taxonID	type	description	language	source
03F487DBFF8F246AFC59FF16FAC2C9B3.taxon	description	K. R. PEDERSEN gen. et sp. nov. P l a n t F o s s i l N a m e s R e g i s t r y N u m b e r. PFN 002665 (for new genus). E t y m o l o g y. From the village of Mira close to where the fossils were collected. D i a g n o s i s. Flowers pedicellate, structurally bisexual, actinomorphic, pentamerous and isomerous. No bracteoles present immediately under calyx. Sepals narrowly triangular, basally fused, persistent. Petals ovate, longer than sepals. Stamens opposite corolla lobes; staminodes alternating with the corolla lobes. Ovary semi-inferior, unilocular, of five carpels; single style long with a wide stylar canal that is five-angled in cross-section. Apical portion of ovary with a slightly raised nectariferous ring and stomata-like openings. Placentation free, central. Ovules numerous and densely spaced on the placenta, but not immersed. Ovules anatropous and bitegmic. Fruit a capsule. Seeds many, minute, angular, with reticulate surface. C o m m e n t s o n t h e g e n u s. The combination of distinctive floral features clearly places Miranthus in the Primulaceae s. l. and all features recorded for the fossil material are known also for the extant primuloid genus Samolus L. (subfamily Theophrastoideae). However, Miranthus also closely resembles flowers of other members of the subfamily, and because not all floral features of the fossil taxon are sufficiently well-known we refrain from assigning the fossils to an extant genus. To our knowledge there are no comparable flowers recorded from the fossil record. Three possible primuloid flowers from the Cainozoic Baltic amber, Berendtia primuloides GÖPP., B. rotata CONW. and Myrsinopsis succinea CONW. are fragmentary, and are preserved only as dispersed, sympetalous corollas with attached stamens (Göppert and Berendt 1845, Conwentz 1886).	en	Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard (2021): Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications. Fossil Imprint 77 (2): 214-230, DOI: 10.37520/fi.2021.016, URL: http://dx.doi.org/10.37520/fi.2021.016
03F487DBFF8F246FFC54FA39FA86C40F.taxon	description	H o l o t y p e. S 170155 (Mira sample 100; figured Textfigs 1 a, 4 c, d, 5 c, d). P a r a t y p e s. S 170106 (Mira sample 99), S 101263, S 101264, S 101266 – S 101268, S 153144 – S 153146, S 154535 – S 154538, S 170123, S 170125, S 266059 (Mira sample 100), S 100732 (Mira sample 105). P l a n t F o s s i l N a m e s R e g i s t r y N u m b e r. PFN 002666 (for new species). R e p o s i t o r y. Palaeobotanical Collections, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. From the elegant shape of the pedicellate flower. T y p e l o c a l i t y. Mira (40 ° 25 ʹ N, 08 ° 44 ʹ 15 ʺ W), about 25 km south of Aveiro, District of Coimbra, Portugal. T y p e s t r a t u m a n d a g e. “ Argilas de Vagos ”, Late Cretaceous (Campanian / Maastrichtian). D i a g n o s i s. As for the genus with the following addition. Epidermis of pedicel and outer (abaxial) epidermis of calyx composed of cells that each have a central pointed papilla. Trichomes scattered and thin. D i m e n s i o n s. Length of flower without pedicel and corolla: 0.8 – 1.0 mm; diameter: 0.7 – 1.0 mm. Length of flower bud with corolla and without pedicel: 1.5 mm. Description and comments on the s p e c i e s. The fossil flowers were briefly described and illustrated by Friis et al. (2010, 2011), but not formally named. More material has subsequently been discovered, including a small, informative flower bud. In most flowers the corolla and androecium have been shed (Text-figs 1 a – g, 2 a – d) and the flowers are probably fossilized at a post-anthetic stage. The flower bud has remains of corolla and stamens preserved (Text-fig. 3 a – f), but unfortunately, internal structures, including the ovary, ovules and parts of the androecium, are not well preserved. These internal structures were partially destroyed during fossilization and are only preserved on one side of the flower, which complicates interpretations of their organisation. The flower bud is assigned to the same species as the post-anthetic flowers based on features of the calyx. A second flower bud from the Mira mesofossil flora is described as a separate species, Miranthus kvacekii, based on epidermal differences from Miranthus elegans. The flowers are borne on a long, slender pedicel (Textfigs 1 a – d, 2 d, 3 a, b) and lack bracts or bracteoles immediately below the calyx. The pedicel is up to about 0.9 mm long (Text-fig. 1 d). All of the fossil flowers are isolated and there is no information on inflorescence structure, or how the pedicel may have been attached to the plant. The flowers are structurally bisexual, actinomorphic, pentamerous and isomerous, about 0.8 – 1.0 mm long, excluding the pedicel and corolla, and 0.7 – 1.0 mm in diameter. Including the corolla, but excluding the pedicel, the flower bud is about 1.5 mm long. The calyx is persistent and consists of five imbricate sepals in a single whorl that are fused at the base (Text-figs 1 a – g, 5 a, b). The sepals are narrowly triangular, with the free portion up to about 0.9 mm long, which tapers into a long tip (Text-fig. 1 a – g). Each sepal is supplied by three distinct vascular bundles, each of which typically appears as a hollow space in transverse sections (Text-figs 5 a, 6 a, b). In some specimens, minor vascular bundles are seen close to the sepal margins. The epidermis of the pedicel, and the abaxial (outer) epidermis of the calyx, is covered by a thick cuticle. The epidermal cells are small, almost equiaxial, thick-walled and typically with a pointed papilla that gives the surface a faint verrucate to spiny appearance. Similar epidermal features are also observed on the outer surface of the young ovaries (Text-figs 1 a – g, 2 f, 3 c – e, 5 a – d, 6 a). Very slender trichomes occur scattered on the outer surface of the sepals (Text-fig. 2 f). The epidermal cells of the adaxial (inner) epidermis of the calyx lobes are small, equiaxial, thin-walled and covered with a thin cuticle (Text-fig. 3 c – e). The calyx of the single flower bud is very similar in shape, size and epidermal features to the calyx post-anthetic flowers (Text-fig. 3 a, b) and the anatomy of all preserved organs is also identical. There is no trace of corolla and androecium in the supposed post-anthetic flowers. Corolla and androecium were most likely shed together after flowering, which would also be consistent with stamens that were fused to the corolla. The corolla is five-lobed, with imbricate lobes that are free for most of their length (Text-fig. 3 c – e). They appear united at the base, but the preservation does not allow a fully secure conclusion that they were sympetalous. The internal structures are preserved on one side of the flower bud, but based on the symmetry of the structures that remain, the androecium can be reconstructed as pentamerous and obhaplostemonous with five antepetalous stamens, which are seen near the base of the flower as stout filaments positioned in front of the petal lobes (Text-fig. 3 c – e) and near the bud apex as remains of thecae. Smaller structures, interpreted as staminodes, are seen alternating with the petals (Text-fig. 3 c – e). Both stamens and staminodes appear to be attached to the corolla near the base (Text-fig. 3 f), which is also consistent with the absence of stamens and stamen bases in the post-anthetic flowers. Normapolles-type pollen, which is very common on the surface of other Mira mesofossils, is also present on Miranthus flowers. However, in addition, two flowers have another kind of pollen on their surface. In both cases, the pollen is about 16 µm long and triaperturate with very long colpi that reach almost to the poles (Text-fig. 2 g). Unfortunately, the grains are folded and it cannot be established whether the grains are tricolpate or tricolporate. The tectum is foveolate. The ovary is semi-inferior and unilocular (Text-figs 2 d, 4 a – d, 5 a, b, 6 a, b, e). The apical portion of the ovary has a slightly protruding ring with stomata-like openings (Text-figs 1 c, 2 b, c). We think it likely that this zone was nectariferous and that the openings were nectar secreting. In some specimens there appear to be remains of septa in the apical part of the ovary, but these may be invaginations of the ovary wall and they do not reach the centre of the ovary (Text-fig. 5 a). The style is long, slender (Text-figs 1 e, 4 c), and distinctly five-angled in cross-section with a vascular bundle that extends for the length of the style in each of the angles (Text-fig. 6 c, d). The style is hollow and the stylar canal is wide along its full length (Text-fig. 6 d), except near the tip where it is closed (Text-fig. 6 c). In some specimens, the style is broken, but the specimens available give no indication of different style lengths and the flower is interpreted as homostylous. Placentation is free and central, with a dome-shaped, almost globose placenta, alternating with the corolla lobes; note verrucate abaxial surface of calyx lobes (ca) and ovary wall (ow) partly distorted and displaced to one side. f: Longitudinal section (orthoslice yz 0567) of flower bud showing ovary wall (ow) and insertion of calyx (ca), corolla (co) and stamens (st) on a hypanthium rim. Specimen, Mira 100 - S 266059 (a – f). Scale bars = 600 µm (a, b), 300 µm (c – f). borne on a central column (Text-figs 4 a – d, 5 a – b, 6 a, b, e). Numerous ovules are densely spaced on the placenta, but not immersed in its surface (Text-figs 2 e, 4 a – d, 5 a – d, 6 b, e). Ovules are bitegmic and anatropous, about 0.14 mm long, with a reticulate seed surface (Text-fig. 2 e). Among the post-anthetic flowers, none has a fully developed fruit. However, some are clearly five-parted, have five valves and are interpreted as young capsules (Text-fig. 1 e, f). In one specimen the ovary is split open along one of the valves (Text-fig. 2 d). The epidermal cells of the young capsules are small, equiaxial, and often with tiny papillate projections. Stomata occur scattered over the surface of the calyx lobes and pedicel together with smaller openings that are probably trichome bases. Larger openings are irregularly scattered and are interpreted as schizogenous secretory cavities that are sometimes burst (Text-figs 1 b, c, 4 b).	en	Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard (2021): Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications. Fossil Imprint 77 (2): 214-230, DOI: 10.37520/fi.2021.016, URL: http://dx.doi.org/10.37520/fi.2021.016
03F487DBFF852461FF2AF9F5FBCFCA62.taxon	description	H o l o t y p e. S 170157 (Mira sample 100; figured Textfig. 7 a – e). P l a n t F o s s i l N a m e s R e g i s t r y N u m b e r. PFN 002673 (for new species). R e p o s i t o r y. Palaeobotanical Collections, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. In commemoration of Zlatko Kvaček and his outstanding contributions to Cretaceous and Cainozoic angiosperm palaeobotany. T y p e l o c a l i t y. Mira (40 ° 25 ʹ N; 08 ° 44 ʹ 15 ʺ W), about 25 km south of Aveiro, District of Coimbra, Portugal. T y p e s t r a t u m a n d a g e. “ Argilas de Vagos ”, Late Cretaceous (Campanian / Maastrichtian). D i a g n o s i s. As for the genus with the following addition: Epidermis of pedicel and outer (abaxial) epidermis of calyx with closely spaced, robust, stiff trichomes. D i m e n s i o n s. Length of flower excluding pedicel and corolla: 0.8 – 1.0 mm; diameter: 0.7 – 1.0 mm. Length of flower bud with corolla, but excluding the pedicel: 1.5 mm. D e s c r i p t i o n a n d c o m m e n t s o n t h e s p e c i e s. The species is based on a single flower bud discovered with Miranthus elegans in the Mira mesofossil flora (Text-fig. 7 a – e). The individual floral parts are strongly appressed and somewhat distorted, which impedes a full understanding of the floral structure and organization (Textfig. 7 b – e). However, the androecium clearly consists of five stamens. The stamens also apparently alternate with the sepals and are obhaplostemonous (Text-fig. 7 d, e). The stamen filaments appear to be fused to the corolla for part of their length, while the anthers are free from the corolla and from each other. Alternating with the filaments are filament-like structures. These are not as distinct as the stamen filaments and interpreted as staminodes (Text-fig. 7 e). The flower bud is similar to Miranthus elegans in its characteristic free, dome-shaped placenta (Text-fig. 7 b, c) and semi-inferior ovary (Text-fig. 7 b, c), but differs in having short, densely spaced trichomes on the outer surface of the calyx lobes, particularly along the margins (Text-fig. 7 a). Although scattered trichomes and trichome basis are observed in Miranthus elegans, the trichomes are much finer and less densely spaced.	en	Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard (2021): Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications. Fossil Imprint 77 (2): 214-230, DOI: 10.37520/fi.2021.016, URL: http://dx.doi.org/10.37520/fi.2021.016
03F487DBFF862465FF98F8BFFF1FCDAD.taxon	description	Extensive studies on extant primuloids by Anderberg and colleagues (e. g., Anderberg and Ståhl 1995, Anderberg et al. 1998 a, b, 2000 a, b, 2007, Källersjö et al. 2000, Källersjö and Ståhl 2003, Wanntorp and Anderberg 2011), as well as by other researchers (e. g., Caris et al. 2000, Ma and Saunders 2003, Caris and Smets 2004, Morozowska et al. 2020), have been pivotal for our comparative studies and for recognizing the affinities of the fossil flowers. Relationships among the main primuloid lineages are well supported based on phylogenetic analyses of both morphological and molecular data. The monogeneric Maesoideae is resolved as the sister lineage to the remaining taxa, with Theophrastoideae resolved as sister to a well-supported Primuloideae- Myrsinoideae clade (Källersjö et al. 2000, Bremer et al. 2002, Schönenberger et al. 2005). Within Theophrastoideae, the monogeneric Samoleae are sister to all other members of the subfamily. Shared features for Miranthus and the Primulaceae include the pentamerous and isomerous organisation of the flowers, persistent calyx, haplostemonous androecium and the highly characteristic free central and dome-shaped placenta. Among Ericales, a free central placenta is known only for the Primulaceae and is a clear synapomorphy for the group (Schönenberger et al. 2005). Few floral features distinguish the various primuloid lineages and a separation of the lineages based on floral morphology is only poorly supported. As noted by Caris and Smets (2004) “ it seems that no unambiguous morphological synapomorphies can be found that justify a position of Samolus in either Primulaceae or Theophrastaceae ”, but a position as sister to Theophrastoideae is well supported by molecular data (Källersjö and Ståhl 2003). Nevertheless, the semi-inferior ovary of Miranthus flowers, and the presence of staminodes alternating with the corolla lobes, are important distinctive floral features. Furthermore, among extant primuloids, a semi-inferior ovary is known only for Maesa FORSSK. and Samolus L., and staminodes are present only in Maesa, the genera of Theophrastoideae (including Samolus), and Soldanella L. of the Primuloideae (Anderberg and Ståhl 1995, Anderberg et al. 1998 a). Anectariferous zone on the upper part of the ovary, referred to as “ nectarostomata on the flanks of the ovary ” by Caris and Smets (2004), is a further feature shared with Maesa and Samolus that is not present in other primuloid taxa (Caris et al. 2000, Caris and Smets 2004). Miranthus thus appears to be nested securely in the primuloid clade.	en	Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard (2021): Early Flowers Of Primuloid Ericales From The Late Cretaceous Of Portugal And Their Ecological And Phytogeographic Implications. Fossil Imprint 77 (2): 214-230, DOI: 10.37520/fi.2021.016, URL: http://dx.doi.org/10.37520/fi.2021.016
