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Is morphology telling the truth about the evolution of the species rich genus Peperomia (Piperaceae)?

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An Erratum to this article was published on 30 May 2009

An Erratum to this article was published on 30 May 2009

Abstract

Peperomia is with approximately 1,600 species one of the species rich angiosperm genera. Several characters on which current infrageneric classifications are based are influenced by parallel evolution. A well-resolved molecular backbone phylogeny of the genus is needed to address evolutionary questions about morphological traits. Based on separate and combined analyses of a morphological data set and three molecular data sets, phylogenetic relationships within Peperomia are investigated with respect to character evolution. The resulting trees from different datasets are highly congruent. Morphological characters are mapped on a combined molecular tree, visualizing the contrast between previously used homoplastic characters and some newly observed characters, that can be used to delimit monophyletic groups. Length mutational events of the chloroplast dataset are coded and plotted on the respective tree, to test if indels support alternative hypothesis of relationships found in the nuclear datasets as well as the overall performance of indels compared with substitutional mutations. Our findings indicate that length distribution of indels is highest among five and six bp events. Autapomorphic and synapomorphic length mutations are most frequent in both insertions and deletions and are also more frequent independent of the length of the mutation. Concluding, independent of the length, mutations are of phylogenetic importance and should not be disregarded. None of the homoplastic indels turn into synapomorphic indels, supporting the different topology of the nrDNA tree but indicate areas of molecular evolution in favour of length mutations resulting in independent events.

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Acknowledgments

Financial support for this study came from the Research Foundation-Flanders (FWO G.0172.07), the German Science Foundation (DFG NE 681/5-1), the Department of Biology, Ghent University (travel grant to MS), the Special Research Fund (BOF, Ghent University) (grant to LV) and the Friends of the Botanical Garden, Ghent. During the preparation of this manuscript, the last author was supported by the German Academic Exchange (DAAD) Postdoc program, which is gratefully acknowledged. We also gratefully acknowledge material received from the Botanical Gardens Bonn and Berlin-Dahlem (Germany), Robert Maijer (Joure, The Netherlands) and Guillermo Pino (Lima, Peru). Thanks are also due to Pieter Asselman (technical assistant) for assisting with data assembling and Guido Mathieu (Ghent, Belgium) for remarks on the manuscript. Many thanks to Daniel Nickrent (Southern Illinois University) for remarks on 26S. Thanks are also due to Isabelle De Graeve and Ľuboš Majeský for help with the practical work on Peperomia fruits. Thanks to Marcel Verhaegen (BR) for the SEM images and Jan Rammeloo, Director of the National Botanic Garden of Belgium. Finally, we thank James A. Doyle and an anonymous reviewer for their constructive comments.

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Authors and Affiliations

  1. Department of Biology, Research Group Spermatophytes, Ghent University, K.L. Ledeganckstraat 35, 9000, Ghent, Belgium

    Marie-Stéphanie Samain, Liesbeth Vanderschaeve, Peter Chaerle & Paul Goetghebeur

  2. Institut für Botanik, Zellescher Weg 20b, Technische Universität Dresden, 01062, Dresden, Germany

    Christoph Neinhuis

  3. Institut für Botanik, Plant Phylogenetics and Phylogenomics Group, Zellescher Weg 20b, Technische Universität Dresden, 01062, Dresden, Germany

    Stefan Wanke

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  1. Marie-Stéphanie Samain
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Correspondence to Marie-Stéphanie Samain.

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An erratum to this article can be found at http://dx.doi.org/10.1007/s00606-009-0186-4

Appendices

Appendix 1: Morphological characters

All characters are unordered.

  1. 1.

    Life form: (0) perennial; (1) annual.

  2. 2.

    Growth form: (0) terrestrial; (1) epiphytic.

  3. 3.

    Tuber: (0) absent; (1) present.

  4. 4.

    Stem orientation: (0) subterraneous; (1) erect; (2) prostrate; (3) decumbent; (4) pendent.

  5. 5.

    Stem width: (0) tuberous; (1) succulent and thick; (2) succulent and thin; (3) slender; (4) filiform.

  6. 6.

    Stem cross-section: (0) terete; (1) tuberous; (2) angular; (3) ribbed; (4) irregular.

  7. 7.

    Stem vestiture: (0) glabrous; (1) pubescent; (2) pilose.

  8. 8.

    Stem internodes: (0) absent; (1) elongated; (2) shortened.

  9. 9.

    Architecture: (0) no change before flowering; (1) change before flowering.

  10. 10.

    Leaf arrangement: (0) spiral; (1) whorled; (2) basal rosette.

  11. 11.

    Petiole position: (0) at lamina base; (1) peltate + petiole central; (2) peltate + petiole near base.

  12. 12.

    Leaf shape: (0) window-leaf; (1) peltate; (2) peltate window-leaf; (3) ovate; (4) obovate; (5) oblong; (6) orbicular; (7) cordate; (8) lanceolate; (9) rhomboid

  13. 13.

    Leaf texture: (0) membranous; (1) succulent; (2) coriaceous.

  14. 14.

    Leaf venation pattern: (0) palmate; (1) pinnate; (2) obscure. It is difficult to distinguish the leaf venation in very succulent leaves because of the multiple/multiseriate hypodermis.

  15. 15.

    Leaf vestiture adaxial side: (0) absent; (1) present.

  16. 16.

    Leaf vestiture abaxial side: (0) absent; (1) present.

  17. 17.

    Leaf aroma: (0) absent; (1) present.

  18. 18.

    Petiole vestiture: (0) glabrous; (1) pubescent; (2) tomentose.

  19. 19.

    Inflorescence type: (0) spadix; (1) panicle

  20. 20.

    Inflorescence is shed: (0) no; (1) yes. This character is especially interesting in the species with changing architecture before flowering (see character 9). In these species (e.g. Panicularia, the inflorescence together with a major part of the elongated stem is dropped.

  21. 21.

    Inflorescence position: (0) terminal; (1) (pseudo)lateral; (2) terminal and (pseudo)lateral.

  22. 22.

    Inflorescence ramification: (0) always solitary spadix; (1) 1–2 spadices; (2) 2–3 spadices; (3) 3–6 spadices; (4) numerous spadices. States 1, 2 and 3: number of spadices is varying within the same plant.

  23. 23.

    Peduncle vestiture: (0) glabrous; (1) pubescent; (2) pilose.

  24. 24.

    Peduncle bracts: (0) absent; (1) one; (2) two; (3) three. In most Peperomia species a nonpeltate green to brown bract is present on the peduncle of the inflorescence. However, in some species, this bract seems to be absent. In few species, there are two to three nonpeltate bracts at the base of the spadix.

  25. 25.

    Rachis vestiture: (0) absent; (1) present.

  26. 26.

    Distance between flowers: (0) crowded; (1) loose.

  27. 27.

    Floral bract shape: (0) orbicular; (1) lanceolate; (2) rhomboid; (3) obovate.

  28. 28.

    Style: (0) absent; (1) present. The style is absent in some Peperomia species with an apical stigma (see character 33). In all Peperomia species with a subapical stigma we have investigated so far, the style is absent as well.

  29. 29.

    Stigma position: (0) apical; (1) subapical.

  30. 30.

    Fruit shape: (0) globose; (1) ovoid; (2) obovoid; (3) cylindric; (4) oblong.

  31. 31.

    Fruit attachment: (0) sessile; (1) pedicellate; (2) pseudopedicellate. The flowers and the drupes are sessile in most species, pedicellate in some. In other species, the flowers are sessile, but a pseudopedicel develops as an outgrowth of the rachis when the fruit reaches maturity (Sastrapradja 1968).

  32. 32.

    Fruit surface: (0) smooth; (1) rugose; (2) papillose (whole surface); (3) papillose (mainly at fruit base); (4) papillose (mainly at fruit base and apex); (5) rugose with lacunae; (6) verrucose.

  33. 33.

    Pseudocupula: (0) absent; (1) present. A very clear morphological synapomorphy for the subgenus Micropiper sensu Dahlstedt is the so-called pseudocupula at the basis of the drupe (Dahlstedt 1900).

  34. 34.

    Fruit apex: (0) pointed; (1) oblique; (2) rostrated—rostrum shorter than body—not curved; (3) rostrated—rostrum shorter than body—slightly curved; (4) rostrated—rostrum shorter than body—hooked; (5) rostrated—as long as body—hooked.

Appendix 2: Morphological dataset

-: character state unknown

*: character polymorphic

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Samain, MS., Vanderschaeve, L., Chaerle, P. et al. Is morphology telling the truth about the evolution of the species rich genus Peperomia (Piperaceae)?. Plant Syst Evol 278, 1–21 (2009). https://doi.org/10.1007/s00606-008-0113-0

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