Abstract
We present a numerical cladistic (parsimony) analysis of seed plants plus progymnosperms, using characters from all parts of the plant body, outgroup comparison, and a method of character coding that avoids biases for or against alternative morphological theories. The robustness of the results was tested by construction of alternative trees and analysis of subsets of the data. These experiments show that although some clades are strongly supported, they can often be related to each other in very different but nearly equally parsimonious ways, apparently because of extensive homoplasy. Our results support Rothwell’s idea that coniferopsids are derived fromCallistophyton- like platyspermic seed ferns with saccate pollen, but the hypothesis that they evolved fromArchaeopteris- like progymnosperms and the seed arose twice is nearly as parsimonious. Meyen’s division of seed plants into radiospermic and primarily and secondarily platyspermic lines is highly unparsimonious, but his suggestion that ginkgos are related to peltasperms deserves attention. Angiosperms belong among the platyspermic groups, as the sister group of Bennettitales,Pentoxylon, and Gnetales, and this “anthophyte” clade is best related toCaytonia and glossopterids, although relationships with other combinations of Mesozoic seed fern taxa are nearly as parsimonious. These results imply that the angiosperm carpel can be interpreted as a modified pinnate sporophyll bearing anatropous cupules (=bitegmic ovules), while gnetalian strobili are best interpreted as aggregations of highly reduced bennettitalian flowers, as anticipated by Arber and Parkin and Crane. Our most parsimonious trees imply that the angiosperm line (though not necessarily all its modern features) extended back to the Triassic, but a later derivation of angiosperms from some species ofCaytonia or Bennettitales, which would be nearly as parsimonious, should also be considered. These results raise the possibility that many features considered key adaptations in the origin and rise of angiosperms (insectpollinated flowers, rapid reproduction, drought tolerance) were actually inherited from their gymnospermous precursors. The explosive diversification of angiosperms may instead have been a consequence of carpel closure, resulting in increased speciation rates due to potential for stigmatic isolating mechanisms and/or new means of dispersal. DNA sequencing of extant plants and better information on anatomy, chemistry, sporophyll morphology, and embryology of Bennettitales and Caytoniales and the morphological diversity of Mesozoic anthophytes could provide critical tests of relationships.
Résumé
Nous présentons une analyse cladistique numérique (de parcimonie, ou économie d’hypothèses) des Spermatophytes plus Progymnospermes, utilisant des caractères de tous les organes du corps végétal, la comparaison extra-groupe, et une méthode de codification de caractères qui évite des prédispositions en faveur de ou contre les théories morphologiques alternatives. La solidité des résultats a été testé par la construction d’arbres phylétiques alternatifs et l’analyse de sous-ensembles des données. Selon ces expériences, certains phylums sont bien appuyés, mais ils peuvent être reliés de façons très différentes mais presque également économiques, apparemment à cause d’homoplasie répandue. Nos résultats confirment le concept de Rothwell, selon lequel les Coniféropsides sont dérivées de Ptéridospermes platyspermiques à pollen saccate proches deCallistophyton, mais l’hypothèse d’une dérivation de Progymnospermes proches d’Archaeopteris et d’une origine diphylétique de la graine est presque aussi économique. La division de Meyen des Spermatophytes en lignées radiospermiques et primairement et secondairement platyspermiques est très peu économique, mais son concept d’une affinité entre les Ginkgoales et les Peltaspermes mérite de l’attention. Les Angiospermes se situent parmi les groupes platyspermiques, comme groupe-frère des Bennettitales,Pentoxylon, et Gnetales, et le phylum ainsi constitué (“Anthophytes”) est le mieux lié àCaytonia et aux Glossoptérides, bien que des rapports avec d’autres combinaisons de taxons de Ptéridospermes mésozoïques soient presque aussi économiques. Ces résultats indiquent que le carpelle des Angiospermes peut être interprété comme une sporophylle pennée modifiée portant des cupules anatropes (=ovules bitégumentés), puisque les strobiles des Gnetales sont le mieux interprétés comme des agglomérations de fleurs bennettitaliennes fort réduites, comme l’a proposé Arber et Parkin et Crane. Nos arbres phylétiques les plus économiques supposent que la lignée des Angiospermes (mais pas forcément tous ses traits modernes) s’étend jusqu’au Trias, mais une origine plus récente à partir de quelque espèce deCaytonia ou de Bennettitales, hypothèses presque aussi économiques, doit aussi être considérée. Ces résultats suggèrent que plusieurs traits considérés comme des adaptationsclés dans l’origine ou l’expansion des Angiospermes (fleurs entomophiles, rapidité de reproduction, tolérance de sécheresse) étaient en fait hérités de leurs ancêtres gymnospermiques. La diversification explosive des Angiospermes pourrait plutôt être une conséquence de la clôture du carpelle, conduisant à un taux de spéciation élevé dû au potential pour des mécanismes d’isolement stigmatiques et/ou de nouveaux moyens de dispersion. L’étude de séquences d’ADN des végétaux actuels et de meilleurs renseignements sur l’anatomie, la phytochimie, la morphologie des sporophylles, et l’embryologie des Bennettitales et Caytoniales pourraient fournir des testes critiques de rapports phylétiques.
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Doyle, J.A., Donoghue, M.J. Seed plant phylogeny and the origin of angiosperms: An experimental cladistic approach. Bot. Rev 52, 321–431 (1986). https://doi.org/10.1007/BF02861082
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DOI: https://doi.org/10.1007/BF02861082