The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms Friis, Else Marie Crane, Peter R. Pedersen, Kaj Raunsgaard Mendes, Mário Miguel Kvaček, Jiří Fossil Imprint 2022 2022-12-20 78 2 341 424 E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVACEK E. M. FRIIS, P. R. CRANE, K. R. PEDERSEN, M. M. MENDES et J. KVACEK 2022 Magnoliopsida Chloranthaceae Canrightia GBIF Plantae Chloranthales 6 347 Tracheophyta species foveolata sp. nov.   Text-figs 3a–f, 4a–i  Holotype. S174249 (Catefica sample 49; figured Text-fig. 3a–f).  Plant Fossil Names Registry Number. PFN002785 (for new species). Paratypes. S175179, S265998, S266057, S266107 (Catefica sample 49), S266042 (Catefica sample 154), S175178 (Catefica sample 242).   Repository. Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden.  Etymology. From Latin: fovea (pit) referring to the densely pitted surface of the endotesta.   Type locality. Catefica ( 39° 03ʹ 30ʺ N; 09°14ʹ 30ʺ W), between the villages of Cateficaand Mugideira, about 4 kmsouth of Torres Vedras, Portugal. Type stratum and age. Almargem Formation, Early Cretaceous (Aptian-early Albian).  D i a g n o s i s. Fruit obovoid with a broad hypanthium and two pendent seeds. Perianth of six tepals. Contact surface between the two seeds flat; external surface rounded. Crystals evenly distributed in the cells of the endotesta. Surface of endotesta foveolate with shallow foveae arranged in more than 30 closely packed longitudinal rows. Fruit wall particularly thick apically over the seeds. Distinguishing features. The new species is assigned to the extinct genus  Canrightiabased on the berry-like fruit with pendent, orthotropous seeds that have an endotestal-endotegmic seed coat and a crystalliferous endotesta.  Canrightia foveolatais distinguished from  C. resinifera(see above), and from  C. elongatafrom the Torres Vedras mesofossil flora ( Friis et al. 2019a), mainly by the densely pitted and grooved surface of the endotesta. Seeds of  C. foveolataalso have crystals that are of more or less of similar size and that are evenly distributed in the endotestal cells, whereas in  C. resiniferaand  C. elongatalarger crystals are concentrated close to the outer surface of endotesta.  Canrightia foveolatais also two-seeded, as are most specimens of  C. resiniferafrom the Famalicão locality, while fruits of  C. resiniferafrom the Catefica locality typically have three to five seeds and  C. elongatahas three seeds.   Canrightia foveolatais further distinguished from the two other species of  Canrightiaby the well-developed soft tissue of the fruit wall above the seeds.  Canrightia foveolatamay also be distinguished from the two other species by the larger number of perianth parts, but as the perianth is known for only one specimenof  C. foveolata, and only a few specimens of  C. resinifera, the range of tepal numbers in  Canrightiais not fully established. A pitted surface of the endotesta is also present in seeds of  CanrightiopsisE.M.FRIIS, G.W.GRIMM, M.M.MENDES et K.R.PEDERSEN and KvacekispermumE.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, two other extinct genera of chloranthoid affinity ( Friis et al. 2015a, 2018b), but both of these genera have one-seeded fruits and a much thicker endotestal seed coat.  Dimensions. Length of fruit: 1.7 mm; maximum width of fruit: 1.6 mm; length of seed: 0.85–1.05 mm; maximum width of seed: 0.6–0.9 mm.  Description and remarks. The new species is based on a single fruit, containing two seeds ( Text-fig. 3a–f). There are also several isolated seeds ( Text-fig. 4a–i). The fruit and two of the isolated seeds were studied using SRXTM. The fruit is partly abraded, and although the stigmatic region is missing, the fruit is otherwise well preserved in its apical part. There is a swollen rim about halfway up the fruit with six, small poorly developed tepals that are best preserved on one side of the fruit ( Text-fig. 3a). Five vascular bundles are preserved in the hypanthium, each extending to a tepal and their symmetry indicates that a sixth bundle has been lost where the fruit wall is abraded ( Text-fig. 3d). The fruit wall is particularly thick in the region above the seeds and consists mainly of isodiametric cells ( Text-fig. 3a–c). The seeds are broadly elliptical, crescent-shaped in lateral view, slightly pointed at the micropylar end and rounded at the chalazal end ( Text-figs 3b, 4a–d, f). Where the two seeds meet, their faces are flattened, but with a prominent chalaza that projects towards the face where the seeds meet (Textfig. 4b, d). The opposite faces are rounded ( Text-figs 3c, d, 4b, c, e). In the isolated seeds, the outer cells of the seed coat are abraded exposing the surface of the endotesta, which is characterized by numerous small pits arranged in more than 30 shallow, closely-spaced, longitudinal grooves ( Text-fig. 4a–d). In the fruit the exotesta of the seeds is partly preserved and consists of thick-walled, isodiametric cells. The endotesta is thin (about 30 µm) in the region between the two seeds, but thicker (about 55 µm) in the chalazal region and toward the outer surfaces ( Text-fig. 3c, e, f). The exotesta is so tightly appressed to the tissue of the fruit wall that the two tissues are sometimes difficult to delimit. The endotesta consists of palisade-shaped cells that are infilled with fibrous material in which there are abundant casts of cubic crystals. The casts of these crystals are distributed more or less evenly within the cells ( Text-figs 3f, 4e–i).  Text-fig. 3. Synchrotron radiation X-ray tomographic microscopy (SRXTM) images of fruits of  Canrightia foveolata sp. nov.; Catefica locality, Portugal. a) Volume rendering of fruit showing prominent rim around the middle of the fruit with reduced tepals (arrowheads) and partly abraded fruit wall exposing the pitted endotesta surface of one of two seeds (arrow); note two of the vascular bundles (vb) extending from the base of the fruit to the tepals; b) Voltex of fruit showing prominent rim around the fruit (arrowhead) and dense precipitation of crystals in the endothelium cells of one of the two seeds in the fruit; c) Longitudinal section of fruit (orthoslice yz0520) showing the inferred hypanthium rim (arrow head) and two seeds, one with a dense precipitation of crystals; note the prominent endothelium cells (asterisks) of the inner integument and the well-developed fruit wall above the seeds; d) Transverse section through basal part of fruit and seeds close to the micropyle (orthoslice xy0312) showing partly abraded fruit wall with five vascular bundles (vb) and details of the seed coat with endotesta (oi-end) surrounding the tegmen consisting of an outer epidermis (ii-o), middle layer (ii-m) and a distinct inner epidermis (endothelium) consisting of radially elongated cells (asterisk); e) Transverse section (orthoslice xy1680) through apical part of the fruit close to chalaza showing the tips of two seeds; note the endotesta (oi-end) surrounded by thick-walled cells of the exotesta (oi-o); f) Transverse section (orthoslice xy1485) through fruit in the region of the hypanthium rim showing sections through the two seeds close to the chalazal region; note endotesta (oi-end) surrounded by larger cells of exotesta (oi-o) and fruit wall (fr). Specimen, Catefica 49-S174249 ( holotype, a–f). Scale bars = 300 Μm (a–c, e, f), 100 Μm (d). The inner integument is three cell layers thick. It consists of an outer epidermis, a middle layer of thick-walled and slightly longitudinally elongated cells, and an inner epidermis that develops into an endothelium of thin-walled and radially elongated cells ( Text-figs 3c, d, 4e, f, h). The stigmatic area is not preserved and no pollen was observed on the surface of the fruit.  Affinity and other occurrences. The relationships of  Canrightia foveolata, as for  Canrightia resinifera, arelikelyclosetothebaseofextantChloranthaceae (see above).  Canrightia foveolatais currently known only from the Catefica locality. 4013209307 R Sweden Swedish Museum of Natural History Palaeobotanical Collections Department of Palaeobiology 6 347 1 Stockholm 4013209305 Portugal Torres Vedras 39.058334 Catefica 19 -9.241667 Mugideira 6 347 1