Representing supraspecific taxa in higher-level phylogenetic analyses: guidelines for palaeontologists

53 1 January 1 9 10.1111/j.1475-4983.2009.00918.x

BRUSATTE, S. L. 2010. Representing supraspecific taxa in higher-level phylogenetic analyses: guidelines for palaeontologists. Palaeontology53, 1, 1–9.

Stephen L. Brusatte As phylogenetic analyses become larger, one of the greatest methodological difficulties is representing speciose supraspecific clades in higher-level analyses (e.g. trilobites within studies of arthropod phylogeny). Several strategies have been proposed, including using representative single composite terminals or species-level exemplars, and various methods are currently used in the palaeontological literature. However, this is problematic, as simulation studies and empirical arguments in the systematics literature have clearly identified multiple exemplars as the optimal method. The continuing usage of suboptimal strategies in palaeontology may lessen the accuracy of phylogenies and hampers comparison between alternative studies. Here, I outline problems with suboptimal strategies, review arguments in support of multiple exemplars and provide guidelines for palaeontologists undertaking higher-level phylogenetic analyses
  • ANDERSON, J. S., REISZ, R. R., SCOTT, D., FRÖBISCH, N. B. and SUMIDA, S. S. 2008. A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders. Nature, 453, 515–518.
  • BENNETT, S. C. 1996. The phylogenetic position of the Pterosauria within the Archosauromorpha. Zoological Journal of the Linnean Society, 118, 261–308.
  • BENTON, M. J. 1999. Scleromochlus taylori and the origin of dinosaurs and pterosaurs. Philosophical Transactions of the Royal Society of London, Series B, 354, 1423–1446.
  • BENTON, M. J. 2004. Origin and relationships of Dinosauria. 7–19. In WEISHAMPEL, D. B., DODSON, P. and OSMOLSKA, H. (eds). The Dinosauria. Second edition. University of California Press, Berkelely, California, 861 pp.
  • BENTON, M. J. and WALKER, A. D. 2002. Erpetosuchus, a crocodile-like basal archosaur. From the Late Triassic of Elgin, Scotland. Zoological Journal of the Linnean Society, 136, 25–47.
  • BININDA-EMONDS, O. R. P. (ed.). 2004. Phylogenetic supertrees: combining information to reveal the Tree of Life., Computation Biology, Vol. 4. Kluwer Academic Publishers, Dordrecht, the Netherlands.
  • BININDA-EMONDS, O. R. P., BRYANT, H. N. and RUSSELL, A. P. 1998. Supraspecific taxa as terminals in cladistic analysis: implicit assumptions of monophyly and a comparison of methods. Biological Journal of the Linnean Society, 64, 101–133.
  • BLOCH, J. I., SILCOX, M. T., BOYER, D. M. and SARGIS, E. J. 2007. New Paleocene skeletons and the relationship of plesiadapiforms to crown-clade primates. Proceedings of the National Academy of Sciences USA, 104, 1159–1164.
  • BRUSATTE, S. L. and SERENO, P. C. 2008. Phylogeny of Allosauroidea (Dinosauria: Theropoda): comparative analysis and resolution. Journal of Systematic Palaeontology, 6, 155–182.
  • CARROLL, R. L. 2007. The Palaeozoic ancestry of salamanders, frogs, and caecilians. Zoological Journal of the Linnean Society, 150, 1–140.
  • CLARK, J. M., NORELL, M. A. and MAKOVICKY, P. J. 2002. Cladistic approaches to the relationships of birds to other theropod dinosaurs. 31–60. In CHIAPPE, L. M. and WITMER, L. M. (eds). Mesozoic birds: above the heads of Dinosaurs. University of California Press, Berkeley, California.
  • DONOGHUE, P. C. J. and SMITH, M. P. 2001. The anatomy of Turinia pagei (Powrie), and the phylogenetic status of the Thelodonti. Transactions of the Royal Society of Edinburgh: Earth Sciences, 92, 15–37.
  • ESTES, R. K., DE QUIEROZ, K. and GAUTHIER, J. A. 1988. Phylogenetic relationships within Squamata. 119–281. In ESTES, R. and PREGILL, G. K. (eds). Phylogenetic relationships of the Lizard families. Stanford Univ. Press, Palo Alto, California.
  • GATESY, J., MATTHEE, C., DE SALLE, R. and HAYASHI, C. 2002. Resolution of supertree/supermatrix paradox. Systematic Biology, 51, 652–664.
  • GOLOBOFF, P. A. 1999. Analyzing large data sets in reasonable times: solutions for composite optima. Cladistics, 15, 415–428.
  • GOLOBOFF, P. A., FARRIS, J. S. and NIXON, K. C. 2003. T.N.T.: Tree analysis using new technology. Version 1.0. Program and documentation available at http://www.zmuc.dk/public/Phylogeny/TNT.
  • HARRIS, S. R., PISANI, D., GOWER, D. J. and WILKINSON, M. 2007. Investigating stagnation in morphological phylogenies using consensus data. Systematic Biology, 56, 125–129.
  • JENNER, R.A. 2006. Unburdening evo-devo: ancestral attractions, model organisms, and basal baloney. Development Genes and Evolution, 216, 385–394.
  • JUUL, L. 1994. The phylogeny of basal archosaurs. Palaeontologica Africana, 31, 1–38.
  • LANGER, M. C. and BENTON, M. J. 2006. Early dinosaurs: a phylogenetic study. Journal of Systematic Palaeontology, 4, 309–358.
  • MALIA, M. J., LIPSCOME, D. L. and ALLARD, M. W. 2003. The misleading effects of composite taxa in supermatrices. Molecular Phylogenetics and Evolution, 27, 522–527.
  • NESBITT, S. 2007. The anatomy of Effigia okeeffeae (Archosauria, Suchia), theropod-like convergence, and the distribution of related taxa. Bulletin of the American Museum of Natural History, 302, 1–84.
  • NIXON, K. C. 1999. The parsimony ratchet, a new method for rapid parsimony analysis. Cladistics, 15, 407–414.
  • NOVAS, F. E. 1996. Dinosaur monophyly. Journal of Vertebrate Paleontology, 16, 723–741.
  • PARRISH, J. M. 1993. Phylogeny of the Crocodylotarsi, with reference to archosaurian and crurotarsan monophyly. Journal of Vertebrate Paleontology, 13, 287–308.
  • PRENDINI, L. 2000. Phylogeny and classification of the superfamily Scorpionoidea Latreille 1802 (Chelicerata, Scorpiones): an exemplar approach. Cladistics, 16, 1–78.
  • PRENDINI, L. 2001. Species or supraspecific taxa as terminals in cladistic analysis? Groundplans versus exemplars revisited. Systematic Biology, 50, 290–300.
  • PRENDINI, L. 2003. A new genus and species of bothriurid scorpion from the Brandberg Massif, Namibia, with a reanalysis of bothriurid phylogeny and a discussion of the phylogenetic position of Lisposoma Lawrence. Systematic Entomology, 28, 149–172.
  • PRENDINI, L., CROWE, T. M. and WHEELER, W. C. 2003. Systematics and biogeography of the Family Scorpionidae (Chelicerata: Scorpiones), with a discussion of phylogenetic methods. Invertebrate Systematics, 17, 185–259.
  • PRENDINI, L. and WHEELER, W. C. 2005. Scorpion higher phylogeny and classification, taxonomic anarchy, and standards for peer review in online publishing. Cladistics, 21, 446–494.
  • O’LEARY, M. A. 1999. Parsimony analysis of total evidence from extinct and extant taxa and the cetacean-artiodactyl question (Mammalia, Ungulata). Cladistics, 15, 315–330.
  • QUICKE, D. L., TAYLOR, J. J. and PURVIS, A. 2001. Changing the landscape: a new strategy for estimating large phylogenies. Systematic Biology, 50, 60–66.
  • RICE, K. A., DONOGHUE, M. J. and OLMSTEAD, R. G. 1997. Analyzing large data sets: rbcL revisited. Systematic Biology, 46, 554–563.
  • ROSHAN, U. M., MORET, B. M. E., WILLIAMS, T. L. and WARROW, T. 2004. Rec-I-DCM3: a fast algorithmic technique for reconstructing large phylogenetic trees. Proceedings 3rd IEEE Computational Systems Bioinformatics Conference, CSB 2004, 98–109.
  • RUTA, M., COATES, M. I. and QUICKE, D. L. J. 2003. Early tetrapod relationships revisited. Biological Reviews, 78, 251–345.
  • RUTA, M. and COATES, M. I. 2007. Dates, nodes and character conflict: addressing the lissamphibian origin problem. Journal of Systematic Palaeontology, 5, 69–122.
  • SALISBURY, B. A. and KIM, J. 2001. Ancestral state estimation and taxon sampling density. Systematic Biology, 50, 557–564.
  • SANSOM, R. S. 2008. The origin and early evolution of the Osteostraci (Vertebrata): a phylogeny for the Thyestiida. Journal of Systematic Palaeontology, 6, 317–332.
  • SCHOCH, R. R. 2009. Evolution of life cycles in early amphibians. Annual Review of Earth and Planetary Sciences, 27, 15.1–15.28.
  • SCOTT, E. 2005. A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data. Cladistics, 21, 507–574.
  • SERENO, P. C. 1991. Basal archosaurs: phylogenetic relationships or functional implications. Society of Vertebrate Paleontology Memoir, 2, 1–53.
  • SERENO, P. C. and ARCUCCI, A. B. 1990. The monophyly of crurotarsal archosaurs and the origin of bird and crocodile ankle joints. Neues Jahrbuch für Geologie und Palontologie, Abhandlung, 180, 21–52.
  • SIMMONS, N. B. 2001. Misleading results from the use of ambiguity coding to score polymorphisms in higher-level taxa. Systematic Biology, 50, 613–620.
  • TRUEB, L. and CLOUTIER, R. 1991. A phylogenetic investigation of the inter- and intrarelationships of the Lissamphibia (Amphibia: Temnospondyli). 223–313. In SCHULTZE, H.-P. and TRUEB, L. (eds). Origins of the higher groups of tetrapods: controversy and consensus. Cornell Univ. Press, Ithaca, New York.
  • WIENS, J. J. 1998. The accuracy of methods for coding and sampling higher-level taxa for phylogenetic analysis: a simulation study. Systematic Biolology, 47, 397–413.
  • WIENS, J. J. 2003. Incomplete taxa, incomplete characters, and phylogenetic accuracy: is there a missing data problem? Journal of Vertebrate Paleontology, 23, 297–310.
  • WIENS, J. J. 2005. Can incomplete taxa rescue phylogenetic analyses from long-branch attraction? Systematic Biology, 54, 731–742.
  • WIENS, J. J. 2006. Missing data and the design of phylogenetic analyses. Journal of Biomedical Informatics, 39, 34–42.
  • YEATES, D. K. 1995. Groundplans and exemplars: paths to the tree of life. Cladistics, 11, 343–357.
Wiley Online Library