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First published online June 10, 2009; doi:10.3732/ajb.0800236 American Journal of Botany 96: 1361-1371 (2009) © 2009 Botanical Society of America, Inc.

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Perianth evolution in the sandalwood order Santalales¹

² Department of Phanerogamic Botany, Swedish Museum of Natural History, Svante Arrhenius väg 7, P.O. Box 50007 SE-104 05 Stockholm, Sweden ³ Royal Botanic Garden Edinburgh, 20A Inverleith Row, Edinburgh EH3 5 LR, Scotland, UK

ABSTRACT

Flowers of Santalales remain largely unexplored with several questions of homology unanswered despite the large size of the order. Morphological and ontogenetic floral studies have the potential to identify new informative characters. We studied floral development in species of Loranthaceae, Santalaceae, Opiliaceae, and "Olacaceae" with scanning electron microscopy to clarify the origin and evolution of the perianth in Santalales. The perianth is either dichlamydeous or when monochlamydeous interpreted as sepals, petals, or tepals. A girdling calyculus of debatable origin is found in some clades. We show that species of Diogoa and Heisteria in "Olacaceae" have a dichlamydeous perianth, unlike Olax in which the calyx is replaced by a calyculus. The calyculus arises by development of two lateral primordia, supporting the hypothesis of bracteole origin. A calyculus with similar development is present in species of Loranthaceae and possibly of Opiliaceae, suggesting a position of Olax closer to these families than to traditional genera of "Olacaceae". The monochlamydeous perianth in Santalaceae is shown to correspond to petals of other members of Santalales. Flower ontogenetic evidence suggests a repeated loss of the calyx, replacement by a calyculus, and further loss, leading to monochlamydeous perianths in Santalaceae.

Key Words: flower morphology • Loranthaceae • Olacaceae • Opiliaceae • ontogeny • perianth evolution • Santalaceae • Santalales

Received for publication 10 July 2008. Accepted for publication 11 March 2009.

FOOTNOTES

¹ The authors thank F. Christie for assistance with the SEM. They acknowledge Synthesys, support made available by the European Community–Research Infrastructure Action under the FP6 Structuring the European Research Area Program for two grants, one to L.W. (GB-TAF-2700) for a visit to RBG Edinburgh and the other to L.R.D.C. (SE-TAF-4524) for a visit to NRM, Stockholm, and also the Sibbald Trust, RBGE for support to L.W. They are grateful to A. Anderberg and to the Directors of the Herbaria at Kew (K) and Wageningen (WAG) for permission to use the pickled collections, and they thank D. Harris for collecting material of Diogoa. V. Malécot, D. Nickrent, and one anonymous reviewer for valuable comments on the paper.

⁴ Author for correspondence (e-mail: livia.wanntorp{at}nrm.se)

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