Grana ISSN: 0017-3134 (Print) 1651-2049 (Online) Journal homepage: http://www.tandfonline.com/loi/sgra20 Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites-type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae Else Marie Friis, Guido W. Grimm, Mário Miguel Mendes & Kaj Raunsgaard Pedersen To cite this article: Else Marie Friis, Guido W. Grimm, Mário Miguel Mendes & Kaj Raunsgaard Pedersen (2015) Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites-type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae, Grana, 54:3, 184-212, DOI: 10.1080/00173134.2015.1060750 To link to this article: http://dx.doi.org/10.1080/00173134.2015.1060750 © 2015 The Author(s). Published by Taylor & View supplementary material Francis. Published online: 15 Jul 2015. Submit your article to this journal Article views: 298 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=sgra20 Download by: [Statsbiblioteket Tidsskriftafdeling] Date: 18 January 2016, At: 04:18 Grana, 2015 Vol. 54, No. 3, 184–212, http://dx.doi.org/10.1080/00173134.2015.1060750 Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites- type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae ELSE MARIE FRIIS1, GUIDO W. GRIMM2, MÁRIO MIGUEL MENDES3 & KAJ RAUNSGAARD PEDERSEN4 1Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden, 2Department of Palaeontology, University of Vienna, Vienna, Austria, 3CIMA – Centre for Marine and Environmental Research, University of Algarve, Faro, Portugal, 4Department of Geoscience, University of Aarhus, Aarhus, Denmark Abstract Canrightiopsis with three species (C. intermedia, C. crassitesta, C. dinisii) is described from the Early Cretaceous of Portugal based on small, one-seeded berries. The fruits are derived from bisexual flowers with three stamens borne on one side of the ovary. There are no traces of a perianth. Pollen is of the Clavatipollenites-type, monocolpate, semitectate, reticulate- columellate with heterobrochate reticulum and muri with beaded supratectal ornamentation. The ovary is unilocular with a single pendant, orthotropous and bitegmic ovule. The seed is endotestal. The endotesta consists of one layer of palisade- shaped crystal cells with fibrous infillings. The fruit wall has resin bodies or cavities from presumed ethereal oil cells sometimes seen as stomata-like structures on the fruit surface. A phylogenetic analysis resolves Canrightiopsis as a close relative of extant Chloranthaceae, particularly close to extant Chloranthus and Sarcandra. All three taxa share the one-sided position of the stamens on the ovary. An evolutionary sequence from fossil Canrightia to fossil Canrightiopsis and extant Chloranthus and Sarcandra is suggested by loss of perianth, reduction in number of ovules and stamens and displacement of stamens to one side of the ovary. Canrightiopsis also shares several critical features with extant Ascarina including mono- aperturate pollen and beaded supratectal ornamentation of the pollen wall. Keywords: Canrightia, Chloranthaceae, Clavatipollenites, Early Cretaceous, endothelium, fossil flowers, synchrotron radiation X-ray tomographic microscopy (SRXTM) The Chloranthaceae are a small family usually 1960; Walker & Walker 1984)andAsteropollis placed close to the base of the angiosperm phyloge- R.W.Hedl. et G.Norris (compared to pollen of Hedyos- netic tree (APG 2009). The family includes four mum;Walker&Walker1984) occur very early in the extant genera, Ascarina J.R.Forst. et G.Forst., Chlor- history of angiosperms with records extending back to Downloaded by [Statsbiblioteket Tidsskriftafdeling] at 04:18 18 January 2016 anthus Sw., Hedyosmum Sw. and Sarcandra Gardner the Barremian (e.g. Couper 1958;Hedlund&Norris that today occur in warm temperate to subtropical 1968; Walker & Walker 1984;Hughes1994; Nichols and tropical regions (Endress 1987). The family has et al. 2006; Friis et al. 2011; Martínez et al. 2013). In a long geological history with an almost worldwide addition to numerous dispersed pollen grains, the fos- distribution in the Cretaceous, and the present dis- sil record of Chloranthaceae also includes a number of tribution and diversity of the family are clearly relic- Cretaceous floral structures. From the Early Creta- tual. Dispersed pollen grains of Clavatipollenites ceous Hedyosmum-like pistillate and staminate struc- Couper (compared to pollen of Ascarina;Couper tures with Asteropollis-type pollen attached to the ovary Correspondence: Else Marie Friis, Department of Palaeobiology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden. E-mail: [email protected] (Received 8 March 2015; accepted 19 May 2015) © 2015 The Author(s). Published by Taylor & Francis. This is an Open Access article distributed under the terms of the Creative Commons Attribution-NonCommercial-NoDerivatives License (http://creative commons.org/licenses/by-nc-nd/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited, and is not altered, transformed, or built upon in any way. Canrightiopsis, a new Early Cretaceous fossil 185 or in situ in the stamens have been identified from ovule per ovary as in extant Chloranthaceae. Portugal and eastern North America (Friis et al. Further, Canrightiopsis is similar to Chloranthus and 1997, 2011). From the Late Cretaceous of Scania Sarcandra in having the stamens borne in a partly and eastern North America, inflorescences, isolated epigynous position. Particularly the presence of flowers, androecia and fruits closely resembling those three separate stamen scars in Canightiopsis suggests of extant Chloranthus were assigned to three species a derivation of the tripartite androecium in Chlor- of the extinct genus Chloranthistemon P.R.Crane, anthus from three stamens, not from one. Monocol- E.M.Friis et K.R.Pedersen (Crane et al. 1989;Here- pate, reticulate Clavatipollenites-type pollen similar ndeen et al. 1993;Eklundetal.1997). Chloranthiste- particularly to pollen of extant Ascarina was obser- mon-like floral structures are also known from the mid- ved on the stigmatic surfaces or on the ovary wall of Cretaceous of Germany (Friis et al. 2011; Hartkopf- all three Canrightiopsis species and corroborates pre- Fröder et al. 2011). vious suggestions of a chloranthaceous affinity for Floral structures in all four extant genera of Chlor- some of the early dispersed angiosperm pollen. anthaceae are simple (Endress 1987). Never- theless, the interpretation of certain floral features and basic organisation has not been straightforward. Material and methods fl This to some extent may re ect extensive extinction Material within the lineage and missing data from the geo- logical record. Particularly the nature of the unusual The fossil material reported here was collected from tripartite androecium of Chloranthus has been much several localities (Arazede, Buarcos, Catefica, Juncal- debated (Swamy 1953; Endress 1987). In Chloran- Chicalhão, Famalicão, Vale de Água, Vila Verde) in thus, the androecium is typically three-lobed with a the Lusitanian Basin, western Portugal (for informa- median, tetrasporangiate lobe and two lateral, bis- tion on the mesofossil floras see Friis et al. 2011; porangiate lobes (Endress 1987). Endress (1987) Mendes et al. 2014a). offered two hypotheses for the evolutionary origin Based on stratigraphic information for the Lusi- of the androecium, both starting from a single sta- tanian Basin (e.g. Dinis et al. 2008), the plant bear- men with the three lobes formed either by polymer- ing sediments at the Chicalhão site (Juncal) and the isation, concrescence and reduction, or by lobation Vale de Água locality studied here belong to the and fractionation of a single stamen. basal part of the Famalicão Member of the Figueira New fossil fruits from the Early Cretaceous meso- da Foz Formation established by Dinis (2001). fossil floras of Portugal may fill in some of the miss- Buarcos, Vila Verde and Arazede belong to the ing data critical for understanding the evolutionary basal part of the Calvaria Member of the Figueira steps resulting in the tripartite androecium of extant da Foz Formation established by Dinis (2001). The Chloranthus and fossil Chloranthistemon as well as age of the Famalicão and Calvaria members was other floral features in extant Chloranthaceae. Can- indicated as late Aptian–early Albian by Rey et al. rightia resinifera E.M.Friis et K.R.Pedersen com- (2006). The mesofossils from the Famalicão locality prises small berries with two to five seeds. The were collected in a stratigraphic position below the fruits are derived from bisexual flowers with a partly Calvaria Member indicating a late Aptian age (or epigynous, strongly reduced perianth and four or five older) for this flora. According to Jacques Rey (per- stamens in a radial arrangement on the rim of the sonal communication, June 2012), the plant bearing hypanthium. The seeds are orthotropous, bitegmic sediments of the Catefica locality may belong to the Downloaded by [Statsbiblioteket Tidsskriftafdeling] at 04:18 18 January 2016 and endotestal very similar to those of Ascarina, younger part of the Almargem Formation that is of Chloranthus and Sarcandra. There are, however, sev- late Aptian–early Albian age. However, the Catefica eral features in Canrightia E.M.Friis