Systematization of the Annelida: different approaches

Different approaches and different data sets are used to address questions about the phylogenetic relationships of the polychaete family taxa, the position of the Clitellata, the monophyletic nature of the Polychaeta and the Annelida, as well as the position of the Pogonophora and Echiura. In part, the authors of this contribution come to different conclusions regarding these questions: (1) The analysis of G. W. Rouse minimizes assumptions about any process concerning evolution in annelids and makes a detailed series of homology assessments, based on morphology, across the entire range of taxa concerned. A monophyletic taxon Polychaeta is found but there is no evidence of a sister group for the Clitellata among the polychaetes. The Pogonophora are positioned as members of the Polychaeta. A new classification of the polychaete families is provided. Problems with coding morphology for cladistic analysis are outlined and discussed. (2) D. McHugh uses the sequence of a nuclear gene, elongation factor-1α, for phylogenetic analysis by the parsimony and neighbour joining methods. The main message from phylogenetic analyses of molecular data is that there is no evidence for a monophyletic Polychaeta; instead, placement of the clitellates, pogonophorans and echiurans within the polychaete clade is supported, rendering the Polychaeta paraphyletic. (3) The phylogenetic system of W. Westheide and G. Purschke is based on Hennigian reasoning, and their character weighting is based on diverse, mainly functional considerations. Their annelid tree shows the Clitellata as a highly evolved monophylum and the polychaetes as paraphyletic. Special emphasis is placed on the demonstration that the Clitellata are of terrestrial origin. Advantages and disadvantages of the different approaches and open questions are discussed.

[1]  M. Siddall,et al.  Probabilism and Phylogenetic Inference , 1997, Cladistics : the international journal of the Willi Hennig Society.

[2]  G. Purschke,et al.  Anatomy and ultrastructure of the ventral pharyngeal organs of Saccocirrus(Saccocirridae) and Protodriloides(Protodriloidae fam. n.) with remarks on the phylogenetic relationships within the Protodrilida (Annelida: Polychaeta) , 1988 .

[3]  R. Raff,et al.  Evidence for a clade of nematodes, arthropods and other moulting animals , 1997, Nature.

[4]  W. Westheide,et al.  CLSM analysis of serotonin-immunoreactive neurons in the central nervous system of Nais variabilis, Slavina appendiculata and Stylaria lacustris (Oligochaeta: Naididae) , 1999, Hydrobiologia.

[5]  J. Garey,et al.  The position of the Arthropoda in the phylogenetic system , 1998, Journal of morphology.

[6]  W. Westheide,et al.  The direction of evolution within the Polychaeta , 1997 .

[7]  C. Young,et al.  Embryology of vestimentiferan tube worms from deep-sea methane/sulphide seeps , 1996, Nature.

[8]  P. Fauvel Polychètes sédentaires : addenda aux errantes, archiannélides, myzostomaires , 1927 .

[9]  D. Wake,et al.  Phylogenetic reconstruction. , 1978, Science.

[10]  R. Wachter,et al.  Phylogeny of protostome worms derived from 18S rRNA sequences. , 1995, Molecular biology and evolution.

[11]  K. Fauchald Polychaete Phylogeny: A Problem in Protostome Evolution , 1974 .

[12]  Won Kim,et al.  Phylogenetic relationships of annelids, molluscs, and arthropods evidenced from molecules and morphology , 1996, Journal of Molecular Evolution.

[13]  R. Brinkhurst Evolutionary relationships within the clitellata , 1992 .

[14]  W. Westheide,et al.  Ultrastructure of Chrysopetalid paleal chaetae (Annelida, Polychaeta) , 1992 .

[15]  T. Takagi,et al.  The Deep-Sea Tube Worm Hemoglobin : Subunit Structure and Phylogenetic Relationship with Annelid Hemoglobin : Biochemistry , 1989 .

[16]  S. Kojima Paraphyletic status of Polychaeta suggested by phylogenetic analysis based on the amino acid sequences of elongation factor-1 alpha. , 1998, Molecular phylogenetics and evolution.

[17]  O. Hartman Atlas of the sedentariate polychaetous annelids from California , 1968 .

[18]  Won Kim,et al.  Phylogenetic positions of the aberrant branchiobdellidans and aphanoneurans within the Annelida as derived from 18S ribosomal RNA gene sequences , 1996, Hydrobiologia.

[19]  D. Eernisse,et al.  Annelida and Arthropoda are Not Sister Taxa: A Phylogenetic Analysis of Spiralian Metazoan Morphology , 1992 .

[20]  Junhyong Kim,et al.  Separate Versus Combined Analysis of Phylogenetic Evidence , 1995 .

[21]  A. Whittle,et al.  Fine structure of nuchal organs in some errant polychaetous annelids , 1974, Journal of morphology.

[22]  G. Rouse,et al.  Polychaete systematics: Past and present , 1997 .

[23]  A. H. Clark,et al.  Animal evolution , 1981 .

[24]  Mário C. C. Pinna CONCEPTS AND TESTS OF HOMOLOGY IN THE CLADISTIC PARADIGM , 1991 .

[25]  P. Lupetti,et al.  An ultrastructural investigation of Hrabeiella Pizl & Chalupský, 1984 (Annelida). II. The spermatozoon. , 1997, Tissue & cell.

[26]  Thomas Bartolomaeus,et al.  Structure, function and development of segmental organs in Annelida , 1999, Hydrobiologia.

[27]  G. Purschke Terrestrial polychaetes – models for the evolution of the Clitellata (Annelida)? , 1999, Hydrobiologia.

[28]  O. Giere,et al.  A systematic account of the Questidae (Annelida, Polychaeta), with description of new taxa , 1998 .

[29]  J. Day A monograph on the Polychaeta of southern Africa. Part 2. Sedentaria , 1967 .

[30]  W. Westheide Progenesis as a principle in meiofauna evolution , 1987 .

[31]  T. Bartolomaeus CHAETOGENESIS IN POLYCHAETOUS ANNELIDA : SIGNIFICANCE FOR ANNELID SYSTEMATICS AND THE POSITION OF THE POGONOPHORA , 1998 .

[32]  A. Kluge,et al.  Testability and the Refutation and Corroboration of Cladistic Hypotheses , 1997, Cladistics : the international journal of the Willi Hennig Society.

[33]  W. F. Gutmann Relationships Between Invertebrate Phyla Based on Functional-Mechanical Analysis of the Hydrostatic Skeleton , 1981 .

[34]  G. Rouse,et al.  Cladistics and polychaetes , 1997 .

[35]  J. Wägele First principles of phylogenetic systematics, a basis for numerical methods used for morphological and molecular characters , 1996 .

[36]  C. Erséus Phylogenetic analysis of the aquatic Oligochaeta under the principle of parsimony , 1987, Hydrobiologia.

[37]  S. Gardiner,et al.  On the Significance of Larval and Juvenile Morphology for Suggesting Phylogenetic Relationships of the Vestimentifera , 1994 .

[38]  The ancestral cleavage pattern of the clitellates and its phylogenetic deviations , 1999 .

[39]  G. Rouse Rearticulating with extra assumptions: a response to Eibye‐Jacobsen and Nielsen , 1997 .

[40]  A. Minelli,et al.  Myriapod metamerism and arthropod segmentation , 1988 .

[41]  C. O. Hermans The Systematic Position of the Archiannelida , 1969 .

[42]  J. Torrey THE EARLY EMBRYOLOGY OF THALASSEMA MELLITA (CONN) , 1901 .

[43]  G. Purschke,et al.  Morphological reinvestigation and phylogenetic relationship of Acanthobdella peledina (Annelida, Clitellata) , 1993, Zoomorphology.

[44]  G. Purschke Ultrastructure of Nuchal Organs in Polychaetes (Annelida) — New Results and Review , 1997 .

[45]  R. B. Clark Dynamics in Metazoan Evolution: The Origin of the Coelom and Segments. , 1964 .

[46]  T. Bartolomaeus Structure and formation of the uncini in Pectinaria koreni, Pectinaria auricoma (Terebellida) and Spirorbis spirorbis (Sabellida): inplications for annelid phylogeny and the position of the Pogonophora , 1995, Zoomorphology.

[47]  A. Giangrande,et al.  METAMERISM AND LIFE-STYLE WITHIN POLYCHAETES : MORPHO-FUNCTIONAL ASPECTS AND EVOLUTIONARY IMPLICATIONS , 1998 .

[48]  P. Omodeo History of Clitellata , 1998 .

[49]  M. L. Jones,et al.  On the Vestimentifera, new phylum: six new species, and other taxa, from hydrothermal vents and elsewhere , 1985 .

[50]  D. McHugh,et al.  Molecular evidence that echiurans and pogonophorans are derived annelids. , 1997, Proceedings of the National Academy of Sciences of the United States of America.

[51]  F. Pleijel ON CHARACTER CODING FOR PHYLOGENY RECONSTRUCTION , 1995 .

[52]  G. Purschke,et al.  Dorsolateral Ciliary Folds in the Polychaete Foregut: Structure, Prevalence and Phylogenetic Significance , 1996 .

[53]  A. R. Templeton,et al.  PHYLOGENETIC INFERENCE FROM RESTRICTION ENDONUCLEASE CLEAVAGE SITE MAPS WITH PARTICULAR REFERENCE TO THE EVOLUTION OF HUMANS AND THE APES , 1983, Evolution; international journal of organic evolution.

[54]  Greg W. Rouse,et al.  The articulation of annelids , 1995 .

[55]  W. W. Newby embryology of the echiuroid worm Urechis Caupo , 1940 .

[56]  Å. Franzén,et al.  Ultrastructure of Spermatozoa and Spermatids in Bonellia viridis and Hamingia arctica (Echiura) With Some Phylogenetic Considerations , 1992 .

[57]  G. Purschke Structure of the prostomial appendages and the central nervous system in the Protodrilida (Polychaeta) , 1993, Zoomorphology.

[58]  E. Rota Morphology and adaptations of Parergodrilus Reisinger and Hrabeiella Pizl & Chalupský, two enigmatic soil‐dwelling annelids , 1998 .

[59]  E. Southward Development of the gut and segmentation of newly settled stages of Ridgeia (Vestimentifera): implications for relationship between Vestimentifera and Pogonophora , 1988, Journal of the Marine Biological Association of the United Kingdom.

[60]  R. Raff,et al.  Molecular phylogeny of the animal kingdom. , 1988, Science.

[61]  Kristian Fauchald,et al.  The Polychaete Worms: Definitions and Keys to the Orders, Families and Genera , 1977 .

[62]  J. George,et al.  Polychaetes: British Amphinomida, Spintherida and Eunicida , 1985 .

[63]  S. Morris,et al.  Articulated Halkieriids from the Lower Cambrian of North Greenland and their Role in Early Protostome Evolution , 1995 .

[64]  D. Faith Cladistic permutation tests for monophyly and nonmonophyly , 1991 .

[65]  W. Westheide,et al.  CLSM analysis of development and structure of the central nervous system of Enchytraeus crypticus (”Oligochaeta”, Enchytraeidae) , 1999, Zoomorphology.

[66]  O. Giere,et al.  Questidae—Polychaetes with Oligochaetoid Morphology and Development 1 , 1981 .

[67]  R. B. Clark CHAPTER 1 – Systematics and Phylogeny: Annelida, Echiura, Sipuncula , 1969 .

[68]  K. Nixon,et al.  ON OUTGROUPS , 1993, Cladistics : the international journal of the Willi Hennig Society.