Since the time of Darwin1, evolutionary biologists have been fascinated by the spectacular adaptations to insect pollination exhibited by orchids. However, despite being the most diverse plant family on Earth2, the Orchidaceae lack a definitive fossil record and thus many aspects of their evolutionary history remain obscure. Here we report an exquisitely preserved orchid pollinarium (of Meliorchis caribea gen. et sp. nov.) attached to the mesoscutellum of an extinct stingless bee, Proplebeia dominicana, recovered from Miocene amber in the Dominican Republic, that is 15–20 million years (Myr) old3. This discovery constitutes both the first unambiguous fossil of Orchidaceae4 and an unprecedented direct fossil observation of a plant–pollinator interaction5,6. By applying cladistic methods to a morphological character matrix, we resolve the phylogenetic position of M. caribea within the extant subtribe Goodyerinae (subfamily Orchidoideae). We use the ages of other fossil monocots and M. caribea to calibrate a molecular phylogenetic tree of the Orchidaceae. Our results indicate that the most recent common ancestor of extant orchids lived in the Late Cretaceous (76–84 Myr ago), and also suggest that the dramatic radiation of orchids began shortly after the mass extinctions at the K/T boundary. These results further support the hypothesis of an ancient origin for Orchidaceae.
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We thank Y. Goldman for facilitating access to the amber inclusion discussed here, G. Romero for his assistance in the examination of herbarium specimens, and G. Alpert and D. Smith for assistance in the production of the fossil microphotographs. We thank B. Archibald, C. Bell, M. Chase, A. Knoll, R. van der Ham, D. Hewitt, C. Jaramillo, M. Patten, E. Pringle, J. Pringle and T. Quental for useful comments. This research was sponsored by grants from the Barbour Fund (Museum of Comparative Zoology) and the National Science Foundation (DDIG) to S.R.R. and N.E.P., and a grant from the Fulbright Junior Scholar programme to B.G.
Author Contributions S.R.R., B.G. and N.E.P. procured and curated the specimen. S.R.R., B.G. and R.B.S. reviewed herbaria specimens, analysed ancestral floral morphology, and coded and analysed morphological characters. S.R.R. and C.R.M. designed dating approaches and considered their interpretation. S.R.R. wrote the paper. All authors discussed the results and commented on the manuscript.
Reprints and permissions information is available at www.nature.com/reprints. The authors declare no competing financial interests.
This file contains Supplementary Figure 1 with Legend, detailing the fossil-calibrated molecular clock analyses presented in Figure 4; Supplementary Methods 1-6, showing the morphological character codes and the matrix used to explore the phylogenetic placement of M. caribea, the methodology used in the phylogenetic analyses and divergence time estimation, and additional comments on the palaeontology of fossil calibrations; Supplementary Tables 1-3, indicating herbarium specimens examined, age estimates obtained with different methods, and GenBank accession numbers; and Supplementary Notes, indicating additional cited references. (PDF 534 kb)
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Ramírez, S., Gravendeel, B., Singer, R. et al. Dating the origin of the Orchidaceae from a fossil orchid with its pollinator. Nature 448, 1042–1045 (2007). https://doi.org/10.1038/nature06039
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