth M.; Purschke, Günter; Bleidorn, Christoph; Struck, Torsten H. (2014). "Illuminating the Base of the Annelid Tree Using Transcriptomics". Molecular Biology and Evolution. 31 (6): 1391–1401. doi:10.1093/molbev/msu080. ISSN 1537-1719.
  8. Erséus et al. (2008)
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References
  • Erséus, Christer; Wetzel, Mark J. & Gustavsson, Lena (2008): ICZN rules – a farewell to Tubificidae (Annelida, Clitellata). Zootaxa 1744: 66–68. PDF fulltext
  • Reichardt, Anna Katharina (2006): Systematische Zoologie.
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