Phylogeny of basal descendants of cocoon-forming annelids (Clitellata)

Molecular analyses have been unsuccessful in placing the clitellates among annelids. Available morphological characters are ordered into congruent transformation series according to Hennigian principles. Orbiniidae is chosen as the outgroup. I propose the following phylogeny [number of apomorphies within brackets]: (Clitellata sensu lato [6] (Questidae [6] + Apodadrilida, new [2] (Parergodrilidae [3] (Stygocapitella [3] + Parergodrilus [4]) + Oligochaeta [7] (Aphanoneura [2] (Aeolosomatidae [2] + Potamodrilidae [3]) + Dorsopharyngea, new [7] (Hrabeiellidae, new [6] + Euclitellata [5])))). Worms with ring-like clitella surrounding the body almost completely are herein renamed Euclitellata. Clitellata is expanded to include all annelids that, in addition to producing cocoons, have sperm receptacles and a restricted number of gonads. Oligochaeta is established as a valid clade for the hermaphroditic annelids bearing few or no chaetae. Questidae, Parergodrilidae, and Hrabeiella belong to Clitellata. Terrestrial "polychaetes", such as Parergodrilus and Hrabeiella, are actually clitellates. Hrabeiella is the sister group of Euclitellata, despite a secondary loss of spermathecae and clitellum. Aphanoneura belong to Oligochaeta.

Phylogeny of basal descendants of cocoon-forming annelids (Clitellata)

Molecular analyses have been unsuccessful in placing the clitellates among annelids. Available morphological characters are ordered into congruent transformation series according to Hennigian principles. Orbiniidae is chosen as the outgroup. I propose the following phylogeny [number of apomorphies within brackets]: (Clitellata sensu lato [6] (Questidae [6] + Apodadrilida, new [2] (Parergodrilidae [3] (Stygocapitella [3] + Parergodrilus [4]) + Oligochaeta [7] (Aphanoneura [2] (Aeolosomatidae [2] + Potamodrilidae [3]) + Dorsopharyngea, new [7] (Hrabeiellidae, new [6] + Euclitellata [5])))). Worms with ring-like clitella surrounding the body almost completely are herein renamed Euclitellata. Clitellata is expanded to include all annelids that, in addition to producing cocoons, have sperm receptacles and a restricted number of gonads. Oligochaeta is established as a valid clade for the hermaphroditic annelids bearing few or no chaetae. Questidae, Parergodrilidae, and Hrabeiella belong to Clitellata. Terrestrial "polychaetes", such as Parergodrilus and Hrabeiella, are actually clitellates. Hrabeiella is the sister group of Euclitellata, despite a secondary loss of spermathecae and clitellum. Aphanoneura belong to Oligochaeta.

___

  • Aguinaldo, A.M.A., Turbeville, J.M., Linford, L.S., Rivera, M.C., Garey, J.R., Raff , R.A. and Lake, J.A. 1997. Evidence for a clade of nematodes, arthropods and other moulting animals. Nature 387: 489-493.
  • Aiyer, K.S.P. 1926. Notes on the aquatic Oligochaeta of Travancore, II. Ann. Mag. Nat. Hist. Ser. 9 18: 131-142.
  • Aiyar, R.G. 1935. Hermaphroditism in Lycastis indica (Southern). Current Science Bangalore 3: 367-368.
  • Almeida, W.O. and Christoff ersen, M.L. 2001. Análise cladística dos grupos basais de Metameria: uma nova proposta para o posicionamento dos Arthropoda e grupos afi ns entre os poliquetos errantes. Segunda Edição. Serie Teses, Dissertações e Monografi as, 1. Holos, Ribeirão Preto, São Paulo.
  • Almeida, W.O., Christoff ersen, M.L., Amorim, D.S., Garraff oni, A.R.S. and Silva, G.S. 2003. Polychaeta, Annelida and Articulata are not monophyletic: articulating the Metameria (Metazoa, Coelomata). Rev. Bras. Zool. 20: 23-57.
  • Amorim, D.S. 1997. Elementos básicos de sistemática fi logenética (2a edição, revista e ampliada). Holos, Ribeirão Preto, São Paulo.
  • Audouin, J.V. and Milne Edwards, H. 1833. Classifi cation des annélides, et description de celles qui habitent les côtes de la France. Ann.Sci. Nat. Paris Ser. 1 29: 195-269, pls. 13-18.
  • Ax, P. and Bunke, D. 1967. Das Genitalsystem der Aeolosomatidae mit phylogenetisch ursprünglichen Organisationzügen für die Oligochaeten. Naturwissenschaft en 54: 222-225.
  • Baltzer, F. 1931. Entwicklungsmechanische Untersuchungen an Entwicklungsgeschwindigkeit und des Entwicklungsgrades der männlichen Larve von der Dauer des Rüsselparasitismus. Rev. Suisse Zool. 38: 361-371.
  • Bartolomaeus, T., Purschke, G. and Hausen, H. 2005. Polychaete phylogeny based on morphological data—a comparison of current attempts. Hydrobiologia 535: 341-356.
  • Bartolomaeus, T. and Quast, B. 2005. Structure and development of nephridia in Annelida and related taxa. Hydrobiologia 535: 139-165.
  • Beddard, F.E. 1891. Th e classifi cation and distribution of earthworms. Proc. Roy. Phys. Soc. Edinburgh 10: 235-290, pls. 13-14.
  • Beddard, F.E. 1895. A monograph of the order Oligochaeta. Clarendon Press, Oxford.
  • Beddard, F.E. 1901. Earthworms and leeches. In: Th e Cambridge Natural History, vol. 2. Chaetopoda (Eds., S.F. Harmer and A.E. Shipley), MacMillan, New York.
  • Benham, W.B. 1890. An attempt to classify earthworms. Quart. J. Micr. Sci. 31: 201-315.
  • Blake, J.A. 2000. A new genus and species of polychaete worm (Family Orbiniidae) from methane seeps in the Gulf of Mexico, with a review of the systematics and phylogenetic interrelationships of the genera of Orbiniidae. Cahiers de Biologie Marine 41: 435-449.
  • Blanchard, E. 1849. Fauna Chilena. Anulares. In: Historia física y política de Chile. Segun documentos adquiridos en esta republica durante doce ańos de residencia en alla. Zoologia, 3 (ed. P.C. Gay), Paris, pp. 9-52, pls. 1-2 in Atlas.
  • Bleidorn, C. 2005. Phylogenetic relationships and evolution of Orbiniidae (Annelida, Polychaeta) based on molecular data. Zool. J. Linn. Soc. 144: 59-73.
  • Bleidorn, C., Hill, N., Erséus, C. and Tiedemann, R. 2009. On the role of character loss in orbiniid phylogeny (Annelida): Molecular vs. morphology. Mol. Phylogen. Evol. 52: 57-69.
  • Bleidorn, C., Vogt, L. and Bartolomaeus, T. 2003. New insights into polychaete phylogeny (Annelida) inferred from 18S rDNA sequences. Mol. Phylogen. Evol. 29: 279-288.
  • Borchiellini, C., Boury-Esnmault, N., Vacelet, J. and Le Parco, Y. 1998. Phylogenetic analysis of the Hsp70 sequences reveals the monophyly of Metazoa and specifi c phylogenetic relationships between animals and fungi. Mol. Biol. and Evol. 15: 647-655.
  • Bretscher, K. 1901. Beobachtung über Oligochaeten der Schweiz. Revue Suisse de Zoologie 9: 189-223, pl. 14.
  • Brinkhurst, R.O. 1971. Phylogeny and classifi cation. Part 1. In: Aquatic Oligochaeta of the world (Eds., R.O. Brinkhurst and B.G.M. Jamieson), Oliver & Boyd, London, pp. 165-177.
  • Brinkhurst, R.O. 1982. Evolution in the Annelida. Can. J. Zool. 60: 1043-1059.
  • Brinkhurst, R.O. 1994. Evolutionary relationships within the Clitellata: an update. Megadrilogica 5: 109-112.
  • Brinkhurst, R.O. 1999. Retrospect and prospect: refl ections on forty years of study of aquatic oligochaetes. Hydrobiologia 406: 9-19.
  • Brinkhurst, R.O. and Gelder, S.R. 1989. Did the lumbriculids provide the ancestors of the branchiobdellidans, acanthobdellidans and leeches? Hydrobiologia 180: 7-15.
  • Brinkhurst, R.O. and Jamieson, B.G.M. 1971. Aquatic Oligochaeta of the world. Oliver & Boyd, Edinburgh.
  • Brinkhurst, R.O. and Nemec, A.F.L. 1987. A comparison of phenetic and phylogenetic methods applied to the systematics of Oligochaeta. Hydrobiologia 155: 65-74.
  • Brinkhurst, R.O. and Wetzel, M.T. 1984. Aquatic Oligochaeta of the world. Supplement. A catalogue of new freshwater species, descriptions and revisions. Can. Tech. Rep. Hydrogr. Ocean Sci. 44: 1-101.
  • Bullock, T.H. 1965. Annelida. Volume 1. Structure and function in the nervous system of invertebrates (Eds., T.H. Bullock and G.A. Horridge), Freeman and Company, San Francisco, pp. 661-789.
  • Bunke, D. 1967. Zur Morphologie und Systematik der Aeolosomatidae Beddard 1895 und Potamodrilidae nov. fam. (Oligochaeta). Zool. Jb. Abt. Syst. Ökol. Geogr. Tiere 94: 187-368.
  • Bunke, D. 1985. Ultrastructure of the spermatozoon and spermiogenesis in the interstitial annelid Potamodrilus fl uviatilis. J. Morph. 185: 203-216.
  • Bunke, D. 1986. Ultrastructural investigations on the spermatozoon and its genesis in Aeolosoma litorale with considerations on the phylogenetic implications for the Aeolosomatidae (Annelida). J. Ultr. Mol. Struct. Res. 95: 113-130.
  • Bunke, D. 1988. Aeolosomatidae and Potamodrilidae. In: Introduction to the study of meiofauna (Eds., R.P. Higgins and H.Th iel), Smithsonian Institution Press, Washington, D.C. pp.
  • Bunke, D. 1994. Ultrastructure of the metanephridial system in Aeolosoma bengalense (Annelida). Zoomorphology (Berlin) 114: 247-258.
  • Bunke, D. 1998. Ultrastructure of the nephrostome in Enchytraeus albidus (Annelida, Clitellata). Zoomorphology (Berlin) 118: 177-182.
  • Bunke, D. 2000. Ultrastructure of the preseptal part of metanephridia in Nais variabilis and Dero digitata (Annelida, Clitellata). Zoomorphology (Berlin) 120: 39-46.
  • Bunke, D. 2003a. Early development of metanephridia in the caudal budding zone of a clitellate annelid, Dero digitata (Naidida): an electron-microscopical study. Act. Zool. 84: 87-97.
  • Bunke, D. 2003b. Early development of metanephridia in the caudal budding zone of a clitellate annelid (correction of vol. 84, p. 87, 2002). Act. Zool. 84: 317-320.
  • Čekanovskaja, O.V. 1959. On the Oligochaeta of the water-bodies of Central Asia (Fergen Valley and River Murgab). Zool. Zh. 38: 1152-1162.
  • Cerruti, A. 1909. Contributo all’ anatomia, biologia e sistematica delle Paraonidae (Levinseniidae) con particolare riguardo alle species del golfo di Napoli. Mitt. Zool. Stat. Neapel Berlin 19:
  • Chalupsky, J. 1992. Terrestrial Enchytraeidae (Oligochaeta) and Parergodrilidae (Polychaeta) from Sweden, with description of a new enchytraeid species. Zool. Scr. 21: 133-150.
  • Chapman, G. 1965. Th e egg cocoons of Scoloplos armiger O. F. Müller. Biol. Bull. Mar. Biol. Lab. Woods Hole 128: 189-197.
  • Christoff ersen, M.L. and Araşjo-de-Almeida, E. 1994. A phylogenetic framework of the Enterocoela (Metameria: Coelomata). Rev. Nord. Biol. 9: 173-208.
  • Coates, K.A. 1986. Redescription of the oligochaete genus Propappus, and diagnosis of the new family Propappidae (Annelida: Oligochaeta). Proc. Biol. Soc. Wash. 99, 3: 417-428.
  • Colgan, D.J., Hutchings, P.A. and Braune, M. 2006. A multigene framework for polychaete phylogenetic studies. Org. Div. Evol. 6: 220-235.
  • Cook, D.G. 1971. Anatomy. Microdriles. In: Aquatic Oligochaeta of the world. (Eds., R.O. Brinkhurst and B.G.M. Jamieson), Oliver and Boyd, Edinburgh, pp. 7-31.
  • Corrêa, D.D. 1948. A polychaete from the Amazon region. Bol. Fac. Filos. Ciênc. Letr. Univ. Săo Paulo Zoologia 13: 245-257, pls 1-2.
  • Cuvier, G. 1812. Sur un nouveau rapprochement a établir entre les classes qui composent le regne animal. Ann. Mus. Hist. Nat. Paris 19: 73-84.
  • Day, J.H. 1967. A monograph on the Polychaeta of southern Africa, part. 2: Sedentaria. Trustees of the British Museum of Natural History, London, pp. 459-878.
  • Des Châtelliers, M.C., Juget, J., Lafont, M. and Martin, P. 2009. Subterranean aquatic Oligochaeta. Freshw. Biol. 54: 678-690.
  • Dohle, W. 1999. Th e ancestral cleavage pattern of the clitellates and its phylogenetic derivations. Hydrobiologia 402: 267-290.
  • Ehrenberg, C.G. 1828. Symbolae physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegyptum, Nubiam, Dongalam, Syriam, Arabiam et Habessiniam. Pars Zoologia; Animalia evertebrata, exclusis insectis. Berlin.
  • Ehrenberg, C.G. 1831. Symbolae physicae, seu Icones et descriptiones Corporum Naturalium novorum aut minus cognitorum, quae ex itineribus per Libyam, Aegyptum, Nubiam, Dongalam, Syriam, Arabiam et Habessinian. Pars Zoologica; Animalia evertebrata, exclusis insectis. Berlin.
  • Eibye-Jacobsen, D. and Nielsen, C. 1996. Th e rearticulation of annelids. Zool. Scr. 253: 275-282.
  • Eisig, H. 1914. Zur Systematik, Anatomie und Morphologie der Ariciiden nebst Beiträgen zur generallen Systematik. Mitt. Zool. Stat. Neapel 21: 153-600, pls. 10-27.
  • Erséus, C. 1999. Parvidrilus strayeri, a new genus and species, an enigmatic interstitial clitellate from underground waters in Alabama. Proc. Biol. Soc. Wash. 112: 327-337.
  • Erséus, C. 2005. Phylogeny of oligochaeteous Clitellata. Hydrobiologia 535: 357-372.
  • Erséus, C. and Källersjö, M. 2004. 18S rDNA phylogeny of Clitellata (Annelida). Zool. Scr. 33, no. 2: 187-196.
  • Erséus, C., Prestegaard, T. and Källersjö, M. 2000. Phylogenetic analysis of Tubifi cidae (Annelida, Clitellata) based on 18S and rDNA sequences. Mol. Phylogen. Evol. 15: 381-389.
  • Erséus, C. and Rota, E. 1998. Polychaetes (bristle worms) on dry land. Faun. Flor. 93: 97-104.
  • Fauchald, K. 1977. Th e polychaete worms. Defi nitions and keys to the orders, families and genera. Sci. Ser. Nat. Hist. Mus. Los Angeles County 28: 1-188.
  • Fauchald, K. and Rouse, G. 1997. Polychaete systematics: Past and present. Zool. Scr. 26: 71-138.
  • Fauvel, P. 1927. Polychaetes sedentaires. Addenda aux errantes, archiannelides, myzostomaires. In: Faune de France (ed. A. Vandel), Paul Lechevalier, Paris, pp. 413-551.
  • Ferraguti, M. 1982. Clitellata. In: Reproductive biology of invertebrates, vol. 3. Spermatogenesis and sperm function (ed. K. G. Adiyodi) Wiley & Sons, London.
  • Ferraguti, M. 1984. Th e comparative ultrastructure of sperm fl agella central sheath in Clitellata reveals a new autapomorphy of the group. Zool. Scr. 13: 201-207.
  • Ferraguti, M. 2000. Euclitellata. In: Reproductive biology of invertebrates. Volume IX, Part B. Progress in male gamete ultrastructure and phylogeny (Ed. B.G.M. Jamieson), New Delhi, pp. 125-182.
  • Ferraguti, M., and C. Erséus. 1999. Sperm types and their use for a phylogenetic analysis of aquatic clitellates. Hydrobiologia 402:
  • Feuerborn, H.J. 1931. Ein Rhizocephale und zwei Polychaeten aus dem Süsswasser von Java und Sumatra. Verh. Int. Ver. theor. ang. Limnol. Stuttgart 5: 618-660.
  • Franzén, A. 1970. Phylogenetic aspects of the morphology of spermatozoa and spermiogenesis. In: Comparative spermatology (ed. B. Baccetti), Academic Press, New York, pp. 29-55.
  • Franzén, A. 1977. Ultrastructure of spermatids and spermatozoa in Archiannelida. Zoon, Uppsala 5: 97-105.
  • Garraff oni, A.R.S. and Amorim, D.S. 2003. Análise fi logenética de Questidae e Clitellata: o problema da parafi lia de “Polychaeta”. Iheringia 93: 97-109.
  • Gelder, S.R. and Uglow, R.F. 1973. Feeding and gut structure in Nerilla antennata (Annelida: Archiannelida). J. Zool. London
  • Giere, A., Ebbe, B. and Erséus, C. 2007. Questa (Annelida, Polychaeta, Orbiniidae) from Pacifi c regions − new species and reassessment of the genus Periquesta. Org. Div. & Evol. 7:
  • Giere, O.W. and Erséus, C. 1998. A systematic account of the Questidae (Annelida, Polychaeta), with description of new taxa. Zool. Scr. 27: 345-360.
  • Giere, O.W. and Riser, N.W. 1981. Questidae - polychaetes with oligochaetoid morphology and development. Zool. Scr. 10: 95- 103.
  • Glasby, C.J. 1999. Th e Namanereidinae (Polychaeta: Nereididae) Part 1 Taxonomy and phylogeny Part 2 Cladistic biogeography. Rec. Austral. Mus. Suppl. 25: 1-144.
  • Glasby, C.J. 2000. Questidae. In: Fauna of Australia, Vol. 4. Annelida through Tunicata (Eds., C.J. Glasby and P.A. Hutchings) Australian Biological Resources Study, Sydney.
  • Glasby, C.J., Kitching, R.L. and Ryan, P.A. 1990. Taxonomy of the arboreal polychaete Lycastopsis cataractarum Feuerbeoorn (Namanereidinae: Nereididae), with a discussion of the feeding biology of the species. J. Nat. Hist. 24: 341-350.
  • Glasby, C.J. and Timm, T. 2008. Global diversity of polychaetes (Polychaeta; Annelida) in freshwater. Hydrobiologia 595: 107- 115.
  • Glasby, C.J. Timm, T., Muir, A.I. and Gil, J. 2009. Catalogue of non- marine Polychaeta (Annelida) of the world. Zootaxa 2070: 1-52.
  • Gluzman de Pascar, C. 1997. Sperm cells in Aeolosoma marcusi (Annelida, Oligochaeta). Biocell 21: 137-142.
  • Goodrich, E.S. 1945. Th e study of nephridia and genital ducts since 1895. Part I. Quart. J. Micr. Sci. 86: 113-392.
  • Graefe, U. 1975. Adenodrilus Graefe and Parergodrilus Reisinger two aberrant Annelida in forest soils. Resumen II Coloquio Europeo sobre Anélidos del Suelo, Jaca, 1-4 Septiembre de
  • Graefe, U. 1977. Adenodrilus Graefe (in press) and Parergodrilus Reisinger two aberrant Annelida in forest soils. Publ. Cent. Per. Biol. Exp. Jaca 9: 25.
  • Gravier, C. 1901. Sur deux nouvelles espèces du genre Lycastis Savigný, Aud. et Edw. rev., de la Guyane française. Bull. Mus. Nat. Hist. Nat. 7: 397-402.
  • Grube, A.E. 1850. Die Familien der Anneliden. Arch. Naturg. Berlin 16: 249-364.
  • Grube, A.E. 1851. Die Familien der Anneliden. Arch. Naturg., Berlin 16: 249-364.
  • Grube, A.E. 1872. Über die Gattung Lycastis und ein Paar neue Arten derselben. Jahr.-Ber. Schles. Ges. 49: 47-48.
  • Haeckel, E. 1874. Die Gastraea-Th eorie, die phylogenetische Classifi cation des Th ierreichs und die Homologie der Keimblätter. Z. Naturw. 8: 1-55.
  • Hämmerling, J. 1924. Die ungeschlechtliche Fortpfl anzung und Regeneration bei Aeolosoma hemprichii. Zool. Jahrb. Allg. Zool. Phys. Tiere 41: 581-656, pls. 16-19.
  • Hall, K.A., Hutchings, P.A. and Colgan, D.J. 2004. Further phylogenetic studies of the Polychaeta using 18S rDNA sequence data. J. Mar. Biol. Ass. U.K. 84: 949-960.
  • Harman, W.J. and Loden, M.S. 1984. Capilloventer atlanticus gen. et sp. n. member of a new family of marine Oligochaeta from Brazil. Hydrobiologia 115: 51-54.
  • Hartman, O. 1942. Th e polychaetous Annelida. Report on the scientifi c results of the Atlantis expeditions to the West Indies under the joint auspices of the University of Havana and Harvard University. Mem. Soc. Cubana Hist. Nat. 16: 89-104, pls. 8-9.
  • Hartman, O. 1959. Capitellidae and Nereididae (marine annelids) from the Gulf side of Florida with a review of freshwater Nereidae. Bull. Mar. Sci. Gulf Caribb. 9: 153-168.
  • Hartman, O. 1966. Quantitative survey of the benthos of San Pedro Basin, southern California, pt. II. Final results and conclusions. Allan Hancock Found. Pac. Exp. 19: 187-456, 13 pls.
  • Hartmann-Schröder, G. 1996. Annelida, Borstenwürmer, Polychaeta. Die Tierwelt Deutschlands 58 (ed. H. Schumann), Gustav Fischer, Jena.
  • Hausdorf, B. 2000. Early evolution of the Bilateria. Syst. Biol. 49: 130- 142.
  • Hennig, W. 1950. Grundzuge einer Th eorie der phylogenetischen Systematik. Deutscher Zentralverlag, Berlin.
  • Hennig, W. 1966. Phylogenetic systematics. Univ. Illinois Press, Urbana.
  • Herlant-Meewis, H. 1950. Cyst-formation in Aeolosoma hemprichi (Ehr.). Biol. Bull. Mar. Biol. Lab. Woods Hole 99: 173-180.
  • Hessling, R. and Purschke, G. 2000. Immunohistochemical (cLSM) and ultrastructural analysis of the central nervous system and sense organs in Aeolosoma hemprichi (Annelida, Aeolosomatidae). Zoomorphology Berlin 120: 65-78.
  • Hessling, R. and Westheide, W. 1999. CLSM analysis of development and structure of the nervous system of Enchytraeus crypticus (“Oligochaeta”, Enchytraeidae). Zoomorphology Berlin 119: 37-47.
  • Hobson, K.D. 1970. Novaquesta trifurcata, a new genus and species of the family Questidae (Annelida, Polychaeta) from Cape Cod Bay, Massachusetts. Proc. Biol. Soc. Wash. 83: 191-194.
  • Hobson, K.D. and Banse, K. 1981. Sedentariate and archiannelid polychaetes of British Columbia and Washington. Can. Bull. Fish. Aquatic Sci. 209: 1-132.
  • Holt, P.C. 1965. Th e systematic position of the Branchiobdellidae (Annelida: Clitellata). Syst. Zool. 14: 25-32.
  • Hrabĕ, S. 1981. Th e freshwater Oligochaeta (Annelida) of Czechoslovakia. Act. Univ. Carol. Biol. 1-2: 1-167. [in Czech]
  • Huxley, T.H. 1874. On classifi cation of the animal kingdom. Nature
  • Ivanov, A.V. 1949. A new representative of the class Pogonophora]. Zool. Zh. 28: 79-84. [in Russian]
  • Jamieson, B.G.M. 1971. Phylogeny and classifi cation. Part 2. In: Aquatic Oligochaeta of the world (Eds., R.O. Brinkhurst and B.G.M. Jamieson), Oliver & Boyd, London, pp. 177-192.
  • Jamieson, B.G.M. 1980. Preliminary discussion of an Hennigian analysis of the phylogeny and systematics of opisthoporous oligochaetes. Rev. Ecol. Biol. Sol 17: 261-275.
  • Jamieson, B.G.M. 1981. Th e ultrastructure of the Oligochaeta. Academic Press, London.
  • Jamieson, B.G.M. 1983a. Th e ultrastructure of the spermatozoon of the oligochaetoid polychaete Questa sp (Questidae, Annelida) and its phylogenetic signifi cance. J. Ultr. Res. 84: 238-251.
  • Jamieson, B.G.M. 1983b. Spermiogenesis in the oligochaetoid polychaete Questa (Annelida, Questidae). Zool. Scr. 12: 179- 186.
  • Jamieson, B.G.M. 1983c. Spermatozoal ultrastructure - evolution and congruence with a holomorphological phylogeny of the Oligochaeta (Annelida). Zool. Scr. 12: 107-114.
  • Jamieson, B.G.M. 1988. On the phylogeny and higher classifi cation of the Oligochaeta. Cladistics 4: 367-410.
  • Jamieson, B.G.M. 1992. Oligochaeta. Microscopic anatomy of invertebrates. Vol. 7. Annelida (Eds., F.W. Harrison and S.L. Gardiner), Wiley-Liss, New York, pp. 217-322.
  • Jamieson, B.G.M. and Rouse, G.W. 1989. Th e spermatozoa of the Polychaeta (Annelida) - an ultrastructural review. Biol. Rev. Cambridge Phil. Soc. 64: 93-157.
  • Jamieson, B.G.M., Tillier, S., Tillier, A., Justi, J.L., Ling, E., James, S., McDonald, K. and Hugall, A.F. 2002. Phylogeny of the Megascolecidae and Crassiclitellata (Annelida, Oligochaeta) combined versus partitioned analysis using nuclear (28S) and mitochondrial (12S, 16S) rDNA. Zoosystema 24: 707-734.
  • Jamieson, B.G.M. and Webb, R.T. 1984. Th e morphology, spermatozoal ultrastructure and phylogenetic affi nities of a new species of questid (Polychaeta, Annelida). In: Proceedings of the First international polychaete conference, Sydney (Ed. P.A. Hutschings), Th e Linnean Society of New South Wales, Sydney, pp. 21-34.
  • Jans, W. and Römbke, J. 1989. Funde eines terrestrischen Polychaeten (Annelida) in Wäldern Baden-Württembergs. Carolinea 47: 158-162.
  • Jenner, R.A. 2004. Towards a phylogeny of the Metazoa: evaluating alternative phylogenetic positions of Platyhelminthes, Nemertea, and Gnathostomulida, with a critical reappraisal of cladistic characters. Contr. Zool. 73: 3-163.
  • Jenner, R.A. 2006. Challenging received wisdoms: some contributions of the new microscopy to the new animal phylogeny. Int. Comp. Biol. 46: 93-103.
  • Jennings, J.B. and Gelder, S.R. 1969. Feeding and digestion in Dinophilus gyrociliatus (Annelida, Archiannelida). J. Zool. 158: 441-457.
  • Jördens, J., Struck, T. and Purschke, G. 2004. Phylogenetic inference regarding Parergodrilidae and Hrabeiella periglandulata (‘Polychaeta’, Annelida). J. Zool. Syst. Evol. Res. 42: 270-280.
  • Johnson, H.P. 1903. Fresh-water nereids from the Pacifi c coast and Hawaii, with remarks on fresh-water Polychaeta in general. In: Historia física y política de Chile. Segun documentos adquiridos en esta republica durante doce ańos de residencia en alla (Ed., P.C. Gay), Mark Anniversary Volume, New York, pp. 205-222, pls. 16-17.
  • Johnson, H.P. 1908. Lycastis quadraticeps, an hermaphrodite nereid with gigantic ova. Biol. Bull. Woods Hole 14: 371-386.
  • Karling, T.G. 1958. Zur Kenntnis von Stygocapitella subterranea Knöllner und Parergodrilus heideri Reisinger (Annelida). Ark. Zool. Kungl. Sv. Vet. 11: 307-342, pls. 1-4.
  • Knöllner, F. 1934. Die Tiere des Küstengrundwassers bei Schilksee (Kieler Bucht). 5. Stygocapitella subterranea nov. gen. nov. spec. Schr. Naturwiss. Ver. Schles. Holstein 20: 468-472.
  • Kojima, S. 1998. Paraphyletic status of Polychaeta suggested by phylogenetic analysis based on the amino acid sequences of elongation factor - 1α. Mol. Phylogenet. Evol. 9: 255-261.
  • Lamarck, J.B.P.M. 1803. Biologie, ou considérations sur la nature, les facultés, les développements et l’origines des corps vivants. Unpublished sketches for a book.
  • Lamarck, J.B.P.B. 1818. Histoire naturelle de Animaux sans vetèbres présentant les caracteres généraux et particuliers de ces animaux, lur distribution, leurs classes, leurs families, leurs genres, et la citation des principales espèces qui s´y rapportent; précédée d´une Introduction off rant la détermination des caractères essentiels de l´animal, sa distinction du végétal et des autres corps naturels, enfi n, l´exposition des principes fondamenteux de la Zoologie. Ed. 1, t. 3. Verdière, Paris.
  • Lana, P.C. 1987. Padrões de distribuição geográfi ca dos poliquetas errantes (Annelida: Polychaeta) do Estado do Paraná. Cienc. Cult. 39: 1060-1063.
  • Lasserre, P. 1971. Oligochaeta from the marine meiobenthos. Taxonomy and ecology. Smiths. Contr. Zool. 76: 71-86.
  • Lasserre, P. 1975. Clitellata. In: Reproduction of marine invertebrates. Vol. 3: Annelids and echiurans (Ed. A.C. Giese and J.S.P. Pearse), Academic Press, New York, pp. 215-272.
  • Lastochkin, D.A. 1935. New names for the genus Stephensoniella Lastochkin (Oligochaeta). Ann. Mag. Nat. Hist. Ser. 10 16: 488.
  • Levinsen, G.M.R. 1884. Systematisk-geografi sk Oversigt over de nordiske Annulata, Gephyrea, Chaetognathi og Balanoglossi, II. Vid. Medd. Nat. Foren. Kjİb. 9-11: 9-350, pls. 1-7.
  • Livanow, N. 1931. Die Organisation der Hirudineen und die Beziehungen zu den Oligochaeten. Erg. Fortschr. Zool. 7: 284- 290.
  • Manum, S. B., Bose, M.N. and Sawyer, R.T. 1991. Clitellate cocoons in freshwater deposits since the Triassic. Zool. Scr. 20: 347-366.
  • Marcus, E. 1944. Sobre Oligochaeta límnicos do Brasil. Bol. Fac. Filos. Cienc. Letr. Univ. S. Paulo Zool. N. 8 43: 5-102.
  • Marescalchi, O., Gugnali, A. and Falconi, R. 2008. First report on the chromosomes of Aeolosoma viride and Aeolosoma hemprichi (Aeolosomatidae; Annelida). Zool. Sci. 25: 904-906.
  • Marotta, R., Ferraguti, M., Erseus, C. and Gustavsson, L.M. 2008. Combined-data phylogenetics and character evolution of Clitellata (Annelida) using 18s rDNA and morphology. Zool. J Linn. Soc. 154: 1-26.
  • Marotta, R., Ferraguti, M. and Martin, P. 2003. Spermiogenesis and seminal receptacles in Aeolosoma singulare (Annelida, Polychaeta, Aeolosomatidae). Ital. J. Zool. 70: 123-132.
  • Martin, P. 2001. On the origin of the Hirudinea and the demise of the Oligochaeta. Proc. Roy. Soc. London Ser. B Biol. Sci. 268: 1089-1098.
  • Martin, P., Kaygorodova, I., Sherbakov, D.Y. and Verheyen, E. 2000. Rapidly evolving lineages impede the resolution of phylogenetic relationships among Clitellata (Annelida). Morph. Phyl. Evol. 15: 355-368.
  • Martin, P., Martinez-Ansemil, E., Pinder, A., Timm, T. and Wetzel, M.J. 2008. Global diversity of oligochaetous clitellates (“Oligochaeta”; Clitellata) in freshwater. Hydrobiologia 595: 117-127.
  • McHugh, D. 1997. Molecular evidence that echiurans and pogonophorans are derived annelids. Proc. Nat. Acad. Sci. U.S.A. 94: 8006-8009.
  • McHugh, D. 2000. Molecular phylogeny of the Annelida. Can. J. Zool. 78: 1873-1884.
  • McHugh, D. 2005. Molecular systematics of polychaetes (Annelida). Hydrobiologia 535: 309-318.
  • Metschnikoff , E. 1881. Über die systematische Stellung von Balanoglossus. Zool. Anz. 4: 139-143, 153-157.
  • Meyer, A. 1927. Ist Parergodrilus heideri (Reisinger) ein Archiannelide? Zool. Anz. 72: 19-35.
  • Michaelsen, W. 1900. Oligochaeta. Das Tierreich, 10. R. Friedländer and Sohn, Berlin.
  • Michaelsen, W. 1919. Über die Beziehungen der Hirudinea zu den Oligochaeten. Mitt. Zool. Staatsinst. Zool. Mus. Hamburg 36: 51-73.
  • Michaelsen, W. 1926. Schmarotzende Oligochäten nebst Erörterung über verwandschaft liche Beziehungen der Archioligochäten. Mitt. Staatsinst. Zool. Mu. Hamburg 42: 91-103.
  • Michaelsen, W. 1928. Ordnung der Clitellata: Oligochaeta = Regenwürmer und Verwandte. In: Handbuch der Zoologie, Band 2 (2) Leif 8 und Nachtrag (Eds., W. Kükenthal and T. Krumbach), Walter de Gruyter, Berlin.
  • Michaelsen, W. 1930. Oligochaeta - Regenwürmer und Verwandte: Nachtrag. Handb. d. Zool. 2, no. 15, Part 8, Suppl. 1: 116-117 (1930) Suppl. 2: 118 (1932).
  • Michaelsen, W. 1934. Dritte Klasse der Vermes polymera (Annelida) Clitellata = Gürtelwürmer. In: Handb d. Zool., Band 2, Leif 8 (Eds., W. Kükenthal and T. Krumbach), Walter de Gruyter, Berlin.
  • Moon, S.Y., Kim, C.B., Gelder, S.R. and Kim, W. 1996. Phylogenetic position of the aberrant branchiobdellidans and aphanoneurans within the Annelida as derived from 18S ribosomal RNA gene sequences. Hydrobiologia 324: 229-236.
  • Müller, M.C.M. 2006. Polychaete nervous systems: ground pattern and variations - Cls microscopy and the importance of novel characteristics in phylogenetic analysis. Integr. Comp. Biol. 46: 125-133.
  • Müller, O.F. 1774. Vermium terrestrium et fl uviatilum seu animalium infusorium, helminthicorum et testaceorum non marinorum, succincta historia Volume 1 Pars 2. Heineck and Faber, Havniae et Lipsiae.
  • Müller, O.F. 1776. Zoologiae Danicae Prodomus, seu Animalium Daniae et Norvegiae Indigenorum Characters, nomina et synonyma imprimis popularium, Volume 1. Havniae et Lipsiae.
  • Nielsen, C. 2001. Animal evolution: interrelatinships of the living phyla. Second Edition. Oxford Univ. Press, Oxford.
  • Nielsen, C.O. and Christensen, B. 1961. Th e Enchytraeidae. Critical revision and taxonomy of European species Supplement 1. Nat. Jutl. 10: 1-23.
  • Olive, P.J.W. 1983. Annelida − Polychaeta. In: Reproductive biology of invertebrates. Spermatogenesis and sperm function. Vol. 2 (Eds., K.G. Adiyodi and R.G. Adiyodi), Wiley, New York.
  • Omodeo, P. 1998. History of Clitellata. Ital. J. Zool. 65: 51-73.
  • Orrhage, L. 1995. On the innervation and homologues of the anterior end appendages of the Eunicea (Polychaeta), with a tentative outline of the fundamental constitution of the cephalic nervous system of polychaetes. Acta Zool. Stockholm 76: 229-248.
  • Pfl ugfelder, O. 1933. Landpolychaeten aus Niederländlisch-Indien. Ergebnisse ders Sunda-Expedition der Notgemeinschaft der deutschen Wissenschaft 1929-30. Zool. Anz. Leipzig 105: 65- 76.
  • Philippe, H., Derelle, R., Lopez, P., Pick, K., Borchiellini, C., Boury- Esnault, N., Vacelet, J., Renard, E., Houliston, E., Queinnec, E., Da Silva, C., Wincker, P., Le Guyader, H., Leys, S., Jackson, D.J., Schreiber, F., Erpenbeck, D., Morgenstern, B., Worheide, G. and Manuel, M. 2009. Phylogenomics revives traditional views on deep Animal Relationships. Curr. Biol. 19: 706-712.
  • Pižl, V. and Chalupský, J. Jr. 1984. Hrabeiella periglandulata gen. et sp. n. (Annelida) - A curious worm from Czechoslovakia. Vĕst. Čĕs. Společ. Zool. 48: 291-295.
  • Purschke, G. 1986. Ultrastructure of the nuchal organ in the interstitial polychaete Stygocapitella subterranea (Parergodrilidae). Zool. Scr. 15: 13-20.
  • Purschke, G. 1987. Anatomy and ultrastructure of ventral pharyngeal organs and their phylogenetic importance in Polychaeta (Annelida). III. Th e pharynx of the Parergodrilidae. Zool. Jb. Abt. Anat. Ontog. Tiere 115: 331-362.
  • Purschke, G. 1993. Structure of the prostomial appendages and the central nervous system in the Protodrilida (Polychaeta). Zoomorphology Berlin 113: 1-20.
  • Purschke, G. 1997. Ultrastructure of nuchal organs in polychaetes (Annelida): new results and review. Act. Zool. Stockholm 78: 123-143.
  • Purschke, G. 1999. Terrestrial polychaetes - models for the evolution of the Clitellata (Annelida)? Hydrobiologia 406: 87-99.
  • Purschke, G. 2000. Sense organs and central nervous system in an enigmatic terrestrial polychaete, Hrabeiella periglandulata (Annelida) - implications for annelid evolution. Inv. Biol. 119: 329-341.
  • Purschke, G. 2002a. On the ground pattern of Annelida. Org. Div. Evol. 2: 181-196.
  • Purschke, G. 2002b. Male genital organs, spermatogenesis and spermatozoa in the enigmatic terrestrial polychaete Parergodrilus heideri (Annelida, Parergodrilidae). Zoomorphology 121: 125-138.
  • Purschke, G. 2003a. Ultrastructure of phaosomous photoreceptors in Stylaria lacustris (Naididae, ‘Oligochaeta’, Clitellata) and their importance for the position of the Clitellata in the phylogenetic system of the Annelida. J. Zool. Syst. Evol. Res. 41: 100-108.
  • Purschke, G. 2003b. Is Hrabeiella periglandulata (Annelida, “Polychaeta”) the sister group of Clitellata? Evidence from an ultrastructural analysis of the dorsal pharynx in H. periglandulata and Enchytraeus minutus (Annelida, Clitellata). Zoomorphology 122: 55-66.
  • Purschke, G. 2005. Ultrastructure of the nuchal organ in the interstitial polychaete Stygocapitella subterranea (Parergodrilidae). Zool. Scr. 15: 13-20.
  • Purschke, G. and Fursman, M.C. 2005. Spermatogenesis and spermatozoa in Stygocapitella subterranea (Annelida, Parergodrilidae), an enigmatic supralittoral polychaete. Zoomorphology 124: 137-148.
  • Purschke, G. and Hausen, H. 2007. Lateral organs in sedentary polychaetes (Annelida) - ultrastructure and phylogenetic signifi cance of an insuffi ciently known sense organ. Act. Zool. 88: 23-39.
  • Purschke, G. and Hessling, R. 2002. Analysis of the central nervous system and sense organs in Potamodrilus fl uviatilis (Annelida: Potamodrilidae). Zool. Anz. 241: 19-35.
  • Purschke, G., Hessling, R. and Westheide, W. 2000. Th e phylogenetic position of hte Clitellata and the Echiura - on the problematic assessment of absent characters. J. Zool. Syst. Evol. Res. 38: 165-173.
  • Purschke, G. and Jordens, J. 2007. Male genital organs in the eulittoral meiofaunal polychaete Stygocapitella subterranea (Annelida, Parergodrilidae): Ultrastructure, functional and phylogenetic signifi cance. Zoomorphology 126: 283-297.
  • Purschke, G. and Müller, M.C.M. 1996. Structure of prostomial photoreceptor-like sense organs in Protodriloides species (Polychaeta, Protodrilida). Cah. Biol. Mar. 37, no. 3: 205-19.
  • Purschke, G. and Müller, M.C.M. 2006. Evolution of body wall musculature. Int. Comp. Biol. 46: 497-507.
  • Purschke, G. and Tzetlin, A.B. 1996. Dorsolateral ciliary folds in the polychaete foregut: structure, prevalence and phylogenetic signifi cance. Act. Zool., Stockholm 77: 33-49.
  • Purschke, G., Westheide, W., Rohde, D. and Brinkhurst, R.O. 1993. Morphological reinvestigation and phylogenetic relationships of Acanthobdella peledina (Annelida, Clitellata). Zoomorphology, Berlin 113: 91-101.
  • Reisinger, E. 1925. Ein landbewohnender Archiannelide (Zugleich ein Beitrag zur Systematik der Archianneliden). Z. Ökol. Tiere A 3: 197-254.
  • Reisinger, E. 1929. Die systematische Stellung von Parergodrilus heideri Reisinger. Zool. Anz. 80: 12-20.
  • Reisinger, E. 1960. Die Lösung des Parergodrilus Problems. Z. Morph. Ökol. Tiere 48: 517-544.
  • Reynolds, J.W. and Cook, D.G. 1976. Nomenclatura Oligochaetologica. A catalogue of names, descriptions and type specimens of the Oligochaeta. Th e University of New Brunswick, Fredericton.
  • Reynolds, J.W. and Cook, D.G. 1981. Nomenclatura Oligochaetologica, Supplementum Primum.. A catalogue of names, descriptions and type specimens of the Oligochaeta. Th e University of New Brunswick, Fredericton.
  • Reynolds, J.W. and Cook, D.G. 1989. Nomenclatura Oligochaetologica, Supplementum Secundum. A catalogue of names, descriptions and type specimens of the Oligochaeta. Monographic Series of the New Brunswick Museum, Vol. 8. Th e University of New Brunswick, Fredericton.
  • Reynolds, J.W. and Cook, D.G. 1993. Nomenclatura Oligochaetologica, Supplementum Tertium. A catalogue of names, descriptions and type specimens of the Oligochaeta. Monographic Series of the New Brunswick Museum, Vol. 9. Th e University of New Brunswick, Fredericton.
  • Riser, N.W. 1980. Th e aberrant polychaet Stygocapitella from some American beaches. Wasman J. Biol. 38: 10-17.
  • Riser, N.W. 1984. General observations on the intertidal interstitial fauna of New Zealand. Tane 30: 239-250.
  • Rota, E. 1997. First Italian record of the terrestrial polychaete Parergodrilus heideri Reisinger, with anatomical and ecological notes. Ital. J. Zool. 64: 91-96.
  • Rota, E. 1998. Morphology and adaptations of Parergodrilus Reisinger and Hrabeiella Pižl & Chalupský, two enigmatic soil-dwelling annelids. Ital. J. Zool. 65: 75-84.
  • Rota, E. and Lupetti, P. 1997. An ultrastructural investigation of Hrabeiella Pižl & Chalupský, 1984 (Annelida). II. Th e spermatozoon. Tiss. Cell 29: 603-609.
  • Rota, E., Martin, P. and Erséus, C. 2001. Soil-dwelling polychaetes: enigmatic as ever? Some hints on their phylogenetic relationships as suggested by a maximum parsimony analysis of 18S rRNA gene sequences. Contr. Zool. 70: 127-138.
  • Rouse, G.W. 1998. Recent views on the status, delineation and classifi cation of the Annelida. Amer. Zool. 12: 1132-1137.
  • Rouse, G.W. and Fauchald, K. 1995. Th e articulation of annelids. Zool. Scr. 24: 269-301.
  • Rouse, G.W. and Fauchald, K. 1997. Cladistics and polychaetes. Zool. Scr. 2: 139-204.
  • Rouse, G.W. and Fauchald, K. 1998. Recent views on the status, delineation and classifi cation of the Annelida. Amer. Zool. 38:
  • Rouse, G.W. and Jamieson, B.G.M. 1987. An ultrastructural study of the spermatozoa of the polychaetes Eurythoe complanata (Amphinomidae), Clymenella sp and Micromaldane sp (Maldanidae), with defi nition of sperm types in relation to reproductive-biology. J. Submicr. Cyt. Path. 19: 573-584.
  • Rouse, G.W. and Pleijel, P. (Eds.). 2001. Polychaetes. Oxford Univ. Press, New York.
  • Rouse, G.W. and Pleijel, F. 2003. Problems in polychaete systematics. Hydrobiologia 496: 175-189.
  • Rouse, G.W., Pleijel, F. and McHugh, D. 2008. Annelida. Segmented worms: Bristleworms, ragworms, earthworms, leeches and their allies. Pp. 1-9. http://tolweb.org/tree?group=Annelida&c ontgroup=Bilateria
  • Rousset, V., Pleijel, F., Rouse, G.W., Erseus, C. and Siddall, M.E. 2007. A molecular phylogeny of annelids. Cladistics 23: 41-63.
  • Rousset, V., Plaisance, L., Erseus, C., Siddall, M.E. and Rouse, G.W. 2008. Evolution of habitat preference in Clitellata (Annelida). Biol. J. Linn. Soc. 95: 447-464.
  • Runganadhan, V. 1943. A review of the mode of breeding and development in brackish and fresh water polychaetes. Proceedings of the Th irtieth Indian Science Congress, Calcutta Pt. 3, pp. 72.
  • Ruttner-Kolisko, A. 1955. Rheomorpha neiswestnovae und Marinellina fl agellata zwei phylogenetisch interessante Würmtypen aus dem Süsswasserpsammon. Österr. Zool. Z. 6: 55-69.
  • Saint-Joseph, M. 1900. Sur quelques invertébrés marins des cotes du Sénégal (annélides polychètes, nématoide endoparasite d´annélide polychète et crustacé décapode parasite). Ann. Sci. Nat. Zool. Pal. 12: 217-248, pls. 8-9.
  • Sars, M. 1863. Geologiske og zoologiske lagttagelser, anstillede paa en Reise i en deel af Trondhjems Stift i Sommeren 1862. Nytt Mag. Naturw. 12: 253-340.
  • Sayers, C.W., Coleman, J. and Shain, D.H. 2009. Cell dynamics during cocoon secretion in the aquatic leech, Th eromyzon tessulatum (Annelida: Clitellata: Glossiphoniidae). Tiss. Cell 41 1: 35-42.
  • Schmidt, H. and Westheide, W. 2000. Are the meiofaunal polychaetes Hesionides arenaria and Stygocapitella subterranea true cosmopolitan species? Results of RAPD-PCR investigations . Zool. Scr. 29: 17-27.
  • Schmidt, P.J. 1897. Zur Kenntnis der Gattung Aeolosoma. C.R. Soc. Imp. Nat. St. Petersbourg 27: 161-163.
  • Semernoi, V.P. 1982. New species of oligochaetes from Lake Baikal. New data on the fauna of Lake Baikal (Ed., G.I. Galazii). Nauka, Novosibirsk, pp. 58-85. [in Russian]
  • Siddall, M.E., Apakupakul, K., Burreson, E.M., Coates, K.A., Erséus, C., Gelder, S.R., Källerjö, M. and Trapido-Rosenthal, H. 2001. Validating Livanow: Molecular data agree that leeches, branchiobdellidans and Acanthobdella peledina form a monophyletic group of oligochaetes. Mol. Phylogen. Evol. 21: 346-351.
  • Siddall, M.E. and Burreson, E.M. 1996. Leeches (Oligochaeta?: Euhirudinea), their phylogeny and the evolution of life-history strategies. Hydrobiologia 334: 277-285.
  • Solis-Weiss, V. and Fauchald, K. 1989. Orbiniidae (Annelida: Polychaeta) from mangrove root-mats in Belize, with a revision of protoariciin genera. Proc. Biol. Soc. Wash. 102: 772-792.
  • Southern, R. 1921. Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Mem. Indian Mus. 5: 563-659, pls. 19-31.
  • Stephenson, J. 1930. Th e Oligochaeta. Oxford University Press. Oxford.
  • Stolte, H.A. 1969. Oligochaeta. In: Klassen u. Ord. Tierreichs (Ed., H.G. Bronn), Bd IV, Abt. III: Annelides, 3 Buch: Oligochaeta. Lief 6 (1962): 984-1141; Lief 7 (1968): 1143-1218; Lief 8 (1960): 1284-1414. Akad. Verlags, Leipzig.
  • Storch, V. and Welch, U. 1972. Ultrastructure and histochemistry of air breathing polychaetes from mangrove swamps of Sumatra. Marine Biology 17: 137-143.
  • Struck, T.H. 2006. Progenetic species in polychaetes (Annelida) and problems assessing their phylogenetic affi liation. Int. Comp. Biol. 46: 558-568.
  • Struck, T., Hessling, R., and Purschke, G. 2002. Th e phylogenetic position of the Aeolosomatidae and Parergodrilidae, two enigmatic oligochaete-like taxa of the ‘Polychaeta’, based on molecular data from 18S rDNA sequences. J. Zool. Syst. Evol. Res. 40: 155-163.
  • Struck, T.H., Nesnidal, M.P., Purschke, G. and Halanych, K.M. 2008. Detecting possibly saturated positions in 18s and 28s sequences and their infl uence on phylogenetic reconstruction of Annelida (Lophotrochozoa). Mol. Phylogen. Evol. 48: 628-645.
  • Struck, T.H. and Purschke, G. 2005. Th e sister group relationship of Aeolosomatidae and Potamodrilidae (Annelida: “Polychaeta”) − a molecular phylogenetic approach based on 18S rDNA and cytochrome oxidase I. Zool. Anz. 243: 281-293.
  • Suschenko, D. and Purschke, G. 2009. Ultrastructure of pigmented adult eyes in errant polychaetes (Annelida): implications for annelid evolution. Zoomorphology 128: 75-96.
  • Taylor, J.L. and Gathof, J.M. 1984. Family Questidae Hartman 1966. In: Taxonomic guide to the polychaetes of the northern Gulf of Mexico, vol. 1 (Eds., J.M. Uebelacker and P.G. Johnson). Barry A. Vittor, Mobile, Alabama, pp. 1-85.
  • Timm, T. 1981. Origin and evolution of aquatic Oligochaeta. Eesti NSV Tead Akad. Toim. Biol. 30: 174-181.
  • Timm, T. 1984. Potential age of aquatic Oligochaeta. Hydrobiologia 115: 101-104.
  • Timm, T. 1987. Annelids have emerged from the sea on more than one occasion. In: Aquatic Oligochaeta (Eds., O.L. Kachalova and E.A. Parele). Latvian Acad. Sci., Riga. [in Russian]
  • Treadwell, A.L. 1926. A new polychaetous annelid from Kartabo, British Guyana, genus Namaonereis. Zoologica New York 7:
  • Tzetlin, A. and Purschke, G. 2005. Pharynx and intestine. Hydrobiologia 535: 199-225.
  • Van der Land, J. 1971. Family Aeolosomatidae. In: Aquatic Oligochaeta of the wld (Eds., R.O. Brinkhurst and B.G.M. Jamieson). Oliver & Boyd, Edinburgh, pp. 665-706.
  • Vejdovský, F. 1879. Beiträge zur vergleichenden Morphologie der Anneliden. I. Monographie der Enchytraeiden. 1-61, 14 pls. Friedrich Temspky, Prag.
  • Vejdovský, F. 1880. Vorläufi ger Bericht über die Turbellarien der Brunnen von Prag, nebst Bemerkungen über einige einheimische Arten. Sitzungsber. K. Böhm. Ges. Wiss. 1879: 501-515. [in Russian]
  • Vejdovský, F. 1884. System und Morphologie der Oligochaeten. F. Rivnáe, Prague.
  • Von Bertalanff y, L. 1968. General system theory. Foundations, developments, applications. Braziller, New York.
  • Von Nordheim, H. 1991. Ultrastructure and functional morphology of male genital organs and spermatophore formation in Protodrilus (Polychaeta, Annelida). Zoomorphologie Berlin 111: 81-94.
  • Wesenberg-Lund, E. 1958. Lesser Antillean polychaetes, chiefl y from brackish water, with a survey and bibliography of fresh and brackish water polychaetes. Stud. Faun. Curaçao Oth. Caribb. Isl. 8: 1-41.
  • Westheide, W. 1997a. Th e direction of evolution within the Polychaeta. J. Nat. Hist. 31: 1-15.
  • Westheide, W. 1997b. Th e reading direction of the annelid evolution. Bull. Mar. Sci. 60: 626.
  • Westheide, W. and Bunke, D. 1970. Aeolosoma maritimum nov. sp. Die erste Salzwasserart aus der Familie Aeolosomatidae (Annelida, Oligochaeta). Helgoländer Wiss. Meeres. 21: 134
  • Westheide, W., McHugh, D., Purschke, G. and Rouse, G.W. 1999. Systematization of the Annelida: diff erent approaches. Hydrobiologia 402: 291-307.
  • Westheide, W. and Müller, M.C. 1996. Cinematographic documentation of enchytraeid morphology and reproductive biology. Hydrobiologia 334: 263-267.
  • Wilkens, V. and Purschke, G. 2009. Central nervous system and sense organs, with special reference to photoreceptor-like sensory elements, in Polygordius appendiculatus (Annelida), an interstitial polychaete with uncertain phylogenetic affi nities. Inv. Biol. 128: 46-64.
  • Wilson, G.D.F. and Hessler, R.R. 1987. Speciation in the deep-sea. Ann. Rev. Ecol. Syst. 18: 185-207.
  • Wilson, W.H. 1991. Sexual reproductive modes in polychaetes: classifi cation and diversity. Bull. Mar. Sci. 48: 500-516.
  • Winnepenninckx, B.M.H., Van der Peer, Y. and Backeljau, T. 1998. Metazoan relationships on the basis of 18S rRNA sequences: A few years later. Am. Zool. 38: 888-906.
  • Worsfold, T.M. 2006. British records of the interstitial polychaete Stygocapitella subterranea (Annelida: Parergodrilidae). JMBA2 – Biod. Rec. Assessed Oct. 21, 2008 at: http://www.mba.ac.uk/ jmba/pdf/5429.pdf
  • Zrzavý, J., Riha, P., Piálek, L. and Janouskovec, J. 2009. Phylogeny of Annelida (Lophotrochozoa): total-evidence analysis of morphology and six genes. BMC Evolutionary Biology, 9: 189. Assessed Sep. 30, 2009 at: http: www.biomedcentral.com/1471- 2148/9/189.