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Molecular and morphological phylogeny of Menispermaceae (Ranunculales)

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Abstract

The phylogeny of Menispermaceae (Ranunculales) is reconstructed here using both morphological and molecular data from a broad sample of species. Morphological data include characters of leaves, wood, flowers, fruits, seeds, pollen and phytochemistry from 73 species representing the different subgroups recognized within the family. This dataset allowed us to study the morphological evolutionary trends in Menispermaceae. The molecular data focused on cpDNA sequences: rbcL and atpB. Maximum parsimony (MP) and Bayesian methods were used to reconstruct the phylogeny of Menispermaceae. The results obtained from the three datasets are partly incongruent. The morphology indicates that Menispermaceae can be divided into two major groups, one including the tribes Fibraureae and Tinosporeae, the other one including the tribes Anomospermeae, Pachygoneae and Menispermeae. Only Burasaia Thouars was not included in any of these groups. The characters of fruits and seeds appeared to be most useful to differentiate these groups whereas convergences were found for the androecium.

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Acknowledgments

We thank the Bogor Botanic Garden, the Kepong Forest Reasearch Institute of Malaysia, the Queen Sirikit Botanical Garden and the National Botanic Garden of Belgium for their assistance in specimen collection. We thank Sophie Nadot, Dario De Franceschi and two anonymous reviewers for useful comments on former versions. This study was partly funded by LVMH.

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  1. F. M. B. Jacques

    Present address: Kunming Institute of Botany, No. 132 Lanhei Road, Kunming, 650204, Yunnan, People’s Republic of China

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  1. Muséum National d’Histoire Naturelle, Dpt Histoire de la Terre CP 38, UMR 5143, USM 0203 Paléobiodiversité et paléoenvironnements, 75231, Paris, France

    F. M. B. Jacques

  2. Laboratoire Ecologie Systématique Evolution, UMR 8079, Université Paris-Sud, Orsay, France

    P. Bertolino

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Correspondence to F. M. B. Jacques.

Appendix 1: character coding

Appendix 1: character coding

Habit. 1. Port: 0 = liana (all climbing habit), 1 = small tree. Stem. 2. Tubercle: 0 = absent, 1 = present; 3. Young stem pubescent: 0 = no, 1 = yes; 4. Long trichomes longs: 0 = no, 1 = yes; 5. Old stems pubescent: 0 = no, 1 = yes; 6. Old bark verrucose: 0 = no, 1 = yes; 7. Old bark lenticellate: 0 = no, 1 = yes; 8. Old bark parcheminate: 0 = no, 1 = yes; Leaf. 9. Petiole insertion cupular: 0 = no, 1 = yes; 10. Petiole pubescent: 0 = no, 1 = yes; 11. Petiole striation: 0 = no, 1 = yes; 12. Petiole sulcate: 0 = no, 1 = yes; 13. Petiole enlarged at base: 0 = no, 1 = yes; 14. Petiole swollen at base: 0 = no, 1 = yes; 15. Petiole geniculate at base: 0 = no, 1 = yes; 16. Petiole swollen at apex: 0 = no, 1 = yes; 17. Petiole geniculate at apex: 0 = no, 1 = yes; 18. Petiole length: 0 = <leaf length, 1 = ≥leaf length; 19. Compound leaf: 0 = no, 1 = yes; 20. Lobed leaf: 0 = no, 1 = yes; 21. Lamina shape: 0 = length/width < 1, 1 = length/width ≥1; 22. Leaf acuminate: 0 = no, 1 = yes; 23. Leaf apex mucronate: 0 = no, 1 = yes; 24. Lamina base: 0 = convex, 1 = straight, 2 = concave; 25. Leaf peltate: 0 = no, 1 = yes, petiole insertion < 10% of leaf length, 2 = yes, petiole insertion ≥ 10% of leaf length; 26. Venation type: 0 = palmate, 1 = pinnate; 27. If leaf pinnate, distal venation pinnate: 0 = no, 1 = yes; 28. Number of basal veins: 0 = 3, 1 = 5, 2 = >5; 29. Veins suprabasal: 0 = no, 1 = yes; 30. Tertiary external veins from the lateral basal veins: 0 = no, 1 = yes; 31. Veins fusing with margin: 0 = no, 1 = yes; 32. Margin type: 0 = entire, 1 = toothed, 2 = sinuous; 33. Veins prominent on abaxial face: 0 = no, 1 = yes; 34. Upper face pubescent: 0 = no, 1 = yes (even only few hairs); 35. Location of pubescence on upper face: 0 = everywhere, 1 = only on veins; 36. Lower face pubescent: 0 = no, 1 = yes (even only few hairs); 37. Location of pubescence on lower face: 0 = everywhere, 1 = only on veins; 38. Domatia: 0 = no, 1 = yes; 39. Leaf texture: 0 = membranous, 1 = chartaceous, 2 = coriaceous; 40. Shape of upper epidermis cells: 0 = polygonal, 1 = irregular; 41. Anticlinal wall of those cells: 0 = straight, 1 = sinuous, 2 = sinuolate; 42. Shape of lower epidermis cells: 0 = polygonal, 1 = irregular; 43. Anticlinal wall of those cells: 0 = straight, 1 = sinuous, 2 = sinuolate; 44. Stomata type: 0 = anomocytic, 1 = staurocytic, 2 = cyclocytic, 3 = actinocytic. Wood. 45. Wood with successive cambia: 0 = no, 1 = yes; 46. Wood rays lignified: 0 = no, 1 = yes. Inflorescence. The highly variability in inflorescence between species does not allow a simple coding of inflorescence type, the inflorescences characteristics were therefore separated in several characters. 47. Flower in inflorescence: 0 = no, 1 = yes; 48. Composition degree of inflorescence: 0 = 1, 1 = 2, 2 = >2; 49. Type of highest level of inflorescence: 0 = raceme, 1 = cyme; 50. Type of lowest level of inflorescence: 0 = raceme, 1 = cyme; 51. Inflorescences fasciculate: 0 = no, 1 = yes; 52. Inflorescence position: 0 = axillary, 1 = ramiflorous, 2 = terminal; 53. Inflorescence on old wood: 0 = no, 1 = yes; 54. Pauciflorous inflorescence: 0 = no, 1 = yes; 55. Inflorescence pubescent: 0 = no, 1 = yes; 56. Inflorescence length: 0 = <leaf length, 1 = ≥leaf length. Flower. As Menispermaceae species are all dioic and male and female flowers as sometimes different structures, all male and female characters were handled separately. 57. Flower type: 0 = hermaphrodite, 1 = monoecious, 2 = dioecious; 58. Merosity: 0 = 3, 1 = 2, 2 = 5. 59. Sepal number in male flower relatively to merosity: 0 =  >3, 1 = 1, 2 = 2, 3 = 3; 60. Sepal number in female flower relatively to merosity: 0 = >3, 1 = 1, 2 = 2, 3 = 3; 61. Outer sepal pubescent: 0 = no, 1 = yes; 62. Inner sepals pubescent on outer face: 0 = no, 1 = yes; 63. Relative size of inner sepals: 0 = small, 1 = equal, 2 = large; 64. Sepals in tube: 0 = no, 1 = yes; 65. Sepals fleshy: 0 = no, 1 = yes; 66. Sepal type: 0 = valvate, 1 = imbricate; 67. Petal number in male flower relatively to merosity: 0 = 0, 1 = 1, 2 = 2, 3 = >3; 68. Petal number in female flower relatively to merosity: 0 = 0, 1 = 1, 2 = 2, 3 = ≥3, 4 = 1 petal; 69. Fused petal: 0 = no, 1 = yes; 70. Petal size relatively to inner sepals: 0 = minute, 1 = smaller than sepals, 2 = equal, 3 = larger. 71. Petals surrounding stamens: 0 = no, 1 = yes; 72. Petal texture: 0 = membranous, 1 = fleshy; 73. Stamen number relatively to merosity: 0 = >3, 1 = 1, 2 = 2, 3 = 3; 74. Stamen fusion: 0 = no, 1 = only inners, 2 = all; 75. Synandrium type: 0 = no synandrium, 1 = central column, vertical anthers, 2 = central column, horizontal anthers, 3 = stalked pyramid, 4 = sessile pyramid; 76. Connective extended over the anthers: 0 = no, 1 = yes; 77. Filament enlarging: 0 = no, 1 = yes; 78. Connective highly visible: 0 = no, 1 = yes; 79. Anther shape: 0 = globular, 1 = ellipsoid; 80. Orientation of outer anthers: 0 = introrse, 1 = extrorse; 81. Orientation of inner anthers: 0 = introrse, 1 = extrorse; 82. Orientation of dehiscence slit: 0 = transversal, 1 = longitudinal; 83. Pistillodes: 0 = no, 1 = yes. 84. Hairs in the center of male flower: 0 = no, 1 = yes; 85. Staminodes number relatively to merosity: 0 = 0, 1 = 1, 2 = 2, 3 = ≥3. 86. Anthers on the staminodes: 0 = no, 1 = yes. 87. Carpel number relatively to merosity: 0 = >2, 1 = 1, 2 = 2, 3 = 3, 4 = 1 carpel; 88. Ovule number per carpel: 0 = 1 or 2, 1 = >2; 89. Style: 0 = no, 1 = yes; 90. Stigma recurved: 0 = no, 1 = yes; 91. Stigma shape: 0 = entire, 1 = bifid, 2 = trifid; 92. Carpel pubescent: 0 = no, 1 = yes. Fruit. 93. Fruits apocarpous, 0 = no, 1 = yes; 94. Fruit shape: 0 = spheroid, 1 = ellipsoid; 95. Fruit colour: 0 = red, 1 = white, 2 = black; 96. Carpophore: 0 = no, 1 = yes; 97. Stylar scar position: 0 = basal, 1 = lateral, 2 = terminal; 98. Fruit pubescent: 0 = no, 1 = yes; 99. Mesocarp sclerified: 0 = no, 1 = yes; 100. Endocarp sclerified: 0 = no, 1 = yes; 101. Endocarp straight: 0 = no, 1 = yes; 102. Endocarp with excavated faces: 0 = no, 1 = yes; 103. If not straight, endocarp larger than wide: 0 = no, 1 = yes; 104. Endocarp with dorsal ridges: 0 = no, 1 = yes; 105. Number of lateral ridges on each face; 106. Ridges wing-shaped: 0 = no, 1 = yes; 107. Endocarp with lateral radial ridges: 0 = no, 1 = yes; 108. Endocarp surface reticulate: 0 = no, 1 = yes; 109. Type of reticulum: 0 = as hollow, 1 = as bump; 110. Endocarp surface with spines: 0 = no, 1 = yes; 111. Condyle present: 0 = no, 1 = yes; 112. Condyle outer conspicuous: 0 = no, 1 = yes; 113. Condyle type: 0 = single, 1 = double; 114. Condyle perforate: 0 = no, 1 = yes; 115. Condyle in prominent ventral chamber: 0 = no, 1 = yes; 116. If endocarp not straight, condyle parallel to symmetry plane: 0 = no, 1 = yes; 117. Embryo shape: 0 = straight, 1 = curved; 118. Endosperm: 0 = no, 1 = yes; 119. Endosperm ruminate: 0 = no, 1 = yes; 120. Cotyledon type: 0 = leafy, 1 = fleshy; 121. Cotyledons divaricate: 0 = no, 1 = yes; 122. Radicle length: 0 = small, 1 = equal. Pollen. 123. Pollen with colpus: 0 = no, 1 = yes; 124. Pollen with pore: 0 = no, 1 = yes; 125. Position of apertures: 0 = fosaperturate, 1 = angulaperturate, 2 = spiraperturate; 126. Pollen shape: 0 = P/E < 1, 1 = P/E = 1, 2 = P/E > 1; 127. Exine thickness: 0 = <1.5 nm, 1 = 1.5 nm, 2 = >1,5 nm; 128. Reticulation coarse: 0 = no, 1 = yes; 129. Wall section: 0 = cylindrical, 1 = triangular. 130. Endoaperture shape: 0 = elliptic, 1 = circular; 131. Lumina regular: 0 = no, 1 = yes; 132. Lumina hexagonal: 0 = no, 1 = yes. Phytochemistry. 133. Benzylisoquinoline: 0 = no, 1 = yes; 134. Bisbenzylisoquinoline: 0 = no, 1 = yes; 135. Proaporphine: 0 = no, 1 = yes; 136. Aporphine: 0 = no, 1 = yes; 137. Tropoloneisoquinoline: 0 = no, 1 = yes; 138. Azafluoranthene: 0 = no, 1 = yes; 139. Isooxaporphine: 0 = no, 1 = yes; 140. Protoberberine: 0 = no, 1 = yes; 141. Morphinane: 0 = no, 1 = yes; 142. Hasubanane: 0 = no, 1 = yes; 143. Acutimine: 0 = no, 1 = yes; 144. Isoquinoline: 0 = no, 1 = yes; 145. Phentylcinnamide: 0 = no, 1 = yes.

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Jacques, F.M.B., Bertolino, P. Molecular and morphological phylogeny of Menispermaceae (Ranunculales). Plant Syst Evol 274, 83–97 (2008). https://doi.org/10.1007/s00606-008-0038-7

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