Abstract
The exine morphology, with emphasis on the ultrastructure, of six genera and species of Ecliptinae was studied with transmission electron microscopy and scanning electron microscopy. A caveate exine with a single layer of columellae and predominantly circular to sub-circular internal foramina (Helianthoid pattern of structure) is present in all the studied taxa and most probably in all the Ecliptinae. The morphologies of the internal foramina and foot layer are consistent within the Ecliptinae, and their potential taxonomic and phylogenetic relevance was discussed. The existence, for almost all the investigated characters and ratios, of a continuous gradient among the taxa, exists in the wedelioid group and constitutes a quite well defined parallelism between the exine data and those of the major clade of Ecliptinae (wedelioid group). The exine sculpture of the worldwide weed Eclipta prostrata (L.) L. presents some variation, which is, most probably, correlated to the existence of geographical clines and/or variations in its chromosome number. A mesoaperture concerning the foot layer and the upper part of the endexine exists in all the taxa studied and, presumably, in all the Ecliptinae, probably constituting a synapomorphy for all the Asteroideae and possibly for all the Asteraceae.



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Acknowledgments
We are deeply grateful to the Director of the Laboratory of Transmission Electron Microscopy of the Institute for Biomedical Imaging and Life Sciences (IBILI) of the Faculty of Medicine of the University of Coimbra, and especially to Dr. Mónica Zuzarte, for the very helpful technical assistance. We also thank very much the curators of Oporto (PO) and Oeiras (LISE) Herbaria and the late curator of the Coimbra Herbarium (COI), Dr. Maria Teresa Almeida for having allowed the pollen removal of the vouchers.
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Handling editor: Jürg Schönenberger.
To the memory of Dr. John J. Skvarla.
Appendix
Appendix
Studied specimens
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Clibadium armanii
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Brazil: Bahia, Ilhéus, 2 Aug 1979, AM. Carvalho 124 (COI).
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Eclipta prostrata
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Portugal: Alto Alentejo, Álcacer do Sal, 18 Sep 1960, J. Matos 7611 (PO 26527); Beira Baixa, Vila Velha de Rodão, margens do Tejo, 30 Sep 1955, Bento-Rainha 3204 (LISE 48840); Ribatejo, Várzea de Samora, Coutos de D. João, 5 Sep 1956, Vasconcellos 56018 (COI).
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Lasianthea podocephala
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Mexico: Chihuahua, rocky hills near Chihuahua, 29 Aug 1885, C.G. Pringle s.n. (COI).
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Melanthera integrifolia
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Hawaii: Punalu Court, Kau, 16 Jul 1976, O. Degener & I. Degener s.n. (COI); Punalu Court, Kau, 15 Jul 1976, O. Degener & I. Degener 33603 (COI).
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Perymenium parvifolium
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Mexico: State of Chihuahua, rocky hills near Chihuahua, 29 May 1885, C.G. Pringle 152 (COI).
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Wedelia prostrata
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Japan: Honshu, Shiraama in Awa, 29 Oct 1957, N. Maruyama & K. Okamoto 1604 (COI); Kyushu, Kagoshima Pref., Kimotsuki-gun, Sata.-cho, Odowari, 4 Jul 1974, H. Ohba & J. Marata 201 (COI).
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Coutinho, A.P., Valente, M., França, R. et al. A contribution to the knowledge of the exine, with special reference to the ultrastructure, in subtribe Ecliptinae (Heliantheae, Asteraceae). Plant Syst Evol 302, 1207–1216 (2016). https://doi.org/10.1007/s00606-016-1326-2
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DOI: https://doi.org/10.1007/s00606-016-1326-2


