Search

Canrightiopsis sp. R e m a r k s. The three species of Canrightiopsis described from the Early Cretaceous of Portugal are closely similar to each other in morphology and in the organization of the fruits and seeds. The three species are mainly distinguished by details of their seed coat (Friis et al. 2015a). The Catefica mesofossil flora includes several Canrightiopsis fruits that cannot be placed securely in any of the existing species due to the lack of information on their internal structure. These seeds are collectively referred to here as Canrightiopsis sp., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 351, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Grimm, G. W., Mendes, M. M., Pedersen, K. R. (2015 a): Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites - type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae. - Grana, 54: 184 - 212. https: // doi. org / 10.1080 / 00173134.2015.1060750"]}

Genus Elasmostemon E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK gen. nov. T y p e. Elasmostemon paisii E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK 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. PFN0002792 (for new genus). E t y m o l o g y. From Greek elasma for lamina and stemon for stamen. G e n e r i c d i a g n o s i s. Stamen laminar. Anthers tetrasporangiate, dithecate. Thecae positioned close to the stamen margin and separated by a massive, broad connective. Stamen apex rounded without an apical extension. Dehiscence longitudinal. Staminal tissue with larger cells, probably ethereal oil cells. Pollen monocolpate, circular in equatorial view, semitectate-reticulate. Reticulum loosely attached, heterobrochate. Aperture long, reaching the equator, with distinct margins. Columellae short. C o m m e n t s o n t h e g e n u s. The broad and flattened stamen and monoaperturate pollen strongly suggest a position among basal grade angiosperms. Laminar or laminar-like stamens that have pollen sacs positioned at, or close to, their margin occur among some extant ANA-grade angiosperms (Austrobaileya C.T.WHITE and Nymphaeaceae) and Magnoliales, and ethereal oil cells like those seen in the fossil stamens are also present in stamens of Austrobaileya and Magnoliales (e.g., Canright 1952, Endress and Hufford 1989). Stamens of Austrobaileya are flattened as in the Catefica fossils and Austrobaileya pollen is also reticulatecolumellate. However, in Austrobaileya the two thecae are close together on either side of the mid-line of the stamen with little connective tissue between them (Canright 1952, Endress and Hufford 1989). Pollen of Austrobaileya is also much larger, between 51 and 100 µm, than the pollen of the Catefica material (Halbritter 2016). Stamens of Nymphaeaceae lack oil cells and the pollen is non-reticulate and often zona-aperturate (see discussion of “Stamen with zona-aperturate pollen” above). In Magnoliales stamens of Degeneria I.W.BAILEY et A.C.SM. (Degeneriaceae) and Galbulimima F.M.BAILEY (Himantandraceae) are broad as in the Catefica stamen, but pollen in Degeneria is psilate with granular infratectal layer and pollen in Galbulimima is atectate (Endress and Hufford 1989). Eupomatia (Eupomatiaceae) also has laminar-like stamens, but the pollen is zona-aperturate. In Magnoliaceae, the thecae are laminar or marginal and pollen is typically psilate. Semitectate-reticulate pollen is known among other Magnoliales, including species of Horsfieldia WILLD., Myristicaceae (Sauquet and Le Thomas 2003) and several Annonaceae (Walker 1971), but stamens in these two families are not laminar., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 375, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Canright, J. (1952): The comparative morphology and relationships of the Magnoliaceae. I. Trends of specialization in the stamens. - American Journal of Botany, 39: 484 - 497. https: // doi. org / 10.1002 / j. 1537 - 2197.1952. tb 13058. x","Endress, P. K., Hufford, L. D. (1989): The diversity of stamen structures and dehiscence patterns among Magnoliidae. - Botanical Journal of the Linnean Society, 100: 45 - 85. https: // doi. org / 10.1111 / j. 1095 - 8339.1989. tb 01709. x","Halbritter, H. (2016): Austrobaileya scandens. - PalDat - A palynological database. [accessed May 23, 2021] https: // www. paldat. org / pub / Austrobaileya _ scandens /","Sauquet, H., Le Thomas, A. (2003): Pollen diversity and evolution in Myristicaceae (Magnoliales). - International Journal of Plant Sciences, 164: 613 - 628. https: // doi. org / 10.1086 / 375424","Walker, J. W. (1971): Pollen morphology, phytogeography, and phylogeny of the Annonaceae. - Contributions from the Gray Herbarium of Harvard University, 202: 3 - 130. https: // doi. org / 10.5962 / p. 272704"]}

Pennicarpus tenuis E.M.FRIIS, K.R.PEDERSEN et P.R.CRANE, 2000 D e s c r i p t i o n a n d r e m a r k s. Two fruits, about 0.8–1.3 mm long and 0.5 mm broad, were recovered from Catefica sample 50. The fruits (not figured) are strongly flattened and elliptical in outline with a very thin fruit wall and thin, longitudinal ridges, probably from vascular bundles, that extend for the full length of the fruits. A f f i n i t y a n d o t h e r o c c u r r e n c e s.The fruits are closely similar in size, shape and texture to those of Pennicarpus tenuis described from the Vale de Água and Buarcos mesofossil floras (Friis et al. 2000). The fruits from Catefica have not been studied using SEM and it is unknown whether they have adhering pollen of Pennipollis E.M.FRIIS, K.R.PEDERSEN et P.R.CRANE as is known for Pennicarpus tenuis from Vale de Água and Buarcos. Pennicarpus and the associated Pennistemon E.M.FRIIS, K.R.PEDERSEN et P.R.CRANE and Pennipollis, collectively referred to as the Pennipollis plant (Friis et al. 2011), were placed in the monocots mainly based on the distinct acolumellate pollen wall (Friis et al. 2000), although an affinity with Chloranthaceae has also been suggested (see Doyle and Endress 2014). So far, only two fruits have been recovered from Catefica and Pennipollis grains have not been recognized in the palynological preparations. In the Vale de Água and Buarcos mesofossil floras remains of the Pennistemon plant are abundant. Dispersed Pennipollis pollen has also been reported in early Aptian to middle Albian palynological assemblages from coastal sections in Portugal (Heimhofer et al. 2007) and from the dispersed palynoflora of Casal do Borracho (Torres Vedras) (Mendes et al. 2018a). Pennipollis pollen is also widespread in Early Cretaceous palynofloras from other regions (see Friis et al. 2000)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 383, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Pedersen, K. R., Crane, P. R. (2000): Fossil floral structures of a basal angiosperm with monocolpate, reticulate-acolumellate pollen from the Early Cretaceous of Portugal. - Grana, 39: 226 - 245. https: // doi. org / 10.1080 / 00173130052017262","Doyle, J. A., Endress, P. K. (2014): Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: ANITA lines and relatives of Chloranthaceae. - International Journal of Plant Sciences, 175: 555 - 600. https: // doi. org / 10.1086 / 675935","Heimhofer, U., Hochuli, P. A., Burla, S., Weissert, H. (2007): New records of Early Cretaceous angiosperm pollen from Portuguese coastal deposits: Implications for the timing of the early angiosperm radiation. - Review of Palaeobotany and Palynology, 144: 39 - 76. https: // doi. org / 10.1016 / j. revpalbo. 2005.09.006","Mendes, M. M., Barron, E., Dinis, P., Rey, J., Batten, D. J. (2018 a): A new palynoflora from upper Barremian-lower Aptian rocks at Casal do Borracho, Torres Vedras, western Portugal, and its palaeoecological significance. - Cretaceous Research, 90: 363 - 374. https: // doi. org / 10.1016 / j. cretres. 2018.06.012"]}

Appomattoxia sp. Text-fig. 20a–d D e s c r i p t i o n a n d r e m a r k s. The material comprises three strongly compressed and lignitised fruits. SRXTM of one of the specimens did not provide any information on internal structure and there is no information on how the fruits were attached to the plant. The fruits are minute, elliptical to ovate in outline, 0.5–0.6 mm long and 0.3–0.35 mm wide (Text-fig. 20a, b). The fruit wall is thin with its surface covered by a thick cuticle bearing densely arranged, short, sometimes hooked trichomes, each with a broad base (Text-fig. 20c, d). The stigmatic area is indistinct, sessile and lacks trichomes. No pollen grains have been observed on the stigmatic region or on the fruit surface. A f f i n i t y a n d o t h e r o c c u r r e n c e s. The fossils are similar to fruits assigned to the fossil genus Appomattoxia in general shape, nature of the stigmatic region, thin fruit wall and the prominent trichomes that are sometimes coiled. The type species, A. ancistrophora E.M.FRIIS, K.R.PEDERSEN et P.R.CRANE from the Early Cretaceous mesofossil flora from Puddledock, Virginia, USA, has much larger fruits with trichomes that are longer and more distinctly and more regularly hooked (Friis et al. 1995). The fruits from Catefica are more similar to fruits of Appomattoxia minuta E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN from the Torres Vedras mesofossil flora (Friis et al. 2019a), but the trichomes are shorter, less prominently hooked and more densely arranged. Pollen grains associated with Appomattoxia ancistrophora and A. minuta are identical to pollen of Goczania E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN. No pollen was found attached to the Appomattoxia fruits from Catefica, but Goczania stamens and pollen occur with the fruits in the Catefica mesofossil flora (see below). A piperalean affinity is inferred for Appomattoxia based on the combined pollen, fruit and seed characters (Friis et al. 1995, for further discussion see also Friis et al. 2019a)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 368, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Pedersen, K. R., Crane, P. R. (1995): Appomattoxia ancistrophora gen. et sp. nov., a new Early Cretaceous plant with similarities to Circaeaster and extant Magnoliidae. - American Journal of Botany, 82: 933 - 943. https: // doi. org / 10.1002 / j. 1537 - 2197.1995. tb 15710. x","Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013"]}

Anacostia sp. D e s c r i p t i o n a n d r e m a r k s. Three specimens (S266205, S266208, S266218) assignable to Anacostia (not figured) were recovered from the Catefica mesofossil flora. Cells of seed and fruit surface are distinctive for Anacostia, but the preservation does not allow a species level assignment. A f f i n i t y a n d o t h e r o c c u r r e n c e s. Anacostia was first described based on fruits, seeds and associated pollen from the Early Cretaceous of Maryland and Virginia, USA (Anacostia marylandensis E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, A. virginiensis E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN) and from Portugal (Anacostia portugallica E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, Anacostia teixeirae E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN) (Friis et al. 1997). The genus is characterized by its one-seed fruits and exotestal seeds that have a crystalliferous seed coat and the inner layer of testa with strongly undulate walls. The four species also share the regular occurrence of trichotomocolpate, and occasionally monocolpate, pollen on the stigma and fruit surface. The monocolpate pollen indicates a relationship to non-eudicot angiosperms and the presence of an embryo with two cotyledons allows a monocot affinity to be rejected (Friis et al. 2015b). A possible affinity with Austrobaileyales was suggested by Friis et al. (1997) and has also been inferred based on several phylogenetic analyses by Doyle and Endress (e.g., Doyle and Endress 2014). However, there are critical features of Anacostia, such as the crystalliferous exotesta and the trichotomocolpate pollen, that are not consistent with the characters of extant taxa of Austrobaileyales, and relationship to other early diverging angiosperm lineages, for example among magnoliids (e.g., Canellales) cannot be ruled out. Only three specimens of Anacostia have been recovered in the Catefica mesofossil flora, so far. This relative rarity contrasts with occurrences of the genus in the Buarcos, Famalicão and Vale de Água mesofossil floras where fruits and seeds of Anacostia are abundant., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 370, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (1997): Anacostia, a new basal angiosperm from the Early Cretaceous of North America and Portugal with trichotomocolpate / monocolpate pollen. - Grana, 36: 225 - 244. https: // doi. org / 10.1080 / 00173139709362611","Friis, E. M., Crane, P. R., Pedersen, K. R., Stampanoni, M., Marone, F. (2015 b): Exceptional preservation of tiny embryos documents seed dormancy in early angiosperms. - Nature 528, 551 - 554. https: // doi. org / 10.1038 / nature 16441","Doyle, J. A., Endress, P. K. (2014): Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: ANITA lines and relatives of Chloranthaceae. - International Journal of Plant Sciences, 175: 555 - 600. https: // doi. org / 10.1086 / 675935"]}

Serialis communis E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2019 Text-fig. 17a D e s c r i p t i o n a n d r e m a r k s. The material includes several fruits with three to five permanently adhering seeds assignable to Serialis communis (Text-fig. 17a). The fruit wall is thin and typically almost entirely lost by abrasion. The seeds are anatropous, and bitegmic with a thick mesotestal-endotestal seed coat. The micropyle is formed from the inner integument and the micropylar region is seen on the seed surface as a transverse slit in the testa (Text-fig. 17a). In all details the seeds are comparable to the type material from the Famalicão mesofossil flora (Friis et al. 2019c). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Fruits and seeds assigned to the extinct genus Serialis are among the most diverse fossils in Early Cretaceous mesofossil floras from Portugal and nine different species have been recognized (Friis et al. 2019c). Phylogenetic analysis indicates a close relationship of Serialis to Magnoliales, but the genus cannot be placed confidently in any extant taxon within the order. The type material of Serialis communis is from the Famalicão mesofossil flora where S. communis is the most abundant species and more than 2,230 specimens are known. Serialis communis is also reported from the Arazede and Vale de Água localities (Friis et al. 2019c) and is also present in the Chicalhão mesofossil flora (“Fruits with co-adhering seeds in row type 1”; Mendes et al. 2014)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 366, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 c): Extinct diversity among Early Cretaceous angiosperms: mesofossil evidence of early Magnoliales from Portugal. - International Journal of Plant Sciences, 180: 93 - 127. https: // doi. org / 10.1086 / 701319","Mendes, M. M., Dinis, J., Pais, J., Friis, E. M. (2014): Vegetational composition of the Early Cretaceous Chicalhao flora (Lusitanian Basin, western Portugal) based on palynological and mesofossil assemblages. - Review of Palaeobotany and Palynology, 200: 65 - 81. https: // doi. org / 10.1016 / j. revpalbo. 2013.08.003"]}

Proencistemon sp. Text-fig. 9h–j D e s c r i p t i o n a n d r e m a r k s. The material includes a single staminate inflorescence fragment consisting of about seven tightly packed, stamens that lack a well-developed filament and have anthers that are almost sessile (Text-fig. 9h). Anthers are elongate, narrowly obtriangular, and 0.45 mm broad. Their full length is not preserved, but they are more than 0.8 mm long. Pollen grains are trichotomocolpate, about 16 µm in diameter, and semitectate-reticulate (Text-fig. 9i, j). These grains are very similar to pollen found in situ in the stamens of Proencistemon portugallicus. A f f i n i t y a n d o t h e r o c c u r r e n c e s. Pollen grains of Proencistemon sp. are closely similar in size, shape, aperture configuration and details of pollen wall to those found in situ in Proencistemon portugallicus, but the anthers are longer and differ in their narrow obtriangular shape. More material is needed to determine if a new species should be recognized formally. We have not observed similar stamens and pollen in other Early Cretaceous mesofossil floras from Portugal., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 357, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

Stamen with Asteropollis - type pollen sp. 2 Text-fig. 15a–e D e s c r i p t i o n a n d r e m a r k s. The species is based on a single stamen with numerous Asteropollis- type pollen grains in situ. The stamen is about 0.9 mm long and 0.4 mm broad, obovate in shape, with a pointed base and a domeshaped sterile extension of the connective with short, stiff trichomes at the apex (Text-fig. 15a). There are no remains of a filament and the anthers may have been sessile or almost sessile. The anther is tetrasporangiate with two pairs of pollen sacs (Text-fig. 15a). Pollen is pentachotomocolpate, rarely tetrachotomocolpate, circular in equatorial outline and about 15–18 µm in diameter (Text-fig. 15b–d). The arms of the aperture are short and do not reach to the equator. The aperture margins are poorly defined and the aperture membrane is covered by irregular verrucae (Text-fig. 15b– d). The exine is semitectate-reticulate, columellate, with short, densely spaced columellae, about 0.8 µm long that diminish in thickness towards the thick foot layer (Text-fig. 15b–e). Lumina are irregular in shape, up to about 1.2 µm in diameter. Muri are about 0.4 µm wide with a rounded profile and are ornamented by minute verrucae aligned in two longitudinal rows that form poorly defined transverse ridges. A f f i n i t y a n d o t h e r o c c u r r e n c e s. The in situ pollen grains of Asteropollis- type pollen sp. 2 differ from the pollen associated with Hedyflora crystallifera, and also pollen of Asteropollis- type pollen sp. 1 (above), in their smaller size and in having a mainly pentachotomocolpate aperture configuration in contrast to the typical tetrachotomocolpate, or sometimes trichotomocolpate, pollen of Hedyflora crystallifera. Stamens producing the two Asteropollis pollen types (sp. 1 and sp. 2) are also distinct in shape and size. In aperture configuration the pentachotomocolpate grains are more comparable to pollen of Asteropollis asteroides and pollen of extant Hedyosmum (e.g., Walker and Walker 1984)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 362-363, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Walker, J. W., Walker, A. G. (1984): Ultrastructure of Lower Cretaceous angiosperm pollen and the origin and early evolution of flowering plants. - Annals of the Missouri Botanical Garden, 71: 464 - 521. https: // doi. org / 10.2307 / 2399035"]}

Hedyflora crystallifera E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2019 Text-fig. 7a–f D e s c r i p t i o n a n d r e m a r k s. Hedyflora crystallifera was established based on floral structures with adhering pollen from the Buarcos mesofossil flora (Friis et al. 2019b). The specimens from Catefica are often strongly compressed and lignitised. Internal details are known from only a few specimens. It is possible that the material represents more than one species, but the information currently available does not warrant recognition of several taxa. Differences in size may be attributed to differences in maturity as the floral structures appear to be preserved at different post-anthetic stages. The fruits/ovaries are obovate in longitudinal view, triangular in transverse section with rounded to sharp edges (Text-fig. 7a, b, d), and have the remains of three apical tepals (Text-fig. 7a, b). The hypanthium is thickened over the edges of the fruits with lateral depressions between the angles (Text-fig. 7a, b). The outer surface of the hypanthium over the lateral depressions shows polygonal cells each with a central papilla (Text-fig. 7c). Fruits of Hedyflora are one-seeded with an orthotropous, pendent and endotestal seed. The endotesta is distinctly crystalliferous and the outer tegmen is sclerified (Text-fig. 7d). Pollen grains attached to the surface of several fruits are circular in equatorial view, about 22 µm in diameter, and have an irregular branched polar aperture. The aperture is typically tetrachotomocolpate with a poorly defined aperture membrane that has irregular verrucate ornamentation (Text-fig. 7e). The tectum is reticulate with narrow muri, about 0.3 µm wide, ornamented by two poorly defined rows of minute verrucae and supported by long, scattered columellae (Text-fig. 7f). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Hedyflora is closely similar to the pistillate flowers and fruits of extant Hedyosmum SW. (Chloranthaceae) from which it is mainly distinguished by the more elaborate seed coat in the fossil material. Extant Hedyosmum has an unspecialized seed coat that lacks a crystalliferous endotesta and also lacks sclerified cells in the tegmen, both features that are shared by Hedyflora and other extinct and extant Chloranthaceae (Friis et al. 2019b). Fossils assigned to Hedyflora are known from several mesofossil floras from Portugal, including Arazede, Buarcos, Torres Vedras and Vale de Água (Friis et al. 2019a, b). Pollen grains are similar to dispersed fossil grains typically assigned to the pollen genus Asteropollis R.W.HEDL. et G.NORRIS. Closely similar grains are also found in situ in an isolated stamen from Catefica (“Stamen with Asteropollis - type pollen sp. 1”; Text-fig. 14) and similar grains also occur in situ in stamens from the Vale de Água locality (Friis et al. 2019b). Dispersed pollen grains of the Hedyflora type are also present in the Catefica palynoflora, but are rare., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 351-353, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 b): Hedyosmum - like fossils in the Early Cretaceous diversification of angiosperms. - International Journal of Plant Sciences, 180: 232 - 239. https: // doi. org / 10.1086 / 701819","Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013"]}

Serialis crassitesta E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2019 Text-fig. 17b D e s c r i p t i o n a n d r e m a r k s. The material includes several fruits with permanently adhering seeds that are assignable to Serialis crassitesta (Text-fig. 17b). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Conclusions on the relationships of Serialis crassitesta are similar to those on Serialis communis (see above). Serialis crassitesta is common in the Famalicão mesofossil flora with about 375 specimens, but less common than S. communis. Serialis crassitesta is also common in the Vale de Água mesofossil flora (Friis et al. 2019c) and is present in the Chicalhão (“Fruits with co-adhering seeds in row type 2”; Mendes et al. 2014) and Nossa Senhora da Luz mesofossil floras (“Seeds in row”; Mendes and Friis 2018)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 366, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 c): Extinct diversity among Early Cretaceous angiosperms: mesofossil evidence of early Magnoliales from Portugal. - International Journal of Plant Sciences, 180: 93 - 127. https: // doi. org / 10.1086 / 701319","Mendes, M. M., Dinis, J., Pais, J., Friis, E. M. (2014): Vegetational composition of the Early Cretaceous Chicalhao flora (Lusitanian Basin, western Portugal) based on palynological and mesofossil assemblages. - Review of Palaeobotany and Palynology, 200: 65 - 81. https: // doi. org / 10.1016 / j. revpalbo. 2013.08.003","Mendes, M. M., Friis, E. M. (2018): The Nossa Senhora da Luz flora from the Early Cretaceous (early Aptian-late Albian) of Juncal in the western Portuguese Basin. - Acta Palaeobotanica, 58: 159 - 174. https: // doi. org / 10.2478 / acpa- 2018 - 0015"]}

Canrightia resinifera E.M.FRIIS et K.R.PEDERSEN, 2011 Text-fig. 2c–g D e s c r i p t i o n a n d r e m a r k s. Fruits and seeds of Canrightia resinifera are the most common angiosperm fossils in the Catefica mesofossil flora with several hundred specimens recorded so far including the type material on which the genus was established (Friis and Pedersen 2011). The fruits are elliptical to spherical in outline, contain two to five seeds, and have abundant resin-bodies in the fruit wall. showing the long colpus and coarse reticulum; g) Transverse section (orthoslice xy0705) through a fruit showing four seeds all with radially elongated endothelium cells formed from the inner epidermis of the tegmen (asterisks). Specimens, Catefica 150-S174254 (a, b), Catefica 49-S170377 (c), Catefica 49-S170372 (d), Catefica 50-S170401 (e), Catefica 50-S170404 (f), Catefica 50-S174906 (g). Scale bars = 300 Μm (a–e, g), 6 Μm (f). The fruits are interpreted as berries (Text-fig. 2c, d). The fruits develop from bisexual flowers with a semi-inferior ovary and about four staminal scars in a radially symmetrical arrangement on the rim of the hypanthium (Text-fig. 2c). The many resin bodies in the fruit wall, combined with the often-wrinkled fruit surface, sometimes make the precise position of the hypanthium and the staminal scars difficult to distinguish. The stigma at the apex of the fruit is lobed. The seeds are orthotropous, pendent and endotestal, with a distinct, finely crystalliferous, endotesta (Text-fig. 2d, e) and with the inner epidermis of the tegmen developed as a distinct endothelium (Text-fig. 2g). Pollen grains are common in the stigmatic region. They are monocolpate, 15.8–21.0 μm with a long extended colpus, and a coarsely reticulate tectum. Muri are smooth, with a high and sharp profile, and are supported by long and scattered columellae (Text-fig. 2f). The grains are similar to pollen assigned to the extinct genus Piercipollis E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN (Friis et al. 2019a). Dispersed pollen of this type has traditionally been assigned to the extinct genus Retimonocolpites R.L.PIERCE, but in the type species, Retimonocolpites dividuus R.L.PIERCE, the colpus extends from the distal surface over to the proximal surface of the grain dividing the grain in two halves (Pierce 1961). In contrast, in Piercipollis the colpus is restricted to the distal half of the grain (Friis et al. 2019a). A second species of Canrightia, Canrightia foveolata sp. nov., is formally described below from the Catefica mesofossil flora. It is distinguished from C. resinifera by its finely pitted endotesta. Canrightia elongata E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN described from the Torres Vedras mesofossil flora (Friis et al. 2019a) is distinguished from both of the Catefica species by its more elongated fruits that have a shorter hypanthium. A f f i n i t y a n d o t h e r o c c u r r e n c e s. The original phylogenetic analysis of Canrightia placed the genus close to the base of the Chloranthaceae (Friis and Pedersen 2011), a position that has been corroborated by several subsequent analyses (Doyle and Endress 2014, Friis et al. 2015a). Canrightia resinifera is one of the most common angiosperm fossils in the Early Cretaceous floras of Portugal with numerous specimens recorded from the Arazede, Buarcos, Catefica, Famalicão and Vale de Água mesofossil floras (Friis and Pedersen 2011), as well as from the Chicalhão and Nossa Senhora da Luz mesofossil floras (Mendes et al. 2014, Mendes and Friis 2018). In the Catefica mesofossil flora Canrightia resinifera is recorded from all samples. Pollen similar to that associated with Canrightia resinifera has not been observed in situ in any of the dispersed stamens from the Catefica mesofossil flora (Tab. 1). Pollen grains similar to those observed on the Canrightia fruits are present, however, in palynological samples analyzed from the Catefica locality and in other palynofloras from the Early Cretaceous of Lusitanian Basin in western Portugal., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 345-347, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Pedersen, K. R. (2011): Canrightia resinifera gen. et sp. nov., a new extinct angiosperm with Retimonocolpites - type pollen from the Early Cretaceous of Portugal: missing link in the eumagnoliid tree? - Grana, 50: 3 - 29. https: // doi. org / 10.1080 / 00173134.2011.559728","Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013","Pierce, R. L. (1961): Lower Upper Cretaceous plant microfossils from Minnesota. - Minnesota Geological Survey Bulletin, 42: 1 - 86.","Doyle, J. A., Endress, P. K. (2014): Integrating Early Cretaceous fossils into the phylogeny of living angiosperms: ANITA lines and relatives of Chloranthaceae. - International Journal of Plant Sciences, 175: 555 - 600. https: // doi. org / 10.1086 / 675935","Friis, E. M., Grimm, G. W., Mendes, M. M., Pedersen, K. R. (2015 a): Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites - type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae. - Grana, 54: 184 - 212. https: // doi. org / 10.1080 / 00173134.2015.1060750","Mendes, M. M., Dinis, J., Pais, J., Friis, E. M. (2014): Vegetational composition of the Early Cretaceous Chicalhao flora (Lusitanian Basin, western Portugal) based on palynological and mesofossil assemblages. - Review of Palaeobotany and Palynology, 200: 65 - 81. https: // doi. org / 10.1016 / j. revpalbo. 2013.08.003","Mendes, M. M., Friis, E. M. (2018): The Nossa Senhora da Luz flora from the Early Cretaceous (early Aptian-late Albian) of Juncal in the western Portuguese Basin. - Acta Palaeobotanica, 58: 159 - 174. https: // doi. org / 10.2478 / acpa- 2018 - 0015"]}

Endressistemon cateficensis E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-fig. 28a–g H o l o t y p e. S107778 (Catefica sample 49; figured Text-fig. 28a, b, e, f). 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. PFN002797 (for new species). P a r a t y p e s. S107751, S107768, S107769, S266022 (Catefica sample 49). R e p o s i t o r y. Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. From the Catefica locality where the fossils were collected. Ty p e l o c a l i t y. Catefica (39° 03ʹ30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. Almargem Formation, Early Cretaceous (Aptian-early Albian). S p e c i f i c d i a g n o s i s.As for the genus. D i m e n s i o n s. Stamens up to about 0.9 mm long and 0.2 mm in broad. D e s c r i p t i o n a n d r e m a r k s. The material includes isolated staminal structures each consisting of two lateral stamens and in some specimens a median axislike structure. The two stamens and axis-like structure are borne on a short, common stalk (Text-fig. 28a–e). The anthers are basifixed, tetrasporangiate and dithecate (Text-fig. 28a, b)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 381, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

cf. Endressistemon sp. 3 Text-fig. 29e–g D e s c r i p t i o n a n d r e m a r k s. The material comprises a single stamen from which the stamen base is missing. The anthers are basifixed, tetrasporangiate and dithecate (Text-fig. 29e). The stamens are up to about 0.7 mm long, including the apical projection, and about 0.3 mm broad over the anther. Apically the stamen has a prominent, coriaceous projection that is peltate to wing-like (Text-fig. 29e). Dehiscence is longitudinal and the anther wall is rolled back indicating valvate dehiscence (Text-fig. 29e). Pollen grains in situ are monocolpate, semitectatereticulate, circular in equatorial view, 13 µm in diameter, with a homobrochate reticulum (Text-fig. 29f, g). The grains are folded and the extent of the aperture is not fully exposed. A f f i n i t y a n d o t h e r o c c u r r e n c e s. The stamen is similar to stamens of some extant Annonaceae (see comments on Endressistemon above), but also to the individual stamens of Endressistemon cateficensis and it is possible that it was originally part of a similar staminate structure. The pollen grains in situ are also similar to those of Endressistemon cateficensis, but are larger and the two taxa are probably not conspecific. Clade Monocotyledons, Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 383, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

Saportanthus parvus E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2017 Text-fig. 18c, d D e s c r i p t i o n a n d r e m a r k s. The species was described based on well-preserved flowers from the Catefica mesofossil flora (for a full description see Friis et al. 2017). The flowers are small, actinomorphic, with six to eight broadly ovate tepals, five to seven stamens, and a unicarpellate, uniovulate, semi-inferior ovary (Text-fig. 18c, d). The pollen is 8–12 µm in diameter. The aperture configuration of the pollen is not securely established for the material from Catefica, but the tectum ornamentation is finely striate and forms a fingerprint-like pattern (Friis et al. 2017: fig. 13A–C) similar to that known for the trichotomocolpate and dicolpate pollen of the two other species of Saportanthus recorded from other Early Cretaceous mesofossil floras from Portugal. A f f i n i t y a n d o t h e r o c c u r r e n c e s. A phylogenetic assessment of Saportanthus suggests that the genus is sister to, or embedded within, core Laurales (Friis et al. 2017). The genus is widely distributed among the Early Cretaceous mesofossil floras from Portugal. Three species are currently recognized; S. brachystemon E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, S. dolichostemon E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN and S. parvus. Currently, S. parvus is known only from Catefica. “Flower sp. 2” from the Chicalhão site near Juncal is closely similar and may also belong to this species, but only one specimen is known and there are no details of internal features (Friis et al. 2017). Flowers of Saportanthus parvus are common in the Catefica mesofossil flora, but the characteristic, finely striate pollen grains produced by these flowers have so far not been observed in the Catefica dispersed palynoflora, probably due to their very thin and poorly preservable pollen wall., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 367-368, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2017): Saportanthus, an extinct genus of Laurales from the Early Cretaceous of Portugal. - International Journal of Plant Sciences, 178: 650 - 672. https: // doi. org / 10.1086 / 693108"]}

Ibericarpus cuneiformis E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-figs 21–23 H o l o t y p e. S115851 (Catefica sample 49; figured Text-fig. 21a–c). 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. PFN002791 (for new species). P a r a t y p e s. S115852–S115856, S118683–S118685, S265996, S266012 (Catefica sample 49), S170413– S170417, S174907, S266037, S266135 (Catefica sample 50); P0477 (Catefica sample MM75). R e p o s i t o r y. Holotype: Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S). Paratypes: Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S) and Geological Museum of Lisbon, Lisbon, Portugal (P). E t y m o l o g y. From the wedge-shaped fruits. Ty p e l o c a l i t y. Catefica (39° 03ʹ 30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. Almargem Formation, Early Cretaceous (Aptian-early Albian). S p e c i f i c d i a g n o s i s.As for the genus. D i m e n s i o n s. Carpel bearing axis up to about 1.7 mm long and 1.1 mm in diameter; individual carpels up to 1.05 mm long and 0.4 mm broad. D e s c r i p t i o n a n d r e m a r k s. The species is known from two pistillate structures bearing carpels (Textfigs 21a–c, 22a, b) as well as detached fruits that occur either isolated (Text-fig. 22c) or in groups (Text-figs 22d, 23a–c). One group of detached fruits (S174907; Text-fig. 22d) was studied for internal details using SRXTM. The holotype consists of a central axis with numerous carpels borne in a spiral arrangement (Text-fig. 21a–c). Although the specimen has lost some of its carpels the total original number is estimated to have been about 70, based on those still attached to the axis and the scars from the detached carpels. This specimen was probably preserved early in development before the carpels were shed. Another carpel-bearing specimen is thought to be at fruiting stage and the few carpels still attached to the axis when it was recovered were only loosely attached and fell off as the axis was mounted for SEM (Text-fig. 22a, b). The infructescence axis of this second specimen is about 1.9 mm long and based on the scars on the axis, there were about 70 carpels as also in the holotype. The diameter of the axis (ca. 0.3 mm) is more or less uniform from base to apex (Textfig. 22a). There are no traces of other floral organs or bracts associated with the individual carpels or with the carpelbearing axis. Specifically, there is no distinct joint between the infructescence stalk and the portion of the inflorescence axis that has the carpel scars. There are also no scars from bracts, perianth parts or stamens associated with the carpel scars. The carpels are densely spaced on the axis. Each carpel is about 0.35 mm long and 0.25 mm wide, with the carpels of the second specimen (Text-fig. 22) larger than those of the holotype. Carpels are obconical to pyriform in lateral view, and angular in transverse section as a result of their dense packing on the axis. Each carpel contains a single ovate ovule/seed with micropyle pointing towards the base and in one specimen with remains of embryo preserved (Textfig. 23a–c). The epidermis of the carpel wall consists of small, bulging, isodiametric cells covered by a thick cuticle (Textfigs 21a, 22b–d, 23a–c). The cell outlines are particularly distinct in the protected regions where adjacent carpels meet, but less so in the apical portion of the carpel that is free (Text-figs 21c, 22b–d). The stigmatic area of each carpel is seen as a small apical swelling (Text-figs 21c, 22b–d). No pollen grains have been observed in the stigmatic region or on other parts of the structure. A f f i n i t y a n d o t h e r o c c u r r e n c e s. For comments on the possible relationships of Ibericarpus see comments on the genus (above). Ibericarpus cuneiformis is common in the Catefica mesofossil flora, where it is characteristic for the basalmost layers of the outcrop. There are also fruitlets of Ibericarpus cuneiformis in the Buarcos mesofossil flora., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 373-374, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

Genus Ibericarpus E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK gen. nov. T y p e. Ibericarpus cuneiformis E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK 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. PFN0002790 (for new genus). E t y m o l o g y. From the Iberian Peninsula where the fossil was collected. G e n e r i c d i a g n o s i s. Pistillate structure with numerous, densely spaced carpels borne in a spiral arrangement along a slender axis, with no remains of perianth parts or stamens. Carpels obconical to pyriform, sessile, uniovulate. Style lacking, stigmatic region slightly bulging. Fruit indehiscent. Epidermal cells of fruit with isodiametric facets. Ovule/seed obovate with micropyle pointing towards the base of the carpel. Embryo tiny. Seed coat unspecialized. C o m m e n t s o n t h e g e n u s. There are no scars from bracts, perianth parts or stamens on the axis below the carpels, and there are no traces of a perianth or stamens associated with the individual carpels. The structure of the carpel is uncertain, but its shape and the lack of an obvious suture suggests that it is ascidiate. Interpreting the floral structure is not straightforward. One possibility is that the carpel-bearing axis of Ibericarpus cuneiformis is a simple, unbranched inflorescence bearing numerous ebracteate pistillate flowers, each consisting of only a single carpel. Under this interpretation, Ibericarpus cuneiformis shows some similarity to floral structures of Chloranthaceae. Flowers of Chloranthaceae have simple, typically naked flowers, that are borne in elongated inflorescences and the carpels are ascidiate and uniovulate without a style. Among extant Chloranthaceae, Hedyosmum and Ascarina also have unisexual flowers. However, in extant Chloranthaceae the flowers are typically in the axil of a distinct bract and only the staminate flowers of Hedyosmum are ebracteate. Because no bracts are present associated with the individual carpels in Ibericarpus cuneiformis, we regard the inflorescence interpretation as unlikely. This conclusion is also supported by the unspecialized seed coat of Ibericarpus. In all chloranthoid seeds so far described from the Cretaceous, the seed coat is endotestal with crystalliferous endotestal cells. An alternative interpretation of Ibericarpus is that the fruiting structure is derived from a pistillate, perhaps naked, flower with an apocarpous gynoecium of numerous free carpels. Among extant angiosperms, taxa with an apocarpous gynoecium of many carpels arranged spirally along a long, slender floral axis occur in Kadsura and Schisandra (Schisandraceae, Austrobaileyales), in Magnoliaceae (Magnoliales), and also in Galbulimima F.M.BAILEY (Himantandraceae, Magnoliales). Flowers of Magnoliaceae differ from those of Ibericarpus cuneiformis in being bisexual, and typically with well developed, often leathery, perianth parts that leave distinct scars after flowering. The carpels also have a distinct style, and each contains two or more ovules. Flowers of Galbulimima also differ from those of I. cuneiformis in being bisexual, but they are more like the fossils in being naked and in having uniovulate carpels that lack a style. Fruits of Galbulimima are drupes, while those of Ibericarpus are nuts or one-seeded berries. Carpels in both Magnoliaceae and Himantandraceae are plicate or intermediate plicateascidiate. If the carpels of Ibericarpus cuneiformis are correctly interpreted as ascidiate then in this feature they are more similar to the carpels of Austrobaileyales. Flowers of Kadsura and Schisandra are similar to Ibericarpus cuneiformis in their unisexual organization as well as having carpels that lack a style, but flowers of both extant genera have a distinct perianth and also have one to several ovules per carpel. Against this background, while we think that Ibericarpus cuneiformis is most likely an elongated receptacle bearing numerous fruitlets, and while a relationship to extant Austrobaileyales seems the most likely possibility, I. cuneiformis cannot be included securely in any extant angiosperm family or order. Among the fossil floral structures that have a multicarpellate and apocarpous gynoecium, species of Atlantocarpus E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN from the Early Cretaceous floras of Puddledock, Virginia, USA, and also Buarcos and Vale de Água, Portugal (Friis et al. 2020a), are the most similar to Ibericarpus. Atlantocarpus has a very long receptacle and apparently ascidiate carpels that are uniovulate and lack a style. However, fossils of Atlantocarpus have distinct remains of floral organs below the carpels and the receptacle is obconical, rather than slender and stalk-like as in Ibericarpus. Floral structures of Choffaticarpus compactus E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN first described from the Torres Vedras mesofossil flora (Friis et al. 2019a), and Anacostia ? sp. from the Puddledock flora of eastern North America (Friis et al. 2020a), are also similar to Ibericarpus in having tightly packed carpels spirally arranged along an elongated receptacle. However, Anacostia ? sp. differs in having a distinct joint between pedicel and flower with remains of other floral parts below the carpels and Choffaticarpus compactus differs having strongly compressed carpels with a distinct ventral depression (see above)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 370-373, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2020 a): Multiparted, apocarpous flowers from the Early Cretaceous of eastern North America and Portugal. - Fossil Imprint, 76: 279 - 296. https: // doi. org / 10.37520 / fi. 2020.023","Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013"]}

Canrightiopsis intermedia E.M.FRIIS, G.W.GRIMM, M.M.MENDES et K.R.PEDERSEN, 2015 Text-fig. 6d–f D e s c r i p t i o n a n d r e m a r k s. A single specimen from the Catefica mesofossil flora can be assigned with confidence to Canrightiopsis intermedia. It is a small, single-seeded berry with remains of a hypanthium and scars from three stamens on the probable abaxial side of the fruit (Text-fig. 6d). In this respect the specimen is closely similar to Canrightiopsis crassitesta (see above) as also is the seed in being orthotropous, pendent and endotestal with a pitted outer surface of the endotesta. However, the Catefica specimen of C. intermedia differs from C. crassitesta in its much thinner endotesta (Text-fig. 6e, f). Pollen grains have not been observed attached to the C. intermedia specimen from Catefica, but Clavatipollenites - type pollen grains were reported on specimens of C. intermedia from Famalicão (Friis et al. 2015a). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Canrightiopsis is placed in the Chloranthaceae close to Ascarina, Sarcandra and Chloranthus clade (seeabove). Canrightiopsis intermedia is based on fruits and seeds from the Famalicão mesofossil flora where the species is very common (Friis et al. 2015a) and is also reported from the Buarcos and Vale de Água localities. The species is distinguished from C. crassitesta mainly by its much thinner endotesta., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 351, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Grimm, G. W., Mendes, M. M., Pedersen, K. R. (2015 a): Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites - type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae. - Grana, 54: 184 - 212. https: // doi. org / 10.1080 / 00173134.2015.1060750"]}

Genus Valvidistemon E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK gen. nov. T y p e. Valvidistemon globiferus E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK 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. PFN0002794 (for new genus). E t y m o l o g y. From Greek valvida for valve and stemon for stamen. G e n e r i c d i a g n o s i s. Stamen small with a poorly differentiated base. Anther tetrasporangiate, dithecate. Dehiscence valvate by two laterally hinged vales over each theca. Connective between thecae massive. Apical extension of the connective prominent, only slightly shorter than the anther, globose, overhanging the thecae. Pollen small, semitectate-reticulate. C o m m e n t s o n t h e g e n u s. Similarstamenswith valvate dehiscence,laterally hinged valves, poor differentiation between stamen base and anther, and a prominent globose apical extension of the connective, are characteristic of several extant members of Annonaceae (e.g., Endress and Hufford 1989, Van Heusden 1992). Comparable stamens also occur in extinct and extant members of Platanaceae (Friis et al. 1988, Endress and Hufford 1989), but in Platanaceae the stamens have a distinct filament, and the pollen is tricolpate. The stamen is assigned here to a new genus. We are not aware of similar stamens that have been described in the fossil record and because information on other floral parts are missing detailed comparison with extant angiosperms is currently not possible., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 379, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Endress, P. K., Hufford, L. D. (1989): The diversity of stamen structures and dehiscence patterns among Magnoliidae. - Botanical Journal of the Linnean Society, 100: 45 - 85. https: // doi. org / 10.1111 / j. 1095 - 8339.1989. tb 01709. x","Van Heusden, E. C. H. (1992): Flowers of Annonaceae: morphology, classification, and evolution. - Blumea, Supplement, 7: 1 - 218.","Friis, E. M., Crane, P. R., Pedersen, K. R. (1988): Reproductive structure of Cretaceous Platanaceae. - Biologiske Skrifter, det Kongelige Danske Videnskabernes Selskab, 31: 1 - 56."]}

Mugideiriflora portugallica E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2020 Text-fig. 2a, b D e s c r i p t i o n a n d r e m a r k s. Mugideiriflora portugallica is based on a single small, partly abraded, early anthetic flower (Text-fig. 2a) that has a multiparted perianth, androecium and gynoecium and was recovered from the Catefica mesofossil flora (for a full description and discussion of the species see Friis et al. 2020a). Additional specimens are preserved at very early developmental stages. The receptacle is slightly concave, but with a short conical apex in the gynoecial region (Text-fig. 2a). There are about 50 laminar tepals, 50 stamens and more than 50 carpels, all apparently in a spiral arrangement (Text-fig. 2b). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Phylogenetic assessment suggests that Mugideiriflora portugallica is closely related to members of extant Austrobaileyales, although a possible affinity with members of extant Magnoliales cannot be excluded (Friis et al. 2020a). Mugideiriflora portugallica is currently known only from the Catefica mesofossil flora where it is recorded from samples collected near the base of the exposure., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 345, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2020 a): Multiparted, apocarpous flowers from the Early Cretaceous of eastern North America and Portugal. - Fossil Imprint, 76: 279 - 296. https: // doi. org / 10.37520 / fi. 2020.023"]}

Proencistemon portugallicus E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-figs 8a–f, 9a–g H o l o t y p e. P0341 (Catefica sample MM282; figured Text-fig. 8a, c, d). 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. PFN002789 (for new species). P a r a t y p e s. S266015, S266016 (Catefica sample 49), S170393, S170394 (Catefica sample 50), S174257 (Catefica sample 150), S122086 (Catefica sample 342). R e p o s i t o r y. Holotype: Geological Museum of Lisbon, Portugal (P). Paratypes: Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. From Portugal where the fossils were recovered. Ty p e l o c a l i t y. Catefica (39° 03ʹ30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. Almargem Formation, Early Cretaceous (Aptian-early Albian). S p e c i f i c d i a g n o s i s. As for the genus with the following addition: anthers very narrowly elliptical and of the same width from base to apex. D i s t i n g u i s h i n g f e a t u r e s. Proencistemon portugallicus is distinguished from the staminate inflorescence fragment from Catefica described here as Proencistemon sp., in having shorter stamens in which the pollen sacs are of equal width from base to apex. In Proencistemon sp. the pollen sacs are broader towards the apex. D i m e n s i o n s. Staminate structure (measured on holotype) about 1.5 mm in diameter; stamens 0.55 mm long and 0.25 mm broad; pollen diameter 12.5–16 µm. D e s c r i p t i o n a n d r e m a r k s. The material includes two spherical staminate inflorescences that are three dimensionally preserved and have several series of stamens radiating from the central axis (Text-fig. 8a, b). There are also strongly compressed, lignitised inflorescence fragments (Text-fig. 9a, b), isolated stamens and pollen clumps. The inflorescences are unisexual, about 1.5 mm in diameter and appear to be more or less spherical. They consist of up to 30 densely packed stamens. Bracts observed on the abaxial surface of an inflorescence fragment are poorly preserved (Text-fig. 9a) and it is unclear whether each bract subtends one or several stamens. The arrangement of the stamens in the three-dimensionally preserved specimens is also not conclusive. In specimen P0341 stamens appear to be arranged singly (Text-fig. 8a), while in specimen S174257 some of the stamens appear to occur in pairs (Text-fig. 8b). This is also the case for several of the compressed specimens (Text-fig. 9a, b). Stamens are narrowly elongate, elliptical to rectangular in shape, about 0.55 mm long, 0.25 mm broad, and lack a well-developed filament. Anthers are sessile or almost sessile, dithecate, tetrasporangiate and with a short flattened apical connective that is mostly abraded (Text-fig. 8a, b). Stamens in the lignitised specimens are flattened and elliptical (Text-fig. 9b). They are slightly longer than those in the charcoalified stamens, but are treated here as the same species based on the identical pollen. Pollen grains are small, circular in outline, 12.5–16 µm in diameter and have a trichotomocolpate aperture in which the arms are short and do not reach to the equator (Text-figs 8c, e, 9c–f). The aperture margin is indistinct and the aperture membrane is irregularly verrucate (Text-figs 8c, e, 9c–f). The pollen wall is semitectate-reticulate with a homobrochate reticulum (Text-figs 8c–f, 9c–f). Muri are about 0.2 µm broad with a rounded profile and a supratectal ornamentation of minute verrucae that are aligned in two to three longitudinal rows and form poorly defined transverse ridges over the muri (Text-figs 8d, f, 9g). Muri are supported by medium sized and widely spaced columellae (Text-figs 8d, 9g). Lumina are irregular in shape and up to about 0.8 µm in diameter. Tiny, spherical orbicules, about 0.5 µm in diameter, which are ornamented by fine verrucae-spinules, are present on the surface of some pollen grains (Text-fig. 8f). Pollen morphology and ultrastructure was described in detail for specimen P0341 (Tekleva et al. 2021), which we designate here as the holotype of Proencistemon portugallicus. A f f i n i t y a n d o t h e r o c c u r r e n c e s. For comments on the relationships to extant chloranthoids see discussion of the new genus above. A phylogenetic analysis was performed by Tekleva et al. (2021) based on specimen P0341, which suggested that “despite some uncertainty … phylogenetic analyses are most consistent with a position attached to the stem lineage of Hedyosmum.” This conclusion may be correct, but because other relevant specimens were not considered, the full significance of the material requires further analysis. In specimen P0341 the bracts are not obvious and the stamens appear to be borne singly. The flowers were therefore interpreted as ebracteate and unistaminate (Tekleva et al. 2021). However, in another specimen bracts are clearly present (Text-fig. 9a), and in several inflorescence fragments stamens appear to occur in pairs (Text-figs 8b, 9a). Together with the trichotomocolpate pollen, these points of similarity with extant Ascarina, rather than with extant Hedyosmum, need to be considered. Densely crowded stamens very similar to those of Proencistemon portugallicus, and also with similar in situ trichotomocolpate pollen, have been described from the Torres Vedras mesofossil flora (Friis et al. 2019a: text-fig. 21). The two taxa are clearly closely related, but stamens of the Torres Vedras specimens are larger and more crowded and also have larger pollen grains (about 18–22 µm in diameter compared to 12.5–16 µm in diameter in Proencistemon portugallicus). Whether the stamens in the Torres Vedras specimens are in pairs, and whether the flowers were bracteate or ebracteate is unknown. In the Catefica palynoflora similar trichotomocolpate pollen grains are rare, but have been reported from coastal exposures in Portugal that are of Early Cretaceous age as Asteropollis cf. asteroides, Asteropollis sp. 3 and Asteropollis sp. 4 (Heimhofer et al. 2007). These trichotomocolpate grains are similar to those of Proencistemon portugallicus in general morphology, but are larger. The specimen illustrated and assigned to Asteropollis as Asteropollis cf. asteroides (Heimhofer et al. 2007: pl. III, figs 1, 2) differs more significantly in being tetrachotomocolpate., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 355-357, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Tekleva, M., Mendes, M. M., Kvacek, J., Endress, P. K., Doyle, J. A. (2021): Morphology, ultrastructure, and evolutionary significance of pollen in a chloranthaceous staminate structure from the Early Cretaceous of Portugal. - International Journal of Plant Sciences, 182: 817 - 832. https: // doi. org / 10.1086 / 716778","Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013","Heimhofer, U., Hochuli, P. A., Burla, S., Weissert, H. (2007): New records of Early Cretaceous angiosperm pollen from Portuguese coastal deposits: Implications for the timing of the early angiosperm radiation. - Review of Palaeobotany and Palynology, 144: 39 - 76. https: // doi. org / 10.1016 / j. revpalbo. 2005.09.006"]}

Kempia longicolpites E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2019 Text-fig. 30a–f D e s c r i p t i o n a n d r e m a r k s. The material comprises two adhering stamens (only one cut stamen illustrated) with dithecate, tetrasporangiate anthers (Text-fig. 30a) and in situ pollen. The stamens are about 1 mm long and 0.25 mm broad with a distinct, triangular apical extension of the connective. One stamen was cut transversely into two pieces to expose the pollen for SEM. The other stamen was removed for TEM. Stamens and anthers are closely similar to Kempia longicolpites described from the Torres Vedras locality (Friis et al. 2019a) and the specimen is assigned here to the same species. Pollen grains are small, about 11– 12 µm long, monoaperturate and with the colpus extending beyond the full length of the grains. The exine is semitectatereticulate, columellate (Text-fig. 30b–f) with the reticulum and columellae only loosely attached to the foot layer (Text-fig. 30f). The muri have a rounded profile and a smooth surface (Text-fig. 30c). The foot layer is thick, and the endexine is restricted to the apertural region (Text-fig. 30e). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Kempia longicolpites was first described from the Torres Vedras locality (Friis et al. 2019a) and is currently known from only the Torres Vedras and Catefica mesofossil floras., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 385, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013"]}

Elasmostemon paisii E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-figs 25a–h, 26a–c H o l o t y p e. S105281 (Catefica sample 151; figured Text-fig. 25d–h). 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. PFN002793 (for new species). P a r a t y p e s. S115859, S172560 (Catefica sample 49). R e p o s i t o r y. Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. In honor of Professor João Pais (1949 – 2016) for his contribution to the palaeobotany and geology of Portugal. Ty p e l o c a l i t y. Catefica (39° 03ʹ30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. Almargem Formation, Early Cretaceous (Aptian-early Albian). S p e c i f i c d i a g n o s i s.As for the genus. D i m e n s i o n s. Stamen fragments 0.6–1.7 mm long (full length unknown); 0.5–0.7 mm broad. D e s c r i p t i o n a n d r e m a r k s. The material comprises two small stamen fragments (S105281 and S115859) that are about 0.6–0.8 mm long and 0.5 mm broad (Text-fig. 25a, d) and a larger fragment (S172560), about 1.6 mm and 0.7 mm broad (Text-fig. 26a–c). The three fragments preserve different parts of the stamen, and apparently also slightly different developmental stages. They are treated here as a single species based on the stamen shape, the orientation and positioning of the narrow pollen sacs, and the shared in situ monocolpate, semitectatereticulate pollen (Text-figs 25a–h, 26a–c). The stamens are broad, tetrasporangiate and dithecate, and abaxially-adaxially flattened. The stamen apex, preserved in specimens S115859 and S172560, is rounded without an apical extension (Text-figs 25a, 26a). The stamen base is not preserved in any of the specimens and the full length of the stamen is unknown. In specimen S115859 the marginal tissue appears to be abraded. The pollen sacs are arranged in two pairs on one surface of the stamen close to the stamen margins. It is unknown whether the pollen sacs are on the abaxial or adaxial stamen surface. The two pairs of pollen sacs are separated from each other by a broad zone of connective tissue but are oriented such that they converge and meet near the stamen apex (Text-fig. 25a, d). Dehiscence of the pollen sacs is longitudinal. In the two smaller fragments the thecae are not open, while in the larger specimen the thecae are dehisced with their walls curved back (Text-fig. 26a). Larger cells, interpreted as ethereal oil cells, are closely spaced in the staminal tissue and particularly well-preserved in specimen S115859 as shallow depressions surrounded by several cells that produce rounded swellings (Text-fig. 25a, b). In the other two specimens these cells are obscured by poor preservation. Mature pollen grains are exposed by fractures in the undehisced, smaller specimens. In the larger, dehisced specimen most of the pollen had been shed, but a group of grains, perhaps immature, remained attached to the inside of the anther wall. Pollen grains of specimen S115859 were described and figured earlier as Pollen type D.8 (Friis et al. 1999). Grains from specimen S105281 are very similar but folded, which obscures the apertures. The pollen is circular in equatorial view, about 15–17 µm in diameter, and monocolpate. The exine is semitectate-reticulate with a heterobrochate, loosely attached reticulum (Text-fig. 25c, e–h). The aperture is long, reaches to the equator, and has distinct margins (Text-fig. 25e). Lumina are rounded to angular, with larger lumina up to about 1.6 µm in diameter and smaller lumina about 0.2–0.5 µm in diameter. Muri are narrow, about 0.2 µm wide, with a flattened profile and smooth surface (Text-fig. 25h). Columellae are short, about 0.2 µm long (Text-fig. 25h). Pollen grains in S172560 vary markedly in size and may be immature but are also partly obscured by residual organic material. They show a gradation, from grains that are almost smooth, to grains with a very weakly developed reticulum (Text-fig. 26b, c). Pollen grains in specimen S172560 are also smaller than in the two other specimens, about 12 µm in diameter, and in some grains the reticulum is denser. The inner surface of the anther wall in the dehisced specimen is finely granular, probably reflecting the presence of tiny orbicules. A f f i n i t y a n d o t h e r o c c u r r e n c e s. For the possible systematic relationships of Elasmostemon paisii see comments on the genus above. The stamen fragments are closely similar to the specimen described below as “Laminar stamen with monocolpate reticulate pollen”. However, the pollen in the two stamen types differ in the details of their wall structure. Similar stamens have not been encountered in other mesofossil floras from Portugal. Melloniflora E.M.FRIIS, P.R. CRANE et K. R.PEDERSEN, and several different isolated stamens from the Early Cretaceous Puddledock flora of Virginia, USA, have pollen sacs that are embedded in the staminal tissue in a non-marginal position, but they differ in their larger size and their more elongate, scale-like form (Friis et al. 2020b)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 376-378, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Pedersen, K. R., Crane, P. R. (1999): Early angiosperm diversification: the diversity of pollen associated with angiosperm reproductive structures in Early Cretaceous floras. - Annals of the Missouri Botanical Garden, 86: 259 - 296. https: // doi. org / 10.2307 / 2666179","Friis, E. M., Crane, P. R., Pedersen, K. R. (2020 b): Melloniflora, a new extinct multiparted flower from the Early Cretaceous of Virginia. - International Journal of Plant Sciences, 181: 887 - 897. https: // doi. org / 10.1086 / 710490"]}

Paisia -like follicles Text-figs 34a–h, 35a–e, 36a–e D e s c r i p t i o n a n d r e m a r k s. The material includes several isolated follicles probably derived from apocarpous gynoecia. The follicles vary considerably in size and shape, and range from narrow elongated-ellipsoidal to obovate. Follicles are 0.7–1.7 mm long, 0.35–0.7 mm broad in the dorsi-ventral direction, and 0.3–6.2 mm wide (Textfigs 34a–h, 35a–e, 36a–e). The follicles have a distinct ventral suture with a decurrent stigma that extends along the full length of the follicle. In some specimens there is a distinct papillate zone that extends along both sides of the ventral suture from the follicle base to the apex (Text-fig. 34a–d). This papillate zone is possibly stigmatic. In many specimens the follicles have a distinct apical cleft (Text-figs 34d, 35a, d, 36b, d). The follicles contain many anatropous ovules/seeds that are borne on placentae that extend on either side of the ventral suture for the full length of the follicle. In most specimens the ovules/seeds are arranged in two distinct rows (Text-figs 34e–h, 35e), but in one specimen (Text-fig. 36a–e) this is less distinct and the ovules/seeds are more crowded. This specimen is larger than the other follicles and clearly more mature. There are smaller undeveloped ovules in its lower part (Text-fig. 36c) but there are larger, probably mature, seeds in the upper part (Text-fig. 36c). Whether the crowding of the seeds is due to the stage of maturity of the follicles, or because this larger specimen represents another species, is uncertain. The epidermal cells of the ovules/seeds have slightly raised anticlinal walls that give the surface a striate-reticulate appearance (Text-figs 34e, f, 36c). The follicle wall is thick. It consists of an inner layer of transversely aligned fibres (Text-fig. 34e), a middle layer that is one to two cell layers deep and an outer epidermis of smaller, thin-walled cells (Text-figs 34g, 35e). The mesocarp is composed of large, isodiametric, thick-walled cells that have a rounded cell lumen (Text-figs 34e, f, h, 35e). The follicle is supplied by one dorsal and two ventral bundles (Text-figs 34g, h, 35e, 36e). A f f i n i t y a n d o t h e r o c c u r r e n c e s. The follicles are closely similar to the follicles seen in flowers of Paisia pantoporata in their elongate shape and in the anatomical details of the follicle wall. However, the carpels in the Paisia flowers are immature and the stigmatic zone is indistinct. Other features seen in the isolated follicles, such as the apical cleft, and a papillate zone extending along the margins of the ventral suture, are also not seen in Paisia pantoporata. It is possible that the isolated follicles represent different developmental stages of a single species, perhaps Paisia pantoporata. However, it is also possible that they represent additional species of Paisia or a closely related genus. Isolated follicular fruits occur in other mesofossil floras from Portugal, but Paisia -like follicles like those described here are known only from Catefica. Isolated eudicot stamens and pollen clumps with tricolpate pollen in situ Five different kinds of eudicot have been identified from the Catefica mesofossil flora based on isolated stamens and pollen clumps with different kinds of tricolpate pollen. Dispersed tricolpate pollen grains are typically assigned to species of dispersed pollen genera such as Foveotricolpites R.L.PIERCE, Psilatricolpites HAMMEN ex HAMMEN et WYMSTRA, Retitricolpites HAMMEN ex HAMMEN et WYMSTRA, Rhoipites WODEHOUSE, Rousea SAT.K.SRIVAST., or Striatopollis KRUTZSCH and Tricolpites COOKSON ex COUPER. However, the application of these pollen genera is typically very broad and their type specimens have been studied and illustrated mainly using light microscopy. Light microscope resolution is insufficient for detailed comparison with specimens studied using scanning electron microscopy, including the specimens described here from Catefica. New genera are probably warranted for the Catefica material, but for present purposes we simply refer to the five different taxa as “Stamen/pollen clump with tricolpate pollen sp. 1 – sp. 5”. Stamen with tricolpate pollen sp. 1 Text-fig. 37a–f D e s c r i p t i o n a n d r e m a r k s. The material comprises a single fragmentary stamen, about 2.3 mm long and 0.5 mm wide. The anther is tetrasporangiate and dithecate with long narrow pollen sacs (Text-fig. 37a). Pollen grains in situ are small, almost spherical, about 21 µm in diameter and tricolpate (Text-fig. 37b–d). The colpi are long, reaching almost to the poles, and have a distinct margin (Text-fig. 37b– d). The grains are semitectate-reticulate with a heterobrochate reticulum that is coarse in the mesocolpium zones but finer over the poles and along the margins of the colpi (Textfig. 37b–d). The muri are smooth, about 0.4 µm wide, with a rounded to flattened profile. Columellae supporting the muri are short and densely spaced (Text-fig. 37f). Orbicules are densely-scattered on the inner surface of the anther wall and over the surface of the pollen grains (Text-fig. 37e). Orbicules are up to about 1 µm long, irregular in shape and have a solid base of laterally fused spheres with rod-like projections (Text-fig. 37e). A f f i n i t y a n d o t h e r o c c u r r e n c e s. The triaperturate pollen grains clearly indicate a relationship to eudicots, but relationships to extant taxa within the group are uncertain. In the mesofossil floras from Portugal this taxon is currently known only from Catefica. The in situ pollen grains are identical in size, shape and most features of the reticulum to the dispersed pollen Retitri-Liliret described by Penny (1991) from the Aptian of Egypt, but in that material the polar regions are foveolate to psilate rather than finely reticulate., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 388-390, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Penny, J. H. J. (1991): Early Cretaceous angiosperm pollen from the borehole Mersa Matruh 1, North West Desert, Egypt. - Palaeontographica, Abt. B, 222: 31 - 88."]}

Canrightia foveolata E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-figs 3a–f, 4a–i H o l o t y p e. S174249 (Catefica sample 49; figured Text-fig. 3a–f). 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. PFN002785 (for new species). P a r a t y p e s. S175179, S265998, S266057, S266107 (Catefica sample 49), S266042 (Catefica sample 154), S175178 (Catefica sample 242). R e p o s i t o r y. Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden. E t y m o l o g y. From Latin: fovea (pit) referring to the densely pitted surface of the endotesta. Ty p e l o c a l i t y. Catefica (39° 03ʹ 30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. 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. D i s t i n g u i s h i n g f e a t u r e s. The new species is assigned to the extinct genus Canrightia based on the berry-like fruit with pendent, orthotropous seeds that have an endotestal-endotegmic seed coat and a crystalliferous endotesta. Canrightia foveolata is distinguished from C. resinifera (see above), and from C. elongata from the Torres Vedras mesofossil flora (Friis et al. 2019a), mainly by the densely pitted and grooved surface of the endotesta. Seeds of C. foveolata also have crystals that are of more or less of similar size and that are evenly distributed in the endotestal cells, whereas in C. resinifera and C. elongata larger crystals are concentrated close to the outer surface of endotesta. Canrightia foveolata is also two-seeded, as are most specimens of C. resinifera from the Famalicão locality, while fruits of C. resinifera from the Catefica locality typically have three to five seeds and C. elongata has three seeds. Canrightia foveolata is further distinguished from the two other species of Canrightia by the well-developed soft tissue of the fruit wall above the seeds. Canrightia foveolata may also be distinguished from the two other species by the larger number of perianth parts, but as the perianth is known for only one specimen of C. foveolata, and only a few specimens of C. resinifera, the range of tepal numbers in Canrightia is not fully established. A pitted surface of the endotesta is also present in seeds of Canrightiopsis E.M.FRIIS, G.W.GRIMM, M.M.MENDES et K.R.PEDERSEN and Kvacekispermum E.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. D i m e n s i o n s. 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. D e s c r i p t i o n a n d r e m a r k s. 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). 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. A f f i n i t y a n d o t h e r o c c u r r e n c e s. The relationships of Canrightia foveolata, as for Canrightia resinifera, arelikelyclosetothebaseofextantChloranthaceae (see above). Canrightia foveolata is currently known only from the Catefica locality., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 347-349, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013","Friis, E. M., Grimm, G. W., Mendes, M. M., Pedersen, K. R. (2015 a): Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites - type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae. - Grana, 54: 184 - 212. https: // doi. org / 10.1080 / 00173134.2015.1060750","Friis, E. M., Crane, P. R., Pedersen, K. R. (2018 b): Rightcania and Kvacekispermum: Early Cretaceous seeds from eastern North America and Portugal provide further evidence of the early chloranthoid diversification. - Fossil Imprint, 74: 65 - 76. https: // doi. org / 10.2478 / if- 2018 - 0006"]}

Goczania rugosa E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2019 Text-fig. 20e–h D e s c r i p t i o n a n d r e m a r k s. Goczania rugosa is represented in the Catefica mesofossil flora by two anthers and an isolated pollen sac with pollen grains in situ (Textfig. 20e–h). Identical pollen also occurs in pollen clumps and coprolites. The anther is short and broad, about 0.6 mm long and 0.55 mm wide, dithecate and tetrasporangiate (Text-fig. 20e). As in the type material from Torres Vedras, the inner wall of the anthers of the Catefica specimen and the in situ pollen grains show numerous small, spherical orbicules with a finely spiny surface ornamentation (Text-fig. 20f). The pollen grains are oblate, circular to elliptical in equatorial outline, about 17 µm in diameter and monocolpate (Text-fig. 20f–h). The colpus is short with an irregular margin (Text-fig. 20h). The exine is tectate with the tectum covered with densely spaced microechinae that occur singly without merging with their neighbors (Text-fig. 20f–h). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Pollen of the Goczania type has been found on the stigma and surface of Appomattoxia fruits in the Torres Vedras mesofossil flora and also on fruits of Appofructus E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN from Torres Vedras (Friis et al. 2019a). Goczania- type pollen has also been found on the stigma and surface of Appomattoxia fruits from the Puddledock mesofossil flora of eastern North America. Appomattoxia and Appofructus are both thought to be related to Piperales (Friis et al. 1995, 2019a). Goczania rugosa was first described from the Torres Vedras mesofossil flora (Friis et al. 2019a) and the anthers with in situ pollen from Catefica are closely similar to the type material. Small differences, such as the slightly larger size of the pollen grains and slightly smoother pollen wall in the Catefica specimens, may be related to differences in preservation, with the Torres Vedras material being slightly more shrunken. Two other species of Goczania occur with Goczania rugosa at Torres Vedras, but they differ in details of the supratectal ornamentation of the pollen wall (Friis et al. 2019a). Pollen grains of Goczania rugosa have also been observed in palynological strew preparations of the Catefica microfossil assemblages., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 368-370, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013","Friis, E. M., Pedersen, K. R., Crane, P. R. (1995): Appomattoxia ancistrophora gen. et sp. nov., a new Early Cretaceous plant with similarities to Circaeaster and extant Magnoliidae. - American Journal of Botany, 82: 933 - 943. https: // doi. org / 10.1002 / j. 1537 - 2197.1995. tb 15710. x"]}

Canrightiopsis crassitesta E.M.FRIIS, G.W.GRIMM, M.M.MENDES et K.R.PEDERSEN, 2015 Text-fig. 6a–c, g, h D e s c r i p t i o n a n d r e m a r k s. Canrightiopsis crassitesta was established based on fruits, seeds and adhering pollen from the Catefica mesofossil flora (Friis et al. 2015a). The fruits are elliptical to spherical in outline and are interpreted as berries with a single seed (Text-fig. 6a, b). They are derived from bisexual flowers and remains of a hypanthium, as well as scars from stamens, are present on the probable abaxial face of the fruit, about one third to two thirds of the distance from the base (Text-fig. 6a, d). The seeds are orthotropous, pendent and endotestal with a distinct, thick and finely crystalliferous endotesta (Textfig. 6b, c). The outer surface of endotesta is characterized by relatively large pits arranged in longitudinal rows that are also visible where the fruit wall is compressed or poorly preserved (Text-fig. 6a). The tegmen is three cell layers thick. In some specimens, remains of an endothelium are seen as slightly elongated cells, but the distinct endothelium seen in other species of Canrightiopsis has not been observed. Pollen grains attached to the fruits are similar to dispersed pollen assigned to the extinct pollen genus Clavatipollenites COUPER (Text-fig. 6g, h). Grains are 12–14 µm in equatorial diameter, monocolpate, semitectate-reticulate with a long, extended colpus with an irregular margin. The reticulum is composed of narrow, beaded muri supported by long, scattered columellae (Text-fig. 6g, h). The embryo is minute and surrounded by a nutritive tissue of thin-walled, isodiametric cells (Text-fig. 6b, c). A f f i n i t y a n d o t h e r o c c u r r e n c e s. Analysis of the phylogenetic relationships of Canrightiopsis placed the genus in the Chloranthaceae as part of the Ascarina J.R.FORST. et G.FORST. - Sarcandra GARDNER- Chloranthus SW. clade, particularly close to Sarcandra and Chloranthus (Friis et al. 2015a), a result also supported by a subsequent analysis (Doyle and Endress 2018). Fruits and seeds of Canrightiopsis are common in Early Cretaceous mesofossil floras from Portugal. In addition to C. crassitesta, two other species have been recognized including C. intermedia and C. dinisii E.M.FRIIS, G.W.GRIMM, M.M.MENDES et K.R.PEDERSEN. Only C. crassitesta and C. intermedia are present in the Catefica mesofossil flora. C. crassitesta is distinguished from C. intermedia by its much thicker endotesta, but the two species are similar in fruit morphology and without internal details, the fossils are difficult to separate. All Canrightiopsis specimens from Catefica studied using SEM are typical C. crassitesta, while only one specimen is a distinct C. intermedia. Other specimens from Catefica for which internal features are unknown are referred to as Canrightiopsis sp. (Friis et al. 2015a). Fruits and seeds of Canrightiopsis are particularly common in the mesofossil flora from Famalicão, but are also reported from the Arazede, Buarcos, Chicalhão, Vale de Água and Vila Verde mesofossil floras (Friis et al. 2015a). Currently C. crassitesta is reported only from the Catefica mesofossil flora. Pollen grains found on fruits of Canrightiopsis crassitesta are similar in size and general morphology to those found in situ in isolated stamens and inflorescence fragments from Catefica with Clavatipollenites -type pollen (Text-figs 10– 13, Tab. 1), but the reticulum of the pollen associated with 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). Canrightiopsis crassitesta is more open and the grains are smaller., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on pages 349-351, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Grimm, G. W., Mendes, M. M., Pedersen, K. R. (2015 a): Canrightiopsis, a new Early Cretaceous fossil with Clavatipollenites - type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae. - Grana, 54: 184 - 212. https: // doi. org / 10.1080 / 00173134.2015.1060750","Doyle, J. A., Endress, P. K. (2018): Phylogenetic analyses of Cretaceous fossils related to Chloranthaceae and their evolutionary implications. - The Botanical Review, 84: 156 - 202. https: // doi. org / 10.1007 / s 12229 - 018 - 9197 - 6"]}

Valvidistemon globiferus E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK sp. nov. Text-fig. 27a–d H o l o t y p e. S107779 (Catefica sample 49; figured Text-fig. 27a–d). 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. PFN002795 (for new species). R e p o s i t o r y. Palaeobotanical Collections, Department of Palaeobiology, Swedish Museum of Natural History, Stockholm, Sweden (S). E t y m o l o g y. From Latin in Greek combinations globifer, with spherical organ. Ty p e l o c a l i t y. Catefica (39° 03ʹ30ʺ N; 09°14ʹ 30ʺ W), between the villages of Catefica and Mugideira, about 4 km south of Torres Vedras, Portugal. T y p e s t r a t u m a n d a g e. Almargem Formation, Early Cretaceous (Aptian-early Albian). S p e c i f i c d i a g n o s i s.As for the genus. D i m e n s i o n s. Stamens up to about 0.9 mm long and 0.2 mm in broad. D e s c r i p t i o n a n d r e m a r k s. The material comprises a single stamen, about 0.6 mm long with semitectate-reticulate pollen in situ (Text-fig. 27a–d). The stamen consists of a short base that is poorly differentiated from the basifixed anther. The anther is tetrasporangiate and dithecate with an almost spherical apical extension of the connective that overhangs the thecae (Text-fig. 27a, b). Epidermal cells of the connective tissue between the thecae are slightly elongate and arranged in longitudinal rows (Textfig. 27c). The four pollen sacs are arranged in two pairs in a lateral position and are separated by a broad connective that is equally thick on both adaxial and abaxial sides (Textfig. 27a, b). Dehiscence is valvate by laterally hinged valves that result from a distal and proximal bifurcation of the stomium (Text-fig. 27c). Valves are preserved on one side of the stamen (Text-fig. 27a, c), but are broken off on the other side exposing the distinct, quadrangular cells of the endothecium (Text-fig. 27b). The thecae are dehisced. Most pollen has been shed and only a few grains remain adhering to the inside of the anther wall (Text-fig. 27d). Pollen grains are about 15 µm in diameter. The grains are folded and while the apertures of all the grains are poorly exposed the pollen appears monoaperturate, probably monocolpate. The pollen wall is semitectate-reticulate with a homobrochate reticulum (Textfig. 27d). There is no trace of orbicules on the inside of the anther wall. A f f i n i t y a n d o t h e r o c c u r r e n c e s. Detailed consideration of possible systematic affinities is not possible given the limited material and information available. Similar stamens have not been observed in other Early Cretaceous mesofossil floras from Portugal and North America., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 379, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

cf. Endressistemon sp. 1 Text-fig. 29a, b D e s c r i p t i o n a n d r e m a r k s. The material includes a pair of compressed stamens resembling the staminate structures of Endressistemon cateficensis described above.The stamens are up to 0.8 mm long including the apical projection, and about 0.3 mm broad. The apical projection is longer than the thecae, about 0.45 mm long, and tapers to a long, pointed tip (Text-fig. 29a). The two stamens adhere together closely, but their bases are missing, and whether the stamens had separate distinct bases or a shared base, or whether the base was lost during fossilization, is not known. Anther dehiscence is longitudinal. Pollen grains observed in situ (Text-fig. 29b) are monocolpate, semitectate-reticulate, circular in equatorial view, about 9 µm in diameter. The aperture is long, reaching almost to the equator, and the aperture margin is distinct. The reticulum is homobrochate (Text-fig. 29b). A f f i n i t y a n d o t h e r o c c u r r e n c e s. The stamens and in situ pollen are closely similar to those of Endressistemon cateficensis described above, but the pollen sacs are more rounded and the apical projection is much longer. Because of the missing stamen base it is uncertain whether the two taxa are closely related., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 383, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

OrderChloranthales R.BR. ex SIMS, 1821 R e m a r k s. Chloranthoid fossils are the most diverse group of angiosperms in the Catefica mesofossil flora. Eighteen taxa are recognized based on inflorescences and flowers as well as isolated fruits, seeds and stamens (Textfigs 2–16, Tab. 1)., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 345, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

Catanthus dolichostemon E.M.FRIIS, P.R.CRANE et K.R.PEDERSEN, 2020 Text-fig. 18a, b D e s c r i p t i o n a n d r e m a r k s. The species was described based on several flower buds and open flowers preserved as charcoalified or lignitic specimens from the Catefica mesofossil flora (for a full description see Friis et al. 2021; taxon names valid from effectively published online version in 2020, see Friis et al. 2020c). The flowers are whorled with nine tepals in three whorls of three and many stamens in several successive whorls. The stamens have long, broad and fleshy bases that continue into the short anthers without a joint. In lignitised specimens, the stamens are flattened, but their original bulky, three-dimensional shape is particularly well-preserved in charcoalified specimens (Text-fig. 18a, b). The pollen is circular in polar view, about 12 µm in diameter, monoaperturate and trichotomocolpate (Friis et al. 2021: fig. 4A–C). The gynoecium is superior, apocarpous and consists of six, or rarely five, carpels. A f f i n i t y a n d o t h e r o c c u r r e n c e s. A phylogenetic assessment of Catanthus dolichostemon suggests a relationship to extant members of Canellales and Magnoliales (Friis et al. 2021). The species is currently known only from the Catefica and Vale de Água mesofossil floras. Pollen grains of Catanthus dolichostemon have also been observed in palynological strew preparations from the Catefica microfossil assemblages., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 367, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2021): Catanthus, an extinct magnoliid flower from the Early Cretaceous of Portugal. - International Journal of Plant Sciences, 182: 28 - 45. https: // doi. org / 10.1086 / 711081","Friis, E. M., Crane, P. R., Pedersen, K. R. (2020 c): Catanthus, an extinct magnoliid flower from the Early Cretaceous of Portugal. - International Journal of Plant Sciences, 182 (online): 18 pp. [online version of Friis et al. (2021), which meets the requirements of ICN for effectively published publication] https: // doi. org / 10.1086 / 711081"]}

Genus Proencistemon E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK gen. nov. T y p e. Proencistemon portugallicus E.M.FRIIS, P.R.CRANE, K.R.PEDERSEN, M.M.MENDES et J.KVAČEK 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. PFN002788 (for new genus). E t y m o l o g y. In honor of Pedro Proença e Cunha for his contributions to understanding the stratigraphy and geology of the Early Cretaceous of Portugal and stemon (Greek for stamen). G e n e r i c d i a g n o s i s. Staminate inflorescences spherical with closely packed, unisexual flowers radiating from the central axis. Each flower delimited by a bract subtending one or two stamens. Anthers sessile, or almost sessile, narrowly elongate, dithecate and tetrasporangiate, lacking a well-developed filament but with a short flattened apical connective. Anther dehiscence lateral by longitudinal slits. Pollen small, trichotomocolpate, circular in outline, semitectate-reticulate, columellate, with long, scattered columellae and a homobrochate reticulum. Muri with a rounded profile and supratectal ornamentation composed of minute verrucae aligned in two to three longitudinal rows that create poorly defined transverse ridges over the muri. Aperture margin indistinct. Aperture membrane irregularly verrucate. Orbicules spherical with fine verrucate-spiny ornamentation. D i s t i n g u i s h i n g f e a t u r e s. Among extant angiosperms, species of Ascarina (Chloranthaceae) are comparable to Proencistemon in having staminate inflorescences consisting of simple flowers with one to three stamens. Pollen grains of extant Ascarina, as well as extant Hedyosmum, are also closely similar to those found in situ in Proencistemon. The grains have an indistinctly delimited polar aperture with poorly defined aperture margins, an aperture membrane covered by irregular verrucae, and a semitectate-reticulate pollen wall with finely beaded supratectal ornamentation. The trichotomocolpate pollen of Proencistemon is most similar to the monocolpate or sometimes trichotomocolpate pollen of Ascarina. Pollen of Hedyosmum is pentachotomocolpate or more rarely tetra-or hexachotomocolpate. Proencistemon flowers also differ from those of Hedyosmum in being delimited by a bract. In extant Hedyosmum the flowers are ebracteate and presumed to be unistaminate. There are several records of staminate inflorescences, inflorescence fragments and isolated stamens from the Early Cretaceous of Portugal that are similar to Proencistemon and that also contain chloranthoid pollen, but so far none of them has been named. Hedyosmum -like staminate inflorescences from the Torres Vedras locality (Friis et al. 2019a: text-fig. 20h) and from the Vale de Água locality (Friis et al. 2011: figs 8.13D, E, 16.2B) that have five whorls of staminate flowers differ in being ebracteate with smaller stamens, and have a larger number of stamens in each whorl than in Proencistemon. In addition, while the form of the pollen aperture in the single Torres Vedras specimen of a Hedyosmum -like staminate inflorescence is unknown, it is tetra- to pentachotomocolpate in the Vale de Água specimen. The tetra- to pentachotomocolpate pollen grains of the Vale de Água specimen are comparable to grains of Hedyosmum and to dispersed grains assigned to the extinct pollen genus Asteropollis. In contrast, Proencistemon has trichotomocolpate pollen. Although trichotomocolpate pollen grains are sometimes assigned to Asteropollis, the type material for Asteropollis asteroides R.W.HEDL. et G.NORRIS from the Early Cretaceous (Albian) of Oklahoma, USA, only includes forms with pentachotomocolpate, or rarely tetrachotomo- or hexachotomocolpate apertures. Trichotomocolpate grains are not recorded from the type locality (Hedlund and Norris 1968). Fossil pollen grains with a trichotomocolpate aperture similar to that of Proencistemon have sometimes been assigned to the extinct pollen genus Clavatipollenites COUPER (Doyle and Robbins 1977) or treated as aff. Clavatipollenites (Walker and Walker 1984). Dispersed trichotomocolpate pollen grains from the Early Cretaceous (Aptian) of Patagonia described as Jusinghipollis ticoensis M.LLORENS et LOINAZE (Llorens and Perez Loinaze 2015) are closely similar to the in situ pollen of Proencistemon, but the reticulum is looser, the muri are narrower, and the ornamentation of the tectum is coarser. The type species of Jusinghipollis JANSON. et HILLS, J. microreticulata (JUHÁSZ et GÓCZÁN) JANSON. et HILLS, which is based on dispersed pollen from the Albian of Hungary (Juhász and Góczán 1985), also has a trichotomocolpate aperture, but differs from pollen of Proencistemon in having longer aperture arms that almost reach to the equator. The aperture margins are also more well defined than the diffuse aperture margins in Proencistemon pollen grains., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 353, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Crane, P. R., Pedersen, K. R. (2019 a): The Early Cretaceous mesofossil flora of Torres Vedras (NE of Forte da Forca), Portugal: a palaeofloristic analysis of an early angiosperm community. - Fossil Imprint, 75: 153 - 257. https: // doi. org / 10.2478 / if- 2019 - 0013","Hedlund, R. W., Norris, G. (1968): Spores and pollen grains from Fredericksburgian (Albian) strata, Marshall Coun- ty, Oklahoma. - Pollen et Spores, 10: 129 - 159.","Doyle, J. A., Robbins, E. I. (1977): Angiosperm pollen zonation of the continental Cretaceous of the Atlantic coastal plain and its application to deep wells in the Salisbury embayment. - Palynology, 1: 41 - 78. https: // doi. org / 10.1080 / 01916122.1977.9989150","Walker, J. W., Walker, A. G. (1984): Ultrastructure of Lower Cretaceous angiosperm pollen and the origin and early evolution of flowering plants. - Annals of the Missouri Botanical Garden, 71: 464 - 521. https: // doi. org / 10.2307 / 2399035","Llorens, M., Perez Loinaze, V. S. (2015): Late Aptian angiosperm pollen grains from Patagonia: Earliest steps in flowering plant evolution at middle latitudes in southern South America. - Cretaceous Research, 57: 66 - 78. https: // doi. org / 10.1016 / j. cretres. 2015.07.019","Juhasz, M., Goczan, F. (1985): Comparative study of Albian monosulcate angiosperm pollen grains. - Acta Biologica Szegediensis, 31: 147 - 172."]}

Paisia pantoporata E.M.FRIIS, M.M.MENDES et K.R.PEDERSEN, 2018 Text-fig. 33a–c D e s c r i p t i o n a n d r e m a r k s. Paisia pantoporata is based on about 60 specimens of fossil flowers and isolated floral parts recovered from the Catefica mesofossil flora (for a full description and discussion see Friis et al. 2018a). Flowers are small, about 1 mm long and 1.2 mm in diameter, actinomorphic and pentamerous with a perianth consisting of a single whorl of five tepals, an androecium consisting of a single whorl of stamens, and a gynoecium of five free carpels (Text-fig. 33a, c). The organs of the three whorls are arranged on the same radius with the incurved margins of each tepal embracing a stamen (Text-fig. 33c). Stamens have a short, stout filament that merges above into the anther. Anthers are tetrasporangiate with two pairs of pollen sacs borne laterally and separated by a massive connective. Pollen grains are small, spheroidal, about 11–14 µm in diameter and pantoporate with six to eight pores. The pollen wall is tectate-punctate with spiny supratectal ornamentation (Text-fig. 33b). The carpels are follicular, narrowly elliptical to obovate with a ventral slit that extends for the full length of the carpel. Each carpel has one dorsal and two ventral bundles and contains about 20–30 seeds borne in two rows along the full length of the ventral suture. A f f i n i t y a n d o t h e r o c c u r r e n c e s. Pantoporate pollen occurs scattered among all major groups of angiosperms. However, the characters of the pollen combined with the pentamerous organisation indicate that Paisia pantoporata is most likely an extinct lineage of basal eudicots, probably most closely related to extant Ranunculales. Flowers of Paisia pantoporata have so far been recovered only from the Catefica locality. Pollen of Paisia pantoporata is also encountered in the palynological strew preparations from Catefica, where it occurs in clumps of several grains. Pantoporate pollen grains of the kind produced by Paisia pantoporata have not been reported from other palynofloras. Pantoporate pollen grains are not uncommon in Early Cretaceous palynofloras (for references see Ibrahim et al. 2015, Friis et al. 2018a), but grains similar to pollen of Paisia pantoporata have not been reported from other mesofossil floras or dispersed palynofloras., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 388, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801, {"references":["Friis, E. M., Mendes, M. M., Pedersen, K. R. (2018 a): Paisia, an Early Cretaceous eudicot angiosperm flower with pantoporate pollen from Portugal. - Grana, 57: 1 - 15. https: // doi. org / 10.1080 / 00173134.2017.1310292","Ibrahim, M. I. A., Zobaa, M. K., El-Noamani, Z. M., Tahoun, S. S. (2015): A review of the angiosperm pollen genus Cretacaeiporites Herngreen, with one new species from the Upper Cretaceous of Egypt. - Palynology, 41: 101 - 116. https: // doi. org / 10.1080 / 01916122.2015.1093561"]}

cf. Endressistemon sp. 2 Text-fig. 29c, d D e s c r i p t i o n a n d r e m a r k s. The material comprises a single stamen fragment, about 0.8 mm long including the apical projection, and about 0.3 mm broad. The apical projection is compressed covering the distal portions of the thecae (Text-fig. 29c). Pollen grains in situ are poorly exposed and the aperture configuration is uncertain, although it is clear that the grains are not tricolpate and we assume that the pollen is monoaperturate. Pollen is semitectate-reticulate, circular in equatorial view, about 9 µm in diameter. The reticulum is homobrochate and only loosely attached to the main body of the pollen (Text-fig. 29d). A f f i n i t y a n d o t h e r o c c u r r e n c e s. The stamen fragment is closely similar in general morphology to the individual stamens of Endressistemon cateficensis, but slightly larger and the reticulum of the pollen grains is more open and only loosely attached to the main body of the grains., Published as part of Friis, Else Marie, Crane, Peter R., Pedersen, Kaj Raunsgaard, Mendes, Mário Miguel & Kvaček, Jiří, 2022, The Early Cretaceous Mesofossil Flora Of Catefica, Portugal: Angiosperms, pp. 341-424 in Fossil Imprint 78 (2) on page 383, DOI: 10.37520/fi.2022.016, http://zenodo.org/record/7522801

info:eu-repo/semantics/publishedVersion

Premise of research. Accumulating data from phylogenetic analyses of living taxa and from paleobotany (pollen, leaves, and floral structures) has greatly improved our understanding of the Cretaceou...