Dating the origin of the Orchidaceae from a fossil orchid with its pollinator

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The antiquity of this relationship is demonstrated by the remarkable fossil shown on the cover. Preserved in 15–20-million-year-old amber from the Dominican Republic, a worker stingless bee carries an orchid pollinarium. The pollinarium is the male reproductive structure, transferred as a single unit during pollination, complete with a mass of pollen grains. The find, surprisingly, is the first definitive fossil from the Orchidaceae family. The pollinarium's morphology locates the plant in the subtribe Goodyerinae. By combining this information with dates from related fossil plants, a new calibration of the molecular phylogeny of the Orchidaceae has been obtained. The analysis rejects the common assumption of a recent (Tertiary) origin for orchids, and suggests, instead, that they originated during the Late Cretaceous, 80 million years ago. The antiquity of the close relationship between orchids and their pollinators is demonstrated by a 15–20 million years old fossil of an orchid pollinarium attached to the business end of a stingless bee, preserved in amber from the Dominican Republic. This spectacular find is used to constrain the evolution of orchids, suggesting that the most recent common ancestor of orchids lived in the Late Cretaceous. 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. 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The antiquity of this relationship is demonstrated by the remarkable fossil shown on the cover. Preserved in 15–20-million-year-old amber from the Dominican Republic, a worker stingless bee carries an orchid pollinarium. The pollinarium is the male reproductive structure, transferred as a single unit during pollination, complete with a mass of pollen grains. The find, surprisingly, is the first definitive fossil from the Orchidaceae family. The pollinarium's morphology locates the plant in the subtribe Goodyerinae. By combining this information with dates from related fossil plants, a new calibration of the molecular phylogeny of the Orchidaceae has been obtained. The analysis rejects the common assumption of a recent (Tertiary) origin for orchids, and suggests, instead, that they originated during the Late Cretaceous, 80 million years ago. The antiquity of the close relationship between orchids and their pollinators is demonstrated by a 15–20 million years old fossil of an orchid pollinarium attached to the business end of a stingless bee, preserved in amber from the Dominican Republic. This spectacular find is used to constrain the evolution of orchids, suggesting that the most recent common ancestor of orchids lived in the Late Cretaceous. 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. 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The antiquity of this relationship is demonstrated by the remarkable fossil shown on the cover. Preserved in 15–20-million-year-old amber from the Dominican Republic, a worker stingless bee carries an orchid pollinarium. The pollinarium is the male reproductive structure, transferred as a single unit during pollination, complete with a mass of pollen grains. The find, surprisingly, is the first definitive fossil from the Orchidaceae family. The pollinarium's morphology locates the plant in the subtribe Goodyerinae. By combining this information with dates from related fossil plants, a new calibration of the molecular phylogeny of the Orchidaceae has been obtained. The analysis rejects the common assumption of a recent (Tertiary) origin for orchids, and suggests, instead, that they originated during the Late Cretaceous, 80 million years ago. The antiquity of the close relationship between orchids and their pollinators is demonstrated by a 15–20 million years old fossil of an orchid pollinarium attached to the business end of a stingless bee, preserved in amber from the Dominican Republic. This spectacular find is used to constrain the evolution of orchids, suggesting that the most recent common ancestor of orchids lived in the Late Cretaceous. 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. 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itemscope itemtype="https://schema.org/ListItem"> <span itemprop="name">article</span><meta itemprop="position" content="3"></li> </ol> </div> </nav> </div> <div class="u-container u-mt-32 u-mb-32 u-clearfix" id="content" data-component="article-container" data-container-type="article"> <main class="c-article-main-column u-float-left js-main-column" data-track-component="article body"> <article lang="en"> <div class="c-article-header"> <header> <ul class="c-article-identifiers" data-test="article-identifier"> <li class="c-article-identifiers__item" data-test="article-category">Letter</li> <li class="c-article-identifiers__item">Published: <time datetime="2007-08-30">30 August 2007</time></li> </ul> <h1 class="c-article-title" data-test="article-title" data-article-title="">Dating the origin of the Orchidaceae from a fossil orchid with its pollinator</h1> <ul class="c-article-author-list c-article-author-list--short" data-test="authors-list" data-component-authors-activator="authors-list"><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Santiago_R_-Ram_rez-Aff1" data-author-popup="auth-Santiago_R_-Ram_rez-Aff1" data-author-search="Ramírez, Santiago R." data-corresp-id="c1">Santiago R. Ramírez<svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-mail-medium"></use></svg></a><sup class="u-js-hide"><a href="#Aff1">1</a></sup>, </li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Barbara-Gravendeel-Aff2" data-author-popup="auth-Barbara-Gravendeel-Aff2" data-author-search="Gravendeel, Barbara">Barbara Gravendeel</a><sup class="u-js-hide"><a href="#Aff2">2</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Rodrigo_B_-Singer-Aff3" data-author-popup="auth-Rodrigo_B_-Singer-Aff3" data-author-search="Singer, Rodrigo B.">Rodrigo B. Singer</a><sup class="u-js-hide"><a href="#Aff3">3</a></sup>, </li><li class="c-article-author-list__item c-article-author-list__item--hide-small-screen"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Charles_R_-Marshall-Aff1-Aff4" data-author-popup="auth-Charles_R_-Marshall-Aff1-Aff4" data-author-search="Marshall, Charles R.">Charles R. Marshall</a><sup class="u-js-hide"><a href="#Aff1">1</a>,<a href="#Aff4">4</a></sup> & </li><li class="c-article-author-list__show-more" aria-label="Show all 5 authors for this article" title="Show all 5 authors for this article">…</li><li class="c-article-author-list__item"><a data-test="author-name" data-track="click" data-track-action="open author" data-track-label="link" href="#auth-Naomi_E_-Pierce-Aff1" data-author-popup="auth-Naomi_E_-Pierce-Aff1" data-author-search="Pierce, Naomi E.">Naomi E. Pierce</a><sup class="u-js-hide"><a href="#Aff1">1</a></sup> </li></ul><button aria-expanded="false" class="c-article-author-list__button"><svg width="16" height="16" focusable="false" role="img" aria-hidden="true" class="u-icon"><use xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="#icon-eds-i-chevron-down-medium"></use></svg><span>Show authors</span></button> <p class="c-article-info-details" data-container-section="info"> <a data-test="journal-link" href="/" data-track="click" data-track-action="journal homepage" data-track-category="article body" data-track-label="link"><i data-test="journal-title">Nature</i></a> <b data-test="journal-volume"><span class="u-visually-hidden">volume</span> 448</b>, <span class="u-visually-hidden">pages </span>1042–1045 (<span data-test="article-publication-year">2007</span>)<a href="#citeas" class="c-article-info-details__cite-as u-hide-print" data-track="click" data-track-action="cite this article" data-track-label="link">Cite this article</a> </p> <div class="c-article-metrics-bar__wrapper u-clear-both"> <ul class="c-article-metrics-bar u-list-reset"> <li class=" c-article-metrics-bar__item" data-test="access-count"> <p class="c-article-metrics-bar__count">4494 <span class="c-article-metrics-bar__label">Accesses</span></p> </li> <li class="c-article-metrics-bar__item" data-test="citation-count"> <p class="c-article-metrics-bar__count">211 <span class="c-article-metrics-bar__label">Citations</span></p> </li> <li class="c-article-metrics-bar__item" data-test="altmetric-score"> <p class="c-article-metrics-bar__count">27 <span class="c-article-metrics-bar__label">Altmetric</span></p> </li> <li class="c-article-metrics-bar__item"> <p class="c-article-metrics-bar__details"><a href="/articles/nature06039/metrics" data-track="click" data-track-action="view metrics" data-track-label="link" rel="nofollow">Metrics <span class="u-visually-hidden">details</span></a></p> </li> </ul> </div> </header> </div> <div class="c-article-body"> <section aria-labelledby="Abs3" data-title="Abstract" lang="en"><div class="c-article-section" id="Abs3-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Abs3">Abstract</h2><div class="c-article-section__content" id="Abs3-content"><p>Since the time of Darwin<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 1" title="Darwin, C. On the Various Contrivances by which British and Foreign Orchids are Fertilised by Insects, and on the Good Effects of Intercrossing (J. Murray, London, 1862)" href="/articles/nature06039#ref-CR1" id="ref-link-section-d281937300e361">1</a></sup>, evolutionary biologists have been fascinated by the spectacular adaptations to insect pollination exhibited by orchids. However, despite being the most diverse plant family on Earth<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 2" title="Dressler, R. L. The Orchids: Natural History and Classification (Harvard Univ. Press, Cambridge, Massachusetts, 1981)" href="/articles/nature06039#ref-CR2" id="ref-link-section-d281937300e365">2</a></sup>, 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 <i>Meliorchis caribea</i> gen. et sp. nov.) attached to the mesoscutellum of an extinct stingless bee, <i>Proplebeia dominicana</i>, recovered from Miocene amber in the Dominican Republic, that is 15–20 million years (Myr) old<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 3" title="Iturralde-Vinent, M. E. & MacPhee, R. D. E. Age and paleogeography of Dominican amber. Science 273, 1850–1852 (1996)" href="/articles/nature06039#ref-CR3" id="ref-link-section-d281937300e375">3</a></sup>. This discovery constitutes both the first unambiguous fossil of Orchidaceae<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 4" title="Schmid, R. & Schmid, M. J. in Orchid Biology: Reviews and Perspectives Vol. 1 (ed. Arditti J.) 17–45 (Cornell Univ. Press, London, 1977)" href="/articles/nature06039#ref-CR4" id="ref-link-section-d281937300e380">4</a></sup> and an unprecedented direct fossil observation of a plant–pollinator interaction<sup><a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 5" title="Grimaldi, D. & Engel, M. S. Evolution of the Insects (Cambridge Univ. Press, New York, 2005)" href="/articles/nature06039#ref-CR5" id="ref-link-section-d281937300e384">5</a>,<a data-track="click" data-track-action="reference anchor" data-track-label="link" data-test="citation-ref" aria-label="Reference 6" title="Peñalver, E., Engel, M. S. & Grimaldi, D. Fig wasps in Dominican amber (Hymenoptera: Agaonidae). Am. Mus. Novit. 3541, 1–16 (2006)" href="/articles/nature06039#ref-CR6" id="ref-link-section-d281937300e387">6</a></sup>. By applying cladistic methods to a morphological character matrix, we resolve the phylogenetic position of <i>M. caribea</i> within the extant subtribe Goodyerinae (subfamily Orchidoideae). We use the ages of other fossil monocots and <i>M. caribea</i> 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.</p></div></div></section> <noscript> <div class="c-nature-box c-nature-box--side " data-component="entitlement-box"> <div class="js-access-button"> <a href="https://wayf.springernature.com?redirect_uri=https%3A%2F%2Fwww.nature.com%2Farticles%2Fnature06039" class="c-article__button" data-test="ra21"> <svg class="u-icon" width="18" height="18" aria-hidden="true" focusable="false"><use href="#icon-institution"></use></svg> <span class="c-article__button-text">Access through your institution</span> </a> </div> <div class="js-buy-button"> <a href="#access-options" class="c-article__button c-article__button--inverted" data-test="ra21"> <span>Buy or subscribe</span> </a> </div> </div> </noscript> <div class="js-context-bar-sticky-point-mobile" data-track-context="article body"> </div> <div class="c-article-access-provider u-mb-32 u-mt-0"> <p class="c-article-access-provider__text u-sans-serif">This is a preview 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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.</p><p><b>Author Contributions</b> 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.</p></div></div></section><section aria-labelledby="author-information" data-title="Author information"><div class="c-article-section" id="author-information-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="author-information">Author information</h2><div class="c-article-section__content" id="author-information-content"><h3 class="c-article__sub-heading" id="affiliations">Authors and Affiliations</h3><ol class="c-article-author-affiliation__list"><li id="Aff1"><p class="c-article-author-affiliation__address">Museum of Comparative Zoology, Harvard University, 26 Oxford St., Cambridge, Massachusetts 02138, USA, </p><p class="c-article-author-affiliation__authors-list">Santiago R. Ramírez, Charles R. Marshall & Naomi E. Pierce</p></li><li id="Aff2"><p class="c-article-author-affiliation__address">Nationaal Herbarium Nederland, Universiteit Leiden, P.O. Box 9514, Leiden, The Netherlands, </p><p class="c-article-author-affiliation__authors-list">Barbara Gravendeel</p></li><li id="Aff3"><p class="c-article-author-affiliation__address">Depto Botânica, Instituto de Biociências, Universidade Federal do Rio Grande do Sul, Av. Bento Gonçalves 9500, RS 91501-970, Porto Alegre, Brasil, </p><p class="c-article-author-affiliation__authors-list">Rodrigo B. Singer</p></li><li id="Aff4"><p class="c-article-author-affiliation__address">Department of Earth and Planetary Sciences, Harvard University, 20 Oxford St., Cambridge, Massachusetts 02138, USA, </p><p class="c-article-author-affiliation__authors-list">Charles R. Marshall</p></li></ol><div class="u-js-hide u-hide-print" data-test="author-info"><span class="c-article__sub-heading">Authors</span><ol class="c-article-authors-search u-list-reset"><li id="auth-Santiago_R_-Ram_rez-Aff1"><span class="c-article-authors-search__title u-h3 js-search-name">Santiago R. 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Pierce</span><div class="c-article-authors-search__list"><div class="c-article-authors-search__item c-article-authors-search__list-item--left"><a href="/search?author=Naomi%20E.%20Pierce" class="c-article-button" data-track="click" data-track-action="author link - publication" data-track-label="link" rel="nofollow">View author publications</a></div><div class="c-article-authors-search__item c-article-authors-search__list-item--right"><p class="search-in-title-js c-article-authors-search__text">You can also search for this author in <span class="c-article-identifiers"><a class="c-article-identifiers__item" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=search&term=Naomi%20E.%20Pierce" data-track="click" data-track-action="author link - pubmed" data-track-label="link" rel="nofollow">PubMed</a><span class="u-hide"> </span><a class="c-article-identifiers__item" href="http://scholar.google.co.uk/scholar?as_q=&num=10&btnG=Search+Scholar&as_epq=&as_oq=&as_eq=&as_occt=any&as_sauthors=%22Naomi%20E.%20Pierce%22&as_publication=&as_ylo=&as_yhi=&as_allsubj=all&hl=en" data-track="click" data-track-action="author link - scholar" data-track-label="link" rel="nofollow">Google Scholar</a></span></p></div></div></li></ol></div><h3 class="c-article__sub-heading" id="corresponding-author">Corresponding author</h3><p id="corresponding-author-list">Correspondence to <a id="corresp-c1" href="mailto:sramirez@oeb.harvard.edu">Santiago R. Ramírez</a>.</p></div></div></section><section data-title="Ethics declarations"><div class="c-article-section" id="ethics-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="ethics">Ethics declarations</h2><div class="c-article-section__content" id="ethics-content"> <h3 class="c-article__sub-heading">Competing interests</h3> <p>Reprints and permissions information is available at <a href="http://www.nature.com/reprints">www.nature.com/reprints</a>. The authors declare no competing financial interests.</p> </div></div></section><section data-title="Supplementary information"><div class="c-article-section" id="Sec3-section"><h2 class="c-article-section__title js-section-title js-c-reading-companion-sections-item" id="Sec3">Supplementary information</h2><div class="c-article-section__content" id="Sec3-content"><div data-test="supplementary-info"><div id="figshareContainer" class="c-article-figshare-container" data-test="figshare-container"></div><div class="c-article-supplementary__item" data-test="supp-item" id="MOESM31"><h3 class="c-article-supplementary__title u-h3"><a class="print-link" data-track="click" data-track-action="view supplementary info" data-test="supp-info-link" data-track-label="supplementary information" href="https://static-content.springer.com/esm/art%3A10.1038%2Fnature06039/MediaObjects/41586_2007_BFnature06039_MOESM31_ESM.pdf" data-supp-info-image="">Supplementary Information</a></h3><div class="c-article-supplementary__description" data-component="thumbnail-container"><p>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. 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Dating the origin of the Orchidaceae from a fossil orchid with its pollinator | Nature