Vol. 7, No. 16, p. 119-147 - Aug. 31, 2020
Character investigation and homology, with a brief discussion on the phylogenetic relationships of Annelida and Polychaeta within Metazoa
José Eriberto De Assis and Martin Lindsey Christoffersen
Abstract
The intense production of information demands coherent treatment of available bodies of theoretical knowledge. A critical revision of methods in use within research programs is essential to maintain an adequate ontology and to guarantee the particular epistemological position required by each research area. The theory of Phylogenetic Systematics was developed in multiple forms, resulting in hypotheses that may be widely incongruent. This reflects the crisis of present paradigm, and illustrates failures in present phylogenetic thinking. The aim of the study is to discuss characters and homology hypotheses based on Hennigian principles. We present samples from the most unresolved groups within the evolutionary history of the Metazoa: Annelida and Polychaeta. The main phylogenetic proposals appearing in the history of the Annelida, from morphological (living and fossilized organisms) to molecular approaches are discussed. We do not consider annelids and polychaetes to be monophyletic entities. Both taxa need more detailed comparisons with others groups, such as deuterostomes. According to our published opinions, Annelida and Polychaeta are placed at the base of the deuterostomes, and these worm-like organisms are ultimately the ancestors of the Enterocoela. Thus, in order to systematize the Metameria, we have regarded Annelida, Pleistoannelida, Lophotrochozoa and Protostomia as being paraphyletic taxa.
Keywords
Phylogenetic principles; Annelids; Synapomorphies; Morphology; Molecular Systematics.
DOI
10.21472/bjbs(2020)071601
Full text
PDF
References
Adrianov, A. V.; Malakhov, V. V.; Maiorova, A. S. Development of the tentacular apparatus in sipunculans
(Sipuncula): I. Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920).
Journal of Morphology, v. 267, p. 569-583, 2006. https://doi.org/10.1002/jmor.10423
Almeida, W. O.; Christoffersen, M. L.; Amorim, D. S.; Elloy, E. C. C. Morphological support for the position
of Pentastomida and related fossils. Biotemas, v. 21, p. 81-80, 2008. https://doi.org/10.5007/2175-7925.2008v21n3p81
Almeida, W. O.; Christoffersen, M. L.; Amorim, D. S.; Garraffoni, A. R. S; Sene, S. G. Polychaeta, Annelida,
and Articulata are not monophyletic: Articulating the Metameria (Metazoa, Coelomata). Revista Brasileira
de Zoologia, v. 20, p. 23-57, 2003. https://doi.org/10.1590/S0101-81752003000100006
Almeida, W. O; Christoffersen, M. L. Análise cladística dos grupos basais de Metameria: uma
nova proposta para o posionamento dos Arthropoda e grupos afins entre os poliquetas errantes. Ribeirão
Preto: Holos Editora, 2001.
Álvarez-Campos, P.; Sergio, T.; San Martín, G.; Leiva, C.; Riesgo, A. Phylogenetic relationships
and evolution of reproductive modes within flattened syllids (Annelida: Syllidae) with the description of a
new genus and six new species. Invertebrate Systematics, v. 32, p. 224-251, 2018. https://doi.org/10.1071/IS17011
Amorim, D. S. Elementos da Sistemática Filogenética. Ribeirão Preto: Holos Editora,
2002.
Arendt, D.; Nübler-Jung, K. Common ground plans in early brain development in mice and flies. BioEssays,
v. 18, p. 255-259, 1996. https://doi.org/10.1002/bies.950180314
Arendt, D.; Nübler-Jung, K. Comparison of early nerve cord development in insects and vertebrates. Development,
v. 126, p. 2309-2325, 1999.
Arendt, D.; Nübler-Jung, K. Dorsal or ventral: Similarities in fate maps and gastrulation patterns in annelids,
arthropods and chordates. Mechanisms of Development, v. 61, p. 7-21, 1997. https://doi.org/10.1016/S0925-4773(96)00620-X
Arendt, D.; Nübler-Jung, K. Inversion of dorsoventral axis? Nature, v. 371, p. 26, 1994. https://doi.org/10.1038/371026a0
Arendt, D.; Tessmar-Raible, K.; Snyman, H.; Dorresteijn, A. W.; Wittbrodt, J. Ciliary photoreceptors with a vertebrate-type
opsin in an invertebrate brain. Science, v. 306, p. 869-871, 2004. https://doi.org/10.1126/science.1099955
Assis, L. C. S. Are homology and synapomorphy the same or different? Cladistics, v. 29, p. 7-9, 2012.
https://doi.org/10.1111/j.1096-0031.2012.00431.x
Assis, L. C. S.; Santos, L. M. Phylogenetics is not phylogenomics. Cladistics, v. 30, p. 8-9, 2014.
https://doi.org/10.1111/cla.12028
Bartolomaeus, T. Structure and formation of the uncini in Pectinaria koreni, Pectinaria auricoma
(Terebellida) and Spirorbis spirorbis (Sabellida): Implications for annelid phylogeny and the position
of the Pogonophora. Zoomorphology, v. 115, p. 161-177, 1995. https://doi.org/10.1007/BF00403171
Bartolomaeus, T. Ultrastructure and formation of hooked setae in Owenia fusiformis Delle Chiaje, 1842:
Implications for annelid phylogeny. Canadian Journal of Zoology, v. 74, p. 2143-2153, 1996.
https://doi.org/10.1139/z96-243
Bartolomaeus, T.; Purschke, G.; Hausen, H. Polychaete phylogeny based on morphological date: A comparison of
current attempts. Hydrobiologia, v. 535/536, p. 341-356, 2005. https://doi.org/10.1007/s10750-004-1847-5
Beaty, J. The evolutionary contigency thesis. Proceedings of the II Pittsburg-Konstanz Colloquium in the
Philosophy of Science, 1993, Pittsbugh, University of Pittsburghy Press, 1995.
Blake, A. J. The Annelida. Part 1. Introduction to Polychaeta. In: Black, A. J.; Hilbig, B.; Scott, P. H.
(Eds.). Taxonomic atlas of the benthic fauna of the Santa Barbara Channel. Santa Barbara, California,
1997. p. 39-133.
Bleidorn, C.; Podsiadlowski, L.; Bartolomaeus, T. The complete mitochondrial genome of the orbiniid polychaete
Orbinia latreillii (Annelida, Orbiniidae): A novel gene order for Annelida and implications for annelid
phylogeny. Gene, v. 370, p. 96-103, 2006. https://doi.org/10.1016/j.gene.2005.11.018
Bleidorn, C.; Vogt, L.; Bartolomaeus, T. New insights into polychaete phylogeny (Annelida) inferred from 18S
rDNA sequences. Molecular Phylogenetics and Evolution, v. 29, p. 279-288, 2003. https://doi.org/10.1016/S1055-7903(03)00107-6
Borkent, A. The state of phylogenetic analysis: Narrow visions and simple answers: Examples from the Diptera
(flies). Zootaxa, v. 4374, p. 107-143, 2018. https://doi.org/10.11646/zootaxa.4374.1.7
Bright, M.; Eichinger, I.; von Salvini Plawen, L. The metamorphose of a deep-sea hydrothermal vent vestimentiferan
(Polychaeta: Sibodlinidae). Organisms, Diversity & Evolution, v. 13, p. 163-188, 2013. https://doi.org/10.1007/s13127-012-0117-z
Brooks, D. R.; McLennan, D. A. The nature of diversity: An evolutionary voyage of discovery. Chicago:
The University of Chicago Press, 2002.
Brooks, D. R.; Wiley, E. O. Theories and methods in different approaches to phylogenetic systematics. Cladistics,
v. 1, p. 1-11, 1985. https://doi.org/10.1111/j.1096-0031.1985.tb00407.x
Brower, A. V. Z.; de Pinna, M. C. C. Homology and errors. Cladistics, v. 1, p. 1-10, 2012. https://doi.org/10.1111/j.1096-0031.2012.00398.x
Brower, A. V. Z. The meaning of "phonetic". Cladistics, v. 28, p. 113-114, 2012. https://doi.org/10.1111/j.1096-0031.2011.00374.x
Brown, F. D.; Prendergast, A.; Swalla, B.J. Man is but a Worm: Chordate origins. Genesis, v. 46, p. 605-613, 2008.
https://doi.org/10.1002/dvg.20471
Bubko, O. V.; Minichev, Y. S. The nevous system in Oweniidae. Zoologichesckii Zhurnal, v. 51, p. 1288-1299, 1972.
Budaeva, N.; Schepetov, C.; Zanol, J.; Neretina, T.; Willassen, E. When molecules support morphology: Phylogenetic
reconstruction of the Family Onuphidae (Eunicida, Annelida) based on 16S rDNA and 18S rDNA. Molecular Phylogenetics
and Evolution, v. 94, p. 791-801, 2015. https://doi.org/10.1016/j.ympev.2015.10.011
Carnap, R. The methodological character of theoretical concepts. In: Feigl, H.; Scriven, M. (Eds.). The foundations
of Science and the concepts of Psychology and Psychoanalysis. Minneapolis: University of Minnesota Press, 1956.
Carpenter, J. M. Choosing among multiple equality parsimonious cladograms. Cladistics, v. 2, p. 291-296, 1998.
https://doi.org/10.1111/j.1096-0031.1988.tb00476.x
Chen, H.; Parry, L. A.; Vinther, J; Zhai, D.; Hou, X.; Ma, X. A Cambrian crown annelid reconciles phylogenomics and
the fossil record. Nature, v. 583, p. 249-252, 2020. https://doi.org/10.1038/s41586-020-2384-8
Christoffersen, M. L. Clitellate evolution and leech diversity: Glossifoniidae excl. Helobdella (Annelida: Hirudinea:
Hhynchobdellida) from South America. Gaia Scientia, v. 1, p. 131-140, 2007.
Christoffersen, M. L.; Ara&uacte;jo, M. E.; Moreira, M. A. M. A molecular method for a qualitative analysis of potentially
coding sequences of DNA. Brazilian Journal of Biology, v. 64, p. 383-398, 2004. https://doi.org/10.1590/S1519-69842004000300003
Christoffersen. M. L.; Araújo-de-Almeida. E. A phylogenetic framework of the Enterocoela (Metameria: Coelomata).
Revista Nordestina de Biologia, v. 9, p. 173-208, 1994.
Clark, C.; Curran, D. J. Outgroup analysis, homoplasy, and global parsimony: A response to Maddison, Donoghue, and Maddison.
Systematic Zoology, v. 35, p. 422-426, 1986. https://doi.org/10.1093/sysbio/35.3.422
Clark, R. B. Dynamics in metazoan evolution. The origin of the coelom and segments. Oxford: Clarendon Press; 1964.
Clark, R. B. Systematics and Phylogeny: Annelida, Echiura, Sipuncula. Chemical Zoology 4. New York: Academic Press,
1969.
Colgan, D. J.; Hutchings, P. A.; Braune, M. A multigene framework for polychaete phylogenetic studies. Organisms,
Diversity & Evolution, v. 6, p. 220-235, 2006. https://doi.org/10.1016/j.ode.2005.11.002
Conway-Morris, S. Once we were worms. New Scientist, v. 179, p. 34-37, 2003.
De Assis, J. E.; Christoffersen, M. L. Pogonophora (Siboglinidae) are not polychaetes: A systemic view of metamerian
animals. Revista Nordestina de Biologia, v. 19, p. 55-76, 2010.
De Carvalho, M. R.; Ebach, M. C. Death of the specialist, rise of the machinist. History and Philosophy of the
Life Sciences, v. 31, p. 467-470, 2009.
De Pinna, M. C. C. Concepts and test of homology in the cladistic paradigm. Cladistics, v. 7, p. 367-394, 1991.
https://doi.org/10.1111/j.1096-0031.1991.tb00045.x
De Queiroz, K. The ontogenetic method determining character polarity and its relevance to phylogenetic systematics.
Systematics Zoology, v. 34, p. 280-299, 1985. https://doi.org/ 10.1093/sysbio/34.3.280
Donoghue, M. J.; Cantino, P. D. The logic and limitations of the outgroup substitution approach to cladistic analysis.
Systematics Botany, v. 9, p. 19-202, 1984.
Dretske, F. I. Laws of nature. Philosophy of Science, v. 44, p. 248-268, 1977. https://doi.org/10.1086/288741
Dzik, J.; Krumbiegel, G. The oldest 'onychophoran' Xenusium: A link connecting phyla? Lethaia, v. 22,
p. 169-181-1989. https://doi.org/10.1111/j.1502-3931.1989.tb01679.x
Eibye-Jacobsen, D.; Nielsen, C. Point of view: the rearticulation of annelids. Zoologica Scripta, v. 25, p. 275-282,
1996. https://doi.org/10.1111/j.1463-6409.1996.tb00166.x
Fang, F. C.; Casadevall, A. Reductionistic and holistic science. Infection and Immunity, v. 79, p. 1401-1404, 2011.
https://doi.org/10.1128/IAI.01343-10
Farris, J. A successive approximations approach to character weighting. Systematics Zoology, v. 18, p. 374-385, 1969.
https://doi.org/10.2307/2412182
Farris, J. Methods for computing Wagner trees. Systematics Zoology, v. 19, p. 83-92, 1970. https://doi.org/10.2307/2412028
Fauchald, K. Polychaeta phylogeny: A problem in protostome evolution. Systematics Zoology, v. 23, p. 493-506, 1974.
https://doi.org/10.1093/sysbio/23.4.493
Fauchald, K. The polychaete worms: Definitions and keys to the orders, families and genera. Natural History Museum of
Los Angeles County, Science Series, v. 28, p. 1-190, 1977.
Fitzhugh, K. Species as explanatory hypotheses: Refinements and implications. Acta Biotheoretica, v. 57, p. 201-248,
2009. https://doi.org/10.1007/s10441-009-9071-3
Fitzhugh, K. The philosophical basis of character coding for the inference of phylogenetic hypotheses. Zoologica Scripta,
v. 35, p. 261-286, 2006. https://doi.org/10.1111/j.1463-6409.2006.00229.x
François, V.; Bier, E. Xenopus chordin and Drosophila short gastrulation genes encode homologous
proteins functioning in dorsal-ventral axis formation. Cell, v. 80, p. 19-20, 1995. https://doi.org/10.1016/0092-8674(95)90446-8
Gardiner, S. L. Fine structure of Owenia fusiformis (Polychaeta: Oweniidae) and its significance for annelid
phylogeny. Chapel Hill: University of North Carolina, 1979. (Ph.D. Thesis).
Gardiner, S. L. Fine structure of the ciliate epidermis on the tentacles of Owenia fusiformis (Polychaeta, Oweniidae).
Zoomorphologie, v. 91, p. 37-48, 1978. https://doi.org/10.1007/BF00994152
Garraffoni, A. R. S.; Amorim, D. S. Análise filogenética de Questidae e Clitellata: o problema da parafilia
de "Polychaeta". Iheringia, Série Zoologia, v. 93, p. 97-109, 2003. https://doi.org/10.1590/S0073-47212003000100011
Glasby, C. J.; Hutchings, P. A.; Fauchald, F.; Paxton, H.; Rouse, G. W.; Watson, R. C.; Wilson, R. S. Class Polychaeta.
In: Beesley, P. L.; Ross, G. J. B.; Glasby, C. J. (Eds.). Polychaetes & allies: The Southern Synthesis.
Fauna of Australia. Vol. 4A. Polychaeta, Myzostomida, Pogonophora, Echiura, Sipuncula. Melbourne: CSIRO Publishing,
2000, p. 1-296.
Goloboff, P. A. Methods for faster parsimony analysis. Cladistics, v. 12, p. 199-220, 1996.
https://doi.org/10.1111/j.1096-0031.1996.tb00009.x
Goloboff, P. A.; Farris, J. S.; Nixon, K. C. TNT, a free program for phylogenetics analysis. Cladistics, v. 24,
p. 774-786, 2008. https://doi.org/10.1111/j.1096-0031.2008.00217.x
Grandcolas, P.; Guilbert, E.; Robillard, T.; D'Haese, A.; Murienne, J; Legendre, F. Mapping characters on tree
with or without the outgroups. Cladistics, v. 20, p. 579–582, 2005. https://doi.org/10.1111/j.1096-0031.2004.00037.x
Gustus, R. M.; Cloney, R. A. Ultrastructural similarities between chaetae of brachiopods and polychaetes. Acta
Zoologica, v. 53, p. 229-233, 1972. https://doi.org/10.1111/j.1463-6395.1972.tb00590.x
Halanych, K. M. The new view of animal phylogeny. Annual Review of Ecology, Evolution, and Systematics, v. 35,
p. 229-256, 2004. https://doi.org/10.1146/annurev.ecolsys.35.112202.130124
Hall, B. K. Homology: The hierarchical basis of comparative Biology. New York: Academic Press, 1994.
Haszprunar, G. The types of homology and their significance for evolutionary biology and phylogenetics. Journal
of Evolutionary Biology, v. 5, no. 1, p. 13-24, 1992. https://doi.org/10.1046/j.1420-9101.1992.5010013.x
Hempel, C. G. Aspects of scientific explanation, and other essays in the philosophy of science. New York:
Free Press, 1965.
Hempel, C. G. Oppenheim, P. Studies in the logic of explanation. Philosophy of Science, v. 15, p. 135-175,
1948.
Hennig, W. Phylogenetic Systematics. Urbana: University of Illinois Press, 1966.
Holland, L. Z.; Kene, M.; Williams, N. A.; Holland, N. Sequence and embryonic expression of the amphioxus engrailed
gene (AmphiEn): The metameric pattern of transcription resembles that of its segment-polarity homolog in Drosophila.
Development, v. 124, p. 1723-1732, 1997.
Holley, S. A.; Jackson, P. D.; Sasai, Y.; Lu, B.; Robertis, E. M. Hoffmann, F. M.; Ferguson, E. L. Conserved system
for dorsal–ventral patterning in insects and vertebrates involving sog and chordin. Nature, v. 376, p. 249-253,
1995. https://doi.org/10.1038/376249a0
Hyman, L. H. The invertebrates: Smaller coelomate groups. New York: McGraw-Hill, 1959.
Ivanov, A. V. Analysis of the embryonic developmental of Pogonophora in connection with the problems
of phylogenetics. Journal of Zoological Systematics and Evolutionary Research, v. 26,
p. 161-185, 1998. https://doi.org/10.1111/j.1439-0469.1988.tb00308.x
Ivanov, A. V. The main features of the organization of Pogonophora. Systematics Zoology, v. 4,
p. 171-174, 1955a. https://doi.org/10.2307/2411669
Ivanov, A. V. On external digestion in Pogonophora. Systematics Zoology, v. 4, p. 174-176,
1955b. https://doi.org/10.2307/2411670
Ivanov, A. V. On the assignment of class Pogonophora to a separate phylum of Deuterostomia: Brachiata.
A. Ivanov, Phyl. Nov. Systematics Zoology, v. 4, p. 177-178, 1955c. https://doi.org/10.2307/2411671
Ivanov, A. V. Embranchement des Pogonophores. In: Grassé, P. P. (Ed.). Traité de Zoologie.
Paris: Masson, 1960. p. 1521-1622. v. 5, 2.
Ivanova-Kazas, O. M. Molecules, morphology, and phylogeny. Paleontology Journal, v. 50, p. 1474-1476,
2016.
Ivanova-Kazas, O. M. On the problem of the origin of Pogonophora. Russian Journal of Marine Biology,
v. 33, p. 338-342, 2007. https://doi.org/10.1134/S1063074007050124
Jennings, R. M.; Halanych, K. M. Mitocondrial genomes of Clymenella torquata (Maldanidae) and
Riftia pachyptila (Siboglinidae): Evidence for conserved gene order in Annelida. Molecular
Phylogenetics and Evolution, v. 22, p. 210-222, 2005. https://doi.org/10.1093/molbev/msi008
Johansson, K. E. Über Lamellisabella zachsi und ihre systematische Stellung. Zoologischer Anzeiger,
v. 117, p. 23-26, 1937.
Jones, C. M.; Smith, J. C. Revolving vertebrates. Current Biology, v. 5, p. 474-476, 1995.
Jones, M. L. On the Vestimentifera, new phylum: Six new species, and other taxa, from hydrothermal
vents and elsewhere. Bulletin of the Biological Society of Washington, v. 6, p. 117-158, 1985.
Jones, M. L. Vestimentiferan pogonophores: Their biology and affinities. In: Morris, S. C.; George, J. D. R.;
Platt, H. M. G. (Eds.). The origin and relationships of lower invertebrates. Cambridge: Oxford
University Press, 1985. (The Systematic Association, Special Vol. 28). p. 327-342.
Källersjö, M.; Albert, V. A.; Farris, J. S. Homoplasy increases phylogenetic structure. Cladistics,
v. 15, p. 91-93, 1999. https://doi.org/10.1111/j.10960031.1999.tb00400.x
Kara, J.; Santos, C. S. G.; Macdonald, A. H. H.; Simon, C. A. Resolving the taxonomic identities and genetic
structure of two cryptic Platynereis Kinberg species from South Africa. Invertebrate Systematics,
v. 34, p. 618-636, 2020. https://doi.org/10.1071/IS19072
Kluge, A. G.; Farris, J. S. Quantitative phyletics and the evolution of anurans. Systematic Zoology, v. 18,
p. 1-32, 1969. https://doi.org/10.2307/2412407
Kluge, A. G.; Strauss, R. E. Ontogeny and systematic. Annual Review of Ecology, Evolution, and Systematics,
v. 16, p. 247-268, 1985. https://doi.org/10.1146/annurev.es.16.110185.001335
Kojima, S. Paraphyletic status of Polychaeta suggested by phylogenetic analysis based on the amino acid sequences
of elongation factor X. Molecular Phylogenetics and Evolution, v. 9, p. 255-261, 1998. https://doi.org/10.1006/mpev.1997.0472
Kück, P.; Wägele, W. Plesiomorphic character states cause systematic errors in molecular phylogenetic
analyses: A simulation study. Cladistics, v. 32, p. 461-478, 2016. ttps://doi.org/10.1111/cla.12132
Kuhn, T. S. The structure of scientific revolutions. 2. ed. Chicago: University Press, 1970.
Kvist, S.; Siddall, M. E. Phylogenomics of Annelida revisited: A cladistic approach using genome-wide expressed
sequence tag data mining and examining the effects of missing data. Cladistics, v. 29, p. 435-448, 2013.
https://doi.org/10.1111/cla.12015
Laubichler, M. D. Homology in development and the development of the homology concept. American Zoologist,
v. 40, p. 777-788, 2000. https://doi.org/10.1093/icb/40.5.777
Lawrence, P. A. Compartments in vertebrates? Nature, v. 344, p. 382-383, 1990. https://doi.org/10.1038/344382a0
Lester, S. M. Cephalodiscus sp. (Hemichordata: Pterobranchia): Observations of functional morphology, behaviour
and occurence in shallow water around Bermuda. Marine Biology, v. 85, p. 263-268, 1985. https://doi.org/10.1007/BF00393246
Lüter, C.; Bartolomaeus, T. The phylogenetic position of Brachiopoda: A comparison of morphological and molecular
data. Zoologica Scripta, v. 26, p. 245-253, 1997. https://doi.org/10.1111/j.1463-6409.1997.tb00414.x
Malakhov, V. V.; Popelyaev, I. S.; Galkin, S. A. On the position of Vestimentifera and Pogonophora in the system of
the Animal Kingdom. Zoologichesckii Zhurnal, v. 76, p. 1336-1347, 1997.
Manton, S. M. The Arthropods: Habits, functional morphology and evolution. Oxford: Clarendon Press, 1977.
Martínez, S. F. Reducionismo em Biologia: uma tomografia da relação Biologia-sociedade. In: Abrantes,
P. C. (Ed.). Filosofia da Biologia. Porto Alegre: Artmed, 2011. p. 37-59.
McGinnis, W.; Hart, C. P.; Gehring, W. J.; Ruddle, F. H. Molecular cloning and chromosome mapping of a mouse DNA sequence
homologous to homeotic genes of Drosophila. Cell, v. 38, p. 675-680, 1984. https://doi.org/10.1016/0092-8674(84)90262-9
McHugh, D. Molecular evidence that echiurans and pogonophorans are derived annelids. PNAS, v. 94, p. 8006-8009, 1997.
https://doi.org/10.1073/pnas.94.15.8006
McHugh, D. Molecular phylogeny of the Annelida. Canadian Journal of Zoology, v. 78, p. 1873-1884, 2000.
Mickevich, M. F. Transformation series analysis. Systematic Zoology, v. 31, p. 461-478, 1982.
Mickevich, M. F.; Lipscomb, D. L. Parsimony and the choice between alternative transformations for the same data set.
Cladistics, v. 7, p. 111-139, 1991. https://doi.org/10.1111/j.1096-0031.1991.tb00028.x
Mooi, R. D.; Gill, A. C. Phylogenies without synapomorphies: A crisis in fish systematics. Zootaxa, v. 2450,
p. 26-40, 2010.
Moore, J. M.; Nishi, E.; Rouse, G. W. Phylogenetic analyses of Chaetopteridae (Annelida). Zoologica Scripta,
v. 46, p. 596-610, 2017. https://doi.org/10.1111/zsc.12238
Nelson, G.; Platnick, N. Systematics and Biogeography: Cladistics and vicariance. New York: Columbia University
Press, 1981.
Nelson, G. Outgroup and ontogeny. Cladistics, v. 1, p. 29-45, 1985. https://doi.org/10.1111/j.1096-0031.1985.tb00409.x
Nielsen, C. Animal evolution: Interrelationships of the living phyla. 3. ed. New York: Oxford University, 2012.
Nixon, K. C.; Carpenter, J. M. On outgroup. Cladistics, v. 9, p. 413-462, 1993.
https://doi.org/10.1111/j.1096-0031.1993.tb00234.x
Nixon, K. C; Carpenter, J. M. On homology. Cladistics, v. 28, p. 160-169, 2011.
https://doi.org/10.1111/j.1096-0031.2011.00371.x
Nixon, K. C.; Carpenter, J. M. More on homology. Cladistics, v. 28, p. 225-226, 2012.
https://doi.org/10.1111/j.1096-0031.2011.00388.x
Nübler-Jung, K. N.; Arendt, D. Dorsoventral axis inversion: Enteropneust anatomy links invertebrates to chordates
turned upside down. Journal of Zoological Systematics and Evolutionary Research, v. 37, p. 93-100, 1999.
https://doi.org/10.1046/j.1439-0469.1999.372106.x
O'Grady, R. T.; Deets, G. B. Coding multistate characters, with special reference to the use of parasites as
characters of their hosts. Systematic Zoology, v. 36, p. 268-279, 1987. https://doi.org/10.2307/2413066
Orrhage, L. Light and electron microscope studies of some brachiopod and pogonophoran chaetae. With a discussion of the
'annelid chaetae' as a phylogenetic-systematic character. Zeitschrift für Morphologie der Tiere,
v. 74, p. 253-270, 1973. https://doi.org/10.1007/BF00636879
Osorio, D. Evidence that a larval brachiopod has ciliary photoreceptors that are directionally selective, and therefore
may function as eyes, bears on an enduring puzzle about photoreceptor evolution in animals. Nature, v. 472,
p. 300-301, 2011. https://doi.org/10.1038/472300a
Parry, L. A.; Edgecombe, G. D.; Eibye-Jacobsen, D.; Vinther, J. The impact of fossil date on annelid phylogeny inferred
from discret morphological characters. Proceedings of the Royal Society B, Biological Sciences, v. 283, p. 1-9,
2016. https://doi.org/10.1098/rspb.2016.1378
Pavlinov, I. Ya. The contemporary concepts of homology in Biology: A theoretical review. Biology Bulletin Reviews,
v. 2, no. 1, p. 36-54, 2012. https://doi.org/10.1134/S2079086412010057
Pigliucci, M. On the different ways of 'doing theory' in Biology. Biological Theory, v. 7, p. 287-297,
2012. https://doi.org/10.1007/s13752-012-0047-1
Pimentel, R. A.; Riggins, R. The nature of cladistics data. Cladistics, v. 3, no. 3, p. 201-209, 1987.
https://doi.org/10.1111/j.1096-0031.1987.tb00508.x
Pleijel, F. On character coding for phylogenetic reconstruction. Cladistics, v. 11, p. 309-315, 1995.
https://doi.org/10.1111/j.1096-0031.1995.tb00092.x
Pleijel, F.; Dahlgren, T; Rouse, G. W. Progress in systematic: From Siboglinidae to Pogonophora and Vestimentifera
and back to Siboglinidae. Comptes Rendus Biologies, v. 332, p. 140-148, 2009. https://doi.org/10.1016/j.crvi.2008.10.007
Purschke, G. On the ground pattern of Annelida. Organisms Diversity & Evolution, v. 2, p. 181-196, 2002.
https://doi.org/10.1078/1439-6092-00042
Purschke, G.; Bleidorn, C.; Struck, T. Systematics, evolution and phylogeny of Annelida: A morphological perspective.
Memoirs of Museum Victoria, v. 71, p. 247-269, 2014.
Rouse, G. W. A cladistic analysis of Siboglinidae Caullery, 1914 (Polychaeta, Annelida): Formerly the Phyla Pogonophora
and Vestimentifera. Zoological Journal of the Linnean Society, v. 132, p. 55-80, 2001.
https://doi.org/10.1006/zjls.2000.0263
Rouse, G. W.; Fauchald, K. The articulation of annelids. Zoologica Scripta, v. 24, p. 269-301, 1995.
https://doi.org/10.1111/j.1463-6409.1995.tb00476.x
Rouse, G. W.; Pleijel, F. Annelida. Zootaxa, v. 1668, p. 245-264, 2007.
Rouse, G. W.; Pleijel, F. Polychaetes. Oxford: Oxford University Press, 2001.
Rouse, G. W.; Fauchald, K. Cladistics and Polychaetes. Zoologica Scripta, v. 26, p. 139-204, 1997.
https://doi.org/10.1111/j.1463-6409.1997.tb00412.x
Rousset, V.; Pleijel, F.; Rouse, G. W.; Erséus, C.; Siddall, M. E. A molecular phylogeny of annelids.
Cladistics, v. 23, p. 41-63, 2007. https://doi.org/10.1111/j.1096-0031.2006.00128.x
Saglam, S.; Saunders, R.; Lang, S. A.; Shain, D. H. A new species of Hirudo (Annelida: Hirudinidae):
Historical biogeography of Eurasian medicinal leeches. BMC Zoology, v. 1, p. 1-12, 2016.
https://doi.org/10.1186/s40850-016-0002-x
Salvini-Plawen, L. V. What is convergent/homoplastic in Pogonophora? Journal of Zoological Systematics and
Evolutionary Research, v. 38, p. 133-147, 2000. https://doi.org/10.1046/j.1439-0469.2000.383143.x
Schulze, A. Phylogeny of Vestimentifera (Siboglinidae, Annelida) inferred from morphology. Zoologica Scripta,
v. 32, p. 321-342, 2003. https://doi.org/10.1046/j.1463-6409.2003.00119.x
Scotlan, R. W.; Olmstead, R. G.; Bennett, J. R. Phylogeny reconstruction: The role of morphology. Systematic
Biology, v. 54, p. 539-548, 2003.
Siewing, R. Thoughts about the phylogenetic: Systematic position of Pogonophora. Zeitschrit für Zoologische
Systematik und Evolutionsforchung, v. 12, p. 127-138, 1975.
Smart, T. I.; von Dassow, G. Unusual development of the mitraria larva in the Polychaete Owenia collaris.
The Biological Bulletin, v. 217, p. 253-268, 2009. https://doi.org/10.1086/bblv217n3p253
Smith, P. R.; Ruppert, E. E.; Gardiner, S. L. A deuterostome-like nephridium in the mitraria larva of Owenia
fusiformis (Polychaeta, Annelida). The Biological Bulletin, v. 172, p. 315-323, 1987. https://doi.org/10.2307/1541711
Southward, E. C. Fine structure and phylogeny of the Pogonophora. Symposia of the Zoological Society of London,
v. 36, p. 235-251, 1975.
Southward, E. C.; Andersen, A. C.; Hourdez, S. Lamellibrachia anaximandri n. sp., a new vestimentiferan tubeworm
(Annelida) from the Mediterranean, with notes on frenulate tubeworms from the same habitat. Zoosystema, v. 33,
p. 245-279, 2011. https://doi.org/10.5252/z2011n3a1
Southward, E. C. Development of Perviata and Vestimentifera (Pogonophora). Hydrobiologia, v. 402, p. 185-202, 1999.
https://doi.org/10.1023/A:1003796626851
Steiner, T. M.; Amaral, A. C. Z. Two new species of Marphysa Quatrefages, 1865 (Eunicidae, Polychaeta) from
intertidal sandy beaches of the São Sebastião Channel, State of São Paulo (Brazil). Bulletin
of Marine Science, v. 67, p. 479-490, 2000. https://doi.org/10.11646/zootaxa.4377.2.3
Storch, V.; Welsch, U. Über den Bau und Entstehung der Mantelrandstacheln von Lingula unguis L. (Brachiopoda).
Zeitschrift für Wissenschaftliche Zoologie, v. 183, n. 3/4, p. 181-189, 1972.
Struck, T. H. Direction of evolution within Annelida and the definition of Pleistoannelida. Journal of Zoological
Systematics and Evolutionary Research, v. 49, p. 340-345, 2011. https://doi.org/10.1111/j.1439-0469.2011.00640.x
Struck, T. H.; Paul, C.; Hill, N.; Hartmann, S.; Hoesel, C.; Kube, M.; Lieb, B.; Meyer, A.; Tiedemann, R.; Purschke, G.;
Bleidorn, B. Phylogenomic analyses unravel annelid evolution. Nature, v. 471, p. 95-98, 2011. https://doi.org/10.1038/nature09864
Sweeney, L. B.; Luo, L. Fore brain: A hint of the ancestral cortex. Cell, v. 142, no. 5, p. 679-681, 2010.
https://doi.org/10.1016/j.cell.2010.08.024
Szucsich, N.; Wirkner, C. S. Homology: A synthetic concept of evolutionary robustness of patterns. Zoologica
Scripta, v. 36, p. 281-289, 2007. https://doi.org/10.1111/j.1463-6409.2007.00275.x
Tautz, D. Segmentation. Developmental Cell, v. 7, p. 301-312, 2004. https://doi.org/10.1016/j.devcel.2004.08.008
Temereva, E. N. New phoronid species Phoronopsis malakhovi (Lophophorata, Phoronida) from the South China
Sea. Zoologichesckii Zhurnal, v. 79, p. 1088-1093, 2000.
Tessemar-Raible, K.; Raible, F.; Christodoulou, F.; Guy, K.; Rembold, M.; Hausen, H.; Arendt, D. Conserved
sensory-neurosecretory cell types in annelid and fish forebrain: Insights into hypothalamus evolution.
Cell, v. 129, p. 1389-1400, 2007. https://doi.org/10.1016/j.cell.2007.04.041
Tomer, R.; Denes, A. S.; Tissamar-Raible, K.; Arendt, D. Profiling by image registration reveals common
origin of annelids mushroom bodies and vertebrate pallium. Cell, v. 142, p. 800-809, 2010.
https://doi.org/10.1016/j.cell.2010.07.04.041
Wägele, J. W. Foundations of Phylogenetic Systematics (erweiterte und übersetzte Auflage des Buches
von 2000). 2005.
Wägele, J. W. Hennig's Phylogenetic Systematics brought up to date. In: Williams, D. M.; Forey, P. L. (Eds).
Milestones in systematics. London: CRC Press, 2004. (The Systematics Association, Special Volume Series, 67).
p. 101-125.
Wagner, G. P. The character concept in evolutionary Biology. San Diego: Academic Press, 2001.
Walton, L. B. The polychaete ancestry of the insects. The American Naturalist, v. 61, no. 674, p. 226-250, 1927.
Weigert, A.; Helm, C.; Meyer, M.; Nickel, B.; Arendt, D.; Hausdorf, B.; Santos, S. R.; Halanych, K.; Purschke, G.;
Bleidorn, C. Illuminating the base of the annelid tree using transcriptomics. Molecular Phylogenetics and
Evolution, v. 31, p. 1391-1401, 2014. https://doi.org/10.1093/molbev/msu080
Westheide, W. The direction of evolution within the Polychaeta. Journal of Natural History, v. 31, no. 1,
p. 1-15, 1997. https://doi.org/10.1080/00222939700770011
Westheide, W.; McHugh, D.; Purschke, G.; Rouse, G. Systematization of the Annelida: Different approaches.
Hydrobiologia, v. 402, p. 291-307, 1999. https://doi.org/10.1023/A:1003713230485
Wheeler, Q. D. Losing the plot: DNA "barcodes" and taxonomy. Cladistics, v. 21, p. 405-407, 2005.
https://doi.org/10.1111/j.1096-0031.2005.00075.x
Wiley, E. O. Phylogenetics: The theory and practice of phylogentic systematics. New York: John Wiley &
Sons, 1981.
Wiley, E. O.; Lieberman, B. S. Phylogenetics: Theory and practice of phylogenetic systematic. 2. ed.
New York: Wiley-Blackwell, John Wiley & Sons, 2011.
Wilkinson, M. A comparison of two methods of character construction. Cladistics, v. 11, p. 297-308, 1995.
https://doi.org/10.1111/j.1096-0031.1995.tb00091.x
Williams, D. M. Homologues and homology, phenetics and cladistics: 150 years of progress. In: Williams, D. M.;
Forey, P. L. (Eds.). Milestones in systematics. London: CRC Press, 2004. (The Systematics Association,
Special Volume Series, 67). p. 191-124.
Williams, D. M.; Ebach, M. C. Confusing homologs as homologies: A reply to "On homology". Cladistics,
v. 28, p. 223-224, 2012. https://doi.org/10.1093/sysbio/syw094
Williams, D. M.; Ebach, M. C. Drowning by numbers: Rereading Nelson's "Nullius in Verba". The
Botanical Review, v. 71, p. 415-447, 2005.
Winchell, C. J.; Sullivan, J.; Cameron, C. B.; Swalla, B. J.; Mallatt, J. Evaluating hypotheses of deuterostome
phylogeny and chordate evolution with new LSU and SSU ribosomal DNA data. Molecular Phylogenetics and Evolution,
v. 19, p. 762-776, 2002. https://doi.org/10.1093/oxfordjournals.molbev.a004134M
Zeppelini, D. Phylogeny of Arrhopalites s.l. (Collembola: Symphypleona: Arrhopalitidae): Testing the monophyly
of the recently erected genera Arrhopalites s.s. and Pygmarrhopalites. Invertebrate Systematics,
v. 25, no. 2, p. 91-105, 2011. https://doi.org/10.1071/IS10044
Zeppelini, D. Relações filogenéticas do gênero Arrhopalites (Collembola: Symphypleona:
Arrhopalitidae). São Paulo: Universidade de São Paulo, 2001. (Doctorat thesis).
ISSN 2358-2731